Pollen morphology, exine structure and systematics of Acalyphoideae (Euphorbiaceae), Part 41

Review of Palaeobotany and Palynology 121 (2002) 231^336 www.elsevier.com/locate/revpalbo

Pollen morphology, exine structure and systematics of Acalyphoideae (Euphorbiaceae), Part 41 Tribes Acalypheae pro parte (Erythrococca, Claoxylon, Claoxylopsis, Mareya, Mareyopsis, Discoclaoxylon, Micrococca, Amyrea, Lobanilia, Mallotus, Deuteromallotus, Cordemoya, Cococceras, Trewia, Neotrewia, Rockinghamia, Octospermum, Acalypha, Lasiococca, Spathiostemon, Homonoia), Plukenetieae (Haematostemon, Astrococcus, Angostyles, Romanoa, Eleutherostigma, Plukenetia, Vigia, Cnesmone, Megistostigma, Sphaerostylis, Tragiella, Platygyna, Tragia, Acidoton, Pachystylidium, Dalechampia), Omphaleae (Omphalea), and discussion and summary of the complete subfamily Joan W. Nowicke a; , Masamichi Takahashi b a b

Department of Botany, NHB 166, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560, USA Department of Environmental Science, Faculty of Science, Niigata University, 8050, Ikarashi 2-no-cho, Niigata City 950-2181, Japan Received 22 December 2000; received in revised form 13 November 2001; accepted 18 January 2002

Abstract This is the fourth, last and largest installment of a palynological study of the Acalyphoideae that examined in light, scanning electron and transmission electron microscopy, the pollen of a grand total of 460 collections representing 372 species from 112 of the 116 assigned genera. Part 4 describes the pollen of tribe Acalypheae pro parte, subtribes Claoxylinae (Erythrococca, Claoxylon, Claoxylopsis, Mareya, Mareyopsis, Discoclaoxylon, Micrococca, Amyrea), Lobaniliinae (Lobanilia), Rottlerinae (Mallotus, Deuteromallotus, Cordemoya, Cococceras, Trewia, Neotrewia, Rockinghamia, Octospermum), Acalyphinae (Acalypha), Lasiococcinae (Lasiococca, Spathiostemon, Homonoia), tribe Plukenetieae, subtribe Plukenetiinae (Haematostemon, Astrococcus, Angostyles, Romanoa,

1 This study of pollen morphology and exine structure is dedicated to the original Euphorbiaceae palynologist, Willem Punt, who, using only light microscopy of a 1960 vintage, produced a lasting and remarkably accurate assessment of pollen of the entire family. * Corresponding author. Tel.: +1-202-786-2994; Fax: +1-202-786-2563. E-mail address: [email protected] (J.W. Nowicke).

0034-6667 / 02 / $ ^ see front matter 6 2002 Elsevier Science B.V. All rights reserved. PII: S 0 0 3 4 - 6 6 6 7 ( 0 2 ) 0 0 0 8 7 - 8

PALBO 2467 11-10-02

232

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Eleutherostigma, Plukenetia, Vigia); subtribe Tragiinae (Cnesmone, Megistostigma, Sphaerostylis, Tragiella, Platygyna, Tragia, Acidoton, (Pachystylidium); subtribe Dalechampiinae (Dalechampia) and tribe Omphaleae (Omphalea). None of the above multigeneric tribes or subtribes have uniform morphology or exine structure, but, with the notable exception of Plukenetieae, neither do they have any genera with outstanding palynological distinctions. Most members of Cloaxylinae have pollen of a generalized type, although the data support the separation of Mareyopsis from Mareya. Pollen of Amyrea is distinguished by a striate sculpture. Pollen of Mallotus is uniform and representative of the subtribe Rottlerinae: punctate-microspinulose tecta, thin foot layers, small columellae and thick continuous tecta. Pollen of Acalypha is small, brevicolp(or)ate, and mostly angulaperturate in polar view. One genus of subtribe Lasiococcinae, Spathiostemon, has a supratectal sculpture of lirae with cross striations, while a second, Homonoia, has a tectum composed of elongate, vertically oriented individual rods with acute tips. The exine structure of the large tribe Plukenetieae is strikingly diverse with most variants unique in the Acalyphoideae if not the family. Pollen of most members of Plukenetiinae is of either the Conophora type (punctate tecta, prominent columellae, and granular area under the tectum) or the Loretensis type (reticulate exines, crenate muri, unremarkable structure). Pollen of subtribe Tragiinae is more diverse than Plukenetiinae. Three Asian genera, Cnesmone, Megistostigma and Pachystylidium, have poorly defined apertures, and reduced exines. Inaperturate pollen is present in Platygyna, two species of Acidoton and Tragia sellowiana, all of which have exines consisting mostly of columellae. Three pollen types are recognized in Tragia: (1) the Ramosa type is three-colpate, intectate, and has exines of large columellae and thick foot layers; (2) the Lukafuensis type is three-colpate and has reticulate exines with arched colpal margins at the equator; (3) the Urens type has poorly defined apertures, is tectate with exines lacking foot layers. Pollen of Tragiella is very similar to the Lukafuensis type of Tragia. The large grains of the monogeneric subtribe Dalechampiinae are characterized by two prominent equatorial bands (costae). In Plukenetieae, the pollen data support: the reduction of three monotypic genera, Eleutherostigma, Romanoa and Vigia to species of Plukenetia; a close relationship among Cnesmone, Megistostigma and Pachystylidium; a congeneric treatment of Platygyna and Acidoton microphyllus and A. urens, or at least the restriction of Acidoton to these two species; separate generic status for Tragia sect. Tragia which has the Ramosa pollen type; a congeneric treatment of Tragiella and those species of Tragia (T. sect. Tagira) with the Lukafuensis pollen type; separate tribal status for Dalechampia. Pollen of Omphaleae is three-colpate, has irregular columellae and foot layers, and a non-apertural endexine that, after acetolysis, separates from the ectexine like the aperturate pollen of Crotonoideae. The results of the pollen study of the entire Acalyphoideae are discussed and Parts 1, 2 and 3 are summarized. Of the multigeneric tribes and subtribes, there are significant pollen differences among the taxa of Ampereae, of Chrozophoreae (subtribe Ditaxinae), of Pycnocomeae (subtribe Pycnocominae), and of Adelieae, as well as taxa of Plukenetieae. The nearly identical pollen of Ricininae and Adrianinae would support their treatment as one subtribe. The subtribes Mercurialinae and Dysopsidinae share similar pollen that is sufficiently distinct from the remaining Acalypheae to merit their treatment as one subtribe. Most pollen of Acalyphoideae is three-colporate with a lalongate endoaperture and punctate-microspinulose or punctate sculpture. Most exines consist of thin foot layers, modest columellae and continuous, mostly thick tecta. 6 2002 Elsevier Science B.V. All rights reserved. Keywords: Acalyphoidea; exine structure; pollen; systematics

1. Introduction This is the fourth and last paper of an extensive study of pollen morphology and exine structure of Acalyphoideae combining data from light, scanning electron and transmission electron microscopy (LM, SEM and TEM). Part 4 describes the pollen of Acalypheae pro parte (subtribes Claoxylinae, Lobaniliinae, Rottlerinae, Acalyphinae,

and Lasiococcinae) and tribes Plukenetieae and Omphaleae. Parts 1 (Nowicke et al., 1998), 2 (Nowicke et al., 1999) and 3 (Takahashi et al., 2000) were published in earlier volumes of the Review of Palaeobotany and Palynology. Part 2 and Part 3 summarized the results of the previous paper(s) as a convenience to the reader. Because this is the ¢nal paper, there is a discussion and summary of the results for the entire subfamily.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

For this reason, summaries of Parts 1, 2 and 3 are not given in the Introduction.

2. Materials and methods Pollen samples for all preparations, LM, SEM and TEM, were ¢rst acetolysed according to Erdtman (1966). The methodologies for LM, SEM and TEM are su⁄ciently common that we do not repeat them in Part 4. For greater detail the reader is advised to consult Parts 1 (Nowicke et al., 1998), 2 (Nowicke et al., 1999), or 3 (Takahashi et al., 2000). Accurate measurements of polar axes, equatorial diameters, colpus length, and to a lesser extent endoaperture dimensions, depend on expanded, intact grains. Pollen grains can be mechanically compromised by acetolysis ^ partial collapse and/or rupture of exines ^ making measurements suspect at best and erroneous at worst. In grains in which the organization of the apertures establishes polar axis and equatorial diameter, both measurements can be made in equatorial view. However, oblate-shaped grains tend to orient in polar view in glycerin jelly mounts, making the polar dimension unmeasurable. In many members of Plukenetieae, the frequently depauperate samples and collapsed/ruptured grains made measurements impossible for certain genera. In comparing our polar and equatorial dimensions of pollen of the tribe Plukenetieae with those of Gillespie (1994), ours were always smaller, but understandably so since she rehydrated £owers in 3% Aerosol-OT and then the isolated pollen grains were mounted in Hoyer’s medium for examination in LM. For three species, Astrococcus cornutus, Plukenetia conophora, and Tragiella natalensis, Gillespie also made an acetolysed preparation from the same collection. In all three species, the acetolysed grains were smaller, and the measurements did not even overlap except in the equatorial diameter of the second species and the polar axis of the third species, where the largest acetolysed measurement matched the smallest unacetolysed one. Another example of size disparity was found between grains mounted in silicone oil versus grains mounted in glycerin jelly of the

233

same collection of Mallotus eriocarpus (Table 2). The average polar dimension of grains was 17.5 Wm in silicon oil and 22.4 Wm in glycerin jelly. In those taxa of Acalyphoideae that have pollen with compound apertures, the endoapertures are frequently lalongate with di¡use lateral margins, and the equatorial measurements should be viewed with reserve. For all of the above reasons, pollen measurements, especially of Plukenetieae, should be viewed with reserve. It has come to our attention (Punt, personal communication) that the present authors’ concept of costa (ectocolpi and endocolpi) is more restricted than the de¢nition in Glossary of Pollen and Spore Terminology (Punt et al., 1994). The present authors do not consider any extension of the endexine into the mesocolpium as evidence of costae. It has been our experience that apertural endexines do not end abruptly at the margin of the colpus, instead, the endexine thins or tapers as it extends into the mesocolpium (e.g., Plates II, 12; XIX, 143; XX, 152). See also examples taken from convenient publications: Nowicke and Skvarla (1977, ¢gs. 148, 149, 152, 153, 157), Dickison et al. (1982, ¢gs. 36, 38, 65, 67^69), Nowicke (1994, ¢g. 95). This is not to say that we do not recognize costa, but rather that our experience has been with certain groups, e.g., Gronovioideae (Loasaceae) for one (Poston and Nowicke, 1994, ¢gs. 12, 18, 23, 30, 33, 37) and Panax L. (Araliaceae) for another (Wen and Nowicke, 2000, ¢gs. 26, 31, 34, 66^68) in which the pollen has the thickest endexine not under the apertures but under the margins. In LM, these particular grains of Loasaceae and Araliaceae have unmistakable and conspicuous dark streaks, costae ectocolpi, to each side of the colpus. Very few taxa of Acalyphoideae have such costae. The species examined, voucher data and ¢gure numbers, if illustrated, are given in Table 1. Pollen size data are given in Table 2. The distribution of selected pollen characteristics is given in Table 3. In the descriptions, characteristics and measurements based on LM and SEM are given ¢rst, those based on TEM are given second. The measurements given in the descriptions represent the range of dimensions as given in Table 2. The ter-

PALBO 2467 11-10-02

234

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Table 1 Species examined, voucher data and ¢gure numbers if illustrated Taxon

Collection

Plates and ¢gures

S.G. Adam 29225 (MO), French Guinea Witte 11704 (MO), Zaire Bequaert 5282, Zaire Du«mmer 1413, Uganda Graham 2123 (MO), Kenya Gereau 1491 (MO), Tanzania Schlieben 7595, South Africa Bequaert 3209, Zaire Troupin 6263, Zaire Schweinfurth and Riva 1037, Eritrea

Plate I, 1^3 Plate I, 4^7

Tribe 30. Acalypheae Subtribe 30g. Claoxylinae 168. Erythrococca E. africana Prain E. bongensis Pax E. hirta Pax E. kirkii (Mueller Argoviensis) Prain E. menyharthii Prain E. oleracea Prain E. welwitschiana Prain Erythrococca species indet. 169. Claoxylon C. albicans Merrill C. australe Baillon C. cf. bakerianum Baillon C. hainanense Pax et K. Ho¡mann C. insulanum Mueller Argoviensis

Plate I, 8, 9

Plate Plate Plate Plate

II, 10^12 II, 13^15 III, 16^18 III, 19^21

C. longifolium (Blume) Miquel C. lutescens Pax et K. Ho¡mann C. sitibundum Croizat

Ramos and Edano 2-107, Philippines White 12725 (AA), Australia Schatz et al., 1687 (MO), Madagascar Tak 190 (MO), China Balansa 269 (MO), New Caledonia McKee 196 (AA), New Caledonia Rahmat Si Toroes 4652, Sumatra Brass 31619 (AA), New Guinea Smith 7524, Fiji

Plate Plate Plate Plate Plate Plate Plate

170. Claoxylopsis C. purpurescens A. Radcli¡e-Smith

Perrier de la Bathie 17400 (K), Madagascar

Plate VI, 40^42, 42 inset

171. Mareya M. micrantha (Bentham) Mueller Argoviensis M. spicata Baillon

Baldwin 9239 (MO), Liberia Cooper 404, Liberia

Plate VII, 43^45 Plate VII, 46^48

M. oligogyna

Zenker 4228 (MO), Cameroon Zenker 35, Cameroon McPherson 16174, Gabon

Plate VIII, 49, 51 Plate VIII, 50, 52^55, Plate IX, 56 Plate IX, 57

172. Discoclaoxylon D. hexandrum (Mueller Argoviensis) Pax et K. Ho¡mann

Baldwin 6696 (MO), Liberia Nemba and Mamba 780 (MO), Cameroon

Plate IX, 58, 61 Plate IX, 59, 60

173. Micrococca M. capensis Prain M. holstii (Pax) Prain M. mananarenta Leandri M. oligandra (Mueller Argoviensis) Prain

Gerstner 2403 (MO), South Africa Agnew et al. 9600 (MO), Kenya D’Alleizette s.n. (L), Madagascar Kostermans 26309 (L), India

Plate Plate Plate Plate

Miller 3702 (MO), Madagascar Humbert 6599, Madagascar Capuron s.n. 27/10/1954 (MO), Madagascar

Plate XI, 73, 74, 76, 77 Plate XI, 72, 75 Plate XI, 78^80

Benoist 1302 (K), Madagascar Humbert 18324 (K), Madagascar

Plate XII, 81^84 Plate XII, 85^89

171a. Mareyopsis longifolia Pax

174. Amyrea A. humbertii Leandri A. sambiranensis Leandri

Plate IV, 22^25 IV, 26^28 IV, 29, 30 V, 31 V, 32 V, 33^35 V, 36 VI, 37^39

X, X, X, X,

62^64 65 66^68 69^71

Subtribe 30h. Lobaniliinae 175. Lobanilia L. bakeriana (Baillon) A. Radcli¡e-Smith L. claoxyloides A. Radcli¡e-Smith

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336 Table 1 (Continued). Taxon

Collection

Plates and ¢gures

Dr. King’s Collector s.n. x/x/1884, Andaman Islands Yao 10366 (MO), China Tsang 27858 (AA), China Nicoll et al. 550 (MO), Madagascar Henry 9525B (AA), China Sumithraarach-chi s.n. 9/18/1974 (MO), Sri Lanka Kostermans 25522 (L), Sri Lanka Fosberg 37783, Ryukyu Islands Alvarez 23743, Philippines Achmad 1208 (L), Sumatra Britton and Winder69 (AA), Tasmania Tsang 28024 (AA), China Petelot 1945, Vietnam SAN (D.I.N.) 34757 (L), Malaysia Wilson 3292 (AA), China Brass 19761 (AA), Australia Fang 16497 (AA), China Dickason 6641 (MO), Burma Ridsdale 33560 (L), Papua New Guinea W. Beer’s Collectors 6823 (L), Solomon Islands Nemba and Thomas 357 (MO), Cameroon Main 1903 (L), Borneo

Plate XIII, 90^92

Subtribe 30i. Rottlerinae 176. Mallotus M. acuminatus Mueller Argoviensis M. apelta Mueller Argoviensis M. baillonianus Mueller Argoviensis M. barbatus (Wallich) Mueller Argoviensis M. eriocarpus (Thwaites) Mueller Argoviensis M. M. M. M. M. M. M. M. M. M. M. M. M. M. M. M.

fuscescens (Thwaites) Mueller Argoviensis japonicus Mueller Argoviensis korthalsii Mueller Argoviensis leucodermis Hooker f. leucophyllus Mueller Argoviensis microcarpus Pax et K. Ho¡mann mollissimus (Geiseler) Airy Shaw muticus (Mueller Argoviensis) Airy Shaw nepalensis Mueller Argoviensis nesophilus Mueller Argoviensis oreophilus Mueller Argoviensis philippinensis (Lamarck) Mueller Argoviensis polyadenos F.v. Mueller puber Bollendor¡ subulatus Mueller Argoviensis sumatranus (Miquel) Airy Shaw

177. Deuteromallotus D. acuminatus (Baillon) Pax et K. Ho¡mann Deuteromallotus species indet. 178. Cordemoya C. integrifolia Baillon

179. Coccoceras 180. Trewia T. nudi£ora L.

181. Neotrewia N. cumingii (Mueller Argoviensis) Pax et K. Ho¡mann

182. Rockinghamia R. angustifolia (Bentham) Airy Shaw R. brevipes Airy Shaw 183. Octospermum O. pleiogynum (Pax et K. Ho¡mann) Airy Shaw

Plate Plate Plate Plate Plate

XIII, XIII, XIII, XIII, XIV,

93 94 95 96^98 99^101

Plate Plate Plate Plate Plate Plate Plate

XIV, 102, 103 XIV, 104^106 XIV, 107 XV, 108, 109 XV, 113 XV, 110^112 XV, 114

Plate XV, 115 Plate XVI, 116^119 Plate XVI, 120

Plate XVI, 121, 122 Plate XVI, 123

Nicoll et al. 500 (MO), Madagascar Decary 16884, Madagascar Schatz and Modesle 3133 (MO), Madagascar

Plate XVII, 125 Plate XVII, 126^128 Plate XVII, 124

Lorence 2231 (MO), Mauritius Goode et al. 3952 (K), Mauritius

Plate XVII, 129^131 Plate XVII, 132

No material

Road s.n. 4/4/1929 (MO) (Country unknown) Wenzel 71 (MO), Philippines Fernandes 31 (K), India

Plate XVIII, 133 Plate XVIII, 135, 137 Plate XVIII, 134, 136

Elmer 17641 (MO), Philippines

Plate XVIII, 139, 141

Sulit 22901 (K), Philippines

Plate XVIII, 138, 140

Gray 376 (K), Australia Hyland 7778 (K), Australia

Plate XIX, 142^146 Plate XIX, 147^150

Streimann and Katik NGF 28607 (K), Papua New Guinea Kostermans 287 (K), Papua New Guinea

Plate XX, 151^153

PALBO 2467 11-10-02

Plate XX, 154^156

235

236

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Table 1 (Continued). Taxon

Collection

Plates and ¢gures

Wilson 1987, China Schulz 15213 (MO), Argentina Alverson and DiStilio 2581, Costa Rica

Plate XXI, 157 Plate XXI, 158, 159

Subtribe 30j. Acalyphinae 184. Acalypha A. acmophylla Hemsley A. communis Mueller Argoviensis A. costaricensis (Kuntze) Knoblauch ex Pax et K. Ho¡mann A. diversifolia Jacquin A. grantii Baker et Hutchinson A. lanceolata Willdenow

Hamilton et al. 3299 (MO), Panama Dummer 5435 (AA), Uganda Wheeler 12945, Sri Lanka Reeder 876, New Guinea Tharp 2275, USA Synnott 831 (MO), Uganda Braun s.n., 27 Aug 1917, USA Bayliss 7191 (AA), South Africa Bayliss 3015 (AA), South Africa Tenorio 1555 (MO), Mexico Wenzel 1507? (MO), Philippines Yuncker 3107, Hawaii

Plate Plate Plate Plate

185. Lasiococca L. symphylliifolia Hooker f. (Kurz)

H.D.C. s.n., 6-4-1887 (L), India

Plate XXV, 189, 190

186. Spathiostemon S. javensis Blume

Ramos 17661 (MO), Philippines Beaman et al. 8860 (MO), Borneo

Plate XXV, 196, 197, Plate XXVI, 198 Plate XXV, 191^195

Huang 2459 (MO), Taiwan McGregor 247, Philippines

Plate XXVI, 199, 200 Plate XXVI, 201^203

188. Haematostemon H. guianensis Sandwith

Fanshawe 2869, British Guiana

Plate XXVII, 204^207

189. Astrococcus A. cornutus Bentham

Liesner 8693 (MO), Venezuela

Plate XXVII, 208, 209

190. Angostyles A. longifolia Bentham

Ducke 23528, Brazil

Plate XXVIII, 210^213

Trinta 533 and Fromm 1609, Brazil Harshberger 833, Brazil

Plate XXVIII, 214, 215, 217 Plate XXVIII, 216

Cazalet and Pennington 5089, Ecuador Lawrence 276 (MO), Colombia

Plate XXIX, 218 Plate XXIX, 219^222

Vargas 18799, Peru Zenker 3311, Cameroon Liesner 3788 (MO), Venezuela Schunke 3686, Peru McPherson 8461 (MO), Panama

Plate Plate Plate Plate Plate

A. A. A. A. A. A. A. A.

monococca (A. Gray) L. Miller et K.N. Gandhi ornata Hochstetter ex A. Richard ostryaefolia Riddelsdell peduncularis (S. Moore) Meissner ex Krauss schinzii Pax setosa A. Richard stipulacea Klotzsch wilkesiana Mueller Argoviensis

Plate XXI, 160^162 Plate XXI, 163, 164 Plate XXII, 165^167 Plate XXII, 168^170 Plate XXII, 171^173

XXIII, XXIII, XXIII, XXIV,

174^176 177^179 180^182 183^188

Subtribe 30k. Lasiococcinae

187. Homonoia H. riparia Loureiro

Tribe 31. Plukenetieae Subtribe 31a. Plukenetiinae

191. Romanoa R. tamnoides (A. Jussieu) A. Radcli¡e-Smith

192. Eleutherostigma E. lehmannianum Pax et K. Ho¡mann

193. Plukenetia P. brachybotrya Mueller Argoviensis P. conophora Mueller Argoviensis P. loretensis Ule P. penninervia Mueller Argoviensis

PALBO 2467 11-10-02

XXIX, 223, 224 XXX, 227^230 XXX, 232, 234 XXX, 231, 233, 235 XXXI, 236^238

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

237

Table 1 (Continued). Taxon

Collection

Plates and ¢gures

P. polyadenia Mueller Argoviensis

Mexia 6470 (MO), Peru Ducke 20619, Brazil Liesner 2068 (MO), Costa Rica Asplund 14129, Peru

Plate Plate Plate Plate

Pederson 8790 (K), Brazil Thomas et al., 10221, Brazil

Plate XXXI, 243, 244

Stone and Sidek 12536 (MO), Malaya Pe¤telot 6521, Vietnam

Plate XXXII, 245, 247 Plate XXXII, 246, 248

P. volubilis L.

194. Vigia V. serrata Vellozio

XXIX, XXIX, XXXI, XXXI,

225 226 240, 242 239, 241

Subtribe 31b. Tragiinae 195. Cnesmone C. javanica Blume C. tonkinensis (Gagnepain) Croizat 196. Megistostigma M. burmanicum (Kurz) Airy Shaw M. cordatum Merrill M. malaccense (Hooker f.) Pax

197. Sphaerostylis 198. Tragiella T. friesiana (Prain) Pax et K. Ho¡mann T. natalensis (Sonder) Pax et K. Ho¡mann

199. Platygyna P. dentata Alain P. hexandra (Jacquin) Mueller Argoviensis P. parvifolia Ala|¤n 200. Tragia T. bailloniana Mueller Argoviensis T. cf. cordifolia Bentham T. gardneri Prain T. lukafuensis Willdenow T. mitis Hochst ex A. Richard T. pungens F. Mueller T. ramosa Torrey T. sellowiana (Klotzsch) Mueller Argoviensis T. smallii Shinners T. spathulata Bentham T. stylaris Mueller Argoviensis T. urens L. T. urticaefolia Michaux T. volubilis L. 201. Acidoton A. microphyllus Urban A. nicaraguensis (Hemsley) Webster

Maxwell 89-1034 (L), Malaysia Argent et al., 108380 (L), Malaysia Chan 5/9/71 (L), Malaysia Rahmat Si Toroes 1975, Sumatra Sinclair 8025, Malaya

Plate XXXII, 249, 249 inset, 250 Plate XXXIII, 251

No material

Richards 13208 (K), Rhodesia Kemp 741 (MO), Swaziland Mearns 255, Kenya

Plate XXXIII, 252^255 Plate XXXIII, 256, 258 Plate XXXIII, 257

Morton and Acu‹na 3692, Cuba Curtiss s.n. 12/8/1904 (MO), Cuba Morton 10253, Cuba Shafer 1427, Cuba

Plate Plate Plate Plate

XXXIV, XXXIV, XXXIV, XXXIV,

Stevens 24383 (MO), Costa Rica Schlieben 7556, South Africa Rutherford-Smith 228 (K), Zimbabwe Angus 718 (K), Zimbabwe Gibert and Thulin 689 (MO), Ethiopia Pappi 1065, Eritrea Herrick 233, USA Nee 39396, Bolivia Palmers 7705, USA Breteler 7203, Togo Sperry 74, USA Merrill 584, USA Thomas 104979 (MO), USA Webster and Wilbur 3231, USA Klein 5707, Brazil

Plate Plate Plate Plate Plate Plate Plate Plate

XXXV, 268^270 XXXV, 271^273 XXXV, 274^276 XXXVI, 277^280 XXXVI, 281, 282 XXXVII, 287^290 XXXVI, 283^286 XXXVII, 293^295, 295 inset

Plate Plate Plate Plate Plate Plate

XXXVII, 291, 292 XXXVII, 296 XXXVIII, 300^303 XXXVIII, 297, 299 XXXVIII, 298 XXXVIII, 304, 305

Plate Plate Plate Plate

XXXIX, XXXIX, XXXIX, XXXIX,

Liogier 13627, Dominican Republic Ortiz 1104, Nicaragua Standley and Valerio 46475, Costa Rica Moreno 17228 (DAV), Nicaragua Espinoza 188 (DAV), Costa Rica

PALBO 2467 11-10-02

259 261 260 262^267

309^311 313 314, Plate XL, 316 312, Plate XL, 315

238

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Table 1 (Continued). Taxon

Collection

Plates and ¢gures

A. urens Swartz

Proctor 36826, Jamaica Alexander s.n., Jamaica Woytkowski 7527, Peru

Plate XXXIX, 306^308

Ramos 40929 (MO), Philippines Van Balgooy 2974 (L), Celebes

Plate XL, 320 Plate XL, 317^319

Pederson 12236 (MO), Brazil Bayliss 5294, South Africa Steyermark 38062, Guatemala Humbert II692, Madagascar Allen 5342, Costa Rica Liesner 2108, Costa Rica Hernger and Rizzini 17544, Brazil Grijalva 752 (MO), Nicaragua Pederson 8333 (MO), Argentina Humbert 7080, Madagascar Antonio 3296 (MO), Panama

Plate Plate Plate Plate Plate Plate

XLI, 321 XLI, 322^325 XLI, 326 XLI, 327^330 XLII, 331 XLII, 332^336

Plate Plate Plate Plate

XLII, 337^339 XLIII, 340, 341 XLIII, 342^345 XLIII, 346^348

Acidoton species indet. 202. Pachystylidium P. hirsutum (Blume) Pax et K. Ho¡mann

Subtribe 31c. Dalechampiinae 203. Dalechampia D. brevipes Mueller Argoviensis D. capensis Sprengel f. D. cissifolia Poeppig et Endlicher D. clematidifolia Bojer ex Baillon D. dioscoreifolia Poeppig et Endlicher D. heteromorpha Pax et K. Ho¡mann D. humilis Mueller Argoviensis D. scandens L. D. stenosepala Mueller Argoviensis D. subternata Mueller Argoviensis D. tiliifolia Lamarck Tribe 32. Omphaleae 204. Omphalea O. bracteata Merrill O. diandra L. O. oleifera Hemsley O. oppositifolia (Willd.) L.J. Gillespie O. palmata Leandri O. triandra L. O. trichotoma Mueller Argoviensis

Violal 2702, Laos Palacios 6823 (MO), Ecuador Irwin et al. 48498, Brazil Manriquez 2334 (MO), Mexico Manriquez 443 (MO), Mexico Gillespie 4014 (MO), Madagascar Malcomber 2832 (MO), Madagascar Phillipson 2266 (MO), Madagascar Holdridge 2077, Haiti Britton et al. 6238, Cuba

Plate XLIV, 349 Plate XLIV, 351 Plate XLIV, 350 Plate XLIV, 352 Plate Plate Plate Plate

XLIV, 353 XLIV, 354 XLV, 355^357 XLV, 358^360

All collections USA unless otherwise noted.

minology follows, for the most part, that of Punt et al. (1994), but with occasional recourse to that of Erdtman (1966). The sequence of tribes, subtribes and included genera as well as geographical distributions are taken from Webster (1994).

3. Results Tribe 30. Acalypheae pro parte (Plates I^XXVI) Subtribe 30g. Claoxylinae (Plates I^XI) 168. Erythrococca Bentham, 50 species in Africa. (Plates I^III, 1^21). Species examined: E. africana (Plate I, 1^3) ; E. bongensis (Plate I, 4^7); E. hirta (Plate I, 8, 9);

E. kirkii; E. menyharthii (Plate II, 10^12); E. oleracea (Plate II, 13^15); E. welwitschiana (Plate III, 16^18); Erythrococca species indet. (Plate III, 19^ 21). Pollen prolate spheroidal to subprolate in equatorial view; PUE = (15.0^24.7)U(13.6^22.1) Wm; three- to ¢ve-colporate, the colpi 7.8^15.6 Wm long, mostly narrow, costa ectocolpi weakly developed, margo absent, operculum absent; the endoaperture mostly lalongate, PUE = ( 6 1.0^ 3.9)U(3.9^11.7) Wm, sometimes the lateral margin unclear, in one species (E. welwitschiana) the endoaperture endocingulate or nearly so; the tectum punctate-microspinulose (e.g., Plate I, 2; Plate II, 11, 14) or more rarely weakly microscabrate (Plate III, 19). In thin section, the mesocolpial

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

239

Table 2 Pollen size data based on an average of 10 acetolysed grains mounted in glycerin jelly Taxon

Pollen size

Colpus length

Polar

Equatorial

P/E

(17.5)-18.5-(21.3)

(15.5)-17.3-(19.4)

1.07

No LM preparation (15.0)-15.5-(16.0) (13.6)-15.3-(15.5) (19.5)-21.1-(22.1) (18.2)-19.2-(20.8) No LM preparation No LM preparation (18.2)-19.6-(20.8) (18.2)-20.7-(22.1) (22.1)-23.1-(24.7) (15.6)-17.1-(18.2)

Endoaperture Polar

Equatorial

(9.7)-12.8-(15.5)

(1.3)-2.7-(3.9)

(6.5)-7.8-(9.1)

1.01 1.09

(7.8)-10.1-(11.6) (13.6)-14.7-(5.5)

(2.6)-2.6-(2.6) less than 1.0

(7.8)-9.8-(11.7) unclear margin 4-col.

