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Pollen morphology of the Bignoniaceae from asouth Brazilian Atlantic forestClaudia P. Bove aa Universidade Federal do Rio de Janeiro, Instituto de Biologia. Departamento deBotânica, Laboratório de Palinologia , CEP 21949-900, Rio de Janeiro, BrasilPublished online: 01 Sep 2009.

To cite this article: Claudia P. Bove (1993) Pollen morphology of the Bignoniaceae from a south Brazilian Atlanticforest, Grana, 32:6, 330-337, DOI: 10.1080/00173139309428960

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Grana 32: 330-337, 1993

Pollen morphology of the Bignoniaceae from a south Brazilian Atlantic forest

CLAUDIA P. BOVE

Bove, C. P. 1993. Pollen morphology of the Bignoniaceae from a south Brazilian Atlantic forest. - Grana 32: 330-337. ISSN 0017-3134.

The pollen morphology of 33 species (19 genera) of Bignoniaceae native to the south Brazilian Atlantic forest was examined using light- and scanning electron microscopy. The following pollen types were established: 1. inapenurate (Acletiocn/jrtirtin. Cljiosioitin and Piihecocreriiiori) 2. triapertu- rate (A) psilate (Jacaranda) (B) microreticulate (Arrtrbickreo, Friclericin. Litndin. hlocfo&enn, h!e//o(i. Pnrtigoriia. Sclilegelin and nriiinrii/iirs) ( C ) reticulate (C jb i s i s , Pjrosiegicr and Tabebiria), 3. stepha- nocolpate (~~rti /) / i i lo/)/ i ifrt t i , Aiieriio/Jnegrtin and ~rb(iiInl~~/)/iiiriti), 4. perisyncolpate (htnrisnn).

Clniulio P. Bore, Uriirersitlotle Fetlernl do Hio ile Jnrieiro. lrisiiiitro cle Biologitr. Depcrriairieriio cle Boiriiiicn, hbornidrio ile Pn/iriolo.qin. CEP 21919-900. Rio cle Jiuieiro. Brnsil.

(htmiirscripi received 31 htnrs 1992: revised rersioti occepied 6 hrl j 1993)

The Bignoniaceae are a large pantropical family with few representatives in the temperate zone. It is predominantly neotropical and its greatest diversity occurs in Brazil. Schu- mann (1895) divided the family into five tribes: Bignonieae, Tecomeae, Eccremocarpeae, Crescentieae and Tourrettieae. Much later Gentry (1980) recognized eight tribes, adding Oroxyleae, Coleeae, Schlegelicae. In the south Brazilian Atlantic forest only Tecomeae, Bignonieae and Schelege- lieae occur.

From a palynological perspective, the family is known as heterogeneous. However each genus shows great uniformity. Generic diagnoses based on current taxonomic criteria are notoriously difficult. making the stable pollen pattern a use- ful additional character for identification.

The palynology of the Bignoniaceae was studied at an early stage (e.g., Mohl 1835, Schumann 1895, Urban 1916) and by many subsequent authors: Erdtman (l952), Comes (1955. 1957). Natarajan (1957). Palacios (1966), Mitra (1968), Heusser (1971), Huang (1972), ikuse (I956), Fergu- son & Santisuk (l973), Salgado-Labouriau (l973), Surya- kanta (l973), Buurnian (l977), Markgraf & d'Antoni (1978), Andrade & Miranda (1979), Gentry & Tonib (1979), Carreira & Barth (1987), Silvestre & hlelhern (1989) and Bove & Barth (1992).

This paper describes the pollen morphology of native Bignoniaceae from !he Atlantic forest. Due to the arboreal habit (Cjbistrrx, Jmtcnt-rriida and 7h/ iehi i r t ) and iianescent (Arletiocaljttitii~i, A i t i ~ ? ~ i i ~ ~ ~ ~ ? ~ ~ i f f t t i , Aiietiropnegiiin, Arrnbi- rlrrecr. Cljlostottia Frirlericin, Lrrtich, hfmfndjetin. hfntisorr, AIelloa, Purogotiia, Pirhecoc~etiirrtti. Schlegelin, f i i i t inr i th i is

and UrDatiolophiirtii), the pollen niorphology of the family has value as a paledecological indicator. The characteriza- tion and recognition of the Atlantic forest ecosystem \vill assist palynologists in identifying pollen of fossil and recent. deposition in the Bignoniaceae. Furthermore, the present

paper includes the first pollen descriptions of At~~pltilop/ii~itt~ potiicirlntiriiz and Urbntiolop/iii~tii drrsetiiotiirtii, the latter is the type-species of the genus.

MATERIAL AND METHODS The pollen material from dried specimens u'as supplied by the following herbaria: HBR, R, RFA and GUA.

