SMITHSONIAN CONTRIBUTIONS TO 

BOTANY 

NUMBER 6 

Wallace R. Ernst Floral Morphology 

and Systematics of 

Lamourouxia 

(Scrophulariaceae: 

Rhinanthoideae) 

SMITHSONIAN INSTITUTION PRESS 

CITY OF WASHINGTON 

1972 
ABSTRACT 

Ernst, Wallace R. Floral morphology and systematics of Lamourouxia (Scrophulariaceae : Rhinanthoideae) . Smithsonian Contributions to Botany, number 6, 63 

pages, 37 figures. 197 1 .-Relationships among the twenty-six species, having an 

overall range from northern Mexico to central Peru, are analyzed in regard to 

pollinators and for insight into the differentiation of the three taxonomic sections 

of the genus. Four new names are proposed, including a section, two species, and 

a combination. Two new chromosome numbers are reported. Descriptions for each 

of the species and a dichotomous key are included. 

Errata. Page 43, col. 2: first paragraph under Section I1 

should read "Corollae tubus plus minusve infundi buliformis, 

non ventricosus subroseus vel lavandulaceus vel magenteus, 

non rubro-aurantiacus. 

inferiora, longiora et majora." Page 49, col. 2: transfer 

first two paragraphs to col. 1, preceding "16. L. viscosa." 

Stamina dimorpha, omnia fertilia, 

Library of Congress Cataloging in Publication Data 

Ernst, Wallace Roy, 1928-1971. 

Floral morphology and systematics of Lamourouxia (Scrophulariaceae : Khinanthoideae) 

(Smithsonian contributions to botany, no. 6) 

Bibliography: p. 

1. Lamourouxia. 2. Botany-Latin America. I. Title. 11. Series: Smithsonian Institution. 

Smithsonian contributions to botany, no. 6. 

QKlS274.7 no. 6 [QK495.S43] 581'.08s [583'.81] 72-39820 

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

in the Institution's annual report, Smithsonian Year. 

For sale by the Suoerintendent of Documents, US. Government Printing OWce 

Washington, D.C. 20402 - Price $1.26 (paper cover) 
Contents 

Page 

INTRODUCTION ............................................. 1 

MATERIALS AND METHODS ... ............................ 2 

3 

ACKNOWLEDGMENTS ............................................ 3 

SYSTEMATIC POSITION ....................................... 4 

GEOGRAPHICAL DISTRIBUTION ........................................ 4 

HEMIPARASITISM .............................................. 7 

VEGETATIVE CHARACTERISTICS ..................... ........... 8 

Stems and Inforescences ................................... 8 

Leaves 9 

FLORAL CHARACTERISTICS ................................... 9 

Calyx ......................................................... 9 

Corolla ...................................................... 9 

Stamens ..................................... 

Ovary Capsule ..................................... 

Seeds ................................................... 12 

Hairs ...................................... .... ..... 13 

COMPARATIVE MORPHOLOGY ...................... ............ 15 

Section Lamourouxia .......................................... 15 

Section Adelphidion ....................................... 17 

Section Hemispadon ......................................... 18 

FLORAL FUNCTION AND POLLINATION ............................... 19 

The Red Corollas in Mexico and Central AmericaSections Lamourouxia and Hemispadon ........................ 21 

The Non-red Corollas in South America and MexicoTypification ................................................ 2 *. Variation ..................................................... 

................................................... 

Section Adelphidion ....................................... 22 

SYNOPSIS OF VEGETATIVE CHARACTERS ............................ 22 

DISCIJSSIOX ..................................................... 24 

Systematics and Differentiation of the Sections ............ ...... 24 

Sections Lamourouxia and Hemispadon ......................... 25 

Sections Adelphidion and Hemispadon .......................... 25 

Sections Lamourouxia and Adelphidion .......................... 26 

SYNOPSIS AND CONCLUSIONS .................................. 26 

TAXONOMY, KEY, AND DESCRIPTIONS ............................ 28 

BIBLIOGRAPHY .................................. ..... 61 

INDEX TO THE SPECIES ............................................. 63 

Wallace R. Ernst Floral Morphology 

Introduction 

Some confusing patterns of similarities and differences 

among the species of Lamourouxia attracted me to 

this genus. The distribution of floral and vegetative 

characters seemed almost to be random so that relationships among the species were obscured. Since none 

of the previous authors dealing with this genus has 

attempted an explanation, the purpose of this paper 

is to provide a systematic restructuring of the genus to 

account for morphological relationships among the 

species. The main obstacle proved to be that floral 

and vegetative characters were diffcult to correlate 

for systematic purposes. This forced the decision to 

concentrate on floral criteria, as these provided the 

lower morphological denominator for clustering the 

species into coordinate sections. The flowers of Lamourouxia exhibit two distinctive pollination syndromes. This differentiation brings to mind the observation by Ornduff (1969) that ‘‘in genera having 

a diversity of breeding systems, a taxonomy which 

emphasizes floral characters will be strikingly different from a taxonomic treatment which stresses 

characters not associated with reproductive methods.” 

Thus, we are confronted with a challenge to compare and evaluate the various merits of systems based 

primarily on vegetative or on floral characters, In 

the case of Lamourouxia, the contest resolves to 

choice between characters of low selective value, 

Wallace R. Ernst, Department of Botany, Smithsonian Institution, Washington, D.C. 20560 and The University of 

Kansas, Lawrence. Dr. Ernst’s colleagues will be sorry to 

learn of his death on October 8, 1971, at which time this 

study was in press. 

and Systematics of 

Lamourouxia 

(Scrophulariaceae: 

Rhinanthoideae) 

since Lamourouxia apparently does not depend upon 

vegetative reproduction, and characters of high selective value. 

This paper is based primarily upon comparative 

morphology of the flowers of Lamourouxia and the 

goal is an objective systematic arrangement of the 

species which‘ should also facilitate their identification. 

The necessary first step to its completion was the 

taxonomic revision comprising the latter portion of 

this report. With the taxonomy untangled and the 

species arranged in systematic order, it also seems 

possible to predict some of the circumstances that 

were important in the differentiation of the three 

sections of this genus. To this end, I propose to 

evaluate in the first portion of this study the taxonomic and the biological significance of floral and 

vegetative characters by analyzing their distributions 

among the species and by considering their role in 

reproduction and evolution. In defense of my taxonomic framework, I shall attempt to show that the 

most important taxonomic characters also are biologically significant. My conclusions regarding evolution within Lamourouxia, as judged from morphological relationships among the species, can be 

summarized best by the postulates of Ehrlich & 

Raven (1969) that “selection is both the primary 

cohesive and disruptive force in evolution . . . ,” 

that “populations will differentiate if they are subjected to different selective forces and will tend to 

remain similar if they are not . . .,” and that “it is 

the local interbreeding population and not the 

species that is clearly the evolutionary unit of importance.” 

1 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

The second portion of this paper includes a new 

sectional name ( Adelphidion) , two new species names 

(L. dispar and L. parayana), one new combination 

(L. dasyantha) , two new chromosome numbers (L. 

longiflora and L. viscosa), descriptions for the 26 

species, and a key to their identification. The generic 

name Lamourouxia is a later homonym which only 

recently was conserved (Ernst, 1968; McVaugh, 

1969). Although this genus previously has been revised three times (Bentham, 184-6; Robinson & Greenman, 1895; Paray, 1950), none of these revisions 

represents the diversity now known. Bentham’s generic 

framework was established on the basis of 18 binomials of which only 12 are now recognized as 

species. Four of the 14 species described since 1846 

were collected for the first time between 1952 and 

1958. 

Strictly herbarium studies provide useful information for taxonomic purposes and insight into the 

ranges of variability for commonly accepted taxonomic criteria. For curatorial purposes, monographs 

stressing morphology help to bring forth the correct 

name for the appropriate specimens. In most problem instances, however, the monographer who deals 

only with comparative morphology is not in a favorable position to decide whether a species is a good 

one from a behavioral point of view, and he is at 

an even greater disadvantage when it comes to guessing at phylogeny. Often as not, his studies must 

end where the most interesting part of the subject 

begins. I could readily subscribe to the hypothesis 

that an understanding of the variation patterns in 

Lamourouxia would depend in large measure upon 

an understanding of the morphology and behavior 

of the pollinators. As summarized by Slobodkin for 

another context (see Ehrlich & Holm, p. 248, 1963), 

we are left to wonder what the real world is like. 

Materials and Methods 

This study is based upon 1178 separate collections 

representing 3025 herbarium specimens for which 

all label data have been recorded for comparison. 

These specimens have been studied at, or borrowed 

from, the following 26 institutions, which are abbreviated according to Lanjouw & Stafleu (1964) : A, 

BH, BM, C, CAS, COL, DS, ENCB, F, G, GH, K, 

MEXU, MICH, MO, MSC, NY, P, PENN, PH, 

POM, RSA, TEX, UC, US, WIS. The Escuela 

National de Ciencias Biologicas, Mexico, D. F., is 

abbreviated as ENCB (see Taxon 15: 334. 1966). 

All types and other specimens of Lamourouxia at 

these institutions have been examined and annotated 

up to 1968. Samples from 133 collections, representing all taxa, were cleared in lactic acid and microscopically compared. Seeds, hairs, and the venation 

patterns of leaves and flowers were examined. Meiosis 

was examined in two species (L. longiflora and L. 

viscosa) using buds fixed by Dr. D. E. Breedlove 

in acetic ethanol. The anthers were squashed in 

acetocarmine and observed in bright field and phase 

contrast. 

TYPIFICATION 

The generic name Lamourouxia is a later homonym, 

but was proposed for conservation (Ernst, 1968) and 

conserved (McVaugh, 1969) since its first application was as a superfluous name in algae and there is 

no alternative in Scrophulariaceae. The first seven 

species were published in Humboldt, Bonpland, & 

Kunth (1818). The authorship of the binomials in 

this work, Nova Genera et Species Plantarum, is a 

matter of contention among those favoring Kunth 

( Stearn, 1968) and those supporting Humboldt, Bonpland, & Kunth (Morton, 1969; Stafleu, 1967). The 

necessity at hand is the bibliographical reference to 

the place and date of publication of the earliest 

names in Lamourouxia. Since the generic name and 

constituent binomials were validly published for the 

purpose of establishing priority within Scrophulariaceae, the precise authorship of the names seems to 

me of much less importance. I prefer the widely recognized and understood “H.B.K.” which readily 

identifies the important volumes of the Nova Genera 

et Species Plantarum. Little additional glory can now 

accrue to Kunth by attributing these binomials only 

to him, or to insist upon the more awkward reference 

to Kunth in Humboldt, Bonpland, & Kunth. 

When Bentham (1846), in the first revision of the 

genus, created a section Lamourouxia, I believe this 

narrowed the choice for generic lectotype to one of 

the species in this section. The only original species 

of the genus included in this section was L. multifida 

HBK., and it does not matter now what the rationale 

for the sectional name was in 1846. The only species 

which can at the same time typify the genus Lamourouxia and the section Lamourouxia is L. multifida. 

The Report of the Committee for Spermatophyta 

(McVaugh, 1969) found that the only acceptable 
NUMBER 6 3 

lectotype for the conserved generic name is L. multifida and I have proceeded on the strength of that 

recommendation. 

The proposal by Thieret (1954) that L. uirgata 

HBK. serve as lectotype for the genus was a reasonable choice for a representative species; however, this 

choice introduced an awkward problem in regard to 

sectional names and there is neither morphological 

nor nomenclatural reason why this species must be 

accepted for lectotype. I advocated ( Ernst, 1968) 

acceptance of Thieret’s choice of lectotype because 

the original generic description in Humboldt, Bonpland, & Kunth seemed to deal more directly with 

the kind of staminal development represented in that 

species. The staminal characteristics of L. multifida 

also were covered but rather maladroitly, as the name 

“L. pinnatifida” was inadvertently substituted for L. 

multifida although the latter name was used in the 

formal description of the species. If L. virgata were 

retained as lectotype, then Bentham’s section Lamourouxia would have to be discarded and a totally different section Lamourouxia, new in description and 

new in constitutent species, would have to be created 

to accommodate L. virgata and its nearest allies. 

When Robinson & Greenman (1895) combined 

Bentham’s sections Lamourouxia and Euphrasioides, 

they chose the second name to represent the section; 

however, priority must rest with Lamourouxia. Selection of lectotype for section Euphrasioides must be 

from among those species included under that name 

by Bentham and, of course, this cannot be L. multifida. I propose that L. xalapensis HBK. serve as 

lectotype species for section Euphrasioides Bentham. 

In my system, section Euphrasioides, following the 

example of Robinson & Greenman, is combined with 

section Lamourouxia. The remaining species of the 

genus comprise two sections. I propose that L. uiscosa 

HBK. serve as lectotype species for section Hemispadon and that L. virgata HBK. serve as type species 

for a new section Adelphidion. 

Following the binomial references in the taxonomic 

portion of this paper, I have quoted the author’s own 

published phrases or words concerning the type 

locality, and, when possible, that portion stating the 

collector and the collector’s serial number. This information is the only tangible means for tying the 

published, and presumably edited, account of the 

species to a voucher specimen or specimens. Methods 

of expression have changed and, often as not, these 

data differ somewhat from those appearing on the 

presumed type-specimen label or from current practice. Serial numbers, sometimes supplied later or by 

someone other than the collector, especially on older 

collections, are not always indicative of a single 

gathering, but may be found on specimens taken on 

different days or from different regions. The collections of Galeotti illustrate this difficult situation and 

other examples are not rare, but tedious to discuss. 

Even so, I believe that most botanists base their 

descriptions on all available material at hand rather 

than on a single sheet, and that it is unnecessary to 

designate individual specimens as lectotypes for species 

names when the type collections are not taxonomically 

mixed. I have tried to avoid the much abused term 

“holotype,” as I believe this status can be rightfully 

conferred on a specimen only by the author of a new 

taxon in his original description. All specimens mentioned have been examined and the duplicates of type 

collections which I have cited are taxonomic equivalents. 

VARIATION 

Measurements in the species descriptions are taken 

from mature organs, the smallest at the left, an 

average in parentheses, followed by the largest. Averages are from 10 or more measurements. Notable 

extremes are given in parentheses to the left or right. 

Miscellaneous herbarium specimens gathered at random seldom are adequate for a standardized measuring technique. Diversity within species probably would 

seem less extreme and differences between species 

more distinctive if the measurements could have been 

taken from strictly equivalent organs in exactly the 

same state of development. The dichotomous keys to 

the species, accordingly, are regrettably difficult. 

The distributions of the species are given by country, 

state, or department. Complete data are given for 

rare, unusual, or problem specimens. For the more 

common species, only the number of separately numbered collections examined for that area is given in 

parentheses, excluding duplicates. All specimens mentioned and plotted have been examined. 

Acknowledgments 

I am very grateful to the curators and directors of 

institutions where I have studied or borrowed specimens of Lamourouxia. Among the many others who 

have helped me, the following have my special thanks: 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

D. E. Breedlove, K. V. Flannery, M. E. Hale, Jr., 

H. Heine, C. V. Morton, A. Skovsted, J. H. Thomas, 

H. J. Thompson, and J. Roberts. 

Systematic Position 

The position of Lamourouxia in subfamily Rhinanthoideae ( "Rhinanthideae" ) has remained unchanged 

since Bentham's ( 1846) classification of Scrophulariaceae. In a systematic sense this is near the end of the 

family among genera such as Bartsia, Castilleja, Euphrasia, Odontites, Pedicularis, and Rhinanthus. The 

criterion for establishing the subfamily, which is 

clearly evident in the buds of all species of Lamourouxia, is the ontogeny of the corolla in which the 

lower lip enfolds the upper. 

Tribal status is less settled, Euphrasieae (Bentham, 

1846; Thieret, 1954), Rhinantheae (Wettstein, 

1891), and cannot be resolved here without a comprehensive survey of many other genera. It is clear, 

however, that if Lamourouxia and Rhinanthus are 

sufficiently similar to be accommodated within the 

same tribe the name of the tribe must be Rhinantheae. The significant characters appear to be the 

galeate shape of the upper lip of the corolla, the 

frequently connivent anthers, and the hemiparasitic 

nature of the plants. The term galeate is ellusive 

in application and, in this case, probably is of little 

importance. The upper lip of the corolla is conspicuously galeate in the buds of all species but 

remains so only in L. brachyantha. In the others, the 

upper lip may seem more or less inverted naviculiform, or once the lobes have reflexed, the corolla, 

even through sheltering the anthers, may seem little 

or no more galeate than sympetalous corollas of other 

families. In five or six species the anthers are exserted 

and spreading; in most of the others the fully fertile 

anthers are connivent in pairs by means of shaggy 

pubescence, and are displayed near the mouth of the 

corolla. The similarity of the seeds to those of other 

genera known to be hemiparasitic suggests that Lamourouxia also may make use of rootgrafts for 

nutritional supplements. 

Lamourouxia does not seem to have any very close 

relationship with other Scrophulariaceae in Mexico 

and Central America which might give rise to a 

taxonomic confusion. In South America, however, 

L. sylvatica somewhat recalls the genus Bartsia L., 

and L. uirgata somewhat recalls the genus Virgularia 

Ruiz & Pavon. Taxonomic characters are available 

to distinguish these genera, however, and nothing 

is as yet known about biosystematic relationships. The 

peculiar seed coats which are among the unifying 

features of Lamourouxia are very similar to those of 

Virgularia, suggesting a relatively close relationship 

while the very different seeds of Bartsia suggest a 

more remote relationship (Figure 1 ) , 

Geographical Distribution 

The distribution of Lamourouxia extends from 

northern Mexico at about 30" N, 109' W, in the 

state of Sonora, to western Panama at about 8" 30' 

N, 80" 30' W. It begins again in Colombia near 

the Venezuelan border at about 7' 30' N, 73" W, 

and extends southward to central Peru, near 12" S, 

75" W (Figures 2, 3) , Twenty-four species occur 

between northern Mexico and central Panama and 

FIGURE 1.-Seed coats of Lamourouxia and 

Bartsia, x 30. Center, Bartsia breviflora, Ewan 

16454, US, showing speroidal shape and ridged 

surface. Left, Lamourouxia nelsonii, Pringle 

6777, US, showing characteristic surface, the 

polygonal depressions are closed by a thin 

wall. Right, L. viscosa, Mexia 1321, US, showing open, polygonal perforations of seed coat. 
NUMBER 6 5 

FIGURE 2.-Distributions of twenty-four species of Lamourouxia in Mexico and Central America, 

showing overlapping ranges and discontinuities, compared with distributions of two species in 

South America spanning Colombia, Ecuador, and Peru. 

two species occur in northwestern South America. 

The most common habitat, at least in Mexico, is 

openings in or margins of oak and pine forests. 

At least six species have been collected in or near 

cultivated areas such as maize fields and hedges; eight 

species have been collected on limestone or calcareous 

soil; at least four species have been noted in association with Acacia and Yucca. 

Most collections are from above 1000 m altitude 

(Figure 4) so that low lands, such as the Isthmus 

of Tehuantepec and the Isthmus of Panama are important geographical discontinuities (Figure 2) . The 

most common and widespread species also have the 

greatest altitudinal extremes. The lowest record for 

the genus is for L. viscosa at 450 m in Durango; 

however, this species also is recorded at 2800 m in 

Michoacan. The highest record for the genus is L. 

multifida at 3950 m in the state of Mexico, but this 

species also was collected at 800 m in Sinaloa, indicating an altitudinal range of at least 3000 m. The 

low altitude of 500 m for L. colimae is notable. 

The range of flowering dates (Figure 5) is greater 

for the more common and widespread species. Three 

of these, L. multifida, L. virgata, and L. viscosa, apparently flower throughout the year. For the genus as 

a whole flowering would appear to be most abundant 

from July through October. 

The geographical distributions (Figures 2, 3), 

particularly for the 20 species in Mexico north of 

the Isthmus of Tehuantepec, overlap considerably, at 

least on the scale of this map. Compilations of data 

from herbarium specimens show, however, that seldom has a collector taken more than one species on 

the same day and, in these cases, the localities usually 

appear to be different. A documented exception has 

been brought to my attention by Dr. D. E. Breedlove 

who found L. multifida (section Lamourouxia) and 

L. viscosa (section Hemispadon) at a locality in 

Nayarit and observed the two species being visited 

by the same pollinators. It is difficult to perceive 
6 

SPEC 1 ES 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

’- 300N lstmo 

*2I0N Oaxaca Wan- 1os0w tepec 

I M E x I c o ICENTRALAMERICA~ SOUTHAMERICA 

’ 

111 --*LIT 

brochyonfho 

colirnoe 

dosyontho 

dependens 

dispar 

grocilis 

gutiermzii 

in fegerrimo 

joliscona 

knceoloto 

bngifloro 

macrontho 

microphyllo 

muif ifido 

nelsonii 

ov0fo 

poroyono 

pringlei 

rhinonthifolio 

smifhii 

stemglossa 

sylvofico 

tenuitb/o 

virgofo 

/iscosa 

’OTALSPECIES 

ENDEMICS 

Guats- :hiapas 

I 

6 14 13 (4) 5 7 3 I 2 1 

1-2 6 4 2 I I I 

FIGURE J.-Analysis of geographical distribution of Lamourouxia, showing concentration of 

species in Mexico between 21”N and the Isthmus of Tehuantepec, the four species that cross 

this Isthmus, and the two geographically separated species in South America. 

correlations between geographical distribution patterns and taxonomic characters for the species and 

sections of this genus. Except for South America, the 

species and the sections appear to be geographically 

compatible. Theoretical prerequisites for coexistence 

of closely related species in the same area have been 

outlined by Harper et al. (1960). Comprehensive 

tabulations of altitudinal ranges and of flowering 

dates (Figures 4, 5) are not helpful for understanding the circumstances in individual situations where 

plants of different species of Lamourouxia might 

occur side by side. The few species that I have seen 

in Oaxaca consisted of solitary or a few clustered 

individuals in widely separated localities. Other collectors sometimes have indicated that the plants were 

abundant, and Pringle, near the turn of the century, 

made from 16 to 23 generous duplicates of some 

rather uncommon species, presumably from single 

localities. 