(15.6)-17.1-(18.2) (16.9)-18.2-(19.5) (19.5)-20.3-(22.1) (16.9)-18.0-(19.5)

1.14 1.14 1.14 0.95

(13.0)-14.0-(15.6) (10.4)-12.2-(13.0) (10.4)-12.9-(14.3) (7.8)-9.1-(10.4)

(2.6)-2.9-(3.9) (1.3)-2.5-(2.6) (1.3)-2.1-(2.6) (1.3)-1.3-(1.3)

(3.9)-4.5-(5.2) (6.5)-6.8-(7.8) zonorate? (3.9)-4.4-(5.2)

Tribe 30. Acalypheae Subtribe 30g. Claoxylinae 168. Erythrococca E. africana E. bongensis Bequaert 5282 Witte 11704* E. hirta E. kirkii Gereau 1491 Graham 2123 E. menyharthii E. oleracea E. welwitschiana Erythrococca species indet. 169. Claoxylon C. albicans C. australe C. cf. bakerianum C. hainanense* C. insulanum Balansa 269* McKee 196 C. longifolium C. lutescens C. sitibundum

(21.3)-23.2-(25.2) (19.4)-20.7-(21.3) (21.3)-21.7-(23.2) (19.5)-20.0-(20.8)

(19.4)-21.2-(23.2) (17.5)-18.5-(19.4) (19.4)-20.4-(21.3) (19.5)-20.3-(20.8)

1.09 1.12 1.06 0.98

(15.5)-17.3-(19.4) (15.5)-16.7-(17.5) (7.8)-13.4-(15.5) (7.8)-9.8-(11.7)

(2.0)-2.9-(3.9) less than 1.0 (1.3)-2.1-(2.6) (1.3)-2.1-(2.6)

(5.8)-7.2-(9.7) unclear margin (10.4)-12.0-(13.0) (5.2)-7.7-(10.4)

(15.6)-17.3-(18.2) (17.5)-18.6-(19.4) (15.6)-16.0-(16.9) (17.5)-18.6-(21.3) (19.4)-22.5-(23.2)

(15.6)-16.6-(16.9) (13.6)-17.1-(19.4) (16.9)-17.4-(18.2) (13.6)-16.5-(17.5) (19.4)-21.9-(23.2)

1.04 1.09 0.92 1.12 1.03

(11.6)-12.2-(14.0) (11.6)-13.4-(15.5) (7.8)-9.0-(9.7) (11.6)-13.4-(15.5) (13.6)-15.5-(17.5)

(1.5)-1.8-(2.0) (1.5)-2.6-(3.9) (1.6)-1.8-(2.5) (1.9)-2.3-(3.9) (1.2)-1.8-(2.2)

(3.9)-4.9-(7.8) (3.9)-6.8-(7.8) (3.9)-5.7-(9.7) (5.8)-7.2-(9.7) (3.9)-5.8-(9.7)

170. Claoxylopsis C. purpurescens (smaller grains, n = 7) (larger grains, n = 8)

(16.9)-17.8-(18.2) (20.8)-22.6-(23.4)

(14.3)-16.9-(19.5) (16.9)-18.2-(19.5)

1.05 1.25

(10.4)-13.0-(14.3) (15.5)-16.7-(17.5)

less than 1.0 less than 1.0

less than 1.0 less than 1.0

171. Mareya M. micrantha* M. spicata

(14.3)-14.8-(15.6) (15.6)-16.0-(16.9)

(14.3)-14.6-(15.6) (14.3)-15.3-(16.9)

1.01 1.04

(9.7)-10.3-(11.6) (10.4)-12.0-(13.0)

less than 1.0 unclear

less than 1.0

171a. Mareyopsis M. longifolia Zenker 35 Zenker 4228* M. oligogyna

(18.2)-19.1-(19.5) (14.3)-14.4-(16.9) (16.9)-17.7-(18.2)

(19.5)-20.8-(22.1) (14.3)-15.5-(16.9) (18.2)-18.9-(19.5)

0.92 0.93 0.94

(7.8)-9.6-(10.4) (9.7)-10.1-(11.6) (10.4)-11.2-(11.7)

(1.3)-1.3-(1.3) (1.3)-1.3-(1.3) less than 1.0

(7.8)-8.3-(10.4) (6.5)-6.9-(7.8) (5.2)-6.5-(7.8)

172. Discoclaoxylon D. hexandrum Baldwin 6696* (15.6)-16.8-(16.9) Nemba and Mamba 780 (15.6)-16.6-(16.9)

(15.6)-16.8-(16.9) (16.9)-17.4-(18.2)

1.00 0.95

(7.8)-9.7-(11.6) (7.8)-8.6-(9.1)

less than 1.0 (1.3)-1.7-(2.6)

(5.8)-7.0-(7.8) (3.9)-5.2-(6.5)

173. Micrococca M. capensis* M. holstii* M. mananarenta M. oligandra

(16.9)-17.9-(19.5) (16.9)-17.7-(18.2) (15.6)-16.8-(18.2) (20.8)-23.1-(26.0)

(15.6)-17.2-(18.2) (16.9)-18.1-(19.5) (15.6)-17.1-(18.2) (19.5)-21.3-(23.4)

1.04 0.98 0.98 1.08

(9.7)-12.2-(13.6) (11.6)-12.4-(13.6) (7.8)-12.0-(13.0) (14.3)-16.1-(19.5)

less than 1.0 less than 1.0 (1.3)-2.5-(2.6) (2.6)-5.1-(6.5)

(5.8)-8.0-(9.7) 4-col. (5.8)-6.4-(7.8) (6.5)-7.9-(9.1) (9.1)-11.0-(14.3)

(16.9)-17.9-(19.5) (grains collapsed)

(15.6)-16.9-(18.2)

1.06

(13.0)-13.8-(15.6)

less than 1.0

less than 1.0

174. Amyrea A. humberti Humbert 6599

PALBO 2467 11-10-02

240

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Table 2 (Continued). Taxon

Pollen size

Colpus length

Endoaperture

Polar

Equatorial

P/E

Miller 3702 A. sambiranensis

(22.1)-22.7-(24.3) (19.5)-20.7-(22.1)

(20.8)-21.3-(23.4) (19.5)-19.6-(20.8)

Polar

Equatorial

1.06 1.06

(13.6)-16.1-(17.5) (11.7)-13.3-(14.3)

less than 1.0 (1.3)-1.7-(2.6)

less than 1.0 (3.9)-4.7-(5.2)

(23.4)-25.3-(27.3) (19.4)-22.1-(23.2)

(20.8)-21.5-(23.4) (17.5)-19.5-(21.3)

1.17 1.13

(16.9)-18.8-(23.4) (13.6)-16.9-(19.4)

(2.6)-3.9-(5.2) (1.8)-2.1-(2.5)

(6.5)-8.8-(11.7) (3.9)-5.8-(7.8)

(18.2)-19.5-(20.8)

(18.2)-20.1-(23.4)

0.97

(10.4)-12.2-(13.0)

(1.3)-1.4-(2.6)

(2.6)-3.6-(5.2)

(17.5)-19.4-(21.3) (18.2)-19.2-(19.5) (16.9)-17.8-(18.2) (21.3)-23.2-(25.2)

(19.4)-20.7-(23.2) (19.5)-20.2-(22.1) (18.2)-19.8-(22.1) (23.2)-24.8-(27.2)

0.94 0.94 0.89 0.94

(11.6)-13.4-(15.5) unclear (7.8)-8.1-(11.6) (15.5)-18.6-(21.3)

(1.5)-3.5-(4.2)

(^)-9.7-(^)

less than 1.0 (2.5)-3.8-(4.2)

(5.8)-6.8-(7.8) (9.7)-12.6-(15.5)

(15.5)-17.5-(19.4)

(15.5)-17.5-(19.4)

1.00

(11.6)-13.0-(13.6)

(1.5)-2.0-(3.9)

(5.8)-7.8-(9.7)

(19.5)-22.4-(23.4) (16.9)-18.6-(19.5) (21.3)-23.0-(23.2) (15.6)-17.3-(19.5) (many grains collapsed) (20.8)-23.4-(24.7) (17.5)-19.8-(21.3) (17.5)-19.3-(21.3) (20.8)-21.7-(22.1) (many grains collapsed) (20.8)-25.5-(28.6) (23.2)-25.2-(27.2) (15.5)-16.1-(17.5) (24.9)-26.3-(27.3) (11.6)-13.4-(15.5) (24.7)-26.3-(28.6) (16.9)-18.1-(19.5) (26.0)-27.7-(31.2) (many small, collapsed grains) (23.4)-25.1-(28.6)

(19.5)-21.6-(22.1) (18.2)-19.1-(19.5) (23.2)-25.3-(27.2) (16.9)-17.7-(19.5)

1.03 0.97 0.91 0.98

(13.0)-13.8-(14.3) (7.8)-9.4-(10.4) (13.6)-15.3-(17.5) (9.1)-9.9-(11.7)

(1.3)-2.6-(3.9) (2.6)-3.0-(3.9) (1.0)-1.8-(2.5) (1.3)-2.0-(2.6)

(5.2)-6.8-(7.8) (7.8)-8.8-(10.4) (9.7)-10.7-(13.6) (2.6)-3.6-(3.9)

(22.1)-24.7-(26.0) (19.4)-20.5-(21.3) (17.5)-20.2-(21.3) (20.8)-22.1-(23.4)

0.95 0.97 0.96 0.98

(7.8)-10.1-(11.7) (11.6)-13.8-(15.5) (9.7)-12.4-(13.6) (10.4)-11.3-(13.0)

(1.3)-2.1-(3.9) (1.9)-3.3-(4.2) less than 1.0 (1.3)-1.3-(1.3)

(3.9)-6.5-(9.1) (7.8)-9.2-(13.6) (5.8)-8.0-(9.7) (5.2)-6.9-(9.1)

(24.7)-27.4-(29.9) (21.3)-23.5-(27.2) (15.5)-16.5-(17.5) (23.4)-25.8-(26.0) (13.6)-15.5-(17.5) (23.4)-27.0-(28.6) (16.9)-18.8-(20.8) (26.0)-27.3-(28.6)

0.93 1.07 0.98 1.02 0.86 0.97 0.96 1.01

(9.1)-10.1-(11.7) (17.5)-19.2-(21.3) (9.7)-10.9-(13.6) (13.6)-17.5-(19.4) (5.8)-7.2-(9.7) (9.1)-9.5-(10.4) (7.8)-8.8-(9.1) (19.4)-20.0-(23.2)

(2.6)-3.2-(5.2) less than 1.0 less than 1.0 (1.9)-2.6-(3.9) less than 1.0 (2.6)-3.4-(3.9) (1.3)-2.5-(2.6) (3.9)-4.0-(4.5)

(5.2)-6.9-(9.1) less than 1.0 less than 1.0 (9.1)-10.8-(14.3) (4.5)-5.8-(6.2) (5.2)-6.2-(6.5) unclear (5.8)-9.1-(13.6)

(24.7)-25.6-(27.3)

0.98

(7.8)-9.2-(11.7)

(5.2)-5.6-(6.5)

(6.5)-7.5-(9.1)

(14.3)-19.2-(22.1) (13.6)-15.7-(17.5) (17.5)-19.1-(19.4)

0.89 0.94 0.85

(5.2)-7.1-(10.4) (7.8)-8.6-(9.7) (5.8)-8.6-(11.6)

(1.3)-1.7-(2.6) less than 1.0 (1.5)-2.1-(2.5)

(6.5)-8.3-(10.4) (3.9)-7.0-(9.7) (5.8)-8.4-(11.6)

(20.8)-21.6-(22.1) (15.5)-16.9-(17.5)

0.91 0.85

(7.8)-9.6-(13.0) (5.8)-6.3-(7.8)

(2.6)-2.6-(2.6) (1.5)-1.7-(2.2)

(7.8)-9.1-(10.4) (5.8)-7.4-(9.7)

Subtribe 30h. Lobaniliinae 175. Lobanilia L. bakeriana L. claoxyloides Subtribe 30i. Rottlerinae 176. Mallotus M. acuminatus M. apelta Tsang 27858 Yao 10366* (n = 4) M. baillonianus M. barbatus M. eriocarpus Sumithraarachchi 9/18/ 74* Sumithraarachchi 9/18/74 M. fuscescens M. japonicus* M. korthalsii

M. M. M. M.

leucodermis leucophyllus microcarpus mollissimus

M. M. M. M. M. M. M. M.

muticus nepalensis nesophilus oreophilus philippinensis* polyadenos puber subulatus

M. sumatranus

177. Deuteromallotus D. acuminatus Decary 16884 (14.3)-17.1-(22.1) Nicoll et al. 500* (13.6)-14.7-(15.5) Deuteromallotus species in- (15.5)-16.1-(17.5) det. 178. Cordemoya C. integrifolia Goode et al. 3952 Lorence 2231*

(18.2)-19.6-(20.8) (13.6)-14.4-(15.5)

179. Coccoceras

No material

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336 Table 2 (Continued). Taxon

Pollen size

Colpus length

Polar

Equatorial

P/E

180. Trewia T. nudi£ora Fernandes 31 Road s.n. 4/4/1929* Wenzel 71*

(24.7)-26.7-(29.9) (19.4)-20.7-(21.3) (15.5)-15.7-(17.5)

(24.7)-27.4-(29.9) (21.3)-24.0-(27.2) (13.6)-15.1-(17.5)

0.97 0.86 1.04

181. Neotrewia N. cumingii Elmer 17641 Sulit 22901

No LM preparation (18.2)-19.0-(20.8) (18.2)-19.4-(20.8)

182. Rockinghamia R. angustifolia R. brevipes

(21.3)-23.6-(25.2) (23.2)-25.0-(27.2)

(17.5)-18.3-(19.4) (17.5)-19.2-(19.4)

183. Octospermum O. pleiogynum Kostermans 287 Streimann and Katik 28607

Endoaperture Polar

Equatorial

(10.4)-13.8-(15.6) (9.7)-10.1-(11.6) (9.7)-10.7-(11.6)

(2.6)-3.9-(5.2) (1.9)-3.0-(3.9) (1.5)-2.0-(2.2)

unclear margins (1.8)-3.9-(5.8) (3.9)-5.4-(7.8)

0.98

(9.1)-11.2-(13.0)

(1.3)-1.9-(2.6)

(6.5)-8.3-(10.4)

(19.4)-20.2-(21.3) (21.3)-24.0-(29.1)

1.17 1.04

(17.5)-19.0-(21.3) (17.5)-19.8-(23.2)

less than 1.0 less than 1.0

less than 1.0 less than 1.0

(17.5)-18.8-(19.4) (17.5)-20.0-(21.3)

0.97 0.96

(7.8)-9.5-(11.6) (7.8)-10.1-(11.6)

less than 1.0 less than 1.0

less than 1.0 less than 1.0

Subtribe 30j. Acalyphinae 184. Acalypha (colpi measured from SEM ; endoaperture too small to measure) A. acmophylla (14.3)-15.1-(15.6) (15.6)-15.9-(16.9) 0.95 A. communis* (7.8)-9.3-(9.7) (7.8)-9.7-(11.6) 0.96 A. costaricensis (9.1)-10.4-(11.7) (9.1)-11.5-(13.0) 0.90 A. diversifolia* (9.7)-9.8-(10.5) (9.7)-11.1-(11.6) 0.88 A. grantii (13.6)-15.7-(17.5) (15.5)-16.9-(19.4) 0.92 A. lanceolata Reeder 876 (11.7)-13.1-(14.3) (13.0)-14.6-(16.9) 0.90 Wheeler 12945 No LM preparation A. monococca (11.7)-13.0-(14.3) (13.0)-14.6-(15.6) 0.89 A. ornata* (9.1)-9.7-(10.4) (10.4)-10.7-(11.7) 0.90 A. ostryaefolia (9.7)-11.2-(11.6) (11.6)-12.2-(13.6) 0.92 A. peduncularis (15.5)-16.0-(17.5) (19.4)-20.0-(21.3) 0.80 A. schinzii (13.6)-15.3-(17.5) (17.0)-17.7-(19.4) 0.86 A. setosa* (7.8)-7.9-(8.5) (7.8)-9.0-(9.7) 0.88 A. stipulacea* (9.7)-10.6-(11.6) (11.6)-12.8-(13.6) 0.83 A. wilkesiana tetrads, longest dimension (28.6)-33.8-(36.4)

unclear unclear unclear (2.4)-2.4-(2.4) (4.4)-4.8-(5.0)

4-aperturate

4-aperturate 4-aperturate

unclear unclear unclear (2.3)-2.6-(2.7) unclear (4.0)-4.3-(4.8) (1.5)-1.5-(1.5) (3.0)-3.0-(3.0)

Subtribe 30k. Lasiococcinae 185. Lasiococca L. symphylliifolia

(18.2)-20.8-(22.1)

(18.2)-19.1-(20.8)

1.08

(13.0)-12.9-(14.3)

unclear

186. Spathiostemon S. javensis Beaman et al. 8860 Ramos 17661*

(22.1)-23.2-(24.7) (18.2)-19.0-(20.8)

(19.5)-20.9-(22.1) (16.9)-18.7-(19.5)

1.09 1.02

(11.7)-12.6-(13.0) (10.4)-12.3-(14.3)

(1.3)-2.2-(2.6) (2.6)-2.6-(2.6)

(6.5)-7.7-(7.8) (6.5)-8.6-(10.4)

187. Homonoia H. riparia Huang 2459* McGregor 247

(18.2)-19.9-(22.1) (23.4)-24.7-(27.3)

(20.8)-22.7-(24.7) (24.7)-26.8-(29.9)

0.88 0.88

(7.8)-10.1-(11.6) (6.5)-9.2-(10.4)

(2.6)-2.6-(2.6) (2.6)-2.7-(3.9)

(9.1)-12.3-(14.3) (9.1)-10.3-(11.7)

Tribe 31. Plukenetieae Subtribe 31a. Plukenetiinae Subtribe threecolpate

PALBO 2467 11-10-02

241

242

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Table 2 (Continued). Taxon

188. Haematostemon H. guianensis

Pollen size Polar

Equatorial

P/E

(33.0)-36.9-(38.8)

(36.9)-39.6-(46.6)

0.93

189. Astrococcus A. cornutus* 190. Angostyles A. longifolia (n = 9)

191. Romanoa R. tamnoides Harshberger 833 Trinta 533 and Fromm 1609

Colpus length

31.2, 33.8, 40.3 (n = 3)

(45.5)-47.3-(49.4)

(42.9)-44.8-(46.8)

(41.6)-43.2-(44.2) (n = 4)

(25.2)-28.5-(33.0)

Endoaperture Polar

Equatorial

less than 1.0

less than 1.0

No pollen in equatorial view

1.05

(19.5)-22.5-(28.6) (n = 4)

3,4-colpate

No pollen in equatorial view

No LM preparation

192. Eleutherostigma E. lehmannianum Cazalet and Pennington 39.0 (n = 1) 5089 Lawrance 276 (n = 2) 41.6, 44.2

31.2, 32.5, 32.5 (n = 3) 41.6, 46.8

193. Plukenetia P. brachybotrya P. conophora P. loretensis Liesner 3788 Schunke 3686 (n = 2) P. penninervia* P. polyadenia Ducke 20619 Mexia 6470 P. volubilis Asplund 14129 (n = 2) Liesner 2068*

45.5, 48.1 45.5, 49.4 Depauperate sample

194. Vigia V. serrata Pederson 8790 Thomas et al. 10221

27.3 (n = 1) 32.0 (n = 1)

unclear margins

33.8 (n = 1) (41.6)-44.5-(46.8) (n = 4) No LM preparation No LM preparation 44.9, 48.1 52.0, 52.0 20.8, 20.8 (n = 2) (22.1)-23.1-(24.7) (n = 5) 49.4, 52.0, 52.0 (n = 3)

unclear margins 9.1 no pollen in equatorial view

No LM preparation unclear margins

(31.2)-31.8-(32.5) (n = 4) 33.8, 37.7, 40.3 (n = 3)

Subtribe 31b. Tragiinae 195. Cnesmone C. javanica* (n = 2) C. tonkinensis

poorly de¢ned apertures 31.0, 34.9 29.0, 33.0 Depauperate sample

196. Megistostigma M. burmanicum Depauperate sample M. cordatum Depauperate sample M. malaccense Chan 5/9/71 No LM preparation Rahmat Si Toroes 1975 No LM preparation Sinclair 8025 SEM only 197. Sphaerostylis

poorly de¢ned apertures

No material

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336 Table 2 (Continued). Taxon

198. Tragiella T. friesiana T. natalensis Kemp 741 Mearns 255 199. Platygyna P. dentata P. hexandra Curtiss s.n.* Morton 10253 P. parvifolia

Pollen size Polar

Equatorial

P/E

(27.3)-33.1-(35.1)

(29.9)-31.5-(33.8)

1.05

(15.6)-21.5-(26.0)

No LM preparation (31.0)-37.6-(40.7) (27.2)-33.6-(36.9)

1.12

(23.2)-29.5-(33.0)

Ortiz 1104 Standley and Valerio 46475 A. urens Alexander s.n. Proctor 36826 Acidoton species indet. 202. Pachystylidium P. hirsutum Ramos 40929* Van Balgooy 2974

Endoaperture Polar

Equatorial

genus inaperturate (longest dimension given ¢rst) (27.3)-32.2-(36.4) (27.3)-32.0-(35.1) 29.1 (n = 1) (31.2)-36.8-(41.6) (28.6)-31.7-(35.1)

200. Tragia T. bailloniana* (34.9)-39.2-(42.2) T. cf. cordifolia (n = 4) (28.6)-30.2-(32.5) T. gardneri (19.5)-20.7-(28.6) T. lukafuensis (29.9)-32.1-(33.8) T. mitis* Depauperate sample T. pungens (26.0)-27.7-(28.6) T. ramosa Depauperate sample T. sellowiana (36.4)-39.5-(45.5) T. smallii Depauperate sample T. spathulata (n = 3) 28.6-29.9-31.2 T. stylaris Depauperate sample T. urens (25.2)-28.7-(33.0) T. urticaefolia Thomas 104979 Depauperate sample Webster and Wilbur 3231 (26.0)-27.4-(28.6) T. volubilis (28.6)-30.6-(32.5) 201. Acidoton A. microphyllus A. nicaraguensis Espinoza 188 Moreno 17228

Colpus length

(29.9)-34.0-(36.4)

27.2, 33.0 (n = 2) (31.2)-35.3-(39.0) (27.3)-29.5-(32.5)

(38.8)-44.1-(48.5) (26.0)-29.3-(31.2) (22.1)-27.6-(31.2) (27.3)-30.0-(32.5)

0.89 1.03 0.75 1.07

(19.4)-23.7-(27.2) (9.7)-11.6-(15.5) (13.0)-14.8-(16.9) (19.5)-22.6-(26.0)

(28.6)-30.1-(32.5)

0.92

unclear margins

(36.4)-40.0-(44.2)

1.01

26.0-28.6-31.2

1.04

inaperturate poorly de¢ned apertures (15.6)-16.4-(16.9)

(23.2)-28.3-(29.1)

1.01

poorly de¢ned apertures

(28.6)-33.5-(35.1) (24.7)-28.3-(32.5)

0.82 0.82

(18.2)-19.2-(22.1) 9.7 (n = 1)

(28.6)-30.5-(33.8)

inaperturate

(27.3)-29.5-(32.5) (31.2)-32.2-(35.1) (26.0)-28.9-(31.2) (26.0)-26.8-(28.6) (many grains collapsed) No LM preparation (23.4)-26.7-(28.6) (26.0)-27.0-(29.9) (grains collapsed)

0.91 1.07

(29.9)-32.6-(36.4) (27.3)-30.0-(35.1) No LM preparation (29.1)-31.4-(36.9) (31.0)-32.7-(34.9)

1.09

(16.9)-20.5-(22.1) 15.6 (n = 1)

0.99 inaperturate

0.96

(19.4)-22.3-(29.1) Poorly de¢ned pores

(25.2)-28.1-(31.0) (27.2)-28.3-(31.0) No LM preparation

0.99

(46.6)-55.7-(62.0) (61.1)-65.0-(67.6)

1.24 1.05

Subtribe 31c. Dalechampiinae 203. Dalechampia D. brevipes* D. capensis (n = 5)

(40.7)-44.7-(48.5) (52.0)-61.4-(66.3)

(^)-25.2-(^)

PALBO 2467 11-10-02

(5.8)-8.2-(9.7)

zonorate? grains aberrant, see text

243

244

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Table 2 (Continued). Taxon

Pollen size

Colpus length

Endoaperture

(4.2)-6.1-(8.3)

Polar

Equatorial

P/E

D. D. D. D. D.

cissifolia clematidifolia dioscoreifolia heteromorpha (n = 9) humilis

(55.9)-63.7-(72.8) (63.7)-71.1-(74.1) (78.0)-84.5-(93.6) (79.3)-86.8-(93.6) (66.3)-72.1-(75.4)

(46.8)-59.2-(67.6) (54.6)-56.8-(58.5) (71.5)-75.3-(78.0) (71.5)-73.7-(80.0) (53.3)-57.6-(63.7)

1.07 1.25 1.12 1.17 1.25

(24.0)-28.1-(31.0) (29.9)-33.7-(39.0) (35.1)-57.4-(44.2) (26.0)-27.0-(28.6) (29.9)-32.8-(35.1)

Polar

D. D. D. D.

scandens* stenosepala* subternata tiliifolia*

(56.3)-60.1-(64.0) (42.7)-46.6-(50.4) (66.3)-75.8-(79.3) (93.2)-99.0-(104.8)

(42.7)-46.1-(48.5) (36.9)-40.1-(42.7) (50.7)-57.5-(65.0) (66.0)-70.3-(78.8)

1.30 1.16 1.31 1.41

(23.2)-27.0-(29.1) (19.3)-20.9-(27.2) (37.7)-40.8-(45.5) (28.8)-39.1-(48.5)

three-colpate (26.0)-27.6-(29.9)

(22.1)-24.3-(26.0)

1.14

(15.6)-17.3-(18.2)

28.6, 28.6 (n = 2) (26.0)-30.3-(35.1) (n = 10) 0.94 No LM preparation

13.0, 13.0 (n = 2)

Equatorial

(9.1)-10.9-(13.0) (5.2)-7.1-(9.1) endoaperture unclear (5.8)-8.0-(9.7) (3.9)-4.5-(5.8) (3.9)-5.2-(6.5) (9.7)-10.7-(13.6)

Tribe 32. Omphaleae 204. Omphalea O. bracteata O. diandra Irwin et al. 48498 Palacios 6823 O. oleifera Manriquez 443 Manriquez 2334* O. oppositifolia Gillespie 4014 (n = 3) Malcomber 2832 O. palmata* O. triandra O. trichotoma

No LM preparation (21.3)-22.7-(23.2) (25.2)-28.1-(29.1)

15.5 (n = 1) (23.4)-26.9-(29.9)

(28.6)-29.0-(29.9) (22.1)-24.7-(28.6) (n = 8) (19.4)-20.8-(21.3) (31.2)-32.4-(33.8) (23.4)-27.3-(32.5) (n = 6)

0.81

(9.7)-10.6-(11.6) no grains in equatorial view no grains in equatorial view

0.75 0.83

(16.9)-18.7-(20.8) no grains in equatorial view

An asterisk denotes pollen samples mounted and measured in silicon oil. If there are more E measurements than P, or only E measurements, then some E were made in polar view. For inaperturate grains, longer axis is given ¢rst.

exine 0.8^1.2 Wm thick; the apertural endexine ca. two thirds as thick as the mesocolpial ectexine (e.g., Plate I, 5; Plate III, 16), sometimes with ectexinous granules at the endoaperture (e.g., Plate I, 7; Plate II, 15), the non-apertural endexine thin (e.g., Plate I, 3, 5; Plate II, 12); the foot layer thin (e.g., Plate I, 1, 3; Plate III, 18, 21), sometimes irregular (e.g., Plate I, 5; Plate II, 12, 13); the columellae variable, either short at the equator and elongate at the poles (e.g., Plate I, 1; Plate II, 13; Plate III, 18), or the columellae more uniform in length (e.g., Plate I, 5), frequently variable in shape (Plate II, 10 arrowheads, 13 arrowheads; Plate III, 18), occasionally appearing fused (e.g., Plate II, 10 arrowheads, 13 arrowheads; Plate III, 18); the tectum almost continuous (e.g., Plate I, 1; Plate II, 10, 13: Plate III, 16, 18) to slightly less so (e.g., Plate I, 5, 7, 9), minutely spinulose (e.g., Plate II, 13; Plate III, 20. Discussion : The pollen of three species of Erythrococca, E. africana (Plate I, 1, 2), E. oleracea

(Plate II, 13, 14) and E. welwitschiana (Plate III, 17, 18) have columellae noticeably elongated at the poles but shorter near the equator. The remaining taxa/collections examined have more uniform columellae. See discussion at end of subtribe Claoxylinae (after Amyrea, genus 174). 169. Claoxylon A. Jussieu, 75 species in the Old World tropics but absent from Africa. (Plates IV, V; Plate VI, 37^39). Species examined : C. albicans (Plate IV, 22^ 25); C. australe; C. cf. bakerianum (Plate IV, 26^28); C. hainanense (Plate IV, 29, 30); C. insulanum (Plate V, 31, 32); C. longifolium (Plate V, 33^35); C. lutescens (Plate V, 36); C. sitibundum (Plate VI, 37^39). Pollen oblate spheroidal to prolate-spheroidal in equatorial view; PUE = (15.6^25.2)U(13.6^ 23.2) Wm; (three-) four- (¢ve-) colporate (Plate IV, 29; Plate V, 33), the colpi 7.8^19.4 Wm long,

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

245

Table 3 Distribution of selected pollen characteristics Exine

Tectum

Muri Columellae

Foot layer

thick. (Wm)

sculpt.

0.8^0.9

1.2 No TEM 1.0

Aperture

thick.

xs

leng.

branc. fused

thick.

morph. margo operc.

PMS

TK

^

VAR

^

?

TN

CON

^

^

PMS

AV

^

AV

^

^

AV

IRR

PMS

AV

^

AV

^

^

TN

IRR

^ ^ ^

^ ^ ^

1.0 PMS SEM only PMS 1.0 PMS ^ PMS

AV

^

AV

^

^

TN

IRR

TK TN

^ ^

AV VAR

^ ^

+ +

AV TN

IRR IRR

^ ^ ^ ^

^ ^ ^ ^

1.2

PMS

AV

^

VAR

^

+?

TN

CON

^

^

SCAM

AV

^

AV

^

^

TN

IRR

^

^

1.0 No TEM 1.2 0.8^1.0

PMS PMS PMS PMS

AV/TK ^

SH

^

^

AV

IRR

^

^

AV AV

^ ^

SH SH

^ ^

^ ^

AV AV

IRR IRR

^ ^

^ ^

No TEM 1.2 1.0

AV TK

^ ^

SH SH

^ ^

^ ^

? AV

IRR IRR

1.2

PMS PMS PMS PMS PMS

TK

^

SH

^

^

AV-TK CON

^ ^ ^ ^ ^

^ ^ ^ ^ ^

170. Claoxylopsis C. purpurescens

1.2

PMS

TK

^

AV

^

^

ABS-TN ^

^

^

171. Mareya M. micrantha M. spicata

1.0 1.2

PMS PMS?

TK TK

^ ^

SH SH

^ ^

^ ^

AV AV

CON ?

^ ?

^ ^

171a. Mareyopsis M. longifolia Zenker 35 Zenker 4228 M. oligogyna

1.2 No TEM No TEM

SCAM RETM RETM

TN

^

LG

?

^

TN

IRR

^

^

^

^

172. Discoclaoxylon D. hexandrum Baldurin 6696 1.2 Nemba and Mamba 780 1.1

SCAM SCAM

AV-TK ^ AV-TK ^

AV AV

^ ^

^ ^

AV AV

CON CON

^ ^

^ ^

173. Micrococca M. capensis M. holstii

PMS PMS

AV

^

LG

^

^

AV

CON

^ ^

^ ^

SCAM PMS

AV AV

^ ^

AV-LG ^ AV ^

^ ^

AV-TN ? AV-TN IRR

^ ^

^ ^

Remarks

Tribe 30. Acalypheae Subtribe 30g. Claoxylinae 168. Erythrococca E. africana E. bongensis Bequaert 5282 Witte 11704 E. hirta E. kirkii Gereau 1491 Graham 2123 E. menyharthii E. oleracea E. welwitschiana

Erythrococca species indet. 1.2 169. Claoxylon C. albicans C. australe C. cf. bakerianum C. hainanense C. insulanum Balansa 269 McKee 196 C. longifolium C. lutescens C. sitibundum

M. mananarenta M. oligandra

3^5-aperturate columellae elongate at poles

columellae elongate at poles columellae elongate at poles

minimal TEM data

inner surface of tectum minutely granular

3^4-colporate 1.2 minimal TEM 1.2 1.2

PALBO 2467 11-10-02

246

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Table 3 (Continued).

174. Amyrea A. humbertii Humbert 6599 Miller 3702 A. sambiranensis

Exine

Tectum

Muri Columellae

Foot layer

thick. (Wm)

sculpt.

thick.

xs

leng.

thick.

1.3 1.3 1.2

STR STR STR

TK TK AV

^ ^ ^

SH ^ SH-AV ^ AV ^

^ ^ ?

AV CON AV-TK IRR AV-TN IRR

^ ^ ^

^ ^ ^

1.2 1.6

PMS PMS

AV ^ AV-TK ^

AV SH

^ ^

^ ^

TN IRR AV-TN IRR

^ ^

^ ^

1.0 1.0 No TEM 0.9 1.1 LM only 1.2 1.0 No TEM 0.9 0.9 2.2 No TEM 1.0 No TEM 1.4 0.9 LM only LM only 1.0^1.2 No TEM

PMS PMS SCAM? PMS PMS

TK TK

^ ^

SH SH

^ ^

^ ^

TN TN

CON CON

TK TK

^ ^

SH SH

^ ^

^ ^

TN TN

CON CON

PMS PMS PMS PMS PMS PMS PMS PMS PMS SCAM PMS

TK TK

^ ^

SH SH

^ ^

^ ^

TN TN

IRR CON

TK TK TK

^ ^ ^

SH SH AV

^ ^ ^

^ ^ ^

TN CON TN CON TN-AV CON

TK ^ TK TK

^ ^ ^ ^

SH

^

^

TN

CON

^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ?

^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^

SH SH

^ ^

^ ^

TN TN

CON CON

PMS PMS

TK

^

SH

^

^

TN

CON

? ^ ^ ^ ^ ^

^ ^ ^ ^ ^ ^

SCAM SCAM SCAM

TK TK TK

^ ^ ^

SH SH SH

^ ^ ^

^ ^ ^

TN IRR TN-AV IRR AV IRR

^ ^ ^

^ ^ ^

SCAM SCAM

TK TK

^ ^

SH SH

^ ^

^ ^

TN TN

CON CON

^ ^

^ ^

PMS PMS PMS

TK

^

SH

^

^

TN

CON

TK

^

SH

^

^

TN

CON

^ ^ ^

^ ^ ^

branc. fused

Aperture

morph. margo operc.

