SMITHSONIAN CONTRIBUTIONS TO 

BOTANY 

NUMBER 2 

Harold Robinson A Monograph on 

Foliar Anatomy of 

the Genera Connellia, 

Cottendorfia, and Navia 

(Bromeliaceae) 

SMITHSONIAN INSTITUTION PRESS 

CITY OF WASHINGTON 

1 969 
ABSTRACT 

Harold Robinson. A Monograph on Foliar Anatomy of the Genera Connellia, Cottendorfia, and Navia (Bromeliaceae) . Snithsonian Contributions to Botany, 2 : 1-41. 

1969.-The 102 presently known species of the pitcairnioid genera Connellia, Cottendorfia, and Navia are discussed, illustrated, and keyed on the basis of cross sections 

and epidermal peels of leaves. Comparison with other genera of the bromeliad subfamily Pitcairnioideae indicates Connellia, Cottendorjia, and Navia belong in a series 

with Ayensua, Brocchinia, Fosterella, Pitcairnia, and Puya that have sharply demarcated chlorenchyma and water-storage layers in the mesophyll. Abromeitiella, 

Deuterocohnia, Dyckia, Encholirium, and Hechtia seem to be another related series 

without such sharp demarcation. Connellia is considered related to a series of Cottendorfia species having elongate stomata. Many Cottendorjia species show abaxial waterstorage tissue of a structure similar to Puya and some Amazonian species of Pitcairnia. 

In Navia, the species with paniculate or racemose inflorescences are found to represent 

three rather distinct groups, one of which seems close to Cottendorfia. Species of Navia 

with vascular bundles more completely enclosed in chlorenchyma, with scales replaced 

by trichomes, with sessile or glomerate inflorescences, or with connate and partially 

reduced sepals are considered specialized. A great number of substomatal variations 

are recorded in Navia that distinguish species or groups of species. Navia lopezii is 

shown to have a very modified epidermal structure with simple stomata and no scales, 

characters unlike any other bromeliad presently known. 

Oficial publication date is handstamped in a limited number of initial copies and is recorded 

in the Institution’s annual report, Smithsonian Year. 

UNITED STATES GOVERNMENT PRINTING OFFICE 

WASHINGTON : 1969 

For sale by the Superintendent of Documents, U.S. Government Printing Omce 

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Harold Robinson A Monograph on 

The pitcairnioid genera, Connellia, Cottendorfia, and 

Navia, are restricted almost entirely to the remote 

and relatively inaccesible part of South America known 

as the Guayana Highland. While one species of Cottendorfia and two species of Navia were described as 

early as 1830 and a few additional species from 1846 

to 1929, the vast majority have been described during 

the years since 1930, mostly by Lyman B. Smith working with collections of Tate, Steyermark, Maguire, 

and others. The most complete treatments for the 

genera may be found in the recent summary of the 

Bromeliaceae of the Guayana Highland (Smith, 

1967a) . About eight species have been described since 

the 1967 work. 

As presently accepted, the three genera, Connellia, 

Cottendorfia, and Navia, are distinguished as broad 

but apparently natural groups on the basis of floral 

and seed characters. However, rather marked characters have been observed within genera, such as the 

lack of a scape in Cottendorfia navioides (in a genus 

where it is normally well developed) , and the presence 

of a long scape in various species of Navia (a genus in 

which it is usually lacking). In the single species, 

Navia parvula, the anterior sepal is totally aborted, a 

character that might be considered very significant in 

other groups of plants. One species, Cottendorfia 

porida, is unique in its distribution, being found in 

eastern Brazil, far from the Guayana Highlands and 

all other species of the group. 

One of the primary concerns of this study was a 

Harold Robinson, Department of Botany, Smithsonian Institution, Washington, D.C. 20560. 

Foliar Anatomv of 

the Genera Connellia, 

Cottendorfia, and Navia 

(Bromeliaceae) 

search for anatomical characters that might support 

or refute suspected relationships derived from floral 

characters alone. In the process any evidence of natural or related groups has been sought to decrease 

the reliance on the artificial though workable key 

characters that are now prevalent. It is hoped that 

the series of keys and notes produced in the process 

will be useful in the determination of sterile or otherwise incomplete material. To these ends, Dr. Lyman 

B. Smith of the Smithsonian Institution staff, with 

his continuing interest, has provided time, material, 

and constant encouragement. Dr. Edward S. Ayensu, 

also of the Smithsonian, who has concerned himself 

increasingly with the anatomy of the Bromeliaceae 

has likewise provided much needed assistance. 

A general anatomical treatment of the family Bromeliaceae is available in the work of Solereder and 

Meyer (1929); however, the work does not include 

any of the species of Connellia, Cottendorfia, or Navia. 

I have seen, in manuscript form, some parts of the 

treatment of the family by P. B. Tomlinson for the 

series on the Anatomy of the Monocotyledons edited 

by C. R. Metcalfe, Oxford University Press. This latter work does include a few of the species in the above 

complex. 

Methods 

Many characters are available in studies of bromeliad 

leaf anatomy, but the present effort emphasizes epidermal and stomata1 structure along with the leaf 

structures seen in cross section. Scales and trichomes 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

are used to distinguish some species, but no critical 

review of variations in scale structure has been 

attempted. 

The leaf sections and epidermal preparations for 

this study were all made free hand and mounted in 

Hoyer’s Solution (Anderson, 1954) . All specimens 

were obtained from fragments of dried herbarium material soaked in tap water, The final slides used in the 

study were unstained temporary mounts. Preparations 

kept more than two years clearly show the original 

limits of the chlorenchyma and the resinous deposits. 

In taking epidermal strips and leaf sections, the 

leaf base has been strictly avoided. In the leaf base 

the chlorenchymatous mesophyll is poorly differentiated and air chambers are highly developed. The vascular bundles are surrounded with irregular groupings 

of small parenchyma cells, and the epidermal layers are 

greatly thickened. It is at a higher level that stomata 

begin to appear and finally are densely organized into 

definite bands underlying the smaller bundles or intervening mesophyll regions. In some species internal 

features at this higher level may still show some disorganization characteristic of the leaf base, and for 

critical observations only those sections from midleaf 

or above have been relied on. 

The present study constitutes a preliminary effort 

with definite limitations. Even in the case of Cottendorfia, where sections of leaves have often been taken 

at various levels, conclusions must be drawn with care. 

Rarely are more than two levels sampled or more than 

one leaf, and few of the species are even known from 

more than one collection. In Navia, with its far greater 

number of species, observations are based on far less 

thorough sectioning, and the stomata1 characters, 

which happen to be of particular interest in the genus, 

are emphasized. 

For a list of specimens examined during this study, 

see page 19. 

Generic Limitations 

Of the three genera, Navia, Cottendorfia, and Connellia, the first, Navia, is distinguished almost entirely 

on the basis of the unappendaged seeds. Connellia is 

close to Cottendorfia but distinguished by the subbasifixed anthers and showier flowers. 

This study has involved only the most cursory examination of flower parts, but what has been seen tends 

to indicate uniformity of these structures in such genera 

as Pitcairnia, Brocchinia, Cottendorfia, and Connellia. 

Only the last, a small genus of four species, shows such 

uniformity in leaf anatomy. All the other genera listed 

show two or more distinct types of leaf structure. Some 

of the types that appear in more than one genus might 

indicate relationships, but some are probably the result 

of parallel development. There is not, at present, any 

reason to question that present genera represent natural groups, but there is considerable reason to regard 

them as very broad concepts that may be subject to 

some subdivision. 

After observing transverse sections of the leaves of 

over twenty species selected by Dr. Lyman B. Smith 

to represent the variation in the genus Pitcairnia, and 

of nearly all species of Brocchinia, these genera seem 

relatively uniform with only two very distinct types in 

Pitcairnia and three types in Brocchinia. This is contrary to the impression that is given by the macroscopic leaf variation in Pitcairnia where the basic anatomical unity is obscured by broad bladed and 

slenderly petiolate, narrow bladed and deciduous, and 

heterophyllous variants. The variation in transverse 

sections in Cottendorfia seems at least as great, although there are certain types totally lacking, reflecting a basic uniformity in the genus. The genus Navia 

is, from all indications, the most richly varied in almost all structures, patterns of leaf cross section, stomata, and scales. One might conclude that although 

restricted to a climatic and topographic area lacking 

any great antiquity, Navia may represent one of the 

oldest lines of development in the subfamily. 

The leaf anatomy does not clearly demarcate the 

existing generic concepts of the genera Connellia, Cottendorfia, and Navia, but: it does show distinctive 

trends within the latter two genera. These trends are 

discussed in the following section but they are summarized as follows. 

Characters sometimes found in Navia, not in Connellia or Cottendorfia: (1) Vascular bundles separated from adaxial water-storage cells by a layer of 

chlorenchyma; (2) substomatal pores with large or 

long intrusions or reduced to a slit; (3) leaves bearing 

radially symmetrical uniseriate trichomes. 

Character sometimes found in Cottendorfia, not in 

Connellia or Navia: Vascular bundles in contact with 

well-developed areas of water-storage tissue both abaxially and adaxially. 
NUMBER 2 3 

Key to Basic Distinctions of the Pitcairnioid Genera (adapted from Smith, 196713) 

1. Epigynous tube absent. 

2. Plants monoecious with perfect flowers or if rarely polygamodioecious (Dyckb maritima 

and D. selloa) then petals yellow or orange and plants native of southernmost Brazil and 

Uruguay. 

3. Bases of the filaments separate from each other, but sometimes individually adnate to 

the petals and sepals. 

4. Seeds appendaged. 

5. Petals not tightly coiled after anthesis. 

6. Ovary wholly superior; petals regular. 

7. Petals naked. 

8. Seeds with a falciform or broad apical appendage; plants of northeastern 

Brazil ......................... Encholirium 

8. Seeds bicaudate-appendaged. 

9. Anthers basifixed, linear, coiled at anthesis; inner filaments adnate to base 

of petals; leaf blades thin, more or less contracted at base; mesophytic 

plants of Mexico to Argentina .............. Fosterel& 

9. Anthers subbasifixed to equitant, stout, straight; filaments usually free; 

leaf blades firm, not contracted at base; plants of the Guayana Highland 

and northeastern Brazil. 

10. Anthers subbasifixed; petals brightly colored, more or less massed 

together after anthesis but not twisted; sepals large and firm . Connsllia 

10. Anthers equitant; petals white, separate after anthesis; sepals not over 

7 mm long, thin, flat ............... Cottadorfia 

7. Petals appendaged with a single basal scale each; plants of the southern Andes 

from Peru to Chile and Argentina. 

11. Scape evident; inflorescence usually well developed; plants ring-forming 

so far as known ................... Deuterocohnia 

1 1. Scape lacking ; inflorescence one-few-flowered; plants polster-forming 

........................... Abromsitielb 

6. Ovary partially to wholly inferior, or if superior then the petals zygomorphic 

(Pitcairnia mirabilis). 

12. Petals large, naked or appendaged, usually zygomorphic and forming a hood 

over the anthers; sepals convolute with the left side of each overlapping the 

right of the next one; plants of Mexico and the West Indies to Argentina and 

Brazil ........................... Pitcairnia 

12. Petals minute, regular; sepals cochlear with both posterior ones overlapping 

the anterior; plants of the Guayana Highland ........ Brocchiniu 

5. Petals tightly coiled together after anthesis; Andean plants of open summits from 

Costa Rica and Venezuela to Chile and Argentina ............ Puya 

13. Sepals convolute; petals zygomorphic (Pitcairnia aphelandrifolia) . , Pitcairnia 

13. Sepals cochlear; petals regular, naked; ovary superior (in most species) to nearly 

inferior; plants of the Guayana Highland ............... Navia 

3. Bases of the filaments forming a tube and adnate to the petals; petals yellow or orange; 

sepals cochlear; plants of eastern South America .............. Dyckia 

2. Plants dioecious with functionally unisexual flowers; petals rose or white; plants of Texas, 

Mexico, and northern Central America .................. H6chtia 

1. Epigynous tube present; plants of the Guayana Highland ........... Ayacsuo 

4. Seeds naked at maturity. 

In addition to these general trends there are many 

individual characters by which species can be distinguished, as indicated in the keys to the species of 

each genus. There are many species, however, that 

possess an intermediate type of leaf anatomy that cannot be placed in the proper genus without referring to 

floral characters. 

Anatomical Structures 

ZONATION.-AS seen in transverse section, all the species of Navk Cottendorfia, and Connek except the 

very aberrant N. lopezii, have the chlorenchymatous 

mesophyll and the adaxial water-storage tissue sharply 

demarcated from each other (Figures 1, 4, 9, 12, 15, 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

the Pitcairnioideae in the recently described genus, 

Ayensua (Ayensu, 1969; Smith, 1969), in at least two 

species of Fosterella, and in Brocchinia uestita, B. 

maguirei, B. delicata, B. melanacra, and B. cowanii. In 

Brocchinia the covered bundles are highly characteristic of the particular series, but the probable primitive 

type, B. serrata, and the series including the type, B. 

paniculata, have bundles not covered with chlorenchyma adaxially. 

In what I would call the Cottendorfia-type cross 

section, the chlorenchyma may form large masses extending upward between the vascular bundles, but 

with rare exceptions there is a group of water-storage 

cells reaching the adaxial surface of each bundle 

(Figures 1, 4, 9, 15, 61, 64). This pattern is found 

in all species of Cottendorfia and Connellia. In Cottendorfia the appearance can become greatly accentuated with the development of large abaxial waterstorage canals that also reach the vascular bundles 

(Figure 43). This extreme type of Cottendorfia cross 

section seems unlikely to have evolved very often, if 

more than once, and I find it significant that similar 

sections occur in Puya as represented by P. santosii 

and P. sodiroana, and in a small section of rather distinctive Pitcairnia species as represented by P. nuda, 

P. bulbosa, and P. armata. The latter three Pitcairnia 

species are from the part of the extensive range of their 

genus in which Cottendorfia is found. 