A total of 33 species (from 71 specimens) belonging to 19 genera has been investigated using light and scanning electron microscopy. The pollen grains were acetolyzed according to Erdtman (1953, except that the maximum temperature in the water bath was 70°C. Slides for light microscopy were mounted in glycerin jelly and sealed with paraffin. hfeasurements of the polar and equatorial axes were made from 25 grains randomly selected. The pollen types and subtypes are based on characters discernible in light microscopy.

The palynological terminology follows Erdtman, 1952, 1969 and Praglowski 8: Punt, 1973, except for stephanocolpate and peri- syncolpate terms (sensu Faegri 8: lversen 1950). A new term macro- reticulate is defined for a coarsely reticulate exine with a largest lumina diameter of inore than 5 pm or exceeding 10% of the largest axis.

A brcrioiiotis: A = apocolpium diameter, E = equatorial diameter; E(PV) = equa- torial diameter in polar view; L = lumina diameter, Lhl = light microscopy, hl = muri, N = nexine thickness, P = polar axis, S = sexine thickness. SEhl = scanning electron microscopy.

GENERAL POLLEN DESCRIPTION

Pollen in monads, medium to large sized (24.0-99.0 x 28.0- 94.5 pn), isopolar or apolar, radially symmetrical, spheroi- dal, subspheroidal or prolate, inaperturate or 3-(4-5)6-7-8- (9-l0)-aperturate. Amb rounded to rounded-triangular. Sex- h e seniitectate to tectate-perforate, thicker than nexine, mac- roreticulate to microreticulate or punctate. Foot layer absent. Nexine uniformly thick (<1.0 pn).

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POLLEN TYPES

The pollen morphology of the Bignoniaceae from a south Brazilian Atlantic forest can be divided segregated into four distinctive pollen types based on apertures, and subtypes based on tecta (Table I).

I . Irinpertirrnte type (Figs. I , 2, 19, Tcble IIA)

A hlacroreticulate subtype Taxa included: Arleiiocnlyiiiiiin rlirseriii, A. riinrgirintririi, Cly- tostoiiin sciirri~~obirliri~i. Pitliecncteiiiirrii clolichoirles and t? eclririntiirii.

Pollen grains apolar, spheroidal, (41 .0-)61.7(-80.0) pm, inaperturate, macroreticulate, heterobrochate, curvimurate, simplibaculate, lumina with granules. Sexine varying in thickness, usually more than four times as thick as nexine.

These species are very similar but may be distinguished by the pollen size, exine structure and sculpturing. The pollen grains of Clytostoiiin sciirripbirliiiii are smaller than the other species of this .type. as are the lumina. The pollcn grains of Pithecocteriiiriii doliclioirles and t? ecliiiintrrrii can be, distinguished by their larger lumina diameter and muri wider than the Arleriocnlyrniiin species.

2. Triopertitrote rype

A. Psilate subtype (Figs. 3, 4, 20, Table IIB) Taxa included: Jocnrniidn riiicmiitho and J. pirberirlo

Pollen grains isopolar, oblate spheroidal to prolate, (34.5-)33.0(-50.5) x (24.8-)38.0(48.0) pn, tricolpate or tricolporoidate with one or more ruptures on the colpus membrane. psilate in LM, in SEM tectate-perforate. Sexine more than two times thicker than nexine, thinning nearer the colpi.

Due to the continuous range of minute variations in shape and size of the pollen grain, and the homogeneity of the other characteristics, the two species cannot be separated by the pollen grains.

B. Microreticulate subtype (Table IlC)

Pollen grains isopolar, suboblate to prolate, (24.0-)10.0(- 63.0) x ( 19.2-)38.0(56.0) pm, tricolporate or tricolporoidate,

microreticulate, simplibaculate. Sexine two or more times thicker than nexine. thinning nearer the colpi. This pollcn subtype is divided into two forms on the basis of the integrity of the colpus membrane.

(n) Colprrite jbr-tir (Figs. 5, 6 )

Taxa included: Arrobidneo cliico, A. coroliiin, A. leircopo- goii. A. sririiyrloides, A. selloi, Liiiidici rritiilirln and Schlegelio I i f l l l 11 y oro.

The apertures consist of colpi with lalongate ora; these species cannot be distinguished by their pollen morphology.

(6) Colporoidnte fr,r-iii (Figs. 7, 8, 21-23)

Taxa included: Fridericin speciosn, hlmfiidyeiin cleritotor, hl. iirollis, hl. iiiigrris-criti, hlelloa qiradriud~*is, Pnrngoriia pyr- nrriirlntn and Tyr,iioiitli~i.~ elegmis.

One or more ruptures on the colpus membrane may occur. The pollen grains of hlocforlyerin can be recognized by their prolate shape. 7jxiimitIiirs (monotypic) has the colpus con- stricted in the equatorial region. The other species are indis- tinct palynologically.