The two species in South America appear to be 

mutually exclusive with very little if any geographical 

overlap, and they are closely allied to two different 

species occurring in central Mexico. These four species 

comprise section Adelphidion, which alone has a discontinuous geographical distribution and tends to 

support “Jordan’s Law” pertaining to geographical 

isolation and speciation (Jordan, 1905; Abrams, 1905; 

Stebbins, 1950). 
NUMBER 6 7 

SPECIES 

braebyantha 

colime 

dasyantba 

dependens 

dispar 

gracifis 

gu tierrezii 

iniegerrima 

jofiscana 

fanceofatu 

longiffom 

macrantba 

microphyf fa 

muftifida 

ne fsonii 

ovata 

pamyana 

pringlei 

rbinantbi fo fia 

smitbii 

stenogfossa 

sylvatica 

tenuifo fia 

virgat a 

viscosa 

xa fapensis 

ALTITUDE IN METERS 

500 1000 1500 2000 2500 3( 

I I& 1- I 

- 03 0 

FIGURE 4.-Altitudinal distribution of the species. 

Hemiparasitism 

Observations and taxonomic conclusions in this paper, 

particularly in regard to morphological variation and 

geographical distribution, should be viewed against 

a somewhat uncertain background of hemiparasitism. 

The heterotrophic way of life for Lamourouxia is 

implied by the systematic association of this genus 

with other Rhinanthoid Scrophulariaceae. The similarity of the seed coats (Figure 1) to those of 

Orthocarpus described by Atsatt (1965) and to other 

known hemiparasites in subfamily Rhinanthoideae 

also substantiates this opinion. The flowers are attractive, but as there are no records of this genus in 

cultivation one assumes that the seeds may have 

special requirements for germination and that the 

plants are not suited for ordinary garden conditions. 

The few living plants that I have seen were luxuriantly green and could have been mistaken at first 

glance for Labiatae. This deceptive appearance probably explains why so little information on substrates 

or hosts has been reported on specimen labels. 

A review of hemiparasitism in Scrophulariaceae, 

including illustrations of representative seeds, appears 

in Kuijt ( 1969). Experiments and observations by 

Atsatt (1970b) and by Atsatt & Strong (1970) provide insight into the biology of Orthocarpus which 

also may be applicable, at least in part, to Lamourouxia. The only general characteristics of hemiparasites are the ability to form rootgrafts by means of 

haustoria and the presence of functional chlorophyll ; 

both are assumed to be advantageous. Rootgrafting in 

Orthocarpus can correct some genetic deficiencies, 

such as restoring chlorophyll synthesis in certain kinds 

of albino plants, through physiological buffering by 

host plants (Atsatt, 1970a). Rootgrafts would appear 

to be formed more or less indiscriminantly between 

roots of the same individual, from one to another 

individual of the same species, and to plants of other 

families in the same community. One, therefore, 
8 

bmchyantha 

calimae 

dasyantha 

dependens 

dispar 

gracilis 

gutierrezli 

integerrima 

jaliscana 

fanceofota 

langiflora 

macrontho 

microphylla 

multifido 

nelsonii 

ovata 

pamyana 

pringfei 

rhinonthifalia 

smithii 

sternglossa 

syfvatica 

tenuifalio 

virgata 

viscosa 

xalapensis 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

r 

- 

-- 

- 

- 

- 

-- 

h- SPECIES J~ FEt 

MONT 

FIGURE 5.-Flowering period of the species with more 

abundant records represented by thickened lines. 

might visualize mixed communities or whole populations of plants that are interconnected and sharing 

water, mineral, and organic resources. Genetic differences among the plants of Orthocarpus range from 

those capable of independent autotrophic growth to 

those completely dependent upon hosts. Populations 

may differ in their heterotrophic and autotrophic 

capabilities. Survival and reproductive potential 

mostly are increased when host plants are present. 

Differences in behavior are to be expected between 

inbreeding and outbreeding plants of Orthocarpus. 

Facultative hemiparasitism in Cordylanthus, a genus 

allied to Orthocarpus and Castilleja, has been discussed by Chuang & Heckard (1971). 

The ability to form rootgrafts perhaps may expand opportunities for survival and sympatric occurrences in Lamourouxia and have taxonomic implications. The standing crop of a species might include 

side by side plants that are variously out of phase 

with the regular breeding system and plants that are 

optimally functional. Extension of the life expectancy 

of plants with reduced or marginal fitness, particularly among perennials, probably enriches the genetic 

and the morphological diversity available for sampling 

by pollinators as well as by botanical collectors intent 

upon preparing herbarium specimens. Taxonomists 

might be well advised to accept broader patterns of 

variation in superficial vegetative and floral morphology for hemiparasitic plants and, as suggested 

by Ornduff ( 1969), strengthen their taxonomic judgments by focusing more attention on understanding 

reproductive methods. 

Vegetative Characteristics 

STEMS AND INFLORESCENCES 

The stems arise from gnarled burls and probably are 

somewhat woody below and brittle above, perhaps 

biennial or perennial. Longer stems may require surrounding vegetation for upright support. Most species 

are upright, the branching excurrent, the branches 

diverging at an acute angle, and the inflorescence 

erect or ascending. 

The inflorescences are terminal, indeterminate, 

racemose, bracteate, and pedicellate. The bracts are 

more or less leaflike, opposite, decussate, each subtending a solitary pedicel without bracteoles. The 

flowers thus are paired and often turned so as to 

appear secund. The inflorescences vary from pubescent to glandular, or glabrous, and are of two kinds. 

Twenty-two species have ascending inflorescences with 

erect tips which are associated with excurrent branching. Three species in section Lamourouxia (L. dependens, L. ovata, and L. xalapensis) and one species 

in section Hemispadon (L. lanceolata) have horizontal or dependent inflorescences with depressed tips 

which are associated with deliquescent branching. 

The pedicels in these species are relatively thin and 

wiry, reflexed away from the apex of the inflorescence, and inverted so that the flowers assume a 

more or less normal dorsiventral position. Pressed 

specimens of the latter often appear to have been 

mounted upside down. Flowers in the species with 

erect inflorescences may be protandrous while those 

with dependent inflorescences may be proterogynous 

(Wagner, 1946). These differences in the position of 

the inflorescences probably are correlated with differences in the breeding system. 
NUMBER 6 9 

LEAVES 

The bracts subtending pedicels and the leaves are 

similar, variable in size and shape, but distinctive and 

provide characters that are essential for separating 

the species. The blades of 18 species are rounded or 

tapered to the base with an indistinct petiole of a 

millimeter or less in length, yet these are not truly 

sessile; however, the leaves on the major axes of 

these species, especially those subtending branches, 

frequently are quite sessile. Definite petioles of more 

than a millimeter occur in five species-all of these 

in section Lamourouxia. In strongest contrast, some 

or all of the leaves of six species are broadly ovate or 

deltoid, cordate basally, definitely sessile, and more 

or less clasping. Petiolate and sessile leaves occur in 

each of the three sections. Many exceptions are found 

to any generalization about the margin, shape, and 

size of the leaves. 

The margins of the blades are as difficult to describe as the petioles; mostly, they are serrate or 

nearly so, less often dentate or crenate, and sometimes more or less doubly incised. In many instances 

the incisions are a blend of two or more states on 

one plant or on one leaf. Leaf margins in about four 

species vary from serrate or dentate to crenate; in 

about six species from serrate or dentate to entire or 

subentire; in about three species from twice dentate 

or serrate or crenate; in one species, L. hyssopifolia 

(Figure 16, see L. longiflora), with entire, lanceolate 

blades there are, on rare occasions, from one to three 

very broad teeth. 

The venation is pinnate and craspedodromus with 

the secondary veins mostly leading directly to the 

sinus between marginal incisions. The primary and, 

less often, the secondary veins stand out below 

(abaxially) and the tertiary veins usually are inconspicuous. Prominent reticulate tertiary veins standing out below usually are characteristic of L. viscosa. 

The venation patterns occasionally are recessed on the 

upper (adaxial) surface. 

The most unusual leaves, those of L. multifida, are 

deeply pinnately divided with several nearly linear 

or narrowly lanceolate divisions which sometimes are 

again pinnately parted (Figure 16) , Divided leaves 

also occur in other genera of Scrophulariaceae, as in 

Aureolaria, Leucospora, Macranthera, and Seymeria. 

The parasitic Scrophulariaceae are the only parasitic Angiosperms having variously divided leaves 

(Kuijt, p. 82, 1969). The leaves of 20 species of 

Lamourouxia are 5 or fewer times as long as broad; 

7 species have leaves 6 to 15 times as long as broad. 

Basal leaves usually are missing from herbarium specimens and there is no indication of juvenile conditions. 

The leaves are similar to, and grade into, the bracts 

that subtend the flowers, mostly being larger below 

and smaller above. 

Floral Characteristics 

CALYX 

The calyx is united, cup-shaped below, and fourlobed (Figure 6). The symmetry varies from nearly 

regular to very strongly bilabiate; the surface from 

pubescent, sometimes glandular, to glabrous ; the lobes 

from deltoid to subulate, and from entire to incised. 

A fifth calyx lobe is rare. A median vein leads to 

each lobe and one or two veins lead to the sinus, the 

latter sometimes varying within a species or within 

a single calyx. When there is one vein it branches at 

the sinus; when there are two, the veins join just 

below the sinus and send a branch off to the lobes on 

either side; thus, each lobe of the calyx receives three 

veins. The major calyx veins sometimes appear to be 

overlain externally by a fleshy rib. The axil of the 

sinus sometimes is revolute, protruding outward like 

a small lip and sometimes there is an anomalous, 

small, wartlike appendage situated in the axil (L. 

dasyantha) . Characteristics of the calyx often seem to 

be correlated with those of the leaves and bracts in 

regard to pubescence, shape, and margin, and may 

be useful for separating species. In those instances 

where there are taxonomic problems between similar 

species, the characteristics of the calyx usually are not 

helpful. The calyx is persistent and may enlarge 

somewhat with the maturation of the capsule (Figure 

28). 

COROLLA 

The corollas are horizontal or inclined, bilabiate, 

bilaterally symmetrical tubes which are variously deformed or ventricose below (Figure 8). All are pubescent externally, sometimes also glandular, and the 

color of the hairs may contrast with that of the 

tube; some corollas also may be pubescent internally. 

During aestivation the lower lip enfolds the upper. 

Cleared flowers show that the basic venation pattern 

is 5-merous (Figure 7). Venation in the receptacle 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 6.--Sample diagrams of floral features. a, L. viscosa, seed, 1.5 mm long, showing 

reticulated coat; 6, same, with endosperm containing embryo removed. c, L. macrantha, transverse section of young fruit after abscission of corolla showing placentation; d, same, longitudinal section, x %. e, L. microphylla, calyx, showing simple venation. f, L. dasyantha, calyx, 

showing double and single venation to sinuses with short, anomalous appendages in axiles of 

sinuses. g-i, Introrse views showing diversity in basal mucros of anthers: g, L. multifida; h, L. 

gutierrezii; i, L. dependens. j-m, Extrorse views of anthers showing diversity in connectives: 

j, L. macrantha; k, L. virgata; 1, L. tenuifolia; m, L. nelsonii. Pubescence not shown on anthers. 

n-t, Typical hairs from middorsal portion of corolla tube except “0”: 0-q, gland-tipped hairs: 

n, L. brachyantha; 0, same, except from calyx; p, L. multifida; q, L. viscosa; r, L. virgata; 

s, L. lanceolata; t, L. microphylla. 

below the insertion of the floral appendages has not 

been explored. The lips are dissimilar and somewhat 

different from species to species (Figures 7, 12). The 

upper lip is more or less two-lobed; the lobes usually 

are reflexed upward. Whether or not two-lobed, the 

upper lip always is veined as though two-lobed with 

each lobe receiving a prominent median vein and a set 

of laterial veins, all originating independently from 

the receptacle (Figure 7). Angular, secondary, cross 

venation unites most of the major veins. The upper 

lip at first is galeate, embracing the anthers on the 

longer filaments. In all but three or four species the 

upper lip continues to shelter the anthers after anthesis, and in most species, because of the lobing, the 

upper lip tends to separate the stigma from the 

anthers. After anthesis the corollas of only L. brachyantha remain conspicuously galeate ; the others, 

although always strongly bilabiate and somewhat 

naviculiform, usually are no more geleate than corollas 

of other sympetalous families. The upper surface of 
NUMBER 6 11 

the lower lip usually is centrally creased with a groove 

that leads deep into the corolla chamber. On either 

side of the crease the lower lip is deformed upward 

into the palate which is sufficiently large in L. 

dasyantha and L. syluatica to partially or completely 

occlude the mouth of the corolla. 

STAMENS 

There are two pairs of stamens (Figure 7). Robinson 

& Greenman (1895, p. 165) reported a rudimentary 

fifth stamen but I have never observed one. In some 

species there may be a few extra veins in the median 

position of the upper lip of the corolla, the expected 

position for a fifth stamen; however, these veins appear only to be additional supply to the corolla. The 

filaments are epipetalous, persistent, and inserted at 

approximately the same level near the base of the 

corolla. At their insertion, the filaments usually are 

swollen, pubescent, probably nectariferous, and frequently somewhat geniculate. Each filament receives 

one vein from the receptacle. The upper pair of 

filaments, as the flowers frequently are nearly horizontal, varies from slightly shorter to much shorter 

than the lower filaments. In 10 species (section 

Lamourouxia) the 2 sets of filaments are subequal; 

in 4 species (section Adelphidion) the anthers on the 

1 

shorter filaments are smaller but mostly fertile; in 12 

species (section Hemispadon) the anthers on the 

shorter filaments mostly are rudimentary with 1 or 

2 unequal locules, usually sterile, or absent (Figures 

8, 14). The shorter, upper filaments are inserted 

below the sinuses separating the lower from the upper 

lip of the corolla. The longer, lower filaments often 

are more thickly developed, with somewhat larger 

anthers and broader connectives, and are inserted 

below the sinuses between the lobes of the lower lip 

of the corolla (Figure 7). 

Fully fertile anthers are basifixed, usually shaggy 

pubescent, and bilocular at maturity. The Iocules 

are more or less equal in size, completely separated 

by the connective, laterally dehiscent, and often 

spurred or mucronate below (Figure 7). The fertile 

anthers usually are more or less vertical, turned 

nearly at right angles to the filaments, and face 

inward in pairs so that corresponding locules are 

appressed. The meshing of the pubescence in most 

species appears to hold the introrse anthers pairs 

together and also to hold or direct the pollen downLvard and away from the stigma. The larger anthers 

of most species are usually at least partially visible 

near or just beyond the mouth in the mature flowers 

(Figure 8). In some species, long, uniseriate hairs 

originating on the inside of the corolla become eni 

i 

\ 

\ 

\ 

FIGURE 7.-Diagrams of major venation patterns of corollas, reduced to same size, distorted by 

spreading toward base. 4, L. nultifida, based upon Skutch 580, US. b, L. lanceolata, based 

upon Skutch 4206, US. c, L. sylvatica, based upon Pennell d Ferreyra 14924, US. d, Fully 

developed lower stamens; d', fully developed upper stamens; d", small, fertile, upper stamens; 

d"', rudimentary upper stamens. e, Median traces to lower lip; e', main traces to upper lip. 

f, Palate region, stippled. 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

LAMOUROUXIA HEMISPADON 

”u-/ ADELPHIDION 

tangled in the hairs of the anthers, thus partially 

obstructing the mouth of the corolla and perhaps also 

holding the anthers in a definite place. The spurs on 

the lobes of the anthers probably ensure dorsal contact with pollinators; pressure against the spurs probably tends to separate the anthers and to facilitate 

the downward sifting of the pollen. In three to five 

species the anthers become exserted and spreading 

(Figure 21 ) and lobes of the anthers are more nearly 

obtuse rather than mucronate. In two species, L. 

brachyantha and L. sylvatica, the larger pair of 

anthers always remains included (Figure 8) . 

OVARY CAPSULE 

The ovary (Figure 6) is two-locular with axile 

placentation, ovoid, attenuated apically, slightly compressed laterally, and, as the flower lies nearly horizontally, one locule is above, the other below. The 

ovary sometimes seems to rest upon a solid, short pedicel that at times appears notched and perhaps is 

secretory; or the ovary is sessile with thickened walls 

toward the base. A disk has not been observed. The 

ovary, which only rarely is pubescent, tapers upward 

into a filiform style which often is sparsely pubescent. 

The style is situated above the stamens in the horizontal flower and curves distally, pointing the stigma 

downward just beyond the anthers. Even in the buds 

the stigma exceeds the stamens. The stigma is delicate 

and difficult to study in pressed specimens, appearing, at times, conical, clavate, or unequally two-lobed. 

The delicate, filiform style remains attached to the 

FIGURE 8.-Diagrams of cleared flowers comparing 

development and position of stamens and stigma in 

the three sections of Lamourouxia. Only one anther 

of each pair shown; pubescence omitted; XI, except 

L. brachyantha and L. sylvatica, x 2. Left, section 

Lamourowia, two pairs of fully developed anthers 

with subequal filaments; center, section Adelphidion, 

one pair of large anthers and one pair of smaller 

anthers; right, section Hemispadon, one large pair of 

anthers and one pair of sterile filaments. 

ovary as the corolla disengages and falls away. 

Herbarium specimens frequently show the styles attached to the capsules but this must be of short duration in nature. In some cleared specimens there is a 

change in the density of the tissues which suggests 

a predetermined place where the style breaks from 

the ovary; however, an abscission line has not been 

observed. The capsules are incompletely loculicidally 

dehiscent from above, splitting through the base of 

the style (Figure 24). The capsules are attenuated 

apically into a flattened and usually crooked beak 

which is the base of the style (Figure 28). 

SEEDS 

The ovules are numerous and radiate from the 

two axile placentae (Figure 6). The seeds are tightly 

packed, narrower at the base, broader and truncated 

distally. The shape is more or less irregularly rectangular to oblong, about twice as long as broad 

(Figure 1). The surface is deeply reticulated, with 

a lengthwise suture probably associated with the 

funiculus, and sufficiently translucent to show the 

opaque, narrowly oblong envelope of endosperm 

round the small, straight, dicotyledonous embryo 

(Figure 6). In some but not all samples of seeds 

from L. viscosa the hollows of the reticulated seed 

coat become open, honeycomblike perforations resembling a loose net around the embryo. These seeds 

are similar to those of Orthocarpus densiflorus described by Atsatt (1965) in his account of a “coordinated seed dispersal mechanism” which may be 
NUMBER 6 

applicable in part to Lamourouxia and to some 

other Scrophulariaceae. In this case the seeds of 

Orthocarpus at the study site became impaled on 

the awns of the achenes of Hypochoeris glabra, presumably ensuring that the seeds of the hemiparasite 

and host are together for dispersal and germination. 

Generally more than one seed of Orthocarpus became impaled on the several awns of a given 

achene of Hypochoeris, apparently providing an opportunity for clustering the seeds of the self-incompatible Orthocarpus prior to dispersal and germination. The seeds of other species of Lamourouxia 

vary slightly in appearance and the hollows of their 

reticulated coats are closed by a fragile, porous wall 

rather than being completely open. 

HAIRS 

Pubescence, often one of the most conspicuous features of a specimen, nevertheless is diffcult to characterize quantitatively for any taxon. Qualitatively, 

however, the hairs are uniseriate, sometimes arising 

from a multicellular platform (Figure 6), but otherwise falling into two major categories: simple, and 

branched or tufted. Both kinds of hairs are sometimes also gland tipped. Grandular hairs are terminated by a globose or flattened gland which usually 

is on a narrow neck cell. The glands, which may 

be composed of more than one cell, are difficult to 

study and are generally similar to the one shown 

by Wettstein (Pflanzenfam. IV. 3B. (65) :42, Fig. 

18-C. 1891). Occasionally there are obscure, very 

minute, sessile or very short stalked or “peltate” 

glands, also shown by Wettstein (p. 42, Fig. 18-E 

and F), and by Metcalf & Chalk (p. 980, Fig. 

228-C and F. 1950), which seem different and even 

more difficult to study than the more conspicuous 

hairs. These minute hairs or glands have been observed on corolla, calyx, and leaves in Lamourouxia, 

but otherwise have been ignored in this present 

paper. 

Simple hairs may occur almost everywhere, but 

branched or tufted hairs seem only to occur on the 

corollas. The calyx, bracts, and leaf blades usually 

agree in characters of pubescence (Table 1). Some 

specimens, at least, of almost all taxa exhibit cauline 

pubescence. Those that are abundantly pubescent on 

leaf and calyx often have conspicuously pubescent 

stems; those that are glabrate or glabrous may have 

cauline pubescence confined to paired, narrow, decurrent, internodal strips-the condition that is called 

bifarious. Organs which seem glabrous sometimes are 

microscopically pubescent, often in sheltered places 

such as exiles of serrations on leaf margins, or along 

the inner margins of calyx lobes, or at the union 

of the calyx and pedicel. 

