IAWA Bulletin n.s., Vol. 3 (1),1982 15

SYSTEMATIC WOOD ANATOMY OF DESMOPSIS, SAPRANTHUS AND STENANONA ()

by

Ben J.H. ter Welle and Jan van Rooden Institute of Systematic Botany, Heidelberglaan 2, 3508 TC Utrecht, The Netherlands

Summary The wood anatomy of three related genera record and at the same time description of Sa­ of the Annonaceae has been studied and de­ pranthus as a genus was by Seemann in 1866. scribed as part of a research project on the Afterward authors as e.g. Bentham and Hooker taxonomy of neotropical taxa of the family. (1867) and Prantl (1891), included Sapranthus The wood anatomy of the three genera is char­ in other genera and it was not until 1900 that acterised by the occurrence of many fine apo­ Fries reinstated Saprallthus as a separate genus. tracheal continuous concentric bands of paren­ In the first monographic treatment of Sapran­ chyma. The genera Desmopsis, Sapranthus, and thus (Fries, 1931 a, b) seven species are listed. Stenanona, restricted to Central America, can Desmopsis, consisting of inconspicuous under­ be separated on wood anatomical characters storey treelets, was erected by Safford in 1916, such as element length, ray height and the partly as a segregate from the heterogeneous number of parenchyma bands per mm. A key genus Ullonopsis. He recognised five species. for the genera is presented and the possible Fries (1931 a, b) recognised nine species in his phylogenetic relationships are discussed. Final­ monograph of Desmopsis. The genus Stenallo­ ly, the wood anatomical results are correlated na erected by Standley (1929) is based on Ste­ with data on pollen morphology, flower mor­ nallona pallamellsis, a collection ofG.P. Cooper phology and morphology of fruits and leaves. in the Almirante region of Panama. It is still the only collection of the type species. After­ Introduction wards only one other species was added. More This study forms part of a research project new species were found in Saprtmthus and Des­ on the systematics and wood anatomy of neo­ mopsis. Fries's monograph of the Annonaceae tropical Annonaceae. The genera Desmopsis, (1959) finally included 8 Sapranthus and 16 Sapranthus, Stenanona, Anaxagorea and Rolli­ Desmopsis species. In a forthcoming revision nia have been monographed (to be published in of the three genera treated here, several new the near future) and the wood anatomy of the species will be described (Van Rooden, in three former genera is presented here. Separate prep.). publication seems justified because Desmopsis, Sapranthus and Stenanona form a coherent Wood anatomy group of mutually very closely related genera Since Solereder published his 'Systematische (Fries, 1931a, b; Van Rooden, in prep.). It Anatomie der Dicotyledonen' (1899 & 1908), should be realised that some conclusions pre­ in which he stated that the wood anatomy of sented here might need reconsideration after representatives of the Annonaceae is character­ more genera will have been studied. ised by the presence of fine tangential bands of parenchyma, much research has been carried Historical Survey out. Although many more samples of mature wood have been studied, his general remarks Taxonomy have always been confirmed. So far, Moll and Probably due to their uncommon occurrence, J anssonius (1906) have given the most detailed the genera Desmopsis, Sapranthus, and Stena­ description of Annonaceous woods (21 species nona were recognised as distinct taxa only fair­ belonging to 13 genera). Their descriptions in­ ly recently. In the first monograph of the family dicate that the Javanese Annonaceae are very on a world-wide scale by Dunal (1817) we find similar in their wood anatomy. Nevertheless, the first record of a representative of the pres­ they separated these taxa into four groups. ent genus Sapranthus; the species was collected Hess (1946) studied 21 American genera of this by Mocino and Sesse in Mexico during one of family. He states that th'e woods of all Ameri­ their field trips from 1781 ~ 1803 and was can genera are so similar that it is impossible to­ placed by Dunal in the genus Unolla. The next make a workable key for their identification.