C. Reticulate subtype (Table IID)

Pollen grains isopolar, oblate spheroidal to prolate, (3 I .O-) 43.0(-73.0) x (17.0-)32.0(-55.7) pin, tricolpate or tricol- poroidate, reticulate with lumina diameter smaller towards colpus margin and poles, simplibaculate or duplibaculate. Sexine thicker than nexine (ca. three or four times), thinning nearer the colpi. This pollen subtype is divided in two forms on the basis of the integrity of the colpus membrane.

(o) Colpnte forrii (Figs. 9, IO, 27)

Taxon included: Pyrostegia wiiirstri

duplibaculatc niuri (4-colpate grains also may occur). This fomi has large grains, an entire colpus membrane and

(6) Colporoirlnte foriii (Figs. II-13, 24-26)

Taxn included: C y l i ~ t m oiitisjpliiliticiz, TobeOirin olbrr, T. nwllnrierlne, 7: chr):rotrichn, 1 pirlclierririin and E pnbel - lata.

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332 C. F! Bore

Figs. 1-18. Lhl of Bignoniaceae pollen. AH taken at x500. ( I ) AderiocdjrrirfirrJ riiarginmtrti. (2) A. dirseriii. ( 3 4 ) Jacaranda pitheritla. (3) VE. (4) VP. (5) Arubidrrien cliica. VE. (6 ) Litridin riitiditk7. VE. (7-8) Macfndyeria rrriqitis-cnti. (7) VE. Note the poorly defined endoapertures. (8) VP. (9-10) Fyrostegin reriusfci. (9) VE. Arrow points to duplibaculate muri. (10) VP. Note that the luinina diameter becomes smaller towards the pole. ( I I ) Tobehiria a//7n. VE. (12) T orellmedoe. VE. Note the irregularly ruptured colpus. (13) T: pii/r/ierrirticr. VP. ( 16-15) A r ~ i / ? / i i / ~ / ~ / i i r i r r ~ p i i c i i k t t i t m (14) VP. (15) VE. (16) A. prost~lit/ti. VP. ( 17) U r b a ~ l o / ~ / i i ~ t i ditseriianirni. VP. ( I 8) Afrrnsoci dijjirilis.

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Pollen of Bigrimriincene fioni S. Bm5linri Arkmitic forest 333

Table 11. Morphoriietric clotri of pollen types A = Apocolpium diameter N = Nexine thickness E = Equatorial diameter P = Polar axis E(PV)= Equatorial diameter PIE = Polar axis/equatorial diameter ratio L = Lumina diameter S = Sexine thickness hl. = hlurus width X = mean, S i = standard error, X,,,,-X,,, = range, L(%) = as percentage of grain diameter

A. Inaperturate-macroreticulate type

Taxa Diameter L hl S N L(%)

Acleiioc~iljiiirrio cliiseiiii 68.5 2 0.9 (59.9-80.0) 11.6 I .4 2.9 0.6 16.9 A. niar~qir~ntiiiri 53.7 -e 0.8 (45.4-63.0) 9.2 1.5 2.5 0.4 17.1 CI~IOS~OI I IN s c i i i r i ~ ~ ~ i l ~ r i l r i ~ ~ i 46.920.3 (40.9-54.8) 5. I I .2 2.3 0.6 10.9 Pitlrecocreiiiiini clolichoides 72.k0.7 (63.0-76.9) 17.3 1.9 4.6 0.5 24.0 P. eclriiintiirii 67.4k0.8 (58.0-74.3) 20.4 2.4 4.8 0.5 30.3

B. Triaperturate-psilate type

Taxa P E E(PV) A S N PIE

X+SX (X,,,,,-Xman) Xksn (x,",".-x"ld,.)

Jncnrniidn ~ ~ ~ i c r n ~ i t l ~ n 4 1.720.6 (31.416.4) 45.0203 (40.048.0) 49.9 9.7 I .6 0.7 0.92 J. pitberiilo 43.8k0.5 (36.8-50.6) 30.920.7 (24.8-38.4) 41.0 14.7 I .2 0.5 1.41

C. Triaperturate-microreticulate type

Taxa P E E(PV) A L hl S N PIE L(%)

X-rSR (x",,"-x"lJ X-csn (x">,"-xmn)

A rrn6iclnen cliicn A. cornlirin A. leiicopogori A. snriijcloides A. selloi Friclericin specinsn Lirrl&l rlitidllln

hfhcfnr!verin clerifotn hi. IlIoIlis hi. iirigiiis-coti hfellon qirndriinlvis Pnrngorrin

Schlegelin panifolio Tvrirrniithus elegmis

pyamidoto

27.2k0.4 40.5+0.5 45.7k0.8 51.5+1.5 42.2k0.5 29.3k0.1 32.520.4 47.820.6 44.050.4 50.3k0.6 43.120.8