Excluding the corolla and its internal organs, 10 

or 12 species are glabrate to glabrous; most of the 

pubescent species sometimes are glabrate; 3 species 

(L. multifida, L. syluatica, L. uirgata) seem to run 

the gamut from pubescent to glabrous; 4 species (L. 

brachyantha, L. jaliscana, L. nelsonii, L. uiscosa) are 

conspicuously glandular; 4 or 5 additional species 

are often glandular; 5 species are mostly scabrous; 

and 3 other species sometimes are scabrous (Table 

1). Scabrous hairs appear to be simple, short, often 

curved or appressed, and with heavily thickened 

wall. Inconsistent variation in abundance and quality, 

and whether glandular very seriously limits the systematic and taxonomic usefulness of the hairs, especially on vegetative portions of the plants. 

A little additional information can be gleaned 

from study of the hairs on the corollas when these 

are compared from closely equivalent areas (Table 

1). All corollas are conspicuously, even if not always 

copiously, pubescent. Branched hairs are thought to 

be unique to the corollas, and, depending upon the 

taxon and the place on the corolla examined, 

branched as well as simple hairs and glandular hairs 

may occur side by side. Glandular hairs, when they 

are present, usually are more abundant near the 

mouth of the corolla. Hairs appearing on the vegetative portions of the plants usually appear more or 

less white or colorless, although the glands may be 

dark; however, the corolla hairs usually are darkly 

or reddish colored. 

Even though the hairs on the corolla vary from 

place to place in structure, appearance, abundance, 

and in whether glandular, comparisons of the midupper portion of the surface of the corolla tube 

show some similarities which tend to correlate with 

the clustering of the species, These observations 

apply only to this region of the corolla tube and, 

needless to say, no two specimens are exactly the 

same. The species nearest to L. multifida, section 

Lamourouxia (eg, those above line 2 in Figure 9), 

mostly have short, simple hairs, often rather sparse, 

which frequently terminate in a gland (Figure 6). 

The species closest to L. xalapensis (in the same 

section, e.g., those species below line 2 in Figure 9), 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

TABLE 1,-Distribution of hairs with and without glands on mid-upper portion of 

corolla tube, calyx, and leaves, compared for the three sections of Lamourouxia. 

Exceptions are common in almost every taxon; outstanding exceptions are indicated 

by the open circles. 

HAIRS 

Mid-upper portion of 

COROLLA TUBE 

Hairs mostly simple, 

often glandular 

Hairs mostly branched, 

arms mostly straight, 

cells elongated, apex attenuate, mostly not 

glandular 

Hairs mostly branched, 

arms straight or curved, 

mostly not glandular 

Hairs for cluster, diverse, mixed, often 

glandular 

Hairs mostly intricately 

branched, often appearing tufted, the cells 

short, apical cells mostly 

rounded, mostly not 

glandular 

CALYX 

Pubescent, mostly not 

glandular 

Mostly glandular 

Mostly glabrous or glabrate 

LEAVES 

Pubescent, mostly not 

glandular 

Mostly glandular 

Mostly glabrous or glabrate 

Mostly scabrous 

LAMOUROUXI A 

mom a 

aaaam 

0. a a 

a. a. 0 

a a a 

a. 0 

0 a m 

a. 

ADELPHIDION 

0 

D 

ac 

0 

D 

ac 

DO 

HEMISPADON 

ma00 

aaaaaa 0 

0 

a. a. 

aa0.a. 

a 

a 

a. a. 

aaoaoa 

.O 0 a 
NUMBER 6 15 

mostly tend to have longer hairs with longer, narrow 

cells; branching is common from below with long. 

straight arms; glands are very rare or absent and 

the terminal cell usually is attenuated. It must be 

noted that in L. dispar, section Lamourouxia, the 

hairs are not glandular, repeatedly branched, tending 

to be matted, and the cells are relatively short, thus 

resembling the species below line 2 in Figure 11, 

section Hemispadon. 

The species nearest to L. viscosa, section Hemispadon (e.g., most of those above line 2 in Figure 11) , are mixed or less easy to characterize. Glandular and 

nonglandular, simple and branched hairs often stand 

side by side; branching mostly is from below (Figure 

6). The species nearest to L. Zanceolata, same section 

(e.g., those below line 2 in Figure 1 1 ) , are much more 

distinctive. Glands are absent and the cells tend to be 

about as long as broad, often appearing like a string 

of beads, and the terminal cells usually are rounded. 

These hairs often appear tufted, frequently are intricately branched with short, curving arms, the arms 

often branching a second time (Figure 6). In this 

section, Hemispadon, L. jaliscana is peculiar since the 

hairs are usually simple, long, and narrow, with attenuated apexes, some are branched from below, thus 

recalling species in section Lamourouxia such as L. 

xalapensis. A few hairs also are glandular, somewhat 

as in L. multifida. 

The hairs at equivalent positions of the corolla 

tubes of section Adelphidion lack glands. Those of 

L. sylvatica are straight, relatively long, frequently 

simple, unbranched, and sparse. Those of L. brachyantha are dense, mostly branched from below, and 

have rather long, straight arms. The hairs of L. dasyantha are sparse while those of L. uirgata (Figure 6) 

are dense, but otherwise are similar, intricately 

branched, and the cells and arms are rather short. 

The relative selective value of the kinds, abundance, and color of the corolla hairs is by no means 

clear, especially when simple hairs are contrasted with 

branched hairs. Pigment or color in the corolla hairs, 

depending upon their abundance, undoubtedly affects the corolla color and perhaps the attractiveness 

of the flowers to pollinators. Glandular hairs, especially near the corolla mouth, possibly might discourage some visitors from entering. For systematic purposes, the hairs of the corolla lend support to the 

clustering of the species. In a few instances they are 

contradictory as in L. dispar and L. jaliscana. 

Comparative Morphology 

Independently of vegetative or other floral characteristics, the species can be clustered objectively to form 

three sections according to the development of the 

stamens (Figure 8). The 10 species of section Lamourouxia have 2 pairs of similar anthers; the 12 species 

of section Hemispadon have 1 pair of anthers and a 

pair of sterile stamens; the 4 species of section Adelphidion are intermediate with 2 dissimilar pairs of 

anthers. Shape and color of the corolla are correlated 

with the development of the stamens so that two 

pollination syndromes are represented. The flowers of 

the first two sections have long, narrow, red corollas, 

and are presumed to be pollinated by hummingbirds 

(Figures 16, 21, 28, 34). The flowers of the third 

section have shorter, broader, non-red corollas, and 

are presumed to be pollinated by bees (Figures 8, 

24). The differential development of the stamens 

provides a morphological series (Figure 14) in which 

the bee-flowered species appear to be intermediate 

between the two sections of bird-flowered species and 

suggests an explanation of how there can be two 

sections with flowers that are similar in appearance 

and function. 

SECTION LAMOUROUXIA 

The flowers in this section have two subequal pairs 

of stamens (Figure 8). The four, fully fertile anthers 

are similar in size and shape, inserted on similar filaments, and the lower pair slightly exceeds the upper 

in length. The corollas are vivid red, and suggest the 

syndrome of ornithophily (Faegeri & van der Pijl, 

1966). These are the unifying characters of this 

section. 

The corollas show two extremes in silhouettes which 

are correlated with the position of the anthers and 

with characters of the inflorescences. The tubes of the 

corollas of L. multifida, typical also of L. longiflora, 

L. parayana, and L. pringlei, are more or less narrowly 

funnelform, usually conspicuously ventricose, somewhat narrowed at the mouth, and the flowers are 

displayed in ascending racemes. The larger upper lip 

shelters the mouth and the four partially exposed, 

mutually connivent, shaggy, and strongly mucronate 

anthers. The forward margins of the upper lip are 

slightly lobed or not lobed, slightly reflexed or not 

reflexed, and appear more or less inverted naviculiform. The mouth is somewhat obstructed by the 
16 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

connivent anthers and the narrow, lower lip tends 

to arch downward abruptly (Figure 16). 

The other extreme is exemplified by L. xalapensis, 

which also is typical of L. dependens and L. ovata. 

The corollas in this case have tubes more broadly 

funnelform, not ventricose or only slightly so, sometimes concave below, and they are broadest at the 

mouth. The rounded lobes of the upper lip are reflexed upward, the anthers are exserted, spreading, 

less pubescent, and slightly or not mucronate. The 

broad lower lip usually juts forward with spreading 

lobes, and the gaping mouth is unobstructed (Figures 

8, 21). This latter group of species, including L. 

xalapensis, is remarkable because the inflorescences 

are depressed or dependent rather than ascending. 

This appears to be correlated with deliquescent growth 

and nearly right angle branching. The wiry pedicels 

permit the flowers to invert, thus assuming the normal 

dorsiventral position. It would appear that the species 

closest to L. xalapensis differ in some important way 

from those closest to L. multifida. There is a suggestion in a study of hummingbird behavior (Wagner, 

1946) that the flowers of “L. exserta” (=L. xalapensis) are proterogynous; thus, one might speculate 

that the flowers of the first group, including L. multifida, with their erect inflorescences, may be protandrous. This, of course, ought to be verified. The dependent inflorescence occurs also in L. lanceolata of 

section Hemispadon. 

The intervening species detract from the distinctness of the above two extremes in floral morphology 

which otherwise could have been separated taxonomically into two sections or subsections of coordinate 

rank. The distribution of characters in the intervening 

species (Figure 9) links all of them into a more or 

less continuous morphological series, so that they seem 

more like one section. Various combinations of characters suggest a close alliance among L. macrantha, 

L. ovata, and L. dependens. A taxonomic problem 

unexpectedly links L. macrantha with L. stenoglossa 

in Guatemala. As the name implies, L. dispar is enigmatic because some of its characters suggest a relationship with section Hemispadon while others, especially the four subequally developed stamens, indicate 

a central position within section Lamourouxia. These 

problems are discussed in the taxonomic portion of 

this report following the species descriptions. 

The distribution of some vegetative characters in 

section Lamourouxia is shown in Figure 9. Some 

LAMOUROUXIA 

3 

,,......---. .._ .--__._ ____ ,,?, ; PARAYANA . ..__._~______--_--.--_-_.___-.__

4‘ 

I 1- 

2= 

‘\ , -. - - - . . ~ . . - ~ 

.I ., 

“s,~ 7’ OVATA 1 XALAPENSIS 

\ L--www 3 .--.-. --.__.___ 1. ’-. : .-.-; DEPENDENS 1 

FIGURE 9.-Approximate systematic position of species in 

section Lamourouxia showing distribution of taxonomic 

characters. Above line 1 the anthers tend to be mutually 

‘connivent and barely exposed in the mouth of the corolla; 

below the line they tend to be exserted and spreading. Above 

line 2 the plants are upright with excurrent branching and 

ascending inflorescence tips ; below the line the plants are 

dependent, with deliquescent, often right-angle branching, 

and the inflorescence tips are depressed with correspondingly 

inverted flowers. To the left of line 3 the leaf blades tend 

to be ovate; to the right lanceolate. Line 4 indicates species 

tending to have glandularly pubescent calyx and leaves. 

Species 5 has unique, pinnately divided leaf blades. Species 

6 has broadly cordate leaves on short petioles. Species 7 has 

broadly cordate, sessile leaves. 

vegetative characters tend to reinforce the grouping 

of the species while others only emphasize the diversity 

within the section. Line 1 divides the species on the 

basis of characters of the anthers. Line 2 divides them 

again on the basis of the inflorescence. Line 3 divides 

the species according to whether the leaves are mostly 

ovate or mostly lanceolate. Although individual specimens of any of these species may vary from pubescent 

to glabrate or glabrous, the ones enclosed by line 4 

mostly are conspicuously pubescent and the hairs 

mostly are glandular. The leaves of species 5, L. 

multifida, are exceptional because they have relatively 

long petioles and the blades are pinnately divided into 
NUMBER 6 17 

narrow segments. The leaves of species 6, L. macrantha, are exceptional because they are large, short 

petiolate, broadly ovate, and deeply cordate at the 

base. The leaves of species 7, L. ovata, are very similar to those of L. macrantha, except that they are 

sessile. All other species in this section are petiolate 

with the blades rounded or tapered to the base. 

Although the petioles sometimes are less than a millimeter in length the blades are not truly sessile. Another 

character, not indicated in the figure, is the distinctive 

double or irregular leaf margins, which vary in some 

combinations of serrate-dentate-crenate, for L. macrantha, L. stenoglossa, L. ouata, and L. dependens. 

This character of the leaf margins occurs among 

relatively interrelated species whose geographical distributions center on the margins of the Isthmus of 

Tehuantepec (Figure 22). The margins in the other 

species are simple serrate or dentate or crenate, except 

for the margins of L. longiflora and the margins of 

the narrow division in L. multifida, which are entire. 

The restricted distribution of L. longiflora is contiguous with that of L. multifida, and the two are predicted to be relatively closely related (Figure 17). 

Six species only occur in Mexico northwest of the 

Isthmus of Tehuantepec; two species, L. stenoglossa 

and L. dependens, only occur on the southeast side of 

the Isthmus; and two species, L. multifida and L. 

macrantha, occur on either side of the Isthmus. The 

Isthmus of Tehuantepec is particularly notable in 

regard to the distribution of section Lamourouxia 

(Figure 3). The three species with dependent inflorescences, L. dependens, L. ovata, and L. xalapensis, have distributions more or less neighboring 

around this Ithmus, the species being restricted to the 

one side or to the other (Figure 22). The one species 

in section Hemispadon with the same inflorescence 

character, L. Zanceolata, also shares this distribution 

but occurs on either side of this Isthmus (Figure 30) 

A few specimens of L. dependens from Guatemala 

have unusually small calyxes which also tend to be 

glabrous, thus causing these specimens to be confusingly similar to L. xalapensis which only occurs to 

the west of the Isthmus. West of the Isthmus, L. 

macrantha is among the most distinctive species; to 

the east, especially in Chiapas, L. stenoglossa is remarkably distinctive. Both of these species also Occur 

in Guatemala, where their distinguishing characters 

appear to become blended in some specimens (Figure 22). 

SECTION ADELPHIDION 

The stamens in this section are dimorphic (Figure 8). 

The longer pair have thicker filaments and larger anthers which are pubescent and mucronate. The upper 

pair are half or two-thirds as long, the filaments are 

thinner, the anthers smaller, less pubescent, attentuate 

basally, always included, mostly two-locular at anthesis, and usually fertile. The corollas are pink, lavender, exceptionally magenta, purple or white, and 

suggest the syndrome of melittophily (Faegeri & van 

der Pijl, 1966), The tubes tend to be broadly funnelform, broadest toward the mouth, and slightly or not 

at all ventricose, with the lobes of the lips generally 

broadly rounded, and the mouths relatively large, 

although mostly occluded by large palates in two 

species. These are the unifying characters of this 

section (Figure 24). 

Several characters clearly mark L. brachyantha as 

exceptional. The development of the stamens and the 

color of the corolla are the reasons for placing it in 

the new section Adelphidion where it seems remote 

from the other species. The corollas are conspicuously 

galeate with a large, open, downward-facing mouth. 

The large pair of anthers is included and the stigma 

is slightly or not at all exposed. 

The corolla mouths are relatively large in the 

remaining three species but almost or completely 

obstructed by the large palate in L. sylvatica and 

L. dasyantha, and less so in L. virgata. The large 

pair of anthers and their thickened filaments are 

partially exposed in the mouth of L. virgata, thus 

recalling section Hemispadon. The large anthers usually are less exposed in L. dasyantha and included in 

L. sylvatica. The stigma is exposed in L. virgata and 

L. dasyantha, but little or rarely exposed in L. 9’1- 

vatica. The tendency for included anthers and the nonred corollas of L. brachyantha, L. sylvatica, and L. 

dasyantha suggest the syndrome of melittophily in 

which the pollinators bodily enter the corolla chamber. 

The corollas of L. virgata usually are longer and sometimes approach red in color. The morphology of the 

corollas of L. uirgata, more than any other species, 

suggest that they may be pollinated by bees as well 

as by birds. The small included pair of anthers in 

L. virgata mostly are bilocular and fertile, but occasionally they are rudimentary and sterile, again recalling characters of section Hemispadon. 

The distribution of vegetative characters in section 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Adelphidion (Figure 10) is less instructive because 

there are so few species to examine. Two distinctive 

features characterize L. brachyantha: the leaves are 

broadly ovate, cordate, sessile, and clasping, and the 

plants are densely glandular pubescent. This combination of characters recalls some species in section 

Hemispadon, and probably influenced Greenman 

(1905) to place L. brachyantha next to L. rhinanthifolia. The leaves of the remaining species in section 

Adelphidion mostly are Ianceolate, narrowed to the 

base, mostly glabrate, and not glandular, The leaves 

on the main axis, particularly those subtending 

branches, are sessile. A number of exceptions occur in 

L. sylvatica where the leaves sometimes are broadly 

lanceolate to ovate, occasionally pubescent, and rarely 

the hairs are glandular. Plants of L. dasyantha frequently are well branched from near the base, and 

sometimes are described as tufted in appearance. 

Two quite dissimilar species, L. brachyantha, which 

is uncommon and restricted, and L. dasyantha, more 

common and widely distributed, occur in central 

Mexico (Figure 25). Two other species, L. sylt~atica 

FIGURE 10.-Approximate systematic position of species in 

section Adelphidion showing distribution of taxonomic characters. Above line 1 the leaves are sessile, cordate, deltoid, 

and glandular pubescent; the calyx always is serrate, glandular pubescent; and the corolla always is strongly galeate. 

Below line 1 the leaves are tapered or rounded to a petiolelike base, and mostly glabrous and nonglandular; the calyx 

is variable; the corolla is little or not galeate. Above line 2 

the species occur in central Mexico; below, in South America. To the right of line 3 the stigmas and anthers always 

are included; to the left, the stigmas are exserted and the 

larger pair of anthers usually is at least partially exposed 

in the mouth of the corolla; and often there are small 

appendages in the sinuses of the calyx lobes. Line 4 indicates 

the palates are well developed and more or less occlude the 

mouth of the corolla. 

and L. virgata, occur in northwestern South America, 

are about equally common, and replace one another 

geographically with very little or no overlap in distribution (Figure 27). There is rather close, although 

possibly coincidental, similarity between L. dasyantha 

and L. virgata. 

SECTION HEMISPADON 

The flowers in this section have very strongly dimorphic stamens (Figure 8). The lower pair are 

longer, with broad connectives and thickened filaments. Their anthers are large, shaggy pubescent, 

connivent, short mucronate, and at least partially 

visible beneath the upper lip of the corolla, The upper pair of stamens are short, sterile, and filiform. Infrequently there are deformed anthers of one or two 

unequal locules on these short filaments which rarely 

contain a small amount of pollen. The corollas are 

red, very exceptionally pale orange or pink. The tubes 

tend to be more nearly cylindrical, indistinctly or not 

ventricose, and often slightly narrowed at the mouth 

(Figures 28, 34). Instead of a gaping mouth where 

the lips become separated, the proportionately long 

and narrow lips tend to continue the line of the 

tube. The relatively long, parallel portions of the 

two lips are an inconspicuous but regular characteristic of this section with the single exception of L. jaliscana. The upper lip shelters the mouth and the two 

large, mucronate anthers. The small palate is displayed some distance from the true mouth, usually 

as a slight construction at the base of the short lobes 

of the lower lip. These species have relatively uniform 

flowers which at best could be separated into groups 

with slightly broader and narrower corollas. The 

leaves are of two general kinds, those which are 

sessile and truncated to cordate basally, and those 

which are tapered to the base, thus appearing as 

though petiolate. 

The distribution of vegetative characters in section 

Hemispadon is shown in Figure 11. Line 1 separates 

the species with sessile, mostly cordate and clasping 

leaves from those below the line which appear as 

though petiolate, having blades tapered or rounded 

to the base. Line 2 separates the species which tend 

to be heavily pubescent and mostly glandular from 

those below the line which mostly are glabrate or 

glabrous and not glandular. The leaves, stems, and 

calyx of species 3, L. gutierrezii, are densely scabrous 
NUMBER 6 19 

-1 

=2 

H EM IS PAD 0 N 

:s VISCOSA 

4 

1 ( 

A\. 

RHlNANTHlFOLlA I': 

INTEGERRIMA 

5 LANCEOLATA 

,-_-_- ... . ..__ , 4 *, ' 

COLIMAE GUTlERREZll ?' * . - - . - . - -' 

FIGURE 11 .-Approximate systematic position of species in 

section Hemispadon showing distribution of taxonomic characters. The leaves above line 1 are mostly sessile and clasping; below they are more or less petiolate and tapered or 

rounded to the base. The species above line 2 mostly are 

glandular. Line 3 indicates the species with conspicuous 

pubescence on leaves, stems, and calyx. To the left of line 4 

the leaves mostly are lanceolate, to the right they mostly are 

ovate. Species 5 has dependent inflorescences, all others are 

erect. The species within line 6 mostly have proportionately 

broad corollas; outside the line they are narrower. 

pubescent. This would seem to link this species to 

the L. uiscosa complex; however, the plants of L. 

gutierrezii are totally devoid of glands and the leaves 

are petiolate, which is the reason this species is placed 

in the glabrous-glabrate complex near L. lanceolata, 

L. colimae, and L. microphylla. Line 4 separates the 

species with primarily lanceolate leaves from those 

mostly with ovate leaves. In this section, species 5, 

L. lanceolata, having deliquescent growth, dependent 

inflorescences, and resupinate flowers, is unique ; however, there are three species with similar characters in 

section Lamourouxia. All other species in section 

Hemispadon have excurrent growth and ascending 

inflorescences. Line 6 shows those species which tend 

to have broader corollas in proportion to their lengths. 

