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TWO FOSSIL FLOWERS OF TRICHILIA (MELIACEAE) IN DOMINICAN AMBER PDF

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Preview TWO FOSSIL FLOWERS OF TRICHILIA (MELIACEAE) IN DOMINICAN AMBER

TWO FLOWERS OF DOMINICAN AMBER FOSSIL TRICHILIA (MELIACEAE) IN Chambers George Brown Kenton Alex 0. Poinar, L. Jr. E. and Department ofBotany Plant Pathology Department ofZoology 629 Euclid Ave. Oregon State University Berkeley, California 94708, U.S.A. Oregon Corvallis, 97331, U.S.A. RESUMEN INTRODUCTION & A reconstruction of the Late Oligocene/Early Miocene tropical forests of Hispaniola was provided by Poinar Poinar (1999), based on fossil plant and animal remains in amber derived from resin of the arborescent legume Hymenaea protera. Novel taxa recently described from flowers found in these deposits include species of & Chambers Licania (Chrysobalanaceae) (Poinar 2008a, revised by Poinar 2010), Persea (Lauraceae) et al. (Chambers and Moraceae) (Poinar 2008b). The flowers newly reported et al. 2011), Trochanthera (possibly et al. New here are and commercially important family found throughout the Old and referable to Meliaceae, a large World Although on one one tropics and subtropics, but also penetrating into temperate zones. flower the pistil, we petal, and most of the anthers are missing, very likely through the predation of an insect, believe the flower structure of both in particular the characteristic filament tube with sharply pointed lobes between the fossils, We new anthers, point placement Browne. here propose their description as the species to their in Trichilia P. Trichilia and glaesaria Trichilia antiqua. AND METHODS MATERIALS The Dominican between fossils originated from mines in the Cordillera Septentrional of the Republic, the cit- ies of Puerto Plata and Santiago. Dating of these amber deposits is controversial, with a youngest proposed age & of 20-15 mybp, based on foraminifera (Iturralde-Vinent McPhee 1996) and an oldest of 45-30 mybp based on coccoliths Most amber secondarily deposited in turbiditic sandstones of the (Cfipek in Schlee 1999). of the is Upper Eocene Lower Miocene Mamey Group (Draper 1994). to et al. J 8«t - Res. Inst Texas 463-4 5(2): HISPANIOLA. DOMINICAN Chambers glaesaria K.L. i: Trichilia (holotyph: ner s.n. catalogue : Oregon Corvallis, 97331, ty, number Sd-9-27B, deposited in the 1 mm shallowly cup-shaped, with 0.8 long, Flower total length 4.5 nun; calyx abaxially strigillose, pistillate, mm with incurved broadly acute lobes 0.45 long (Fig. petals 5, free, valvate, lanceolate, acute, tip, distinct, 2), mm mm abaxial surface wide, adaxial surface densely low-papillate, strigil- spreading to revolute, 3.4 long, 1.5 mm mm filament tube cyathiform, glabrous, 1.8 long, 3.3 wide (Fig. 1), anthers 10, sterile, sessile, lanceo- lose, mm anther sacs narrow, indehiscent [or dehisced and empty on some anthers, (Fig. glabrous, 0.7-0.9 long, late, higher and between placement of anthers alternating slightly rim of filament tube with acute lobes anthers, 1)], ovary equaling anther tube in length, stigma capitate, minutely papillose, strigillose. lower, present, style pistil — amber. Etymology. from the Latin “glaesarius,” of & K Brown, 3-5). HISPANIOLA. Dominican antiqua L Chambers, Poinar, A.E. sp. nov. (Figs. type: Trichilia mm mm; long no Functional gender of flower uncertain, total length 3.4 calyx glabrous, rotate, ca. 0.8 (Fig. 4), mm mm lobes evident, remaining petals 5 of free, valvate, lanceolate, acute, strongly recurved, 2.6 long, 0.7 6, mm wide, adaxial and abaxial surfaces densely low-papillate, filament tube cylindrical, glabrous, 1.8 long, 1.6 mm mm wide remaining anther of glabrous, subsessile, globose, diameter 0.47 (Fig. 5), (Fig. 1 10, sterile, 3), mm rim of filament tube with acuminate lobes 0.4 long alternating with basal stubs of 9 missing anthers, alter- nate anthers positioned lower on the rim (Fig. pistil apparently damaged due to insect predation. 