38.9k0.9 33.k0.5 34.2k0.5

(24.0-32.0) (35.246.4) (39.0-57.6) (40.8-61.0) (37649.6) (24.6-32.0) (29.1-36.8) (40.8-52.8) (39.748.0)

(32.848.8)

(30.446.4)

(43.2-51.4)

(27.9-38.6) (28.8-38.4)

32. I-eO.4 44.220.4 39.6k0.7 49.220.8 49.0k0.4 34.020.4 40.920.7 30.620.9 28.950.7 35.520.5 39.521. I

43.720.7 36.120.7 26.220.5

(28.0-35.2) (40.048.0) (30.441.0) (43.2-56.0) (45 6-54.4) (30.4-37.9) (33.046.4) (23.8-39.2) (22.9-38.4) (30.430.8) (23.218.0)

(36.849.3) (29.412.0) ( 19.2-30.2)

32.4 5.6 0.5 0.3 1.4 0.4 0.85 1.6 44.3 7.3 0.7 0.4 1.7 0.4 0.91 1.6 42.8 7.5 0.7 0.3 1.6 0.4 1.15 1.5 50.0 9.4 0.5 0.5 1.8 0.4 1.10 0.9 48.5 8.6 0.8 0.4 1.7 0.4 0.86 1.6 33.8 8.4 0.5 0.4 0.8 0.4 0.86 1.5 39.8 5.2 0.6 0.6 1.6 0.4 0.79 1.5 43.1 7.4 0.6 0.5 1.1 0.5 1.56 1.2 38.1 6.4 0.7 0.5 1.0 0.6 1.52 1.6 45.2 3.7 0.7 0.4 0.9 0.4 1.42 1.4 39.2 8.8 0.7 0.4 1.4 0.4 1.09 1.6

42.1 6.4 0.6 0.4 1.0 0.5 0.89 1.4 34.5 10.1 0.7 0.6 1.4 0.7 0.94 1.9 33.1 6.0 0.6 0.5 0.8 0.4 1.30 1.7

D. Triaperturate-reticulate type

T3.U P E E(PV) A L hf S H PIE L(%)

X2SR (xm,"-xmJ X+Si (X,,,,.-X,,,J

Cjb i s t~r ciritisjpliiliticcr 3.5.220.6 Pymtegin veiiiistn 65.350.8 Tiibebirin nl6a 36.520.5 7: civellairedne 38.650.6 7: rhnsotricho 3 8.420.4 'I: pirlcherrinin 39.050.6 7: lrl~rbellntn 49.420.8

(30.941.3) 39.6k0.4 (33.443.2) 38.0 3.8 3.9 0.7 2.0 0.5 0.89 9.8 (56.0-73.1) 44.0+1.1 (32.3-54.7) 61.4 3.6 4.2 0.8 1.7 0.5 1.48 6.4 (32.0-12.4) 25.3202 (20.3-28.8) 32.6 4.5 1.5 0.5 1.3 0.3 1.44 4.1 (33.646.9) 29.620.8 (21.6-37.6) 33.0 6.1 1.9 0.4 1.2 0.5 1.30 4.9 (33.132.0) 22.320.6 (17.0-28.8) 30.9 4.4 1.6 0.5 1.1 0.3 1.72 4.2 (33.644.5) 29.620.6 124.0-33.4) 38.7 6.6 1.2 0.6 1.3 0.4 1.31 3.1 (41.656.8) 31.420.6 (23.2-36.0) 47.2 7.0 1.4 0.7 1.4 0.3 1.57 2.8

cont.

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334 C. l? B o w

Table II. (corit.)

E. Stephanocolpate-macropreticulate type ~

Taxa P E E(PV) A L hl S H P/E L(%)

R2SR (xm,"-xmJ XASR (x,","-x"J A I t Iphilc~ph il 1 I t I

po/iicii/[itii/ti 59.620.9 (50.4-68.0) 63.120.8 (56.7-74.3) 70.5 27.5 8.9 2.4 5.1 0.2 0.91 14.1 rlricriiopoegrtin

chnriiberlriyii 61.520.2 (48.5-72.5) 59.521.0 (50.4-66.1) 69.5 31.5 8.3 1.7 2.4 0.6 1.03 13.5 A. prostcifiirii 57.550.8 (48.0-61.8) 61.3k0.9 (52.8-67.2) 75.7 26.6 9.6 1.5 2.8 0.5 0.94 15.7 Urbnriolophitirii

ilii.ye/iin/iiitii 80.921.8 (63.0-99.5) 78.7~1.8 (59.8-94.5) 86.7 51.9 13.1 2.5 6.6 0.9 1.02 16.2

F. Perisyncolpate-reticulate type

Taxon Diameter L h l S N L(%)

X2SR (x,,,,"-X",,J

hlansoa difficilis 50.1+0.8 (44.0-58.4) 3.0 0.9 1.5

This latter form have pollen grains with one or more ruptures on the colpus membrane. The pollen grains of Cy- bistcix nritisjplrilirico can be distinguished by their larger lumina and muri wider than the Ttrbcbirin species.