The two most common and most widely distributed 

species are L. uiscosa, mostly with narrow corollas and 

glandular calyx, and L. rhinanthifolia, mostly with 

broader corollas and nonglandular calyx (Figure 3). 

These species normally are quite distinctive but at 

times have overlapping characteristics, perhaps indicating hybridization, and they also appear to be 

closely allied with L. nelsonii and L. smithii, both 

rather rare and restricted (Figures 29, 30). Various 

combinations of characters mark L. gutierrezii, L. 

jaliscana, and L. lanceolata as exceptional. The last 

two, although clearly linked to Hemispadon, have 

some characters in common with section Lamourouxia. Three species, L. gracilis, L. tenuifolia, and 

L. integerrima, although not appearing very closely 

similar, have relatively large and broad corollas. 

These last three species share elongated or lanceolate 

leaves with L. lanceolata, and this last species seems 

to intergrade with L. integerrima in Guatemala. Although there are strong differences in the leaves, 

L. lanceolata, L. colimae, L. gutierrezii, and L. microphylla tend to have similar, relatively narrow corollas. 

Eight species occur only in Mexico north of the 

Isthmus of Tehuantepec; L. lanceolata and L. viscosa 

cross this Isthmus; L. integerrima and L. gutierrezii 

are restricted to Central America (Figure 3). 

Floral Function and Pollination 

Floral morphology exhibits a myriad of subtile to 

conspicuous variations, for example, in the degree of 

lobing of the lips of the corollas (Figure 12) and 

whether or how the lobes are reflexed. The abundance 

of these minor variations among and especially within 

species negates their value for taxonomic purposes 

and also suggests that they are of little or no evolutionary significance. Characters of the calyx, like some 

vegetative characters, such as shape and marginal 

incisions of the leaves or lengths of petiole, sometimes 

are helpful taxonomically but probably also are of 

little or neutral selective value. Those floral characters directly involved in the process of pollination, on 

the other hand, such as color of the corolla and 

development of the stamens provide the criteria for 

clustering the species into sections and reaffirm the 

observation that the process of pollination gives meaning to floral morphology (Straw, 1956; Thompson & 

Ernst, 1967; see also Carlquist, 1970). The most 

important taxonomic characters in Lamourouxia also 
20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 12.-Sample diagrams of opened corollas of Lamourouxia comparing characters of the 

upper and lower lips which are considered to be of relatively low selective value, section 

Lamourouxia to the left (omitting L. ovata and L. parayana), section Adelphidion central and 

stippled (L. brachyantha shown folded because it is so strongly galeate), section Hemispadon 

to the right. 

are biologically significant. 

The corollas are bilabiate, horizontal or inclined, 

bilaterally symmetrical tubes. The lips are dissimilar 

and somewhat different from species to species (Figure 

12) ; however, the upper lip tends to shelter both the 

mouth and the anthers in most species. The corolla 

tapers to a narrow cylinder around the ovary and 

the stamina1 filaments are inserted at the same level 

on the corolla within a few millimeters of its base. 

At their insertion the filaments are thickened, geniculate, frequently densely pubescent, and probably secretory. The corolla wall at the insertion of the stamens 

frequently is pubescent. The base of the ovary sometimes also appears as though secretory. The young 

ovary more or less is blocked off from the corolla 

chamber by the hairs and swollen bases of the filaments. The deformed or ventricose corollas and the 

secretorylike bases of the filaments suggest mechanisms for presenting nectar. The palate on the lower 

lip is centrally creased with a groove that leads deep 

into the corolla chamber toward the nectar. 

The stigma is above and beyond the anthers and 

pointed downward by the curved style; both are 

spatially removed from the ovary and nectariferous 

region by the length of the elongated corolla, and by 

the elongated filiform, style, and filaments. The stigma 

and anthers are near the roof or above the mouth 

of the corolla. The downward direction of the stigma, 

the downward restriction of the movement of the 

pollen, the overhanging upper lip of the corolla, and 

the recessed or smaller lower lip, suggests that visitors 

arrive straight on or on the rise from below. The 

dorsal portion of the visitor’s body would automatically contact the stigma first and then the anthers. 

The structure of the flowers would appear to separate the anthers from the stigma in such a way as to 

favor out-crossing over automatic self-pollination. In 

a few specimens the anthers seem to touch the stigma, 

providing an opportunity for selfing, but this may 

be an artifact of pressing or the flowers may have 

been immature. Another opportunity for selfing occurs 

as the corollas abscise and slip along the persistent 

style, carrying along the epipetalous stamens so that 

the anthers would seem to rub against the stigma. 

Whether this is significant, of course, depends upon 

compatibility relationships and the period of receptivity of the stigmas. The corollas appear to adjust the 

flowers to specific pollinators and to regulate the 

amount of inbreeding and outbreeding (Thompson & 

Ernst, 1967). 
NUMBER 6 

THE RED COROLLAS IN MEXICO AND CENTRAL 

AMERICA-SECTIONS LAMOUROUXIA AND HEMISPADON 

The position of the stigma and the anthers distally 

and near or beyond the mouth of the corolla removes 

pollen as a potential lure from deep within the red 

corollas. That pollen probably is not the primary 

attraction is emphasized by the retention of the presumed secretory bases of the short, sterile filaments 

in section Hemispadon even though their anthers 

have been lost. The narrow corolla mouth, the relatively large anthers, their fixed position, shaggy pubescence, and often formidable mucros, along with 

the cobwebby interior of the corollas would seem 

especially well suited for the purposes of hovering 

visitors intent upon gathering nectar by means of a 

long, thin proboscis. The same characters would seem 

to discourage bodily entry into the corolla chamber 

by larger insects. A larger organism, such as a bumble 

bee, grappling with the distal portion of the corolla, 

as in gathering pollen, probably would be tipped off; 

nevertheless, there would be a good chance for such a 

visitor to brush against the stigma as well as the 

stamens. Insects small enough to enter these corollas 

for nectar, pollen, or shelter probably would not 

automatically contact the stigmas. 

The literature and relevant topics concerning hummingbird pollination have been reviewed so thoroughly 

by Grant & Grant (1968), that one can assume, almost axiomatically, that tubular, red, scentless, nectariferous flowers, at least in the southwestern United 

States, are pollinated by hummingbirds. These authors 

have documented hummingbird pollination in 42 species representing 19 families (Grant & Grant, 1970), 

and predict that the total number of hummingbirdpollinated species in western North America will 

exceed 129 species. The red corollas of sections Lamourouxia and Hemispadon, by extrapolation, also 

present the syndrome of ornithophily. One of the 

factors substantiating this hypothesis is that bees are 

insensitive to red while red has been suggested to be 

the color most visible to vertebrates (see Sprague, 

p. 197, 1962). Close examination probably will show 

that in many cases in Lamourouxia the net effect is 

a vivid red or red-orange corolla with the mouth or 

tube being yellowish or orange while the abundant 

pubescence is dark red. Hummingbirds are assumed 

to occur throughout the range of Lamourouxia, but 

only one published account links Lamourouxia to 

hummingbirds (Wagner, 1946) ; however, other obser21 

vations have been brought to my attention. Near 

Tepic, Nayarit, Dr. D. E. Breedlove (unpublished) 

observed three species of hummingbirds, the Berylline 

(Amazilia beryllina) , the White-eared (Hylocharis 

leucotis), and the Rufous (Selasphorus rufus) visiting 

a variety of plants without much discrimination, 

among them L. multifida (section Lamourouxia) and 

L. viscosa (section Hemispadon). There would seem 

no reason to doubt that the red corollas of Lamourouxia are pollinated by hummingbirds. 

Among the red corollas the broader, open-mouthed 

L. xalapensis (Figure 8) and the few species closest 

to it in section Lamourouxia (Figures 9, 21) provide 

a strong contrast to the narrow-mouthed corollas of 

the remainder of section Lamourouxia and all of 

section Hemispadon. The open-mouthed corollas have 

exserted stigma, exserted and spreading anthers, and 

undoubtedly freely admit smaller insects into the 

corolla chamber. The exserted anthers would seem 

to place the pollen beyond the reach of any small 

insects sheltering in the corolla chamber, and there 

are few or no glandular hairs surrounding the corolla 

mouth. These exserted anthers do not have conspicuously eldngated spurs. 

Wagner ( 1946) appropriately observed the Mexican 

Violet-eared hummingbird, Coulibri t. thallassinus, 

with pollen-dusted heads, visiting the flowers of L. 

xalapensis (,,L. exserta”) in the mountains surrounding the Valley of Mexico. He believed these flowers 

to be proterogynous, the nectar being produced only 

during the first few days, after which the hummingbirds were attracted by small insects sheltering in the 

corollas. The racemes of L. xalapensis are dependent, 

the tips pointing downward. The flowers are displayed 

on longer, more flexible pedicels which allow the 

flowers to invert to their normal dorsiventral position. 

Maturation of the flowers is acropetal but in a 

descending direction. Thus, it would seem that the 

flowers on erect inflorescences may be protandrous 

while those on dependent inflorescences may be proterogynous. 

The modifications of the corolla which appear to 

be accommodations to certain pollinators imply a 

mutual dependency between pollinator and flower for 

stability in proportion and shape. It is, therefore, 

surprising that variation in lengths of corollas in four 

species with red corollas can have extremes differing 

by a factor of at least two. These extremes for corolla 

length in section Lamourouxia are L. ZongifEora, 26-56 

mm; L. macrantha, 30-60 mm; L. multifida, 22-55 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

mm; and in section Hemispadon, L. ui.scosa, 30-60 

mm. This suggests that these species may be responding to different pollinators or that their pollinators are 

indiscriminate, pointing out the difficulty in attempting to correlate taxonomic and biologic concepts for 

these species. 

THE NON-RED COROLLAS IN SOUTH AMERICA 

AND MEXICO-SECTION ADELPHIDION 

The corollas of section Adelphidion collectively are 

shorter in length, relatively more broadly funnelform, 

and they suggest the syndrome of melittophily (Faegeri & van der Pijl, 1966). Perhaps one of the more 

important reasons for this conclusion is the absence 

of bright red color as the corollas tend instead to be 

pink, lavender, or magenta, and possibly have other 

components which are visible to bees (see Eisner, 

Silberglied, et al., 1969; Macior, 1971). The corolla 

mouths are at the broadest portion of the tubes as 

in L. xalapensis of section Lamourouxia. The large 

mouths, however, are not necessarily open and tend 

to be obstructed by the large, pillowlike palate in 

L. sylvatica, less so in L. dasyantha, and partially 

obstructed by the large anthers and their broad filaments in L. virgata, open but looking downward in 

L. brachyantha. The larger pair of anthers is included 

within the corollas of L. brachyantha and L. sylvatica, 

but partially exposed beneath the upper lip in L. 

dasyantha and L. uirgata. The smaller, upper pair of 

anthers mostly are fertile and bilocular, but always 

deeply included within the corolla chamber. If these 

included anthers are to have a function, the pollinators must bodily enter the corolla chamber. Macior 

( 1971 ) cautions that “coadaptation between pollination mechanisms and pollinators cannot be accurately 

inferred on morphological grounds alone.” 

These corollas seem more like bee- or gullet-type 

flowers. There is no reason to doubt that insects other 

than bees or that hummingbirds also visit these 

flowers. Probing of these corollas by hummingbirds 

for nectar or for very small insects, however, would 

seem to require awkward or purposeful maneuvering 

to accomplish pollen transfer to the appropriate 

stigma. Very small insects entering these corolla 

chambers for shelter, nectar, or to gather pollen, 

probably would not automatically touch the stigma. 

These flowers probably are pollinated by beelike organisms approximately as large as the corolla mouth 

so that they automatically touch the stigma and the 

anthers while struggling to enter the corolla chamber, 

perhaps each species of section Adelphidion requiring 

a different pollinator. 

The morphology of the flowers of L. uirgata seems 

somewhat ambivalent in that the corollas are longer, 

sometimes magenta or reddish in color, and the 

larger pair of anthers and their thickened filaments 

are displayed in the corolla mouth. These flowers 

would seem sufficiently similar in appearance to those 

of sections Lamourouxia and Hemispadon that they 

ought to attract hummingbirds; but, at the same 

time, the smaller, upper pair of anthers usually is 

fertile and always included, suggesting that pollinators 

are expected to enter the corolla chamber. In this 

species we may have a model to illustrate the situation 

suggested by Baker & Hurd (1968, p. 397) in which 

a flower type may be adapted to more than one kind 

of pollinator. Assuming that characters of the flower 

involved with pollination are labile, it is easy to 

visualize that a shift in emphasis to one or the other 

pollinator could lead, on the one hand, to ornithophilous flowers like those of section Hemispadon and, 

on the other hand, to more melittophilous flowers like 

the others of section Adelphidion without an “interregnum” period. 

Synopsis of Vegetative Characters 

Vegetative morphology exhibits a wide diversity of 

distinctive characters which are useful for taxonomic 

purposes. With few exceptions, the vegetative differences among and within species seem trivial and 

probably of relatively low selective value. Perhaps 

this is an indication that most vegetative characters 

are mutually efficient in this genus and that they 

have become variously recombined so that there no 

longer are whole blocks of characters to indicate 

distinct phylogenetic lines. This recalls a situation in 

the Asclepiadaceae called reticulate variation (Ehrlich 

& Holm, p. 258, 1963) which may be the result of 

complex, obligate insect pollination mechanisms in 

these plants. It is difficult to predict the advantage 

of a petiole of less than a millimeter in length over 

that of a sessile leaf blade. One might easily assume 

that photosynthetic activity is about as good in one 

kind of leaf shape and margin as another. If this be 

true, vegetative characters might be expected to be 

less stringently affected by the whims of selection 
NUMBER 6 23 

and to indicate important, long-standing phylogenetic 

relationships. Similar species with relatively many 

vegetative characters in common might represent 

rather recent or contemporary biosystematic contacts. 

Dissimilar species seeming to have some common 

vegetative similarity may represent vestiges of ancient 

phylogenetic alliances which no longer have any biosystematic relevance-or they might be products of 

random convergences. It is difficult to accept a taxonomy which places taxa with opposite floral extremes 

side by side merely because their leaves are similar. 

On face value, two plants with very similar flowers 

potentially seem more likely to be closely related than 

two plants having only similar leaves in common. 

Some of the vegetative characters help to support 

the clustering of the species within sections; however, 

vegetative characters alone do not appear to offer a 

general explanation of relationships in Lamourouxia. 

For graphic representation of vegetative relationships 11 categories of characters were coded and compared for matching similarities. For an example of 

similar coding technique see Ernst, 1967. Each character exhibits from two to five contrasting states which 

collectively exhaust the supply of traits for distinguishing the species of Lamourouxia vegetatively. 

The characters vary in many instances beyond the 

limits of the coded states, necessitating value judgments as to the most representative condition for the 

species. Some kinds of characters, such as size of 

blade or length of petiole, may be similar among the 

leaves of some species; however, this is an abstract 

comparison since the texture, margin, or pubescence 

may be so different as to negate the association as 

a similarity when visually compared. 

The purpose of the circular graph (Figure 13) is 

to show the strength of intersectional vegetative similarities among 36 independent pairs of species sharing 

a maximum of 8 to 10 similarities out of a total of 

11 categories. The species names are arranged alphabetically around the circle and grouped by sections. 

Fully half the lines connect species from different 

sections ; the scarcity of lines connecting species within 

sections suggests the lack of correlation between floral 

and vegetative characters. Adelphidion appears to 

have more vegetative characters in common with 

Hemispadon than with section Lamourouxia ; this 

supports my argument that Hemispadon has been 

derived from Adelphidion. That section Lamourouxia 

appears to have more vegetative characters in common with Hemispadon than with Adelphidion is in 

FIGURE 13.-Table comparing various strengths of intersectional vegetative similarities (explanation in text) ; broad 

line indicating 10 similarities, narrow line 9, and broken line 

8 ; species names arranged alphabetically by section, Hernispadon to the left, Adelphidion below central, section Lamourouxia to the right. 

direct contradiction of my view that sections Lamourouxia and Hemispadon have had different phylogenetic pathways. In Adelphidion, where there are 

the fewest species to compare, there is less internal 

similarity than in sections Lamourouxia and Hemispadon where there are more species to compare. 

The relationship between sections Lamourouxia and 

Hemispadon is mentioned again in the discussion. 

A few of the lines on the circular graph identify 

close similarities among these species. The line between 

L. parayana and L. pringlei correctly identifies the 

strong similarity between these species. The intersectional lines connecting L. brachyantha to the arc 

between L. nelsonii and L. viscosa recalls to mind that 

Greenman ( 1905), disregarding floral morphology, 

originally placed L. brachyantha in Hemispadon near 

L. rhinanthifolia. If similar reasoning were to prevail, 

with vegetative morphology considered more important than floral, it would be just as valid to associate 

L. longiflora (Lamourouxia) with L. integerrima 

(Hemispadon) ; L. parayana (Lamourouxia) with 

L. gutierrezii (Hemispadon) ; L. dasyantha (Adelphidion) with L. tenuifolia (Hemispadon) ; and L. xalapensis (Lamourouxia) with L. lanceolata (Hemispadon). 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Discussion 

SYSTEMATICS AND DIFFERENTIATION OF THE SECTIONS 

Floral characters seem more useful than vegetative 

characters for evaluation of species relationships in 

Lamourouxia. The features of the flowers on which 

the sections are founded, as well as those demarcating 

species, are simple modifications in color, proportion, 

position, or shape which do not involve reorganization 

of floral vascular structure. Conformity within the sections centers on sets of characters which have functional significance in the process of pollination (see 

Ornduff, p. 121, 1969). While some characters of 

the flowers probably have relatively low selective value 

(Figure 12) those characters directly associated with 

the pollination system (Figure 8) probably are labile 

and have high selective value. This suggests that pollinators may have bred some corolla shapes that are 

deceptively similar in function or appearance from 

genetically rather different stocks. 

Differences in color and shape of the corolla and in 

the development of the stamens appear to record 

changes in the pollination system of Lamourouxia. 

Aside from purely attracting and protective functions 

(see Straw, 1956), the net effect of these differences 

almost certainly ensures that pollen is precisely applied 

to and withdrawn from the bodies of the established 

pollinators. In this respect, the proposed arrangement 

of the species appears to be an inadvertent classification of the pollinators. Since the morphological reasoning for the clustering has been explained, it now 

is appropriate to consider what circumstances might 

have surrounded the differentiation of these groups 

as a kind of test of the systematic value of this 

arrangement of the species. For the purpose of this 

discussion, it is not so important that the pollinators 

are assumed to be bees and hummingbirds but, rather, 

that there are two distinctive floral morphologies-the 

one adapted to pollinators that bodily enter the corolla 

chamber, and the other adapted to pollinators that 

only insert a narrow proboscis into the corolla 

chamber. 

In most flowers of section Adelphidion the pair of 

short stamens with smaller, deeply included anthers 

mostly produce pollen, suggesting that these anthers 

are functional in the process of pollination. The fact 

that these anthers are smaller and sometimes sterile 

or rudimentary perhaps indicates that they are of 

secondary importance. One presumes, however, there 

must be some selective advantage to their continuing 

presence or they surely would be lost as in section 

Hemispadon (see discussion of staminode in Penstemon, Straw, p. 117, 1956). The equivalent pair of 

short, deeply included stamens in Hemispadon mostly 

have sterile or rudimentary anthers but, in a few 

flowers, these anthers produce a little pollen, suggesting their potential for full restoration were there a 

sustained function for this pollen. Pollinators, it may 

be inferred, bodily enter the corolla chamber and 

contact the small stamens in Adelphidion, but in 

Hemispadon pollinators hover outside, inserting only 

a narrow proboscis into the corolla chamber and do 

not contact the small, sterile or rudimentary anthers. 

This suggests that anthers, having lost their role in 

reproduction, cease to produce pollen and become 

rudimentary. 

Not only do the stamens provide the means for 

separating the species into plausible sections, they also 

illustrate a natural morphological sequence (Figure 

14) showing the upper pair of anthers in full development in section Lamourouxia, in an intermediate 

condition in Adelphidion, and reduced to sterile rudiments in Hemispadon. This open-ended sequence can 

be read in either direction and, theoretically, is reversible from the morphological point of view. The 

possibility of morphological reversals seldom has been 

acknowledged (see Eyde, 1971). Other reversible and 

irreversible situations were mentioned by Carlquist 

( 1970). In potentially reversible sequences it is difficult to arrive at a clear understanding of primitive 

LAMOUROUXIA HEM IS PAD0 N 

ADELPHIDION 

FIGURE 14.-Diagrams comparing stamina1 development in 

the three sections of Lamourouxia, only one of each pair of 

stamens shown. Left, two pairs of fully developed anthers, 

the filaments subequal; center, one pair of large anthers, one 

pair of smaller anthers; right, one pair of fertile anthers, one 

pair of sterile filaments. 
NUMBER 6 25 

conditions. In Lamourouxia the pollination syndrome 

and the evolution of the stamens appear to be inseparable. This provides an opportunity to consider 

change from one morphological state to another as 

change from one selection regime to another. 

Color and shape of the corolla are presumed relatively labile from generation to generation. Within 

the flexibility of the genotype, this lability probably 

ensures the continued efficiency of the flowers in 

regard to pollination. The readiness with which these 

changes may be effected can be realized from the simple nature of the extremes. The extremes for color 

range from red to not red; for shape the extremes 

range from permitting the pollinator to enter bodily 

the corolla chambers to only allowing the insertion 

of a narrow proboscis. The stamens, by comparison, 

seem more complicated and are assumed to be somewhat less labile than the corolla because, in addition 

to the primary necessity of pollen and nectar, the 

anthers must be accurately placed, correctly shaped, 

and chronologically coordinated for meiosis and anthesis with the remainder of the flowers. Anthers having no purpose, as suggested by the flowers of Hemispadon, are lost, probably by default. 

