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Pollen morphology of the Bignoniaceae from a south Brazilian Atlantic forest

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DOI: 10.1080/00173139309428960

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 Grana

ISSN: 0017-3134 (Print) 1651-2049 (Online) Journal homepage: http://www.tandfonline.com/loi/sgra20

Pollen morphology of the Bignoniaceae from a

south Brazilian Atlantic forest

Claudia P. Bove

To cite this article: Claudia P. Bove (1993) Pollen morphology of the Bignoniaceae from a south
Brazilian Atlantic forest, 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 j b i s i s , Pjrosiegicr and Tabebiria),3. stepha-
Aiieriio/Jnegrtinand ~rb(iiInl~~/)/iiiriti),
nocolpate (~~rti/)/iilo/)/iifrtti, 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.
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(htmiirscripi received 31 htnrs 1992: revised rersioti occepied 6 hrlj 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/iehiirt) and iianescent
(Arletiocaljttitii~i,A i t i ~ ? ~ i i ~ ~ ~ ~Aiietiropnegiiin,
?~~ifftti, Arrnbi-
rlrrecr. Cljlostottia Frirlericin, Lrrtich, hfmfndjetin. hfntisorr,
AIelloa, Purogotiia, Pirhecoc~etiirrtti.Schlegelin, f i i i t i n r i t h i i s
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~
and Urbntiolop/iii~tiidrrsetiiotiirtii, the latter is
potiicirlntiriiz
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).

Gniriit 32
 microreticulate, simplibaculate. Sexine two or more times
POLLEN TYPES
thicker than nexine. thinning nearer the colpi. This pollcn
The pollen morphology of the Bignoniaceae from a south subtype is divided into two forms on the basis of the integrity
Brazilian Atlantic forest can be divided segregated into four of the colpus membrane.
distinctive pollen types based on apertures, and subtypes
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based on tecta (Table I). (n) Colprrite jbr-tir (Figs. 5, 6 )

Taxa included: Arrobidneo cliico, A. coroliiin, A. leircopo-

I . Irinpertirrnte type (Figs. I , 2, 19, Tcble IIA)
goii. A. sririiyrloides, A. selloi, Liiiidici rritiilirln and Schlegelio
A hlacroreticulate subtype I i f l l l yoro.
11

Taxa included: Arleiiocnlyiiiiiin rlirseriii, A. riinrgirintririi, Cly- The apertures consist of colpi with lalongate ora; these
Pitliecncteiiiirrii clolichoirles and t?
tostoiiin sciirri~~obirliri~i. species cannot be distinguished by their pollen morphology.
eclririntiirii.
Pollen grains apolar, spheroidal, (41 .0-)61.7(-80.0) pm, ( 6 ) Colporoidnte fr,r-iii (Figs. 7, 8, 21-23)
inaperturate, macroreticulate, heterobrochate, curvimurate,
Taxa included: Fridericin speciosn, hlmfiidyeiin cleritotor, hl.
simplibaculate, lumina with granules. Sexine varying in
iirollis, hl. iiiigrris-criti,hlelloa qiradriud~*is,
Pnrngoriia pyr-
thickness, usually more than four times as thick as nexine.
nrriirlntn and Tyr,iioiitli~i.~ elegmis.
These species are very similar but may be distinguished by
One or more ruptures on the colpus membrane may occur.
the pollen size, exine structure and sculpturing. The pollen
The pollen grains of hlocforlyerin can be recognized by their
grains of Clytostoiiin sciirripbirliiiii are smaller than the
prolate shape. 7jxiimitIiirs (monotypic) has the colpus con-
other species of this .type. as are the lumina. The pollcn
stricted in the equatorial region. The other species are indis-
grains of Pithecocteriiiriii doliclioirles and t? ecliiiintrrrii can
tinct palynologically.
be, distinguished by their larger lumina diameter and muri
wider than the Arleriocnlyrniiin species.
C. Reticulate subtype (Table IID)
2. Triopertitrote rype 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-
A. Psilate subtype (Figs. 3, 4, 20, Table IIB)
poroidate, reticulate with lumina diameter smaller towards
Taxa included: Jocnrniidn riiicmiitho and J. pirberirlo
colpus margin and poles, simplibaculate or duplibaculate.
Pollen grains isopolar, oblate spheroidal to prolate,
Sexine thicker than nexine (ca. three or four times), thinning
(34.5-)33.0(-50.5) x (24.8-)38.0(48.0) pn, tricolpate or
nearer the colpi. This pollen subtype is divided in two forms
tricolporoidate with one or more ruptures on the colpus
on the basis of the integrity of the colpus membrane.
membrane. psilate in LM, in SEM tectate-perforate. Sexine
more than two times thicker than nexine, thinning nearer the
colpi. (o) Colpnte forrii (Figs. 9, IO, 27)
Due to the continuous range of minute variations in shape Taxon included: Pyrostegia wiiirstri
and size of the pollen grain, and the homogeneity of the other This fomi has large grains, an entire colpus membrane and
characteristics, the two species cannot be separated by the duplibaculatc niuri (4-colpate grains also may occur).
pollen grains.
( 6 ) Colporoirlnteforiii (Figs. II-13, 24-26)
B. Microreticulate subtype (Table IlC)
Taxn included: C y l i ~ t moiitisjpliiliticiz, TobeOirin olbrr, T.
Pollen grains isopolar, suboblate to prolate, (24.0-)10.0(- nwllnrierlne, 7: chr):rotrichn, 1 pirlclierririin and E p n b e l -
63.0)x ( 19.2-)38.0(56.0) pm,tricolporate or tricolporoidate, lata.
Gnrtra 32
 332 C. F! Bore
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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 / ~ / ~ / ipi ri i rcri ~i k t t i t m (14) VP. (15) VE. (16)A. prost~lit/ti.VP. ( 17) U r b a ~ l o / ~ / iditseriianirni.
i~ti VP. ( I 8)
Afrrnsoci dijjirilis.
 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~iljiiirriocliiseiiii 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~O s cI Ii Ii iNr i ~ ~ ~ i l ~ r i l r i ~46.920.3
~i (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
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Taxa P E E(PV) A S N PIE