The abaxial water-storage canals occur in a part 

of the leaf in which they are often replaced by a fibrous 

region connecting the vascular bundle to the abaxial 

epidermis. Such extensions of the sclerenchyma occur 

in three species of Cottendorfia-C. florida (Figure 4), 

C. thyrsoidea (Figure 24), and C. minor (Figure 

25)-and in Nauia immersa (Figure 136), N. serrulata 

(Figure 182), and various species of the Nauia reflexa 

group (Figure 79). In these two genera there is no 

indication that any of the species are particularly 

closely related except for those in the Nauia reflexa 

group. In neither of these genera do the fibrous bundle groups extend adaxially to reach or nearly reach 

the epidermis as sometimes happens in Pitcairnia or 

Brocchinia. 

cies of Navia the central portion of the leaf may be 

rather distinctly thickened, but there is none of the 

marked alteration in arrangement or shape of the 

vascular bundles that is so prominent in the costalike central portion of the leaves of many species of 

PLACEMENT OF VASCULAR BUNDLES.-In SOme Spe253, 256, 261, 266). This is in contrast to the genera 

Hechtia, Dyckia, Encholirium, Deuterocohnia, and 

Abromeitiella, which have an area of poorly differentiated cells between, forming a transition zone. In dried 

material this can be very difficult to observe since water-storage cells can collapse and become obliterated 

or compacted into a layer that looks like a demarcation 

line. I would refer to sections lacking a sharp demarcation as a Dyckia type, which in its most typical form is 

very thick-leaved with vascular bundles remote from 

the abaxial surface, and the abaxial surface deeply 

ridged with stomata restricted to the deep furrows. 

There has not been much tendency to relate the South 

American genera of this group with the Mexican and 

northern Central American Hechtia because of their 

very divergent and rather remote distributions. On the 

basis of leaf anatomy, however, these genera together 

seem to form a related series well out of the Nauia, 

Cottendorfia, Connellia relationship. 

which the chlorenchyma surrounds the vascular bundles seems to 'be a variable factor that can differ in 

closely related groups, at least in Navia (Figures 232, 

239, 242, 243, 253, 256, 261, 266). In Pitcairnia, at 

least two species, P. punicea and P. oaxacana, have 

chlorenchyma covering the smaller bundles adaxially, 

while most of the species, including many seemingly 

closely related, have bundles all emergent against or 

into the adaxial water-storage tissue. 

In spite of such variation between related species, 

the overall trends indicate rhe exposure of the vascular 

bundles is of considerable significance. The extreme 

situations are quite contrasting and are not found in 

related groups. Also, in some cases whole groups do 

show uniformity in the character. The bundles are 

covered adaxially with chlorenchyma in most species 

of Nauia or show a tendency toward such cover with 

only the larger bundles being exposed (Figures 92, 

100, 106, 109, 116, 117, 124, 128, 132, 133). I would 

refer to such sections as the Navia type. A marked exception in Nauia is a group of perhaps primitive species 

having a paniculate inflorescence of the N. reflexa 

(Figure 70), N. hohenbergioides (Figure 79), N. 

ramosa (Figure 69) relationship, which all have bundles definitely exposed. This group is so marked that 

the few species that have paniculate inflorescences but 

with bundles covered with chlorenchyma adaxially 

seem only distantly related. 

Such distinctly covered bundles appear elsewhere in 

EXPOSURE OF VASCULAR BuNDLEs.-The extent to 
NUMBER 2 

Pitcairnia. The differentiation of the middle part of 

the leaf in Navia is so slight that there is no reason 

to assume any close relationship to that in Pitcairnia. 

In the latter genus this pseudocostate or what I would 

call the Pitcairnia-type arrangement has been present 

to some extent in most species seen, except those in 

the previously mentioned group including P. nuda, P. 

bulbosa, and P. armata. I have seen nothing I would 

call a Pitcairnia-type section in any other genus, and it 

seems to have evolved completely within the genus 

Pitcairnia. 

The review of the various Pitcairnioideae has left the 

impression that the Cottendorfia-type leaf section, at 

least in its less specialized form, is primitive within the 

group. In Brocchinia, Pitcairnia, and Navia the less 

specialized species all or mostly seem to show such 

sections, and the Navia-type section seems to be only 

in ubviously derived groups of species. 

STOMATA.-The guard cells and subsidiary cells, except for those of the very distinctive Navia lopezii, 

are relatively uniform. They vary in relative width in 

both Nauia and Cottendorfia. In Cottendorfia the 

genus seems divisible into natural groups on this basis 

(Figures 5, 7, 10, 13, 16, 19, vs. 22, 27, 29, 31, 34, 

37, 41, 44, 46). No such pattern is evident in Nauia 

though obviously related species tend to be alike. 

In Navia loperii the guard cells are very narrow 

and have similarly shaped lateral subsidiary cells that 

are broadly exposed on the outer surface (Figures 275, 

277). This pattern of narrow guard cells and exposed 

subsidiary cells is not seen elsewhere in Navia, but 

resembles the amazingly uniform stomatal type found 

in most species of Pitcairnia. In Pitcairnia, only three 

species observed, P. nuda, P. bulbosa, and P. armata, 

which have other cottendofioid characters, tend to 

have broader guard cells and more included subsidiary 

cells. Variation between stomata of the Navia lopezii 

type with only the two lateral subsidiary cells (paracytic) and those such as seen in other bromeliads with 

both lateral and polar subsidiary cells (tetracytic) has 

been noted by Stebbins and Khush (1961) in various 

families, but not previously between species of a single 

genus. 

It is notable that the developmental influence of the 

stomata of most bromeliads extends laterally to the 

cells of the adjacent rows to the same extent it does 

to the polar cells, usually resulting in a true total of six 

subsidiary cells. In some cases the true inner lateral 

subsidiary cells seem to become completely included 

5 

(Figures 215, 216, 218, 219), and the outer laterals 

directly contact the guard cells becoming subsidiary 

cells in the strictest sense of the term. 

SUBSTOMATAL INmusIoNs.-The pore immediately 

inside the stoma is usually rounded or only slightly misshapen in most genera of the Pitcairnioideae. It has 

slight intrusions in some Hechtia or Pitcairnia species 

and is rather reduced in Cottendorfia florida (Figure 

6). It is in the genus Navia, however, that the pore is 

subject to incredibly rich variation in shape. One 

totally unique variant is the special case of N. lopezii 

in which the pore does not occur because an inner 

layer of subepidermal cells is not formed (Figure 275). 

Other variations in Navia concern the types of intrusions projecting inward from surrounding cells. 

These variations include N. breweri (Figure 236) and 

N. involucrata (Figure 108) types in which the surrounding cells are thick-walled and intrude leaving 

only a small slitlike opening or no opening that can be 

seen from face view. In the N. reflexa type (Figures 

72, 75) hooked thick-walled intrusions reduce the 

opening to a double-anchor shape, The N. subpetiolata type (Figure 204) shows similar but thin-walled 

projections. The N. cretacea type (Figure 161) is similar to that of N. subpetiolata but bears many extra 

lobes or papillae. The N. parvula type (Figure 102) 

consists of four separate instrusions that reduce the 

opening to a cruciate form. The N. abysmophila type 

(Figure 153) has the opening divided into a number 

of separate round pores. There are in addition a large 

number of species of Navia such as N. myriantha 

(Figure 264) with the simple structure most often 

found in other genera. 

In Navia there are strong indications that stomatal 

types are stable not only in species but species groups. 

The stomatal type in N. reflexa is rather characteristic 

of the group of species, and the Navia subpetiolata 

group seems to be a natural one showing stomatal uniformity. In most of the species related to N. myriantha 

the substomatal pore is simple. The distinctive substomatal pore of N. parvula is correlated with the distinctive aborted sepal in the flower but N. lepidota with 

a reduced sepal has a stomatal type that is as near 

as any seen to that in N. parvula. Still, a case of obvious failure of the character is in the unmodified opening of N. ramosa, a species rather obviously related to 

N. reflexa and its relatives with their otherwise distinctive pore type. The similarity of the substomatal 

pore in N. breweri and N. cardonae might indicate 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

close relationship, but here the character is in conflict 

with the occurrence of trichomes, and in the present 

treatment I have given the latter character priority. 

In many species (Figures 6, 141, 166, 252) I have 

observed a series of stomata in a row in which an 

extra cell is interposed in the substomatal layer that 

forms a bridge across the pore. The fact that this seems 

to occur erratically without regard to species and that 

it appears in a series of six to eight stomata in direct 

line, indicates that it is a common aberration that acts 

in the manner of a small chimera. 

EPIDERMAL CELLS.-On the basis of material examined the epidermal cells of the Pitcairnioideae 

seem to hold amazingly to a normal type with oblong 

shape and sinuous walls. The walls tend to be 

straighter around the stomata and more sinuous in the 

part of each wall nearer the outer surface. There is 

considerable variation in size as can be seen in the 

various figures which, except for Navia lopezii, are 

drawn to the same scale. Among the smallest cells 

are those of N. abysmophila (Figure 152) while the 

largest epidermal cells of normal structure occur in 

Navia intermedia (Figure 134). 

It is remarkable that all marked departures from 

this basic pattern, even though stable within a species, 

are presently each known only from a single species. 

In Cottendorfia fEorida only, the walls of the epidermal cells appear to be straight throughout. In a species outside the immediate study, Brocchinia paniculata, one end of each epidermal cell is much enlarged 

and thick-walled and sinuous walls are mostly lacking. In this species a very distinctive pattern of interlocking fish-shaped cells is formed. Other closely related species of Brocchinia lack this pattern. 

The most distinctive form of epidermal cell is that 

of the unique species, Nauia lope.& (Figures 275-277). 

Here the cells are extremely large, individual cells 

reaching nearly 1 mm in length. These cells are visible 

under a dissecting microscope. Similar large epidermal 

cells are seen in the outer floral bracts and the character is reflected to some extent in the carpel wall in the 

fruit. I have concluded on the basis of the epidermal 

structure that these cells are the result of the failure 

of protoderm cells to subdivide fully before maturing. 

As a result whole layers of tissues fail to differentiate. 

As illustrated in surface view (Stebbins and Khush, 

1961) , members of some genera in other families, 

Juncus (Juncaceae) and Anigosanthus (Haemodoraceae), have similar structure and may have arisen in 

a similar way. A similar process was probably responsible for the peculiar sporophyte development in the 

moss genus Archidium, where there are only eight 

large spores and there is no central columella. The 

evolution of fern sporangia and of leaves of Hymenophyllaceous ferns have also involved a reduction in 

numbers of cells, but these examples involve more 

long term trends than the abrupt development of 

Navia lopezii. 

A central silica body is characteristic of the epidermal cells of most Bromeliaceae as well as some other 

monocot families. As seen in the Pitcairnioideae this is 

generally distinct in such genera as Dyckia, Hechtia, 

and Navia, and it tends to be weaker or lacking in 

species of Cottendorfia, Pitcairnia, and Brocchinia. 

This is only a trend, however, with many exceptions. 

In both Navia lopezii (Figures 276, 277) and N. 

cataractarum (Figure 95) the silica body seems to be 

entirely lacking. 

SaLEs.-The scales are one of the primary characteristics of the family Bromeliaceae, being found 

somewhere on the plant in all but one species. The 

most common type is similar to those seen in Cottendorfia argentea (Figure 40), Navia serrata (Figure 

183), and N. caulescens (Figure 194), platelike with 

elongate cells at least toward the margin. The scales 

may be reduced to a plug as seen in N. garcia-barrigae 

(Figure 89) where it sits in the center of a pit covering 

a circle of stomata. Vestigial scales as shown in Cottadorfia savannensis (Figure 58) are undoubtedly 

common in the family, but only evident under the 

compound microscope. The scale type in Navia arida 

with its thin-walled rounded cells and partially resorbed margin of small cells is unique among the species in the three-genus complex, but somewhat similar 

ones have been seen in Pitcairnia and Brocchinia. 

The most distinctive type of emergence encountered 

in the study is the trichome (Figures 244,262,265,271, 

274) seen only in certain species of Navia. The radially 

symmetrical uniseriate structure seems very distinct 

from all scales seen, but might represent a single 

branch of some of the deeply cleft types occurring in a 

few other species. For the present it seems reasonable 

to treat species so armed as a related complex, but like 

other characters this may vary in expression within the 

part of the genus where it occurs. 

Even in species with leaves that seem glabrous, 

microscopic examination will usually reveal vestigial 

scales with distinct basal plugs set in the leaf. In one 
NUMBER 7. 7 

species, Navia lopezii, the highly modified epidermis 

of the leaf has resulted in the elimination of all traces 

of scales. This species is the one bromeliad now known 

in which there seems to be no trace of scales on any 

part of the plant. 

Comparative Evaluation of Floral Characters 

Some floral characters used in previous keys, such as 

shape of sepals, have been admittedly artificial. Three 

characters used by Smith (1967a) in his treatment 

of the genus Navia, however, are worthy of special 

consideration: (1 ) connate versus free sepals; (2) inferior versus superior ovary; and (3) paniculate versus 

glomerate inflorescence. 

Connate sepals, as seen in the Smith key, are characteristic of a series of Colombian species, including 

Navia heliophila and N. graminifolia, a fact called to 

my attention by Dr. Smith. In spite of some differences 

in general aspect, all these Colombian species show 

similar basic anatomy and three, N. heliophila, N. 

acaulis, and N. bicolor, are unquestionably closely related. The connate sepals prove to be characteristic of 

another group of closely related species, the Navia 

subpetiolata group. Here, the cross section, substomatal 

intrusions, and general aspect all agree and close relationship is unquestionable. Only N. caricifolia of 

this series seems particularly different and it differs 

only by the adaxial exposure of the vascular bundle 

and the somewhat more sinuous walls of the epidermal 

cells. There is no reason now to believe all species of 

Navia with connate sepals are closely related, but I 

have nevertheless grouped them together in my survey 

of the genus. 

The inferior ovary is an important character used to 

distinguish Pitcairnia and Brocchizia from most other 

genera of the Pitcairnioideae. The character vanes 

considerably, however, in the genus Navia. The species 

of Navia having inferior or partially inferior ovaries 

include a diverse lot of which only N. parvula and N. 

lepidota seem particularly closely related. Others include N. aloifolia which has open substomatal pores 

and trichomes, N. arida which has open substomatal 

pores and very distinctive scales, N. immersa with 

closed pores and vascular bundles fused to the abaxial 

epidermal layers, and N. scopulorutn which has closed 

pores and free bundles. Navia fontoides has closed 

pores, but, like only N. parvula and N. lepidota of the 

series, has chlorenchyma completely covering the vas348-752 0-69-2 

cular bundles adaxially. In addition, the degree of 

inferiority of the ovary in the group varies. No basic 

unity is evident between the species involved, and I 

have been inclined to relegate them to various groups 

on the basis of other characters. It is significant, of 

course, that while these species show no close relationship among themselves, neither do they show close 

relationship to any other species in the genus. They 

are an assemblage of some of the most singularly distinctive species in the genus. 