3. Ste~~hariocol~~ate-rIlncr-cir-eticrrlnte t jpe (Figs. 14-1 7,

Taxa included: ~~rr i / ) l i ; / c ) /? l i i i i l i i pririiciilntiirri. Alierliopaegrrin charriberlrryiii, A. prostmirii and Ur-~ril i[)l[)/?li i i irri cliisciiin-

Pollen grains isopolar, oblate spheroidal to prolate sphe- roidal, (48.0-)65.0(-99.5) x (50.5-)65.5(-91.5) pin, 5-10- colpate, macroreticulate, heterobrochate, curvimurate, sim- plibaculate, lumina with granules. Sexine much more thicker than nexine. Ariiplrilophiiirli pririiciilritiilli pollen is (7-)8-9(-10) - col-

pate, Arielriopoeqrrin chnrliberlriyriii and A. pi-ostntirrri (5-)6 (-7)-colpate and Urbmolopliiirr~r diiseiiinriiirii (7-)8(-10)- colpate. The latter species is distinguished from the others by having the largest pollen grains and the widest reticulum. Arii/?lii/o/?liiiirri pariiciilntirrri differs from Aricrrio~inegrliri clirir~iberlcijriii and A. prostcrtiuli in the thickness of the exine. The former species has a sexinc much thicker (almost dou- ble) than the others.

28-31, Tnb. IIE)

1111111.

4. Perisjricoli~ritr-r-et;ciilcrte tjpe (Figs. IS. 32, Eib. IIF)

Taxon included: hlnrrsori clifjicilis Pollen grains apolar, spheroidal, (44.0-)50.0(-58.5) pm,

perisyncolpate, reticulate, heterobrochate, simplibaculate. The sexine varies in thickness, being usually more than three times as thick as the nexine. The anastoniosed colpi sur-

Grm:,i 32

0.5 6.0

rounding the grain delimit slight pentagonal areas. A single grain has 12 to IS of these areas.

KEY T O GENERA

Based mainly on light microscopy observations of acetolyzed pollen grains of a limited number of species.

I . Pollen grains inaperturate .......................... 2

2. Largest lutnina diameter not reaching 15.0 pm; exine

Largest lumina diameter greater than 15.0 pm; exine about 5.0 piii thick ............................. Pithecocferiitori

3. Triaperturate ..................................... 5 6

4. Largest lumina diameter about 10.0 p m .... AdeIiocnlwirtio Largest luniina diameter about 5.0 pin ........ Cljtosfoirin

............. Jcicarcirih Reticulate or microreticulate ........................ 7

6. Perisyncolpate ............................... Alnrison Stcphanocolpate .................................. 8

7. hlicroreticulate ................................... 9 Reticulate ....................................... 10

8. Largest Iumina diameter about 13.0 pin .... Urb[iIic)lc~iJ/iiiiiIi

Largest luniina diameter about 9.0 p m ............... I I 9. Colporate

Colporoidate ..................................... 12

Colporoidate, mur i simplibaculate ................... 13 I I . Sexine about 5.0 pm thick ............... Ai/ i /J / i i / c~/ ) / i i i i t t i

Sexine about 3.0 p m thick ............... AIieniopoegrtio

Subspheroidal .................................... I4

2.0 p m .................................... C.vbistn.r

2.0 pni .................................... Tribebitin 14. Colpi constricted i n the equatorial region ...... T\ririmithiis

.............. Friilericin, Alelloci. Piimgoriicc or Sclilegelin

Pollen grains aperturate ............................ 3

about 3.0 pin thick ............................... 4

hlore than three apertures ..........................

5. Psilate (tectate-perforate in SEh1)

....................... A rriiDiiliiea or Liiiitlin

10. Colpate; muri duplibaculate .................. Pyosfegici

12. Prolate .................................. Alncfn&e/io

13. Exine about 2.5 pni thick; largest lumina diameter greater than

Exine about 1.5 p i thick; largest lumina diameter not reaching

Colpi Lvithout constriction .............................

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folleri of Bigrioriicrceere from S. Broziliori Atlciritic forest 335

Figs. 19-32. SEhl of Bignoniaceae pollen. ( 19) fithecocteriiirru rlolichoicles. x750. Note the macroreticulate tectum. (20) Jncoroiido pberrrki. ~7.50. (2 1-22) fnro~onio~~roniidnta. (21) TIVO pollen grains in oblique view showing some ruptures along the colpus membrane. x550. (22) High magnification of tectum. Portion of grain including part of an aperture. x2400. (23) ~ v ~ i m i ~ i t / i i i s elegciris. XI 000. Note the equatorial constriction of one colpus. (24) Tobebirici chrysotricho. Slightly oblique equatorial view shoa.ing some ruptures along the C O ~ P U S

membrane. XI OOO(25-26) CjDhar ontisyiliiliticci. XI 000. (25) VP. (26) VE. (27) fjrostegicz reriirsfo. x750. (28, 29) ~ ~ ~ j i / i / i i l ~ ~ / ~ l i ; i f r j i

pci~iicctltitrrrri. (28) VP. x550. (29) High magnification of tectum. Portion of grain including part of the apertures. X I 600. (30. 31) Arierriolioegriin clioriiberlcfyiii (30) VP. ~ 5 5 0 . (3 I ) High magnification of the tectum. Note the lumina with granules. x I 600. ( 3 2 ) MorlSocJ clifficilis. x750.