The ornithophilous flowers of section Lamourouxia 

and of section Hemispadon, and the melittophilous 

flowers of L. brachyantha and L. sylvatica (section 

Adelphidion) appear to be too strongly committed 

by their adaptive modifications to their respective 

pollinators to be readily susceptible to major modifications. As long as the established pollination regime 

does not change, the floral morphology in these taxa 

will remain relatively stabile. The kinds of flowers 

most pliable to change probably are sufficiently flexible 

to be pollinated by more than one kind of visitor as 

suggested by Baker & Hurd (p. 397, 1968). The 

morphology of L. virgata, section Adelphidion, suggests that this species may be pollinated by bees as 

well as by birds. The color of these corollas, although 

usually reported to be pink or rose, sometimes varies 

to magenta or red. The larger anthers and their 

broad filaments are partially exposed in the corolla 

mouth ; the smaller included anthers usually are fertile 

but sometimes they are rudimentary or sterile. The 

mouth is relatively open and the palate is only moderate in size. The general appearance as well as some 

technical characters cause L. virgata to resemble 

section Hemispadon. 

There is no particularly urgent reason to conclude 

that L. virgata is an ancestor of section Hemispadon 

but this species does seem like a useful evolutionary 

model for the purpose of illustrating how section 

Hemispadon may have been derived. Assuming that a 

population of L. virgata were pollinated both by bees 

and by hummingbirds, a change in emphasis of pollinators probably would have a disruptive effect on 

floral morphology. A surge in the frequency of visits 

by one kind of pollinator, as there are no intermediates between bees and birds, probably would provide 

an opportunity for the flowers to become better adjusted to the more frequent kind of pollinator. Natural 

selection would lead to flowers more conspicuously 

adapted to their pollinators than were their more 

ambivalent predecessors. Stability in the major characters of floral morphology presumes stability in the 

pollination regime. On the basis of these generalizations we may now consider how differences in floral 

morphology exhibited by the sections might be correlated with changes in the pollination regimes. 

SECTIONS LAMOUROUXIA AND HEMISPADON 

The red corollas of section Lamourouxia, with two 

pairs of fully developed anthers, and the red corollas 

of Hemispadon, with only one pair of fertile anthers 

(Figure 8), are sufficiently similar in appearance and 

function that one would assume them to have shared 

much of the same phylogenetic pathway. The sterile 

filaments of Hemispadon, our only clear landmark, 

must be derived but it is improbable that Hemispadon 

could have been derived directly from section Lamourouxia because no selective advantage can be 

visualized to account for the loss of a pair of anthers 

between functionally identical ornithophilous flowers. 

The rudimentary stamens in Hemispadon, although 

mostly sterile, nevertheless, are potentially functional 

since, from time to time, they bear small, one- or 

two-locular anthers with a little pollen. Their permanent decline and the continuing differences between 

Hemispadon and section Lamourouxia rest upon the 

stability of the pollination system and the unlikely 

occurrence of a pollinator capable of restoring these 

rudimentary anthers to full service. 

SECTIONS ADELPHIDION AND HEMISPADON 

Derivation of Hemispadon from Adelphidion (Figures 

8, 14) seems more likely and can be explained as 

the consequence of differentially labile floral organs 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

and a relatively sudden reversal in an established 

selection regime, such as ornithophilous capture of 

dominance in a previously ambivalent pollination 

system. Such a shift in emphasis of pollinators would 

favor redder, longer, narrower corollas. The pace of 

the change of the corollas probably exceeded the 

ability of the smaller, upper pair of stamens to adjust. 

The included anthers of the upper stamens became 

isolated from bees through elongation of the corollas 

and narrowing of the corolla mouths. Once their 

pollen was no longer involved in pollination these 

anthers became sterile rudiments. The filaments probably continue to exist because of their nectariferous 

contributions. 

Difference in number of floral organs, particularly 

organs of the reproductive system, often delimit major 

groups of flowering plants. The adaptive significance 

of change from one number to another is difficult to 

appreciate by comparing families or genera (see 

Cronquist, p. 118, 1968) because these taxa are too 

remote from the circumstances surrounding the change 

and the transitional stages are lost or not recognized. 

Ultimately, such changes must be traced to the effects 

of selective forces operating among closely related 

individuals within populations. The stamens in Lamourouxia provide an opportunity to examine intermediate stages in a morphological series. The normally 

fertile, upper pair of stamens in Adelphidion sometimes is sterile; the normally sterile, upper pair of 

stamens in Hemispadon sometimes is fertile. This 

provides insight into how pollinators acting as selective 

forces on potentially variable, closely related plants, 

might have initiated the change from two pairs of 

fertile anthers in Adelphidion to one pair of fertile 

anthers in Hemispadon. 

SECTIONS LAMOUROUXIA AND ADELPHIDION 

Early specialization in this genus is assumed to have 

led, on the one hand, to flowers like those of section 

Lamourouxia favoring bird pollination and, on the 

other hand, to flowers like those of Adelphidion favoring bee pollination (Figures 8, 14). The morphological separation of these sections probably represents a 

prior differentiation in selection regimes than the 

one involving Hemispadon because a switch from 

melittophilous to ornithophilous dominance of pollination in flowers like those of Adelphidion is predicted to have led to flowers like those of Hemispadon. 

The nearness of these relationships could be tested 

by cytological analyses of interspecific hybrids. 

Within section Lamourouxia there are two lines 

of development : one with dependent inflorescences, 

open-mouthed, proterogynous corollas, and exserted, 

spreading stamens as in' L. xalapensis; and the other 

with ascending inflorescences, more closed-mouthed, 

protandrous corollas, and mutually connivent anthers, 

as in L. multifida (Figure 8). The flowers of L. ouata 

and L. macrantha, two species more or less centrally 

located in section Lamourouxia, are phenotypically 

similar and are suggestive of evolutionary models 

showing how the above extremes in section Lamourouxia may have been derived. The inflorescences are 

erect in L. macrantha but dependent in the very 

similar L. ovata. 

The depressed inflorescences in three species of 

section Lamourouxia and the same character in L. 

lanceolata of Hemispadon may be a coincidental convergence, or may be a clue to an extinction in section 

Adelphidion. If sections Lamourouxia and Adelphidion were derived from similar ancestors while Hemispadon was derived from Adelphidion, the genes for 

depressed inflorescence may have passed to Hemispadon through a now extinct species having the 

characteristics of Adelphidion (Figure 15). 

Synopsis and Conclusions 

Only a few floral characters are useful for appraising 

systematic relationships within Lamourouxia and these 

LAMOUROUXIA 

birds 

3d 7e 

\I 

H E M IS PADON 

birds 

ADE LPH ID ION 

bees 

4e [d,?lost] 

FIGURE 15.-Evolutionary implications based upon the major 

morphological characters of the flowers and the inflorescences for the three sections of Lamourouxia. The number 

of species with dependent (d) and erect (e) inflorescences 

indicated. 
NUMBER 6 27 

are concluded to have high selective value. In addition to the characters which are common to all 

species of the genus, conformity within the sections 

rests upon three criteria which are functionally correlated with the pollination system: color and shape 

of the corolla and the morphology of the stamens. 

Were it not for the stamens, the species would separate into two groups representing different selection 

regimes, the red-flowered, ornithophilous species, and 

the non-red, melittophilous species. Disparate development of the stamens, however, proclaims a more complicated evolutionary history for the red-flowered species, as some have four fertile anthers and others only 

have two fertile anthers. The major differences in 

floral morphology among the sections are attributed 

to changes in the pollination regimes. 

The somewhat scattered distribution of vegetative 

similarities suggests that the characters of the ancient 

species may have become recombined through some 

kind of reticulate pattern of evolution so that phylogenetic significance is obscured. Those instances where 

floral and vegetative morphology seem to be correlated 

might be examples of random convergences or examples of more recent or secondary speciations. Lack of 

recognizable patterns in geographical distributions corresponding to patterns in floral or vegetative morphology returns us to the proposition that pollinators 

may have bred some flowers that are deceptively 

similar from genetically rather different stocks. 

Comparative floral morphology, particularly of the 

stamens, shows the relationship of the sections to be 

Lamourouxia - Adelphidion - Hemispadon, and by 

implication, these plants are, respectively, bird-beebird, in regard to pollination. The only cytological 

information available is one diploid count for section 

Lamourouxia (L. Zongiflora, n = 7) and one tetraploid 

count for Hemispadon (L. viscosa, n = 14). In general, 

this sequence appears to read from left to right (Figures 8, 14). Since the characters of the corolla directly 

involved in the process of pollination are concluded 

to be labile, there is no reason why this morphological 

sequence, given the appropriate pollinators and circumstances, could not proceed in either direction or 

reverse itself. Most of the present species, however, 

suggest states of terminal specializations for their 

respective pollinators and for these, a major reversal 

in the pollination system probably would have to 

overcome an enormous amount of inertia. 

The flowers of L. virgata, section Adelphidion, are 

singled out as being less specialized and perhaps pollinated by bees as well as by birds. This species is 

used as an example to show how a change in the 

regular pollination regime could lead, on the one 

hand, to flowers like those of section Hemispadon and, 

on the other hand, to flowers more specialized for 

bee pollination. Although section Hemispadon and 

section Lamourouxia are superficially similar, comparative morphology of the stamens suggests that 

Hemispadon is derived from Adelphidion. The differences between sections Lamourouxia and Adelphidion 

are attributed to a similar but earlier differentiation 

in the pollination regimes. 

Some authors (see Percival, p. 117, p. 154, 1965) 

have observed that the foraging behavior of bees and 

of at least one species of Old World Sun-birds tends 

to be from bottom to top in protandrous, racemose 

inflorescences. If generally true, this would favor outcrossing of the lower, older flowers. Proterogyny in 

dependent, racemose inflorescences, such as those of 

L. ovata, L. dependens, and L. xalapensis, therefore, 

might be a compensation to facilitate out-crossing of 

the lower, younger flowers. One might easily surmise 

from observing hummingbird behavior that these organisms are much too ingenious to restrict their foraging activities to so narrow a pattern as that characteristic of bees. Among others, Baker & Hurd (1968, 

p. 394) and Grant & Grant (1968, 1970) suggest 

that bee pollination existed before bird pollination. 

The differences in the inflorescences of section Lamourouxia perhaps represent ancient adaptations to 

the foraging behavior of bees that predate ornithophily in this genus. 

The foregoing discussion explains, primarily on 

the basis of comparative morphology of the flowers, 

the objectivity of clustering the species of Lamourouxia into and within sections. The nature and distribution of the floral characters in this arrangement 

of the species suggests that adaptations to hummingbird pollination are superimposed on earlier adaptations to bee pollination. This provides insight into 

the evolution and survival value of the sets of characters which distinguish the sections and implies that 

stability in floral morphology, hence conformity within 

the sections, depends upon natural selection and stability in the pollination regime. From this position, 

we may rationalize that the evolutionary units of 

importance are the local, interbreeding populations 

where the principal selection regime is the pollination 
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

regime. A disruption of the established selection 

regime, such as switch from bee to bird, or a change 

in emphasis from bee and bird to bee or bird pollination, probably would initiate differential reproduction 

of genotypes and provide the impetus for the changes 

in floral characters which distinguish the sections of 

Lamourouxia. This interpretation appears to strengthen support for the proposed arrangement of the 

species and for the taxonomic conclusions expressed 

in this paper; however, it also confronts us with a 

morphological series which, depending upon the circumstances of natural selection, might proceed in 

either direction, and the phylogenetic dilemma that 

the sections of Lamourouxia, and perhaps’ also some 

of the species, at least in part, might represent the 

independent products of convergent evolution. 

Taxonomy, Key, and Descriptions 

Lamourouxia H.B.K., Nov. Gen. 2: ed. folio 269, ed. 

quarto 335. “1817,” 1818. Nom. cons. 

Plants herbaceous, perennial, and probably hemiparasitic, the leafy stems somewhat woody below, from a 

gnarled burl; stems excurrent and ascending, or deliquescent and/or arching over other vegetation. 

Stems, leaves, and calyxes pubescent, sometimes glandular, or glabrous ; corolla always pubescent, sometimes 

glandular. Hairs uniseriate, sometimes terminated by a 

gland ; branched hairs mostly confined to the corolla, 

often mixed with simple hairs. Leaves and floral bracts 

similar, opposite throughout, sessile to petiolate, pinnately veined, craspedodromus, margins sometimes 

entire but mostly incised, the secondary veins usually 

terminating at the sinus. Flowers many, pedicelled 

and solitary in axiles of frequently foliaceous bracts; 

bracteoles absent. Inflorescences terminal, racemose, 

simple or branched from below. Calyx tubular campanulate, four-lobed, often somewhat bilabiate, persistent. Corolla tubular, elongated, pentamerous, bilabiate, the larger upper lip morphologically twolobed, although often appearing as one, strongly 

galeate in bud, but inverted naviculiform or mostly 

little or obscurely galeate after anthesis; the dissimilar 

lower lip three-lobed and partially enclosing upper 

lip before anthesis. The corolla with the epipetalous 

stamens deciduous. Stamens diadelphous, inserted 

near base of corolla, but one pair above, one below 

as the flowers are ascending Dr horizontal; stamens 

homoeomorphous or dimorphous, the upper pair 

shorter and sometimes with vestigial or smaller anthers ; fully fertile anthers two-locular at maturity, 

mostly shaggy pubescent, frequently connivent, mostly 

sheltered by the upper lip of corolla, or exserted and 

spreading. Locules of fully fertile anthers more or 

less equal, laterally dehiscent, nonconfluent, often 

mucronate basally. Pollen tricolpate. Ovary bilocular, 

style narrowly attenuate, stigma more or less twolobed, mostly exceeding anthers and usually exserted. 

Ovules numerous, multiseriate, radially inserted on 

two axile placentae. Capsules mostly ovoid, apically 

attenuated into crooked base of style, IoculicidaI from 

above. Seeds small, tightly packed, about twice as 

long as wide, truncated, sides somewhat angular, coat 

with uniform, polygonal reticulations. Dicotyledonous 

embryo orthotropous, contained in a small envelope 

of endosperm (Figures 1, 6, 8). 

LECTOTYPE SPECIES.-L. multifida H.B.K. (Taxon 

18: 480. 1969). 

DISTRIBUTION. - Twenty-four species, northern 

Mexico to western Panama, and two species, Colombia to central Peru (Figure 2). 

Key to Lamourouxia 

1. Stamens subequal, the 4 anthers more or less equal in size, fertile, mutually connivent near 

I. SECTION LAMOUROUXIA 

Species 1-10 

the mouth of the corolla or exserted and spreading; corolla red . 

I. SECTION LAMOUROUXIA 

Species 1-10 

2. Plants upright, excurrent, branches ascending at acute angle, inflorescence tips ascending. 

3. Leaves 1- or 2-pinnately, deeply, and narrowly divided; widespread, Mexico and Guatemala ................................................................................................ ............ 1. L. multifida 

3. Leaves not divided. 

4. Leaves lanceolate or oblanceolate; length 3~ (or more) width. 
NUMBER 6 29 

Key to Lamourouxia--continued 

Species I-IO-continued 

I. sECTION LAMOUROUXIA-cOntinUed 

5. Petioles less than 1 mm long; blades tapered to base, margins T entire; calyx 

lobes subulate, 1 mm or less wide; infrequent, Chihuahua, Durango, and Jalisco 

2. L. longiflora 

5. Petioles 1-2 mm long, margins incised; calyx lobes triangular, 3 mm or more 

wide. 

6. Base of blade obtuse to truncate, margins irregularly doubly dentate-serrate6. Base of blade acute, margins evenly dentate-serrate; Guerrero ..... 5. L. dlspor 

crenate; Chiapas ( ?Guatemala) ............................................... .7. L. stsnoglossu 

4. Leaves ovate to obovate or length 3 X (or less) width. 

7. Leaves elliptical to obovate, 9 mm or less wide, rounded or tapered to base, 

petioles 1 mm or less long. 

8. Blades membranous, ovate, evenly crenate, sparsely pubescent; calyx 5-6 mm 

long; southwestern State of Mexico ........ 

8. Blades coriaceous, elliptical, obovate to spatulate, margins entire to irregularly 

few-dentate or -crenate, densely scabrous pubescent; calyx 9-10 mm long; 

Hidalgo, Oaxaca .............................................................................. 4. L. pringki 

7. Leaves broadly ovate, cordate, 15 mm or more wide; margins irregularly, 

doubly, multi-serrate-dentate-crenate, petioles 2-8 mm long ; Mexico ( ?Guate2. Plants dependent, deliquescent, branching frequently at nearly right angles, inflorescence 

tips arching downward, flowers resupinate. 

9. Leaves sessile, ovate, cordate, 22-37 mm wide; Oaxaca ......................... 8. L. ovuta 

9. Leaves with petioles 1-7 mm long, blade 6-23 mm wid$. 

10. Leaves ovate, calyx and leaves mostly pubescent (sometimes glabrous), axil of 

10. Leaves lanceolate, calyx and leaves mostly glabrous or glabrate; Mexico, north of 

1. Stamens strongly unequal, the anthers dimorphic, the longer pair of filaments thickened, 

their anthers large, usually connivent near the mouth of the corolla or barely exposed; the 

shorter pair of filaments filiform, their anthers smaller, rudimentary, or absent, always deeply 

included. 

11. Fertile anthers 4, the smaller pair rarely sterile; corolla lavender, pink, or magenta, not 

red ...................................................................... 11. SECTION ADELPHIDION 

Species 11-14 

............... .3. L. porayana 

mala) ......... .......................... 6. L. macrantha 

sinus of calyx revolute, protruding; Guatemala ................................ 9. L. dsfiendsns 

Isthmus of Tehuantepec .................................................................... 10. L. xahpensk 

11. SECTION ADELPHIDION 

Species 11 -1 4 

12. Plants from Mexico. 

13. Leaves ovate, cordate, glandual pubscent ............................ 11. L.. brachyantha 

13. Leaves lanceolate to subulate, tapered to base, glabrous, not glandular 

12. L. dasyunthu 

12. Plants from South America. 

14. Corollas 13-24 mm. long, mouth occluded by palate, anthers included 

14. Corollas 29-45 mm long, mouth open, lower pair of anthers exposed 

13. L. sylvutica 

14. L. virgata 

111. SECTION HEMISPADON 

Species 15-26 

1 1. Fertile anthers 2, the rudimentary pair rarely fertile; corolla red 

111. SECTION HEMISPADON 

Species 15-26 

15. Calyx and leaves definitely pubescent, sometimes glandular. 

16. Leaves sessile, rounded, truncate, or cordate at base. 

17. Calyx glandular; average corolla 6-7 mm wide; leaves ovate to lanceolate. 
30 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Key to Lamourouxia-continued 

111. SECTION HEM~spmN-continued 

Species 15-26-continued 

18. Leaves to 15 mm wide, young and old glandular-pubescent, not scabrous, margins to 14 dentate on a side, not reticulate below, calyx lobes 

4-27 mm long. Infrequent, south-central Mexico ............ 15. L. nelconii 

18. Leaves 15 mm or more wide, densely pubescent, often scabrous, young 

leaves and bracts glandular, margins 14 or more dentate on a side, 

reticulate venation standing out below, calyx lobes 2-4 mm long. Wide17. Calyx weakly or not glandular, bracts and leaves not glandular; average 

corolla 9-10 mm wide; leaves mostly ovate, pinnate venation standing out 

below. 

19. Leaves to 50 mm long, to 28 mm wide, sometimes scabrous; calyx lobes 

17. L. rhinanthifolia 

19. Leaves 50 mm or more long, 29 mm or more wide, membranous, not 

spread, Mexico and Central America ................................ 16. L. uiscosu 

frequently irregular to dentate. Widespread in Mexico. 

scabrous; calyx lobes entire; rare, Oaxaca ........................ 18. L. nnithii 

late, glandular; Jalisco ............................................................ 19. L. jalircataa 

......................................... 26. L. gutierrexii 

16. Leaves not sessile, tapered to the base. 

20. Leaves lanceolate, 50 mm or more long; calyx lobes 10-12 mm long, subu20. Leaves ovate, to 25 mm long; calyx lobes 2-5 mm long, not glandular; 

glandular; leaves tapered to base, neither 

Costa Rica and Panama 

15. Calyx and leaves glabrous or glabr 

cordate, nor sessile. 

2 1. Leaves ovate, elliptical to obovate. 

22. Leaves 18-28 mm long. 

23. Leaves elliptical to obovate; calyx lobes to 2 mm long corolla 45-49 mm 

long, 8-10 mm wide; Guerrero, Morelos ..................... 

23. Leaves ovate, calyx lobes 4-6 mm long, corolla to 35 

wide; Colima ............... ................................................. 25. L. colimaa 

22. Leaves less than 10 mm Ion yx lobes to 1 mm long; Oaxaca. 

21. L. microphylla 

2 1. Leaves lanceolate to subulate. 

24. Inflorescence tips ascending; calyx lobes 6 mm or less long; branching ascending. 

25. Blades subulate, irregularly dentate to 3 rnm wide; Oaxaca. 

22. L. tenuifolia 

25. Blades lanceolate, sub-entire, 3-7 mm wide; Guatemala ( ?intergrading 

24. Inflorescence tips dependent; calyx lobes 6-15 mm long, spreading or re24. L. lanceolata 

with lanceolata) ................ ................................ 23. L. integerrima 

flexed; branching at nearly right angles; Oaxaca and Central America. 