Subtribe 30h. Lobaniliinae 175. Lobanilia L. bakeriana L. claoxyloides Subtribe 30i. Rottlerinae 176. Mallotus M. acuminatus M. apelta M. baillonianus M. barbatus M. eriocarpus M. fuscescens M. japonicus M. korthalsii M. leucodermis M. leucophyllus M. microcarpus M. mollissimus M. muticus M. nepalensis M. nesophilus M. oreophilus M. philippinensis M. polyadenos M. puber M. subulatus M. sumatranus

177. Deuteromallotus D. acuminatus Decary 16884 1.5 Nicoll et al. 500 1.3 Deuteromallotus species in- 1.5 det. 178. Cordemoya C. integrifolia Goode et al. 3952 Lorence 2231

1.6 1.2

179. Coccoceras

No material

180. Trewia T. nudi£ora L. Fernandes 31 Road s.n. 4/4/1929 Wenzel 71

1.2 No TEM 0.8^1.0

PALBO 2467 11-10-02

Remarks

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

247

Table 3 (Continued). Exine

Tectum

thick. (Wm)

sculpt.

181. Neotrewia N. cumingii Elmer 17641 Sulit 22901

0.9 1.0

PMS PMS

182. Rockinghamia R. angustifolia

0.9^1.3

R. brevipes 183. Octospermum O. pleiogynum Kostermans 287 Streimann and Katik 28607

Muri Columellae

Foot layer

thick.

xs

leng.

branc. fused

thick.

morph. margo operc.

TK TK

^ ^

SH SH

^ ^

^ ^

TN TN

CON CON

^ ^

^ ^

+

AV

^

^

TN

IRR

^

^

1.5

PCTD andAV RETM PCTD TK

+

AV

^

^

TK

CON

^

^

0.9^1.2 1.2

PMS PMS

^ ^

SH SH

^ ^

^ ^

TN TN

CON CON

^ ^

^ ^

TK TK

Aperture

Remarks

Subtribe 30j. Acalyphinae 184. Acalypha A. acmophylla A. communis A. costaricensis A. diversifolia A. grantii A. lanceolata Reeder 876 Wheeler 12945 A. monococca A. ornata A. ostryaefolia A. peduncularis A. schinzii A. setosa A. stipulacea A. wilkesiana

3^4^5-aperturate LM only No TEM minimal TEM 0.8 0.5^0.7

ACA ACA

^ ^

^ ^

ACA ACA

AV ^ TK-AV ^

AV-LG ^ SH ^

^ ^

TN TN

IRR IRR

^ ^

^ ^

0.7^0.8 No TEM 0.7^0.8 0.6^0.7 LM only 0.9 0.9 0.6 1.4 0.9^1.2

ACA ACA ACA ACA

TK-AV ^

SH

^

^

TN-AV IRR

TK-AV ^ TK ^

AV SH

^ ^

^ ^

AV IRR TN-AV IRR

ACA ACA ACA ACA ACA

TK TK TK TK TK

SH SH SH-AV SH-AV SH

^ ^ ^ ^ ^

^ ^ ^ ^ ^

AV TN TN TN-AV TN

^ ^ ^ ^ ^ ^ ^ ^ ^ ^

^ ^ ^ ^ ^ ^ ^ ^ ^ ^

No TEM

PMS

^

^

^ ^ ^ ^ ^

IRR CON CON IRR IRR

tetrads?

Subtribe 30k. Lasiococcinae 185. Lasiococca L. symphylliifolia

186. Spathiostemon S. javense Beaman 8860 Ramos 17661 187. Homonoia H. riparia Huang 2459 McGregor 247

segmented lirae 0.9 0.9

SPEC SPEC

AV AV

^ ^

SH SH

^ ^

^ ^

TN TN

CON CON

^ ^

^ ^ densely microechinate

1.0

SPEC SPEC

TK TK

^ ^

SH SH

^ ^

^ ^

Tribe 31. Plukenetieae Subtribe 31a. Plukenetiinae

TN

CON

^ ^

^ ^

minimal TEM

three-colpate

PALBO 2467 11-10-02

248

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Table 3 (Continued). Exine

Tectum

thick. (Wm)

sculpt.

188. Haematostemon H. guianensis

1.5

CROM

189. Astrococcus A. cornutus

No TEM

CROM

190. Angostyles A. longifolia

1.5^1.7

RUGM

Muri Columellae

Foot layer

thick.

xs

leng.

branc. fused

thick.

AV

^

LG

+

AV-TK CON

AV-TK ^

SH

^

^

^

TK

Aperture

morph. margo operc.

CON

^

^

^

^

^

^

191. Romanoa R. tamnoides Harshberger 833 Trinta 533 and Fromm 1609

2.4 2.2

PCT PCT

AV AV

^ ^

LG LG

+ +

+ +

AV AV

IRR IRR

^ ^

granular area under tectum PCT

AV-TK ^

LG

+

?

TN-AV CON

^

^

PCT

AV

^

LG

+

?

TN-AV CON

^

^

193. Plukenetia P. brachybotrya P. conophora

No TEM 1.5

RET PCT

AV

^

LG

+

?

TN

^ ^

^ ^

No TEM 3.0 limited TEM

RET RET RET

TN

^

AV-LG ^

^

TN-AV IRR

^ ^ ^

^ ^ ^

2.2

PCT

AV

^

LG

+

TN-AV IRR

^

^

Mexia 6470 P. volubilis Asplund 14129

No TEM

PCT

^

^

2.2

PCT

^

^

Liesner 2068

No TEM

PCT

^

^

No TEM No TEM

RET RET

^ ^

^ ^

194. Vigia V. serrata Pederson 8790 Thomas et al., 10221

AV-TK ^

LG

+

?

+

IRR

TN-AV IRR

Subtribe 31b. Tragiinae

granular area under tectum

granular area under tectum

granular area under tectum

three-colpate unless otherwise noted apertures poorly de¢ned

195. Cnesmone C. javanica C. tonkinensis

endexine^foot layer irregularly thickened along apertural margin

^ ^

E. lehmannianum Cazalet and Pennington 2.0 5089 Lawrence 276 2.5

P. polyadenia Ducke 20619

granular area under tectum; endexine^foot layer irregularly thickened along apertural margin

granular area under tectum

192. Eleutherostigma

P. loretensis Liesner 3788 Schunke 3686 P. penninervia

Remarks

No TEM 0.9^1.0

PMS PMS

AV

^

IRR

^

^

PALBO 2467 11-10-02

ABS-TN IRR

^ ^

^ ^

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

249

Table 3 (Continued).

196. Megistostigma M. burmanicum M. cordatum M. malaccense Chan 5/9/71 Rahmat Si Toroes 1975 Sinclair 8025 197. Sphaerostylis 198. Tragiella T. friesiana T. natalensis Kemp 741 Mearns 255

Exine

Tectum

thick. (Wm)

sculpt.

No TEM No TEM

PMS? PMS?

1.0

PMS? PMS? PMS?

TN

^

IRR

^

^

2.0

RET

AV

^

AV-LG ^

No TEM 1.9

RET RET

AV

^ ^

AV-LG ^

thick.

Muri Columellae

Foot layer

xs

thick.

leng.

branc. fused

Aperture

morph. margo operc. apertures poorly de¢ned

No TEM

^ ^

^ ^

ABS-TN IRR

^ ^ ^

^ ^ ^

^

TN

CON

?

^

^

TN

CON

^ ?

^ ^

No material

199. Platygyna P. dentata P. hexandra Curtiss s.n. Morton 10253 P. parvifolia

Remarks

inaperturate. Endexine of 1^2 threads No TEM

PCT

No TEM 1.5 1.5

PCT PCT INT

minimal TEM 1.2 1.8 2.0 No TEM 2.0 1.2 1.0^1.2 No TEM 1.8 1.2 1.0

RETF

TN ^

^ ^

LG LG

+ +

+ +

^ ^

^ ^

^

^

^ ^ ^

^ ^ ^

^

^

^ ? ? ^ ? ^ ^ ^ ? ^ ^

^ ^ ^ ^ ^ ^ ^ ^ ^ ^ ^

^

^

surface columellate

intectate

200. Tragia T. bailloniana

T. T. T. T. T. T. T. T. T. T. T.

cf. Cordifolia gardneri lukafuensis mitis pungens ramosa sellowiana smallii spathulata stylaris urens

T. urticaefolia Thomas 104979 No TEM Webster and Wilbur 3231 1.3 T. volubilis 1.0 201. Acidoton A. microphyllus A. nicaraguensis Espinoza 188 Moreno 17228 Ortiz 1104 Standley and Valerio 46475

SCAM RETC RETC RET RETC INT PMS SCAM RETC INT SCAM

TK ^ AV ^ AV-TK ^

AV AV SH

^ ^ ^

^ ^ ^

TN-AV CON AV CON TN CON

AV-TK ^ ^ ^ TN ^

AV LG LG

^ ^ ^

^ ^ ^

TN TK TN

IRR CON IRR

AV ^ TK

AV LG AV/ IRR

^ ^ ^

^ ^ ^

TN TK ^

CON CON ^

^ ^ ^

INT INT INT

surface columellate inaperturate

surface columellate

^ ^

^ ^

LG LG

^ ^

^ ^

TK TK

CON CON

^ ^

^ ^

surface columellate surface columellate

1.6

‘‘RUGM’’ ^

^

LG

IRR

IRR

^

^

^

^

intectate ? inaperturate 3-colpate

No TEM 1.8 No TEM

CROM CROM CROM

AV

^

AV

^

^

ABS/TN IRR

^ ^ ^

^ ^ ^

1.8

CROM

AV-TK ^

SH

^

^

TN

^

^

PALBO 2467 11-10-02

IRR

250

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Table 3 (Continued). Exine

Tectum

Muri Columellae

Foot layer

thick. (Wm)

sculpt.

xs

leng.

branc. fused

thick.

morph. margo operc.

A. urens Alexander s.n. Proctor 36826 Acidoton species indet.

1.5 No TEM LM only

‘RUGM’ ^ ‘RUGM’

^

LG

IRR

IRR

^

^

202. Pachystylidium P. hirsutum Ramos 40929 Van Balgooy 2974

No TEM 0.8^1.0

PMS No SEM TN

1-

SH

^

^

ABS/TN IRR

No TEM 4.0^4.5 3.0 5.5 No TEM 4.0 minimal TEM minimal TEM No TEM 7.5 No TEM

RET RETF RET PCT RET RET RET

AV AV AV

^ ^ ^

LG AV AV

^ ^ ^

^ ^ ^

AV TN TN

CON CON CON

TN

^

LG

^

^

TN

IRR ^

No TEM

PMS

2.0

PMS

No TEM

thick.

Aperture

^ ^ ^

^ ^ ^

^ ^

^ ^

^ ^ ^ ^ ^ ^ ^

^ ^ ^ ^ ^ ^

^

^

^ ^ ^

^ ^ ^

^

^

^

^

PMS

^

^

No TEM No TEM

PMS PMS

^ ^

^ ^

No TEM No TEM No TEM 2.3 1.7

PMS PMS PMS PMS PMS

^ ^ ^ ^ ^

^ ^ ^ ^ ^

Remarks

intectate ?

Subtribe 31c. Dalechampinae 203. Dalechampia D. brevipes D. capensis D. cissifolia D. clematidifolia D. dioscoreifolia D. heteromorpha D. humilis D. scandens D. stenosepala D. subternata D. tiliifolia

RET RET RETF RET

TN-AV ^

SH-LG ^ +

^

TN

CON

Tribe 32. Omphaleae 204. Omphalea O. bracteata O. diandra Irwin et al. 48498 Palacios 6823 O. oleifera Manriquez 443 Manriquez 2334 O. oppositifolia Gillespie 4014 Malcomber 2832 O. palmata O. triandra O. trichotoma

TK

^

TK ^ AV-TK ^

IRR

AV AV

^

^ IRR

^

^ IRR

AV-TK IRR

AV IRR AV-TK IRR

nonapertural endexine separates from ectexine

All taxa have three-colporate pollen unless noted otherwise under ‘Remarks’. Exine thickness is based on measurements of the center(s) of radial sections (TEM) of mesocolpi(a)um. Tectum sculpture or surface: ACA = Acalypha type; CROM = microcrotonoid; ECHM = microechinate; INT = intectate; IRR = irregular; PCT = punctate; PCTD = deeply punctate; PMS = punctate-microspinulose; RET = reticulate; RETC = reticulate/crenate muri; RETF = ¢nely reticulate; RETM = microreticulate; RUGM = microrugulate; SCAM = microscabrate; SPEC = special and de¢ned under remarks; STR = striate. The thickness (at center of mesocolpium) of the tectum and of the foot layer are expressed as a fraction of the total thickness of the ectexine: ThiN 6 1/3, AVerage if 1/3, ThicK if s 1/3. Columellae length: SHort if the length is 6 1/3 of the ectexine thickness, AVerage if 1/3, LonG if s 1/3. Presence/absence of a characteristic is based on all micrographs, not only the ones presented here in ¢gs. 1^360. + = presence of a characteristic; ? = characteristic unclear or inconsistent; ^ = characteristic not observed in the collection(s) examined. Columellae: branched or unbranched; fused or not fused. Foot layer morph: CON = consistent or uniform; IRR = irregular; ABS = absent. Aperture: margo present or absent; operculum present or absent. Muri: triangular in xs or not.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

narrow ; costa ectocolpi poorly developed, margo absent, operculum absent; the endoaperture lalongate, PUE = ( 6 1.0^3.9)U(3.9^13.0) Wm, occasionally with ¢ne granules on the membrane ; the tectum complete (e.g., Plate IV, 23, 27; Plate V, 35) or nearly so, punctate-microspinulose (e.g., Plate IV, 25, 29; Plate V, 35). In thin section, the mesocolpial exine ca. 0.8^1.2 Wm thick; the apertural endexine about equal (e.g., Plate IV, 22) to slightly less (e.g., Plate V, 32, 36) than the thickness of the mesocopial ectexine ; the non-apertural endexine thin (e.g., Plate IV, 26; Plate V, 32; Plate VI, 39), sometimes granular; the foot layer irregular (e.g., Plate IV, 22, 28; Plate VI, 39), occasionally discontinuous (e.g., Plate V, 32); the columellae mostly small (e.g., Plate IV, 22, 30), occasionally thick (e.g., Plate V, 32; Plate VI, 39); the tectum continuous (e.g., Plate IV, 22, 26, 28) or nearly so (e.g., Plate V, 36; Plate VI, 37), minutely spinulose. Discussion : Pollen of Claoxylon is of a generalized type except for the presence of more than three apertures. See discussion after Amyrea (genus 174). 170. Claoxylopsis Leandri, three species endemic to Madagascar (Plate VI, 40^42). Species examined : C. purpurescens (Plate VI, 40^42). Pollen subprolate in equatorial view; PUE = (16.9^23.4)U(14.3^19.5) Wm (but see Table 2); three-colp(or)ate, the colpi 10.4^17.5 Wm long, costa ectocolpi present, margo absent, operculum absent; the endoaperture 6 1.0 Wm in diameter ; the tectum complete, punctate-microspinulose (Plate VI, 40, 41). In thin section, the mesocolpial exine ca. 1.0^1.2 Wm thick; the apertural endexine about one/half the thickness of the mesocolpial ectexine ; the non-apertural endexine thin (Plate VI, 42); the foot layer seemingly absent (Plate VI, 42) to thin and irregular; the columellae average, stout; the tectum continuous, the most prominent part of the ectexine, the inner surface minutely granular. Discussion : Only one collection of Claoxylopsis was available for this study. The existence of two size classes of pollen (Table 2) should be con¢rmed by additional collections. The exine struc-

251

ture ^ threadlike to absent foot layer, short stout columellae, and the minutely granular or fringed inner surface of the tectum ^ would argue against a congeneric treatment of Claoxylopsis and Claoxylon, but not strongly so. 171. Mareya Baillon, two or three species in Africa. (Plates VII, VIII; Plate IX, 56, 57). Webster’s concept of Mareya included the genus Mareyopsis Pax and Ho¡mann, the latter based on Mareya longifolia Pax. The pollen of the two genera is di¡erent, and is described separately. See discussion after Mareyopsis pollen type. 171. Mareya pollen type (punctate-microspinulose tectum, thin columellae). Species examined and included: M. micrantha (Plate VII, 43^45); M. spicata (Plate VII, 46^48). Pollen prolate spheroidal in equatorial view; PUE = (14.3^16.9)U(14.3^16.9) Wm; three-colpate or three-colporate, the colpi 9.7^13.0 Wm long, narrow, the membrane ¢nely granular, costa ectocolpi present (M. spicata) or seemingly absent (M. micrantha); margo absent, operculum absent; the endoaperture poorly de¢ned (M. spicata), or very small ( 6 1.0 Wm P) (M. micrantha); the tectum complete, sparsely punctate-microspinulose (Plate VII, 45). In thin section, the mesocolpial exine ca. 1.0^1.2 Wm thick; the apertural endexine prominent (e.g., Plate VII, 46), twice as thick as the mesocolpial ectexine; the non-apertural endexine thin (Plate VII, 47) to seemingly well-developed (e.g., Plate VII, 44); the foot layer mostly uniform, becoming irregular near the colpi ; the columellae mostly small, slender (e.g., Plate VII, 44, 46, 47), sometimes densely spaced; the tectum continuous (Plate VII, 44, 47), the most prominent component of the ectexine, minutely spinulose. Discussion: In Breteler et al. (1997), Punt illustrates the presence of endoapertures in the pollen of Mareya, but in the collection of M. micrantha that we examined (Baldwin 9239), endoapertures were not apparent in this silicon oil mounting. In the acetolysed preparation of M. spicata (Cooper 404), very narrow endoapertures were observed. See also discussion after Mareyopsis.

PALBO 2467 11-10-02

252

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate I.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

171a. Mareyopsis Pax et K. Ho¡mann, two or three species in Africa. 171a. Mareyopsis pollen type (microreticulate or microscabrate tecta, large columellae). Species examined and included: M. longifolia (Plate VIII, Plate IX, 56); M. oligogyna (Plate IX, 57). Pollen oblate spheroidal to prolate spheroidal in equatorial view; PUE = (14.3^19.5)U(14.3^ 22.1) Wm; three-colporate, the colpi 7.8^11.7 Wm long, costa ectocolpi weakly developed, margo absent, operculum absent; the endoaperture sometimes poorly de¢ned, PUE = ( 6 1.0^1.3)U(5.2^ 10.4) Wm; the tectum almost complete (e.g., Plate VIII, 54; Plate IX, 56), microreticulate (e.g., Plate VIII, 49, 51), or microscabrate (e.g., Plate VIII, 54). In thin section, the mesocolpial exine ca. 1.2 Wm thick; the apertural endexine almost as thick as the mesocolpial ectexine (Plate VIII, 53, 55); the non-apertural endexine thin but continuous (e.g., Plate VIII, 52, 53, 55); the foot layer variable to seemingly absent (Plate VIII, 50, 53, 55); the columellae well developed (e.g., Plate VIII, 52), frequently with an expanded base extending into the endexine (Plate VIII, 50, 53, 55); the tectum slightly discontinuous (Plate VIII, 50, 52, 53, 55), sometimes with minute spinules. Discussion : Pollen from the collection of Zenker 35 has columellae that appear to extend into the endexine (Plate VIII, 50, 53, 55) and the foot layer appears to be absent. The tectum is also di¡erent, microscabrate (Plate VIII, 54) as opposed to microreticulate (Plate VIII, 49, 51) in the collection of Zenker 4228.

253

Although Webster (1994) does not recognize Mareyopsis as a valid genus, the di¡erences in exine structure and tectum would support their separation (see also Breteler et al., 1997). However, there are other genera in Acalyphoideae, e.g., Plukenetia (Plates XXIX, 223^226; XXX; XXXI, 236^242) and Tragia (Plates XXXVXXXVIII), which have much more disparate pollen. 172. Discoclaoxylon (Mueller-Argoviensis) Pax et K. Ho¡mann, three species mostly in west Africa. (Plate IX, 58^61). Species examined: D. hexandrum (Plate IX, 58^61). Pollen spheroidal to oblate spheroidal in equatorial view; PUE = (15.6^16.9)U(15.6^18.2) Wm; three-colporate, the colpi 7.8^11.6 Wm long, costa ectocolpi present; margo absent, operculum absent ; the endoaperture lalongate, PUE = ( 6 1.0^ 2.6)U(3.9^7.8) Wm; the tectum almost complete, microscabrate (Plate IX, 58). In thin section, the mesocolpial exine ca. 1.1^1.2 Wm thick ; the apertural endexine ca. half as thick as the mesocolpial ectexine; the non-apertural endexine thin but consistent, the inner surface slightly granular; the foot layer well-developed, mostly uniform (e.g., Plate IX, 59); the columellae stout, well de¢ned, the length about equal to the tectum thickness; the tectum thick, slightly discontinuous, with numerous tiny spinules. See discussion at end of subtribe after the genus Amyrea (174).

Plate I. SEM and TEM of pollen of tribe Acalypheae, subtribe Claoxylinae, Erythrococca. 1^3. E. africana. 1. Longitudinal section. Note marked elongation of columellae at the pole (top). TEM. 2. High magni¢cation of tectum near the pole. SEM. 3. Equatorial section of 4-aperturate grain. TEM. 4^7. E. bongensis. 4. Tectum along colpus. Note irregular surface level. SEM. 5. Radial section across mesocolpium. The outer surface of the tectum agrees with that in ¢g. 4. TEM. 6. Slightly oblique polar view. SEM. 7. Slightly oblique longitudinal section. Compare this grain with that in ¢g. 1. TEM. 8, 9. E. hirta. 8. Mesocolpium-centered equatorial view. SEM. 9. Longitudinal section along aperture. TEM. Unless otherwise noted all scale bars for all 45 plates equal 1 Wm.

PALBO 2467 11-10-02

254

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate II.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

173. Micrococca Bentham, 12 species in the Paleotropics (Plate X). Species examined : M. capensis (Plate X, 62^ 64); M. holstii (Plate X, 65); M. mananarenta (Plate X, 66^68); M. oligandra (Plate X, 69^71). Pollen spheroidal to prolate spheroidal in equatorial view; PUE = (15.6^26.0)U(15.6^23.4) Wm; three- to four-colporate (Plate X, 63), the colpi 7.8^19.5 Wm long, narrow, costa ectocolpi present, margo absent, operculum absent; the endoaperture small, mostly lalongate, PUE = ( 6 1.0^ 6.5)U(5.8^14.3) Wm; the tectum almost complete, punctate-microspinulose (Plate X, 63, 65) to irregularly so (Plate X, 67, 70). In thin section, the mesocolpial exine ca. 1.2 Wm thick ; the apertural endexine about half as thick as the mesocolpial ectexine (e.g., Plate X, 66); the non-apertural endexine thin, irregular; the foot layer thin (e.g., Plate X, 62, 66, 71) to average (Plate X, 64); the columellae average (Plate X, 66) to more prominent (e.g., Plate X, 62, 64), slightly longer than the thickest part of the tectum, sometimes narrowed at the middle (Plate X, 71); the tectum slightly discontinuous, irregular (e.g., Plate X, 64, 71). Discussion : Ferna¤ndez-Gonza¤lez and LobreauCallen (1996) illustrate two species of Micrococca, one of which, M. scariosa Prain, has a striate sculpture. Examination of US collections suggests that this species is very distinctive relative to the remaining members of Micrococca as identi¢ed. 174. Amyrea Leandri, two species endemic to Madagascar (Plate XI).

255

A. humbertii (Plate XI, 72^77); A. sambiranensis (Plate XI, 78^80). Pollen prolate spheroidal in equatorial view; PUE = (16.9^24.3)U(15.6^23.4) Wm; three-colporate, the colpi 11.7^17.5 Wm long, the margins thin, the membrane slightly granular, costa ectocolpi absent, margo absent, operculum absent; the endoaperture small in A. humbertii and 6 1.0 Wm in diameter, or lalongate in A. sambiranensis and PUE = (1.3^2.6)U(3.9^5.2) Wm; the tectum complete or nearly so, the sculpture striate (Plate XI, 73, 75, 77, 79). In thin section, the mesocolpial exine ca. 1.2^1.3 Wm thick ; the apertural endexine as thick (e.g., Plate XI, 72) as the mesocolpial ectexine or less so (e.g., Plate XI, 80); the non-apertural endexine irregular in thickness, granular (Plate XI, 74, 78); the foot layer appearing thick and uniform in low magni¢cation (e.g., Plate XI, 72) to irregular in high magni¢cation (e.g., Plate XI, 74, 76, 78), or thin and irregular (Plate XI, 80); the columellae short and thin (e.g., Plate XI, 72, 74) to stout (e.g., Plate XI, 80); the tectum slightly discontinuous (e.g., Plate XI, 72, 80), thick in A. humbertii (Plate XI, 72, 74, 76), thinner in A. sambiranensis (Plate XI, 78, 80). Discussion: Pollen of Amyrea has a striate surface, but it is only a super¢cial sculpture of the tectum (Plate XI, 74, 80). However, the rarity of this sculpture in Acalyphoideae plus the fact that Amyrea is the only genus in the subtribe to have bi¢d styles (Webster, 1994) suggests that Amyrea may not be as closely related to the remaining members of this subtribe as has been thought. Pollen of Spathiostemon javense (Plate XXV,

Plate II. SEM and TEM of pollen of tribe Acalypheae, subtribe Claoxylinae, Erythrococca. 10^12. 10. 11. 12. 13^15. 13. 14. 15.

E. menyharthii. Tangential section. Note variable shape and size of columellae in cross section, and that some columellae appear partially fused (arrowheads); compare with columellae in radial section, ¢g. 12. TEM. Mesocolpium-centered equatorial view of whole grain. SEM. Radial section across mesocolpium. Note irregular foot layer and apparent partial fusion of columellae (arrowhead); see also legend of ¢g. 10. TEM. E. oleracea. Somewhat oblique longitudinal section. Note irregularity of columellae, their elongation at the poles (upper left and right), and their occasional apparent fusion (arrowheads). TEM. Mesocolpium-centered equatorial view. SEM. Section through aperture. TEM.

PALBO 2467 11-10-02

256

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

191^197) has a linear sculpture in which lirae have cross striations. The subtribe Claoxylinae includes Erythrococca, Claoxylon, Claoxylopsis, Mareya, Mareyopsis, Discoclaoxylon, Micrococca and Amyrea (Plates I^XI). Most species of this subtribe have generalized pollen-colporate apertures, punctate-micro-

spinulose tecta (Plate I, 2, 4, 6; Plate II, 11; Plate IV, 25, 27, 29; Plate VI, 38, 41; Plate VII, 43, 45; Plate IX, 56; Plate X, 63, 65, 67, 70), or microscabrate (e.g., Plate IX, 58, 61), or more rarely microreticulate (Plate VIII, 49, 51) tecta, and typically strati¢ed exines (e.g., Plate I, 5; Plate II, 12; Plate III, 20; Plate IV, 22; Plate VII, 44; Plate

Plate III. SEM and TEM of pollen of tribe Acalypheae, subtribe Claoxylinae, Erythrococca. 16^18. 16. 17. 18. 19^21. 19. 20. 21.

E. welwitschiana. Equatorial section of 5-aperturate grain. Note small columellae. See legend of ¢g. 18. TEM. High magni¢cation of tectum along colpus near the pole. SEM Section across pole. Note variation in columellae diameter and length. TEM. Erythrococca species indet. Microscabrate tectum. SEM. Section across mesocolpium. TEM. Slightly oblique equatorial section of whole grain. In both SEM and TEM, pollen of this collection (Table 1) is somewhat di¡erent from the rest. TEM.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate IV (for description see p. 258).

PALBO 2467 11-10-02

257

258

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

IX, 59; Plate XI, 72). The most notable exceptions to the above characterization are found in the striate sculpture of pollen of Amyrea (Plate XI, 73, 75, 77, 79), and the exine structure (no foot layer and stout columellae) of Claoxylopsis (Plate VI, 42). The occasionally four- and ¢veaperturate grains found in Erythrococca and Claoxylon, and elongate polar columellae found in some species of Erythrococca are not unusual character states. Subtribe 30h. Lobaniliinae (Plate XII) 175. Lobanilia Radcli¡e-Smith, seven species endemic to Madagascar (Plate XII). Species examined : L. bakeriana (Plate XII, 81^ 84); L. claoxyloides (Plate XII, 85^89). Pollen prolate spheroidal in equatorial view; PUE = (19.4^27.3)U(17.5^23.4) Wm; three-colporate, the colpi 13.6^23.4 Wm long, narrow, costa ectocolpi absent (L. bakeriana) or weakly developed (L. claoxyloides), margo absent, operculum absent; the endoaperture lalongate, and PUE = (1.8^5.2)U(3.9^11.7) Wm; the tectum complete (e.g., Plate XII, 81, 83, 85), punctate-microspinulose (Plate XII, 83, 87). In thin section, the mesocolpial exine ca. 1.2^1.6 Wm thick; the apertural endexine ca. two/thirds as thick as the mesocolpial ectexine (L. claoxyloides, Plate XII, 86, 89) or seemingly not thickened (L. bakeriana, Plate XII, 84); the non-apertural endexine thin (Plate XII, 84, 89); the foot layer thin, irregular (e.g., Plate XII, 82, 88, 89); the columellae small and

thin (Plate XII, 86, 89), or slightly larger and more sparse (e.g., Plate XII, 82, 84); the tectum almost continuous (e.g., Plate XII, 82, 84) to less so (e.g., Plate XII, 86, 88). Discussion: Although Lobanilia is treated as a monogeneric tribe, no aspect of the pollen morphology or exine structure would reinforce this distinction. Subtribe 30i. Rottlerinae (Plates XIII^XXVI) 176. Mallotus Loureiro, 150 species in the Old World tropics, all but two are in Asia and Australia (Plates XIII^XVI). Species examined: M. acuminatus (Plate XII, 90^92); M. apelta (Plate XIII, 93, 94); M. baillonianus (Plate XIII, 95); M. barbatus (Plate XIII, 96^98); M. eriocarpus (Plate XIV, 99^101); M. fuscescens ; M. japonicus (Plate XIV, 102, 103); M. korthalsii (Plate XIV, 104^106); M. leucodermis (Plate XIV, 107); M. leucophyllus (Plate XV, 108, 109); M. microcarpus (Plate XV, 113); M. mollissimus (Plate XV, 110^112); M. muticus (Plate XV, 114) ; M. nepalensis ; M. nesophilus (Plate XV, 115); M. oreophilus (Plate XVI, 116^ 119) ; M. philippinensis (Plate XVI, 120); M. polyadenos; M. puber; M. subulatus (Plate XVI, 121, 122) ; M. sumatranus (Plate XVI, 123). Pollen oblate spheroidal to prolate spheroidal in equatorial view; PUE = (11.6^31.2)U(13.6U 29.9) Wm; three- (four-) colporate, the colpi 5.8^ 23.2 Wm long, costae ectocolpi present (e.g., Plate XIII, 90; Plate XIV, 104), sometimes poorly

Plate IV. SEM and TEM of pollen of tribe Acalypheae, subtribe Claoxylinae, Claoxylon (see p. 257). 22^25. 22. 23. 24. 25. 26^28. 26. 27. 28. 29, 30. 29. 30.

C. albicans. Radial section across mesocolpium and adjacent apertures. Note prominent apertural endexine. TEM. Mesocolpium-centered equatorial view of whole grain. SEM. Longitudinal section along colpus and near endoaperture. TEM. Tectum near the pole. SEM. C. cf. bakerianum. Longitudinal section. Compare relatively uniform length of columellae of this grain with columellae of Erythrococca. africana grain in Plate I, 1. TEM. Slightly oblique mesocolpium-centered equatorial view. SEM. Tangential section. Note that columellae are mostly cylindrical/ovate in cross section but variable in diameter. TEM. C. hainanense. Polar view of 4-aperturate grain. Note short colpi. SEM. Mostly radial section across mesocolpium. TEM.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

developed, margo absent, operculum absent; the endoaperture lalongate, PUE = ( 6 1.0^6.5)U ( s 1.0^15.5) Wm, costa endocolpi present, sometimes poorly developed; the tectum complete (e.g., Plate XIII, 91, 97; Plate XV, 108), punctate-microspinulose (e.g., Plate XIII, 91, 93, 95;

259

Plate XIV, 103, 107; Plate XV, 112, 115; Plate XVI, 117, 123). In thin section, the mesocolpial exine ca. 0.9^2.2 Wm thick; the apertural endexine equally (Plate XVI, 116) to almost twice (e.g., Plate XIII, 90; Plate XIV, 104; Plate XV, 113) as thick as the mesocolpial ectexine, sometimes

Plate V. SEM and TEM of pollen of tribe Acalypheae, subtribe Claoxylinae, Claoxylon. 31, 32. 31. 32. 33^35. 33. 34. 35. 36.

C. insulanum. Aperture-centered equatorial view. SEM. Mostly radial section across mesocolpium. Note variable diameter of columellae and irregularity of foot layer. TEM. C. longifolium. Polar view of four-colporate grain. SEM. Tangential section. Note variable shape and size of columellae in cross section. TEM. High magni¢cation of tectum at pole. SEM. C. lutescens. Longitudinal section along aperture. TEM.