DISCUSSION AND CONCLUSIONS shape is subspheroidal to prolate. The anib is rounded to rounded-triangular. The apertures are usually three in num- -

The pollen grains are medium to large. The sizes range from 24.0 x 28.0 prn in Arrobidoen chica to 99.0 x 91.5-pm in (Irbarzolopliirriii dirseriiririirrii (extreme values). The grains are radially symmetrical, isopolar or apolar. The general

ber, less often none or many, always equatorially placed with one exception perisyncolpak in hlarisntr tlirficili.y.

Pjrostegio wnirsta was first recorded to bc 1-colpnte by Gornes (1955) and Gentry (1973) or spirapcrlurate (Mitra

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336 C. f? Bore

1968). However Gentry & Tonib (1979) reported i t as 3(-4) colpate, an aperture condition confirmed in the present study. The pollen grains are colpate, colporoidatc or colporate, showing a transformation series, but the boundary between them is sometimes obscure. The colporoidate grains have poorly defined endoapertures, forming irregular patterns. When unacetolyzed and acetolyzed pollen slides were com- pared the ora just represent weak points rather than real endoapertures.

The sexine ornamentation ranges from microreticulate, reticulate and macroreticulate; even the psilate type when observed in detail shows perforations or a reticulate pattern as reported by Silvestre & Melhem (1989: 215). In the macroreticulate type there are granules in the lumina. In the triaperturate type the luniina become smaller nearer the col- pus margin and at the apocolpiuni. The sexine also becomes thinner at the colpus margin. The nexine is always thin and even. The muri are uniform in width, smooth and siniplibac- d a t e with the exception of the grains of Pjrusfegicr reiziistci, which are duplibaculate.

hlitra (1968) pointed out that the presence of both prim- itive and advanced pollen types indicates that the family is polyphyletic. According to the methodology of phylogenetic systematics (Hennig 1966), the polarization of character states of.included taxa is only the first step to a phylogenetic analysis. Monophyly must be identified by shared derived states'(synapomorphies) and the presence of both plesio- morphies and apomorphies does not represent polyphyly. The evolutionary trends presented by Buurman (1977) are fairly supported by the degrees of complexity because con- vergence, parallel evolution or regression cannot be ex- cluded. In order to supplement a phylogenetic revision of the Bignoniaceae, the establishment of closely related groups would be desirable, thus allowing the recognition of plesio- inorphies and synapomorphies.

SPECIMENS EXAMINED (All taxa froni Brazil) Arlenocrrlyinin clirsenii Kraenzlin - R. Reitz 8: R. Klein 9839

(HBR). - A . ninrgimmnr (Cham.) DC. L. B. Smith 9763 (HBR), - L. B. Smith 8: R. Klein 11721 (HBR), Brade 9104 (R).

f t i i i / > / z i / ~ ~ / ~ / i i i r ~ ~ i peiiiicirkrrirur (L.) Kunth. - L. B. Smith & R. Klein 14082 (HBR).

Aiieniopaegnin clrciniberlnyiii (Sinis.) Bur. 8: K. Schuni. - Brade I1353 (R), D. Aralijo 4716 (GUA), E. Rocha et al. 1055 (GUA). D. Araljjo 8: R. Henriques 4886 (GUA), A. Castellanos 24443 (GUA), R. Klein 91 I (HBR). -A. prosrnrrtrti DC. - L. B. Smith 8: R. Klein 7447 (R).

Arrobiclnea chicn Verl. - L. B. Smith 8: R. Klein 12541 (R), L. B. Smith 8: R. Reitz 9648 (R), R. Reitz 8: R. Klein 5785 (HBR). - A . curciliiiri (Jacq.) Sandw. L. B. Smith & R. Klein 11155 (R). - A . lerrcopogori (Cham.) Sandw. - R. Klein 2187 (HBR), B. Lutz 1786 (R), E. Pereira 4250 (RFA). - A. sariiyloides (Chain.) Sandw. - L. B. Smith 8r R. Klein 10680 (R). - A . selloi (Spreng.) Sandw. - L. B. Sniith 8: R. Klein 14099 (R).