3), — Etymology. from the Latin “antiquus,” old. The two species, perhaps contemporaneous in the e Mid-Tertiary amber deposit on Hispaniola, are distin- sa; guishable by several floral characteristics. Difference are seen in the pubescence of the abaxial surfaces of the calyx and petals, in the shape and lobing of the caly and in the shape of the filament tubes and depth of zs, placement of the lower versus higher anthers on rim. Minor differences are the incurved petal tips in tl Trichilia glaesaria, which seem not to be as promineni n T. antiqua, the sessile anthers of the first species versus :cond and acuminate lobes of the staminal tube (Fig. is 5), tl (Figs.] we As shows damage which previously indicated, the flower of Trichilia antiqua to reproductive organs, its may believe be due to an insect feeding inside the filament tube on the devouring most of the anthers, pistil, and removing one petal. The single small and malformed anther on the holotype seems to be a developmental abnormality, rather than the result of insect damage. The ovary of a partly damaged can be seen well pistil down The in the filament tube. absence of normal anthers and prevent us from assigning a gender to this pistil | The flower. perianth parts, filament tubes, and anther arrangement of both fossils favor their chosen assign- [ ment to Trichilia. The alternation of sharply pointed lobes with the sessile or subsessile anthers on the rim of & the staminal tube in both species is characteristic of many neotropical taxa of this genus (Pennington Styles j Pennington 1975, et al. 1981). Also, the placement of the anthers in higher and lower positions on the rim of the tube found some is in Trichilia species, example and DC. and magni- for T. appendiculata (Triana Planch.) C. T. Pennington foliola (Pennington C. et al. 1981). Adaxially papillose petals are found in glabra claussenxi T. L., T. ] DC., pallida Swartz, and among T. C. DC., and ovary T. pittieri others, while abaxially strigillose calyx, petals, common New are World in species of the genus (Pennington et 1981). al. Due to the Mid-Tertiary age of the two holotypes, and to the lack of information on features of critical their vegetative habit and male we an flowers, favor their description as novel species rather than proposing 465 Two flowers of Trichilia in Dominican amber Chambers et fossil al.. and Fig. 1. Trichilia glaesaria. Flower in lateral view, showing alternating anthers and acute lobes on rim of filament tube. Note incurved tip of petal = mm. empty anther Bar 0.8 capitate stigma visible within filament tube. Arrows indicate dehisced, sacs. Chambers et aL, Two fossil flowers of Trichilia in Dominican amber 467 Fig. 5. Trichilia antiqua. Apical view of flower, showing rim of filament tube with remnant of abnormally developed anther. Anther tube obscured by from = mm. debris possible insect herbivore. Bar 0.6 & uncertain placement in particular extant species of this large and diverse genus (Pennington Styles 1975). The critical differences between the two are in the shape and pubescence of the calyx and the abaxial fossils pubescence of the corolla. The relatively cup-shaped staminal tube of T. glaesariq versus the cylindrical one of T. antiqua might suggest only intraspecific variability, as for example in the difference between staminate and pistillate flowers in moschata Swartz, as illustrated by Pennington et (1981, 194). Variation in calyx T. al. p. shape and pubescence also occurs in some modern species, e.g. T. schomburgkii C. DC. (Pennington et al. 1981, pp. 162-167), in which the calyx may be “patelliform with 4-5 broadly ovate or triangular lobes. .or [the] . margin ± truncate, sparsely minutely puberulous or glabrate.” As we have only the two examples at hand and cannot examine species variability, we have chosen to describe them as separate taxa, hoping this will assist in modem their comparison with and South America. species of Central The fossil record of limited to a report from the Lower Miocene palynoflora of Cameroon as- Trichilia is named cribed to this genus (Salard-Cheboldaeff 1978, pollen as Psilastephanocolporites [sic!] grandis n. sp.). In DNA a paper by Muellner and other Meliaceae together with chloroplast data, are the et (2006), this fossils, al. DNA basis for estimates of divergence times in the family. The rbcL cladogram presented by these authors in- cludes a monophyletic clade of Trichilia and 11 other genera, which is assigned an origin in the Oligocene. Apart from Trichilia, members of this clade are presently limited to the Old World tropics, from Africa to Gondwanan Madagascar, India, Indo-China, Malesia, and Austroasia. Muellner et al. (2006) propose a West Cretaceous and between and North America origin for the