PALBO 2467 11-10-02

260

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

granular on the inner surface (e.g., Plate XIII, 96; Plate XVI, 116); the non-apertural endexine granular, frequently irregular (e.g., Plate XIII, 98; Plate XIV, 100, 102; Plate XV, 110; Plate XVI, 116, 121); the foot layer thin (e.g., Plate XIII, 92,

98; Plate XIV, 102; Plate XV, 110, 113; Plate XVI, 116, 121); the columellae short, thin (e.g., Plate XIII, 90, 92, 94; Plate XIV, 100, 102, 106; Plate XV, 113; Plate XVI, 116, 121); the tectum continuous, thick, the predominant layer of the

Plate VI.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

ectexine (e.g., Plate XIII, 90, 92; Plate XIV, 100, 102; Plate XV, 109, 113; Plate XVI, 119, 121), ¢nely channeled and minutely spinulose. Discussion : Pollen of Mallotus can be characterized by a combination of thin foot layers, small columellae and thick tecta. See discussion after Octospermum (genus 183) at the end of the subtribe. 177. Deuteromallotus Pax et K. Ho¡mann, two species endemic to Madagascar (Plate XVII, 124^128). Species examined: D. acuminatus (Plate XVII, 125^128); Deuteromallotus species indet. (Plate XVII, 124). Pollen oblate spheroidal in equatorial view; PUE = (13.6^22.1)U(13.6^22.1) Wm; three-colporate, the colpi short and narrow, 5.2^11.6 Wm long, costa ectocolpi present, margo absent, operculum absent; the endoaperture lalongate, PUE = ( 6 1.0^2.6)U(3.9^11.6) Wm, costa endocolpi poorly developed; the tectum complete, microscabrate (Plate XVII, 125, 127). In thin section, the mesocolpial exine ca. 1.3^1.5 Wm thick; the apertural endexine ca. half to almost as thick as the mesocolpial ectexine; the non-apertural endexine thin, irregular, sometimes ¢nely granular; the foot layer thin, seemingly thicker near the apertures (Plate XVII, 126, 128); the columellae short ; the tectum mostly continuous, thick, and with tiny spinules. See discussion at end of subtribe (genus 183). 178. Cordemoya Baillon, three or four species in Asia (Plate XVII, 129^132).

261

Species examined : C. integrifolia (Plate XVII, 129^132). Pollen suboblate to oblate spheroidal in equatorial view; PUE = (13.6^20.8)U(15.5^22.1) Wm; three-colporate, the colpi short, 5.8^13.0 Wm long, costa ectocolpi present, margo absent, operculum absent; the endoaperture lalongate, PUE = (1.5^2.6)U(5.8^10.4) Wm; the tectum complete, microscabrate (e.g., Plate XVII, 129, 131). In thin section, the mesocolpial exine ca. 1.2^1.6 Wm thick; the apertural endexine ca. half as thick as the mesocolpial ectexine; the non-apertural endexine ¢nely lamellate (Plate XVII, 130), sometimes discontinuous; the foot layer thin (Plate XVII, 130), mostly consistent but sometimes thickened near the aperture (Plate XVII, 132 top); the columellae short, thin; the tectum continuous (Plate XVII, 132), variable in thickness. Discussion: The only di¡erence between pollen of Mallotus and that of Deuteromallotus and Cordemoya is a slight modi¢cation in the tectum sculpture^punctate-microspinulose in the former, and microscabrate (with short striae) in the latter two. 179. Coccoceras Miquel, three or four species in tropical Asia. No material. 180. Trewia L., one species ranging from India to southern China and Indonesia (Plate XVIII, 133^ 137). T. nudi£ora (Plate XVIII, 133^137). Pollen grains oblate spheroidal to prolate spheroidal in equatorial view; PUE = (15.5^ 29.9)U(13.6^29.9) Wm; three-colporate, the colpi

Plate VI. SEM and TEM of pollen of tribe Acalypheae, subtribe Claoxylinae, Claoxylon (37^39) and Claoxylopsis (40^42). 37^39. 37. 38. 39. 40^42. 40. 41. 42. 42 inset.

Claoxylon sitibundum. Section across equator of 5-aperturate grain. TEM. Slightly oblique aperture-centered equatorial view of whole grain. SEM. Radial section across mesocolpium. TEM. Claoxylopsis purpurescens. High magni¢cation of tectum in mesocolpium. SEM. High magni¢cation of tectum at aperture. The sculpture is not as distinct on this grain as the one in ¢g. 40 although both are from the same collection (Table 1). SEM. Mostly equatorial section of whole grain. Note very thin, or in some areas even absence of the foot layer, the stout columellae and the ‘fringed’ or slightly granular inner surface of tectum. TEM. Mesocolpium-centered equatorial view of whole grain. SEM.

PALBO 2467 11-10-02

262

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate VII. SEM and TEM of pollen of tribe Acalypheae, subtribe Claoxylinae, Mareya. 43^45. 43. 44.

45. 46^48. 46.

47. 48.

M. micrantha. Polar view of whole grain. SEM. Radial section. The very thick endexine (innermost, less electron-dense layer) indicates the section was close to and/or parallel with the long axis of a colpus. Note short columellae, well-developed and uniform foot layer, and thick continuous tectum. TEM. High magni¢cation of tectum. SEM. M. spicata. Mostly radial sections of two grains. Both grains show prominent endexines, unusual in Acalyphoideae, although the angle of section may have enhanced these layers in grain at right; at left, section is across colpus but above or below the endoaperture. TEM. Somewhat oblique equatorial section of whole grain. Note prominent foot layer and thick, almost continuous tectum. TEM. Tangential section. The columellae appear more similar to those in ¢g. 46 than in ¢g. 47. TEM.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

9.7^15.6 Wm long, costa ectocolpi present, margo absent, operculum absent; the endoaperture lalongate, PUE = (1.9^5.2)U(1.8^7.8, or lateral margins unclear) Wm; the tectum complete (Plate

263

XVIII, 133, 137), punctate and microspinulose. In thin section, the mesocolpial exine ca. 1.2 Wm thick; the apertural endexine equal in thickness to the mesocolpial ectexine ; the non-apertural en-

Plate VIII. SEM and TEM of pollen of tribe Acalypheae, subtribe Claoxylinae, Mareyopsis longifolia from two collections (Table 1). 49. 50. 51. 52. 53. 54. 55.

Mesocolpium-centered equatorial view of whole grain. SEM. Tangential section. Note that here and in ¢gs. 53 and 55, the columellae appear to have an expanded base that extends into the endexine and a foot layer is not identi¢able. TEM. High magni¢cation of tectum. SEM. Section across mesocolpium. In this grain, there is a continuous foot layer and a thin but continuous nonapertural endexine. TEM. Longitudinal section. See legend of ¢g. 50. TEM. High magni¢cation of tectum. Compare tectum of this collection with that from another collection illustrated in ¢gs. 49 and 51. SEM. Longitudinal section of whole grain. See legend of ¢g. 50. TEM.

PALBO 2467 11-10-02

264

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

dexine thin, granular on the inner surface, occasionally discontinuous; the foot layer very thin; the columellae short or diminutive (Plate XVIII, 134, 136) ; the tectum continuous, thick, predominant, with microspinules. See discussion after Octospermum (genus 183). 181. Neotrewia Pax et K. Ho¡mann, one species

in the Philippines and Celebes (Plate XVIII, 138^ 141). N. cumingii (Plate XVIII, 138^141). Pollen oblate spheroidal in equatorial view; PUE = (18.2^20.8)U(18.2^20.8) Wm; three-colporate, the colpi 9.1^13.0 Wm long, costa ectocolpi present, margo absent, operculum absent; the endoaperture lalongate, PUE = (1.3^2.6)U(6.5^

Plate IX. SEM and TEM of pollen of tribe Acalypheae, subtribe Claoxylinae, Mareyopsis (56, 57) and Discoclaoxylon (58^61). 56. 57. 58^61. 58. 59. 60. 61.

M. longifolia. Polar view of whole grain. Note short colpi. SEM. M. oligogyna. High magni¢cation of tectum over endoaperture. SEM. D. hexandrum. High magni¢cation of tectum. Note close similarity of this sculpture to that of one collection of Mareyopsis longifolia, Plate VIII, 54. SEM. Radial section across mesocolpium. All units, tectum, columellae and foot layer are well-developed and uniform. TEM. Mostly equatorial section of whole grain. Note slight outward arch of tectum in aperture margin (bottom). TEM. Mesocolpium-centered equatorial view of whole grain. SEM.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate X (for description see p. 268).

PALBO 2467 11-10-02

265

266

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XI (for description see p. 268).

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XII (for description see p. 268).

PALBO 2467 11-10-02

267

268

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate X. SEM and TEM of pollen of tribe Acalypheae, subtribe Claoxylinae, Micrococca (see p. 265). 62^64. 62. 63. 64. 65. 66^68. 66. 67. 68. 69^71. 69. 70. 71.

M. capensis. Longitudinal section. TEM. Polar view of 4-aperturate grain. SEM. Radial section across mesocolpium. Note elongate, mostly uniform columellae. TEM. M. holstii. Polar view of whole grain. SEM. M. mananarenta. Equatorial section of whole grain. Note outward arch of tectum in aperture at bottom. TEM. High magni¢cation of tectum near the pole. SEM. Slightly oblique section parallel with colpus. TEM. M. oligandra. Tangential section. Compare irregular outer surface of foot layer with that in the mostly radial section, ¢g. 71; note also that in cross section some columellae have very narrow diameters (arrowheads). TEM. Tectum of mesocolpium. SEM. Somewhat oblique radial section across mesocolpium. Note that columellae appear to be narrowed at midlength and/or expanded at the base (arrowhead); this modi¢cation agrees with the location of small cross sections in ¢g. 69. TEM.

Plate XI. SEM and TEM of pollen of tribe Acalypheae, subtribe Claoxylinae, Amyrea (see p. 266). 72^77. 72. 73.

74.

75. 76. 77. 78^80. 78. 79. 80.

A. humbertii from two collections (Table 1). Equatorial section of whole grain. Note prominent, albeit detached, apertural endexine and short columellae. TEM. Two whole grains. The upper is in oblique polar view and the lower in oblique equatorial view. Amyrea is the only member of Acalyphoideae to have a striate sculpture (but see also ‘‘segmented strands’’ of pollen of Spathiostemon, Plate XXV, 191^197, Plate XXVI, 198). SEM. Radial section across mesocolpium. Note that: striate surface is a supratectal (and super¢cial) sculpture of a thick continuous tectum; columellae are short and sparse, although the latter characterization is compromised by the outer surface of the foot layer, which in certain places, suggests stalagmite columellae; the foot layer is, for Acalyphoideae, well developed albeit irregular; the non-apertural endexine is sparsely and loosely granular. TEM. High magni¢cation of striate sculpture. SEM. Tangential section. The very irregular outer boundary of the foot layer agrees with its depiction in radial section, ¢g. 74; the outer extensions of the tectum are the lirae. TEM. Striate sculpture along colpus. TEM. A. sambiranensis. Tangential section. Compare with the more shallow tangential section in ¢g. 76. See legend of ¢g. 80. TEM. Oblique equatorial view of whole grain. SEM. Somewhat oblique equatorial section of whole grain. Although the striate sculpture of this species is clearly demonstrated in all three electron micrographs, the pollen di¡ers from that of A. humbertii in having a thinner less continuous tectum and thinner foot layer. TEM.

Plate XII. SEM and TEM of pollen of tribe Acalypheae, subtribe Lobaniliinae, Lobanilia (see p. 267). 81^84. 81. 82. 83. 84. 85^89. 85. 86. 87. 88. 89.

L. bakeriana. Mesocolpium-centered equatorial view of whole grain. SEM. Tangential section. The shape of the section indicates that its long axis is parallel with the long axis of the grain, and the elongation of the foot layer (arrows) is an artifact of this angle of section. TEM. High magni¢cation of tectum. SEM. Equatorial section of whole grain. Note sparse columellae and thin foot layer. TEM. L. claoxyloides. Aperture-centered equatorial view of whole grain. Note undulate surface; arrowhead points to raised area that probably corresponds to the thickened area in ¢g. 86 (arrowhead). SEM. Equatorial section of whole grain near equator. Note: diminutive, irregular columellae; the slight prominence of one aperture margin (arrowhead) indicate the section is across a raised area (¢g. 85). TEM. High magni¢cation of tectum. SEM. Tangential section. TEM. Radial section across mesocolpium. See legend of ¢g. 88. TEM.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

10.4) Wm; the tectum complete (Plate XVIII, 139), punctate-microspinulose, occasionally the surface undulate (Plate XVIII, 139). In thin section, the mesocolpial exine ca. 0.9^1.0 Wm thick; the apertural endexine slightly thicker than the mesocolpial ectexine, granular on the inner surface ; the non-apertural endexine irregular; the foot layer very thin (Plate XVIII, 140), consistent ; the columellae minute (Plate XVIII, 140); the tectum continuous, the predominant component (it is a reduction in tectum thickness that produces the depressed areas seen in SEM in 139), very sparsely microspinulose. See discussion of subtribe Rottlerinae after Octospermum (genus 183). 182. Rockinghamia Airy Shaw, two species in tropical Australia (Plate XIX). R. angustifolia (Plate XIX, 142^146); R. brevipes (Plate XIX, 147^150). Pollen prolate spheroidal to subprolate in equatorial view; PUE = (21.3^27.2)U(19.4^29.1) Wm; three-colporoidate, the colpi 17.5^23.2 Wm long, costa ectocolpi poorly developed if at all, margo absent, operculum absent; the endoaperture indistinct, 6 1.0 Wm in diameter ; the tectum incomplete (e.g., Plate XIX, 142, 150), microreticulate (Plate XIX, 146) to deeply punctate (e.g., Plate XIX, 142, 144). In thin section, the mesocolpial exine ca. 1.2 Wm thick; the apertural endexine ca. half to three quarters as thick as the mesocolpial ectexine ; the non-apertural endexine thick (Plate XIX, 149) to thin (Plate XIX, 143); the foot layer thick (Plate XIX, 149) to thin and irregular (Plate XIX, 143) ; the columellae stout (Plate XIX, 149) to less so (inset); the tectum discontinuous, the muri triangular in cross section (Plate XIX, 149) or not. See discussion of subtribe after Octospermum immediately below. 183. Octospermum Airy Shaw, one species in New Guinea (Plate XX). O. pleiogynum (Plate XX, 151^156). Pollen oblate spheroidal in equatorial view; PUE = (17.5^19.4)U(17.5^21.3) Wm; three-colp(or)ate, the colpi 7.8^11.6 Wm long, costa ectocolpi present, margo absent, operculum absent;

269

endoaperture 6 1.0 Wm in diameter; the tectum complete (Plate XX, 151, 154, 156), punctate-microspinulose (Plate XX, 154). In thin section, the mesocolpial exine ca. 0.9^1.2 Wm thick ; the apertural endexine ca. half as thick as the mesocolpial ectexine; the non-apertural endexine thin; the foot layer thin but consistent (e.g., Plate XX, 153) ; the columellae short ; the tectum continuous, thick, with minute spinules. Discussion: Pollen of the subtribe Rottlerinae, Mallotus, Deuteromallotus, Cordemoya, Trewia, Neotrewia, Rockinghamia and Octospermum, is characterized by thin foot layers, short columellae, and thick prominent tecta (e.g., Plate XIV, 102, 104; Plate XVIII, 136, 140; Plate XX, 153, 156), which are mostly punctate-microspinulose (e.g., Plate XV, 108, 114; Plate XVI, 123) or occasionally microscabrate (e.g., Plate XVII, 127, 129). Punt (1962) assigned a large number of genera to his Mallotus pollen type which he characterized, in part, as tricolporate or stephanocolporate, and having small endoapertures, a psilate or scabrate tectum, small columellae and an oblate shape. Besides Mallotus, he included Coelodiscus ( = Mallotus), Avellanita, Cordemoya, Trewia, Coccoceras, Deuteromallotus, Wetria, Adriana, Cleidion and Macaranga. The exine structure of Adriana and of most species of Cleidion is di¡erent from that of Mallotus, but not markedly so (Takahashi et al., 2000). Subtribe 30j. Acalyphinae (Plates XXI^XXIV) 184. Acalypha L., as many as 450 species with all but a few widespread in the tropics (Plates XXI^ XXIV). Species examined: A. acmophylla; A. communis (Plate XXI, 157); A. costaricensis (Plate XXI, 158, 159) ; A. diversifolia (Plate XXI, 160^162); A. grantii (Plate XXI, 163, 164); A. lanceolata (Plate XXII, 165^167); A. monococca (Plate XXII, 168^ 170) ; A. ornata (Plate XXII, 171^173); A. ostryaefolia; A. peduncularis; A. schinzii (Plate XXIII, 174^176); A. setosa (Plate XXIII, 177^179); A. stipulacea (Plate XXIII, 180^182); A. wilkesiana (Plate XXIV, 183^188). Pollen suboblate to oblate spheroidal in equatorial view (mostly, if not all, in tetrads in

PALBO 2467 11-10-02

270

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XIII (for description see p. 273).

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XIV (for description see p. 273).

PALBO 2467 11-10-02

271

272

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

A. wilkesiana, Plate XXIV, 184, see Table 2); PUE = (7.8^17.5)U(7.8^21.3) Wm; three- (¢ve-) colpate or colporate, the colpi very short, 1.5 Wm to 5.0 Wm long (measurements taken mostly from SEM), frequently the margins slightly pro-

truding (e.g., Plate XXI, 163; Plate XXII, 166, 168; Plate XXIII, 179), costa ectocolpi present but not always endexinous (Plate XXIV, 184, 185), operculum absent; endoaperture too small to measure; the tectum complete, the sculpture of

Plate XV (for description see p. 273).

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

273

Plate XIII. SEM and TEM of pollen of tribe Acalypheae, subtribe Rottlerinae, Mallotus (see p. 270). 90^92. 90. 91. 92. 93, 94. 93. 94. 95. 96^98. 96. 97. 98.

M. acuminatus. Equatorial section of whole grain. The endexine appears to be more electron-dense than the ectexine, but this may be an artifact of the staining preparation. TEM. High magni¢cation of tectum along colpus. SEM. Radial section. This ectexine structure ^ thin foot layer, short columellae and very thick tectum ^ characterizes most members of Mallotus (e.g., 96; Plate XIV, 100, 102; Plate XV, 113; Plate XVI, 116, 121). TEM. M. apelta. Equatorial view of portion of grain. The short colpus is shown almost full length. SEM. Longitudinal section parallel with colpus. TEM. M. baillonianus. Equatorial view of portion of grain. SEM. M. barbatus. Mostly equatorial section of whole grain. TEM. Tectum along colpus. SEM. Radial section across mesocolpium. The prominent and irregular non-apertural endexine and lower magni¢cation mask the similarity between this section and, e.g., the one in ¢g. 92. TEM.

Plate XIV. SEM and TEM of pollen of tribe Acalypheae, subtribe Rottlerinae, Mallotus (see p. 271). 99^101. 99. 100. 101. 102, 103. 102. 103. 104^106. 104. 105. 106.

107.

M. eriocarpus. Aperture-centered equatorial view of whole grain. SEM. Radial section of mesocolpium. Note thin foot layer, short columellae, and thick continuous tectum. TEM. High magni¢cation of tectum along colpus. SEM. M. japonicus. Radial section across mesocolpium. See legend of ¢g. 100. TEM. Aperture-centered equatorial view of part of grain. SEM. M. korthalsii. Equatorial section of whole grain. Note prominent apertural endexine, the electron density of which may be enhanced by staining. TEM. Oblique equatorial view of whole grain. SEM. Tangential section. To judge from the overall shape and the extensive foot layer-endexine, this longitudinal section could have started above the one end of a colpus and passed along the lateral margins, and exited the grain below the other end of the colpus. TEM. M. leucodermis. High magni¢cation of tectum. SEM.

Plate XV. SEM and TEM of pollen of tribe Acalypheae, subtribe Rottlerinae, Mallotus (see p. 272). 108, 109. 108. 109. 110^112. 110. 111.

112. 113. 114. 115.

M. leucophyllus. High magni¢cation of tectum along colpus. SEM. Tangential section. Note prominence of the tectum and irregular shape of many columellae in cross section. TEM. M. mollissimus. Radial section across mesocolpium. TEM. Tangential section. One interpretation of this complex section is that the orientation is slightly oblique to the polar axis in the area of an aperture: (from the top) the section started above the end of the colpus, then passed more deeply (in part because of the curvature of the grain) shallowly grazing the foot layers of two mesocolpia (dark included areas) making the foot layer appear more extensive than it is, then exiting where the margins are no longer separate. TEM. High magni¢cation of tectum at equator. The ‘bulge’ is the bridge of ectexine over the endoaperture. See also legend of ¢g. 111. SEM. M. microcarpus. Equatorial section of whole grain. TEM. M. muticus. High magni¢cation of tectum along colpus. SEM. M. nesophilus. Two grains in oblique view. SEM.

PALBO 2467 11-10-02

274

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XVI (for description see p. 277).

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

.

Plate XVII (for description see p. 277).

PALBO 2467 11-10-02

275

276

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

small irregularly shaped psilate areas separated by punctate-microspinulose areas (e.g., Plate XXII, 168, 173; Plate XXIII, 179, 181; Plate XXIV,

.

188). In thin section, the mesocolpial exine ca. 0.6^1.4 Wm thick; the apertural endexine thin, sometimes a fastigium of sorts delimited distally

Plate XVIII (for description see p. 277).

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

277

Plate XVI. SEM and TEM of pollen of tribe Acalypheae, subtribe Rottlerinae, Mallotus (see p. 274). 116^119. 116. 117. 118. 119.

120. 121, 122. 121. 122. 123.

M. oreophilus. Equatorial section of whole grain. The variable contour of the tectum re£ects the irregularity of the surface depicted in ¢gs. 117 and 118; note prominent non-apertural endexine. TEM. High magni¢cation of the tectum. See legend of ¢g. 116. SEM. Mesocolpium-centered equatorial view. SEM. Tangential section. The appearance of the various layers agrees with the radial section: at right the section passed through a thick area of the tectum; the small proportion of the area occupied by columellae re£ects their short length; the irregular, very thin foot layer is di⁄cult to delineate from the endexine. TEM. M. philippinensis. Polar view of whole grain. The surface of the tectum is uniform compared to the grain in ¢g. 118. SEM. M. subulatus. Radial section. Note thin foot layer. TEM. Aperture-centered equatorial view of part of grain. SEM. M. sumatranus. High magni¢cation of tectum along colpus. SEM.

Plate XVII. SEM and TEM of pollen of tribe Acalypheae, subtribe Rottlerinae, Deuteromallotus (124^128) and Cordemoya (129^ 132) (see p. 275). 124. 125^128. 125. 126. 127. 128. 129^132. 129. 130. 131. 132.

Deuteromallotus species indet. Tangential section. Note cylindrical shape of columellae in cross section, and irregular outer margin of foot layer. TEM. D. acuminatus. Polar view of whole grain. The short colpi scarcely extend into the apocolpium. SEM. Radial section across mesocolpium. Note thick tectum, irregular foot layer, and thin, but continuous nonapertural endexine. TEM. High magni¢cation of tectum. SEM. Section across aperture. TEM. C. integrifolia. Polar view of whole grain. The colpus length and sculpture are very similar to grains of Deuteromallotus, ¢g. 125. SEM. Section of mesocolpium near aperture. TEM. High magni¢cation of tectum. SEM. Somewhat oblique section of whole grain. Note: thickened foot layer near apertures (top and lower left), diminutive columellae, and almost continuous endexine; that the variable thickness of the tectum agrees with the surface depicted in ¢gs. 129 and 131. TEM.

Plate XVIII. SEM and TEM of pollen of tribe Acalypheae, subtribe Rottlerinae, Trewia (133^137) and Neotrewia (138^141) (see p. 276). 133^137. 133. 134. 135. 136. 137. 138^141. 138. 139. 140. 141.

T. nudi£ora. Polar view of whole grain. SEM. Radial section across mesocolpium. Note very short columellae, thin foot layer and very thick, continuous tectum. TEM. Aperture-centered equatorial view of whole grain. SEM. Longitudinal section of whole grain. TEM. High magni¢cation of punctate-microspinulose tectum. SEM. N. cumingii. Longitudinal section parallel with long axis of colpus. TEM. Aperture-centered equatorial view of whole grain. Note undulate surface of tectum. SEM. Equatorial section of whole grain. See legend of ¢g. 134. Grain appears partially attached (top) to another. TEM. Tectum. SEM.

PALBO 2467 11-10-02

278

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XIX.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

by the outward arching of the ectexine and proximally by the endexine extending across the gap (e.g., Plate XXI, 158, 162; Plate XXIII, 180; Plate XXIV, 185); the non-apertural endexine irregular, lamellate-granular (e.g., Plate XXI, 160; Plate XXII, 165, 169; Plate XXIII, 180, 182); the foot layer irregular, thin to sometimes thicker within a grain (e.g., Plate XXI, 160; Plate XXII, 169; Plate XXIII, 182), sometimes more uniformly thin (e.g., Plate XXIII, 176, 178), sometimes thickened at the aperture (e.g., Plate XXIII, 180; Plate XXIV, 185); the columellae mostly thin (Plate XXI, 158, 160, 162; Plate XXII, 169; Plate XXIII, 178, 180), variable in length but mostly short (e.g., Plate XXI, 164; Plate XXII, 167, 171; Plate XXIII, 174, 176, 178); the tectum continuous, of variable thickness within a grain (e.g., Plate XXI, 165; Plate XXIII, 180, 182; Plate XXIV, 187), with minute spinules. Discussion : Pollen of Acalypha can be distinguished from all remaining Acalyphoideae by a combination of oblate shape, small size (only one species had grains with a dimension greater than 20 Wm), and very small protruding apertures (e.g., Plate XXI, 159, 161, 163; Plate XXII, 166, 168, 172; Plate XXIII, 175, 179). The well-delimited fastigium-like chambers are formed by the protrusion of the entire ectexine, unlike the poorly delimited fastigia found in pollen of some Plukenetieae (e.g., Plate XXIX, 218, 226). The tectum sculpture (e.g., Plate XXII, 168, 173; Plate XXIII,

279

179; Plate XXIV, 188) is not found elsewhere in the subfamily. Pollen of Acalypha may have the shortest colpi in the subfamily, although grains of Cephalomappa (Takahashi et al., 2000) and some species of Cleidion (Takahashi et al., 2000) also have short colpi. The pollen tetrads of A. wilkesiana (Plate XXIV) are maintained by a common tectum (Plate XXIV, 183) between tetrad-members. This sample also contained ‘monads’, but, at least in LM, they appeared to have tears or mechanical damage suggesting they were from a disrupted tetrad. Subtribe 30k. Lasiococcinae (Plates XXV, XXVI) 185. Lasiococca Hooker f., three species in Asia (Plate XXV, 189, 190). Species examined: L. symphylliifolia (Plate XXV, 189, 190). This was a depauperate sample and a TEM preparation was not made. Pollen prolate spheroidal in equatorial view; PUE = (18.2^22.1)U(18.2^20.8) Wm; three-colporate, the colpi 13.0^14.3 Wm long, narrow, costae absent; margo absent, operculum absent; endoaperture small, poorly de¢ned, P = (1.0) Wm; the tectum complete, punctate-microspinulose (Plate XXV, 189, 190). Discussion: The tectum of Lasiococca symphylliifolia is not like that of Spathiostemon (Plate XXV, 195, 197), but neither does it have the densely spaced microechinae found in pollen of Homonoia (Plate XXVI, 202). The high magni¢cation SEM

Plate XIX. SEM and TEM of pollen of tribe Acalypheae, subtribe Rottlerinae, Rockinghamia. 142^146. 142. 143. 144. 145. 146. 147^150. 147. 148. 149. 150.

R. angustifolia. High magni¢cation of tectum. SEM. Equatorial section of whole grain. TEM. Mesocolpium-centered equatorial view of whole grain. SEM. Tangential section. Note irregularity of tectal perforations (larger to left) and irregularity of the outer surface of foot layer. SEM. High magni¢cation of tectum. Both this con¢guration and the one in ¢g. 142 were found in the collection of Gray 376. TEM. R. brevipes. Longitudinal section along colpus. TEM. Mesocolpium-centered equatorial view of whole grain. SEM. Slightly oblique radial section across mesocolpium. Note di¡erences between this exine structure and that of R. angustifolia: a thick non-apertural endexine, a thick foot layer and triangular shaped muri in cross section. TEM. High magni¢cation of deeply punctate tectum. Compare muri with those in ¢gs. 142 (similar) and 146 (di¡erent). SEM.

PALBO 2467 11-10-02

280

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

(Plate XXV, 190) looks like traces of pollenkitt may be obscuring some projecting elements, but this SEM is representative of the many of Lasiococca (Nowicke, unpublished data). See discussion of Lasiococcinae after Homonoia (genus 187). 186. Spathiostemon Blume, two species in Asia,

from Thailand to the Philippines and New Guinea (Plate XXV, 191^197; Plate XXVI, 198). Species examined : S. javensis (Plate XXV, 191^ 197; Plate XXVI, 198). Pollen prolate spheroidal in equatorial view, the poles slightly £attened; PUE = (18.2^24.7)U(16.9^ 22.1) Wm; three-colporate, the colpi 10.4^14.3 Wm

Plate XX. SEM and TEM of pollen of tribe Acalypheae, subtribe Rottlerinae, Octospermum pleiogynum. 151. 152. 153. 154. 155. 156.

Oblique view of whole grain. SEM. Radial section across colpus. Note slightly more elongate and irregular columellae near aperture. TEM. Radial section across mesocolpium. The exine structure is very similar to that of Mallotus: very thin foot layer, short columellae, and thick continuous tectum. TEM. High magni¢cation of tectum. SEM. Aperture-centered equatorial view of whole grain. Note short, narrow colpus. SEM. Mostly radial section of whole grain. TEM.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

281

Plate XXI. SEM and TEM of pollen of tribe Acalypheae, subtribe Acalyphinae, Acalypha. 157. 158, 159. 158. 159. 160^162. 160. 161. 162.

163, 164. 163. 164.

A. communis. Polar view of three-aperturate grain. SEM. A. costaricensis. Section across aperture. TEM. Portion of polar view. SEM. A. diversifolia. Slightly oblique radial section of mesocolpium. TEM. Equatorial view of whole grain. Very short or even porelike colpi are a characteristic of pollen of Acalypha (e.g., ¢gs. 163 and 166; Plate XXIII, 175, 179). SEM. Radial section of aperture. Note fastigium-like cavity formed by the slight protrusion of the actoapertural margins and an intact/continuous endexine (see also, e.g., Plate XXIII, 178, 180, Plate XXIV, 185). Note that granules on endexine have the same electron density as the ectexine. TEM. A. grantii. Two grains. Grain at left is in equatorial view, the one at right is in slightly oblique polar view. Note short, protruding apertures. SEM. Slightly oblique radial sections of two grains. TEM.

PALBO 2467 11-10-02

282

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

long, narrow, costa ectocolpi poorly developed, margo absent, operculum absent; the endoaper-

ture lalongate, PUE = (1.3^2.6)U(6.5^10.4) Wm, costa endocolpi poorly developed; the tectum

Plate XXII.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

complete, the supratectal sculpture of narrow lirae having shallow cross-striations, appearing minutely segmented (e.g., Plate XXV, 193, 195, 197; Plate XXVI, 198). In thin section, the mesocolpial exine ca. 0.9 Wm thick; the apertural endexine thin (Plate XXV, 192); the non-apertural endexine thin but consistent ; the foot layer very thin (e.g., Plate XXV, 194); the columellae very short, sparse (Plate XXV, 192, 194, 196); the tectum continuous, with scattered punctae (Plate XXV, 192, 194, 196), the outer surface having ridges (Plate XXV, 191). Discussion : Pollen of Spathiostemon ^ a complete tectum with a supratectal sculpture of narrow lirae having numerous super¢cial cross-striations (Plate XXV, 191, 194, 196) ^ has no counterpart in any remaining examined taxa, including Homonoia, of Acalyphoideae. Although Ferna¤ndezGonza¤lez and Lobreau-Callen (1996) show grains with this tecta as belonging to Homonoia javanensis, the collection they examined, Ramos 42809, is actually Spathiostemon javensis according to van Welzen (1998). But see discussion after Homonoia immediately below. 187. Homonoia Loureiro, two species in Asia, one of which is endemic to India (Plate XXVI, 199^ 203). Species examined : H. riparia (Plate XXVI, 199^ 203). Pollen oblate spheroidal to prolate spheroidal in equatorial view, the poles slightly £attened; PUE = (18.2^27.3)U(20.8^29.9) Wm; three-

283

(four-) colporate, the colpi 6.5^11.6 Wm long, sometimes with a granular membrane, costae ectocolpi present, margo absent, operculum absent; the endoaperture lalongate, PUE = (2.6^3.9)U (9.1^14.3) Wm, costa endocolpi present; the tectum complete, the sculpture of densely spaced, uniform microechinae (elements), randomly arranged (Plate XXVI, 201, 202). In thin section, the mesocolpial exine ca. 1.0 Wm thick; the apertural endexine at least as thick as the mesocolpial ectexine, granular on the inner surface (Plate XXVI, 200, 203); the non-apertural endexine well-developed, consistent (Plate XXVI, 199); the foot layer very thin; the columellae very short, sparse (Plate XXVI, 199, 200); the tectum continuous, thick, the predominant component (Plate XXVI, 199, 203), composed of elongate, individual pointed rods or elements (Plate XXVI, 199, 200, 203), vertically oriented. Discussion: The pointed tectal projections of pollen of Homonoia have a very di¡erent origin from the microspinules found in so many taxa in Acalyphoideae, and for this reason we designated them microechinae. Some problematical results were clari¢ed by the exsiccate in van Welzen’s recent (1998) publication. As part of a revision of the subtribe Lasiococcinae, he recognized two species of Homonoia, H. riparia, common in Southeast Asia, and H. retusa Mueller Argoviensis, found only in Central India. One of the collections we examined, McGregor 247, is cited as H. riparia. Another collection, Ramos 17661, identi¢ed at US as H. riparia, is listed by van

Plate XXII. SEM and TEM of pollen of tribe Acalypheae, subtribe Acalyphinae, Acalypha. 165^167. 165. 166. 167. 168^170. 168. 169. 170. 171^173. 171. 172. 173.