The paniculate inflorescence is certainly primitive in 

the Pitcairnioideae compared to the condensed and 

usually sessile inflorescence of most Navia species. One 

of the basic aims of this study involved the analysis 

of the few Navb species having a paniculate inflorescence. As indicated in the discussion of the chlorenchyma distribution, these species of Navia are divided 

into two probably not too closely related groups. The 

group including N. ramosa, N. repexa, N. hohenbergioides, and N. garcia-barrigae has Cottendorfiatype cross sections and no tendency toward the Navia 

type. This group seems to be a side branch of Navia 

evolution that, but for the unappendaged seeds, would 

be called Cottendorfia. An additional character suggesting closer relationship to Navia is the development 

of marked substomatal instrusions in many but not all 

of the species. In the remaining species of Navia having paniculate inflorescences, the cross sections are of 

the Navia type. This series of three species, N. lindmanioides, N. rupestris, and N. cataractarum, may represent a more primitive group near the main line of 

Navia development. 

Systematic Review 

Floral characters in the following generic descriptions 

are taken partially from Smith ( 1967a). 

Genus Connellia 

Flowers showy; sepals free, much shorter than the 

petals; petals free with a narrow claw, the blades suborbicular, brightly colored, spreading at anthesis, not 

twisted together afterward; stamens included, free, 

the anthers subbasifixed ; ovary superior, glabrous, the 

style slender, elongate; placentae elongate, nearly the 

height of the locule; capsule septicidal; seeds narrow, 

bicaudate. 

Leaves thin with reduced scales (except in C. quel- 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

chii) , margins incurved; stomata distinctly longer than 

wide with narrow guard cells; vascular bundles contiguous with adaxial water-storage tissue, sometimes 

with poorly differentiated abaxial water-storage canals. 

As presently recognized the genus contains four 

species which occur in the eastern Guayana Highlands 

from Ptari-tepui in Venezuela to Roraima. 

The leaf anatomy of the species of Connellia appears quite uniform (Figures 1-3) and seems basically 

indistinguishable from the group of Cottendorfia species having elongate stomata. Among these latter species epidermal scales are generally reduced, abaxial 

water-storage canals are usually shallow and often indistinct, and leaves in some species have distinct cartilaginous margins, all these characters being found 

among the species of Connellia. One species, Cottendorfia longipes, from the group with longer stomata, 

is possibly a Connellia. This species is known only from 

fruiting material, and flowers will be necessary for 

final generic placement. 

Vegetative Key to Species of Connellia 

1. Upper leaf margins with a dense fringe of scales , . . . . , . . . . . . . . C. quelchii 

1. Leaf margins without noticeable scales. 

2. Leaf blade with prominent white cartilaginous margins . . . . . . . . . C. curicifoliu 

2. Leaf blade without differentiated white cartilaginous margins . . C. ungusfae, C. nutuns 

I have not discovered any foliar characters distinguishing C. angustae and C. nutans. The abaxial 

water-storage canals are most highly developed in the 

distal part of the leaves of C. quelchii (Figures 1-3). 

They are most rudimentary in C. caricifolia. In all 

species the vascular bundles are covered by chlorenchyma abaxially in at least the lower part of the leaf. 

Genus Cottendorfia 

Floral bracts minute; flowers small; petals free with 

a narrow claw, the blades elliptical, white, separate 

and not twisted after anthesis; anthers versatile; filaments usually free ; ovary superior, glabrous, the style 

slender; placentae short, subbasal ; capsule septicidal; 

seeds narrow, bicaudate. 

Leaves thin to rather thickened, often with dense 

cover of scales, margins incurved or recurved, sometimes essentially flat; stomata elongate with narrow 

guard cells or wider than long with broad guard cells; 

vascular bundles contiguous with adaxial water-storage 

tissue, sometimes contiguous with abaxial water-storage 

canals. 

As presently recognized the genus contains 24 species, mostly in the eastern Guayana highlands. Cottendorfia thyrsoidea and some of the Lindmania group extend into central and southern Venezuela. 

The majority of Cottendorfia species show the 

abaxial water-storage canals in the leaf. In one large 

group of species these canals are highly developed and 

the vascular bundles very high and narrow giving a 

very characteristic appearance to the leaf cross section 

not seen in Nauia or Connellia (Figure 43). These 

species have short broad stomata (Figures 22, 27, 29, 

31, 34, 37, 41, 44, 46, 48, 50, 52, 54, 56, 59) and often 

have a dense cover of epidermal scales on the leaf surface. Another group of species with less distinct waterstorage canals, more rounded vascular bundles, more 

elongate stomata (Figures 5, 7, 10, 13, 16, 19), without 

obvious scales, and in a few cases bearing a narrow 

pale border, shows a strong relationship to Connellia. 

The two other very divergent elements in the genus 

show, each in its own way, resemblance to Nauia, Cottendorfia florida by the partially closed substomatal 

orifice (Figure 6), and C. nauioides by the lack of a 

scape in the inflorescence. 

Vegetative Key to Species of Cottendorfia 

1. Epidennal cells of leaf with straight walls (Figure 5) ; stomata having inner opening usually 

narrow, partially closed by outgrowth of surrounding cells (Figure 6) . . , . . . C. florida 

1. Epidermal cells of leaf with sinuous walls (Figure 7) ; stomata with inner pore oval or circular, 

fully or almost fully open (Figure 8). 

2. Leafy stems elongate, usu&y much longer than leaves; leaves in cross section with upper 

surface contoured over the massive subdermal cells (Figure 64); leaves often glaucous 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. nuvioider 
NUMBER 2 9 

2. Stems short; leaves in section with upper surface showing contours of epidermal cells only 

3. Stomata elongate, longer than wide (Figures 16, 19) ; leaf surface never covered with 

(Figure 6 1 ) ; leaves not glaucous. 

obvious scales; leaves often with narrow cartilaginous border. 

4. Leaves mostly plane or narrowly revolute. 

4. Leaves strongly involute .................... C. steyermurkii 

5. Leaves with a layer of resinous cells internally, at least adaxially (Figure 12). 

6. Vascular bundles deeply recessed in the chlorenchyma (Figure 9) . C. longipes 

6. Adaxial surface of major vascular bundles even with upper level of chlorenchyma 

(Figure 12) ........................ C. dyckioides 

7. Leaves distinctly toothed near base .............. C. wurduckii 

5. Leaves without layers of resinous cells (Figure 15). 

7. Leaves nearly or completely entire margined. 

8. Most leaves 2 cm or more broad at the base, apex broadly 

acute .......................... C. brachyphylla 

8. Most leaves about 1 cm broad at the base, apex very narrowly 

acute .......................... C. tillandsioides 

3. Stomata short, as wide as long or wider (Figures 22, 27, 29); leaf surface often densely 

covered with scales; leaves without distinct border. 

9. Vascular bundles (as seen in cross section) continuous with abaxial epidermal layers 

or separated by a single row of thinner walled cells (Figures 24, 25). 

10. Leaves mostly over 20 cm long, leaf margins plane or recurved ... C. thyrsoidea 

10. Leaves less than 20 cm long, margins slightly incurved ......... C. minor 

9. Vascular bundles separated from lower epidermis by at least two rows of enlarged or 

thin-walled cells (Figures 33, 36, 39, 43). 

11. Vascular bundles (as seen in cross section) covered by chlorenchyma abaxially 

(Figures 33, 36). 

12. Leaves completely without abaxial water-storage canal (Figure 36) ; upper leaf 

margin with few or no teeth ................ C. subsimplex 

12. Leaves with small abaxial water-storage canals (Figure 33); leaf margins distinctly and closely serrate throughout ....... C. serrulata, C. gracillima 

11. Abaxial water-storage canals reaching the vascular bundles, the bundles in contact 

13. Leaf margins incurved ............ C. guianensis, C. stenophylla 

13. Leaf margins mostly recurved. 

14. Leaves with prominent scales on the adaxial surface ..... C. paludosa 

14. Leaves without distinct scales, adaxially, scales often prominent on abaxial 

surface .......... C. argentea, C. cylindrostachya, C. dendritica, 

C. geniculata, C. mrrguirei, C. nubigena, C. phelpsiae, and C. savannensh 

with water-storage tissue both abaxially and adaxially (Figure 43). 

The genus Cottendorfia can be subdivided into the 

GROUP A : Inflorescence scapose ; epidermal cells 

with straight walls; stomata elongate with narrow 

guard cells ; substomatal orifices partially closed ; 

occurring in northeastern Brazil. 

Subgroup I: One species with leaf margins entire 

or nearly entire; vascular bundles connected to 

abaxial epidermis; sepals cochleate (anterior sepal 

included). 

Cottendorfia fZorida (Figures 4-6), the type species 

of the genus, seems unique in the group studied by the 

straight walls of the epidermal cells. These are often 

difficult to see but can be demonstrated best by viewing 

torn edges of the epidermis. The cochleate sepals and 

following four groups and eight subgroups, 

the partially closed substomatal pores are reminiscent 

of Navia where there are a few species with scapose 

inflorescence and half-buttressed vascular bundles as 

in C. porida. Cochleate sepals and partially buttressed 

vascular bundles are found elsewhere in the genus 

Cottendorfia, but not combined as in the paniculate 

Nuvia species. It is on the basis of the appendaged 

seeds that the present species is most clearly distinct 

from Navia. 

Inflorescence scapose; epidermal cells 

with sinuous walls; stomata elongate with narrow 

guard cells; substomatal orifices oval or rounded; 

occurring in the eastern Guayana Highlands. 

One species with leaf margins involute, not cartilaginous, entire ; vascular bundles 

GROUP B : 

Subgroup 11: 
I0 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

not buttressed abaxially, separated by chlorenchyma from abaxial water-storage canals; sepal imbrication not determined. 

In anatomical structure C. steyermarkii is very similar to C. tillandsioides (Figures 13-15) but the relatively short, incurved, and involute leaves seem very 

distinct. Cottendorfia tillandsioides along with other 

species of the following subgroup seem very consistent 

in their flattened or revolute leaves. 

Five species with leaf margins usually plane or reflexed, often cartilaginous, entire 

or sometimes toothed near the base; vascular 

bundles not buttressed abaxially, separated by 

chlorenchyma from abaxial water-storage canals; 

sepals where determined are convolute. 

The five species share many characters with the 

genus Connellia and close relationship seems likely. 

The generic status of one species, Cottendorfia longipes 

(Figures 7-9), remains uncertain until flowering material is discovered. All species of the group including 

C. longipes, C. dyckioides (Figures 10-12), C. tillandsioides (Figures 13-15), C. brachyphylla (Figures 

16-18), and C. wurdackii (Figures 19-21) have distinct abaxial water-storage canals. 

Inflorescence scapose; epidermal cells 

with sinuous walls; stomata as wide as long or 

wider, with broad guard cells; substomatal orifices 

oval or rounded; occumng in the central and 

eastern Guayana Highlands. 

Two species with leaf margins often 

distinctly but remotely serrate; major vascular 

bundles buttressed abaxially ; sepal arrangement 

not determined. 

Though Cottendorfia thyrsoidea (Figures 22-24) 

and C. minor (Figures 25-28) share the half-buttressed 

vascular bundles, they may not be particularly closely 

related. This condition occurs also in C. florida and in 

a few species of Nauia, species of rather remote relationship. In C. minor the half-buttressed condition 

seems to be present only in the upper half of the leaf. 

In other species showing the character, it seems less 

restricted. 

Two species with leaf margins closely 

and finely serrate; vascular bundles covered by 

chlorenchyma abaxially; abaxial water-storage 

canals present; sepal arrangement not determined. 

There can be no question of the close relationship 

Subgroup I11 : 

GROUP C : 

Subgroup IV: 

Subgroup V: 

of C, serrulata (Figures 29-30) and C. gracillima 

(Figures 31-32). They seem to agree in all structures 

other than size of inflorescence parts. In addition, they 

seem closely related to the members of the following 

two subgroups of Cottendorfia, being distinguished by 

the arrangement of chlorenchyma. and water-storage 

tissue in the abaxial half of the leaf. 

Subgroup VI: One species with leaf margins 

nearly entire; vascular bundles completely enclosed in chlorenchyma abaxially; without abaxial 

water-storage canals; sepal arrangement not 

determined. 

The single species, C. subsimplex (Figures 34-36) , has all the characters of the following subgroup except 

for the dense and continuous chlorenchyma found in 

the abaxial half of the leaf. 

Eleven species with leaf margins 

entire or with teeth restricted to near the base; 

vascular bundles contiguous abaxially with welldeveloped water-storage canals; sepal imbrication where determined usually convolute, rarely 

cochleate. 

The subgroup contains Cottendorfia guianensis (Figures 44-45), type-species of the old genus Lindmania. 

This widely distributed and variable species, along 

with C. argentea (Figures 40-43), C. cylindrostachya 

(Figures 48-49), C. dendritica (Figures 46-47), C. 

geniculata (Figures 50-51 ) , C. maguirei (Figures 52- 

53), C. nubigena (Figures 54-55), C. paludosa (Figures 59-60), C. phelpsiae (Figures 61-63), C. sauannensis (Figures 56-58), and C. stenophylla (Figures 

37-39), have the rather thick leaves with high contiguous abaxial water-storage canals (Figure 43) that 

I have come to think of as most characteristic of Cottendorfia. I have not seen such sections in either Navh 

or Connellia, though they occur in those Puya species I have seen, and in a small group of Pitcairnia species. The species of this subgroup seem very closely related and most of them cannot be distinguished from 

each other by vegetative characters. 

Inflorescence nonscapose, corymbose; 

epidermal cells with sinuous walls; stomata as 

wide as long or wider, with broad guard cells; substomata1 orifices oval or rounded; occurring in 

the eastern Guayana Highlands. 

One species with leaf margins 

entire; vascular bundles somewhat buttressed 

abaxially ; sepal arrangement not determined. 

Subgroup VII: 

GROUP D : 

Subgroup VIII. 
NUMBER 2 11 

Cottendorfia navioides (Figures 64-66) is named for 

the Navia-like short, nonscapose inflorescence, a 

structure not otherwise known in Cottendorfia. The 

species is distinguished from Navia by the presence of 

appendaged seeds. The species seems unique and can 

easily be determined by the contour of the adaxial leaf 

surface as seen in cross section. 

Genus Navia 

Inflorescence of one or more distinct or agglomerated 

heads or in a few species paniculate; flowers large or 

small, sessile or pedicellate; sepals cochlear, the anterior covered by the two posterior or rarely the anterior wholly aborted; petals free with a claw, broad, 

naked, connate in a slender tube, blades spreading at 

anthesis, often cucullate with an inflexed acute apex; 

anthers linear, filaments usually free; ovary usually superior, sometimes partially or wholly inferior; style 

slender; placentae linear; ovules naked or nearly SO; 

capsule septicidal; seeds naked or with a vestigial 

wing, reticulate or corrugated. 