C!\iosroiiin scirrripabrrlirni Bur. 8: K. Schuni. - R. Reitz & R. Klein 16670 (HBR).

Cybismn.r cttrrisyphiliricn (hlart,) hlart. - R. Reitz 8: R. Klein 7351

(HBR), P. Occhioni 132 (RFA). A. Sarnpaio 1939 (R), Glaziou 41 15 (R).

Fridericio speciosa hlart. - J. C. Lindeman 8: J. H. Haas 3161 (HBR).

Jncnrmrcln ~iiicrcirrflra Cham. - R. Reitz 8: R. Klein 5690 (HBR), H. P. Veloso 15-11-1913 (R), B. Lutz 1035 (R). - J . prberrrln Cham. - L. B. Smith 8: R. Klein 7378 (R), L. B. Smith, R. Reitz & R. Klein 7956 (R), L. B. Smith 8: Reitz 12532 (R), L. B. Smith & R. Reitz 12442 (R), L. B. Smith 8: R. Reitz 12813 (R), B. A. Lutz 1151 (R).

Lirriclicr rririeltrlo DC. - R. Klein, A. Bresolin & P. Occhioni 7990 (HBR); G. Pabst 8365 & E. Pereira 9176 (R).

Mocfndye/rn tlerifnro K. Schuni. - L. B. Smith & R. Reitz 9729 (R). - M. riiollis (Sonder) Seem. - Glaziou 18377 (R). - rll. rrngrtis- cciri (L.) A. Gentry. - L. B. Smith & R. Reitz 12572 (R).

hlarisoo clifficilis (Cham.) Bur. & K. Schuni. - R. Klein 1354 (HBR), L. B. Smith 8: R. Reitz 12798 (HBR), R. Reitz & R. Klein 14353 (HBR), L. B. Smith 8: R. Klein 13170 (HBR), L. B. Smith 8: R. Reitz 12681 (HBR), L. B. Smith 8: R. Reitz 12917 (HBR), R. Klein 9195 (HBR), R. Klein & A. Bresolin 9396 (R), L. B. Smith 8: R. Klein 14039 (HBR).

hlelloo qrmclrivnhis (Jacq.) A. Gentry - L. B. Smith 8: R. Reitz 12176 (HBR).

forcigonin />yrmirie/nfn (L. C. Rich.) Bur. - R. Klein 7101 (HBR), A. D u c k 2301 (R).

Pirliecocteiiiiriu clolicboicles (Cham.) Bur. ex. K. Schum. - Glaziou 18375 (R) - F! echirmfctrir (Jacq.) Bail]. - L. B. Smith 8: R. Klein 8178 (HBR).

f w x r e g i a wilturn (Ker-Gawl.) hliers. - L. B. Smith & R. Reitz 5939 (R), R. Klein & A. Bressolin 9234 (HBR).

Schlegelin poniflorn (Oerst.) hlonachino. - R. Klein 2399 (HBR). Eibebtrin all7a (Cham.) Sandw. - L. B. Smith 8: R. Reitz 12477 (R). - T. owllnrietlne Lorentz ex Griseb. - R. Reitz 5919 (HBR), R. Reitz 6381 (HBR), G. Hatschbach 16632 (RFA), G. Hatschbach 8: €I. Haas 16588 (RFA). - T. chiyofriclro (hlart. ex DC.) Standley. - R. Reitz 8: R. Klein 7105 (HBR), P. Occhioni 4640 (RFA). - T. ptrlcherrinin Sandw. - L. B. Smith. R. Reitz 8: R. Klein 7.263 (R). - T. piibellnm (Sond.) Sandw. - R. Reitz 8: R. Klein 7152 (HBR), Glaziou 11255 (R).

7jwinrrrhrts elegnris (Cham.) hliers. - R. Klein I051 (HBR). Urbotiolophirtiir clirseriinriirni (Kraenzlin) hlelchior - R. Reitz 8: R.

Klein 18126 (HBR).

ACKNOWLEDGMENTS Special thanks are due to Dr. hlonika Barth for her guidance. I am grateful to the Directors and Curators of the various herbaria for providing polliniferous material and to Dr. Wilson Costa for his constant encouragement and valuable suggestions. The ,manuscript benefited from comments by Drs. hlonika Barth, Wilson Costa, Therezinha hlelhem, Sitvert Nilsson, Jean Ybert and an anonynious reviewer. Finally I would like to express my gratitude for the technical assistance of hlarco Rodrigues and to the Instituto de Biofisica, Universidade Federal do Rio de Janeiro for the access to a scanning electron microscope. Financial support was given by Con- selho Nacional de Pesquisa (CNPq) and Coordenag5o de Aperfei- qoaniento de Pessoal do Ensino Superior (CAPES).

REFERENCES

Andrade, A. P. de 8: hliranda, hl. hl. B. de. 1979. P6len das plantas silvestres do Ceari. IV. Famflia Bignoniaceae. - Rev. Brasil. Farm-60: 155-160.