A. lanceolata. Section across mesocolpium with proximity to aperture (top). Note irregularity of all layers although the angle of section is only slightly oblique. TEM. Tectum and aperture. Arrowheads mark what could be the upper and lower margins of the endoaperture. SEM. Section of whole grain including one aperture. See legend of Plate XXI, 162. TEM. A. monococca. Partial polar view including protruding aperture. SEM. Slightly oblique radial section. Note irregularity and poor separation of endexine and foot layer. TEM. Polar view of 4-aperturate grain. SEM. A. ornata. Section of whole grain. TEM. Polar view of three-aperturate grain. SEM. High magni¢cation of tectum. SEM.

PALBO 2467 11-10-02

284

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XXIII.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Welzen as Spathiostemon javensis, and the pollen agrees with his determination (Plate XXV, 196, 197). Ferna¤ndez-Gonza¤lez and Lobreau-Callen (1996) show grains (¢gs. 141, 149) from an Indian collection identi¢ed as H. retusa, including a fracture in which the tectum appears to be composed of discrete elements. This collection, Hooker and Thomson 1589, was, apparently, not seen by van Welzen. The specimen of H. riparia examined by Ferna¤ndez-Gonza¤lez and Lobreau-Callen (1996) and cited by them as Bensekom et al., 14, is probably van Beusekom et Phengkhlai 14, and van Welzen (1998) con¢rms this identi¢cation. Pollen of this collection is illustrated in three SEMs (1996, ¢gs. 150^152), of which only the grain in ¢g. 152 appears to have the tectal elements in a random arrangement like pollen of Homonoia. The pollen of Homonoia and Spathiostemon is a curious mixture of similarities and distinctions. These grains share exines with small, very sparse columellae, thin or even threadlike foot layers, and consistent non-apertural endexines. Both genera have a tectal sculpture of minute elements ^ but in Spathiostemon, the elements are formed by shallow cross striations on very narrow, supratectal lirae (e.g., Plate XXV, 191, 194, 197) and thus appear in rows, whereas in Homonoia, each surface element is the apex of an individual rod, randomly arranged (Plate XXVI, 199, 200, 202) and whose length is the thickness of the tectum. The

285

tectal elements of pollen of Homonoia are very di¡erent in origin, shape and organization from those of Spathiostemon, and from all other taxa of Acalyphoideae which have punctate-microspinulose tecta. For this reason, the pointed elements of pollen of Homonoia were designated as microechinae. None of the high magni¢cation SEMs of Lasiococca (Nowicke, unpublished data) show the uniformly microechinate tectum of Homonoia. Punt (personal communication) suggested that the tectum of Lasiococca (Plate XXV, 189, 190) is similar to that of Deuteromallotus and Cordemoya (Plate XVII, 127, 129, 131). Data from TEM could resolve the relationships of Lasiococca. Unfortunately, the material of Lasiococca was insuf¢cient for all three preparations. Tribe 31. Plukenetieae (Plates XXVII^XLIII) The 16 genera of Plukenetieae are segregated into three subtribes, Plukenetiinae (seven genera), Tragiinae (eight) and Dalechampiinae (one). The subtribes Plukenetiinae and Tragiinae have a wide diversity of exine structure, apertures, and tectum morphology. The distribution of pollen types transgresses existing generic boundaries in Plukenetiinae and Tragiinae ^ some types are present in more than one genus, and some genera have two or more pollen types. One caveat which applies to our pollen descriptions of subtribes Plukenetiinae and Tragiinae is

Plate XXIII. SEM and TEM of pollen of tribe Acalypheae, subtribe Acalyphinae, Acalypha. 174^176. 174. 175. 176. 177^179. 177. 178. 179. 180^182. 180.

181. 182.

A. schinzii. Slightly oblique equatorial section of grain. TEM. Group of pollen grains. Note orientation of apertures in center grain. SEM. Slightly oblique radial section across mesocolpium. The foot layer is very thin, the columellae small and sparse, and the tectum is the predominant layer. TEM. A. setosa. Slightly oblique polar view of three-aperturate grain. SEM. Equatorial section of grain. The columellae are numerous, short and very thin. See also legend of Plate XXI, 162. TEM. Aperture and tectum. The colpus is 6 1.0 Wm. SEM. A. stipulacea. Radial section across equator of grain. Pollen of this species has the thickest exine of all members of Acalypha examined; note also the thin foot layer and thick, lamellate endexine, characteristics best illustrated in the mesocolpium at left. In LM the apertures appear crassimarginate. TEM. Polar view of whole grain. SEM. Radial section across mesocolpium. TEM.

PALBO 2467 11-10-02

286

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

the paucity or even absence of size data. Pollen samples were frequently insu⁄cient for all three preparations and sometimes LM were not made. After acetolysis, grains were collapsed or ruptured, making measurements di⁄cult or impossible and certainly suspect. For some samples, the only dimension given is the equatorial diameter taken from polar view.

We give a brief summary of the palynological disposition of taxa of Plukenetiinae and Tragiinae at the beginning of each subtribe. The pollen types are designated by a speci¢c epithet of one of the included species. Subtribe XXXI)

31a.

Plukenetiinae

(Plates

XXVII^

Plate XXIV. SEM and TEM of pollen of tribe Acalypheae, subtribe Acalyphinae, Acalypha wilkesiana. 183. 184. 185. 186. 187. 188.

Section near center of tetrad showing areas of shared tectum of three tetrad-members. TEM. Rhomboidal tetrad. The short colpi appear weakly crassimarginate because of costae. LM. Radial section across aperture. Note thickened foot layer which agrees with the tangential section (187), although both sections have passed obliquely through the aperture. TEM. Polar view of a 5-aperturate monad or of a disrupted tetrad-member. Tangential section. See legend of ¢g. 185. TEM. High magni¢cation of tectum. SEM.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XXV (for description see p. 288).

PALBO 2467 11-10-02

287

288

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Pollen of seven of the eight genera (no material of Sphaerostyles) was examined. Plukenetia has two pollen types, designated Conophora and Loretensis. Two monotypic genera, Eleutherostigma and Romanoa, have the Conophora pollen type and are described with that group. A third monotypic genus, Vigia, has the Loretensis pollen type and is described with that group. Pollen of Haematostemon and of Angostyles are described separately. The pollen description of Astrococcus is based on LM and SEM. The Conophora type is restricted to subtribe Plukenetiinae. 188. Haematostemon (Mueller-Argoviensis) Pax et K. Ho¡mann, two species in the New World Tropics (Plate XXVII, 204^207). Species examined: H. guianensis (Plate XXVII, 204^207). Pollen oblate spheroidal in equatorial view; PUE = (33.0^38.8)U(36.9^46.6) Wm; three-colpate, the colpi 25.2^33.0 Wm long, the margins (costa?) irregularly thickened (e.g., Plate XXVII, 205 insets), the layers sometimes separating and forming a fastigium of sorts (Plate XXVII, 205 lower inset; 207) ; margo absent, operculum absent, the tectum complete, microcrotonoid. In thin section, the mesocolpial exine ca. 1.5 Wm thick ; the apertural endexine sometimes poorly distinguished from the foot layer and the two ir-

regularly thickened (Plate XXVII, 205 arrowheads); the non-apertural endexine thin, discontinuous (Plate XXVII, 204); in the mesocolpium, the foot layer almost as thick as the tectum (Plate XXVII, 204); the columellae prominent, occasionally branched, a granular area near the tectum (Plate XXVII, 204, 205, 207); the tectum continuous or nearly so, about as thick as the mesocolpial foot layer, sometimes granular on the inner surface. Discussion: The pollen of Haematostemon guianensis is similar in structure to the Conophora type of Plukenetia (genus 193), but it di¡ers in tectal morphology and the presence of irregular thickenings of the endexine^foot layer along the colpal margins. This characteristic is conspicuous in LM (Plate XXVII, 205 insets). Whether these irregular margins should be called costae is debatable. See discussion after Angostyles (genus 190). 189. Astrococcus Bentham, one species in Brazil and Venezuela (Plate XXVII, 208^209). A. cornutus (Plate XXVII, 208, 209). A depauperate sample examined only in LM and SEM. Pollen diameter E = 31.2^40.3 Wm; three-colpate, the colpal margins protruding (Plate XXVII, 208, 209), intact, costae absent, margo absent, operculum absent; the tectum complete, microcrotonoid or scabrate (Plate XXVII, 208).

Plate XXV. SEM and TEM of pollen of tribe Acalypheae, subtribe Lasiococcinae, Lasiococca (189, 190) and Spathiostemon (191^197) (see p. 287). 189, 190. 189. 190. 191^197. 191.

192. 193. 194.

195. 196. 197.

L. symphylliifolia. Equatorial view of whole grain. SEM. High magni¢cation of tectum. SEM. S. javensis. Tangential section. Compare outer and inner area of individual lira (strand) with the surface depicted in ¢gs. 195 and 197 ^ the cross striations (transverse lines) are super¢cial; note also perforations in continuous (solid) area of the tectum ^ they are produced by gaps in the tectum (connecting arrow to ¢g. 194); see legend of ¢g. 194. TEM. Longitudinal section of most of a grain. TEM. Mesocolpium-centered equatorial view of whole grain. SEM. Mostly radial section. Note: that lirae and striae (grooves) are a supratectal sculpture of a thick continuous tectum, and that the cross striations are an even more super¢cial sculpture of the lirae; sparse columellae and very thin, even threadlike foot layer; and substantial non-apertural endexine. TEM. Tectum and part of colpus. SEM. Radial section across mesocolpium. This collection, Ramos 17661, was originally identi¢ed as Homonoia javensis, but the pollen is that of Spathiostemon (see text). TEM. High magni¢cation of tectum. Grain from Ramos 17661. SEM.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

See discussion after Angostyles, immediately below. 190. Angostyles Bentham, one species in Amazonian Brazil (Plate XXVIII, 210^213).

289

A. longifolia (Plate XXVIII, 210^213) Pollen prolate spheroidal in equatorial view; PUE = (45.5^49.4)U(42.9^46.8) Wm; three-colpate or occasionally four-colpate, the colpi 19.5^ 28.6 Wm long, the margins ragged (Plate XXVIII,

Plate XXVI. SEM and TEM of pollen of tribe Acalypheae, subtribe Lasiococcinae, Spathiostemon (198) and Homonoia (199^ 203). 198. 199^203. 199.

200. 201. 202. 203.

S. javensis. Mesocolpium-centered equatorial view of whole grain. (See legend of Plate XXV, 191, 196). SEM. H. riparia. Radial section. The exine is similar to that of Spathiostemon in having short, sparse columellae and thin foot layer but di¡ers markedly in the structure of the tectum, which is composed of individual, cylindrical elements or rods whose length forms the thickness of the tectum. TEM. Section of aperture. Note prominent apertural endexine with granular inner surface, accumulation of ectexinous granules over the endoaperture, and ¢ne channels separating the elongate tectal elements. TEM. Mesocolpium-centered equatorial view of whole grain. SEM. Tectum along colpus. A microechinate sculpture formed by random organization of acutely tipped tectal elements. SEM. Section of whole grain. TEM.

PALBO 2467 11-10-02

290

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XXVII (for description see p. 292).

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XXVIII (for description see p. 292).

PALBO 2467 11-10-02

291

292

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

210, 211), irregularly thickened (costa?) and appearing like pollen of Haematostemon (Plate XXVII, 205, upper inset), the exine layers scarcely separating (no fastigia), operculum absent; the tectum complete or nearly so, microrugulate (Plate XXVIII, 210). In thin section (based on limited TEM data), the mesocolpial exine ca. 1.5^1.7 Wm thick ; the foot layer and endexine di⁄cult to delineate from each other (Plate XXVIII, 212, 213) and irregularly thickened along the margins of the colpi; the columellae short, less than a quarter of the total exine thickness; the tectum continuous (Plate XXVIII, 212), irregu-

larly thickened, the outer surface re£ecting the crenate muri. Discussion: Pollen of Angostyles, of Haematostemon and of Astrococcus share certain characteristics : the ¢rst two genera have an irregular thickening of the foot layer^endexine along the colpal margins, and the last two have a microcrotonoid tectum. But pollen of Angostyles di¡ers from that of Haematostemon in the absence of a granular area under the tectum, the presence of short columellae and microrugulate sculpture. We examined the same collections of Astrococcus cornutus (Liesner 8693) and of H. guianen-

Plate XXVII. SEM and TEM of pollen of tribe Plukenetieae, subtribe Plukenetiinae, Haematostemon (204^207) and Astrococcus (208, 209) (see p. 290) 204^207. 204. 205.

205 inset top. 205 inset bottom. 206. 207. 208, 209. 208. 209.

H. guianensis. Radial section across mesocolpium. Note prominent columellae, thick foot layer, and slightly granular area under the tectum. TEM. Oblique longitudinal section of whole grain. The prolate shape is an artifact of preparation. At upper left, the irregularity and gaps (arrowheads) in the thickened foot layer^endexine indicates that the section has passed parallel with and close to a colpus. TEM. Equatorial view of grain showing irregular thickening (arrowheads) of endexine^foot layer. LM. Polar view. Arrowheads point to out-of-focus thickenings of colpal margins. LM. Mesocolpium-centered equatorial view of whole grain. The appearance of opercula is misleading. SEM. Radial section across aperture. Both here and in 205 inset bottom, the apertures have a fastigium of sorts. TEM. A. cornutus. Colpus and adjacent tectum. SEM. Polar view of whole grain. Note protruding aperture margins, that, in LM, give the appearance of fastigia. SEM.

Plate XXVIII. SEM and TEM of pollen of tribe Plukenetieae, subtribe Plukenetiinae, Angostyles (210^213) and Romanoa (214^ 217) (see p. 291). 210^213. 210. 211. 212. 213.

214^217. 214.

215. 216. 217.

A. longifolia. Aperture and tectum. SEM. Oblique view of whole grain. SEM. Radial section. Note short thin columellae, thick foot layer^endexine, and absence of granular layer at interface of columellae and tectum. TEM. Oblique section of whole grain. Although this particular TEM does not emphasize the irregular thickening of the foot layer^endexine (arrowheads), in LM this characteristic is as conspicuous as in Haematostemon guianensis (Plate XXVII, 205 insets). TEM. R. tamnoides. Conophora pollen type (see legend of Plukenetia conophora, Plate XXX, 231^235). Radial section across mesocolpium. Note elongate, highly irregular columellae, some of which could be interpreted as being fused (much larger); that inner surface of tectum has ¢ne fringe in addition to loose granules. TEM. Tangential section. Although not readily apparent at this low magni¢cation, there is a layer of ¢ne granules just inside the tectum. TEM. Polar view of colpus. SEM. Radial section across aperture. TEM.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

sis (Fanshawe 2869) as Gillespie (1994) did, and the same collection of A. longifolia (Ducke 23528)

293

as Punt (1962). Gillespie (p. 342) describes pollen of both Astrococcus and Haematostemon as hav-

Plate XXIX (for description see p. 295).

PALBO 2467 11-10-02

294

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XXX (for description see p. 295).

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

ing ‘‘unusually thickened aperture margins with the upper and lower exine layers separating to form an elongate chamber (similar to a vestibulum)’’. Our LM preparation of Liesner 8693 consists of six grains, all of which have protruding, intact aperture margins (exactly like those in the SEM, Plate XXVII, 209), but not the separation of exine layers nor the irregular thickenings of foot layer^endexine along the colpal margins. Punt (1962, pl. XIII, 3) demonstrated a slight separation of exine layers in a line drawing of Astrococcus cornutus. The descriptions and interpretations of the pollen by Punt, by Gillespie and by the present authors are not in complete agree-

295

ment, but neither Punt nor Gillespie examined pollen in TEM, Gillespie’s preparation of Fanshawe 2869 was unacetolysed, and our conclusions regarding Astrococcus are based on a depauperate sample. The classi¢cation of the tectum sculpture of Haematostemon and Astrococcus is controversial. Punt (personal communication) believes they are scabrate, but we think that these tecta are a variant of a croton pattern in which most of the triangular distinction has been lost. We do not believe that Haematostemon and Astrococcus are any more closely related to the Crotonoideae than, e.g., Angostyles or Micrococca. It should

Plate XXIX. SEM and TEM of pollen of tribe Plukenetieae, subtribe Plukenetiinae, Eleutherostigma (218^222) and Plukenetia (223^226) (see p. 293). 218^222. 218. 219. 220. 221. 222.

223, 224. 223. 224. 225, 226. 225. 226.

E. lehmannianum. Conophora pollen type (see legend of Plukenetia conophora, Plate XXX, 227^230). Equatorial section of whole grain. Note fastigia-like morphology of colpal margins. TEM. High magni¢cation of punctate tectum and ragged/granular colpus margins. SEM. Radial section across mesocolpium. Note irregularly thickened tectum and prominent granules between it and the elongate irregular columellae. TEM. Oblique view of whole grain. Note irregular margins of colpi. SEM. Tangential section. Compare the layers here with their appearance in radial section (¢g. 220); the tectum is irregularly thickened; the granular area (just inside the tectum) is more extensive at top and right; the columellae have various shapes in cross-section, and the high proportion of the total area they occupy re£ects their elongation. TEM. P. brachybotrya. Loretensis pollen type (see legend of Plukenetia loretensis, Plate XXX, 231^235). Polar view of whole grain. Note short but wide colpi. SEM. Tectum showing crenate muri. SEM. P. polyadenia. Conophora pollen type (see legend of Plukenetia conophora, Plate XXX, 227^230). Polar view of whole grain. SEM. Oblique section of whole grain including two apertures with fastigia-like margins. TEM.

Plate XXX. SEM and TEM of pollen of tribe Plukenetieae, subtribe Plukenetiinae, Plukenetia (see p. 294). 227^230. 227. 228. 229. 230. 231^235. 231. 232. 233. 234. 235.

P. conophora. Conophora pollen type (colpate, punctate tecta, branched columellae and/or granular area under tectum). Equatorial section of whole grain. Note fastigia-like colpi margins. TEM. High magni¢cation of tectum at pole. SEM. Radial section across mesocolpium. TEM. Polar view of whole grain. SEM. P. loretensis. Loretensis pollen type (colpate, reticulate tecta, unremarkable exine structure). Oblique section of mesocolpus. Note that foot layer appears to consist of minute columellae (short arrows), which agrees with the irregular surface depicted in SEM, ¢g. 234 (short arrows). TEM. Polar view of whole grain. SEM. Somewhat oblique radial section of mesocolpium. Arrow connects globule (free columella?) here with corresponding structure in ¢g. 234. See also legend of ¢g. 231. TEM. Tectum. Note weakly crenate muri and scattered globules or free columellae on foot layer. See legend of ¢g. 233. SEM. Radial section across aperture. Compare aperture with those in SEM, ¢g. 232. TEM.

PALBO 2467 11-10-02

296

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

be remembered that the vast majority of Thymelaeaceae have pollen with a croton (not croto-

noid) tectum and the croton structure (Nowicke, 1994 ; Nowicke and Skvarla, unpublished data),

Plate XXXI.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

and this family is not considered to be closely related to Euphorbiaceae on other bases. 191. Romanoa Trevisan, one species in Brazil. (Plate XXVIII, 214^217). See Conophora pollen type under Plukenetia (genus 193). 192. Eleutherostigma Pax et K. Ho¡mann, one species in Colombia and Ecuador. (Plate XXIX, 218^222). See Conophora type under Plukenetia below. 193. Plukenetia L., 10 species in the neotropics, three in Africa and one each in Madagascar and southeast Asia (Plate XXIX, 223^226; Plates XXX, XXXI). Two pollen types, Conophora and Loretensis. 193. Plukenetia, (i) Conophora pollen type (punctate tecta, prominent and sometimes branched columellae, granular area under tectum). Taxa included and examined: Eleutherostigma lehmannianum (Plate XXIX, 218^222); P. conophora (Plate XXX, 227^230); P. polyadenia (Plate XXIX, 225, 226); P. volubilis (Plate XXXI, 239^ 242); Romanoa tamnoides (Plate XXVIII, 214^ 217). Pollen suboblate to oblate^spheroidal to subprolate in equatorial view; PUE = (39.0^48.1)U

297

(31.2^52.0) Wm, see Table 2 ; three-colpate, the margins frequently ragged (e.g., Plate XXVIII, 216; Plate XXIX, 219, 225; Plate XXX, 230), margo absent, costae absent, operculum absent; the tectum complete and punctate (e.g., Plate XXVIII, 216; Plate XXIX, 219, 225; Plate XXX, 230; Plate XXXI, 240, 242). In thin section, the mesocolpial exine 1.5^2.5 Wm thick; the apertural endexine very thin, seemingly separated from the ectexine and sometimes forming a fastigium of sorts (e.g., Plate XXIX, 218, 226; Plate XXX, 227) ; the non-apertural endexine very thin (e.g., Plate XXVIII, 214; Plate XXX, 229) to slightly thicker (e.g., Plate XXXI, 241) ; the foot layer irregular (e.g., Plate XXVIII, 214) to less so (e.g., Plate XXIX, 220), occasionally ¢nely channeled (e.g., Plate XXVIII, 214), sometimes discontinuous (Plate XXX, 229) ; the columellae elongate, the predominant layer, frequently irregular (e.g., Plate XXVIII, 214; Plate XXIX, 220; Plate XXXI, 239, 241), sometimes branched and/or fused (e.g., Plate XXVIII, 214; Plate XXXI, 241), frequently with ‘loose’ granules near the tectum (e.g., Plate XXIX, 220, 222; Plate XXXI, 239, 241); the tectum continuous, sometimes irregularly thickened (e.g., Plate XXIX, 220, 226; Plate XXX, 229; Plate XXXI, 241), ¢nely granular or ‘fringed’ on

Plate XXXI. SEM and TEM of pollen of tribe Plukenetieae, subtribe Plukenetiinae, Plukenetia (236^242) and Vigia (243, 244). 236^238. 236. 237.

238. 239^242. 239.

240. 241.

242. 243, 244. 243. 244.

P. penninervia. Loretensis pollen type (see legend of Plukenetia loretensis, Plate XXX, 231^235). Polar view of whole grain. Note protruding colpal margins. SEM. Part of an oblique section. Note well-de¢ned circles or ovate shapes of columellae in cross section (compare with tangential section in ¢g. 239); note also the highly variable size diameters of columellae, some of which must be free standing (not attached to the tectum) because there are so few cross sections near the tectum. This section emphasizes the poor de¢nition of the foot layer and its separation from the non-apertural endexine. TEM. High magni¢cation of crenate muri. SEM. P. volubilis. Conophora pollen type (see legend of Plukenetia conophora, Plate XXX, 227^230). Tangential section. At inner margin of tectum note ¢nely granular zone; in contrast to ¢g. 237, many columellae have highly irregular shapes in cross section; near the foot layer, the outlines of some cross sections suggest fusion of two or three columellae. TEM. High magni¢cation of punctate tectum. SEM. Radial section. Compare the layers here with those in ¢g. 239 ^ the columellae are prominent and irregular in shape, the foot layer is irregular on the outer margin, the tectum varies in thickness, as does the extent of the granular area. TEM. Mesocolpium-centered equatorial view of whole grain. SEM. Vigia serrata. Loretensis pollen type (see legend of Plukenetia loretensis, Plate XXX, 231^235). Polar view of whole grain. SEM. High magni¢cation of tectum. SEM.

PALBO 2467 11-10-02

298

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

the inner surface (e.g., Plate XXVIII, 214; Plate XXX, 229).

Discussion: The Conophora type is particularly well-de¢ned by both morphology (complete punc-

Plate XXXII.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

tate tecta, e.g., Plate XXVIII, 216; Plate XXIX, 219, 221, 225; Plate XXX, 228, 230) and by structure (granular interface between tectum and columellae and/or ¢nely branched columellae, e.g., Plate XXVIII, 214, 215; Plate XXIX, 220, 222, 226; Plate XXXI, 239, 241). The structural characteristic is variable. In some taxa, e.g., Romanoa, the inner surface of the tectum seems minutely ¢mbriolate (Plate XXVIII, 214), whereas in others, e.g., Eleutherostigma (Plate XXIX, 220), there are discrete granules. Another characteristic, but of perplexing signi¢cance, is the paucity of apertural endexine, but perhaps the elaborate colpal margins, when intact, compensate for thin or reduced endexines. The pollen data, particularly from TEM, support Gillespie’s (1993) reduction of Eleutherostigma lehmannianum to Plukenetia lehmannianum and would support the reduction of a second monotypic genus, Romanoa, which Webster (1994, p. 93) characterized as ‘‘very close to Plukenetia and questionably distinct’’. 193. Plukenetia, (ii) Loretensis pollen type (reticulate exines, mostly crenate muri, unremarkable exine structure). Taxa included and examined: P. brachybotrya (Plate XXIX, 223, 224) ; P. loretensis (Plate XXX, 231^235); P. penninervia (Plate XXXI, 236^238); Vigia serrata (Plate XXXI, 243, 244). Pollen mostly suboblate in equatorial view;

299

PUE = (20.8^48.1)U(22.1^52.0) Wm; three-colpate, the colpi either narrow and with protruding granular margins (Plate XXXI, 236, 243) or wider and with the membrane irregularly granular (e.g., Plate XXIX, 223, sometimes with fastigia (Vigia), costae absent, operculum absent; the tectum incomplete and reticulate (e.g., Plate XXIX, 223, 224; Plate XXX, 232, 234; Plate XXXI, 236, 243, 244), the muri frequently crenate (e.g., Plate XXIX, 224; Plate XXX, 234; Plate XXXI, 238). In thin section, the mesocolpial exine ca. 3.0 Wm thick (P. loretensis); the apertural endexine variable, seemingly poorly developed, the apertural area frequently ¢lled with very ¢ne granules of the same electron-density as the ectexine (Plate XXX, 235); the foot layer thin, ¢nely channeled, sometimes appearing as very short/minute columellae (Plate XXX, 231, 233, 235; Plate XXXI, 237) ; the columella length equal to twice the tectum thickness (e.g., Plate XXX, 233, 235); the tectum discontinuous (e.g., Plate XXX, 231, 233; Plate XXXI, 237), the surface minutely ridged (Plate XXXI, 237). Discussion: Pollen grains of the Loretensis type share reticulate exines, frequently with crenate muri, and an unremarkable structure. Although the SEM of a grain of Vigia serrata shows apertures with a prominent margin (Plate XXXI, 243), other grains in this sample have apertures more similar to that illustrated for P. brachybotrya (Plate XXIX, 223). The SEM of a grain of P.

Plate XXXII. SEM and TEM of pollen of tribe Plukenetieae, subtribe Tragiinae, Cnesmone (245^248) and Megistostigma (249, 250). 245, 247. 245. 247. 246, 248. 246. 248. 249, 250. 249.

249 inset. 250.

C. javanica. Malaccense pollen type (see legend of Megistostigma malaccense, 249, 250). Mesocolpium-centered equatorial view of whole grain. Note large, poorly de¢ned, but colpate-like apertures. SEM. High magni¢cation of tectum. SEM. C. tonkinensis. Malaccense pollen type (see legend of Megistostigma malaccense, 249, 250). Radial section. At top the prominently thickened endexine suggests proximity to aperture. TEM. Section of whole grain. The exine consists mostly of the tectum. TEM. M. malaccense. Malaccense pollen type (poorly de¢ned apertures, thin exines with sparse columellae, threadlike or endexine/foot layers, thin tecta). Section of whole grain including 4 aperture areas (arrows) where £ecks of ectexine rest on a slightly thickened endexine. Every unit of the exine is reduced ^ the columellae are short and very sparse, the foot layer^endexine together are threadlike, and the tectum is irregular and occasionally discontinuous. The intine (not present after acetolysis) could well be the only continuous layer of the pollen wall. TEM. Portion of grain including three circular aperture areas (arrows) whose morphology agrees with the structure. SEM. High magni¢cation of tectum of a grain from a second collection (see Table 1). SEM.

PALBO 2467 11-10-02

300

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

penninervia (Plate XXXI, 236) also shows such apertures, but the appearance of colpi of all pollen is in£uenced, at least in part, by the vagaries of preparations/acetolysis. The pollen data support the reduction of the monotypic genus, Vigia, to P. serrata (Gillespie, 1993). See discussion of subtribe Plukenetiinae below. 194. Vigia Vellozo, one species in South America (Plate XXXI, 243, 244). See Loretensis pollen type under Plukenetia (193). Discussion : All pollen of subtribe Plukenetiinae (Plates XXVII^XXXI) is three-colpate, suggesting that simple apertures are not a derived condition. Most taxa of Plukenetiinae have pollen similar to one of the two types found in Plukenetia. The Conophora type, punctate tecta and branched columellae and/or a granular area beneath the tectum, is found in two monotypic genera, Romanoa (Plate XXVIII) and Eleutherostigma (Plate XXIX), and P. conophora (Plate XXX), P. polyadenia (Plate XXIX) and P. volubilis (Plate XXXI). Thus, the pollen data from TEM reinforce the reduction of Eleutherostigma lehmannianum to a species of Plukenetia (Gillespie, 1993), and support Webster’s (1994) opinion that Romanoa tamnoides is questionably distinct from Plukenetia. The Loretensis pollen type, reticulate exines with crenate muri and an unremarkable exine structure, occurs in P. brachybotrya (Plate XXIX), P. loretensis (Part 4, Plate XXX), P. penninervia (Plate XXXI) and Vigia (Plate XXXI). The pollen data support the reduction of Vigia serrata to Plukenetia serrata by Gillespie (1993). Pollen of Haematostemon is similar to the Conophora type but di¡ers in having a microcrotonoid (see discussion after the genus description) tectum and an irregular thickening of the foot layer^endexine along the margins of the colpi. The monotypic status of Angostyles is supported by the distinction of the pollen-microrugulate tectum, thick foot layer, short unbranched columellae. Pollen of Astrococcus, which has a tectum that could be described as microcrotonoid, was not examined in TEM. Subtribe 31b. Tragiinae (Plates XXXII^XL, 245^ 320)

Pollen of seven of the eight genera assigned to Tragiinae were examined (no material of Sphaerostyles). Cnesmone and Megistostigma have similar pollen, the Malaccense type, and similar staminate £owers, are likely to be actually closely related (Gillespie, 1994) and are described together. Pollen of Tragiella is described separately, as is pollen of Platygyna. The genus Tragia has three pollen types found in more than one examined species, the Ramosa, Lukafuensis and Urens types; pollen of three other species of Tragia is di¡erent from the above types and from each other and are described individually. There are two pollen types in Acidoton, the inaperturate Microphyllus and the aperturate Nicaraguensis. Pollen of Pachystylidium is described separately. 195. Cnesmone Blume, 10 species in Asia (Plate XXXII, 245^248). See Malaccense pollen type under Megistostigma immediately below. 196. Megistostigma Hooker f., ¢ve species in the tropics of southeast Asia (Plate XXXII, 249^250, Plate XXXIII, 251, 251 inset). Malaccense pollen type (poorly de¢ned apertures, reduced exines of small sparse columellae, mostly granular foot layers, thin tecta). Taxa included and examined: Cnesmone javanica (Plate XXXII, 245, 247) ; C. tonkinensis (Plate XXXII, 246, 248); Megistostigma burmanicum; M. cordatum ; M. malaccense (Plate XXXII, 249, 250, 251 inset); Megistostigma species indet. (Plate XXX, 251). Pollen ovate to spheroidal ; PUE = see discussion below; three- to four-aperturate or occasionally inaperturate, the apertures poorly de¢ned, colpus-like (C. javanica Plate XXXII, 245) or more porelike (M. malaccense, Plate XXXII, 249 inset) and irregularly distributed, the aperture membranes covered with islands or £ecks of ectexine (Plate XXXII, 245, 249, 249 inset); the tectum complete, irregularly punctate and microspinulose (Plate XXXII, 245, 247, 249 inset, 250). In thin section, the exine ca. 0.9^1.0 Wm thick; the apertural endexine thickened, sometimes with more electron-dense granules (Plate XXXII, 246, 249) ; the non-apertural endexine threadlike; the foot layer of sparse granules (Plate XXXII, 246,

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

251 inset) to threadlike to seemingly absent (Plate XXXII, 248, 249) ; the columellae very sparse (e.g., Plate XXXII, 248, 249; Plate XXXIII, 251; 251 inset); the tectum mostly continuous (e.g., Plate XXXIII, 251), irregular in thickness (Plate XXXII, 246, 248, 249). Discussion : Exines of the Malaccense pollen type are characterized by a reduced structure : threadlike or discontinuous foot layer^endexines, small sparse columellae and irregular tecta. Although described as inaperturate by Punt (1962), both SEM (Plate XXXII, 245, 249 inset) and TEM (Plate XXXII, 249 arrows; Plate XXXIII, 251) show areas of thickened endexine under islands or £ecks of ectexine. According to Gillespie (1994), the separation of Cnesmone and Megistostigma appears to break down as more collections become available. See also discussion after Tragiella (genus 198). We have pollen size data for one species of Cnesmone (Table 1) and none of Megistostigma. Punt (1962), whose samples were acetolysed, gave a dimension of 55 Wm for C. javanica, and 37^56 Wm for two species of Megistostigma. Gillespie (1994), whose samples were rehydrated and not acetolysed, gave dimensions of ca. 45.4^60 Wm for four species of Cnesmone, and 42.5^58.5 Wm for two species of Megistostigma. 197. Sphaerostylis Baillon, two species endemic to Madagascar. No material. According to Punt (1962), pollen of Sphaerostylis belongs to his Plukenetia verrucosa subtype, which corresponds to our Loretensis type. 198. Tragiella Pax et K. Ho¡mann, ¢ve species in Africa (Plate XXXIII, 252^258). Species examined: T. friesiana (Plate XXXIII, 252^255); T. natalensis (Plate XXXIII, 256^258). Pollen prolate^spheroidal (Plate XXXIII, 254) in equatorial view; PUE = (27.3^40.7)U(27.2^ 36.9) Wm; three-colpate, the colpi 15.6^33.0 Wm long, narrow, the colpal margins (tectum) sometimes outwardly arched near the equator, costae absent, operculum absent, margo absent; the tectum incomplete (e.g., Plate XXXIII, 254, 256), ¢nely reticulate, the muri crenate (e.g., Plate XXXIII, 252, 258). In thin section, the mesocol-

301

pial exine ca. 2.0 Wm thick, sometimes the exine along the colpus expanded or appearing thickened at the equator (T. friesiana, Plate XXXIII, 253, arrows) ; the apertural endexine thin (Plate XXXIII, 253, 255); the non-apertural endexine thin (e.g., Plate XXXIII, 253); the foot layer thin (e.g., Plate XXXIII, 253), sometimes irregular; the columellae irregular in length ; the tectum discontinuous (Plate XXXIII, 253, 255, 257), somewhat variable in thickness, outwardly arched at equator of colpal margins (Plate XXXIII, 253, open arrows). Discussion: Pollen of T. friesiana, and to a lesser extent that of T. natalensis, have arched or expanded tecta at the colpal margins near the equator of the grain, a characteristic best seen in intact grains in polar view in LM. Those TEMs which include a section across the colpus at the equator of the grain show an outward arching of the tectum but with few or no supporting columellae. Pollen of Tragiella shares protruding colpal margins, a nearly identical exine structure (compare the whole grain section in Plate XXXIII, 253 with that in Plate XXXVI, 279), as well as crenate muri with the Lukafuensis type of Tragia (Plate XXXVI, 277^280). Pollen of the Loretensis type of Plukenetia, although somewhat similar, lacks the protruding tectum at the equator and has a channeled foot layer. According to Webster (1994), Tragiella is considered extremely close to Sphaerostylis and doubtfully separable, and Megistostigma has been combined with Sphaerostylis by some authors. The latter reduction would not be supported by the pollen data since, according to Punt (1962), pollen of Sphaerostylis is similar to the Loretensis type of Plukenetia, which is very di¡erent from that of Megistostigma/Cnesmone (Plate XXXII, Plate XXXIII, 251). 199. Platygyna Mercier, seven species endemic to Cuba (Plate XXXIV). Species examined : P. dentata (Plate XXXIV, 259) ; P. hexandra (Plate XXXIV, 260, 261); P. parvifolia (Plate XXXIV, 262^267). Pollen spheroidal to elliptical; inaperturate; longer sideUshorter side = (27.3^41.6)U(27.2^ 39.0) Wm (occasional tetrads in P. parvifolia);

PALBO 2467 11-10-02

302

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XXXIII (for description see p. 304).