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

Jncnrniidn ~ ~ ~ i c r n ~ i t4l 1.720.6

~n (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-microreticulatetype
Taxa P E E(PV) A L hl S N PIE L(%)
X-rSR (x",,"-x"lJ X-csn (x">,"-xmn)

A rrn6iclnen cliicn 27.2k0.4 (24.0-32.0) 32. I-eO.4 (28.0-35.2) 32.4 5.6 0.5 0.3 1.4 0.4 0.85 1.6
A. cornlirin 40.5+0.5 (35.246.4) 44.220.4 (40.048.0) 44.3 7.3 0.7 0.4 1.7 0.4 0.91 1.6
A. leiicopogori 45.7k0.8 (39.0-57.6) 39.6k0.7 (30.441.0) 42.8 7.5 0.7 0.3 1.6 0.4 1.15 1.5
A. snriijcloides 51.5+1.5 (40.8-61.0) 49.220.8 (43.2-56.0) 50.0 9.4 0.5 0.5 1.8 0.4 1.10 0.9
A. selloi 42.2k0.5 (37649.6) 49.0k0.4 (45 6-54.4) 48.5 8.6 0.8 0.4 1.7 0.4 0.86 1.6
Friclericin specinsn 29.3k0.1 (24.6-32.0) 34.020.4 (30.4-37.9) 33.8 8.4 0.5 0.4 0.8 0.4 0.86 1.5
Lirrl&l rlitidllln 32.520.4 (29.1-36.8) 40.920.7 (33.046.4) 39.8 5.2 0.6 0.6 1.6 0.4 0.79 1.5
hfhcfnr!verin clerifotn 47.820.6 (40.8-52.8) 30.620.9 (23.8-39.2) 43.1 7.4 0.6 0.5 1.1 0.5 1.56 1.2
hi. IlIoIlis 44.050.4 (39.748.0) 28.950.7 (22.9-38.4) 38.1 6.4 0.7 0.5 1.0 0.6 1.52 1.6
hi. iirigiiis-coti 50.3k0.6 (43.2-51.4) 35.520.5 (30.430.8) 45.2 3.7 0.7 0.4 0.9 0.4 1.42 1.4
hfellon qirndriinlvis 43.120.8 (32.848.8) 39.521. I (23.218.0) 39.2 8.8 0.7 0.4 1.4 0.4 1.09 1.6
Pnrngorrin
pyamidoto 38.9k0.9 (30.446.4) 43.720.7 (36.849.3) 42.1 6.4 0.6 0.4 1.0 0.5 0.89 1.4
Schlegelin panifolio 33.k0.5 (27.9-38.6) 36.120.7 (29.412.0) 34.5 10.1 0.7 0.6 1.4 0.7 0.94 1.9
Tvrirrniithus elegmis 34.2k0.5 (28.8-38.4) 26.220.5 ( 19.2-30.2) 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,"-xmJX+Si (X,,,,.-X,,,J

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

Gnurrr 32
 334 C. l? B o w

Table II. (corit.)