I. Section Lamourouxia. Species 1- 10. 

Stamens subequal, the filaments similar, the 4 anthers 

more or less equal in size, fertile, shaggy pubescent, 

mutually connivent, long mucronate, near the mouth 

of the corolla ; or exserted, spreading, less pubescent 

and slightly or not mucronate. Corolla red or redorange, the tube ventricose to somewhat funnelform 

(Figures 8, 9, 16, 17, 21). 

TYPE SPECIES.-L. multifida H.B.K. 

DIsTRIsuTIoN.-Ten species, Mexico and Guatemala (Figures 17, 22). 

1. L. multifida H.B.K., Nov. Gen. 2: ed. folio 273, 

ed. quarto 339. “1817,” 1818. 

TYPE LOCALITY. - “Crescit in temperatis Novae 

Hispaniae inter Guanaxuato et Santa Rosa de la 

Sierra, alt. 1200 hex ... Floret Septembri.” 

TYPE sPEcIMEN.-Bonpland 4273, ‘%a rosa,” P 
NUMBER 6 31 

(in General Herbarium but not in Humboldt & Bonpland Herbarium). 

SYNONYMY.- 

L. laciniata Martens & Galeotti, Bull. Acad. Sci. Bruxelles 

12 (2): 32. 1845. Type locality: “Sur les basaltes de 

Cerro de Macuiltepeque, pres Xalapa a 5,000 pieds; dans 

les bois de chenes et sur les rochers de Juguila del Sur 

(c6te au sud d’Oaxaca) a 5,000 pieds . . . Mai-Septembre 

. . . H. Gal. no. 990.” Syntype specimens: Galeotti 990, 

mixed data, BR (2 specimens), G, P, W. 

L. laciniata var. pilosa Martens & Galeotti, Bull. Acad. Sci. 

Bruxelles 12 (2): 34. 1845. Type locality: “Dam les 

bois San Pedro, pres de Real del Monte, a 8,000 pieds . . . Aout . . . Gal. No. 1085.” Type specimen: Galeotti 

1085, BR. 

L. multifida H.B.K. p grandiflora Bentham, in DC. Prod. 

10:540. 1846. Type locality: “pr. Bolanos (Hartweg! n. 

187) . . . (v. s.).” Type specimens: Hartweg 187, BM, 

G, GH, K (Herb. Benth. “Bolanios”; Herb. Hook. “Zacatecu”), P, W. 

L. grandiflora “Benth.” ex. J. J. Linden, “Cat. no. 10, 6 

(1855) .” “Cette magnifique vari6te du L. multifida . . . .” 

Typescript copy examined at PH; original not seen. 

FIGURE 16.-Section Lamourouxia, diagrams of flowers and 

leaves. a, L. multifida, McVaugh 13437, US; a’, Thomas 

14136, US; b, L. longijlora, Hartweg 188, W; b’, (L. hyssopifolia) Palmer 260, US; b“, (L. hyssopifolia) Goldman 

210, US; c, L. parayana, Paray 2242, ENCP; d, L. pringlei, 

Ptingle 4972, US; d’, Gonzales Q. 3127, ENCP; e, L. dispar, 

Straw d Gregory 1062, MEXU. 

FIGURE 17.-Distribution of section Lamourouxia in Mexico and Guatemala. The position of 

L. hyssopifolia is discussed under L. longiflora. 
32 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

L. pringlei “Gandoyer,”=Gandoger, Bull. SOC. Bot. France 

66:217. 1919. Type locality: “Mexico, ad. Sta Fe prope 

Mexico, alt. 8000 ped. (Pringle n. 7982!).” Type specimens: Pringle 7982, F, MO. Not L. pringlei Robinson & 

Greenman ex Pringle. 

L. tepicana (M. E. Jones) Pennell in Morton, Contr. US. 

National Herb. 29: 115. 1945. Basionym: PediculariJ 

tepicana Jones, Contr. Western Bot. 15: 150. 1929. Type 

locality: “no. 23211, Tepic, Nayarit, February 15, 1927.” 

Type specimen: Date mixed, Jones 23211, 10 Feb. 1927, 

POM. 

Stems erect to 2 mm tall. Leaves glabrous to 

pubescent, 21 (39) 57 mm long, 10 (24) 33 mm broad, 

deeply once or twice pinnatifid, 2- to 5-lobed on a 

side, divisions 1-4 mm broad, mostly linear, often 

mucronulate, margins entire, the petiole 6( 13) 23 mm 

long, midrib standing out below. Inflorescence ascending; pedicels 2(3)6 mm long. Calyx pubescent to 

glabrate, 6 (8) 11 mm long, 4( 5) 7 mm road, the lobes 

broadly triangular to subulate, rarely toothed, 2 (4) 7 

mm long. Corolla red (to orange or ?pink), 22 (41) 55 

mm long, 7( 10) 17 mm broad, ventricose, the upper 

lip 4 ( 11 ) 16 mm long, little or not lobed, the lobes of 

the lower lip 4-6 mm long. Stamens subequal, shaggy, 

all fertile, the anther lobes prominently mucronate 

below. Capsules ovoid, 9-1 1 mm long, 5-6 mm broad. 

(Figure 16). Altitude: Mexico, 800-3950 m; Guatemala, 1067-3000 m. Flowering : throughout year, 

peak August to December. 

DISTRIBUTION ANDNUMBER OF COLLECTIONS EXAMINED (Figures 17, 22) .-MEXICO : Aguascalientes (2) ; Chiapas (10) ; Durango (Breedlove 1741, 

DS, MICH) ; Distrito Federal ( 11 ) ; Guariajuato (4) ; 

Guerrero (5) ; Hidalgo (7) ; Jalisco (12) ; State of 

Mexico (26) ; Michoacan ( 11 ) ; Morelos (Halbinger 

78, GH) ; Nayarit (4) ; Oaxaca (10) ; Puebla (10) ; 

San Luis Potosi (4) ; Sinaloa (3) ; Tlaxcala (2) ; 

Veracruz ( 14) ; Zacatecas (2) . GUATEMALA : 

Alta Verapaz (von Tuerckheim 2022, BR, C, F, G, 

GH, MO, US, W) ; Chimaltenango (3) ; Escuintla 

(Eggler 488, F) ; Guatemala (4) ; Huehuetenango 

(5) ; Jalapa (2) ; Quezaltenango (3) ; SacatepCquez 

(5) ; San Marcos (Steyermark 36688, F) ; Santa Rosa 

(Heyde & Lux 149, G, GH, MO, US) ; Solola (4) ; 

Zacapa (Steyermark 29796, F) . The unique, deeply pinnatifid leaves, although extremely variable, distinguish this species from all 

others. The subulate as well as triangular calyx lobes, 

and the extremes of corolla length, varying from 

22-55 mm, are notable. The “var. grandiflora” is 

only one among many extremes and is an impractical 

taxonomic element. This species seems closely allied 

to L. longiflora. 

2. L. longijlora Bentham, Plantae Hartwegianae, p. 

22. 1839. 

TYPE.-Hartweg 188, unlocalized at time of publication, but later (Benth. in DC. Prod. 10: 540. 

1846) identified as “in Mexico pr. Bolanios.” 

TYPE SPECIMENS.-BM, GH, K (Herb. Benth., 

“Bolanios;” Herb. Hook., “Zacatecas” ) , W (Figure 

18). 

SYNONYMY.- 

L. hyssopifolia A. Gray, Proc. Amer. Acad. 21: 404. 1886. 

Type locality: “At the Frailes in the mountains above 

Batopilas, at about 7,000 feet, Palmer, 260.” Type specimens: Palmer 260, GH, MEXU, NY, PH, US (Figure 

18). 

Stems erect, probably about 1 m tall. Leaves pubescent to glabrate, lanceolate to oblanceolate, 28-56 mm 

long, 3-7 mm broad, usually mucronulate apically, 

attenuate to base, entire or rarely 1- to 3-toothed, 

slightly revolute, midrib and a few laterals standing 

out below. Inflorescence ascending; pedicels 2-5 mm 

long. Calyx pubescent, 7-13 mm long, 3-5 mm 

broad, the lobes narrowly subulate, 2-8 mm long. 

Corolla red, 26-56 mm long, 6-13 mm broad, the 

upper lip 6-16 mm long, little or not galeate, the 

forward margins rounded or obtuse, usually not reflexed; the lobes of the lower lip 3-5 mm long. 

Stamens subequal, the anthers all fertile and shaggy, 

the lobes mucronate below. Capsules 7-8 mm long, 

5 mm broad. (Figure 16.) 

DISTRIBUTION AND COLLECTIONS EXAMINED (Figure 

17) .-MEXICO. Chihuahua: Near Batopilas, 5500- 

6500 ft, Oct. 1898, Goldman 210, GH, US; Palmer 

260, Aug.-Sept. 1885 (type for L. hyssopifolia). Jalisco: (see type). Durango: Steep slope with Pinus, 

Quercus, and Arbutus, 46 miles north of Estacion 

Coyotes along a lumber road, just northwest of San 

Luis del Rio, 7900 ft, 5 Nov. 1970, D. E. Breedlove 

18773, GAS, US; steep slope with Pinus, Quercus, 

Arbutus, and Juniperus, along Highway 40, 6 miles 

west of La Ciudad, 8800 ft, 7 Nov. 1970, D. E. 

Breedlove 1884, GAS, US. Unlocalized : “Sierra 

Madre Mex. Bor.,” Seemann, BM, C, GH, K. 

CHROMOSOMES. - n= 7, counted by Ernst from 

D. E. Breedlove 18773 (see above). 

This species is distinguished by its broad, lanceolate 

or oblanceolate, mostly entire leaves, subulate calyx 
NUMBER 6 33 

FIGURE 18.--Section Lamourouxia. a, L. longiflora, type collection, Hartweg 188, K; 

b, L. hyssopifolia, type collection, Palmer 260, GH. 

lobes, and large flowers. Whether L. hyssopifolia is 

conspecific is debatable as it is known from a distant 

locality in Chihuahua where the dimensions of the 

leaves and corolla are smaller and a few of the leaves 

are provided with one or two conspicuous teeth. The 

smaller flowers from this northern locality generally 

resemble those of the more common L. nzultifida, 

a relationship that is further emphasized by the rare 

toothed leaves. The larger flowers of typical L. longiflora particularly recall the larger flowers of the type 

collection of L. multifida p grandiflora, Hartweg 187, 

also presumed to have originated from near Bolanios. 

Six collections including 15 presumed duplicate 

specimens have been examined. If one had only a 

single specimen of the Hartweg collection of L. 

longiflora and only one collection of L. hyssopifolia, 

it would seem indisputable that two distinct species 

were represented. When all of the specimens are 

viewed at one time, the range of variation is apparent and suggests that one species is a better gamble 

until shown otherwise by biosystematic information. 

3. L. parayana W. R. Ernst, sp. nov. 

Herba suff ruticosa, ?erecta. Folia pubescentia, ovata 

vel elliptica, plus minusve crenata et scabra, brevissime petiolata. Calyx glandulopubescens, limbis late 

triangularibus, acutis, mucronulatis. Corolla rubra, 

cylindrica, ventricosa, labio [superiore bifido, lobis] 

rotundatis, non vel minime reflexis ; labio inferiore 

late trifido. Stamina omnia fertilia, subaequalia. 

Affinis sectionis Lamourouxia, verosimiliter L. pringlei 

sed folia ut videtur majores et calyces parviores. 

TYPE Loc.aITY.-Estado de Mexico, Cerro de las 

Viboras, cerca de Sultepec, altitude 2500 m, 2 Nov. 

1956. 
34 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

HoLoTYPE.-Ladidao Paray 2242, ENCB (Figure 6-8 mm broad, crenate with 3-10 teeth on a side, 

19). narrowed to the base with a petiole of 1 mm or less; 

Stems thin, probably erect and 1 m or less long. midrib standing out below, somewhat depressed above, 

Leaves membranous, pubescent, slightly scabrous with 2-4 faint laterals. Inflorescence ascending, the 

above, ovate or broadly elliptical, 15-20 mm long, pedicels 4-6 mm long. Calyx glandular pubescent, 

FIGURE 19.--Section Lamourouxia. L. parayana, holotype, Paray 2242, ENCB. 
NUMBER 6 35 

6 mm long, 5 mm broad, the lobes 3 mm long, 

broadly triangular, mucronulate. Corolla red, 30-37 

mm long, 10-11 mm broad, ventricose in midportion 

of tube, upper lip 7 mm long, little or not galeate, 

the forward margins little or not reflexed, lobes of 

the lower lip 3-4 mm long, 2-3 mm broad, rounded. 

Stamens subequal, anthers shaggy, all fertile, the lobes 

strongly mucronate below. Immature capsules ovoid, 

8 mm long, 5 mm broad. (Figure 16) 

DIsTRIsuTIoN.-State of Mexico, see type (Figure 

17). 

The flowers of this species resemble those of L. 

multifida and L. pringlei since the forward margins 

of the upper lip of the corolla are little or not reflexed 

and the anthers are equal in size. Unfortunately, L. 

parayana is known only from a single sheet, and 

without additional material, it is difficult to pinpoint 

the characters separating it from L. pringlei. The 

flowering stems of L. parayana are thin and wirelike, 

a millimeter or less in diameter, and possibly decumbent. The dried leaves appear membranous, broadly 

ovate or elliptical and the nonrevolute margins are 

nearly crenate. The calyx is 6 mm or less long with 

the cup portion only about 3 mm deep. The flowering 

stems of L. pringlei are 2-4 mm in diameter and 

upright. The leaves are more coriaceous, more irregularly serrate or sometimes essentially entire, and the 

margins tend to be revolute or the blades are conc?ve, 

varying in shape from elliptical to spatulate. The 

more prominent calyx is 7-12 mm long with the cup 

portion about 5-8 mm deep. 

4. L. pringlei Robinson & Greenman ex Pringle, 

Garden and Forest 8: 273, fig. 39. 10 July 1895. 

TYPE LOCALITY.-“. , . rocky and wooded summit 

ridges of San Felipe . . . .” 

ledges, Sierra de San Felipe, 10,000 ft., 25 September 

1894,” A, BM, BR, C, ENCB, F, G, GH, K, MEXU, 

MO, MSC, NY, P, PH, RSA, US, W. Not L. pringlei 

“Gandoyer,” = Gandoger, Bull. SOC. Bot. France 66 : 

217. 1919, =L. multifida. 

Stems erect to nearly 2 m tall. Leaves pubescent, 

probably scabrous, the larger 13 ( 19) 24 mm long, 

3(6)9 mm broad, elliptical to obovate or spatulate, 

apically rounded or acute, attenuate below, the margins serrate to crenate with up to 7 teeth on a side 

or entire and revolute, midrib standing out below 

TYPE SPECIMENS.-P?‘ingk 4972, Oaxaca, “Cold 

with 3 or 4 faint lateral veins; petiole to 1 mm long, 

larger leaves on major stems sometimes sessile. Inflorescence ascending; pedicels 3 (4) 5 mm long. Calyx 

glandular pubescent 7( 10) 12 mm long, 4(5)6 mm 

broad, the lobes 3 (4) 6 mm long, broadly triangular, 

entire or with 2 or 3 small teeth, the upper pair of 

calyx lobes separated by a shallow sinus. Corolla red 

or red-orange, 35(44)55 mm long, 6(10) 15 mm 

broad, the upper lip 9( 12) 17 mm long, rounded to 

apiculate, little or not galeate, the forward margins 

rounded, little or not reflexed; the lobes of the lower 

lip 3-5 mm long, 2-3 mm broad, mostly cuspidate. 

Stamens subsequal, all anthers fertile, shaggy, the 

lobes strongly mucronate at base. Capsules ovoid, 

14 mm long, 7 mm broad. (Figure 16.) Altitude: 

1900-3247 m. Flowering: August to January. 

DISTRIBUTION AND NUMBER OF COLLECTIONS EXAMINED (Figure 17). - MEXICO : Hidalgo (6) ; 

Oaxaca ( 11) ; ?Puebla (Esperanza, Purpus 2796, F, 

GH, MO, UC, US). 

The name L. pringlei probably was inadvertently 

putlished by Pringle in a narrative of his travels in 

Oaxaca in anticipation of the more formal description 

by Robinson & Greenman. An adequate plate with 

detailed enlargement of the flower and mention of 

the type locality by Pringle establishes the validity 

of ihis name 20 days before its publication in the 

revision of the genus by Robinson & Greenman, Amer. 

Jour. Sci. 50: 170, and, simultaneously, Contr. Gray 

Herb. 9: 170. 1 August 1895. 

This species is quite variable with two centers of 

distribution within its limited range. With exceptions, 

the leaves of the plants in Oaxaca seem to have more 

rounded or elliptical leaves, and sometimes much 

larger corollas, while the leaves of the plants in 

Hidalgo tend to be more obovate or spatulate, and 

the margins more irregularly toothed. The proportionately large calyx and cuspidate lobes of the lower 

lip of the corolla are notable and help to distinguish 

this species from the probably closely related L. 

parayana. 

5. L,. dispar W. R. Ernst, sp. nov. 

Herba suffruticosa. Folia glabriuscula, anguste elliptica 

vel !anceolata, serrata, attenuata basin versus. Calyx 

pubmcens, limbis serratis. Corolla rubra, late cylindrica, labio superiore parum galeate, bifido, lobis 

rotundato-reflexis, labio inferiore late trifido. Stamina 

omnia fertilia, plus minusve subaequalia vel superiora 
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

breviora. Affinis sectionis Lamourouxia autem corollae formam sectionis Hemispadon simulans. 

TYPE LocALITy.-Guerrero : Near Omiltemi, 20 

miles west of Chilpancingo; deep pine-oak-fir cloud 

forest; 7500-8000 ft. Sprawling shrub to 3 ft long; 

corolla bright red; scarce. 31 July 1957. 

HOLOTYPE.-R. M. Straw €8 D. P. Gregory 1062, 

MEXU (Figure 20). 

IS OTYPES .-MI CH, RSA. 

Stems to 1 m long. Leaves variable, glabrate to 

sparingly pubescent, the larger 20-28 mm long, 3-8 

mm broad, narrowly elliptical to lanceolate, apically 

attenuate, tapered to base with petiole about 1 mm 

long, with 5-15 serrate on a side; midrib standing 

out below, secondary veins faint. Inflorescence ascending, pedicels 5-6 mm long. Calyx pubescent, the 

major veins standing out, 8-11 mm long, 5-6 mm 

broad, the lobes triangular, spreading to reflexed, 

serrate, 5-7 mm long. Corolla red, broadly cylindrical, 

55 mm long, 9-10 mm broad, little or not ventricose, 

rounded distally and slightly galeate, the upper lip 

20-22 mm long, the lobes rounded, partly reflexed, 

the lobes of the lower lip 5-7 mm long, about 3 mm 

broad. Stamens 4, subequally developed, the upper 

pair shorter, the filaments thickened distally, the 

anthers shaggy pubescent, the lobes short acuminate 

basally (Figure 17) , 

As the name implies, L. dispar is enigmatic. The 

stamens are subsequal, resembling those of the larger 

flowers of L. macrantha. The anthers are densely 

pubescent, relatively large, loosely or not connivent, 

and the upper pair of filaments is somewhat shorter 

than the lower, although both pairs of anthers are 

near the mouth of the corolla. The shape of the 

corolla with the lower lip continuing forward rather 

than arching downward recalls the morphology of 

section Hemispadon. The broader leaves recall some 

of the more dentate examples of L. virgata in section 

Adelphidion ; the narrower leaves resemble those of 

L. tenuifolia in Hemispadon. The long red corollas 

and the four subequally developed stamens are the 

reasons for placing L. dispar in section Lamouraouxia. 

This species is represented by three specimens of a 

single collection. It appears to incorporate floral 

characters of section Hemispadon and section Lamourouxia. The elongated, uncrowded inflorescence 

with long flowers, and the long leaves suggests similar 

characters in L. longiflora as well as L. jaliscana. 

The narrow leaves of the holotype resemble those of 

L. tenuifolia, while the leaves of the isotype at Michigan are more like some examples of L. virgata. The 

long, blunt corollas with their lower lip continuing 

forward rather than abruptly curving downward resemble those of L. rhinanthifolia; however, the subequally developed, fully fertile anthers place this 

species in section Lamourouxia. 

6. L. macrantha Martens & Galeotti, Bull. Acad. 

Sci. Bruxelles 12(2) : 32. 1845. 

TYPE LOCALITY.--(‘AUX bords des ruisseaux du 

Cerro San Felipe, prb d’Oaxaca, de 8,000 a 9,000 

pieds, et sur les versants du pic d’orizaba, de 8,000 

a 9,500 pieds . . . Aouit-Octobre . . . H. Gal. Nos. 944 

et 1104.” 

SYNTYPE SPECIMENS.-Galeotti 994, Oaxaca, BR, 

G, P, W; Galeotti 1104, Pic0 de Orizaba, BR, G, W. 

SYNONYMY.- 

L. betonicaefolia Bentham in DC. Prod. 10: 539. 1846. 

Type locality: “In Mexici monte Orizaba (Linden.! n. 

225, Galeotti! n. 944 et 1104. Ghiesbrecht!) . . . (v. in 

herb. Hook. et Mus. par.).” Syntype specimens: Linden 

225, Pic0 de Orizaba, BR, K; Galeotti 944, BR, G, P, W; 

Galeotti 1104, BR, G, W. See syntypes for L. macrantha. 