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XXXIV (for description see p. 304).

PALBO 2467 11-10-02

303

304

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

the exine sometimes appearing pitted in LM; ‘‘tectate’’ and punctate (e.g., Plate XXXIV, 259, 261) or intectate and the surface microrugulate (Plate XXXIV, 263, 265). In thin section, the exine ca. 1.5 Wm thick; the endexine of one or two threads with occasional granules (Plate XXXIV, 260, 262, 264, 267); the foot layer absent; the columellae very large, prominent, irregular in shape (e.g., Plate XXXIV, 260 arrows, 262, 266),

sometimes appearing to hang from the tectum (P. hexandra, Plate XXXIV, 260), sometimes two (or more?) columellae appearing fused (Plate XXXIV, 262 solid arrowheads), sometimes expanded distally (e.g., Plate XXXIV, 262 open arrowheads, 264 open arrowheads); tectum present (Plate XXXIV, 260) and discontinuous or the tectum (seemingly) absent (Plate XXXIV, 262, 264, 266).

Plate XXXIII. SEM and TEM of pollen of tribe Plukenetieae, subtribe Tragiinae, Megistostigma (251) and Tragiella (252^258) (see p. 302). 251. 251 inset. 252^255. 252. 253. 254. 255. 256^258. 256. 257. 258.

Megistostigma species indet. Malaccense pollen type (see legend of Megistostigma malaccense, Plate XXXII, 249, 250). Section of whole grain. Note four large apertural areas where endexine is thickened. TEM. M. malaccense. Radial section. Note absence of foot layer except for a few electron-dense granules on a threadlike endexine. TEM. T. friesiana. High magni¢cation of tectum along colpus. SEM. Equatorial section of whole grain. Open arrows point to the areas of arched tectum along margin of colpus at equator. TEM. Mesocolpium-centered equatorial view of whole grain. SEM. Oblique section across an aperture above or below endoaperture. Note dearth of endexine here and in ¢g. 253 as well. TEM. T. natalensis. Slightly oblique polar view of whole grain. SEM. Section across aperture, above or below endoaperture. TEM. High magni¢cation of tectum. Note the crennate muri here and in ¢gs. 252, 254 and 256. SEM.

Plate XXXIV. SEM and TEM of pollen of tribe Plukenetieae, subtribe Tragiinae, Platygyna. Pollen inaperturate (see p. 303). 259. 260, 261. 260.

261. 262^267. 262.

263. 264. 265.

266.

267.

P. dentata. Punctate tectum. SEM. P. hexandra. Radial section across exine. In this species, the pollen is considered tectate or semitectate. Note: the irregularity and prominence of columellae (lower arrow) and that some appear to hang from the tectum (upper arrow); the absence of a foot layer; the endexine of two thin lamellae and granules. TEM. Inaperturate whole grain. SEM. P. parvifolia. Slightly oblique radial sections of two (damaged?) grains. The exines consist of large irregular columellae, some of which (solid arrows) appear to have fused. Note also: that some columellae are slightly expanded distally (open arrows), suggesting a partial tectum; the endexine of one or two threads interspersed with granular material. TEM. Inaperturate whole grain. SEM. Radial section. Note that outer surface agrees with that depicted in SEM, ¢g. 265, and that the exine structure agrees with that in the tangential section, ¢g. 266. See legend of ¢g. 262. TEM. High magni¢cation of exine surface. The surface here appears more continuous than in ¢g. 262, but the more open spacing of columellae in those two sections may be the result of an arti¢cial bending of exine during embedding; or, conversely, the more continuous surface in ¢g. 265 may be the result of partial collapse. SEM. Tangential section. It consists almost entirely of irregular columellae cut at various levels ^ even the endexine is represented by only a faint gray ‘smudge’ in the center; but this tangential section agrees perfectly with the radial ones in ¢gs. 262 and 264. TEM. Oblique section of whole grain. The explanation of the appearance of a tectum is the angle of section and that partial collapse of the grain has compressed the columellae. SEM.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XXXV (for description see p. 307).

PALBO 2467 11-10-02

305

306

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XXXVI (for description see p. 307).

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Discussion : Pollen of Platygyna is among the most distinctive of all Acalyphoideae: inaperturate grains with an exine consisting primarily of large, even massive, columellae sporadically linked to one or two threads of endexine (Plate XXXIV, 260, 262, 264). The foot layer is absent, as is a tectum in one species, P. parvifolia. According to Punt (1962, p. 58) ‘‘small circular thickenings occur on the endexine’’. In LM, the exine does appear pitted and sometimes minutely scratched as well. Pollen of Platygyna shares a similar exine structure and absence of apertures with some species of Acidoton (Plate XXXIX, 306^311). See discussion after that genus (201). Webster (1994) mentions the reduction of Platygyna to a synonym of Tragia by Alain, but that

307

treatment has no palynological merit. See discussion at the end of subtribe Tragiinae (after Pachystylidium, genus 202). 200. Tragia Plumier ex L., 125 species in Africa. and America (Plates XXXV-XXXVIII). Three pollen types occur in more than one species in this genus. Three species have pollen which is distinct from these types and from each other, and are described separately. 200. Tragia, (i) Lukafuensis pollen type (three-colpate, arched colpal margins at equator, ¢nely reticulate tecta, crenate muri, unremarkable exine structure). Taxa examined and included: T. gardneri (Plate XXXV, 274^276); T. lukafuensis (Plate XXXVI,

Plate XXXV. SEM and TEM of pollen of tribe Plukenetieae, subtribe Tragiinae, Tragia (see p. 305). 268^270. 268. 269. 270. 271^273. 271. 272. 273. 274^276. 274. 275. 276.

T. bailloniana. Polar view of whole grain. SEM. Tangential section. TEM. High magni¢cation of tectum. SEM. T. cf. cordifolia. Parts of two grains. At left, the extent of the granular area indicates that the section is probably parallel with the long axis of a colpus; at right, the section is from some area of mesocolpium. TEM. Tectum near pole showing parts of two large apertural areas. SEM. Tangential section. Note that the sparse distribution and small size of columellae agree with their appearance in grain to right in ¢g. 271. TEM. T. gardneri. Lukafuensis pollen type (see legend of Tragia lukafuensis, Plate XXXVI, 277^280). High magni¢cation of tectum showing crenate muri. SEM. Mostly radial section. Note continuity of foot layer and endexine. TEM. Mostly equatorial section of whole grain. Note outwardly arched tectum at equator. TEM.

Plate XXXVI. SEM and TEM of pollen of tribe Plukenetieae, subtribe Tragiinae, Tragia (see p. 306). 277^280. 277. 278. 279. 280. 281, 282. 281. 282. 283^286. 283. 284. 285. 286.

T. lukafuensis. Lukafuensis pollen type (colpate, ¢nely reticulate tecta, unremarkable exine structure). Radial section across aperture. TEM. Aperture-centered equatorial view. Note crenate muri and protruding tecta (open arrows) of colpal margin. SEM. Equatorial section of whole grain. Note short thin columellae, thin foot layer, incomplete tectum and arched tectum at colpal margins (open arrows) at equator of grain. TEM. Radial section. Here the magni¢cation is high enough to show the crenate surface of the muri. TEM. T. mitis. Lukafuensis pollen type (see legend of T. lukafuensis, 277^280). High magni¢cation of tectum. SEM. Polar view of whole grain. Note elaboration of exine along colpus, a characteristic that often occurs in pollen of Plukenetieae. SEM. T. ramosa. Ramosa pollen type (colpate, intectate, exines of large stout columellae and thick foot layers). Tangential section. TEM. Surface. Large columellae in frontal view, a few of which appear fused. SEM. Radial section. A highly unusual exine structure ^ a thick channeled foot layer and stout columellae that are sometimes expanded distally. TEM. Oblique section of whole grain. The apertural endexine is very thin. See also legend of ¢g. 285. TEM.

PALBO 2467 11-10-02

308

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

277^280); T. mitis (Plate XXXVI, 281, 282); T. pungens (Plate XXXVII, 287^290); T. spathulata (Plate XXXVII, 291, 292). Pollen oblate spheroidal to prolate spheroidal in equatorial view; PUE = (19.5^33.8)U(22.1^ 32.5) Wm; three-colpate, 13.0^26.0 Wm long, the margins sometimes expanded or arched near the equator (e.g., Plate XXXVI, 278 arrows, 279 arrows); costae absent, margo absent, operculum absent; the tectum incomplete and ¢nely reticulate to microreticulate (e.g., Plate XXXVI, 278, 281; Plate XXXVII, 287, 291), the muri mostly crenate (e.g., Plate XXXV, 274; Plate XXXVII, 289, 291) or minutely spinulose in T. mitis (Plate XXXVI, 281, 282). In thin section, the mesocolpial exine ca. 1.6^2.0 Wm thick, at the equator the tectum along the colpal margins outwardly arched (e.g., Plate XXXVI, 279 arrows; Plate XXXVII, 290 arrows); the apertural endexine thin (e.g., Plate XXXV, 276; Plate XXXVI, 277, 279; Plate XXXVII, 290) ; the non-apertural endexine thin but consistent (e.g., Plate XXXV, 275; Plate XXXVI, 280; Plate XXXVII, 288) ; the foot layer thin (e.g., Plate XXXVI, 279, 280; Plate XXXVII, 288, 290, 292); the columellae small (Plate XXXVI, 277, 279; Plate XXXVII, 290) to somewhat larger (Plate XXXV, 275, 276); the tectum discontinuous (e.g., Plate XXXVI, 279; Plate XXXVII, 288, 290, 292), protruding near the equator, the muri surface re£ecting the crenate sculpture. Discussion : The Lukafuensis pollen type is characterized by three colpi with arched or protruding tecta at the equator, and ¢nely reticulate to microreticulate tecta having crenate muri (T. mitis has minutely spinulose muri, Plate XXXVI, 281). These species of Tragia have pollen almost identical to that of Tragiella (Plate XXXIII, 252^ 258). All ¢ve species with the Lukafuensis pollen type, T. gardneri, T. lukafuensis, T. mitis, T. pungens and T. spathulata, are assigned to sect. Tagira Mueller Argoviensis by Pax and Ho¡mann (1931). This large section is segregated into nine subsections and none of the above species are in the same one. 200. Tragia, (ii) Ramosa pollen type (three-col-

pate, intectate, exines of large columellae and thick foot layers). Taxa included and examined : T. ramosa (Plate XXXVI, 283^286); T. stylaris (Plate XXXVII, 296) ; T. urticaefolia (Plate XXXVIII, 297^299); T. volubilis (Plate XXXVIII, 304, 305). Pollen prolate spheroidal to oblate spheroidal in equatorial view; PUE = (26.0^32.5)U(24.7^ 35.1) Wm; three-colpate, the colpi 9.7^22.1 Wm long, costae absent, margo absent, operculum absent ; intectate, the surface columellate (e.g., Plate XXXVI, 284; Plate XXXVIII, 299, 305), sometimes the columellae with one to three minute spinules. In thin section, the exine ca. 1.1^1.3 Wm thick; the apertural endexine very thin (e.g., Plate XXXVI, 286; Plate XXXVII, 296) ; the nonapertural endexine thin but consistent (Plate XXXVI, 285; Plate XXXVIII, 298) to almost absent (Plate XXXVIII, 304); the foot layer prominent, the thickness one third to half the columellae length, ¢nely channeled and ridged on the outer surface (Plate XXXVI, 283, 285; Plate XXXVIII, 298, 304) ; the columellae stout, even massive (e.g., Plate XXXVI, 285; Plate XXXVIII, 298), frequently widened at the outer margin (e.g., Plate XXXVI, 285; Plate XXXVIII, 298), sometimes appearing to extend to the non-apertural endexine (e.g., Plate XXXVIII, 298). Discussion: The intectate exine of the Ramosa pollen type consists of a thick foot layer and large columellae. This structure is unique within the Acalyphoideae, and thus, except to emphasize a close relationship among the species in which it occurs (all assigned to the large New World sect. Tragia), no other relationships are suggested. We do not agree with the pollen description of T. volubilis by Suarez-Cervera et al. (2001, p. 84) as ‘‘Ectexine tectate, baculate-clavate’’. ‘‘In the infratectum, closely spaced very small inclusions are observed (¢g. 6C^E) which probably represent extremely reduced columellae. The foot layer is absent or not distinguishable.’’ They consider minute perforations at the interface of the ectexine and endexine as (supposedly) delineating extremely reduced columellae. What they regard as a tectum, we think is a thick, channeled foot layer with occasional perforations. Such foot layers occur elsewhere in pollen with unequivocal colum-

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XXXVII (for description see p. 311).

PALBO 2467 11-10-02

309

310

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XXXVIII (for description see p. 311).

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

ellae, e.g., Mercurialinae (Takahashi et al., 2000, Plates XX, XXI), Rockinghamia (Plate XIX, 147, 149), the Conophora pollen type (Plate XXVIII, 214), the Loretensis type (Plate XXX, 231, 233) and other species of the Ramosa type (Plate XXXVI, 285; Plate XXXVIII, 298). 200. Tragia, (iii) Urens pollen type (poorly de¢ned apertures, thick microscabrate tecta, foot layer absent). Taxa examined and included: T. smallii; T. urens (Plate XXXVIII, 300^303). Pollen spheroidal in equatorial view; PUE =

311

(25.2^33.0)U(23.2^29.1) Wm; probably threeaperturate, short colpi or pores with di¡use margins; costae absent; margo absent; operculum absent ; the tectum complete, microscabrate (Plate XXXVIII, 301, 303). In thin section, the mesocolpial exine ca. 1.0 Wm thick; the apertural endexine irregularly thickened, the membrane covered with irregularly shaped £ecks of ectexine (Plate XXXVIII, 300, grain at right); the non-apertural endexine thin, irregular but continuous (Plate XXXVIII, 300, 302); the foot layer seemingly absent (Plate XXXVIII, 300, 302); the columellae

Plate XXXVII. SEM and TEM of pollen of tribe Plukenetieae, subtribe Tragiinae, Tragia (see p. 309). 287^290. 287. 288. 289. 290. 291, 292. 291. 292. 293^295. 293. 294. 295. 295 inset. 296.

T. pungens. Lukafuensis pollen type (see legend of T. lukafuensis, Plate XXXVI, 277^280). Slightly oblique polar view. SEM. Radial section across mesocolpium. All ectexine units ^ columellae, incomplete tectum and foot layer ^ are irregular, but the grain may have been damaged. See also legend of ¢g. 289. TEM. High magni¢cation along colpus. The crenate muri are irregular in shape and slightly arched as compared with T. lukafuensis (Plate XXXVI, 277^280). SEM. Equatorial section across colpus. Open arrows mark protruding tectum of colpal margins at the equator. TEM. T. spathulata. Lukafuensis pollen type (see legend of T. lukafuensis, Plate XXXVI, 277^280). High magni¢cation of tectum along colpus. SEM. Sections of two grains. At left, grain shows prominent endexine with elements of ectexine above it; at right, grain sectioned at an oblique angle. Both have very thin foot layers and thin non-apertural endexines. TEM. T. sellowiana. Radial section. Note here and in ¢g. 295, the thin, even fragile exine structure: short stout columellae, a threadlike foot layer^endexine, and a poorly de¢ned, thin discontinuous tectum. TEM. High magni¢cation of tectum. The irregular surface of SEM agrees with those of the TEMs in ¢gs. 293, 295, 295 inset. SEM. Section of whole inaperturate grain. Exine consists mostly of columellae. TEM. Mostly radial section. See legends of ¢gs. 293, 294. TEM. T. stylaris. Ramosa pollen type (see legend of T. ramosa, Plate XXXVI, 283^286). Section across colpus. TEM.

Plate XXXVIII. SEM and TEM of pollen of tribe Plukenetieae, subtribe Tragiinae, Tragia (see p. 310). 297^299. 297. 298. 299. 300^303. 300. 301. 302. 303. 304, 305. 305.

T. urticaefolia. Ramosa pollen type (see legend of T. ramosa, Plate XXXVI, 283^286). Aperture-centered equatorial view of three-colpate grain. SEM. Radial section. The exine consists mostly of massive columellae that are expanded distally and sometimes extend as such to a thin endexine proximally; note channels and perforations of foot layer. TEM. High magni¢cation of surface. SEM. T. urens. Urens pollen type (apertures poorly de¢ned, tectum thick and continuous, foot layer absent). Radial sections of two grains. At left, note absence of foot layer; at right the irregular ectexine and thick endexine suggest an apertural area. TEM. High magni¢cation of tectum. SEM. Radial section. The foot layer is absent, the non-apertural endexine irregular but prominent, and the columellae appear narrowed nearer the tectum. TEM. Whole grain. Apertures are poorly de¢ned in this species. SEM. T. volubilis. Ramosa pollen type (see legend of T. ramosa, Plate XXXVI, 283^286). This is the same pollen type as in ¢gs. 297^299, but the distribution of the columellae is less dense. TEM. Polar view of whole grain. SEM.

PALBO 2467 11-10-02

312

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

variable in diameter, wider near the base (Plate XXXVIII, 302), appearing to project through the tectum (Plate XXXVIII, 302) ; the tectum more or less continuous, irregular in thickness. Discussion : We segregated these two species as a pollen type even though we lack TEM data for T. smallii because the SEMs of that species are indistinguishable from those of T. urens, and because, according to Gillespie (1994), they are the only two in sect. Leptobotrys. But this is a tenuous recognition and disposition on our part. Pollen of T. bailloniana, of T. cordifolia, and of T. sellowiana does not appear related to each other nor to any other species of Tragia examined here. They are described individually. Tragia bailloniana (Plate XXXV, 268^270) Pollen oblate spheroidal in equatorial view; PUE = (34.9^42.2)U(38.8^48.5) Wm; three-colpate, the colpi 19.4^27.2 Wm long; costae absent, margo absent, operculum absent; the tectum incomplete and ¢nely reticulate to microreticulate (Plate XXXV, 270), the lumina larger at the poles (Plate XXXV, 268). TEM data too limited to justify a description. Tragia cf. cordifolia (Plate XXXV, 271^273) Pollen prolate spheroidal in equatorial view; PUE = (28.6^32.5)U(26.0^31.2); three-colpate, the colpi 9.7^15.5 Wm long; costae absent, margo absent, operculum absent; the tectum complete, microscabrate (Plate XXXV, 272). In thin section, the mesocolpial exine ca. 1.2 Wm thick; the apertural endexine scarcely thickened; the non-apertural endexine very thin but consistent ; the foot layer average; the columellae average; the tectum thick, continuous or nearly so. Discussion : T. cordifolia is assigned to sect. Tagira which has the Lukafuensis pollen type characterized by reticulate exines (e.g., Plate XXXVI, 277^ 280). It may be that this collection, Schlieben 7556, is misidenti¢ed. Schlieben, who provided the determination, obviously, had some misgivings. Tragia sellowiana (Plate XXXVII, 293^295, 295 inset) Pollen spheroidal to slightly elliptical in shape;

LUS = (36.4^45.5)U(36.4^44.2) Wm; inaperturate; the tectum irregular, with punctae and microspinules (Plate XXXVII, 294), the surface uneven. In thin section the exine ca. 1.0^1.2 Wm thick; the endexine poorly developed; the foot layer (and endexine?) threadlike, discontinuous (Plate XXXVII, 293, 295 inset); the columellae the predominant component (Plate XXXVII, 293, 295), stout, appearing acutely tipped; the tectum discontinuous, irregular, thin (Plate XXXVII, 293, 295, 295 inset). Discussion: The inaperturate pollen of T. sellowiana has exines consisting mostly of columellae and threadlike foot layer^endexines. This wall structure is similar to that of the inaperturate grains of Platygyna (Plate XXXIV) and of two species of Acidoton (Plate XXXIX, 307, 309, 311). 201. Acidoton Swartz, six or seven species in the Greater Antilles, southern Central America and northern South America. (Plate XXXIX ; Plate XL, 315, 316). Two pollen types. 201. Acidoton, (i) Microphyllus pollen type (inaperturate, intectate or semitectate, exines of large columellae and threadlike endexines). Taxa examined and included: A. microphyllus (Plate XXXIX, 309^311); A. urens (Plate XXXIX, 306^308). Grains spheroidal or ellipsoid; inaperturate (Plate XXXIX, 306); LUS = (29.9^36.4)U(27.3^ 35.1) Wm; the surface/tectum microrugulate (Plate XXXIX, 306, 308, 310). In thin section, the exine ca. 1.5^1.6 Wm thick; the endexine of one or two threads with some granular material (Plate XXXIX, 307, 309, 311); the foot layer absent; the columellae prominent, in A. urens sometimes irregular (Plate XXXIX, 307) and occasionally slightly expanded and partially fused at the distal margin (e.g., Plate XXXIX, 307 arrowheads), or in A. microphyllus the columellae larger and markedly irregular ^ the ectexine appearing as folds (Plate XXXIX, 311); the tectum not delimited/de¢ned. Discussion: The ectexine structure of pollen of A. microphyllus (Plate XXXIX, 309, 311) is perplexing. It seems doubtful that the ‘‘distortion of columellae’’ is the result solely of an oblique angle of

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

section. However, compare the similarity of this oblique section of the whole grain (Plate XXXIX, 309) with the oblique section of a whole grain of Platygyna parvifolia (Plate XXXIV, 267), which also has exines composed primarily of columellae. 201. Acidoton, (ii) Nicaraguensis pollen type (three-colpate, tectate, unremarkable exine structure). Plate XXXIX, 312^314; Plate XL, 315, 316). Taxa examined and included: A. nicaraguensis (Plate XXXIX, 312^314; Plate XL, 315, 316); Acidoton species indet. Pollen oblate spheroidal to prolate spheroidal in equatorial view; PUE = (23.4^36.9)U(26.0^ 35.1) Wm; three-colpate, the colpi 16.9^29.1 Wm long; costae absent; the tectum almost complete, punctate to microreticulate to microcrotonoid (Plate XXXIX, 313; Plate XL, 315). In thin section, the mesocolpial exine ca. 1.8 Wm thick; the apertural endexine thin; the non-apertural endexine thick, uniform ; the foot layer thin and irregular (Plate XXXIX, 314) or seemingly absent Plate XXXIX, 312); the columellae irregular but not prominent (Plate XXXIX, 312), sometimes thin (Plate XXXIX, 314); the tectum slightly discontinuous, irregular in thickness. Discussion : Pollen of the Nicaraguensis type is three-colpate, unequivocally tectate, and has an unremarkable exine structure, characteristics which distinguish it from pollen of the Microphyllus type (Plate XXXIX, 306^311). 202. Pachystylidium Pax et K. Ho¡mann, one species in tropical Asia except Sumatra and Borneo (Plate XL, 317^320). P. hirsutum (Plate XL, 317^320). Pollen spheroidal in equatorial view; PUE = (25.2^31.0)U(27.2^31.0) Wm; three-aperturate, appearing porate, the pores poorly de¢ned (Plate XL, 320), the aperture membrane covered with £ecks of ectexine (Plate XL, 317, 319, 320); costae absent; margo absent; operculum absent; the tectum complete, punctate-microspinulose. In thin section, the mesoporal exine ca. 0.8^1.0 Wm thick; the apertural endexine thickened, granular distally, partially solid proximally (Plate XL, 317); the non-apertural endexine thin but consistent

313

(Plate XL, 318); the foot layer mostly absent, consisting of sparse granules; the columellae short, sometimes broad or wedge-shaped, sparse (Plate XL, 318); the tectum continuous, thin. Discussion: In exine structure, pollen of Pachystylidium is very similar to that of Cnesmone and Megistostigma (Plate XXXII ; Plate XXXIII, 251). The apertures of pollen of Pachystylidium are better de¢ned than those in Cnesmone and Megistostigma, but the exine structure of the apertures of all three genera is very similar (Plate XXXII, 249 arrows; Plate XXXIII, 251; Plate XL, 317). Pax and Ho¡mann (1919) established Pachystylidium on Tragia hirsuta Blume, and the pollen data support this treatment since no other examined species of Tragia has similar pollen. Discussion of Tragiinae: pollen of this subtribe (Part 4, Plates XXXII^XL) has the greatest diversity of apertures, exine structure, and tectum morphology of all subtribes of the Acalyphoideae. Pollen of the seven examined genera, Cnesmone, Megistostigma, Tragiella, Platygyna, Tragia, Acidoton and Pachystylidium, can be three-colpate (e.g., Plate XXXVI), inaperturate (e.g., Plate XXXIV), or have poorly de¢ned thin areas (e.g., Plate XL, 319, 320); exines can be intectate and the surface columellate (e.g., Plate XXXVIII, 283^286) or microrugulate (e.g., Plate XXXIX, 306, 308, 310); or tectate and punctate (e.g., Plate XXXIV, 259, 261), punctate-microspinulose (e.g., Plate XXXII, 245, 247), or ¢nely to microreticulate (e.g., Plate XXXIII, 252^258; Plate XXXVI, 277^282). In thin section, some exines have small sparse columellae (e.g., Plate XXXII, 246, 248, 249; Plate XL, 318, 319), while others are composed wholly of massive columellae (e.g., Plate XXXIV, 262, 264, 266; Plate XXXIX, 307); the foot layer ranges from the most prominent of the subfamily (e.g., Plate XXXVI, 285, 286; Plate XXXVIII, 298, 304), to thin, threadlike (e.g., Plate XXXII, 249; Plate XL, 318) or absent (e.g., Plate XXXIV, 260, 262, 264; Plate XXXIX, 307, 309, 311); non-apertural endexines can consist of one or two threads and occasional granules (e.g., Plate XXXIV, 260, 262, 264; Plate XXXIX, 307, 309, 311) or thin granular layers; the apertural endexines are generally thin. If the Tragiinae were reorganized according to

PALBO 2467 11-10-02

314

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XXXIX.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

the distribution of the pollen types, the classi¢cation would be radically changed. Two Asian genera, Cnesmone (Plate XXXII, 245^248) and Megistostigma (Plate XXXII, 246^248; Plate XXXIII, 251, 251 inset) have the Malaccense pollen type (poorly de¢ned apertures, thin tecta, sparse short columellae, and no foot layer) could be combined, and with Pachystylidium (Plate XL, 317^320) be segregated as a subtribe of two genera. The Ramosa pollen type found in T. ramosa (Plate XXXVI, 283^286), T. stylaris (Plate XXXVII, 296), T. urticaefolia (Plate XXXVIII, 297^299) and T. volubilis (Plate XXXVIII, 302, 305) is unequivocally intectate, a rare state in the dicotyledons of the euphorbiaceous hierarchy. Gillespie (1994) found this pollen in ¢ve additional species, all nine of which are assigned to sect. Tragia. This group would constitute a smaller, more coherent Tragia. Pollen of Tragiella (Plate XXXIII, 252^258) is very similar to the Lukafuensis pollen type (three-colpate with protruding

315

tectal margins at the equator, and reticulate exines) found in Tragia gardneri (Plate XXXV, 274^ 276), T. lukafuensis (Plate XXXVI, 277^280), T. mitis (Plate XXXVI, 281, 282), T. pungens (Plate XXXVII, 287^290) and T. spathulata (Plate XXXVII, 291, 292). These species would comprise another genus. The three examined species of Platygyna (Plate XXXIV), Acidoton microphyllus (Plate XXXIX, 309^311) and A. urens (Plate XXXIX, 306^308) share highly distinctive pollen and a West Indian distribution, and could be merged into one genus, or at the very least, the genus Acidoton should be restricted to these two species. The exine structure of these ¢ve species is too distinctive and too similar to be due to chance and chance alone. The Central (and South?) American species of Acidoton with three-colpate grains and an unremarkable exine structure would revert to Gitara Pax et K. Ho¡mann. The disposition of another taxon with inaperturate pollen and exines of mostly columellae, the South Amer-

Plate XXXIX. SEM and TEM of pollen of tribe Plukenetieae, subtribe Tragiinae, Acidoton. 306^308. 306. 307.

308. 309^311. 309.

310.

311.

312^314. 312. 313. 314.

A. urens. Microphyllus pollen type (see legend of A. microphyllus, ¢gs. 309^311). Inaperturate whole grain. There are some irregular, short cracks but no apertures. SEM. Radial section. The ectexine appears to be composed wholly of large columellae some of which are partially fused at the distal margin (arrowheads); there are occasional, small surface areas without support (arrow) ^ this could be the loss of the lower part of a not quite perpendicular columella. TEM. High magni¢cation of surface. Note that surface topography agrees the one in 307. SEM. A. microphyllus. Microphyllus pollen type (inaperturate, exine mostly of columellae). Section of whole grain. An oblique angle of section and partial collapse might account for the appearance of a tectum and the apparent absence of columellae (see legend of ¢g. 311). In this whole grain section as well as others (Nowicke, unpublished data), there is no evidence of apertures. TEM. High magni¢cation of surface. Although this SEM is at a higher magni¢cation than the one of A. urens in ¢g. 308, the surface structures of A. microphyllus are coarser/larger ^ compare the TEMs in ¢gs. 307 and 311, which are at similar magni¢cations. The size and shape of the surface elements agree with the structures seen in thin section, ¢g. 311. SEM. Sections of two grains. Even an oblique angle of section would not wholly account for the continuity of the surface (arrowheads) in some areas. The ectexine appears to consist of irregularly arched or folded elements. Here, and in ¢gs. 307 and 309, the endexine consists of a readily de¢ned thread and irregular granules. Note similarity of variation of pollen morphology and exine structure between the above species of Acidoton and the species of Platygyna illustrated in Plate XXXIV ^ each genus has species with intectate grains (A. urens and P. parvifolia) as well as semitectate (A. microphyllus and P. hexandra). TEM. A. nicaraguensis. Nicaraguensis pollen type (three-colpate, tectate, unremarkable exine structure). Slightly oblique section across mesocolpium. Note prominent and homogeneous non-apertural endexine, the very thin or absent foot layer, and the more uniform, smaller (than in A. microphyllus and A. urens) columellae. TEM. Slightly oblique equatorial view. Note that pollen of this species is aperturate. SEM. Oblique section across aperture and adjacent mesocolpium. The angle of section has enhanced an irregular, discontinuous foot layer (arrows); the paucity of apertural endexine suggests that section was not close to the equator. TEM.

PALBO 2467 11-10-02

316

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

ican Tragia sellowiana (Plate XXXVII, 293^295, 295 inset), is more debatable because of a di¡erent surface morphology. Admittedly, this scheme does not account for some individual species of

Tragia with pollen that is distinct from each other and from any of the above groups. Subtribe 31c. Dalechampiinae (Plates XLI^XLIII)

Plate XL. SEM and TEM of pollen of tribe Plukenetieae, subtribe Tragiinae, Acidoton (315, 316) and Pachystylidium (317^320). 315, 316. 315. 316. 317^320. 317. 318.

319. 320.