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Ingle and Dadswell (1953) described the wood Materials and Methods anatomy of 12 genera from the southwest Paci­ A total of 10 species of 3 genera (18 sam­ fic area. Again the wood anatomy of these ples) was examined. Only samples backed by genera is very similar. Nevertheless, some of herbariu~ vouchers, and identified by the sec­ these genera can be separated anatomically, e.g. ond author were used for this study. Sections on the size of the intervascular pits. and macerations were obtained using standard A treatment of the wood anatomy of the techniques. The wood anatomical descriptions Annonaceae, without geographical limitations, are according to the terminology proposed by was made by Vander Wyk and Can right (1956). the Committee on Nomenclature of the lAW A Their main goal was to establish the relation­ (1964). The average element length in samples ships of this family in the Ranalian complex. is based on at least 25 measurements. In the About 400 species (61 genera) were examined. generic descriptions the range of the averages They state: "The wood anatomical features of are given and, between brackets, the minimum the family revealed a remarkable consistency, and maximum values are given. The ray heights which emphasizes the belief that this is a well­ (in cells and !.lm) were obtained by measuring defined natural group. However, there exists a and counting the highest rays in the tangential considerable degree of generic overlap for the sections, exceptions being eliminated. majority of anatomical features, which, there­ The same procedure was followed for the fore, are of doubtful value in attempting to width of the rays in !.lm, i.e., values are based understand the intrafamiliar relationships of on the widest, yet commonly occurring rays. the Annonaceae." This generic overlap has also The bands of tangentially arranged parenchyma been described by Metcalfe and Chalk (1950). are quoted as number per mm. Values were cal­ Asimina is the only genus which can easily be culated from ten countings over distances of 3 distinguished, as also stated by Vander Wyk mm each. The averages of the element lengths and Canright (1956), by the presence of ring­ were also used to calculate the ratio of fibre porous wood. In a study to establish phylo­ length/vessel member length and parenchyma genetic schemes for the , Gottwald strand length/vessel member length, in the de­ (1977) studied 40 genera of the Annonaceae. scription referred to as F /V and P/V ratio. He also concluded that the family forms a very homogeneous structural group in spite of the Results large number of genera and species. Apart from these more or less comprehen­ Desmopsis Safford (Figs. 3, 6) sive studies, various papers, dealing with one or A genus of c. 17 species consisting of small several taxa of the Annonaceae, have been pub­ trees and shrubs up to 7 m and occurring in lished, e.g. Balan Menon (1959), Benoist (1927), Central America. Its distribution centre is Cos­ Jutte (1958), Lindeman and Mennega (\ 963), ta Rica and Panama. The genus has its northern­ Loureiro (1969. 1970, 1971), Loureiro and Da most extension in the state of Veracruz, Mexi­ Silva (1968), Monteiro and Franca (\ 971). co and reaches as far south as the Dept. of Cho­ Normand (1950), Normand and Paquis (1976), c6, Colombia. Nearly all species of Desmopsis and Perez Mogo1l6n (1973). inhabit the shady understorey of moist forests The wood anatomy of Desmopsis, Sapran­ at low altitudes. Only a few species are known thus, and Stenanona has never been described from altitudes up to 1500 m. in detail. Some remarks have been made by Record and Hess (1943) and by Vander Wyk Material studied: D. bibracteata (Robin­ and Canright (1956). A few characters can be son) Safford: Costa Rica, ARG I (Uw 26771), extracted from the tables they present. diam. 6 cm; provo San Jose, Caffrey 159 (Uw At first view. this literature survey makes it 26774), diam. 4 cm; D. stenopetala (Donn. unattractive to embark on a comprehensive Sm.) R.E. Fries: Belize, Crique Negra, Steven­ study of the wood anatomical diversity in An­ son 105 (Uw 24298, ex SJRw 14889), diam. nonaceae. However. in view of the envisaged 6 cm; D. tuxtlensis Van Rooden (ined.): Mexi­ taxonomic and systematic studies at the Insti­ co, region of Los Tuxtlas. slope of the Volcano tute of Systematic Botany in Utrecht, and the San Martin. Edo. Veracruz. Van Rooden 755 availability of pollen morphological data for (Uw 26193), diam. 3 cm; Van Rooden 760 comparison (Walker, 1971) it seemed worth­ (Uw 26195), diam. 3 cm; D. spec.: Panama, while to re-investigate the wood anatomy of region of Almirante. proVo Bocas del Toro, this family in greater detail. in order to com­ Cooper & Slater 47 (Uw 24086, ex SJRw pare various data sets for the clarification of 10146). diam. 4 em. the intrafamily relationships of the Annona­ N.B.: Uw 24455 from Mexico, Los Tuxtlas ceae. (MEXU II). diam. 15 cm. may be identical