Leaves thin to much thickened, often with dense 

cover of scales or trichomes, margins incurved, recurved or flat, entire to strongly toothed; stomata with 

narrow or broad guard cells, substomatal pore often 

with intrusions from surrounding cells; vascular bundles either exposed to adaxial water-storage tissue or 

partially to completely covered by chlorenchyma, 

abaxial water-storage canals sometimes present but not 

reaching vascular bundles. 

As presently recognized the genus contains 74 species, and occurs throughout the Guayana Highlands. 

Many of the species have vascular bundles covered 

with chlorenchyma abaxially (Figures 92, 100, 106, 

109, 116, 117, 124, 128, 132, 133), a character not 

seen in Cottendorfia or Connellia. The character seems 

somewhat variable in its development, but it is completely lacking from the species that seem most like 

Cottendorfia (Figures 69, 70, 79, 88), and is well developed in almost all species with connate sepals or 

trichomes. The species lacking the character include 

most of those with a paniculate inflorescence. Also common in the genus but not found in Cottendorfia or 

Connellia is a rich variation in the structure of the 

substomatal pore (Figures 72, 75, 102, 108, 153, 161, 

204, 236, 264). The types are itemized in the morphological discussion. Distinctive trichomes (Figures 265, 

271, 274) are found in one series of species, most of 

which have large rounded substomatal pores and vascular bundles covered adaxially by chlorenchyma. I 

have not seen such trichomes in other bromeliad genera. Navia is otherwise notable for a great variety of 

individual species or small species groups that apparently all represent diverse rather unrelated lines of 

development. The most distinctive species, N. lopezii 

(Figures 275-277), could not be assigned to the genus 

or the family on the basis of leaf anatomy alone, having 

an outer surface of extremely large epidermal cells, 

no scales, simple paracytic stomata, and a relatively undifferentiated mesophyll. 

Vegetative Key to Species of Navia 

1. Epidermal cells extremely large, up to 8008 long, without separate subepidermal layers 

(Figures 275-277) ............................ N. lopezii 

1. Epidermal cells small, usually near size of stomata, rarely over 50p long, subepidermal layers 

distinct. 

2. Leaves 50 cm or more long, at least 2 cm and usually 3 cm or more wide near the base 

3. Substomatal openings large and rounded (Figure 68) .......... N. rmosa 

3. Substomatal openings nearly closed by large thick-wall intrusions (Figures 75, 83). 

4. Vascular bundles separatd from the abaxial epidermis by many rows of thin-walled 

(inflorescence paniculate). 

cells (Figure 70). 

5. Adaxial leaf surface bearing many stomata (Figures 71-73) ..... N. repSxa 

5. Adaxial leaf surface with few or no stomata .......... N. heclitiokfes 

4. Vascular bundles connected or nearly connected to abaxial epidermis by a region of 

fibers (Figure 79) , . . N. brocchinioidss, N. diffuse, N. grocilis, N. hohhabcrgioides 

6. Stomata clustered in pits, each cluster protected by a small rather conical scale (Figure 

90) (inflorescence paniculate) ................. N. garcia-burrigac 

6. Stomata not grouped in pits with protective scale. 

2. Leaves distinctly less than 50 cm long or less than 2 cm wide. 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Vegetative Key to Species of Navi4ntinued 

7. Leaves bearing distinct erect stiff trichomes, at least on the abaxial surface (Figures 

265, 271, 274). 

8. Leaves with only small cells abaxial to the vascular bundles (Figures 239, 242, 243) ; 

substomatal openings partially to almost completely closed (Figures 236, 238, 241). 

9. Substomatal openings divided into many small pores (Figure 238) . N. colorata 

9. Substomatal openings not completely divided or at most divided in two (Figures 

236, 241). 

10. Chlorenchyma covering the adaxial surface of the vascular bundles (Figure 

232) .................... N. navicularis, N. breweri 

10. Larger vascular bundles exposed to adaxial water-storage tissue (Figures 242, 

243) ..... N. pauciflora, N. steyermarkii, N. wurdockii, N. xyridiflora 

8. Leaves with large cells forming water-storage canals abaxial to the vascular bundles 

(Figures 253, 256, 261) ; substomatal openings large and rounded (Figures 252, 

255, ~, 258). 

11. Larger vascular bundles exposed to water-storage tissue adaxially (Figure 253) ; 

leaves succulent in appearance ................ N. aloijolia 

11. Chlorenchyma covering adaxial surface of vascular bundles (Figures 256, 261) ; 

leaves usually rather thin. 

12. Leaves with only vestigial marginal teeth. 

13. Lower leaf margins densely pubescent .......... N. lanigeru 

13. Lower leaf margin with only scattered hairs or teeth , , , N. myriantha 

14. Leaf margins strongly reflexed . , . , , , , . , , , , . N. phelpske 

14. Leaf margins plane. 

15. Leaves up to 2 cm wide, with long erect spinose marginal teeth 

.......................... N. stenodonta 

15. Leaves 1 cm or less wide. 

16. Leaves stiff, with prominent tufts of trichomes in axils of all the 

marginal teeth .................. N. trichodonta 

16. Leaves lax, without prominent tufts of trichomes in axils of the 

teeth .................... N. crisps, N. viridis 

17. Substomatal opening cruciate (Figure 102) or 4-lobed (Figure 104) ; anterior 

18. Substomatal opening cruciate (Figure 102); leaves wirh only small scales 

abaxially ; anterior sepal aborted ................ N. parvula 

18. Substomatal opening with 4 broad lobes (Figure 104) ; leaves with large round 

scales abaxially; anterior sepal only reduced ........... N. &pidoh 

17. Substomatal opening not cruciate or 4-lobed; anterior sepal not reduced or aborted. 

19. Substomatal openings single, rather rounded, without obvious intrusions, usually 

20. Leaves with large cells abaxial to the vascular bundles (Figures 193, 218, 222, 

2 1. Abaxial subepidermal cells very thick-walled (Figure 21 7) ; large vascular 

bundles exposed to adaxial water-storage tissue (Figure 218) . . N. mima 

21. Abaxial subepidermal cells with relatively thin walls (Figures 226, 229,231) ; 

vascular bundles covered by chlorenchyma adaxially (Figures 221, 222, 

227). 

22. Leaves entire, narrowed in basal half; epidermal cells thin-walled, without 

silica bodies (Figure 95) ; inflorescence racemose ... N. cataractarum 

22. Leaves at least slightly toothed, not narrowed below; epidermal cells with 

silica bodies (Figures 191, 225, 228) ; inflorescence not racemose. 

23. Abaxial water-storage canals with one layer of very high vertical cells 

(Figure 193) ; leaf bearing prominent asymmetric scales (Figure 194) ; 

inflorescence long scapose , , . . , , , , , . , , . N. caulescens 

23. Abaxial water-storage canals with 2 or 3 layers of equal sized cells 

(Figures 222, 227) ; scales various; inflorescence not long scapose. 

12. Leaves with strongly serrate margins. 

7. Leaves with branching, platelike, or vestigial scales (Figures 58, 183, 184, 194). 

sepal of flower much reduced or aborted. 

large (Figures 186, 189, 192,217,220, 226). 

227). 
NUMBER 2 13 

24. Leaves less than 5 cm long .......... N. schultesiana 

layers (Figure 222) ............ N. graminifolia 

layers (Figure 227) .... N. acaulis, N. bicolor, N. heliophila 

24. Leaves mostly over 5 cm long. 

25. Leaves 1-3 mm wide; abaxial water-storage cells usually in 3 

25. Leaves 7-15 mm wide; abaxial water-storage cells usually in 2 

20. Leaves with only small cells abaxial to the vascular bundles (Figures 99, 187, 

26. Leaves bearing large scales with small rather isodiametric cells (Figure 

184) ; vascular bundles narrowly fused to abaxial epidermis (Figure 187) 

............................. N.arida 

26. Leaf scales vestigial or with elongate cells (Figure 183) ; vascular bundles 

variously shaped. 

27. Vascular bundles broadly fused to abaxial epidermis (Figure 182) 

........................... N.serrulata 

27. Vascular bundles separated from the abaxial epidermis by distinct layers 

of small chlorenchymatous cells (Figures 99, 179, 190). 

28. Adaxial water-storage tissue with 1 or 2 layers of cells (Figure 99) ; 

inflorescence paniculate ............ N. lindmanioides 

28. Adaxial water-storage tissue composed of 3 to many layers of large cells 

(Figures 179, 190) ; inflorescence glomerate, sessile. 

29. Chlorenchyma continuous over adaxial surface of vascular bundles 

(Figure 190) ................... N. saxicola 

29. Larger vascular bundles exposed to adaxial water-storage tissue 

(Figure 179) .................. N. umbratilis 

19. Substomatal openings reduced, multiple, or with obvious intrusions (Figures 94, 

30. Substomatal opening multiple, with many separate scattered pores (Figures 

31. Chlorenchyma covering adaxial surface of vascular bundles (Figure 92) ; 

inflorescence racemose .................. N. rupestris 

31. Vascular bundles mostly exposed to adaxial water-storage tissue (Figures 

151, 156) ; inflorescence not racemose. 

32. Stomata about 20p wide; subepidermal cells mostly 10-15p wide (Figure 

152) ....................... N. abysmophila 

32. Stomata about 25p wide; subepidermal cells mostly 15-2Op wide (Figure 

154) ......................... N. duidae 

30. Substomatal opening not multiple, sometimes divided in the middle or com33. Substomatal opening large though sometimes divided in half, intrusions 

34. Abaxial subepidennal cells and the projections into the substomatal cavity 

thick-walled (Figure 2 14) ............... N. pungens 

34. Abaxial subepidermal cells and the projections into the substomatal cavity 

rather thin-walled (Figures 163, 173, 176, 202). 

35. Projections into the substomatal cavity multifid, bearing many papillae 

36. Adaxial water-storage tissue composed of 3 or more layers of large 

cells, leaves thick (Figure 156) . , , N. brachyphylla, N. glauca 

36. Adaxial water-storage tissue composed of 1 or 2 layers of large cells 

(Figure 157) ; leaves thin, broad and flat. 

37. Leaves narrowed near the base, not chalky below , N. nubicola 

37. Leaves not noticeably narrowed near base, often chalky on abaxial 

surface ............... N. cretacea, N. latijolia 

35. Projections into substomatal cavity usually paired, smooth (Figures 

38. Stomata as broad as long or broader (Figures 169, 172, 175) ; sepals 

190). 

126, 135, 153, 163, 173, 176,214). 

94, 153). 

pletely obscured (Figures 126, 135, 163, 173, 176, 202,214). 

usually thin-walled (Figures 163, 173, 176, 202, 214). 

(Figures 163, 168). 

176, 202, 212). 

separate. 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Vegetative Key to Species of Navia-Continued 

39. Leaves 3-5 mm wide; larger vascular bundles exposed to adaxial 

water-storage tissue (Figure 171) ; stomata about 20p wide; distinct rounded scales present ......... N. semiserrata 

39. Leaves about 1 cm wide; adaxial surface of vascular bundles 

covered by chlorenchyma (Figure 174) ; stomata 25p wide or 

more; scales much dissected or vestigial. 

40. Leaf margins with teeth erect to antrorse ... N. octopoides 

40. Leaf margins with teeth close-set, mostly retrorse . N. caurensb 

38. Stomata longer than broad (Figures 205, 207, 209, 211, 213); 

sepals connate. 

41. Larger vascular bundles exposed to adaxial water-storage tissue 

(Figure 197) ; abaxial epidennal cells with extremely sinuous 

walls (Figure 195) .............. N. caricifolia 

41. Vascular bundles usually covered adaxially by chlorenchyma 

(Figure 200) ; walls of abaxial epidermal cells strongly to laxly 

sinuous (Figures 205,207,209,211,213). 

42. Leaf margins entire or with mostly vestigial teeth . N. ofinis, 

N. connata, N. ghonii, N. mtaguirei, N. subpetiohta 

42. Leaf margins sharply serrate. 

43. Leaf blade very narrow, about 2.5 mm wide, with a short 

broad base .................. N. ocelkata 

43. Leaf blade up to 1 cm or more wide, base not markedly 

wider .................. N. sandwithii 

33. Substomatal opening obscure, or reduced to a narrow slit or dumbbell44. Adaxial water-storage tissue composed of 1 or 2 layers of large cells 

(Figure 150) ................... N. angustifolia 

44. Adaxial water-storage tissue composed of many layers of large cells 

(Figures 139, 156). 

45. Larger vascular bundles exposed to adaxial water-storage tissue (Figures 

133, 136, 139). 

46. Vascular bundles broadly fused to abaxial epidermis (Figure 

46. Vascular bundles separated from lower epidermis by distinct layers 

47. Stomata 40-5Op long, about 30p wide; walls of abaxial epidennal 

cells extremely sinuous (Figure 134) , . , . , , N. intermedia 

47. Stomata 20-3Op long; walls of abaxial epidermal cells strongly 

to weakly sinuous (Figures 140, 142). 

48. Stomata as wide as long, about 30p by 30p (Figure 146). 

48. Stomata longer than wide, about 25p long by 2Op wide (Figures 

shaped pore by thick-walled intrusions (Figures 108, 126, 135). 

136) ....................... N. immersu 

of small chlorenchyma cells (Figure 139). 

..................... N. scopulorm 

49. Leaves less than 5 cm long ......... N. pulvinata 

140, 142). 

49. Leaves over 5 cm long , , , , , . . , , , . N. cucullata 

45. Vascular bundles usually covered adaxially by chlorenchyma (Figures 

50. Substomatal opening completely obscured from face view (Figure 

108) ; leaf with branching scales covering adaxial surface. 

....................... N. involucruta 

50. Substomatal opening distinct as a narrow slit (Figures 121, 123, 

126) ; scales on adaxial surface of leaf sparse or vestigial. 

52. Scales not completely obscuring abaxial leaf surface; leaf 

margins strongly incurved with sharp teeth ; leaf interior without 

obvious resinous cells (Figure 117) ...... N. robinsonii 

128, 132). 

51. Abaxial leaf surface densely covered with branching scales. 
NUMBER 2 15 

52. Scales of abaxial leaf surface forming a dense woolly cover; 

leaves very stiff with erect margins and vestigial teeth; leaf interior with numerous resinous cells (Figures 127, 128, 131, 132). 

53. Leaves broadly lanceolate, with many layers of narrow 

thick-walled resinous cells adaxially (Figure 13 1) 

..................... N. incrasata 

53. Leaves narrowly lanceolate, with 2 or 3 layers of rounded 

thick-walled resinous cells adaxially (Figure 127) 

...................... N.hiantha 

54. Leaves narrow with very large teeth at the base; ovary almost 

wholly inferior ............... N. fontoidss 

54. Leaves without larger teeth at the base, ovary superior. 

55. Leaves broad without broader base; marginal teeth very erect 

and close, about as long as the distance between them. 