A. nicaraguensis. High magni¢cation of tectum along colpus. SEM. Slightly oblique polar view of three-colpate grain. SEM. P. hirsutum. Section across aperture. Note irregular £ecks of ectexine that cover a heterogeneous endexine. TEM. Radial section of mesoporium. The foot layer consists of occasional granules on a thin endexine, and the short, very sparse columellae appear to function mostly as a connection between the endexine^foot layer and the tectum. TEM. Slightly oblique equatorial section of whole grain. TEM. Oblique polar view of whole grain. Note poorly de¢ned apertures. SEM.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XLI (for description see p. 319).

PALBO 2467 11-10-02

317

318

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XLII (for description see p. 319).

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

203. Dalechampia Plumier ex L., about 100 species, 90 of which are in the neotropics, the rest in Africa, Madagascar, India and 1 in China (Plates XLI^XLIII). Species examined: D. brevipes (Plate XLI, 321); D. capensis (Plate XLI, 322^325); D. cissifolia (Plate XLI, 326); D. clematidifolia (Plate XLI, 327^330); D. dioscoreifolia (Plate XLII, 331) ; D. heteromorpha (Plate XLII, 332^336); D. humilis; D. scandens (Plate XLII, 337^339); D. stenosepala (Plate XLIII, 340, 341); D. subternata (Plate XLIII, 342^345); D. tiliifolia (Plate XLIII, 346^348).

319

Pollen prolate spheroidal to prolate in equatorial view; PUE = (42.7^104.8)U(36.9^80.0) Wm; three-colporate, rarely four- to ¢ve-aperturate (D. capensis), the colpi 14.3^48.5 Wm long, narrow, costa ectocolpi absent, margo absent, operculum absent; costa endocingula of two prominently thickened equatorial bands (Plate XLI, 327, 328; Plate XLIII, 342), the polar axis of the endoaperture seemingly limited to the distance between the costae ; the tectum incomplete, mostly reticulate (e.g., Plate XLI 321; Plate XLII, 331, 333, 337, 339; Plate XLIII, 341, 342, 346, 348),

Plate XLI. SEM, TEM and LM of pollen of tribe Plukenetieae, subtribe Dalechampiinae, Dalechampia (see p. 317). 321. 322^325. 322. 323. 324. 325. 326.

327^330. 327. 328. 329. 330.

D. brevipes. Colpus and adjacent tectum. SEM. D. capensis. Radial section. Note thin, elongate columellae and well-de¢ned foot layer. TEM. Tectum. This was the only continental African species examined and the only species with this tectum and with three to ¢ve short, narrow ectoapertures. SEM. Tangential section. Note irregularity of outer surface of foot layer, and that, while variable in size, the columellae are circular or ovae in cross section. TEM. Equatorial view of whole grain. Note distinctive tectum. SEM. D. cissifolia. Section including bands or costae. The shape of the two costae suggests that the orientation of this radial section is probably parallel with the long axis of the grain, yet the disrupted ectexine (between the costae) indicates that a colpus was cut at some angle to its long axis, suggesting a more lateral orientation of the section. TEM. D. clematidifolia. Equatorial view of whole grain. Mid-level focus. Light micrographs best illustrate the unique characteristic of pollen of Dalechampia ^ two thick exinous bands or costae around the equator of the grain. LM. Equatorial view of whole grain. Upper-level focus. LM. Radial section with (probably) longitudinal orientation. The equatorial bands or costae have an electron density similar to the ectexine but have a granular surface more similar to an endexine. TEM. Mesocolpium-centered equatorial view of whole grain. Pollen of this Madagascan species can be distinguished by an almost complete tectum. SEM.

Plate XLII. LM, SEM and TEM of pollen of tribe Plukenetieae, subtribe Dalechampiinae, Dalechampia (see p. 318). 331. 332^336. 332. 333. 334.

335. 336. 337^339. 337. 338. 339.

D. dioscoreifolia. Mesocolpium-centered equatorial view of whole grain. SEM. D. heteromorpha. Tangential section. Compare this tangential section with that of D. capensis, Plate XLI, 324. TEM. Colpus and adjacent tectum. SEM. Oblique orientation of a radial section. The section has passed obliquely through an area of the mesocolpium; the smaller lumina and shorter columellae at bottom right suggest proximity to a colpus. Note irregular foot layer. TEM. Aperture-centered equatorial view of whole grain. SEM. Oblique section with longitudinal orientation through mesocolpium. TEM. D. scandens. Aperture-centered equatorial view of whole grain. Note much larger lumina and ‘higher’ muri (the result of elongate columellae) at poles and mesocolpia. SEM. Oblique section. TEM. Aperture-centered equatorial view of whole grain. At this upper level of focus, the coarse reticulation masks the separation of the two equatorial bands. LM.

PALBO 2467 11-10-02

320

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Plate XLIII.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

frequently the lumina larger at the poles and at the centers of mesocolpia, but smaller nearer the colpi (e.g., Plate XLII, 331, 335, 337; Plate XLIII, 344, 348), the muri simplicolumellate (e.g., Plate XLIII, 341), in one species (D. tiliifolia) some columellae appearing to coalesce (Plate XLIII, 346), or in one species (D. capensis) the tectum ¢nely reticulate and the lumina uniform in size over the entire surface of the grain (e.g., Plate XLI, 323, 325), or in one species (D. clematidifolia, Plate XLI, 330) the tectum almost complete and punctate-perforate. In thin section, the mesocolpial exine ca. 3.0^7.5 Wm thick; an apertural endexine not de¢nable; the non-apertural endexine seemingly absent; at the equator, two exinous bands (costa endocingula) having an electron-density similar to the ectexine, the bands sometimes extending shel£ike into the interior of the grain (e.g., Plate XLI, 326, 329; Plate XLII, 336) ; the foot layer mostly thin (e.g., Plate XLI, 322, 326; Plate XLII, 338; Plate XLIII, 345) and sometimes irregular (e.g., Plate XLII, 334); the columellae variable, elongate (e.g., Plate XLI, 322; Plate XLII, 334) to less so (e.g., Plate XLI, 326, 329),

321

in some species (D. dioscoreifolia, D. subternata, and D. tiliifolia), the columellae conspicuously elongated at the center of the mesocolpium (e.g., Plate XLIII, 343, 345, 347) and to a lesser extent elongated along the margins of the colpus (Plate XLIII, 343 arrowheads, 345 arrowheads, 347 arrowheads and arrows); the tectum thin and discontinuous (e.g., Plate XLI, 326; Plate XLII, 334, 338; Plate XLIII, 345) to rarely almost continuous (Plate XLI, 329). Discussion: Pollen of D. capensis, the only species examined from mainland Africa, has a distinctive tectum (Plate XLI, 323, 325) and at least some four- to ¢ve-aperturate grains. This sample consisted mostly of aberrant grains ^ markedly smaller and thicker walled. Pollen of Dalechampia can be easily distinguished from all remaining members of Acalyphoideae by the costa endocingula, equatorial bands, which are conspicuous in LM (Plate XLI, 327, 328; Plate XLIII, 342). The origin of these bands is debatable. In at least some species, e.g., D. cissifolia, D. clematidifolia and D. heteromorpha, the distance between the bands is wider

Plate XLIII. LM, SEM and TEM of pollen of tribe Plukenetieae, subtribe Dalechampiinae, Dalechampia. 340, 341. 340. 341. 342^345. 342. 343. 344. 345.

346^348. 346. 347.

348.

D. stenosepala. Slightly oblique equatorial view of whole grain. SEM. High magni¢cation of reticulate ectexine. SEM. D. subternata. Aperture-centered mesocolpial view of whole grain. Upper-level focus. The two bands are visible but only the colpus is in focus. LM. Polar view of whole grain. Note that columellae are prominently elongated in the center of mesocolpium, and to a lesser extent, to each side of the colpus (arrowheads). LM. Oblique equatorial view of whole grain. Arrowhead points to area of ridge in ¢g. 343. SEM. Oblique radial section of an equatorial orientation. The section passed through the middle of a mesocolpium (top), and then angled downward to pass obliquely through one of the bands; the arrowheads point to areas equivalent to those in ¢g. 343. TEM. D. tiliifolia. High magni¢cation of ectexine. Note that some muri have areas where columellae may have expanded laterally and fused (short arrows). SEM. Polar view of whole grain. The grain is tilted slightly. This species has a series of ridges formed by elongate columellae alternating with short ones: the lowest ridge is nearest the colpus (arrowhead), then a slight depression or level area, another more prominent ridge (arrows), another trough, and then the most elongate columellae at the center of the mesocolpium. LM. Mesocolpium-centered equatorial view of whole grain. The area to each side of the colpus is sharply delimited by much smaller lumina and muri with much shorter columellae; compare with the more uniform reticulation of D. stenosepala (¢g. 340) and the punctate tecta of D. clematidifolia (Plate XLI, 330), both of which are simply rounded triangular in polar view, without ridges. SEM.

PALBO 2467 11-10-02

322

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

at the colpi, suggesting that the ancestral group had lalongate endoapertures, which, over time, elongated laterally until they merged. In LM, acetolysed grains mounted in glycerin jelly have a thin inner, darker-staining layer and a thicker outer, lighter-staining layer. The inner layer between the costae does not appear to be thinner than anywhere else in the grain. Thus, in the strictest sense, the bands are not costa endocingula because they do not border an endoaperture.

Grains of Dalechampia are the largest of the subfamily, polar dimensions range from 43 to 105 Wm, with six of the nine species examined having average polar dimensions s 55 Wm. They have the thickest exines, 3^7.5 Wm. Species of Dalechampia are the only taxa of Acalyphoideae to have coarsely reticulate exines (e.g., Plate XLI, 321; Plate XLII, 337; Plate XLIII, 346, 348), although Panda oleosa Pierre has similar exines (Nowicke et al., 1998, Plate X, 103). Pollen of some species of Chrozophora (Nowicke et al.,

Plate XLIV. SEM and TEM of pollen of tribe Omphaleae, Omphalea. 349. 350, 351. 350.

351. 352. 353. 354.

O. bracteata. Oblique partial view showing colpus. SEM. O. diandra. Radial section of mesocolpium. Note separation of non-apertural endexine (a characteristic shared with those members of Crotonoideae with aperturate pollen; see text), the irregular foot layer and very irregular columellae. TEM. Partial polar view showing aperture with prominently granular membrane. SEM. O. oleifera. Polar view of whole grain. SEM. O. oppositifolia. High magni¢cation of tectum at the pole. SEM. O. palmata. Polar view of whole grain. Note prominently granular aperture membranes. SEM.

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

1999, Plate XII, 95^99; Plates XIII, XIV) can also be large (36.9^66.3U34.9^68.9 Wm), thick-walled (up to 3.9 Wm), have endocingula (two species)

323

and a similar range of tectal variation, punctate to reticulate-heterobrochate. However, neither Panda nor Chrozophora are closely related to Da-

Plate XLV. LM, SEM and TEM of pollen of tribe Omphaleae, Omphalea. 355^357. 355. 356. 357. 358^360. 358. 359. 360.

O. triandra. Radial section. Note irregular foot layer, sparse columellae, thick continuous tectum and separation of endexine from ectexine; see legend of Plate XLIV, 350. TEM. Equatorial view of whole grain showing separation (top and bottom) of non-apertural endexine. LM. Oblique section of whole grain. TEM. O. trichotoma. Section across aperture. Note dearth of apertural endexine here and in ¢g. 357. TEM. Polar view of whole grain. SEM. Radial section across mesocolpium. The columellae are irregular and unevenly distributed as well; note also the irregularity of the inner surface of the tectum and the separation of endexine from ectexine.

PALBO 2467 11-10-02

324

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

lechampia on other bases. Certainly the restriction of such prominent endocingula and coarsely reticulate exines to pollen of Dalechampia suggest no relationships to other genera of tribe Plukenetieae nor to any remaining members of Acalyphoideae. According to Armbruster (1994), the unusual in£orescences of Dalechampia are probably homologous to those of Plukenetia and Tragia, and all three genera have similar viney habit. Although each of the latter two genera have more than one pollen type (Plates XXVIII, 213; XXXI, 242 versus Plates XXXV^XXXVIII respectively), none is remotely similar to pollen of Dalechampia. The pollen data indicate that Dalechampia is an isolated genus and that it should be treated as a monogeneric tribe. Tribe 32. Omphaleae (Plates XLIV, XLV) 204. Omphalea L., 15 species mostly in Greater Antilles and Madagascar (Plates XLIV, XLV). Species examined: O. bracteata (Plate XLIV, 349); O. diandra (Plate XLIV, 350, 351); O. oleifera (Plate XLIV, 352); O. oppositifolia (Plate XLIV, 353) ; O. palmata (Plate XLIV, 354); O. triandra (Plate XLV, 355^357); O. trichotoma (Plate XLV, 358^360). Pollen oblate spheroidal to more rarely prolate spheroidal in equatorial view; PUE = (15.5^ 29.9)U(19.4^35.1) Wm; three-colpate, the colpi 9.7^18.2 Wm long, the aperture membrane granular; costae absent, margo absent, operculum absent; the tectum complete, ¢nely punctate and microspinulose (Plate XLIV, 349, 351, 353). In thin section, the mesocolpial exine ca. 1.7^2.3 Wm thick; the apertural endexine poorly developed; the non-apertural endexine well developed, uniform in thickness, frequently separating from the ectexine (e.g., Plate XLIV, 350; Plate XLV, 355^358); the foot layer irregular (e.g., Plate XLIV, 350; Plate XLV, 355, 360), from a third (e.g., Plate XLV, 355) to two thirds (e.g., Plate XLIV, 350; Plate XLV, 360) as thick as the tectum ; the columellae irregular (e.g., Plate XLV, 355, 360) in diameter, in length and in shape, sometimes markedly so (Plate XLIV, 350) ; the tectum continuous, thick (e.g., Plate XLIV, 350; Plate XLV, 357, 358), the predominant layer.

Discussion: The pollen of Omphalea is of a generalized sculpture, punctate-microspinulose, and common aperture type, three-colpate. The most notable feature of the exine structure is the irregularity of both columellae (Plate XLIV, 350; Plate XLV, 355, 360) and foot layer. The non-apertural endexine separates from the ectexine (Plate XLIV, 350; Plate XLV, 355^358, 360), a phenomenon that also occurs in the grains of colpate pollen of Crotonoideae (Nowicke, 1994 : ¢gs. 11, 14, 20, 23). Of greater interest are the data from Rbcl studies (K. Wurdack, personal communication) that Omphalea forms a clade with Tetrorchidium Grisebach and Endospermum Bentham, two of the aperturate (three-colpate) genera of that subfamily.

4. Discussion of Acalyphoideae In the following lengthy discussion, when referring to illustrations, we cite only the Part number and roman numerals of the relevant plate(s) to save space in the text. To ¢nd the citation in References, Part 1 is Nowicke et al. (1998), Part 2 is Nowicke et al. (1999), Part 3 is Takahashi et al. (2000), Part 4 is this paper. The most common pollen of Acalyphoideae is small in size, three-colporate, has a lalongate endoaperture, frequently with costae, and a complete tectum with unremarkable sculpture. Most exines have thin non-apertural endexines, thin foot layers, small columellae and thick tecta. These grains, or ones with only slight deviations, are characteristic of many of the larger genera, e.g., Macaranga, Mallotus, Cleidion, and smaller ones, e.g., Adriana, Claoxylon, Enriquebeltrania. Small pollen size, low-pro¢le sculpture and reduced £owers are characteristics frequently associated with wind pollination, said to occur in Acalypha, Macaranga and Mallotus (Webster, 1994). However, wind-pollinated plants usually produce a large volume of pollen, and this is certainly not the case in the Acalyphoideae examined here. Pollination of Dalechampia, which has the largest grains of the subfamily, is reported to be by anthidiine and euglossine bees (Armbruster, 1988).

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

4.1. Variation of pollen characteristics of Acalyphoideae 4.1.1. Apertures Most members of Acalyphoideae have threecolporate pollen. More than three apertures occur in Chrozophora, which is six- to nine-colporate, in Argythamnia, which is four-colpate, and sporadically in other genera, e.g., Acalypha, Adriana, Caperonia, Erythrococca, Micrococca. Pollen of Ditaxis has an unequal distribution of the three colpi ^ two are much closer (Part 2, Plate VI). Alternatives to compound apertures include grains with simple apertures, or with poorly de¢ned apertures or without apertures. Colpate pollen characterizes Argythamnia, Ditaxis (Part 2, Plates VI^VIII), Plukenetiinae (Part 4, Plates XXVII^XXXI), Mercurialinae (Part 3, Plates XX, XXI), Dysopsidinae (Part 3, Plate XXII), and Omphaleae (Part 4, Plates XLIV, XLV). Three-colpate grains also occur in most Tragiinae (Part 4, Plates XXXIII, XXXV^XXXVIII, ), almost all Adelieae (Part 2, Plates VII, VIII, X), most species of Chiropetalum (Part 2, Plates IX, X), and a peripheral group of species in various genera in which the endoapertures are too small to measure, e.g., Alchornea (Part 3, Plate XIII). Poorly de¢ned apertures consisting of areas of discontinuous ectexine in the form of irregular islands over thickened endexines, occur in Cnesmone, Megistostigma (Part 4, Plate XXXII), Tragia urens (Part 4, Plate XXXVIII, 300^303), and Pachystylidium (Part 4, Plate XL). Of all Acalyphoideae examined in this study, unequivocally inaperturate pollen occurs only in Platygyna (Part 4, Plate XXXIV), two species of Acidoton (Part 4, Plate XXXIX, 306^311), and Tragia sellowiana (Part 4, Plate XXXVII, 293^296). Both ectoapertures (colpi) and endoapertures can have costae, thickenings of the endexine (Part 1, Plate XIII) that may function to strengthen or provide rigidity to the margins of apertures. The costae could mitigate both premature rupture and inward collapse of this vulnerable area. This interpretation of costae function would not be supported by pollen of Dalechampia in which the ultimate development of costae may occur (Part 4, Plates XLI^XLIII). These grains have

325

narrow colpi and thick exines (3.0^7.5 Wm) and do not appear particularly susceptible to collapse. Ectoapertures vary widely in length, from porelike in Acalypha (Part 4, Plates XXI^XXIII) and Cephalomappa (Part 3, Plate VI), to short, e.g., Blumeodendrinae (Part 2, Plates XXII, XXIII), to more elongate, e.g., many members of subtribe Epiprininae (Part 3, Plates II, IV, V). No syncolpate or syncolporate grains were observed in any material examined. Most colpi are narrow, but exceptions do occur, e.g., in some members of the Loretensis type of Plukenetia (Part 4, Plate XXIX, 228; Plate XXX, 232), some species of Omphalea (Part 4, Plate XLIV), or the poorly de¢ned colpi of Cnesmone (Part 4, Plate XXXII, 245), all of which have granular membranes. Modi¢cations of colpal margins include psilate margos, which occur in most members of Bernardieae (Part 2, Plates XVII^XIX), most species of Monotaxis (Part 1, Plate XI) and of Amperea (Part 1,Plate XII), some species of Clutia (Part 1, Plate I), of Agrostistachys (Part 2, Plate I), and sporadically in other genera. Certain genera, mostly in tribe Plukenetieae, have colpal margins in which the exine layers slightly separate in the region of the equator (Part 4, Plate XXIX, 218; Plate XXX, 227). The resulting fastigium-like cavity is best viewed in intact grains in polar view in LM. The tectum is slightly arched at the equator in pollen of Tragiella and the Lukafuensis pollen type of Tragia (Part 4, Plate XXXIII, 253; Plate XXXVI, 279). Fastigium-like cavities, delimited by protruding colpal margins and intact endexine, are found in some species of Acalypha (Part 4, Plate XXI, 162; Plate XXIII, 178, 180). The distribution of certain types of opercula appears to have taxonomic value. Unstrati¢ed opercula characterize all examined members of the tribe Alchorneae (Part 3, Plates XXII^XXVII), which, in cross section, appear as a solid, electron-dense mass either circular or ovate in shape. Strati¢ed opercula occur in at least four genera, three of which are assigned to Adelieae. Grains of Adelia (Part 3, Plates VII, VIII), Lasiocroton (Part 3, Plate X) and Leucocroton (Part 3, Plates XI, XII) have opercula with tectum and columellae, but appear to lack a foot layer. The largest opercula occur in pollen of Caperonia (Part 2, Plates

PALBO 2467 11-10-02

326

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

IV, V), which was described at one time as stephanocolporate. These opercula have an exine structure very similar to that of the mesocolpium. Pollen of Chiropetalum (Part 2, Plates IX, X) has slender opercula that re£ect the tectal morphology. Limited TEM data suggest that the opercula found in a few species of Galearieae have columellae (Part 1, Plate X). Endoapertures of Acalyphoideae are almost always lalongate, sometimes markedly so, and frequently with poorly de¢ned lateral margins (Part 1, Plate XIII). Endocingula (zonorate) endoapertures occur in some members of Chrozophora (Part 2, Plate XXVI) and probably Dalechampia (Part 4, Plates XLI^XLIII). In pollen of Adenochlaena leucocephala (Part 3, Plate II) the lateral margins of the endoaperture are forked, a modi¢cation also found in occasional grains of Cephalocroton cordofanus, and of Mareya brevipes (Breteler et al., 1997, ¢g. 4). The evolution of simple apertures in Acalyphoideae occurred at least three times. Once was in the ancestor of the closely related genera, Argythamnia and Ditaxis, of the Chrozophoreae. Given their suprareticulate sculptures, prominent columellae and well developed foot layers (Part 2, Plates VI^VIII), as well as the presence of compound apertures in eight of the remaining nine genera of the Chrozophoreae, it seems likely that the three-colpate condition of Argythamnia and Ditaxis is derived. Although Chiropetalum is considered closely related to the above genera, the pollen data, except for the presence of simple apertures, argue against this conclusion. A second origin of colpate apertures occurred in the ancestral line of three genera of Adelieae, Adelia, Lasiocroton and Leucocroton, all of which have pollen with strati¢ed opercula, microcrotonoid tecta, and similar exine structures. The three-colpate pollen of two subtribes, Mercurialinae and Dysopsidinae (Part 3, Plates XX^XXII), while not highly distinctive, is unrelated to any of the above genera, as well as to any remaining members of the Acalypheae. Because grains of all species of Plukenetiinae and most species of Tragiinae are three-colpate, it seems likely that simple apertures are not a derived condition in Plukenetieae.

The origin of inaperturate pollen in Acalyphoideae probably occurred just once. At this level of the dicotyledonous hierarchy, just the rarity of this condition indicates it is derived. Inaperturate pollen was found in six species, all of which belong to Tragiinae, three of Platygyna (Part 4, Plate XXXIV), two of Acidoton (Part 4, Plate XXIX) and Tragia sellowiana (Part 4, Plate XXXVII). The unique exines of these grains, primarily columellae, suggests that they are closely related and that the inaperturate condition evolved just once. In Tragiinae, the distance between the inaperturate condition and the three-colpate condition might have been spanned by pollen with poorly de¢ned apertures (Megistostigma, Cnesmone and Pachystylidium), were it not for great di¡erences of exine structure among these three groups/pollen types. 4.1.2. Tectum/sculpture variation Pollen grains of the vast majority of Acalyphoideae have complete tecta and the most common sculptures are punctate-microspinulose, e.g., Macaranga (Part 3, Plates XXVI, XXVII), Ricinus (Part 3, Plate XVII), many members of Claoxylinae (Part 4, Plates I^X), most species of Mallotus (Part 4, Plates XII^XVI), Octospermum (Part 4, Plate XX), and punctate or deeply punctate, e.g., Bernardieae (Part 2, Plates XVII^XIX), some species of Pera (Part 1, Plate IV), Caryodendreae (Part 2, Plate XV), Caperonia (Part 2, Plate IV), e.g. (Part 2, Plates XVII^XIX), Speranskia (Part 3, Plate I), and some species of Rockinghamia (Part 4, Plate XIX). In Acalypha (Part 4, Plates XXI^XXIV), the microspinules are clustered among minute psilate areas. Microreticulate exines characterize the closely related subtribes Mercurialinae (Part 3, Plates XX, XXI) and Dysopsidinae (Part 3, Plate XXII) of the tribe Acalypheae. Finely reticulate to microreticulate exines with crenate muri are found in Tragiella (Part 4, Plate XXXIII) and the Lukafuensis type of Tragia (e.g., Part 4, Plates XXXV, XXXVI). Coarsely reticulate exines with variably sized lumina (larger at the poles and the centers of mesocolpia) occur in pollen of Panda oleosa (Part 1, Plate X) and in many species of

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Dalechampia (Part 4, Plates XLII, XLIII) ^ but this is the only pollen characteristic that these two taxa share. Tecta with larger perforations/lumina at the poles/mesocolpia occur sporadically in Acalyphoideae, e.g., Pera tomentosa (Part 1, Plate VI), Speranskia (Part 2, Plate III), Doryxylon (Part 2, Plate XI), and some species of Koilodepas (Part 3, Plate III, 21) and of Tragia (Part 4, Plate XXXV, 268). A number of genera have a sculpture that appears similar to the Croton tectum, de¢ned previously (Nowicke, 1994) as a continuous triangular array, and associated with (almost) all members of Crotonoideae. The best published example of a croton-like tectum in Acalyphoideae is probably that of Botryophora (Part 2, Plate XXV). In other genera, e.g., Adelia (Takahashi et al., 2000, pl. VII, 52; VIII, 64), Chondrostylis (Nowicke et al., 1999, pl. III, 24) and Leucocroton (Takahashi et al., 2000, plate XI, 87) the distinction and size of the triangles is reduced, a variation also found in Crotonoideae (Nowicke, 1994, ¢gs. 40, 55 and particularly 56). Occasional species or some grains within a sample show a vague similarity to a crotonoid tectum, e.g., Mareyopsis longifolia (Part 4, Plate VIII, 54) and Micrococca capensis (Part 4, Plate X, 63). But none of the above examples have the Croton exine structure, in which the triangular subunits are attached to a network of muri (Nowicke, 1994, e.g., ¢gs. 6, 8, 14, 21, 37, 41, 47). We do not believe that the existence of a surface morphology described as microcrotonoid necessarily supports or even indicates a close relationship between Crotonoideae and Acalyphoideae. Pollen of Thymelaeaceae has the croton tectum and the croton structure (Nowicke, 1994, ¢gs. 97^102; Nowicke and Skvarla, unpublished data) and Thymelaeaceae are not considered closely related to Euphorbiaceae. Other examples of pollen with triangular arrays that are familiar to the present authors, occur in certain taxa of Berberidaceae, where the surface was described as having a ‘pinwheel con¢guration’ (Nowicke and Skvarla, 1981: ¢gs. 77, 78, 109^114). Triangular arrays should be viewed as a morphological variant, much like spinose, echinate or reticulate tecta. However, we would be remiss if we did not point out that a

327

recent cladistic analyses of molecular data shows ¢ve genera of Crotonoideae with aperturate pollen as the sister group to 23 genera of Acalyphoideae (Wurdack, unpublished data). Exceptions to the above tectal morphologies are usually restricted to one or sometimes two genera. Suprareticulate tecta occur in Argythamnia (Part 2, Plates VII, VIII) and most species of the closely related genus Ditaxis (Part 2, Plate VI). Pollen of Dicoelia (Part 1, Plate IX) is only weakly suprareticulate and the exine structure is not similar to Argythamnia and Ditaxis. The echinate tecta which occur only in Cheilosa and Neoscortechinia (Part 1, Plate VII) re£ects their close relationship. Amyrea (Part 4, Plate XI) is the only genus of Acalyphoideae with a striate sculpture. In the small subtribe Lasiococcinae, the tectum of Spathiostemon (Part 4, Plates XXV, XXVI) has a unique supratectal sculpture of lirae with shallow cross striations, whereas pollen of a second genus, Homonoia (Part 4, Plate XXVI), has a unique tectum of individual, densely spaced elongate elements whose acute tips form a microechinate sculpture. The columellate surface of the intectate Ramosa pollen type of Tragia sect. Tragia (Part 4, Plate XXXVI, 283^286; Plate XXXVIII, 297^ 299) is unknown in the remaining Acalyphoideae. Heterobrochate tecta are found in some species of Chrozophora (Part 3, Plates XIII, XIV). 4.1.3. Exine structure variation The most common exine structure in Acalyphoideae by virtue of its presence in some of the largest genera, e.g., Macaranga and Mallotus, consists of a thin or irregular foot layer, short or small columellae, and a continuous tectum that is usually the predominant component (Part 3, Plate XXVII; Part 4, Plates XIII^XVI). Exines of most of the examined species of Acalypha (Part 4, Plates XXI^XXIV) are similar, except for a few species with more elongate columellae. The foot layer is rarely, if ever, the predominant component, but it reaches maximum thickness in the intectate exines of the Ramosa pollen type of Tragia (Part 4, Plate XXXVI, 285; Plate XXXVIII, 298) ^ perhaps to add rigidity or strengthen the wall in the absence of a tectum. The foot layer is frequently thin, e.g., Cephalo-

PALBO 2467 11-10-02

328

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

mappa (Part 3, Plate VI), Macaranga (Part 3, Plate XXVII), Mallotus (Part 4, Plate XIII), Chiropetalum (Part 2, Plates IX, X), or threadlike, e.g., Adelia (Part 3, Plates VII, VIII), Lasiocroton (Part 3, Plate X), to seemingly absent, e.g., Pycnocoma (Part 2, Plate XX), Claoxylopsis (Part 4, Plate VI). Occasionally the foot layer is unequivocally absent, e.g., Platygyna (Part 4, Plate XXXIV), some species of Acidoton (Part 4, Plate XXXIX). The foot layer can be irregular on the outer surface, e.g., Dicoelia (Part 1, Plate IX, 88), Pseudagrostistachys (Part 2, Plate II, 16), Alchorneopsis (Part 2, Plate XVI, 124), Koilodepas (Part 3, Plate III, 28), Leucocroton virens (Part 3, Plate X, 80), Micrococca (Part 4, Plate X, 69), Amyrea (Part 4, Plate XI, 74, 76, 78) and Rockinghamia angustifolia (Part 4, Plate XIX, 145). Perforations and/or channels are found in the foot layers of most genera of subtribe Mercurialinae (Part 3, Plate XX, 155, 157; Plate XXI, 163, 165) and in the Ramosa pollen type of Tragia. In some taxa, e.g., those species of Tragia with the Ramosa pollen type (Part 4, Plate XXXVI, 286; Plate XXXVIII, 298, 304) and in Plukenetia loretensis (Part 4, Plate XXX, 231, 233, 235), the foot layer appears to be composed of ‘short columellae’. The variation in tectum structure is usually a re£ection of the sculpture, e.g., the variable thickness of the suprareticulate tecta of Argythamnia and Ditaxis (Part 2, Plates VI^VIII). However, in some groups, modi¢cations of the tectum suggest close relationships, e.g., the granular inner surface of the Conophora pollen type of Plukenetia (Part 4, Plate XXVIII, 214, 215; Plate XXIX, 220, 222; Plate XXXI, 241). Many taxa of Acalyphoideae have pollen with short/small columellae, e.g., most species of Pera (Part 1, Plates IV, VI), Erismanthus (Part 1, Plate VIII), Chiropetalum (Part 2, Plates IX, X), Caryodendron (Part 2, Plate XV), Droceloncia (Part 3, Plate XXI), Enriquebeltrania (Part 3, Plate IX), Adriana (Part 3, Plates XVIII, XIX), many species of Cleidion (Part 3, Plates XXIII^XXV), Macaranga (Part 3, Plate XXVII), Mallotus (Part 4, Plate XIV), Trewia (Part 4, Plate XVIII), Octospermum (Part 4, Plate XX), and most species of Acalypha (Part 4, Plates XXII, XXIII). In pollen

of Spathiostemon (Part 4, Plate XXV), Homonoia (Part 4, Plate XXVI), of Cnesmone, of Megistostigma (Part 4, Plate XXXII) and of Pachystylidium (Part 4, Plate XL), the columellae are small and sparsely distributed. In Pycnocoma (Part 2, Plate XX), the columellae are seemingly reduced to proximal points on the tectal elements. At the other extreme are the exines of Platygyna (Part 4, Plate XXXIV) and two species of Acidoton (Part 4, Plate XXXIX) which consist almost wholly of large, even massive columellae. Large columellae are the exception, but do occur, e.g., in Pera distichophylla (Part 1, Plate V), Amperea (Part 1, Plate XII), Ditaxis (Part 2, Plate VI), Argythamnia (Part 2, Plates VII, VIII), Haematostemon (Part 4, Plate XXVII), Romanoa (Part 4, Plate XXVIII), the Conophora type of Plukenetia (Part 4, Plate XXXIX, 218^222) and the Ramosa type of Tragia (Part 4, Plate XXXVI, 283^286). Some members of Dalechampia (e.g., Part 4, Plate XLIII) have elongated columellae in the middle of the mesocolpia, while some species of Erythrococca have conspicuously elongated columellae at the poles (Part 4, Plates I, II). Tangential sections of pollen of Acalyphoideae reveal that many columellae are irregularly shaped (not circular or ovate) in cross section, e.g., Pera distichophylla (Part 1, Plate V), Chiropetalum (Part 2, Plates IX, X), Koilodepas (Part 3, Plate III), Macaranga (Part 3, Plates XXVI, XXVII), Erythrococca (Part 4, Plate II), Claoxylon (Part 4, Plate V, 34), Amyrea (Part 4, Plate XI, 76), Platygyna (Part 4, Plate XXXIV). Modi¢cations of columellae include branching which occurs in Clutia (Part 1, Plates I, II) and some members of Plukenetia (Part 4, Plates XXIX, XXX), and fusion of two or more, e.g., Pogonophora (Part 1, Plate II), Lasiocroton (Part 3, Plate X, 75), Claoxylon (Part 4, Plate II) and Platygyna (Part 4, Plate XXXIV). Elongate columellae near apertures are found sporadically throughout the subfamily, e.g., Bernardia (Part 3, Plate XVII, 135, 137), Botryophora (Part 2, Plate XXV, 193), but are most consistently present in the tribe Alchorneae (Part 3, Plates XII^XVII). Columellar modi¢cations that appear to have taxonomic value above the level of genera occur in the subtribe Alchorneinae, in which all taxa

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

examined in TEM have a modi¢ed infratectum ^ either the columellae are expanded at mid-length, e.g., Alchornea rugosa (Part 3, Plate XIV, 111 arrowheads) or, the infratectum appears to consist of two poorly distinguished layers of columellae, e.g., Or¢lea (Part 3, Plate XII, 96). The majority of taxa of Acalyphoideae have endexines thickened beneath the colpi (e.g., Part 1, Plates II, XII; Part II, Plate XVII ; Part 3, Plates III, XXIV), but not prominently so. The apertural endexines of pollen of Melanolepis (Part 2, Plate XII, 91) are an exception ^ thick and conspicuously granular on the inner surface. The greatly thickened non-apertural endexine of Pycnocoma (Part 2, Plate XX) constitutes the only continuous layer in a highly modi¢ed exine structure. Pollen of Dalechampia (Part 4, Plates XLI^ XLIII) has a unique modi¢cation in which the endocingulum is de¢ned by two prominent costae around the equator of the grain. In pollen with thin or absent foot layers, the non-apertural endexine is, if not thickened, at least consistent, e.g., Epiprinus (Part 3, Plate I), Adelia (Part 3, Plates VII, VIII) and Claoxylopsis (Part 4, Plate VI). There are taxa in which the endexine is poorly developed throughout the grain, e.g., Ditaxis (Part 2, Plate VI), Chiropetalum (Part 2, Plates IX, X), some species of Mercurialis (Part 3, Plate XX) and Dysopsis (Part 3, Plate XXII). The Conophora pollen type, which includes Romanoa (Part 4, Plate XXVIII), Eleutherostigma (Part 4, Plate XXIX), and some species of Plukenetia (Part 4, Plates XXX, XXXI) has a very thin apertural endexine which separates from the ectexine to form fastigium-like cavities at the equator (Part 4, Plates XXVIII^XXX). The inaperturate grains of Platygyna (Part 4, Plate XXXIV), of Acidoton (Part 4, Plate XXXIX, 306^311) and of Tragia sellowiana (Part 4, Plate XXXVII) share exines consisting mostly of columellae and threadlike endexines. The absence of two continuous ‘‘horizontal’’ strata ^ the foot layer and tectum ^ and the presence of weakly linked vertical elements ^ the large columellae ^ may be adaptations for an easily ruptured wall for germination. Unacetolysed pollen should be embedded and sectioned to assess the role of the intine in inaperturate grains.