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with D. tuxtlensis from the same region, but no Van Rooden 820 (Uw 26209), diam. 4 cm; herbarium specimen is available. Costa Rica, provo Cartago, Turria1ba, Poveda 84 (Uw 26773), diam. 9 cm;S. palanga R.E. Fries: Growth rings faint or absent, marked by an Costa Rica, between Sardinal and EI Coco, increase in the number of tangential bands of provo Guanacaste, Van Rooden 868 (Uw 26216), parenchyma or by zones of fibres with a thicker diam. 12 cm; Costa Rica, behind Canas, proVo wall and a smaller lumen than in the rest of the Guanacaste, Van Rooden 874 (Uw 26217), ground tissue. Vessels diffuse, solitary (10-45 diam. 8 cm; Costa Rica, provo Guanacaste, Po­ %) and in irregular clusters and occasionally ra­ veda 300 (Uw 26722), diam. 8 cm; S. veluticar­ dial chains of 2-4 (6), 6 to 21 (4-28) per sq. pus Van Rooden (ined.): Guatemala, La Plata, mm, angular and sometimes round, walls 3-5 Record & Kuylen G 132 (Uw 24299, ex SJRw /lm, diameter 60 to 74 (47-103) /lm, vessel 10083), diam. 5 cm. member length 475 to 570 (290-800) /lm. Per­ forations simple. Intervascular pits alternate, Growth rings faint, but always present, mark­ round, minute, 1-3 (4) /lm, usually with rims. ed by either fibres with a smaller lumen and a Vessel-ray pits very scarce, because the vessels thicker wall than the other fibres or by an in­ are only sporadically in contact with the rays; crease of the number of vessels per sq. mm or if present, identical to the intervascular pits but by a less regular occurrence of the small tan­ half-bordered. Vessel-parenchyma pits similar. gential bands of parenchyma. Vessels diffuse, Fibres non-septate, diameter II to IS /lm, walls solitary (5-22 %) and in radial chains and ir­ 3-5 /lm. Pits simple, equally frequent on the regular clusters of 2-4 (6), 15 to 53 02-63) radial and tangential walls, 1-3 (4) /lm, often per sq. mm, round to sometimes slightly oval, with crossed apertures. Length 1260 to 1520 walls (3) 4-6 /lm, diameter 55 to 92 (32-105) (840-1890) /lm; F/V ratio 2.58-2.95. Rays /lm, vessel member length (290) 340 to 470 homogeneous or mostly weakly heterogeneous, (210-570) /lm. Perforations simple. Intervas­ with few uniseriates and mostly 6-9-seriate, cular pits alternate, round. minute. 2-4 /lm. composed of procumbent and weakly procum­ seldom with rims. Vessel-ray pits are very scar­ bent and sporadically square and sheath cells. ce, because the vessels are only sporadically in Height of the uniseriate rays 8 to 13 cells (= contact with the rays; if present, half-bordered. 310-480 /lm), 18 to 30 /lm wide. Multiseriate similar to the intervascular pits or slightly lar­ rays 70 to 132 cells (= 2200-3400 /lm) high, ger. Vessel-parenchyma pits 4-6 /lm. similar. and 130 to 240 /lm wide; 3 to 5 (2-7), in gen­ Fibres non-septate, diameter 9 to IS /lm, walls eral 3 to 4 per mm. Parenchyma in apotracheal, 3-5 /lm. Pits simple, occasionally indistinctly continuous, concentric tangential bands, 1-2 bordered, on the radial walls and in general cells wide, with 3-5 fibres between two bands. much less frequent on the tangential walls. 1-3 Number of bands 12-·18 per mm. Strands of 4 /lm. Length (700) 940 to 1235 (450-1680) /lm; cells; length 505 to 555 (420-660) {..lI11, P/V ra­ F IV ratio 2.43-3.6\. Ral's homogeneous or tio 0.99-\,19. Deposits are very scarce, if pres­ mostly weakly heterogeneous. with few uniseri­ ent yellowish brown, amorphous, in the rays ates and mostly (3-) 5 -8-seriate rays, composed and vessels. of weakly procumbent, procumbent and spora­ dically square and sheath cells. Height of the Sapranthus Seemann (Figs. 1,4) uniseriate rays 5 to 8 (10) cells (= 180-300 A genus comprising 10 species of trees or /lm), 15 to 20 /lm wide. Multiseriate rays 35 to shrubs of medium size, from a few metres up 60 cells high (= 800-1680 /lm) and 50 to 150 to about 12 m: Only known from Central Ame­ /lm wide; 4 to 7 (2-8) per sq. mm.Parenchyma rica, from the state of Sinaloa in Mexico south­ in apotracheal, continuous, concentric tangen­ ward into Costa Rica and at altitudes from 0- tial bands, 1(-2) cells wide, with 5-8 fibres 1500 m. between two bands. Number of bands 8--11 per mm. Strands of 2-4, mostly 4 cells; length Material studied: S. hirsutus Van Rooden 350 to 475 (270-480) /lm. P/V ratio 0.99- (ined.): Honduras, ravine north of Siguate­ 1.19. Deposits rare, amorphous, yellow to peque, depto. Comayagua, Van Rooden 850 brown, in the vessels. rays and axial parenchy­ (Uw 26214), diam. 10 cm; S. micro carpus ma. (Donn. Sm.) R.E. Fries: Honduras, Las Limas, depto. Comayagua, Edwards 332 (Uw 24088), StenallUlla Standley (Figs. ~. 5) diam. 3 cm; S. nicaraguensis Seemann: Hondu­ A genus of:2 rarc specics. both small trees of ras, Copan Ruinas. depto. Copan, Van Rooden about 3-6 m tall. S. pallalllcllsis was found in 200 (Uw 24194). diam. 3 cm; Guatemala. be­ the understorey of tropical lowland forest at tween Jocotan and Copan, depto. Chiquimula. the Caribbean coast of Panama. S. c()staricellsis