Stems erect to 1 m tall. Leaves glabrate to pubescent, broadly ovate, basally truncate to cordate, the 

larger 25(50)86 mm long, 15(27)38 mm broad, appearing sessile but petioles 2-8 mm long, margins 

doubly serrate to dentate or crenate, the teeth to 

4 mm long, rounded to mucronulate, 25-56 on a side, 

midrib and 4-9 laterals on a side standing out below. 

Inflorescences ascending, pedicels 6 ( 1 1 ) 2 1 mm long. 

Calyx mostly pubescent, 11(16)26 mm long, 6(8) 16 

mm broad, the lobes 7 (9) 12 mm long, usually broad 

and frequently denticulate. Corolla red or red-orange 

( ?partly yellow), 30 (54) 68 mm long, 10 (14) 18 mm 

broad, frequently ventricose, the upper lip 9 (2 1 ) 29 

mm long, terminally apiculate, little or not galeate, 

forward margins rounded or obtuse, reflexed, the lobes 

of the lower lip 4-6 mm long, sometimes 5 mm broad. 

Stamens subequally developed, all fertile and shaggy 

pubescent, the upper pair sometimes with smaller 

anthers; mature stamens probably exserted and 

spreading; basal lobes of anthers barely acuminate. 

Capsules ovoid, about 15 mm long and 10 mm broad. 

(Figure 2 1 ) . Altitude : 2600-3500 m. Flowering : 

August to October and January, peak August to 

September. 

DISTRIBUTION AND NUMBER OF TYPICAL SPECIMENS 
NUMBER 6 37 

EXAMINED (Figure 22) .-MEXICO : Chiapas (in of Mexico (Temascaltepec, Crucero Agua Blanca, 

pine lands, Mt. Tacana, 2000-4038 m, Matuda 2335, Hinton 4926, GH, US) ; Oaxaca (8) ; Puebla (TesGH, MICH, MEXU, PH) ; Durango (2) ; Guerrero malaquilla, Balls 5335, K, UC, US) ; Veracruz (3). 

(Cerro Teotepec, Tlacotepec, Rtedowski 18581, DS, GUATEMALA: San Marcos (frequent in pine-firENCB, F, MEXU, MICH, TEX) ; Jalisco (2) ; State hardwood forest, San Luis west of Ixchiqah, road 

FIGURE 20.--Section Lamourouxia. L. dispar, holotype, Straw B Gregory 1062, MEXU. 
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 2 1.-Section Lamourouxia, diagrams of flowers and 

leaves. a, L. macrantha, Pringle 4854, MEXU; a', Pringle 

4762, PH; b, L. stenoglossa, Breedlove, 15127; c, L. ovata, 

Conzatti 2273, & 2294, F; d, L. dependens, Holway 568, 

US; d', Skutch 154, PHILA; e, L. xalapensis, Pringle 

8886, MSC, showing laciniate calyx; e', Smith 676, US, 

showing regular calyx; e", Hinton 12076, US. 

to Tacana, ca. 3400 m, Beaman 2342, ENCB, GH, 

MSC, TEX, UC, US). 

DISTRIBUTION AND COLLECTIONS EXAMINED WHICH 

APPEAR TO BLEND CHARACTERS OF L. macrantha AND 

L. stenog1ossa.-GUATEMALA : Chimaltenango (6 

miles west of Tecpan, road to Quezaltenango, White 

5226, F, MICH); Huehuetenango (road to San 

Pedro Soloma, 3 miles southwest of San Mateo 

Ixtatan, Breedlove 11556, DS, F, MICH: Sierra de 

10s Cuchumatanes, Skutch 1252, F, GH, PH, US; 

between San Mateo Ixtatan and Cruz de Limon, 

Sierra de 10s Cuchumatanes, Steyermark 48500, F; 

Cruz de Limon between San Mateo Ixtatan and 

Nuca, Steyermark 49813, F; Cerro Chemal, summit 

of Sierra de 10s Cuchumatanes, Steyermark 50295, 

F, PH) ; Jalapa (Soledad Montania Miramundo between Jalapa and Mataquescuintla, Steyermark 32646, 

F) ; Quezaltenango (Volcan Zunil, Steyermark 34922, 

This species is distinguished by its rigidly upright, 

sparingly branched habit often with a single flowering 

raceme, more or less sturdy, ascending pedicels, usually large, broadly ovate, cordate, subsessile leaves 

and bracts with peculiar double marginal incisions, 

relatively large calyx with broad lobes, apiculate corolla, and the absence of mucros on the base of the 

anthers. The distribution is scattered in Mexico, and 

F) * 

the plants must be conspicuous for their bright red 

flowers. Specimens from Oaxaca, including the type, 

have very large corollas, but like the leaves, they are 

variable with two extremes in size, some with corollas 

approximately 30 mm long and others exceeding 60 

mm long, these preshmed to occur in the same vicinity. The larger flowers appear to have exposed 

stigmas while the smaller seem to have included stigmas; however, the samples are small and inconclusive. 

Away from Oaxaca the corollas are more or less 

intermediate in size. 

The leaves and to a certain extent the smaller corollas of L. macrantha resemble L. ouata so that a 

number of specimens have been incorrectly labeled. 

Although poorly known and rare, L. ovata shows some 

characteristics such as reclining stems, nearly right 

angle branching, depressed inflorescences with inverted flowers which clearly distinguish it from L. 

macrantha. The relationship between these two species 

probably is more remote than the relationship between L. macrantha and L. stenoglossa, which is 

much more surprising since these two at first seem 

to have so much less in common. 

Along the southern border of Mexico between 

Chiapas and Guatemala in the vicinity of Mt. Tacana, 

two collections-Matuda 2335, Chiapas, and Beaman 

3245, San Marcos-are indistinguishable from L. 

macrantha and establish this species as one of the 

four that crosses the Isthmus of Tehuantepec, and it 

is one of the two of these exhibiting taxonomic problems in Central America. 

Elsewhere in Guatemala in the departments of 

Chimaltenango, Huehuetenango, Jalapa, and Quezaltenango, several collections appear to combine characters of L. stenoglossa and L. macrantha so that it 

is perplexing to provide a name for these morphologically intermediate plants. From a taxonomic point 

of view, these are neither typical L. macrantha nor 

L. stenoglossa, and yet they do not seem likely to be 

a separate taxon. For the time being, these Guatemalan collections are referred to L. macrantha, but 

it is fully realized that they also share characters with 

L. stenoglossa. 

7. L. stenoglossa F. W. Hunnewell & L. B. Smith, 

Contr. Gray Herb. 124: 4, pl. 1, fig. 11-12. 1939. 

TYPE LocALITY.--('Mexico: Chiapas: . . . Cerro de 

Huitepeque, alt. 2300 m., 1864-70, Ghiesbreght 706 

(GH, TYPE) . . , ." 
NUMBER 6 39 

HoLoTYPE.-Gkiesbregkt 706, GH. 

ISOTYPES.-BM, F, K, MO, NY. 

Stems erect to 1 m long. Leaves glabrate to pubescent, the larger 22(41)57 mm long, 9(11)15 mm 

broad, narrowly ovate to lanceolate or elliptical, mostly 

truncate basally, petiole 1-2 mm long, margins serrate 

or dentate, often doubly so, the teeth 16-38 on a 

side, midrib standing out below, the 7-9 laterals on 

a side faint. Inflorescences ascending; pedicels thin 

and often somewhat contorted, 9 (1 1) 15 mm long. 

Calyx pubescent, 8(10) 12 mm long, 4(5)6 mm 

broad. Corolla red or red-orange, 39 (43) 46 mm long, 

10 ( 1 1 ) 12 mm broad, mostly ventricose, the upper 

lip 12( 17) 20 mm long, apiculate, little or not geleate, 

the forward margins rounded, reflexed; the lobes of 

the lower lip 2-3 mm long, 2-3 mm broad. Stamens 

subequal or the upper pair shorter, anthers equal in 

size, shaggy, probably exserted and spreading, little 

or not mucronate. Capsules ovoid, 11 mm long, 9 mm 

broad, (Figure 21). Altitude 2287-2866 m. Flowering: July to September and January. 

DISTRIBUTION AND COLLECTIONS EXAMINED (Figure 

22). MEXICO : Chiapas (Chamula, Zontehuitz, 9400 

ft, 30 July 1964, Breedlove 6706, DS, ENCB, F, 

MICH; San Cristobal las Casas, Zontehuitz, 8770 ft? 

19 Jan. 1965, Breedloue B Raven 8107, DS, ENCB, 

F, MICH, US; San Cristobal las Casas, Zontehuitz, 

9300 ft, 20 Sept. 1965, Breedlove 12349, DS, F: 

MEXU, MICH; San Cristobal near Rancho Nuevo, 

9000 ft, 20 Aug. 1966, Breedloue 15127, DS, MEXU; 

Tenejapa, Achlum, 9100 ft, 23 Aug. 1966, Breedloue 

15179, DS; Tenejapa, Matsab, 25 Aug. 1966, Breedlove 15282, DS, MEXU; Ghiesbreght “74, 173, 176,” 

GH; Ghiesbreght 706, see type; near San Cristobal. 

7000-8000 ft, 18-22 Sept. 1895, Nelson 3212, GH, 

The collections most closely resembling the holotype 

and paratypes of this species are restricted to Chiapas, 

US). 

FIGURE 22.-Distribution of section Lamourouxia in Mexico and Guatemala. Localities in 

Guatemala of specimens appearing to blend characters of L. macrantha and L. stenoglossa 

shown by open, five-pointed stars, and localities of glabrate L. dependens shown by open squares. 
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

where they are distinct and ought not to be confused 

with any other taxon. The apparently branched habit 

with more than one raceme, the erect inflorescence: 

the apiculate, ventricose corollas on long wiry pedicels, and the often double incisions of the narrow 

and petiolate leaves will help to confirm this opinion. 

This species resembles L. dependens and L. xalapensis, but is separated from them by the erect inflorescence, hence lack of inverted flowers, At first 

glance, one would not expect a taxonomic problem 

with L. macrantha, of scattered distribution in Mexico, but now believed also to occur in Guatemala. 

In general, the better known Mexican collections of 

L. macrantha tend to be less branched with mostly 

a single raceme that is rigidly erect, the pedicels are 

more rigid and erect, the flowers and calyx usually 

are larger, and the leaves broader and more nearly 

sessile or at least their cordate bases concealing the 

short petiole. Most of the several Guatemalan collections (unlike the Mexican) bear a strong resemblance to L. stenoglossa and thereby diminish the discontinuity between these species. See discussion under 

L. macrantha. 

8. L. ovata Martens & Galeotti, Bull. Acad. Sci. 

Bruxelles 12 (2) : 33. 1845. 

TYPE LOCALITY .-“Dans les montagnes calcaires de 

Zeta, pres de Sola (sud d’Oaxaca) , a 8,000 pieds . . . Septembre . . . H. Gal. No. 989.” 

TYPE SPECIMENS.-Galeotti 989, BR, G. 

Stems reclining. Leaves pubescent, broadly ovate, 

33-58 mm long, 22-37 mm broad, cordate at base 

and sessile, obtuse, doubly serrate to crenate with 

25-31 teeth on a side, veining standing out below 

with 5-8 laterals. Inflorescence arching forward, the 

tip depressed, pedicels 6-9( 13) mm long, reflexed 

away from the growing tip, the flowers inverted. 

Calyx sparsely pubescent, 14-18 mm long, 9-10 mm 

broad, the lobes 8-11 mm long and 8 mm broad, the 

sinus folds projecting outward. Corolla red or redorange, 40-42 mm long, 10-11 mm broad, the upper 

lip 13 mm long, little or not galeate, the forward 

margins broadly reflexed. Lobes of the lower lip 6 mm 

long, 4 mm broad, rounded. Stamens subequal, exserted and spreading, the anthers all fertile and pubescent, the lobes little or not mucronate at base. 

(Figures 21, 23). 

DISTRIBUTION AND COLLECTIONS EXAMINED (Figure 

22) .-MEXICO : Oaxaca (Cerro de San Felipe, 

2000 m, 10 Oct. 1897, Conzatti &? Gonzales 508, GH; 

Cerro San Felipe, 2000 m, 18 Oct. 1908, Conzatti 

2273, F; Cerro San Felipe, 2000 m, 20 Sept. 1908, 

Conzatti 2294, F; Galeotti 989, see type). 

Only four collections of this species are known, 

the type having come from a locality that is presumed 

to be south of the city of Oaxaca, while the others 

are from northeast of the city. The leaves are very 

broad, cordate, and sessile; the very large calyx is 

sparsely pubescent with most of the hairs along the 

major veins and the margins of the lobes. The name 

L. ouata frequently has been misapplied to plants of 

L. macrantha, presumably because of the large, ovate, 

cordate leaves of the latter. The branches and inflorescences of L. macrantha are ascending, while 

L. ovata is reclining with nearly right-angle branching, depressed inflorescences, and inverted flowers. 

The closest affinity probably is with L. dependens of 

Guatemala. 

9. L. dependem Bentham in DC. Prod. 10: 539. 

1846. 

TYPE LocALITY.-“in Guatimala (Skinner! . . . 

TYPE SPECIMENS.-skinner, K (two specimens) . SYNONYMY.- 

(v. in herb. Hook.).” 

L. dependens Bentham var. apiculata Hunnewell & L. B. 

Smith, Contr. Gray Herb. 124: 4, pl. 1, fig. 8. 1939. 

Type locality: “Guatemala: Chimaltenango: . . , “Santa 

Elena,”. alt. 2400-2700 m, Jan. 1933, Skutch 154 (US, 

TYPE . . .).” Holotype: Skutch 154, US 1494811. 

Isotypes: A, MICH, PH. 

L. montana Hunnewell & L. B. Smith, Contr. Gray Herb. 

124: 4. 1939. Type locality: “Guatemala: Chimaltenango: Volcan Acatenango, alt. 2450 m., Feb. 1937, 

Hunnewell 14856 (G, TYPE) .I’ Holotype: Hunnewell 

14856, GH. 

Stems weak or reclining, to 3 m long. Leaves mostly 

pubescent, sometimes glabrate, ovate or broadly elliptical, 28(37)56 mm long, 10(17)25 mm broad, 

rounded or acute apically, blade truncated or obtuse 

basally, margins doubly serrate or dentate with 12-25 

mucronulate teeth on a side; veining standing out 

below with 4 or 5 laterals; petiole distinct, 3-7 mm 

long. Inflorescence arching downward, the tip depressed; pedicels 6( 10) 17 mm long, usually reflexed 

away from growing tip, the flowers inverted. Calyx 

mostly pubescent, often glandular, 8( 12) 16 mm long, 

frequently one-third or one-fourth the length of the 

corolla, 5 (7) 10 mm broad, the lobes 5 (8) 13 mm long, 
NUMBER 6 41 

FIGURE 23.-Section Lamourouxia. L. ouata, Conratti 2273, F. 

usually broadly acute, frequently terminally dentate, strongly reflexed; lobes of the lower lip 2-5 mm long, 

the axil of the sinuses revolute. Corolla red or red- about 3 mm broad, mostly obtuse. Stamens subequally 

orange, 36 (40) 44 mm long, 8 ( 10) 11 mm broad, not developed, sparsely pubescent, all fertile, the lobes 

galeate, the upper lip 11 ( 13) 15 mm long, minutely slightly or not acuminate at base, becoming exserted 

apiculate or emarginate distally, the lobes obtuse and and spreading. (Figure 2 1 ) , Altitude : 1950-3300 
42 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

( ?or more) m. Flowering: December to April, peak 

January to February. 

DISTRIBUTION AND NUMBER OF TYPICAL COLLECTIONS EXAMINED (Figure 22). - GUATEMALA : 

Chimaltenango (9) ; Huehuetenango (2) ; Quezaltenango (6) ; QuichC (3) ; SacatepCquez (4) ; San 

Marcos (between San Sebastian and San Marcos, 

Steyermark 35662, F) ; Solo1a (2) ; Totonicapan (between Quezaltenango and Los Encuentros, Molina, 

Burger, B Wallenta 16416, F, W); and three unknown localities. Doubtful : Warsiwicr 9, “Costa Rica 

et Veragua,” G. 

DISTRIBUTION OF SPECIMENS ANNOTATED AS L. 

dependens BUT REQUIRING SPECIAL COMMENT. - 

GUATEMALA: Huehuetenango (near San Juan 

Ixcoy, Breedlove 8554, DS, F, US; near San Mateo 

IxtatAn, Breedlove 8766, DS, F, US; Chemal, Molina, 

Burger, B Wallenta 16569, F) ; San Marcos (near 

Volcan Tacana, Steyermark 36329, F; near Aldea 

Fraternidad, Williams, Molina, & Williams 25650, 

F, NY, US and 26090, BM, F NY, US, W) . This species is recognized by its reclining habit, 

distinctly petiolate and usually pubescent leaves, depressed inflorescences, inverted flowers, relatively large 

and often glandular pubescent calyx with frequently 

dentate lobes, and the exserted and spreading anthers 

which are only slightly mucronate. The calyx and 

corollas are very similar to those of L. ovata, and 

both species have dependent inflorescences. The leaves 

of L. dependens are petiolate and relatively narrow, 

while those of L. ovata are very broad, sessile, and 

clasping. Not only are the leaves very different, but 

these species also are separated geographically by the 

Isthmus of Tehuantepec. 

The several collections listed above as requiring 

comment are closest to L. dependens but differ because they seem to have somewhat narrower leaves 

and smaller, glabrate or glabrous calyx. This is perplexing, because characters of the calyx distinguish 

typical L. dependens from other species, and characters of the calyx also distinguish these few specimens 

from L. dependens. These specimens may represent 

an undescribed taxon; however, for the time being 

their strongest morphological ties appear to be with 

L. dependens, because there appear to be intennediates. They also resemble L. xalapensis; however, if 

this were their closest specific relationship, they would 

be the only examples of that species south of the 

Isthmus of Tehuantepec. 

The main character distinguishing L. dependens 

var. apiculata was the mucronulate rather than obtuse 

lobes of the lower lip of the corolla. The main character separating L. montana was the minutely emarginate, rather than obtysely cuspidate, termination of 

the upper lip of the corolla. Another difference now 

apparent in the latter is the tendency for shorter 

internodes and a more compact habit. Among the 

specimens definitely closely allied to L. dependens, 

these characters appear to be of low taxonomic magnitude. Both taxa are included in L. dependens and 

neither, I believe, affects the enigmatic relationship 

of L. dependens to the glabrate form in Guatemala. 

10. L. xalapensis H.B.K., Nov. Gen. 2: ed. folio 272, 

ed. quarto 338. “1817,” 1818. 

TYPE LocALITY.-“Crescit in declivitate orientali 

montium Mexicanorum prope urbem Xalapa, alt. 680 

hex . . . Floret Februario.” 

TYPE SPECIMENS.-xalapa, P (in Humboldt & 

Bonpland Herbarium and possibly two sheets in 

General Herbarium, e.g., Herb. Al. de. Bunge, Herb. 

E. Cosson). 

SYNONYMY.- 

L. spathacea Bentham in DC. Prod. 10: 539. 1846. Type 

locality: “In monte Orizaba (Linden! n. 226), in quercetis prov. Oaxaca (Galeotti! n. 1007 bis ex parte) . . . (v. in herb. Hook.) .” Syntype specimens: Linden 226, 

BR, K; Galeotti 1007 bis, BR, K. 

L. paruiflora W. B. Hemsley, Biol. Cent.-Amer. 2: 465. 

1882. Type locality: “Mexico, without locality (Tate, 

31; Mairet, 56). Hb. Kew.” Syntype specimens: Tate 

31, K; Mairet 56, K. 

L. exserta Robinson h Greenman, Amer. Jour. Sci. 50: 171. 

1895 (Contr. Gray Herb. 9: 171. 1895). Type locality: 

“Valley of Mexico, Bourgeau, 986, Schaffner, 367; and 

in rich ravines of Sierra de Clavellina, Oaxaca, altitude 

9,000 feet, 18 October, 1894, Pringle, 4995; also on top 

of Sierra Madre near Chilpancingo, altitude 9,000 to 

10,200 feet, 24 December, 1894, Nelson, 2216, and at 

Tlalixtaquilla, Nelson, 2261 .” Syntype specimens: Bourgeau 986, BR, C, G, K, UC, US; Pringle 4995, BR, 

CAS, G, GH, K, MEXU, MO, MSC, NY, PH, RSA, 

US, W; Nelson 2216, US; Nelson 2261, not found; Schaffner 367, K, NY, P. 

Stems 1-4 m long, weak or reclining. Leaves glabrous or glabrate, infrequently pubescent along major 

veins and along margins, narrowly lanceolate to ovate, 

mostly attenuate apically, broadly acute or tapered 

to the base, 27(48) 63 mm long, 6( 12) 20 mm broad, 

the petiole 1 (3)6 mm long, margins mostly finely 
NUMBER 6 43 

serrate with 20-30 often-mucronulate teeth on a side, 

veining standing out below and sometimes also above, 

with 8-1 1 secondary veins. Inflorescences arching 

downward, the tip depressed; pedicels filiform, 2 (6) 12 

mm long, usually reflexed away from growing tip, the 

flowers inverted. Calyx mostly glabrous, 8(9) 12 mm 

long, 4-5 mm broad, the lobes 3 (5) 7 mm long, entire 

and usually triangular or calyx sometimes appearing 

spathiform, 8( 15) 18 mm long, deeply cleft on the 

lower side with about 4 narrowly elongate lobes on 

the upper side. Corolla red or red-orange, 28(40)48 

mm long, 8 ( 1 1 ) 12 mm broad, often appearing slightly 

arched, the upper lip 9 ( 11) 16 mm long, not galeate, 

often emarginate, the forward margins broadly 

rounded and reflexed, 4 mm broad, the lobes of the 

lower lip to 4 mm long. Stamens subequally developed, the anthers shaggy, not mucronate, finally 

exserted and spreading. Capsules spheroidal to ellipsoidal, 9-10 mm long, 7-9 mm, broad. (Figure 21). 