329

5. Summary and commentary of the tribes Tribe 13. Clutieae (Part 1, Plates I, II) In this monogeneric tribe, the exines have branched columellae and angular reticulate tecta (laterally expanded muri partially cover the lumina), the latter characteristic unknown in the remaining Acalyphoideae, except for some species of Amperea. The pollen data support the present concept of Clutieae. Tribe 14. Pogonophoreae (Part 1, Plate II) Pollen of Pogonophora is of a generalized type, but it is not similar to that of Clutia (Webster, 1994). Evidence of relationships of Pogonophora and the validity of this monogeneric tribe should depend on non-palynological characteristics. Tribe 15. Chaetocarpeae (Part 1, Plates II, III) The pollen of both genera, Chaetocarpus and Trigonopleura, is of a generalized type, although the larger columellae of the latter genus could bridge the modest di¡erence between the pollen of Chaetocarpus and that of Pogonophoreae (Part 1, Plate II). Tribe 16. Pereae (Part 1, Plates IV^VI) Three pollen types, two of which are restricted to a single species, were recognized in Pera. The highly distinctive pollen of P. distichophylla supports its treatment as a monotypic section by Pax and Ho¡mann (1931). However, the unique in£orescence of £owers enclosed in bibracteolate involucres suggests that Pera is monophyletic. Tribe 17. Cheiloseae (Part 1, Plate VII) There is no question of the close relationship between Cheilosa and Neoscortechinia nor of their distinction from the remaining Acalyphoideae. Pollen of this tribe has identical exine structures and share a unique tectum, short echinae on a microrugulate surface. The distinction of the pollen is such that no relationships are suggested. Tribe 18. Erismantheae (Part 1, Plate VIII) The two examined genera, Erismanthus and Moultonianthus, of the three assigned to Erismantheae, have di¡erent pollen. It is punctate/micro-

PALBO 2467 11-10-02

330

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

reticulate with almost continuous tecta and short, irregular columellae in the former genus versus ¢nely reticulate tecta with triangular shaped muri and uniform columellae in the latter. Whether these di¡erences are su⁄cient to merit separate tribal status for Erismanthus and for Moultonianthus is questionable. It is possible that pollen of the unexamined third genus, Syndiophyllum Lauterbach and K. Schumann, could bridge the gap. Tribe 19. Dicoelieae (Part 1, Plate IX) Pollen of Dicoelia has irregular columellae and an irregularly thick, weakly suprareticulate tectum. Although the pollen is not highly distinctive, the pollen data do not support a close relationship with Galearieae, much less a merger with that tribe (see below). Tribe 20. Galearieae (Part 1, Plates IX, X) In Galearieae, we originally considered pollen of Panda, which has a coarsely reticulate exine, to be di¡erent from that of Galearia and Microdesmis, which has a deeply punctate tectum and occasional opercula. Except for columellae length, the exine structures of all three genera are similar, including muri that are triangular in cross section. Moreover, along the colpi, the lumina of pollen of Panda are much smaller and not unlike the tecta of Galearia and Microdesmis. Although Webster (1994) considers Galearieae and Dicoelieae to be closely related, even monophyletic, the pollen data would not support this view. Tribe 21. Ampereae (Part 1, Plates XI, XII) The Ampereae comprise two genera, Amperea and Monotaxis, the pollen of which shares incomplete tecta and apertures with conspicuous psilate margos. There are di¡erences in exine structure. The columellae of Amperea may be the most elongate of the subfamily and the tectum is angular reticulate (laterally expanded muri partially cover the lumina), while the columellae of Monotaxis are, at most, only average and the tectum unmodi¢ed. The pollen di¡erences between the two genera are more substantial than the similarities. Grains with angular reticulate tecta occur in species of Clutia (Part 1, Plate I).

Tribe 22. Agrostistachydeae (Part 2, Plates I^III) Of the four genera assigned to Agrostistachydeae, pollen of Agrostistachys and Pseudagrostistachys is of a generalized type: three-colporate, frequently with margos, punctate or ¢nely reticulate tecta and an unremarkable exine structure. Grains of Chondrostylis have a prominent bridge of ectexine over the endoaperture, a microcrotonoid tectum, thin uniform columellae, and a uniformly thick tectum. Pollen of Cyttaranthus (Part 2, Plate III, 20^23) is three-colporate, weakly suprareticulate, has densely spaced columellae and an irregularly thickened tectum. The distinction of the pollen of Cyttaranthus could be acknowledged by treating it as a monogeneric subtribe. Tribe 23. Chrozophoreae (Part 2, Plates III^XIV) The large tribe Chrozophoreae has disparate pollen elements that are partially addressed by the segregation of the 11 genera into four subtribes, two of which are monogeneric, Chrozophorinae and Speranskiinae. Pollen of Chrozophora (Part 2, Plates XII^XIV) is six- to ninecolporate with mostly heterobrochate tecta, although two closely related species have punctate tecta and endocingula (zonorate endoapertures). The treatment of Chrozophora as a monogeneric subtribe is reinforced by the pollen data. Pollen of Speranskia (Part 2, Plate III) is of a generalized type and the validity of this monogeneric subtribe should be based on evidence from £oral morphology/molecular studies. Pollen data argue against a close relationship among the ¢ve genera of the subtribe Ditaxinae (Part 2, Plates IV^X). Pollen of Ditaxis and Argythamnia is similar and distinctive: suprareticulate tecta, three- to four-colpate, thick-walled with prominent columellae and complete, irregularly thickened tecta. Pollen of a third, supposedly closely related genus, Chiropetalum (Part 2, Plates IX, X), has ¢nely reticulate tecta, is three-colpate with slender opercula, and thin-walled with short columellae. Admittedly, the occurrence of simple apertures in Ditaxis, Argythamnia and Chiropetalum is perplexing, but the magnitude of the di¡erences in exine structure and tectum morphology is much greater. Of the two remaining genera in Ditaxinae (Part 2, Plates

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

IV, V), pollen of Caperonia is colporate with large strati¢ed opercula while that of the monotypic Philyra is three-colporate ; the latter has reticulate tecta and an unremarkable exine structure very similar to some species of Caperonia. Three of the four genera assigned to the fourth subtribe, Doryxylinae (Part 2, Plates XI, XII), Doryxylon, Sumbaviopsis and Melanolepis, have similar pollen: ¢nely reticulate or punctate tecta, and thin exines with irregular foot layers, short columellae and irregularly thick tecta, whereas pollen of the fourth genus, Thyrsanthera, has thicker exines with a channeled foot layer, more elongate columellae and a uniformly thick tectum. In the Chrozophoreae, the pollen data suggest these changes : Ditaxinae be restricted to Ditaxis and Argythamnia ; Chiropetalum be treated as a monogeneric subtribe ; Caperonia and Philyra could be accommodated in the same subtribe ; Doryxylinae should be maintained as it is, unless Thyrsanthera is additionally distinguished by some aspect of £oral or fruit morphology. Tribe 24. Caryodendroneae (Part 2, Plates XV, XVI) Pollen of the three genera of tribe Caryodendreae, Caryodendron, Discoglypremna and Alchorneopsis, is similar and of an unremarkable type that does little to elucidate their relationships to each other or to any other genera of Acalyphoideae.

331

gated into two subtribes. The genera of Blumeodendrinae, Blumeodendron, Podadenia, Ptychopyxis and Botryophora, share pollen with short colpi, almost identical sculptures, well developed foot layers and thick continuous tecta. Of the three genera which comprise subtribe Pycnocominae, pollen of Pycnocoma has a unique exine structure : the columellae are reduced to proximal points of poorly connected (as demonstrated in TEM) tectal elements, the foot layer is threadlike to absent, and the only continuous layer is a thickened endexine. Pollen of the monotypic Droceloncia has a psilate, irregularly thickened tectum and poorly de¢ned short columellae, while that of Argomuellera has a ¢nely reticulate tectum, uniform columellae and wide colpi; they share the absence of microspinules. The pollen data support the present concept of Blumeodendrinae, but not that of Pycnocominae. Pycnocoma should, at the very least, be segregated as a monogeneric subtribe. The disposition of Droceloncia and Argomuellera is less clear, but on the basis of three collections examined, the pollen of the two genera is not similar.

Tribe 25. Bernardieae (Part 2, Plates XVII^XIX) Pollen of Bernardia, Necepsia, Paranecepsia, Discocleidion and Adenophaedra, the ¢ve genera that comprise the Bernardieae, is three-colporate with elongate colpi, psilate margos, and punctate or deeply punctate tecta. Individually, these characteristics are not particularly notable, in combination, however, they support the present concept of the tribe. We agree with Webster’s (1994) comments that based on pollen alone, Bernardia is not closely related to Adelia as suggested by Baillon (1858) nor to Macaranga as suggested by Hutchinson (1969).

Tribe 27. Epiprinae (Part 3, Plates I^VI) The Epiprineae comprise nine genera, of which eight, Epiprinus, Symphyllia, Adenochlaena, Cleidiocarpon, Koilodepas, Cladogynos, Cephalocrotonopsis and Cephalocroton, belong to subtribe Epiprininae. Grains of this subtribe have punctate to deeply punctate tecta and well developed columellae (some species of Koilodepas may be excepted). With the exception of Adenochlaena leucocephala, in which the lateral margins of the endoaperture are forked (Part 3, Plate II, 10), none of these genera have distinctive pollen. However, pollen of the monogeneric subtribe Cephalomappinae (Part 3, Plate VI) can be distinguished from all remaining members of Acalyphoideae by a combination of porelike apertures and reticulate exines with broad muri. The disparity between pollen of Cephalomappa and that of subtribe Epiprininae is su⁄ciently great that the former should be treated as a monogeneric tribe.

Tribe 26. Pycnocomeae (Part 2, Plates XX^XXV) The Pycnocomeae comprise seven genera segre-

Tribe 28. Adelieae (Part 3, Plates VII^XII) In the Adelieae, pollen of three genera, Adelia,

PALBO 2467 11-10-02

332

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

Lasiocroton and Leucocroton, has strati¢ed opercula, microcrotonoid tecta and exines with very thin foot layers and small columellae. Grains of Enriquebeltrania and Crotonogynopsis lack opercula, have punctate tecta and a recognizable foot layer. Furthermore, the latter genus (Part 3, Plate IX) has a complex infratectum similar to many members of Alchorneae (see tribe 29, immediately below). If the tribe is maintained, Adelia, Lasiocroton and Leucocroton should comprise one subtribe, and Enriquebeltrania and Crotonogynopsis placed in monogeneric subtribes. Tribe 29. Alchorneae (Part 3, Plates XII^XVII) Seven of the nine genera (no material of Bossera nor Polyandra) in tribe Alchorneae share pollen with unstrati¢ed opercula and elongated columellae near the very small endoapertures. Grains of subtribe Alchorneinae (Part 3, Plates XII^ XVI), Or¢lea, Alchornea, Coelebogyne, Aparisthmium and Bocquillonia, are further distinguished by exines in which the infratectum is modi¢ed ^ two irregular layers of columellae or one layer in which the columellae are expanded midlength, or both. This exine structure was not found in Conceveiba and Gavarretia (Part 3, Plate XVI), two of the three genera of subtribe Conceveibinae. The pollen data support the present concept of Alchorneae. Tribe 30. Acalypheae (Part 3, Plates XVII^ XXVII ; Part 4, Plates I^XXVI) With 30 genera, the Acalypheae could be expected to have greatest pollen diversity of the subfamily. In fact, with limited exceptions, the pollen of much of this tribe is similar. The large genera ^ Macaranga, Mallotus, and Acalypha have uniform pollen. Although the endoapertures of Acalypha are considered to be too small to be seen in LM, and subtribes Mercurialinae and Dysopsidinae have colpate apertures, the remaining taxa have compound apertures. Grains of Ricininae, with only Ricinus (Part 3, Plate XVII), and of Adrianinae, with only Adriana (Part 3, Plates XVIII, XIX), have almost identical tecta and identical exines with thin foot layers, small columellae and thick tecta. The two

genera could be treated in the same subtribe instead of consecutive monogeneric ones. Pollen of Mercurialinae (Part 3, Plates XX, XXI) with Mercurialis and Leidesia (no material of Seidelia) and of Dysopsidinae (Part 3, Plate XXII) with only Dysopsis is su⁄ciently similar for the two subtribes to be merged. These genera share colpate apertures, microreticulate exines and elongate columellae. These grains are among the more distinctive in the Acalypheae. Pollen of Dysopsis is not inaperturate as has been reported. The subtribe Cleidiinae (Part 3, Plates XXII^ XXVI) consists of two monotypic genera, Wetria and Sampantaea, and a third, Cleidion, with 25 species. Grains of all three have punctate or microrugulate tecta, short colpi and thin foot layers. Pollen of Cleidion and Sampantaea has a thick tectum, but the latter di¡ers in having a prominent endexine and very short columellae. Pollen of Wetria is more generalized, but has a tectum very similar to that of Adriana and Ricinus (Part 3, Plates XVII^XIX). Pollen of the monogeneric subtribe Macaranginae (Part 3, Plates XXVI, XXVII) has the most common exine structure in Acalyphoideae ^ thin foot layers, short irregular columellae and thick continuous tecta. Pollen of the remaining taxa of Acalyphoideae is discussed in more detail because they are ¢rst described in this paper. Tribe 30. Acalypheae con. (Part 4, Plates I^ XXVI) Subtribe 30g. Claoxylinae (Part 4, Plates I^XI) This taxon includes seven genera. Pollen of Erythrococca, Claoxylon, Discoclaoxylon, Mareyopsis and Micrococca is similar : punctatemicrospinulose or microcrotonoid sculpture, and unremarkable exine structures except for the latter two genera having longer columellae. Some species of Erythrococca have markedly longer columellae at the poles. The one collection examined of Claoxylopsis has pollen with a threadlike endexine^foot layer, irregular columellae, and a tectum with a minutely ‘fringed’ inner surface. Exines of Mareya have well-de¢ned foot layers, thin short columellae, and a thick continuous tectum. Pollen

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

data would support, albeit not strongly, the reestablishment of Mareyopsis. The seventh genus of Claoxylinae, Amyrea, is the only member of Acalyphoideae to have pollen with a striate surface, but it is a supratectal sculpture, the tectum is not composed of lirae (striae are the grooves). Nevertheless, this striate sculpture and bi¢d styles, also unique in Claoxylinae, and recent molecular data, reinforce Webster’s view (1994) that the position of Amyrea needs to be re-evaluated. Subtribe Claoxylinae, without Amyrea, is palynologically uniform. Subtribe 30h. Lobanilinnae (Part 4, Plate XII) The two examined species of the monogeneric subtribe Lobaniliinae (Part 4, Plate XII) have different pollen. One species, Lobanilia bakeriana, has exines with uniform but thin foot layers, sparse but well-developed columellae and a tectum of uniform thickness; the other, L. claoxyloides, has a thin irregular foot layer, smaller more numerous columellae and a tectum of irregular thickness. The pollen data are equivocal about supporting the treatment of Lobanilia as a monogeneric subtribe. Subtribe 30i. Rottlerinae (Part 4, Plates XIII^XX) Seven of the eight genera (no material of Coccoceras) were examined. The grains of six, Mallotus, Deuteromallotus, Cordemoya, Trewia, Neotrewia and Octospermum, are similar : punctatemicrospinulose or occasionally microscabrate tecta, and an exine structure of thin foot layer^endexines, short thin columellae, and a thick continuous tectum. Pollen of the Australian genus Rockinghamia is somewhat di¡erent ^ microreticulate or deeply punctate tecta, thicker foot layers and more substantial columellae. Whether Rockinghamia should be treated in a monogeneric subtribe is debatable, but without it, the Rottlerinae are a palynologically very uniform subtribe. Subtribe 30j. Acalyphinae (Part 4, Plates XXI^ XXIV) The large genus Acalypha is treated as a monogeneric subtribe (Part 4, Plates XXXI^XXIV) and the pollen supports this distinction. Grains of Acalypha are small, oblate, angulaperturate with

333

very short, slightly protruding colpi and exines with small or thin columellae and a thick continuous tectum. Subtribe 30k. Lasiococcinae (Part 4, Plates XXV, XXVI) This subtribe comprises three genera, Lasiococca, Spathiostemon and Homonoia, of which the latter two have remarkable pollen. Grains of Spathiostemon have a continuous tectum that has a sculpture of narrow lirae, each of which has super¢cial cross striations that, overall, produce a pattern of minutely segmented strings. Exines of Homonoia consist mostly of a tectum of individual, vertically oriented narrow rods whose length determines the thickness and whose random arrangement and acute tips produce a minutely echinate surface. The tectum of Spathiostemon is fundamentally very di¡erent from tectum of Homonoia, and yet exines of both genera have thin foot layer^endexines and very sparse, short columellae. Pollen of a depauperate sample of Lasiococca shows a tectum sculpture more similar to those of some members of Erythrococca (Part 4, Plates I, II) and some species of Mallotus (Part 4, Plates XII^XVI). The pollen data suggest that these three genera may not be as closely related as has been thought, but until additional material can be examined, especially of Lasiococca in TEM, Lasiococcinae should be maintained as is. Tribe 31. Plukenetieae (Part 4, Plates XXVII^ XLII) Pollen grains of this large tribe, 16 genera (no material of Sphaerostylis), have the greatest diversity in exine structure, tectal/surface morphology, and aperture types of the Acalyphoideae, if not the Euphorbiaceae. Of the three subtribes, Plukenetiinae have seven genera, Tragiinae eight, and Dalechampiinae one. Subtribe 31a. Plukenetiinae (Part 4, Plates XXVII^XXXI) All pollen of this subtribe is three-colpate. Pollen of Romanoa tamnoides, Eleutherostigma lehmannianum, Plukenetia conophora, P. polyadenia and P. volubilis (the type species of Plukenetia) is the Conophora type (punctate tecta, elongate

PALBO 2467 11-10-02

334

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

columellae, branched columellae and/or a granular area under the tectum). Pollen of Plukenetia brachybotrya, P. loretensis, P. penninervia and Vigia serrata is the Loretensis type (reticulate exines, crenate muri, unremarkable structure). Pollen of Haematostemon is similar to the Conophora type but has an irregular thickening of the foot layer^endexine along the colpi and a different tectum. Grains of Angostyles also have this irregularly thickened foot layer^endexine, but have short columellae and lack a granular area under the tectum. We defer comments on the relationships of Astrococcus because the pollen was not examined in TEM. To summarize the new relationships of Plukenetiinae suggested by the pollen data: the genus Plukenetia would be circumscribed to include only those taxa with the Conophora pollen type (P. conofora, P. polyadenia, P. volubilis, Eleutherostigma and Romanoa); those species of Plukenetia with the Loretensis pollen type (P. brachybotrya, P. loretensis, P. penninervia and Vigia) would constitute a second genus; Haematostemon and Angostyles should be maintained. Subtribe 31b. Tragiinae (Plates XXXII^XL, 245^ 320) The extent of pollen diversity in the Tragiinae is greater than in any remaining tribes or subtribes of the subfamily. Grains can be three-colpate, unequivocally inaperturate, or with poorly de¢ned apertures. Pollen walls can be tectate with punctate-microspinulose, punctate, variously reticulate or microrugulate sculptures, or intectate with columellate surfaces. Some exines consist almost wholly of columellae, while others have scarcely any. Foot layers range from probably the thickest in the subfamily to threadlike to unequivocally absent. Pollen of Cnesmone, of Megistostigma and of Pachystylidium have poorly de¢ned apertures and exines consisting of threadlike endexines, small sparse columellae and thin tecta. Within the large genus Tragia, several pollen types were recognized. The intectate exines of the Ramosa pollen type of Tragia consist of a thick channeled foot layer and large, stout columellae. These three-colpate grains occur in T. ramosa, T. styla-

ris, T. urticaefolia, and T. volubilis, all of which belong to Tragia sect. Tragia. The Lukafuensis pollen type (three-colpate, arched colpal margins at the equator, reticulate exines and crenate muri) occurs in Tragia gardneri, T. lukafuensis, T. mitis, T. pungens and T. spathulata, all of which belong to Tragia sect. Tagira. Pollen very similar to the Lukafuensis type was found in the two examined species of Tragiella. The two species, Tragia smallii and T. urens, that constitute Tragia sect. Leptobotrys have pollen with poorly de¢ned apertures and exines consisting of short columellae and an irregularly thick tectum. Pollen of T. sellowiana is inaperturate and has an exine consisting of a threadlike foot layer^endexine and stout columellae expanded distally to form an irregular discontinuous, thin tectum. Inaperturate grains also occur in the Cuban endemic Platygyna and the two West Indian species of Acidoton, A. microphyllus and A. urens, all of which have semitectate exines that consist mostly of massive irregular columellae. The absence of two ‘‘horizontal’’ layers, tectum and foot layer, may be adaptations for an easily ruptured wall, thereby facilitating germination. The Central (and South?) American species of Acidoton have three-colpate grains with an unremarkable exine structure. To summarize new relationships in Tragiinae suggested by the pollen data: the three Asian genera, Cnesmone, Megistostigma and along Pachystylidium should be segregated as a subtribe ; the genus Tragia would be restricted to sect. Tragia, the Ramosa pollen type is unique within the Acalyphoideae; Tragia sect. Tagira with the Lukafuensis type would be merged with Tragiella as another genus; Acidoton would be restricted to A. microphyllous and A. urens, and along with Platygyna would be segregated as a subtribe or tribe; the mainland species of Acidoton would revert to Gitara Pax et K. Ho¡mann. Admittedly, this scheme does not account for the disposition of Tragia baillonia, T. cordifolia, T. sellowiana, T. urens and T. smallii. Subtribe 31c. Dalechampiinae (Part 4, Plates XLI^XLIII) Dalechampia is presently treated as a monogeneric subtribe of Plukenetieae. The very large,

PALBO 2467 11-10-02

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

thick-walled grains (see Tables 2 and 3) of this genus can be easily distinguished from all remaining members of Euphorbiaceae by costa endocingula ^ two prominent equatorial bands, probably an extreme enhancement of the margins of costae. Dalechampia should be given tribal status. Tribe 32. Omphaleae (Part 4, Plates XLIV, XLV) The last tribe of Acalyphoideae is the monogeneric Omphaleae with seemingly unspecialized or generalized pollen, although both columellae and foot layer can be very irregular. These three-colpate grains share a reaction to acetolysis with members of Crotonoideae that have threecolpate pollen, Adenocline Turczaninov, Ditta Grisebach, Endospermum Bentham, Tetrorchidium Poeppig (Nowicke, 1994), a separation of a relatively thick non-apertural endexine from the ectexine. And, Wurdack’s molecular data (personal communication) support a close relationship among Omphalea, Ditta, Tetrorchidium and Endospermum. But apart from this similar reaction to acetolysis, the pollen of Omphalea is very di¡erent from pollen of all Crotonoideae. Previous suggestions that Omphalea is related to Plukenetieae (Croizat, 1942; Punt, 1962) are less easily dismissed. Given the extremes of pollen diversity of this tribe, the inclusion of Omphalea would not unduly stretch the boundaries, but, at the same time, the pollen does not match anything in the tribe Plukenetieae. The view of Gillespie and Armbruster (1997) that Omphaleae probably represent an ancient lineage in the uniovulate Euphorbiaceae seems appropriate.

Acknowledgements We thank Willem Punt for his critical reading of the manuscript and Kenneth Wurdack for sharing his results from molecular studies of Euphorbiaceae. We also thank Sylvia Orli and Stanley Yankowski for their technical assistance, and the curators at AA, GH, K, L and MO for allowing us to remove pollen samples from collections of Acalyphoideae. This work was supported by four Short Term Visitor Awards to Masamichi

335

Takahashi from the Smithsonian Institution’s Of¢ce of Fellowships and Grants.

References Armbruster, W.S., 1988. Multilevel comparative analysis of the morphology, function, and evolution of Dalechampia (Euphorbiaceae) blossoms. Ecology 69, 1746^1761. Armbruster, W.S., 1994. Early evolution of Dalechampia (Euphorbiaceae): insights from phylogeny, biogeography, and comparative ecology. Ann. MO Bot. Gard. 81, 302^ 316. Baillon, H., 1858. Etude Ge¤ne¤rale du Groupe des Euphorbiace¤es. Victor Masson, Paris. Breteler, F.J., Mennega, A.M.W., Punt, W., 1997. Novitates gabonenses (29). A new species of Mareyopsis PaxpK. Ho¡m. (Euphorbiaceae) from Gabon with notes on the taxonomic position of the genus. Bull. Jard. Bot. Natl. Belg. 66, 131^148. Croizat, L., 1942. On certain Euphorbiaceae from the tropical Far East. J. Arnold Arbor. 23, 29^54. Dickison, W.C., Nowicke, J.W., Skvarla, J.J., 1982. Pollen morphology of Dilleniaceae and Actinidiaceae. Am. J. Bot. 69, 1055^1073. Erdtman, G., 1966. Pollen Morphology and Plant Taxonomy. Almqvist and Wiksells, Uppsala. Ferna¤ndez-Gonza¤lez, D., Lobreau-Callen, D., 1996. Le pollen de la tribu des Acalypheae (Acalyphoideae, Euphorbiaceae). Grana 35, 266^284. Gillespie, L.J., 1993. A synopsis of Neotropical Plukenetia (Euphorbiaceae) including two new species. Syst. Bot. 18, 575^592. Gillespie, L.J., 1994. Pollen morphology and phylogeny of the tribe Plukenetieae (Euphorbiaceae). Ann. MO Bot. Gard. 81, 317^348. Gillespie, L.J., Armbruster, W.S., 1997. A contribution to the Guianan Flora: Dalechampia, Haematostemon, Omphalea, Pera, Plukenetia, and Tragia (Euphorbiaceae) with notes on the subfamily Acalyphoideae. Smithson. Contrib. Bot. 86, 1^46. Hutchinson, J., 1969. Tribalism in the family Euphorbiaceae. Am. J. Bot. 56, 738^758. Nowicke, J.W., 1994. A palynological study of Crotonoideae (Euphorbiaceae). Ann. MO Bot. Gard. 81, 245^269. Nowicke, J.W., Skvarla, J.J., 1977. Pollen morphology and the relationships of the Plumbaginaceae, Polygonaceae and Primulaceae to the order Centrospermae. Smithson. Inst. Contrib. Bot. 37, 1^64. Nowicke, J.W., Skvarla, J.J., 1981. Pollen morphology and the phylogenetic relationships of the Berberidaceae. Smithson. Contrib. Bot. 50, 1^83. Nowicke, J.W., Takahashi, M., Webster, G.L., 1998. Pollen morphology and exine structure of Acalyphoideae (Euphorbiaceae). Part 1. Tribes Clutieae (Clutia), Pogonophoreae (Pogonophora), Chaetocarpeae (Chaetocarpus, Trigono-

PALBO 2467 11-10-02

336

J.W. Nowicke, M. Takahashi / Review of Palaeobotany and Palynology 121 (2002) 231^336

pleura), Pereae (Pera), Cheiloseae (Cheilosa, Neoscortechinia), Dicoelieae (Dicoelia), Galearieae (Galearia, Microdesmis, Panda) and Ampereae (Amperea, Monotaxis). Rev. Palaeobot. Palynol. 102, 115^152. Nowicke, J.W., Takahashi, M., Webster, G.L., 1999. Pollen morphology and exine structure and systematics of Acalyphoideae (Euphorbiaceae). Part 2. Tribes Agrostistachydeae (Agrostistachys, Pseudagrostistachys, Cyttaranthus, Chondrostylis), Chrozophoreae (Speranskia, Caperonia, Philyra, Ditaxis, Argythamnia, Chiropetalum, Doryxylon, Sumbaviopsis, Thyrsanthera, Melanolepis, Chrozophora), Caryodendreae (Caryodendron, Discoglypremna, Alchorneopsis), Bernardieae (Bernardia, Necepsia, Paranecepsia, Discocleidion, Adenophaedra), Pycnocomeae (Pycnocoma, Droceloncia, Argomuellera, Blumeodendron, Podadenia, Ptychopyxis, Botryophora). Rev. Palaeobot. Palynol. 105, 1^62. Pax, F., Ho¡mann, K., 1919. Euphorbiaceae-Acalypheae-Plukenetiinae. In: Engler, A. (Ed.), Das P£anzenreich IV, 147, IX (Heft 85), pp. 1^62. Pax, F., Ho¡mann, K., 1931. Euphorbiaceae. In: Engler, Prantl (Eds.), Die Natu«rlichen P£anzenfamilien, edn. 2, Vol. 19c, pp. 11^233. Poston, M.E., Nowicke, J.W., 1993. Pollen morphology, trichome types, and relationships of the Gronovioideae (Loasaceae). Am. J. Bot. 80, 689^704. Punt, W., 1962. Pollen morphology of the Euphorbiaceae with special reference to taxonomy. Wentia 7, 1^116. Punt, W., Blackmore, S., Le Thomas, A., Nilsson, S., 1994.

Glossary of pollen and spore terminology. Lab. Palaeobot. Palynol. Contrib. Ser. 1, 1^71. Suarez-Cervera, M., Gillespie, L., Arcalis, E., Le Thomas, A., Lobreau-Callen, D., Seoane-Camba, J.A., 2001. Taxonomic signi¢cance of sporoderm structure of Euphorbiaceae: tribes Plukenetieae and Euphorbieae. Grana 40, 78^104. Takahashi, M., Nowicke, J.W., Webster, G.L., Orli, S.S., Yankowski, S., 2000. Pollen morphology, exine structure and systematics of Acalyphoideae (Euphorbiaceae), Part 3. Tribes Epiprinae (Epiprinus, Symphyllia, Adenochlaena, Cleidiocarpon, Koilodepas, Cladogynous, Cephalocrotonopsis, Cephalocroton, Cephalomappa), Adelieae (Adelia, Crotonogynopsis, Enriquebeltrania, Lasiocroton, Leucocroton), Alchorneae (Or¢lea, Alchornea, Coelebogyne, Aparisthmium, Bocquillonia, Conceveiba, Gavarretia), Acalypheae pro parte (Ricinus, Adriana, Mercurialis, Leidesia, Dysopsis, Wetria, Cleidion, Sampantaea, Macaranga). Rev. Palaeobot. Palynol. 110, 1^66. van Welzen, P.C., 1998. Revisions and phylogenies of Malesian Euphorbiaceae: subtribe Lasiococcinae (Homonoia, Lasiococca, Spathiostemon) and Clonostylis, Ricinus, and Wetria. Blumea 43, 131^164. Webster, G.L., 1994. Synopsis of the genera and suprageneric taxa of Euphorbiaceae. Ann. MO Bot. Gard. 81, 33^144. Wen, J., Nowicke, J.W., 2000. Pollen ultrastructure of Panax (the Ginseng genus, Araliaceae), an Eastern Asian and Eastern North American disjunct genus. Am. J. Bot. 86, 1624^ 1636.

PALBO 2467 11-10-02