56. Abaxial subepidermal cells with very thick walls, lumens 

very small (Figure 11 1 ) ......... N. batbelkta 

56. Abaxial subepidermal cells with rather thin walls (Figure 

113). ................. N.sphdens 

55. Leaves narrow with a very broad base; marginal teeth short, 

57. Lower teeth of leaf margins with prominent tufts of trichomes in axils; interior of leaf with many layers of thickwalled subepidermal cells abaxially (Figure 124) , N. aurea 

57. Lower teeth of leaf margins without tufts of prominent 

trichomes; interior of leaf with only 1 or 2 layers of distinct 

subepidermal cells abaxially (Figure 117) . . N. cardowe 

5 1. Scales of abaxial leaf surface sparse or vestigial. 

shorter than the distance between them. 

The genus Navia can be subdivided into the followGROUP A : Inflorescence paniculate; vascular 

bundles not covered adaxially by chlorenchyma ; 

occurring in the western and central Guayana 

Highlands of Colombia and southern Venezuela. 

One species with substomatal pore 

large, oval or rounded; vascular bundles free from 

abaxial epidermis. 

The single species, N. ramosa (Figures 67-69), can 

be distinguished from Cottendorfia only by the unappendaged seeds. 

Subgroup 11: Six species with the substomatal 

pore reduced to a narrow double-anchor shape 

by pairs of hooked thick-walled intrusions. 

In habit, the six species are like the preceding N. 

ramosa and like many species of Cottendorfia, but 

they can be distinguished by the shape of the substomatal pore. As presently known, N. reflexa (Figures 

70-76) and N. hechtioides (Figures 77-78) have free 

vascular bundles while those of N. hohenbergioides 

(Figures 79-81 ) , N. gracilis (Figures 82-83), N. brocchinioides (Figures 84-85), and N. diffusa (Figures 

86-87) are connected to the lower epidermis by areas 

ing 7 groups and 26 subgroups. 

Subgroup I: 

348-752 0-69-3 

of sclerenchyma. Stomata seem rather common on the 

adaxial leaf surface of N. reflexa where the surrounding cells of the third subepidermal layer show a rather 

distinctive structure (Figure 73). A few adaxial 

stomata have been observed in N. gracilis, but they 

have not been observed in any other bromeliads 

examined. 

One species with stomata clustered in pits around thick pluglike scales; substomata1 pores irregularly rounded to partially 

closed by short thick-walled intrusions. 

The arrangement of stomata in the single species, 

N. garcia-barrigae (Figures 88-91 ) , seems distinctive. 

The single specimen available has only a few separate 

leaves, which may not be representative of the fully 

vegetative parts of the plant. 

Inflorescence racemose to paniculate; 

vascular bundles nearly or completely covered 

adaxially by chlorenchyma. 

Two small species with racemose 

inflorescence ; leaves rather thick ; silica bodies 

small or lacking; occurring in the eastern Guayana 

Highlands in British Guiana. 

Subgroup 111: 

GROUP B : 

Subgroup IV: 
16 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

In N. rupestris (Figures 92-94) the substomatal 

openings are usually divided into three or more small 

rounded pores. In N. cataractarum (Figures 95-96) 

the pores are large and simple. 

One rather large species with large 

panicle bearing racemose branches; leaves thin ; 

silica bodies distinct; occurring in central 

Venezuela. 

Navia lindmanioides (Figures 97-99) has simple 

rounded substomatal pores. The species is quite distinct from others with paniculate or racemose inflorescences, and vegetatively resembles many having 

glomerate inflorescences. 

Inflorescence glomerate ; vascular bundles covered adaxially by chlorcnchyma ; anterior 

sepal reduced or absent; scales platelike or 

branching. 

Subgroup VI: Two species with substomatal pore 

cruciate or 4-lobed; ovary partly inferior; occurring on Cerro Neblina in southern Venezuela. 

Navia parvula (Figures 100-102) shows the extreme 

conditions of the anterior sepal absent and the substomata1 pore cruciate. In N. lepidota (Figures 103- 

106) the anterior sepal is present in reduced form and 

the substomatal pore has only slight intrusions from 

each of the four corners. In both mentioned characters 

the latter species seems transitional, but the species is 

very distinct in other characters such as its large 

rounded scales. 

GROUP D : Inflorescence glomerate ; sepals equal, 

free; scales platelike, branching, or vestigial. 

Subgroup VII : One species with substomatal 

opening completely obscured from face view; 

vascular bundles covered adaxially by chlorenchyma; occurring in the Guayana Highlands of 

central Venezuela. 

The intrusions into the substomatal pore of N. inuolucrata (Figures 107-108, section as in 106) are so 

complex and thick-walled that it is impossible to determine any opening or any pattern. The broad involute 

leaves with a grayish cover of adaxial scales are also 

rather distinctive. 

Subgroup VIII : One species with substomatal 

pore reduced to a short narrow slit; vascular bundles covered adaxially by chlorenchyma ; leaves 

with a few very large teeth near the base; ovary 

Subgroup V: 

GROUP C : 

almost wholly inferior; occurring in the Guayana 

Highlands of Colombia. 

Nauia fontoides (Figures 114-1 16) seems readily 

distinguishable by its narrow leaves bearing large teeth 

near the base. The species also seems rather distinct in 

its inferior ovary and in its geographical distribution. 

Two species with substomatal pore 

reduced to a short narrow slit; vascular bundles 

covered adaxially by chlorenchyma; leaves broad 

and smooth with closely and finely serrulate margins; ovary superior; occurring in the eastern 

Guayana Highlands of British Guiana and adjacent Venezuela. 

Nauia barbellata (Figures 109-1 11 ) and N. splendens (Figures 112-113, section as in 109) are similar 

in most vegetative characters. The material seen of N. 

barbellata has much thicker walled subepidermal cells. 

Five species with substomatal pore 

reduced to a short narrow slit; vascular bundles 

covered adaxially by chlorenchyma; leaves mostly 

with evident scales on abaxial surface; ovary 

superior; occurring in the Jaua-Duida area of 

south-central Venezuela. 

Nauia lnriantha (Figures 125-128) and N. incrnrsata 

(Figures 129-1 32) both have nearly entire-margined 

leaves with a very dense abaxial cover of scales. The 

three remaining species of the group have less prominent scales. Of these three, N. robinsonii (Figures 120- 

121, section as in 117) has broader involute leaves 

while N. cardonae (Figures 117-119) and N. aurea 

(Figures 122-124) both have narrow blades born on 

broad sheathing bases. Navia aurea is quite distinct in 

the many layers of thick-walled convoluted cells 

adjacent to the lower epidermis. 

One species with substomatal pore 

reduced to a pair of small hidden pores; epidermal cells large with very sinuous walls; vascular bundles not all covered adaxially by 

chlorenchyma; ovary superior; occurring on 

Serrania Jaua in south-central Venezuela. 

The epidermal cells of N. intermedia (Figures 133- 

135) are the largest in the genus that are of normal 

structure. The substomatal structure is also rather 

distinctive, but is very closely related to that in the 

immediately preceding and following groups. The 

leaves are very narrow but thick and nearly entiremargined, unlike any others seen in the genus. 

Subgroup IX: 

Subgroup X: 

Subgroup XI: 
NUMBER 2 

Subgroup XI1 : Five species with substomatal pore 

reduced to a short narrow slit; vascular bundles 

not completely covered by chlorenchyma adaxially; occurring in the Guayana Highlands of 

British Guiana and Venezuela. 

Three of the species, N. cucullata (Figures 139-141), 

N. pulvinata (Figures 142-1&), and N. angustifolia 

(Figures 148-150) , have superior ovaries. Two, N. 

scopulorum (Figures 145-147) and N. immersa (Figures 136-138), have partially inferior ovaries. Navia 

immersa is further distinguished by the vascular 

bundles being broadly fused to the lower epidermis. 

The subgroup represents the greatest concentration of 

the leaf type in which the blade is very narrow and 

stiff from a broad rounded sheathing base. Such leaves 

axe also seen in N. cardonae and N. aurea of Subgroup 

X, N. saxicola of Subgroup XVI, and N. graminifolia 

of Subgroup XX. 

Two species with the substomatal 

opening divided into few or many small rounded 

pores; vascular bundles not completely covered 

adaxially by chlorenchyma ; occurring in southern 

Venezuela. 

The two species, N. abysmophila (Figures 151-153) 

and N. duidae (Figures 154-156), have slightly different habits but very similar basic anatomy. The 

epidermal cells are among the smallest seen in the 

genus. 

Five species with substomatal intrusions short but distinct, thin-walled, much 

divided or bearing many papillae; vascular 

bundles usually covered adaxially by chlorenchyma; occurring in the Duida-Neblina area of 

southern Venezuela. 

Subgroup XI11 : 

Subgroup XIV: 

Two of the species, N. brachyphylla (Figures 165- 

166, section as in 156) and N. glnuca (Figures 167-168, 

section as in 156), have rather thick leaves. In N. 

nubicola (Figures 157-159), N. cretncea (Figures 160- 

161, section as in 157), and N. latifolia (Figures 162- 

164, section as in 157) the leaves are very thin. Navia 

cretacea and N. latifolia are obviously closely related 

and cannot be distinguished vegetatively. 

Three species with substomatal 

intrusions rather short but distinct, thin-walled, 

simple ; vascular bundles partially to completely 

covered adaxially by chlorenchyma; occurring in 

Subgroup XV: 

17 

the Rio Caura-Rio Ventuari-Duida area of southcentral Venezuela. 

The subgroup seems unnatural with species differing 

somewhat in both habit and anatomy. Navia semkerrata (Figures 169-171) differs by its stiff leaves and 

vascular bundles not completely covered by chlorenchyma. In N. caurensis (Figures 172-174) and N. 

octopoides (Figures 175-176, section as in 157) the vascular bundles are covered by chlorenchyma adaxially, 

but the former has broad relatively short thick leaves 

and the latter has much longer and thinner leaves. 

Five species with substomatal 

pores rounded or oval, without distinct intrusions; 

occurring in various parts of the Guayana Highlands from British Guiana to Colombia. 

The five species included in the subgroup are sufficiently distinct that each could constitute a separate 

subgroup of its own. Navia umbratilis (Figures 177- 

179), of Cerro Yutaje in south-central Venezula, has 

long slender nearly entire-marginecl leaf blades from 

a rounded sheathing base. The substomatal pores are 

slightly narrowed, and the larger vascular bundles are 

not completely covered adaxially by chlorenchyma. 

The substomatal pores of the other species are not at 

all obstructed. In both N. serrulata (Figures 180-183), 

of southern Venezuela, and N. arida (Figures 184- 

187), of British Guiana, the vascular bundles are not 

completely covered with chlorenchyma. In the former 

species the leaves are narrow from a broad base, but 

are strongly serrulate, and the scalcs have oblong cells. 

In the latter species the leaves are broad and strapshaped, the scales are rounded with very distinctive 

small rounded cells and an eroded margin, and the 

ovary, unlike other species in the subgroup, is partly to 

wholly inferior. In both N. saxicola (Figures 188-190) , of southern Venezuela, and N. caulescens (Figures 

191-194)) of Colombia, the vascular bundles are completely covered adaxially by chlorenchyma. The former 

species has stiff narrow leaf blades from rounded 

sheathing bases, while the latter has broad strap-shaped 

leaves. Nauia caulescens, the type-species of the genus, 

is most distinct by the elongate inflorescence with 

several moniliform heads born on a scape. 

Subgroup XVI : 

GROUP E: 

Subgroup XVII : 

Inflorescence glomerate ; sepals connate ; scales platelike, branching, or vestigial. 

Eight species with substomatal 

pores partly closed by long, paired, thin-walled 

intrusions; vascular bundles covered adaxially by 
18 

chlorenchyma in all but one species; abaxial 

water-storage canals absent ; occurring in the eastern half of the Guayana Highlands from Cerro 

Sipapo and Neblina in Venezuela to Tafelberg in 

Surinam. 

One species, N. caricifolia (Figures 195-197) , is distinct in the vascular bundles not being covered adaxially by chlorenchyma, and in the more sinuous walls 

of the epidermal cells. In other characters such as the 

long strap-shaped usually nearly entire leaves having 

somewhat sheathing bases and the structure of the 

substomatal pore, N. caricifolia is like N. ocellata (Figures 198-200), N. maguirei (Figures 201-202, section 

as in 200) , N. subpetiolata (Figures 203-204, section 

as in 200) , N. gleasonii (Figures 205-206, section as in 

200), N. sandwithii (Figures 207-208, section as in 

200), N. connata (Figures 209-210, section as in 200), 

and N. afinis (Figures 21 1-212, section as in 200). 

Subgroup XVIII : One species with substomatal 

pores partly closed by short, paired, thick-walled 

intrusions ; vascular bundles covered adaxially by 

chlorenchyma, abaxial water-storage canals absent; occurring on Cerro Guanay and Cem 

Yutaje in south-central Venezuela. 

The single species, N. pungens (Figures 2 13-2 15) , is 

distinct among Navia species with a sessile inflorescence 

by the small but distinct thick-walled intrusions. The 

leaves are thick and stiff, and the plant closely resembles N. mima of the following subgroup in habit. 

One species with substomatal 

pore only slightly obstructed ; vascular bundles not 

completely covered adaxially by chlorenchyma ; 

with distinct large cells forming water-storage 

canals abaxially; occurring on Cerro Sipapo in 

south-central Venezuela. 

The one species, N. mima (Figures 216-218), resembles the preceding, N. pungens, in habit, but it has 

the substomatal pores only slightly obstructed by the 

surrounding thin-walled resinous cells. 

Five species with substomatal pore 

rounded, not obstructed ; vascular bundles completely covered adaxially by chlorenchyma; distinct large cells forming abaxial water-storage 

canals; occurring in the western Guayana Highlands in Colombia. 

The five species, N. acaulis (Figures 230-231, section as in 227), N. bicolor (Figures 228-229, section 

Subgroup XIX : 

Subgroup XX: 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

as in 227), N. heliophila (Figures 225-227)) N. graminifolia (Figures 222-224) , and N. schultesiana (Figures 219-221)) all share the same basic anatomy and 

the same rather distinctive geographic distribution. 