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Downloaded from Brill.com09/23/2021 07:58:46PM via free access IAWA Bulletin n.s., Vol. 3 (1),1982 19 occurs in the understorey of tropical lowland the presence of fine, concentric, apotracheal and lower mountain forests of the eastern side tangential parenchyma bands; weakly hetero­ of the Central Cordillera in Costa Rica. geneous to homogeneous rays; fibres with sim­ ple or indistinctly bordered pits; diffuse vessels, Material studied: S. costaricensis R. E. solitary and in irregular clusters and/or radial Fries: Costa Rica, lower slopes ofVo1can Orosi, chains, with small intervascular pits; uniseriate provo Guanacaste, Wibur & Stone 10706 (Uw rays few or absent; fibres non-septate. On the 26231), diam. 8 cm; S. panamensis StandI.: Pa­ transverse section the three genera are charac­ nama, region of Almirante, proVo Bocas del terised by the occurrence of the above describ­ Toro, Cooper 427, type (Uw 24300, ex MADw ed bands of parenchyma, which feature is typi­ 27009), diam. 5 cm. cal for the Annonaceae (Moll & Janssonius, 1906; Vander Wyk & Canright, 1956). There­ Growth rings faint, but always present, indi­ fore these genera match perfectly the Annona­ cated by a slightly thicker wall of the ground ceous wood anatomical character complex. tissue or by an increase in the width of the tan­ However, many characters, especially the gential bands. Vessels diffuse, solitary (11-27 quantitative ones, are rather constant within a %) and in radial chains and sometimes irregular genus, and mostly do not overlap (see Table I) clusters of 2-4 (6), 26 to 37 (17-47) per sq. in the three genera studied. As already pointed mm, angular and occasionally round, walls 3-4 out in the introduction, these three genera con­ ,urn, diameter 55 to 66 (42-88) ,urn, vessel stitute a taxonomic unit based on morphologi­ member length 405 to 425 (240-660) ,urn. Per­ cal characters. Based on a combination of cer­ forations simple. Intervascular pits alternate, tain features, they can be separated on the round, minute, 3 ,urn. Vessel-ray pits are scar­ wood anatomical characters. Whether the sug­ ce, because the vessels are only sporadically in gested key will remain useful after more genera contact with the rays. Vessel-parenchyma pits have been studied wood anatomically is uncer­ similar to the intervascular pits, but half-bor­ tain. A workable key to the three genera exam­ dered and sometimes larger (up to 4.5 ,urn). ined can be constructed as follows: Fibres non-septate, diameter 12 to 15 ,urn, walls 3-4 ,urn. Pits simple, equally frequent on the I a. The highest multiseriate rays over 2200 ,urn, tangential walls and the radial walls, 3 ,urn, in commonly over 70 cells high, and the avo part with crossed apertures. Length 940 to 1045 vessel member length over 475 ,urn (630-1320) ,urn; F/V ratio 2.32-3.61. Rays Desmopsis homogeneous or weakly heterogeneous; exclu­ b. The highest multiseriate rays less than 2200 sively multiseriate, 6 to II cells wide, composed ,urn, commonly less than 70 cells, and the av. mainly of weakly procumbent and some pro­ vessel member length less than 475,um . 2 cumbent and square cells. Height 53 to 72 cells 2a. Vessels angular, multiseriate rays over 160 (= 1850-2150 ,urn), 175 to 280 ,urn wide: 3 ,urn wide, avo number of rays 3 per mm, and (2-4) per mm. Parenchyma in apotracheal, uniseriate rays absent ...... Stenanona continuous, concentric tangential bands, 1-2 b. Vessels round to slightly oval, multiseriate cells wide, with 3-4 (5) fibres between two rays less than 160 ,urn wide, avo number of bands. Number of bands 13-18 per mm. rays 4-7 per mm, and uniseriate rays pres- Strands of 3-4 cells; length 430 to 455 (330- ent ...... Sapranthus 540) ,urn, P/V ratio 1.07. Yellow and brown deposits occur in the ray cells and sporadically The possibility to distinguish between these in the axial parenchyma cells. three genera using avo element length and ray height and width is also demonstrated in Figs. Key to the genera 7 and 8. Although these quantitative values show The wood anatomy of the three genera ex­ some overlap, the tendencies for the genera are amined show many characters in common, e.g. clear. Desmopsis has the highest values for the