E. Stephanocolpate-macropreticulatetype
~

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

R2SR (xm,"-xmJ XASR (x,","-x"J

A I t Iphilc~phil 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 hl S N L(%)

X2SR (x,,,,"-X",,J
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hlansoa difficilis 50.1+0.8 (44.0-58.4) 3.0 0.9 1.5 0.5 6.0

This latter form have pollen grains with one or more rounding the grain delimit slight pentagonal areas. A single
ruptures on the colpus membrane. The pollen grains of Cy- grain has 12 to IS of these areas.
bistcix nritisjplrilirico can be distinguished by their larger
lumina and muri wider than the Ttrbcbirin species.
KEY T O GENERA
Based mainly on light microscopy observations of acetolyzed pollen
grains of a limited number of species.
tjpe (Figs. 14-1 7,
3. Ste~~hariocol~~ate-rIlncr-cir-eticrrlnte
28-31, Tnb. IIE) I . Pollen grains inaperturate .......................... 2
Taxa included: ~ ~ r r i / ) l i ; / c ) / ? l i i i i lpririiciilntiirri.
ii Alierliopaegrrin Pollen grains aperturate ............................ 3
2. Largest lutnina diameter not reaching 15.0 pm; exine
charriberlrryiii, A. prostmirii and Ur-~rili[)l[)/?liiiirri cliisciiin-
about 3.0 pin thick ............................... 4
1111111. Largest lumina diameter greater than 15.0 pm; exine about 5.0
Pollen grains isopolar, oblate spheroidal to prolate sphe- piii thick ............................. Pithecocferiitori
roidal, (48.0-)65.0(-99.5) x (50.5-)65.5(-91.5) pin, 5-10- 3. Triaperturate ..................................... 5
hlore than three apertures .......................... 6
colpate, macroreticulate, heterobrochate, curvimurate, sim- 4. Largest lumina diameter about 10.0 p m .... AdeIiocnlwirtio
plibaculate, lumina with granules. Sexine much more thicker Largest luniina diameter about 5.0 pin ........ Cljtosfoirin
than nexine. 5. Psilate (tectate-perforate in SEh1) ............. Jcicarcirih
Ariiplrilophiiirlipririiciilritiilli pollen is (7-)8-9(-10) - col- Reticulate or microreticulate ........................ 7
pate, Arielriopoeqrrin chnrliberlriyriii and A. pi-ostntirrri (5-)6 6. Perisyncolpate ............................... Alnrison
Stcphanocolpate .................................. 8
(-7)-colpate and Urbmolopliiirr~rdiiseiiinriiirii (7-)8(-10)- 7. hlicroreticulate ................................... 9
colpate. The latter species is distinguished from the others by Reticulate ....................................... 10
having the largest pollen grains and the widest reticulum. 8. Largest Iumina diameter about 13.0 pin .... Urb[iIic)lc~iJ/iiiiiIi
Arii/?lii/o/?liiiirripariiciilntirrri differs from Aricrrio~inegrliri Largest luniina diameter about 9.0 p m ............... I I
9. Colporate ....................... A rriiDiiliiea or Liiiitlin
clirir~iberlcijriiiand A. prostcrtiuli in the thickness of the exine.
Colporoidate ..................................... 12
The former species has a sexinc much thicker (almost dou- 10. Colpate; muri duplibaculate .................. Pyosfegici
ble) than the others. Colporoidate, muri simplibaculate ................... 13
I I . Sexine about 5.0 pm thick ............... A i / i / J / i i / c ~ / ) / i i i i t t i
Sexine about 3.0 p m thick ............... AIieniopoegrtio
12. Prolate .................................. Alncfn&e/io
4. Perisjricoli~ritr-r-et;ciilcrte tjpe (Figs. IS. 32, Eib. IIF) Subspheroidal .................................... I4
13. Exine about 2.5 pni thick; largest lumina diameter greater than
Taxon included: hlnrrsori clifjicilis 2.0 p m .................................... C.vbistn.r
Pollen grains apolar, spheroidal, (44.0-)50.0(-58.5)pm, Exine about 1.5 p i thick; largest lumina diameter not reaching
pni ....................................
perisyncolpate, reticulate, heterobrochate, simplibaculate. 14. 2.0 Tribebitin
Colpi constricted i n the equatorial region ...... T\ririmithiis
The sexine varies in thickness, being usually more than three Colpi Lvithout constriction .............................
times as thick as the nexine. The anastoniosed colpi sur- .............. Friilericin, Alelloci. Piimgoriicc or Sclilegelin
Grm:,i 32
 folleri of Bigrioriicrceerefrom S. Broziliori Atlciritic forest
335
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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 /elegciris.
iiis X I000. 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. XIOOO(25-26) CjDhar ontisyiliiliticci. XI000. (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)
. I ) High magnification of the tectum. Note the lumina with granules. x I 600. ( 3 2 ) MorlSocJ
Arierriolioegriin clioriiberlcfyiii (30) VP. ~ 5 5 0(3
clifficilis. x750.

shape is subspheroidal to prolate. The anib is rounded to

DISCUSSION AND CONCLUSIONS
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
-
rounded-triangular. The apertures are usually three in num-
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
Grurirr 32
 336 C. f? Bore
1968). However Gentry & Tonib (1979) reported it 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-
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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 / ~ ~ / ~ / ipeiiiicirkrrirur
iir~~i (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).
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