Meliaceae, with dispersal across Eurasia Eurasia provided by extending from North South America the Beringian and North Atlantic land bridges, thence to via island chains or direct land connections during the Tertiary (cf. Muellner et al. 2010, with a dense sampling of crown com- Cedreleae). The age of the Dominican commensurate with the Oligocene age of the clade fossils is modem prising and Concerning the crown group age of the genus Trichilia relatives (Muellner et 2006). al. Miocene Trichilia, however, the chronograms of Muellner et (2006, 2007) suggest a origin. al. DNA The placement of in subfamily Melioideae (Harms 1940) has been confirmed by the phylo- Trichilia on was genetic research (Muellner et 2003, 2007, 2008). However, the tribe Trichilieae, based this genus, al. resolved as non-monophyletic in the latter study (Muellner et al. 2008). Although Harms (1940) had divided modem Trichilia into 10 sections, only 5 of these comprise the genus in its circumscription, according to Pennington et and these authors go further in reducing the recognized sections to only two: Sect. al. (1981), and new belong emended Trichilia Moschoxylum DC. Within the species to the Sect. Sect. C. Trichilia, Moschoxylum, the defining being the valvate petals and completely united filaments. traits ACKNOWLEDGMENTS We thank Terence Pennington for his useful comments regarding possible taxonomic affinities of T. antiqua. Much help was provided by Alexandra Muellner and an anonymous reviewer, especially in our interpretation of the distributional history and chronological dating of the clade based on and Trichilia allied genera. REFERENCES Chambers, K.L. and G.O. Poinar, Jr. 2010. The Dominican amber fossil Lasiambix (Fabaceae: Caesalpinioideae?) a is Licania (Chrysobalanaceae). Bot. Res. Texas 4:21 7-218. J. Inst. A Chambers, K.L, G.O. Poinar, Jr., and A.E. Brown. 201 fossil flower of Persea (Lauraceae) in Tertiary Dominican amber. 1 . Texas 5:457-462. Bot. Res. Inst. J. Draper, G., P. Mann, and J.F. Lewis. 1 994. Hispaniola. In: S. Donovan and T.A. Jackson, eds. Caribbean geology: an intro- duction. The University of the West Indies Publishers' Association, Kingston, Jamaica. Pp. 129-150. Harms, H. 1940. Meliaceae. In: H. Harms and Mattfeld, eds. Die nattirlichen Pflanzenfamilien 19bl. Verlag Wilhelm J. Engelmann, 1-172. Leipzig, pp. Iturralde-Vinent, M.A. AND R.D.E. MacPhee. 1966. Age and paleogeographic origin of Dominican amber. Science 273:1850-1852. SA Muellner, A.N., R. Samuel, Johnson, M. Cheek, T.D. Pennington, and M.W. Chase. 2003. Molecular phylogenetics of DNA Meliaceae (Sapindales) based on nuclear and plastid sequences. Amer. Bot. 90:471 -480. J. Muellner, A.N., V. Savolainen, R. Samuel, and M.W. Chase. 2006. The mahogany family divergence time *’out-of-Africa’: DNA estimation, global biogeographic patterns inferred from plastid rbcL sequences, extant, and fossil distribution of diversity. Molec. Phylogen. Evol. 40:236-250. Muellner, and A.N., D.D. Vassiuades, S.S. Renner. 2007. Placing Biebersteiniaceae, a herbaceous clade of Sapindales, a in temporal and geographic context. Syst 266:233-252. PI. Evol. Muellner, A.N., R. Samuel, M.W. Chase, A. Coleman, andT.F. Stuessy. 2008. An evaluation of tribes and generic relation- ships in Melioideae (Meliaceae) based on nuclear ITS ribosomal DNA. Taxon 57:98-108. Muellner, A.N.,T.D. Pennington, A.V. Koecke, and S.S. Renner. 2010. Biogeography of Cedrela (Meliaceae, Sapindales) in and Central South America. Amer. Bot 97:51 1-518. J. and A Pennington, monograph T.D., B.T. Styles. 1975. generic of the Meliaceae. Blumea 22:419-540. Pennington, and DA.H. T.D., B.T. Styles, Taylor. 1981. Meliaceae. Neotropica 28:1-470. FI. Poinar, G.O., Jr. and R. Poinar. 1 999. The amber forest. Princeton University Press, Princeton, NJ. Poinar, G.O., Jr., K.L. Chambers, and A.E. Brown. 2008a. Lasiambix dominicensis gen. and sp. nov., a eudicot flower in Dominican amber showing affinities with Fabaceae subfamily Caesalpinioideae. Bot. Res. Inst. Texas 2:463-471. J. rescence in Dominican amber. Bot. Texas 67-1 J. Inst. 2:1 1 1 73. Salard-Cheboldaeff, M. 1978. Sur la palynoflore Maestrichtienne et Tertiaire du basin s^dimentaire littoral du Cameroun. Pollen and Spores 20:21 5-260. Schlee, D. 1999. Das Bernstein-Kabinett. Stuttgarter Naturk. Beitr. Ser. C, 28.

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