Fig. 1. Cross section of Sapranthus pa/anga (Uw 26216). - Fig. 2. Cross section of Stenanona pa­ namensis (Uw 24300). - Fig. 3. Cross section of Desmopsis spec. (Uw 24086). The difference in the number of parenchyma bands is obvious. Note also the angular vessels in Figs. 2 and 3, and the rounded vessels in Fig. I. - Fig. 4. Tangential section of Sapranthus microcarpus (Uw 24088). - Fig. 5. Tangential section of Stenanona panamensis (Uw 24300). - Fig. 6. Tangential section of Desmopsis spec. (Uw 24086). The ray height increases from Fig. 4 to Fig. 6. Note also the much smaller rays in Fig. 4.

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1600

E 1500 o STENANONA '" .3 • SAPRANTHUS 1400 l:J. DESMOPSIS "'''' 140 ~ '" <:) <: 1300 '" ~ "'''' 130 o STENANONA 1200 • • SAPRANTHUS ct • • ~ '"co ••• i;j 120 6- DESMOP51S c: 1100

200 300 400 500 600 10 11 AV, VESSEL MEMBER LENGTH (J-Im) RAY WIDTH IN CELLS

Fig. 7. Relationship between the average vessel Fig. 8. Relationship between ray height and ray member length, average fibre length and average width (in cells) of the multiseriate rays of three parenchyma length of three genera of the An­ genera of the Annonaceae. nonaceae. av. fibre length, av. vessel member length and The cladistic approach av. parenchyma strand length and also for the The selection of apomorphic character states height of the multiseriate rays. in wood anatomy can be based mainly on the phylogenetic trends established by Bailey Phylogenetic considerations (1944, 1953, 1957) and his students as e.g. In order to establish phylogenetic relation­ Tippo (1946). Their work is based on studies ships between these three genera, as based on of large groups of many families. There­ wood anatomical characters, two approaches fore, a selection of apomorphic character states can be followed. First a cladistic analysis as de­ in a fairly homogeneous family like the Anno­ scribed by Hennig (1966). Essential for this naceae will always be more or less arbitrary, method is the definition of apomorphic (or but keeping this in mind the following charac­ derived) and plesiomorphic (or primitive) char­ ter states might be considered as apomorphic: acter states to various expressions of the char­ I. Short elements (vessel members and paren­ acters. A second method is a purely phenetic chyma strands) length versus long elements. analysis of the wood anatomical characters, 2. Fibre pits more frequent on radial walls than without preference for specialised states. For on tangential walls versus fibre pits equally both methods it is important that variations frequent on both walls. in wood structure as a result of ecological dif­ 3. Round vessels versus angular vessels (cross ferences and/or adaptations as described by section). e.g. Baas (1973) and Carlquist (1975) should 4. Exclusively multiseriate rays versus uniseriate be recognised and eliminated first. However, and multiseriate rays. since Desmopsis. Sapranthus, and Stenanona 5. Low rays versus higher rays. occur under more or less identical ecological In Desmopsis, Saprantilus and Stenanona the conditions, such differences in wood structure number of apomorphic character states is 0, 4 may be ruled out here. and 3 respectively. Character state no. I is the