Still, the habits of the various species in the subgroup 

are quite diverse. The first three species are much 

alike with long, broad, strap-shaped leaves having no 

distinct central thickening. These three species cannot 

be distinguished vegetatively. Navia graminifolia in 

contrast has the very different slender-leaved form 

found most often in Subgroup XII. Navia schultesiana 

has distinctive small rosettes of serrate stiffly spreading 

leaves, bearing little superficial resemblance to the preceding types. In this study of Navia, most attempts to 

use vegetative habit as an indication of relationship 

have failed. Most subgroups established on the basis of 

anatomy have contained many variations in habit. In 

this subgroup there is some basis to suspect real relationship and therefore potentially good evidence of the 

erratic nature of the vegetative habit. 

GROUP F : Inflorescence glomerate ; sepals equal, 

separate ; scales lacking, large erect trichomes 

present. 

Two species with substomatal 

pores reduced to a short narrow slit; vascular bundles covered adaxially by chlorenchyma; without 

distinct water-storage canals abaxially ; ovary 

superior; occurring in the Jaua-Ichun area of 

south-central Venezuela. 

Subgroup XXI : 

The two species, N. navicularis (Figures 232-234) 

and N. breweri (Figures 235-236, section as in 232), 

seem indistinguishable vegetatively. They and the five 

species in the following two subgroups, and N. viridis 

and N. crispa of Subgroup XXV, show a long, thin, 

strap-shaped leaf often with a distinct central thickening. While this rather distinctive leaf type is most 

common among species with trichomes, it is also found 

in N. octopoides of Subgroup XV and N. lindmanioides of Subgroup V. Some species like N. arida of 

Subgroup XVI and N. nubicola of Subgroup XIV are 

similar but lack the distinct central thickening. 

Subgroup XXII : One species with substomatal 

opening divided into many small usually rounded 

pores; with few or no distinct water-storage cells 

abaxially ; vascular bundles covered adaxially by 

chlorenchyma; ovary superior; occurring in the 

Rio Ventuari area of south-central Venezuela. 
NUMBER 2 

The species, N. colorata (Figures 237-239) , agrees in 

habit and in most anatomical characters with the species of the preceding subgroup, but it differs markedly 

in the thin-walled substomatal cells which form multiple openings. This species is probably most closely related to N. ziiridis and N. crispa of Subgroup XXV. 

Four species with substomatal 

opening obstructed by distinct, long, thin-walled, 

paired intrusions; vascular bundles not completely 

covered adaxially by chlorenchyma; with few distinct water-storage cells abaxially ; ovary superior ; 

occurring in the Chimanta-Marahuaca-Duida 

areas of eastern and south-central Venezuela. 

The four species, N. paucifEora (Figures 247-248, 

section as in 243) , N. steyermarkii (Figura 249-250, 

section as in 243) , N. wurdackii (Figures 243-246)) 

and N. xyridifloru (Figures 240-242)) seem indistinguishable vegetatively. The failure of the chlorenchyma to completely cover the adaxial surface of the 

vascular bundles in this series of species seems to be 

of relatively little significance, since the organization 

of the tissues is similar to that in the related subgroups 

that have covered bundles. 

One species with substomatal 

pore large and unobstructed; vascular bundles 

mostly not covered adaxially by chlorenchyma; 

large cells forming distinct water-storage canals 

abaxially; ovary inferior; occurring on Cem 

Neblina in southern Venezuela. 

The species, N. aloifolia (Figures 251-253), seems 

very distinct in the habit, thick leaves, vascular bundles 

distinctly exposed to the adaxial water-storage tissue, 

and the inferior ovary. The trichomes on the backs of 

the leaves are perhaps the best examples of such structures in the genus. 

Subgroup XXIII : 

Subgroup XXIV : 

19 

Subgroup XXV : Seven species with substomatal 

pores large and unobstructed; vascular bundles 

covered adaxially by chlorenchyma; large cells 

usually forming distinct water-storage canals 

abaxially ; ovary superior; occurring in southcentral and southern Venezuela and adjacent 

Brazil. 

Two species, N. viridis (Figures 254-256) and 

N. crisps (Figures 257-258, section as in 256) , have 

the appearance of the species in Subgroups XXI, 

XXII, and XXIII, but in addition to differences in 

stomata they show a greater development of waterstorage cells abaxially. The remaining five species, 

N. stenodonta (Figures 259-262) , N. myriuntha (Figures 263-266) , N. trichodonta (Figures 267-268, section as in 266), N. lanigera (Figures 269-271), and 

N. phelpsiae (Figures 272-274) , have rather diverse 

leaf types, but all have rather well-developed waterstorage canals abaxially. 

Inflorescence glomerate; sepals equal 

and separate; scales completely absent from leaves. 

One species with broad entiremargined leaves; epidermal cells extremely large, 

subepidermal layers absent ; stomata paracytic ; 

mesophyll not forming distinct layers; occurring in 

the central and western Guayana Highlands of 

southern Venezuela and adjacent Brazil and 

Colombia. 

All the most distinctive features of N. lopetii (Figures 275-277) could be related to the lack of subdivision of initial cells. This is most pronounced in the 

epidermal cells of the leaves and floral bracts, but is 

also somewhat reflected in the structure of the walls 

of the fruits. The modified epidermis has resulted in 

complete elimination of scales, a condition that is in 

exception to the generaIly accepted definition of the 

family. 

GROUP G : 

Subgroup XXVI: 

List of Specimens Examined 

(Specimens used for illustrations are marked with an asterisk.) 

Species Collector and Number Locality Herbarium 

A bromcitiella brevifolia (Griseb . ) Cabrera 16333 Argentina: Prov. Jujuy : Tumbaya US 

Aycnsua mipanensis (Maguire) L. B. Steyermark 93940 Venezuela: Bolivar: Auyain-tepil US 

Brocchinia acuminata L. B. Smith Maguirc 32785 Venezuela: Bolivar: Cerro Guai- US 

Brocchinia cowanii L. B. Smith Maguire, Cowan, & Wurdack 30924 Venezuela: Amazonas: Cerro US holotype 

Castellanos 

Smith 

quinima 

Moriche, Rio Ventuari 
20 

Species 

Brocchinia cryptantha L. B. Smith 

Brocchinia delicatuia L. B. Smith 

Brocchinia hcchtioidcs Mez 

L. B. Smith 

Brocchinia maguirci L. B. Smith 

Brocchinia melanacra L. B. Smith 

Brocchinia micrantha (Baker) Mez 

Brocchinia fianiculata Schult. f. 

Brocchinia prismatica L. B. Smith 

Brocchinia rcducta Baker 

Brocchinia secunda L. B. Smith 

Brocchinia scnata L. B. Smith 

Brocchinia steycrmarkii L. B. Smith 

Brocchinia tatci L. B. Smith 

Brocchinia vcstita L. B. Smith 

Connellia angustae (Rich. & SchomConnellia caricifolia L. B. Smith 

Connellia nutam L. B. Smith 

Connellia quclchii N. E. Brown 

Cottmdorja argentea (L. B. Smith) 

Cottmdorja brachyphylla (L. B. Smith) 

Cottmdorja cylindrostachya (L. B. Smith) 

Cottmdorja dmdritica L. B. Smith 

Cottmdorja dyckioides L. B. Smith 

bock) N. E. Brown 

L. B. Smith 

L. B. Smith 

L. B. Smith 

cottmdorjajorida Schultes f. 

Cottmdorja gmiculata (L. B. Smith) 

Cottcndorja gracillima L. B. Smith 

Cottmdorja guiamis (Beer) Klotzsch 

ex Baker var. vcstita L. B. Smith 

Cottmdorja Iongipes L. B. Smith 

Cottmdorja maguirci L. B. Smith 

Cottmdorja minor (L. B. Smith) L. B. 

Cottmdorja navioides (L. B. Smith) 

Cottmdorja nubigma L. B. Smith 

Cottmdwja paludosa (L. B. Smith) 

L. B. Smith 

Smith 

L. B. Smith 

L. B. Smith 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

List of Specimens Examined-Continued 

Collector and Number Locality Herbarium 

Maguirc, Cowan & Wurdack 30709 Venezuela: Amazonas : Cerro 

Maguire, Wurdack & Maguire 42207 Venezuela: Amazonas: Cerro 

US holotype 

US holotype 

Yapacana, Rio Orinoco 

Maguire & Politi 27603 

Phelps & Hitduock 522 

Maguire & Politi 27950 

Maguire, Phclps et al. 31691 

Steyermark 89564 

Schultes & Cabrera 17131 

Maguire, Cowan, & Wurdack 

Maguire 33185 

Steyermark 59863 

Garcia Barriga 15096 

Maguirc, 'Tilktt, & Tillett 43808 

Maguirc 32874 

Maguirc, Wurdack, etc. 42168 

Steyermark & Nilsson 745 

Steyermark 58846 

Sttyermark 59933 

*Im Thurn 315 

*Steyermark & Wurdack 677 

30596 

* Wurdack 34134-B 

*Maguirc 35280 

*Maguirc, Piles, & Maguire 6054.3 

Steyermark 93186 

*Stymark 93188 

*Martius 1938 

*Stcymmark 75774 

*Steyermark & Nilsson 436 

*Stymark & Niisson 567 

*Sttymnark 93790 

*Maguire, Pires, & Maguirc 60478 

*Steyermark & Wurdack 678-A 

*Steyermark & Wurdack 677 

*Maguire, Wurdack, & Magiure 

*Maguirc 33034 

42151 

Neblina, Rio Yatua 

Venezuela: Amazonas: Cerro 

Sipapo 

Venezuela: Amazonas: Serrania 

Paru, Rio Ventuari 

Venezuela: Amazonas: Cerro 

Sipapo 

Venezuela: Amazonas: Rio 

Manapiare, Rio Ventuari 

Venezuela : Bolivar : Sierra de Lema, 

Cabeceras de Rio Chicanan 

Colombia: Vaupes: Rio 

Piraparana 

Venezuela: Amazonas: Cerro 

Yapacana, Rio Orinoco 

Venezuela: Bolivar: Ilu-tepui 

Venezuela: Bolivar: Ptari-tepui 

Colombia: Vaupes: Rio Kubiyu 

British Guiana: Upper Mazaruni 

Venezuela : Bolivar : Cerro 

Guaiquinima, Rio Paragua 

Venezuela : Amazonas : Cerro 

Neblina, Rio Yatua 

Venezuela: Bolivar: Cerro Venamo 

Venezuela: Bolivar: Mt. Roraima 

Venezuela: Bolivar: Ptari-tepui 

Venezuela : Bolivar : Mt. Roraima 

Venezuela: Bolivar: ChimantaVenezuela: Bolivar : Sarven-tepui 

tepui 

Venezuela: Amazonas: Serrania 

Brazil: Amazonas: Serra da Neblina 

Venezuela: Bolivar: Auyan-tepi 

Venezuela: Bolivar : Auyan-tepuii

Brazil: Bahia: Serra do Sincora 

Venezuela: Bolivar: ChimantaVenezuela: Bolivar: Cerro Venamo 

Venezuela: Bolivar: Uarama-tepuli

Venezuela: Bolivar: Auyan-tepi 

Brazil: Amazonas: Serra da Neblina 

Venezuela: Bolivar: Chimantfa-tepui 

Yutaje, Rio Manapiare 

tepi 

Venezuela: Bolivar: ChimantaVenezuela: Amazonas: Cerro de la 

Venezuela: Bolivar : Cerro Guaiquitepui 

Neblina, Rio Yatua 

nima Rio Paragua 

.- 

us 

NY holotype 

us 

us 

us 

us 

us 

us 

F isotype 

us 

us 

us 

us 

us 

F holotype 

F holotype 

US isotype 

US holotype 

US holotype 

US holotype 

NY holotype 

us 

US holotype 

M holotype 

US holotype 

US holotype 

US holotype 

US holotype 

US holotype 

us 

US holotype 

US holotype 

US holotype 
NUMBER 2 

List of Specimens Examined-Continued 

Species 

Cottcndmja phclpsiac (L. B. Smith) 

Cottmdorja savanncnsis L. B. Smith 

L. B. Smith 

Cottcndmja scrrulata (L. B. Smith) 

L. B. Smith 

Cottmdmja smulata var. rcducta 

(L. B. Smith) L. B. Smith 

Cottcndmja stmophylla (L. B. Smith) 

L. B. Smith 

Cottcndmja stcymarkii (L. B. Smith) 

L. B. Smith 

Cottcndmja subsimplex (L. B. Smith) 

L. B. Smith 

Cottmdmja thyrsoidea (L. B. Smith) 

L. B. Smith 

Cottcndmja tillandsioidcs (L. B. Smith) 

L. B. Smith 

Cottmdmja wurdackii (L. B. Smith) 

L. B. Smith 

Dcutcrocohnia longipctala (Baker) Mea 

Dyckia brevifolia Baker 

Dyckia chmistaminea Mez 

Dyckia duckci L. B. Smith 

Dyckia maritima Baker 

D3kia rcitzii L. B. Smith 

Encholirium dmij7otum Ule 

Fostnclla pcndulijma (C. H. Wright) 

Fostnclla schidospnma (Baker) 

Hcchtia integmima M. B. Foster 

Hcchtia lundcllimum L. B. Smith 

Stapf. 

L. B. Smith 

Hcchtia montana Brand. 

Hcchtia tillandsioidcs (Andre) L. B. 

Navia abysmophila L. B. Smith 

Navia acaulis Martius ex Schultes f. 

Smith 

Navia aflnis L. B. Smith 

Navia aloifolia L. B. Smith 

Navia angustifolia (Baker) Mez 

Navia arida L. B. Smith & 

Navia aurca L. B. Smith 

Navia barbcllata L. B. Smith 

Steyermark 

Collector and Number 

*Phclps & Hitchcock 501 

*Mag&, Wurdack, & Maguiie 

Cardona 1587 

*Stcynmark & Wurdack 1167 

*Stcyermark & Wurdack 951 

Stcyermark & Wurdack 827 

*Steyermark 75924 

Maguire & Maguirc 35385 

*Stcynmark 74863 

*Maguirc, Cowan, & Wurdack 29637 

Castellanos 261448 

Rcitz 3979 

Rambos s. n. (PACA 48832) 

Eitm & Eitcn 4659 

L. B. Smith & Rcitz 5824 

Rcitz & Klcin 2690 

Foster 2474 

Vmturi 5474 

Kill$ & Smith 25326 

Foster 3072 

Van Hping 6066 

Ferris 8747 

Fostn June 1950 

*Maguire, Wurdack, & Maguirc 

*Idrob0 & Schultcs 644 

*Maguire, Wurdack, & Maguire 

*Maguire, Wurdack, & Bunting 

*Appun 1055 

*Stcycrmark 89698 

*Stcynmark 58219 * Tillctt 45557 

42167 

42471 

42470 

37146 

LocaliQ 

Venezuela: Bolivar: Cerro Paru 

Venezuela: Bolivar: Cerro de la 

Venezuela: Bolivar: Cerro Apacara, 

Venezuela: Bolivar: Chimanta-tepui 

Venezuela: Bolivar: ChimantaVenezuela: Bolivar : Chimanta 

Venezuela: Bolivar : Chimanta

Venezuela: Amazonas: Serrania 

Venezuela: Bolivar: AbacapaVenezuela: Amazonas: Cerro 

Argentina: San Juan: Quebrada 

Brazil: Santa Catarina: Salto 

Brazil: Rio Grande do Sul : Porto 

Brazil: Maranhao: Mun. de Loreto

Brazil: Rio Grande do Sul: Mun. 