Bow. C. P. 8: Barth, 0. hi. 1992. Catilogo sistemitico dos pdlens das plaritas arbdreas do Brasil hieridional. XXV. Famflia Bigno- niaceae. - Rev. Brasil. Biol. 52: 283-292.

Dow

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nive

rsity

] at

16:

38 1

5 O

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er 2

014

Polleri .f Bigiioiiimeae frorti S. Bro:ilictri Ailniltic filrest 337

Huang, T. C. 1972. Pollen flora of Taiwan. - Nail. Taitvan Univ.

Ikuse, hl. 1956. Pollen grains of Japan. - Hirokawa Publ., Tot& hlarkgraf, V. 8: d'Antoni, H. L. 1978. Pollcn flora of Argentina.

hlodern spore and pollen types of Preridophyta, Gymnospcrniae and Angiosperm. - Univ. Arizona Press, Tucson.

hlitra, K. 1968. Pollen morphology in Bignoniaceae in relation to taxonomy. - Bull. €301. Sun.. India 10: 319-326.

hlohl, H. 1835. Sur la structure el les formes des grains dc pollen. - Ann. Sci. Nat. 2 ser. 3: 301-346.

Natarajan, A. T. 1957. Studies in the morphology of pollen grains. Tubiflorae. - Phyton (Buenos Aires) 8: 2142 .

Palacios, C. R. 1966. hlorfologia de 10s granos de polen de drboles del Estado de hlorelos. - An. Esc. Nac. Cicnc. Biol. hlex. 16: 4 1 - I 69.

Praglowski, J. 8: Punt, \V. 1973. An elucidation of the niicroretic- da te structure of the exine. - Grana 13: 45-50.

Salgado-Labouriau, hl. L. 1973. ContribuigSo h palinologia do Cer- rado. - Acad. Bras. CiSnc. Ed, Rio de Janeiro.

Schuniann, K. 1895. Bignoniaceae. - In: Die Naturlichen Pnan- zenfamilien 4.3b (ed. A. Engler 8: K. Prantl) pp: 189-252. - W. Engelmann, Leipzig.

Silvestre, hi. S. F. 8: hlelhem, T. S. 1989. Flora polinica (Parque Estadual das Fontes do Ipiranga): 150. Bignoniaceac. - Hoehnea

Suryakanta. 1973. Pollen morphological studies in the Bignonia-

Urban, 1. 1916. Uber Ranken und Pollen der Bignoniaceae. - Ber.

Bot. Dcp. Press, Taipei.

16: 21 1-228.

ceae. - J. Palynol. 9: 15-82.

Drsch. Bot. Ges. 31: 728-758.

Buurman, J. 1977. Pollen morphology of the Bignoniaceae. - Pollen spores 19: 417-519.

Carreira. L. hl. hl. 8: Barth. 0. hl. 1987. hlorfologia polinica de plantas cultivadas no Parque do hluscu Eniflio Goeldi. IV. Fami- lia Bignoniaceae. - Bol. hlus. Par. E. Goeldi Sir. Bot. 3: 79-99.

Erdtman, G. 1952. Pollcn morphology and plant taxonomy. Angios- perms. - Alniquist 8: Wiksell, Stockholm.

Erdtman, G. 1969. Handbook of palynology. hlorphology, taxo- nomy, ecology. - Hafner Publ. Co, New York.

Faegri, K. 8: Iversen, J. 1950. Textbook of modern pollen analysis. - hlunksgaard, Copenhagen.

Ferguson, 1. K. 8: Santisuk, T. 1973. Notes on the pollen morpho- logy of some Asiatic Bignoniaceae. - Kew. Bull. 28: 187-191.

Gentry, A. H. 1973. Flora of Panama. Bignoniaceae. - Ann. hlo. Bot. Gard. 60: 781-977.

Gentry, A. H. 1980. Bignoniaceae. Part I: Cresccntieae 8: Tourre- ticae. Flora Neotropica. hlonogr. 25. - N. Y. Bot. Gard., Bronx.

Gentry, A. H. gL Tomb, A. S . 1979. Taxonomic implications of Bignoniaceae palynology. - Ann. hlo. Bot. Gard. 66: 756-777.

Gomes, J. C. Jr. 1955. Contribuigso sisteinitica das Bignoniaceae brasileiras. - Arq. Serv. Florest. 9: 261-296.

Gornes, J. C. Jr. 1957. Flora do Itatiaia. Bignoniaceae. - Rodrigui- sia 20: I 11-129.

Hennig, \V. 1966. Phylogenctic systematics. - Univ. Illinois Press, Urbana.

Heusser, C. J. 1971. Pollen and spores of Chile. hlodern types of Pteridophyta, Gymnosperinae and Angiosperm. - Univ. Arizona Press, Tucson.

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