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only synapomorphy for Sapranthus and Stena­ ies of the Annonaceae are complete". A com­ nona. Although the element length in Desrnop­ prehensive study of the three genera under dis­ sis is higher it may not be considered as very cussion here has been carried out now by Van long and thus the most plesiomorphic character Rooden (in prep.). The identification of the state within the Annonaceae. Nevertheless, for specimens studied by Walker, as far as available, the time being Sapranthus and Stenanona may could be confirmed. Unfortunately no speci­ be considered as sister groups, linked together men of the genus Reedrollinsia, erected by by the shortening of elements and the shorten­ Walker in 1971 and placed near Stenanona and ing of the rays. The loss of uniseriate rays in Sapranthus, was available. According to their Stenanona and the loss of fibre pits on the tan­ pollen morphology the three genera have many gential walls of the fibres in Sapranthus can be characters in common. Some of these charac­ interpreted as autapomorphies. The relation­ ters Walker considers to be derived, viz. aper­ ship of Desrnopsis to either of these two genera tures inaperturate or disuiculate, grains apolar remains uncertain. or isopolar, grains radiosymmetric, the size of the grains, and a globose shape. In the chapter The phenetic approach on phylogeny Walker presents the following Using similarities and differences as a method possible sequence at the beginning of the Uvaria to show phenetic relationships it is obvious tribe: Desrnopsis, Stenanona, Reedrollinsia, Sa­ from the wood anatomical data presented here pranthus. that the three genera have many characters in Sapranthus has disuiculate pollen, while in common, whereas only eight differentiating Desrnopsis and Stenanona the pollen is inaper­ characters can be found (see also Table I). turate. However, a few samples taken from spe­ Four of them link Desrnopsis with Stenanona, cimens not studied by Walker, revealed that viz. angular vessels, width of the rays, fibre pits aperturate pollen sometimes occurs in Desrnop­ equally frequent on radial and tangential walls, sis (Punt, pers. comm.). Further research on and the number of parenchyma bands per mm. the pollen morphology of all taxa of these Three characters link Stenanona with Sapran­ three genera has to be undertaken in the future thus, viz. vessel member length, fibre length, to bring the total variation pattern to light. and the height of the multiseriate rays in cells. Comparing the data given by Walker with Finally, only one character links Desrnopsis those obtained by the investigation of the with Sapranthus, viz. the occurrence of uniseri­ wood anatomy, the following dadogram may ate and multiseriate rays. Thus it might be sug­ be constructed: gested that Stenanona is more or less interme­ diate between Sapranthus and Desrnopsis, be­ Sapranthus Stenanona Desrnopsis cause it has more characters in common with either Sapranthus or Desrnopsis than Desrnop­ sis and Sapranthus have with each other.