Brazil: Santa Catarina: Serra do 

Brazil: Bahia: Amargosa, Milagres 

Argentina: Jujuy: Sierra de Santa 

Peru: Junin: Rio Paucartambo

Mexico? cultivated 

Mexico: San Luis Potosi

Tamazunchale 

Mexico: Sonora: San Carlos Bay, 

near Guayamas 

Mexico: cultivated 

Neblina, Rio Yatua 

Rio Caroni

tepui

Massif 

Massif 

Yutaje 

tepui 

Duida 

del Zonda 

Blumenau 

Alegre 

Torres 

Oratorio S. Joaquim 

Barbara 

Valley 

Venezuela: Amazonas: Cerro de la 

Neblina. Rio Yatua 

21 

Herbarium 

NY holotype 

US holotype 

US holotype 

US holotype 

US holotype 

NY holotype 

US holotype 

NY holotype 

US holotype 

US holotype 

us 

us 

us 

us 

us 

US holotype 

us 

us 

us 

US holotype 

us 

us 

us 

US holotype 

Colombia: Vaupes: Mesa la Lindosa, US 

Venezuela: Amazonas: Cerro de la US holotype 

Venezuela: Amazonas: Cerro de la US holotype 

British Guiana: Marima K holotype 

Venezuela: Bolivar Sierra de Lema 

Ajaju River 

Neblina, Rio Yatua 

Neblina, Rio Yatua 

US holotype 

Venezuela: Amazonas: Cerro Duida F holotype 

British Guiana: Essequibo, US isotype 

Karowtipo, Mazaruni River 
22 

Species 

Navia bicolm L. B. Smith 

Navia brachythyIla L. B. Smith 

Navia brcweri L. B. Smith & 

Navia brocchinioides L. B. Smith 

Navia cardonac L. B. Smith 

Steyermark 

Navia caricifolia L. B. Smith 

Navia cataractarum Sandwith 

Navia caulcsccns Martius ex Schultes f. 

Navia caurnuis L. B. Smith 

Navia colorata L. B. Smith 

Navia connata L. B. Smith & 

Navia nctacca L. B. Smith 

Navia nispa L. B. Smith 

Navia cucuIIata L. B. Smith 

Navia dzyusa L. B. Smith 

Navia duidac L. B. Smith 

Navia fontoidcs L. B. Smith 

Navia garcia-barrigac L. B. Smith 

Navia glauca L. B. Smith 

Navia glcasonii L. B. Smith 

Navia gracilis L. B. Smith 

Navia graminifalia L. B. Smith 

Navia hcchtioidcs L. B. Smith 

Navia hcliophila L. B. Smith 

Navia hohmbergioidcs L. B. Smith 

Navia immersa L. B. Smith 

Navia incrassata L. B. Smith & 

Navia intermedia L. B. Smith & 

Navia involwata L. B. Smith 

Navia Ianigna L. B. Smith 

Steyermark 

Steyermark 

Steyermark 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

List of Specimens Examined-Continued 

*SchuItes 5444 Colombia: Vaupes: Cerro us 

Collector and Number Locality Her barium 

Chiribiquete, Rio Macaya * Tatc 571 Venezuela: Amazonas: Cerro Duida NY holotype 

*Brewer 248 Venezuela: Bolivar: Cerro Jaua, Rio US holotype 

*Maguirc & Politi 28787 Venezuela: Amazonas: Cerro Sipapo US holotype 

Cardona 349 Venezuela: Bolivar Rio Canaracuni US holotype 

fragment 

*Brewer 249 Venezuela: Bolivar: Cerro jaua, Rio US 

*Maguirc & Maguirc 35437 Venezuela: Amazonas: Serrania US holotype 

* Whitton 51 British Guiana: Essequibo, Amatuk K holotype 

*Schultcs 5859 Colombia: Amazonas: Cerro de us 

*Cardona 2985 Venezuela: Bolivar: Cemo Pauo, Rio US holotype 

*Cowan & Wurdack 31417 Venezuela: Amazonas: Serrania US holotype 

*Steyermark 90223 Venezuela: Bolivar: Sierra Ichun US isotype 

*Maguirc, Cowan, & Wurdack 29882 Venezuela: Amazonas: Cerro US holotype 

*Maguirc, Cowan, & Wurdack 30966 Venezuela: Amazonas: Cerro US holotype 

Caura 

Caura 

Yutaje, Rio Manapiare 

Falls, Potaro River 

Cupati, Rio Caqueta 

Caura 

Paru, Rio Ventuari 

Huachamacari 

*Maguire 32766 

*Maguirc, Wurdack, & Maguirc 

* Tatc 404 

*SchuItcs & Cabrera 15391 

*Garcia Barriga & Schultcs 14137 

*Steyermark 58320 

*Maguirc & Fanshawc 32389 

* Wurdack & Addnlcy 43275 

'Schultcs 5492 

*Maguirc & Politi 27574 

'Schultcs 12085 

42537-E 

Moriche, Rio Ventuari 

Guaiquinima, Rio Paragua 

Neblina, Rio Yatua 

Venezuela: Bolivar: Cerro US holotype 

Venezuela: Amazonas: Cerro de la US holotype 

Venezuela: Amazonas: Cerro Duida NY holotype 

Colombia: Amazonas: Rauda US holotype 

Colombia: Amazonas: Araracuara, US holotype 

Venezuela: Amazonas: Cerro Duida F holotype 

Venezuela: Amazonas: Maroa, Rio US holotype 

Colombia: Vaupes: Cerro US isotype 

Venezuela: Amazonas: Cerro US holotype 

Colombia: Amazonas: Raudel de US holotype 

Yayacopi, Rio Apaporis 

Rio Caqueta 

British Guiana: Essquibo us 

Guainia 

Chiribiquete, Rio Macaya 

Sipapo 

Jiriiirimo, Rio Apaporis 

*Maguirc Cowan, & Wurdack 30970 Venezuela: Amazc&s: Cerro US holotype 

Moriche, Rio Ventuari 

*Maguirc, Cowan, & Wurdack 30325 Venezuela: Amazonas: Cerro US holotype 

Huachamacari, Rio Cunucunuma 

*Steyermark 97881 Venezuela: Bolivar: Meseta de US holotype 

Jaua, Rfo Caura 

*Steyermark 97865 Venezuela: Bolivar: Cerro Jaua, US holotype 

Rio Caura 

*Cowan & Wurdack 31359 Venezuela: Amazonas: Serrania US holotype 

Paru, Rio Ventuari 

*Maguirc ct al. 31794 US holotype 

Camani, Rio Ventuari 

Venezuela : Amazonas : Cerro 
NUMBER 2 

Species 

Navia lasiantha L. B. Smith & 

Navia latijolia L. B. Smith 

Navia lcpidota L. B. Smith 

Navia lindmanioidcs L. B. Smith 

Steyermark 

Navia lopezii L. B. Smith 

Navia maguirci L. B. Smith 

Navia maguirci var. minol L. B. Smith 

Navia mima L. B. Smith 

Navia myriantha L. B. Smith 

Navia navicularis L. B. Smith & 

Navia nubicola L. B. Smith 

Steyermark 

Navia occllata L. B. Smith 

Navia octopoidcs L. B. Smith 

Navia parvula L. B. Smith 

Navia paroula var. cxpansa L. B. Smith 

Navia pauci'flora L. B, Smith 

Navia phelpsiac L. B. Smith 

Navia pulvinata L. B. Smith 

Navia pungens L. B. Smith 

Navia ramosa L. B. Smith 

Navia rcJcxa L. B. Smith 

Navia robinsonii L. B. Smith 

Navia rupestris (Gleason) Sandwith 

Navia sandwithii L. B. Smith 

Navia saxicola L. B. Smith 

Navia schultcsiana L. B. Smith 

Navia scopulorum L. B. Smith 

Navia scmismata L. B. Smith 

Navia scrrulata L. B. Smith 

List of Specimens Examined--Continued 

Collector and Number Locality 

*Steyermark 97865-A 

*Maguirc, Cowan, & Wurhk 

*Maguirc, Wurdack, & Bunting 

*Maguirc, Cowan, & Wurdack 30650 

*Schultcs & Lopez 9956 

Maguirc & Politi 27497 

Wurdack & Addcrb 43564 

*Maguirc 24575 

Maguirc 24334 

*Maguirc & Politi 27590 

*Schultes & Lopez 9955 

*Steyermark 90199 

29642 

37288 

Venezuela : Bolivar : Cerro Jaua, 

Venezuela: Amazonas: Cem Duida 

Rio Caura 

Venezuela: Amazonas: Cerro de la 

Venezuela: Amazonas: Cerro 

Brazil : Amazonas : Serra Dimiti 

Venezuela: Amazonas: Cerro 

Venezuela : Amazonas: Rio Siapa 

Surinam: Tafelberg 

Surinam: Tafelberg 

Venezuela: Amazonas: Cerro 

Brazil: Amazonas: Serra Dimiti 

Venezuela: Bolivar: Sierra Ichun 

Neblina, Rio Yatua 

Yapacana, Rio Orinoco 

Sipapo 

Sipapo 

*Maguirc, Wurdack, & Bunting Venezuela: Amazonas: Cerro de la 

*Maguirc & Politi 27500 Venezuela: Amazonas: Cerro 

*Maguirc & Maguirc 29151 Venezuela: Amazonas: Cerro 

*Maguirc, Wurdack, & Bunting Venezuela: Amazonas: Cerro de la 

Maguire, Wurdack, & Maguire 42246 Venezuela: Amazonas: Cerro de la 

37392 Neblina, Rio Yatua 

Sipapo 

Duida 

37326 Neblina, Rio Yatua 

*Maguire & Maguirc 29154 

'Maguirc, Phelps ct al. 31762 

*Maguirc 29111 

*Maguirc ct al. 31652 

*Cardona 2997 

*Schultcs & Lopez 9321 

*Brewer 250 

*Maguirc & Fanshawc 32360 

*Maguirc & Fanshawe 23435 

'Maguirc, Cowan, & Wurdack 

*Schultcs 5655 

30680 

*Steyermark 75613 

*Maguirc, Cowan, & Wurdack 

*Maguirc, Cowan, & Wurdack 

30959 

30521 

Neblina, Rio Yatua 

Marahuaca, Rio Cunucunuma 

Guanay, Rio Ventuari 

Duida 

Guanay, Rio Ventuari 

Rio Caura 

Castillo) Rio Negro 

Rio Caura 

Falls, Kumpung River 

Venezuela: Amazonas: Cerro 

Venezuela: Amazonas: Cerro 

Venezuela: Amazonas: Cerro 

Venezuela: Amazonas : Cerro 

Venezuela: Bolivar: Cerro Pauo, 

Colombia: Vaupes: San Felipe (El 

Venezuela: Bolivar: Cerro Jaua, 

British Guiana: Essequibo Macreba 

British Guiana: Kaieteur Falls 

Venezuela: Amazonas: Cenu YapaColombia: Vaupes: Cerro del 

Venezuela: Bolivar: Chimanta 

Venezuela: Amazonas: Cerro 

Venezuela: Amazonas: Cerro Yapacana, Rio Orinoco 

Castillo, Rio Apaporis 

Massif 

Moriche, Rio Ventuari 

cana, Rio Orinoco 

23 

Herbarium 

US holotype 

US holotype 

US holotype 

US holotype 

US holotype 

us 

us 

US isotype 

US holotype 

US holotype 

US isotype 

US holotype 

US holotype 

US holotype 

US holotype 

US holotype 

US holotype 

US isotype 

US holotype 

US holotype 

US holotype 

US holotype 

US holotype 

us isotype 

us isotype 

of N. 

Juviatilis 

us 

US holotype 

us isotype 

US holotype 

US holotype 

US holotype 
24 

Species 

Nauia splmdens L. B. Smith 

Nauia stenodonta L. B. Smith 

Navia steyermarkii L. B. Smith 

Nauia subpetiolata L. B. Smith 

Navia trichodonta L. B. Smith 

Navia umbratilis L. B. Smith 

Navia uiridis L. B. Smith 

Nauia wurdackii L. B. Smith 

Napia xyridiflora L. B. Smith 

Pitcairnia altmteinii (Link ex K1. & 

Pitcairnia angustijolia Soland. 

Pitcaimia archeri L. B. Smith 

Otto) Lem. 

Pitcairnia armata Maury 

Pitcaimia atrorubens (Beer) Baker 

Pitcaimia brunnescens L. B. Smith 

Pitcaimia brongniartiana Andre 

Pitcaimia brittoniana Mez 

Pitcaimia bulbosa L. B. Smith 

Pitcaimia caricifolia Martius ex 

Pitcaimia gracilis Mez 

Pitcairnia heterophyla (Lindl.) Beer 

Pitcairnia macrochlamys Mez 

Pitcaimia muheliana AndrC 

Schultes f. 

Pitcaimia multtyora L. B. Smith 

Pitcaimia nigra (Carr.) Andre 

Pitcaimia nuda Baker 

Pitcairnia oaxacana L. B. Smith 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

List of Specimens Examined-Continued 

Collector and Number 

Tillett 45841 

*Cowan & Wurdack 31357 

*Steyermark 58012 

"Cowan & Wurdack 31368 

*Maguire, Cowan, & Wurdack 

*Maguire & Maguire 35406 

*Maguire & Maguire 29078 

* Wurdack 34324 

*Steyermark 58152 

Degado 45 

Sargent 443 

Killip 35255 

Maguire, Wurdack, & Bunting 36094 

30228 

Moore & Bunting 8704 

Asplund 19060 

Killip 34835 

Steyermark & Aristeguieta 15 

Maguire, Wurdack, & Bunting 36083 

Cuatrecasas 7520 

Stehle I804 

Wilbur & Wilbur 2295 

Breedlove 6359 

H. E. Mme 8165 

Cuatrecasas 23948 

Asplund 20040 

A. C. Smith 3644 

Moore & Buntina - 8763 

Pitcairniapaniculata (R. & P.) R. & P. 