General discussion Our study shows that it is possible to select wood anatomical characters by which the genera As the morphological data set cannot be inter­ Desrnopsis, Sapranthus, and Stenanona can be preted yet in terms of plesiomorphic and apo­ separated quite well; despite the general im­ morphic character states, this cladogram must pression from the literature that most genera of be regarded as provisional at this moment. Annonaceae overlap in their wood anatomy. The phenetic relationships of the three genera A phylogenetic intrafamily classification can be discussed using morphological, pollen based on pollen morphology of the family was morphological and wood anatomical characters. presented by Walker (1971). His study includes Among the neotropical Annonaceae these Desrnopsis, Sapranthus and Stenanona. In his genera are put together by their leaf-opposite proposed classification these genera are placed inflorescence with 1-2 bracts along the pedicel in the tribe Uvarieae. (As Fries, 1959, did in his combined with the marginal placentation of classification of the family using morphological the ovules. Although difficult to describe in characters.) Most of the material Walker studied terms, species of these genera share leaf mor­ is also at our disposal, although specimens of phological features which make them recognis­ the material collected by himself are lacking. able, even in sterile condition. Various morpho­ According to him "the hierarchical categories logical characters combine two of the three are only informally proposed at this time be­ genera. Desrnopsis and Stenanona share many cause it is desirable to defer erecting a formal characters, e.g. the shape of the petals and the nomenclature until further morphological stud- few-seeded fruitlets. Sapranthus and Stenanona

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Table I. - Wood anatomical characters in Desmopsis, Sapranthus and Stenanona (Annonaceae)

.::1 ::!'" :;::<:! ~ c ~'" :;:: :;:: !:: ~ i:"! '" t::l. .... C)'" ~ V)'"

Solitary vessels (%) 10-45 5-22 11-27 Shape of vessels A R A Vessels per sq. mm 6-21 15-33 26-37 Vessel wall thickness (/-lm) 3-5 4-6 3-4 Vessel member length (/-lm) 475-570 340-470 405-425 Fibre pits r, t r, (t) r, t Fibre length (/-lm) 1260-1520 940-1235 940-1045 Uniseriate rays + + Width of the widest multiseriate rays (/-lm) 130-240 50-ISO 175-280 Height of highest multiseriate rays in cells 70-132 35-60 53-72 Height of highest multiseriate rays (/-lm) 2200-3400 800-1680 1850-2150 Number of rays (mm) 3-5 4-7 3 Number of fibres between parenchyma bands 3-5 5-8 3-4 Number of parenchyma bands per mm 12-18 8-11 13-18 Length of parenchyma strands (/-lm) 505-555 350-475 430-455

Legenda: Shape of the vessels: A = angular, R = round. Fibre pits: r, t = equally frequent on radial and tangential walls, r, (t) = more frequent on the radial than on the tangential walls. Uniseriate rays: + = present, - = absent. Quantitative data are ranges of mean values for specimens.

share less characters, e.g. a maroon coloured for promptly sending some herbarium speci­ perianth (relatively rare in neotropical Annona­ mens. The discussions with Dr. Wim Punt (Lab­ ceae), and the form/texture of the fruitlets. oratory of Palaeobotany and Palynology, State Finally, Desmopsis and Sapranthus have few University of Utrecht) on pollen morphology characters in common, e.g. the similarity of the showed us the possibilities and limitations of connective appendages. Pollen morphologically interpreting the literature of this data set. Dr. Desmopsis and Stenanona are almost identical. Jifke Koek-Noorman, Prof. Dr. A.L. Stoffers, Sapranthus is easily separated from these two and Prof. Dr. E. Hennipman of the Institute of genera in having disulculate, isopolar pollen Systematic Botany, Utrecht, are acknowledged (Walker, 1971). The evidence from the wood for their comments on various parts of the anatomy is comparable, and confirms the re­ manuscript. Finally, we wish to thank our col­ sults from external morphology and pollen league Drs. L.Y.Th. Westra for correcting the morphology, namely that Stenanona is some­ English text. what intermediate between Sapranthus and Desmopsis, and that Sapranthus is easily sepa­ rable from both Stenanona and Desmopsis. References Baas, P. 1973. The wood anatomical range in Acknowledgements Hex (Aquifoliaceae) and its ecological and Thanks are due to Dr. Jeff Burley (Common­ phylogenetic significance. Blumea 21: 193- wealth Forestry Institute, Oxford) and Dr. Jorgo 258. Richter (Bundesforschungsanstalt fUr Forst­ Bailey, I.W. 1944. The development of vessels und Holzwirtschaft, Hamburg) for loan of in angiosperms and its significance in mor­ wood samples, and Dr. L. Diego Gomez P. phological research. Amer. J. Bot. 31: 421- (Herbario Nacional de Costa Rica, San Jose) 428.

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