Pitcairnia pungens H. B. K. 

Pitcairnia punuea Scheidw. 

Pitcairnia ringm K1. ex Link 

Pitcairnia valerii Standley Allen 2414 

Pup santosii Cuatrecasas 

Puya sodiroana Mez Foster 2605 

Macbride 5114 

Pittk 1028 

Breedlove & Raum 13569 

Van Hying 599 

Cuatrecasas & Jaramillo 12042 

Locality 

British Guiana: Essequibo, Utschi 

Venezuela: Amazonas: Serrania 

Venezuela: Amazonas: Cerro Duida 

Venezuela: Amazonas: Serranfa 

Venezuela: Amazonas: Cerro 

Venezuela: Amazonas: Serrania 

Venezuela: Amazonas: Cerro Duida 

Venezuela: Bolivar: Sarven-tepui 

Venezuela : Amazonas : Cerro Duida 

Venezuela 

Falls, Kamarang River 

Paru, Rio Ventuari 

Paru Rio Ventuari 

Huachamacari, Rio Cunucunuma 

Yutaje, Rio Manapiare 

Puerto Rico: Maricao 

Colombia: Choco: Upper Rio San 

Juan 

Venezuela: Amazonas: Great 

Rapids of Orinoco, Palomal 

Mexico: Nayarit: Tepic to 

Jalcocotan 

Ecuador: Pichincha: El Paso to 

El Volante 

Colombia: Valle: Rio Engana 

Venezuela: Bolivar: El Dorado to 

Luepa 

Venezuela: Amazonas: Great 

Rapids of Orinoco, Puerto 

Ayacucho 

Colombia: Vaupes: Rlo Guayabero 

Guadeloupe: Ste. Rose, near Callas 

Mexico: Jalisco: Autlan 

Mexico: Chiapas: Mun. Tenejapa, 

Mexico: Jalisco: Colima to Puente 

Colombia: Valle: Hoya del Rio 

Ecuador: Tungurahua: Rio Verde 

British Guiana: Mt. Iramaikpang, 

Mexico: Michoacan: Arteaga 

Peru: Huanuco: Pampayacu 

Colombia: Cauca: Tacueyo, Rio 

Mexico: Chiapas: Ocozocoautla 

Mexico: Veracruz: La Joya 

Panama: Panama: Cerro Campana 

Colombia: Cundinamarca: Sabana 

Ecuador: Azuay: Cuenca to Ona 

Tanate River 

San Pedro 

Digua, Rio San Juan 

Grande 

Kanuka Mts. 

Pal0 valley 

de Bogoa 

Herbarium 

US isotype 

US holotype 

F holotype 

US holotype 

US holotype 

US holotype 

US holotype 

US holotype 

F isotype 

us 

us 

us 

us 

us 

us 

us 

us 

us 

us 

us 

us 

us 

us 

us 

us 

us 

us 

us 

us 

us 

us 

us 

us 

us 
NUMBER 2 25 

Literature Cited 

Anderson, L. E. 

1954. Hoyer’s Solution as a Rapid Permanent Mounting 

Medium for Bryophytes. The Bryologut, 57 : 242- 

244. 

Ayensu, E. S. 

1969. The Identity of Velloria uaipanensis (Maguire) 

L. B. Smith: Anatomical Evidence. Memoirs of 

The New York Botanical Garden, 18( 2) : 291-298. 

Memoirs 

Smith, L. B. 

1967a. Bromeliaceae of the Guayana Highland. 

of the New York Botanical Garden, 14: 15-68. 

1967b. Notes on Bromeliaceae, XXV. Phytologia, 14: 

457-491. 

1969. Bromeliaceae. In Maguire, Botany of the Guayana 

Highlands. Memoirs of the New York Botanical 

Garden, 18(2) :29-32. 

Solereder, H., and F. J. Meyer. 

1929. Systematische Anatomie der Monokotyledonen: 

Bromeliaceae. IV: 80-1 29. Berlin. 

Stebbins, G. L., and G. S. Khush. 

1961. Variation in the Organization of the Stomata1 

Complex in the Leaf Epidermis of Monocotyledons 

and Its Bearing on Their Phylogeny. American 

lournal of Botany, 48 : 5 1-59. 
26 

1 2 

11 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

7 

12 14 

4 

9 

FIGURES 1-15.-Connellia and Cottendorfia, leaf anatomy. Connellia guelchii: 1, cross section, x 133 ; 2, abaxial epidermis, 

X 267; 3, abaxial subepidermis, X 267. Cottendorfia Porida: 4, cross section, X 133; 5 abaxial epidermis, X 267; 6, abaxial 

subepidermis, X 267. C. longipes: 7, abaxial epidermis, X 267; 8, abaxial subepidermis, X 267; 9, cross section, X 133. C. 

dyckioides: 10, abaxial epidermis, X 267; 11, abaxial subepidermis, X 267; 12, cross section, X 133. C. tillandsioides: 13, 

abaxial epidermis, X 267; 14, abaxial subepidermis, x 267; 15, cross section, x 133. 
NUMBER 2 27 

FIGURES 16-32.-Cottendorfia, leaf anatomy. C. brachyphylla: 16, abaxial epidermis, x 267; 17, abaxial subepidermis, x 267; 

18, cross section, X 133. C. wurdackii: 19, abaxial epidermis, X 267; 20, abaxial subepidermis, X 267; 21, cross section, 

X 133. C. thyrsoidea: 22, abaxial epidermis, X 267; 23, abaxial subepidermis, x 267; 24, cross section, x 133. C. minor: 

25, cross section from near midleaf, X 133; 26, cross section from near lower fourth, x 133; 27, abaxial epidermis, x 267; 

28, abaxial subepidermis, x 267. C. serrulata var. reducta; 29, abaxial epidermis, x 267; 30, abaxial subepidermis, x 267. 

C. gracillima: 31, abaxial epidermis, x 267; 32, abaxial subepidermis, x 267. 
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

36 

44 

33 

39 

FIGURES 3349.-Cottendorfia, leaf anatomy. C. serrulata var. reducta: 33, cross section, x 133. C. subsimplex: 34, abaxial 

epidermis, X 267; 35, abaxial subepidermis, X 267; 36, cross section, X 133. C. stenophylla: 37, abaxial epidermis, X 267; 

38, abaxial subepidermis, X 267; 39, cross section, X 133. C. argentea: 40, scale, X 267; 41, abaxial epidermis, X 267; 42, 

abaxial subepidermis, X 267; 43, cross section, X 133. C. guianensis: 44, abaxial epidermis, x 267; 45, abaxial subepidermis, 

X 267. C. dendritica: 46, abaxial epidermis, X 267; 47, abaxial subepidermis, x 267. C. cylindrostachya: 48, abaxial epidermis, X 267; 49, abaxial subepidermis, x 267. 
09 

69 

’/ 

ss 
30 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

70 

85 

FIGURES 70-87.-Nuia, leaf anatomy. N. reflexa: 70, cross section, x 133; 71, adaxial epidermis, x 267; 72, adaxial subepidermis, X 267; 73, adaxial third cell layer, X 267; 74, abaxial epidermis, x 267; 75, abaxial subepidermis, x 267; 76, 

azMxial third cell layer, X 267. N. hechtioides: 77, abaxial epidermis, x 267; 78, abaxial subepidermis, x 267. N. hohenbergwidcs: 79, cross section, X 133; 80, abaxial epidermis, X 267; 81, abaxial subepidermis, x 267. N. gracilis: 82, abaxial 

epidermis, X 267; 83, abaxial subepidermis, X 267. N. brocchinioides: 84, abaxial epidermis, X 267; 85, abaxial subepidermis 

X 267. N. diffusa: 86, abaxial epidermis, x 267; 87, abaxial subepidermis, x 267. 
NUMBER 2 31 

103 

106 

FIGURES 88-106.-Nauia, leaf anatomy. N. garcia-barrigae: 88, cross section, x 133; 89, scale, x 133; 90, abaxial epidermis, 

X 267; 91, abaxial subepidermis, X 267. N. rupestris: 92, cross section, x 133; 93, abaxial epidermis, x 267; 94, abaxial 

subepidermis, X 267. N. cataractarum: 95, abaxial epidermis, x 267; 96, abaxial subepidermis, x 267. N. lindmanioides: 

97, abaxial epidermis, X 267; 98, abaxial subepidermis, x 267; 99, cross section, x 133. N. parvula: 100, cross section, 

X 133; 101, abaxial epidermis, X 267; 102, abaxial subepidermis, x 267. N. Iepidota: 103, abaxial epidermis, x 267; 104, 

abaxial subepidermis, X 267; 105, abaxial third cell layer, x 267; 106, cross section, x 133. 
2s NVINOSHIIHS SNOI.I,ilB~N03 OL ANVLOS 
NUMBER 2 33 

FIGURES 127-138.-Na~ia, leaf anatomy. N. Iasiantha: 127, cross section, adaxial surface, x 133; 128, cross section, abaxial 

surface, X 133. N. incrassata: 129, abaxial epidermis, X 267; 130, abaxial subepidermis, X 167; 131, cross section, adaxial 

surface, X 133; 132, cross section, abaxial surface, X 133. N. intermedia: 133, cross section, X 133; 134, abaxial epidermis, 

X 267; 135, abaxial subepidermis, X 267. N. immersa: 136, cross section, x 133; 137, abaxial epidermis, x 267; 138, abaxial 

subepidermis, X 267. 
34 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

139 

140 

142 

4 

FIGURES 139-156.-Nuuia, leaf anatomy. N. cucullata: 139, cross section, X 133; 140, abaxial epidermis, x 267; 141, abaxial 

subepidermis, X 267. N. pulvinata: 142, abaxial epidermis, x 267; 143, abaxial subepidermis, x 267; 144, cross section, 

X 133. N. scopulorum: 145, cross section, x 133; 146, abaxial epidermis, x 267; 147, abaxial subepidermis, x 267. N. 

ungustifoliu: 148, abaxial epidermis, X 267; 149, abaxial subepidermis, X 267; 150, cross section, x 133. N. abysmophih: 151, cross section, X 133; 152, abaxial epidermis, X 267; 153, abaxial subepidermis, x 267. N. duiduc: 154, abaxial epidermis, X 267; 155, abaxial subepidermis, X 267; 156, cross section, x 133. 
NUMBER 2 35 

I57 

169 

li 

159 

170 

II 175 

FIGURES 157-178.-Navia, leaf anatomy. N. nubicola: 157, cross section, x 133; 158, abaxial epidermis, x 267; 159, abaxial 

subepidermis, X 267. N. crctacca: 160, abaxial epidermis, X 267; 161, abaxial subepidermis, X 267. N. latifolia: 162, abaxial 

epidermis, X 267; 163, abaxial subepidermis, x 267; 164, abaxial third cell layer, X 267. N. brachyphylla: 165, abaxial 

epidermis, X 267; 166, abaxial subepidermis, X 267. N. glauca: 167, abaxial epidermis, x 267; 168, abaxial subepidermis, 

X 267. N. semiserrata: 169, &axial epidermis, x 267; 170, abaxial subepidermis, X 267; 171, cross section, x 133. N. 

caurcnsis: 172, abaxial epidermis, x 267; 173, abaxial subepidermis, x 267; 174, cross section, x 133. N. octopoides: 175, 

abaxial epidermis, X 267; 176, abaxial subepidermis, X 267. N. umbratilis: 177, abaxial epidermis, x 267; 178, abaxial 

subepidermis, x 267. 
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

186 183 

FIGURES 179-194.-Navia, leaf anatomy. N. umbratilis: 179, cross section, x 133. N. serrulata: 180, abaxial epidermis, 

X 267; 181, abaxial subepidennis, X 267; 182, cross section, x 133; 183, scale, x 133. N. arida: 184, scale, x 133; 185, 

abaxial epidermis, X 267; 186, abaxial subepidermis, X 267; 187, cross section, x 133. N. saxicola: 188, abaxial epidermis, 

X 267; 189, abaxial subepidermis, X 267; 190, cross section, x 133. N. caulescens: 191, abaxial epidermis, x 267; 192, 

abaxial subepidermis, X 267; 193, cross section, x 133; 194, scale, x 133. 
12 E 
8E NVINOSHLIHS SNOI&fMlXLNOD O& ANV&Og 
NUMBER 2 39 

8 

FIGURES 232-250.-Navin, leaf anatomy. N. navicularis: 232, cross section, x 133; 233, abaxial epidermis, x 267; 234, 

abaxial subepidermis, X 267. N. breweri: 235, abaxial epidermis, x 267; 236, abaxial subepidermis, x 267. N. colorata: 237, 

abaxial epidermis, X 267; 238, abaxial subepidermis, X 267; 239, cross section, x 133. N. xyridipora: 240, abaxial epidermis, 

X 267; 241, abaxial subepidermis, X 267; 242, cross section, x 133. N. wurdackii: 243, cross section, x 133; 244, trichome, 

X 133; 245, abaxial epidermis, X 267; 246, abaxial subepidermis, X 267. N. pauciflora: 247, abaxial epidermis, x 267; 248, 

abaxial subepidermis, X 267. N. steyannarkii: 249, abaxial epidermis, x 267; 250, abaxial subepidermis, x 267. 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

251 

252 

253 

256 

FIGURES 251-262.-Navia, leaf anatomy. N. aloifolia: 251, abaxial epidermis, x 267; 252, abaxial subepidermis, x 267; 

253, cross section, x 133. N. viridis: 254, abaxial epidermis, x 267; 255, abaxial subepidermis, x 267; 256, cross section, 

X 133. N. crispa: 257, abaxial epidermis, X 267; 258, abaxial subepidermis, x 267. N. stenodonta: 259, abaxial epidermis, 

X 267; 260, abaxial subepidermis, X 267; 261 cross section, X 133; 262, trichome, x 133. 
NUMBER 2 41 

i 271 

270 

272 

h 

274 

FIGURES 263-277.-Navia, leaf anatomy. N. myriantha: 263, abaxial epidermis, x 267; 264, abaxial subepidermis, x 267; 

265, trichome, X 133; 266, cross section, X 133. N. trichodonta: 267, abaxial epidermis, X 267; 268, abaxial subepidermis, 

X 267. N. lanigera: 269, abaxial epidermis, X 267; 270, abaxial subepidermis, x 267; 271, trichome, x 133. N. phelpsiae: 

272, abaxial epidermis, X 267; 273, abaxial subepidermis, x 267; 274, trichome, x 133. N. lopezii: 275, cross section, 

X 133; 276, abaxial epidermis, X 133; 277, abaxial epidermis, X 133. 

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