Altitude: 1220-3350 m. Flowering: July to April, 

peak November to December. Frequently mentioned 

as occurring in damp thickets or mixed forests. 

DISTRIBUTION AND NUMBER OF COLLECTIONS EXAMINED; “s” INDICATES SPATHIFORM CALYX (Figure 

22) .-MEXICO : Distrito Federal (5) ; Guerrero 

(11) ; Hidalgo (2s) ; Jalisco (5); State of Mexico 

(11); Michoacan (Coalcoman, 2500 m, 20 Dec. 

1938, Hinton 12798, BR, DS, G, MICH, RSA, TEX, 

UC, US) ; Morelos ( 1 + Is) ; Oaxaca (8+ 5s) ; Puebla 

(3 +4s) ; Veracruz (1 +8s). See also L. dependens 

for unlikely extension of range to Guatemala. 

This species is distinguished by its clambering habit, 

wide- to nearly right-angle branching, depressed inflorescence tips with reflexed and inverted flowers, 

usually long lanceolate, glabrous leaves, and glabrous 

calyx. The calyx is notable for exhibiting two distinctive morphological extremes. In the one, the calyx 

is more or less equally four-lobed and barely bilabiate; 

in the other, the calyx tube is narrowly elongated, 

deeply cleft on the lower side, and more or less laciniate on the upper side. The latter situation appears 

to be the basis for Bentham’s L. spathacea. There 

may be a weak correlation between longer leaves and 

the longer, spathiform calyxes. 

The spathiform calyx is distinctive and perhaps 

should receive infraspecific taxonomic recopition ; 

however, there is intergradation with the more regular 

form and sometimes variation on a single specimen. 

The calyxes on the presumed type specimen at Paris 

from “Xalapa” are not well preserved, but they appear to be more or less equally lobed. This species is 

believed to be restricted to Mexico west of the Isthmus 

of Tehuantepec. See discussion of unusual specimens, 

cited under L. dependens, which resemble L. xalapensis. 

11. Section Adelphidion W. R. Ernst, sect. nov. 

Species 11-14. 

Corolla tuba plus minusve infundibuliformis, non 

ventricosa, subrosea vel lavandulacea vel magentea, 

non rubro-aurantiaca. Stamina dimorphus, omnia 

fertilis, inferiora longiores et majores. 

Stamens strongly unequal, the anthers dimorphic, 

the lower pair with somewhat thickened filaments and 

two large, shaggy pubescent anthers near the mouth 

of the corolla or barely exposed, the upper pair of 

stamens shorter, always included, with filiform filaments and smaller, but fertile anthers. Corolla lavender, pink, or magenta (not red), the tube more or 

less funnelform (Figures 8, 10, 24). 

TYPE SPECIES.--L. dasyantha (Cham. & Schlecht.) 

W. R. Ernst. 

DISTRIBUTION: Two species in Mexico and two 

species in northwestern South America (Figures 25, 

27). 

11. L. brachyantha Greenman, Proc. Amer. Acad. 

41 : 248. 1905 (Contr. Gray Herb. 31 : 248 1905). 

TYPE LOCALITY.-“State of Mexico : Rocky hills, 

Lecheria, Valley of Mexico, altitude 2285 m. 1 July, 

1904, C. G. Pringle, no. 8815 (hb. Gr.) .” 

HoLoTYPE.-Pringle 8815, GH. 

ISOTYPES.-BM, C, CAS, F, G, K, MEXU, MO, 

MSC, NY, PH, UC, US. 

Stems to 1 m tall. Leaves and bracts coriaceous, 

glandular pubescent, 18(25) 35 mm long, 8( 13) 18 mm 

broad, ovate, truncate to cordate, sessile, clasping; 

margins coarsely 9-22 dentate on a side; secondary 

veins 3-5 (9), prominent below. Inflorescence ascending; pedicels 2-4 mm long. Calyx glandular pubescent, 5(8) 11 mm long, 5(6) 7 mm broad, the lobes 

broadly triangular, 3 (4) 6 mm long, conspicuously 

dentate. Corolla pink or violet ( ?white), conspicuously galeate, 15 (18) 22 mm long, 8-1 1 mm broad; 

upper lip 5(7)9 mm long, the lobes obtuse, reflexed; 

lobes of lower lip about 3 mm long, 2-3 mm broad, 

obtuse. Stamens dimorphic, anthers included, upper 
44 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

pair smaller, all fertile. Style pubescent, the stigma 

about 1.5 mm broad, rounded, barely exserted. Capsules 10-12 mm long, 8 mm broad. Altitude: 2200- 

2800 m. Flowering: July to September. Occasionally 

on calcareous soil. (Figure 24). 

DISTRIBUTION AND NUMBER OF COLLECTIONS EXAMINED (Figure 25) .-MEXICO : Distrito Federal 

(5) ; Hidalgo (near Tepiji del Rio, Strare, &3 Gregory 

1170, RSA) ; State of Mexico (7) ; Quer6taro (2). 

This species is distinguished by the conspicuously 

galeate corollas which are about as long as broad 

with broad, open mouth. The anthers remain included, and the relatively large stigma is barely exserted. Greenman (1905) placed this species in section 

Hemispadon next to L. rhinanthifolia, presumably 

because of the similarity of the leaves and calyx. This 

species probably is not very closely related to any 

other and is placed with section Adelphidion because 

of the development of the stamens and the non-red, 

short corolla. 

12. L. dasyantha (Chamisso & Schlechtendal) W. 

R. Ernest. comb. nov. Basionym: Gerardia dasyantha Chamisso & Schlechtendal, Linnaea 5 : 104. 

1830. 

TYPE LoCALITY.--“Tierra fria. Sept.” 

PRESUMED HoLoTYPE.-schiede 165, B, destroyed 

in war. Pre-war photos F, PH, US (Figure 26). 

Fragment of original type, PHIL (see Pennell, F. W., 

Proc. Acad. Sci. Philadelphia 80: 345. 1928; ibid., 

81: 246. 1929). 

SYNONYMY.- 

Lamourouxia linearis Bentham in DC., Prod. 10: 540. 1846. 

Type locality: “In planitie Mexicana (G.J. Graham! 

Berlandier! Alaman! Mendez! Galeotti! n. 1007 . . .v.s.).” 

FIGURE 24.-Section Adelphidion, diagrams of flowers and 

leaves. a, L. brachyantha, Pringle 13593, US; b, L. dasyantha, Pringle 7370, PH; b‘, Langman 3047, PH; c, L. virgata, 

Haught 3285, US; c’, Killip d Smith 17350, US; c“, Dryander 1775, US, capsule; d, L. sylvatica, Tovar 164, US; 

d‘, Ochoa 348, US. 

Syntype specimens: Graham 285, plains of Papetonga, 

BM, K; Berlandier, GH, K; Alaman (not seen) ; Mendez 

(not seen) : Galeotti 1007, BR. G, K, W. 

Gerardia punctata Robinson, Proc. Amer. Acad. 26: 172. 

1891 (Contr. Gray Herb. 1: 172. 1891). Type locality: 

“Hills of Zacatecas; October, 1888 (n. 2183).” Type 

specimen : Pringle 2 138, GH. 

Lamourouxia tenuifolia Martens & Galeotti var. mkrantha 

Greenman, Proc. Amer. Acad. 35: 313. 1900 (Contr. 

Gray Herb. 18: 313. 1900). Type locality: “Collected 

on the west side of Valley of Cuicatlan, Oaxaca, altitude 

2,000 to 2,100 m, by E. W. Nelson, 10 November, 1894, 

no. 1891. Types in herb. Gray and herb. US. Nat. 

Museum.” Isotypes: Nelson 1891, GH, US=L. dasyantha, not L. tenuifolia. 

Stems erect to 1 m tall, frequently branching from 

below. Leaves glabrous to glabrate, 15(29)32 mm 

long, 1 (3) 4 mm broad, subulate to lanceolate, acute 

to obtuse apically, tapered or rounded to the base, 

sessile or petiole less than 1 mm long, margins entire 

to subserrate with 5-9 incisions on a side, sometimes 

revolute ; midrib standing out below. Inflorescences 

ascending, pedicels 2 (3) 4 mm long. Calyx glabrous 

to glabrate, 9 ( 11 ) 13 mm long, 3 (4) 5 mm broad, 

the major veins standing out, the lobes broadly subulate, 5( 7) 10 mm long; occasionally with a short, 

fleshy appendage in the axil of the sinus. Corolla 

pink, lavender, mauve, purplish, 27 (32) 35 mm long, 

8 (9) 11 mm broad, little or not galeate, the upper lip 

8 (10) 13 mm long, the lobes obtuse, strongly reflexed: 

the lobes of the lower lip obtuse, 3-5 mm long and 

broad. Stamens dimorphic, all anthers fertile, the 

smaller pair always included, the larger pair sometimes barely exposed, loosely connivent or spreading. 

Capsules 10 mm long, 6 mm broad. Altitude: 1200- 

3000 m. Occasionally on limestone. Flowering: July 

to December, peak October to November. Figures 

24, 26.) 

DISTRIBUTION AND NUMBER OF COLLECTIONS EXAMINED (Figure 25) .-MEXICO : Coahuila (Carneros Pass, Pringle 3682, GH) ; Distrito Federal (12) ; 

Hidalgo (12) ; State of Mexico (17) ; Michoacan (3) ; 

Nuevo Le6n (14 miles north of Matehuala, Ripley €3 

Barneby 13292, CAS, NY); Oaxaca (12); Puebla 

(10) ; QuerCtaro (mountains of QuerCtaro, Bru8 

1085, MEXU) ; San Luis Potosi (10) ; Tlaxcala (2) ; 

Zacatecas (6). 

This species is distinguished by the tufted or branching habit, the long, narrow, essentially glabrous and 

entire, or obscurely serrated leaves, and the large 

palate which nearly occludes the corolla mouth. One 
45 NUMBER 6 

FIGURE 25.-Distribution of section Adelphidim in Mexico. 

of the extremes is the lush biotype that Bentham 

named L. linearis which has larger flowers and 

broader leaves and recalls L. virgata of South America. The plants described by Robinson as Gerardia 

punctata usually have smaller flowers with dark, 

speckled calyx and almost linear, terete leaves in the 

dried condition. Approximately intermediate are those 

plants matching the fragment and the photo of the 

type of Gerardia dasyantha. These variations are conspicuous, but I believe they are best combined until 

there is biosystematic evidence to the contrary. Specimens have been misidentified frequently in herbaria 

as L. tenuifolia, presumably because the leaves are 

similar, but this is a valid name for a distinctive 

species in section Hemispadon. Greenman’s trinomial 

L. tenuifolia var. micrantha is an example of this 

confusion and his type specimen for the varietal name 

is equivalent to L. dasyantha. 

13. L. sy2vatica H.B.K., Nov. Gen. 2: ed. folio 272, 

ed. quarto 337. “1817,” 1818. 

TYPE LOCALITY.-“CreSCit in sylvis prope pagum 

Ayavaca Peruvianorum alt. 1400 hex . . . Floret 

Augusto.” 

TYPE SPECIMEN.-Ayavaca, n. 3465, P (in Humboldt & Bonpland Herbarium, and in General Herbarium). 

L. breviflora Bentham in DC. Prod. 10: 540. 1846. 

SYNONYMY.- 

Type 

locality: “In Peruviae prov. Chachapoyas (Mathews ! 

n. 2119 et 3138 . . . (v.s.).” Syntype specimens: Mathews 2229, K; Mathews 3238, BM, G, GH, K. 

L. subincisa Bentham in DC. Prod. 10: 540. 1846. Type 

locality: “In Andibus Quitensium (Jamison!) . . . (v. in 

herb. Hook.) .” Type specimens: Herb, Hook. “Andes 

Peru McLean,” K; Herb. Benth. “Andes Hooker Jamison 

1845,” K. 

Stems to 70 cm tall, perhaps longer. Leaves pubescent, sometimes scabrous or rugose, to glabrate and 

smooth, the larger 29(33) 44 mm long, 6(9) 14 mm 

broad, often smaller, elliptical, lanceolate or oblanceolate, broadly attenuate basally, the margins shallowly 

to deeply 3-17 serrate or dentate on a side, the teeth 

obscure to 2 mm long; midrib and/or 3 veins standing 
46 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 26.--Section Adelphidion. a, L. dasyantha (presumed holotype of Gerardia dasyantha, 

Schiede s.n., destroyed in Berlin, photo distributed by Field Museum) ; b, similar equivalent, 

C. L. Lundell 12560, US. 

out below, sometimes reticulate. Inflorescences ascending, pedicels 2-3 mm long. Calyx glabrate to scabrous 

pubescent, 7(9) 10 mm long, 4(5)6 mm broad, the 

lobes triangular, 3 (5) 7 mm long, sometimes serrate. 

Corolla rose-pink to purple or white, 13(20)24 mm 

long, 5(8)9 mm broad, the upper lip 5( 7) 10 mm 

long, little or not galeate, the forward margins 

rounded, reflexed, the lobes of the lower lip 2-4 mm 

long, 4 mm broad, rounded. Stamens dimorphic, the 

upper pair shorter, the anthers always included, 

smaller, usually fertile, the lower pair longer and 

larger, sometimes exserted. Capsule ovoid, 9-12 mm 

long, 7-9 mm broad. (Figure 24). Altitude: 2080- 

3450 m. Flowering: February to July and October, 

peak April to May. 

DISTRIBUTION AND NUMBER OF COLLECTIONS EXAMINED (Figure 27) .-COLUMXA: (Lobb, “Columbia,” K, doubtful). ECUADOR: Azuay (3) ; 

Caniar (5-8 km NE of Azogues 1 Apr. 1945, Camp 

2521, NY). Loja (2). PERU: Amazonas (9); Ancash (9) ; Cajamarca (11) ; Huancavelica (Parapuquio below Conaica, 3300-3350 m, 13 Mar. 1951, 

Tovar 164, US); Junin (4); La Libertad (5 km 

south of Otuzco, 2600 m, 24 Apr. 1957, Sagastegui 

58, US) ; Pasco (2). 

This species probably is closest to L. uirgata but 

is distinguished by the corollas which are consistently 

smaller with proportionately larger palates. The leaves 

are exceedingly variable and, although mostly smaller, 

the larger ones exceed those of L. virgata. In habit, 

some specimens resemble the South American species 

of the genus Bartsia; however, this probably is only 
NUMBER 6 

FIGURE 2 7.-Distribution of section Adelphidion in South America, showing range of L. virgata 

in Colombia and Ecuador, and L. sylvatica in southern Ecuador and Peru. 

coincidental as the latter have very different seed 

characteristics. 

The name L. sylvatica has been ignored because 

the original description did nat seem convincing, the 

species was not illustrated, and the type is incomplete. 

Both “subvolubilis” and “volubilis” appeared in the 

description, but I believe this was an exaggeration. 

The stems of some species of Lamourouxia are not 

strong and longer ones, including those of L. sylvatica, probably are supported by surrounding vegetation. The stems of at least three species in North 

America are so weak as to arch through and over 

other vegetation, but even these do not twine. The 

original description concluded with “Specimina suppetentia carie valde destructa.” The presumed type 

in Paris has capsules, but there are only a few fragments of bracts or leaves and no flowers. The geographic locality, “Prope pagum Ayavaca” is in Peru, 

and I believe south of the range of L. virgata. The 

mostly pubescent calyx suggests the identity of this 

taxon as equivalent to Bentham’s L. breviflora and 

L. subincisa. 

Although Bentham placed his L. breviflora and L. 

subincisa in two different sections, I find the type 

specimens sufficiently similar to be conspecific with 

L. sylvatica, hence, originating from Peru or extreme 

southern Ecuador. Whether this version of L. sylvatica 

includes more than one taxon cannot be answered 

from herbarium studies at this time, as any noteworthy characters appear to be recombined in other 

specimens. Pennell had the advantage of collecting 

Lamourouxia in Peru and used at least three additional manuscript names besides Bentham’s L. subincisa for plants that I refer to L. sylvatica. The important point is that Lamourouxia exhibits two 

classes of flowers in South America: the larger are 

L. virgata in Colombia and Ecuador, and the smaller 

are L. sylvatica, in southern Ecuador and Peru. 

Further subdivision of either species will not change 

this relationship. 
48 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

14. L. virgata H.B.K., Nov. Gen. 2: ed. folio 270, 

ed quarto 335, pl. 167. “1817,” 1818. 

TYPE LocALITY.-“Crescit prope urbem Quito, alt. 

1500 hex . . . Floret Majo.” 

TYPE SPECIMEN.-Quito, n. 3009, P. (in Humboldt 

& Bonpland Herbarium and in General Herbarium). 

SYNONYMY.- 

L. serratifolia H.B.K., Nov. Gen. 2: ed. folio 271, ed. quarto 

336, pl. 168. “1817,” 1818. Type locality: “Crescit 

juxta Santa Fe de Bogota et catarractam Tequendamae 

in Regno Novo-Granatensi, alt. 1360 hex , , . Floret 

Junio.” Type specimens: Santa Fe de Bogota, Bonpland, 

P (in General Herbarium, three specimens, not in Humboldt & Bonpland Herbarium), F, US. 

L. loxensis Bentham, Plantae Hmtwegianae, p. 147. 1844. 

Type locality: “In montibus Loxa.” Type specimens : 

Hartweg 824, K (two specimens). 

Stems to 2.5 m tall. Leaves glabrous to pubescent, 

19(26)31 mm long, 2(5)8 mm broad, lanceolate to 

elliptical, mostly acute, broadly attenuate basally, 

mostly without distinct petiole; finely to coarsely 

2-9 serrate on a side, the teeth to 2 mm long; midrib 

standing out below, the laterals inconspicuous. Inflorescence ascending, the pedicels 3 (4) 6 mm long, 

sometimes secund. Calyx glabrous to finely pubescent, 

9( 11) 14 mm long, 4(6) 8 mm broad, major veins 

often standing out, the lobes triangular, 5 (7) 10 mm 

long, occasionally serrate. Corolla pink or violet to 

magenta, sometimes “red” or purple, 29 (38) 45 mm 

long, 8 ( 11 ) 13 mm broad, the upper lip 8 ( 12) 17 mm 

long, little or not galeate, the forward margins 

rounded and reflexed, the lobes of the lower lip to 

5 mm long, 4-5 mm broad. Stamens dimorphic, the 

upper pair shorter, always included, the anthers small 

but mostly fertile, the lower pair usually partly exposed, shaggy, the lobes short mucronate below, the 

filaments thickened. Capsules ovoid, to 15 mm long, 

about 8-9 mm broad. (Figure 24). Altitude: 1800- 

3821 m. Flowering: throughout the year, peak April 

to October. 

DISTRIBUTION AND NUMBER OF COLLECTIONS EXAMINED (Figure 27) .-COLOMBIA: Boyaca (2) ; 

Cauca (18) ; Cundinamarca (18) ; Narinio (17) ; 

Norte de Santander (2) ; Santander (3). ECUADOR: Azuay (2) ; Bolivar (2) ; Cafiar (5) ; Carchi 

(6); Chimborazo (7); Imbabura (2) ; Loja (see 

type for L. loxensis) ; Pichincha (17) ; Tungurahua 

(3). PERU: Very doubtful, “Pavon, 1827, Peru,” G. 

This species is recognized in South America by the 

large size of the flowers, and by the usually broad, 

glabrate leaves and calyx. Plants from the northern 

part of the distribution, especially in Colombia, often 

have more conspicuously serrate leaves, and were 

described in H.B.K. as L. serratifolia. There appears 

to be no consistent difference to mark L. serratifolia 

or L. loxensis Bentham as separate species. In a 

general way, L. virgata probably is closely related 

to L. sylvatica, but the flowers of the former are 

about twice as long, and the leaves seem to have 

their own patterns of variation. It is doubtful that 

a taxonomic problem will arise with these two species, 

since their ranges of distribution probably do not, or 

only barely, overlap. 

The similarity between L. virgata and L. dasyantha 

is difficult to evaluate. Were the labels removed from 

some specimens, it would at first be difficult to 

decide whether their origins were Mexico or South 

America. The similarities are manifest in the appearance of the flowers and the leaves. The plants 

of the Mexican species, however, are shorter, frequently tufted or branched from below; the flowers 

are shorter, the palate larger, the lower pair of 

anthers smaller; the leaves mostly are narrower. 

Whether the similarity is one of close genetic similarity or morphological convergence is not clear. 

111. Section Hemispadon Bentham in DC. Prod. 10: 

540. 1846. Species 15-26. 

Stamens strongly unequal, the anthers dimorphic, the 

lower pair with thickened filaments, and the 2 large 

anthers connivent by means of shaggy pubescence, 

barely exposed at mouth of corolla, not spreading, 

short mucronate below; the upper pair of stamens 

shorter, always included, with filiform filaments and 

rudimentary, usually glabrate, sterile anthers, or the 

anthers very small, 1- or 2-celled, rarely producing a 

little pollen. Corolla red or red-orange, rarely pink or 

white, the tube more or less cylindrical, sometimes 

slightly doubly ventricose (Figures 8, 11, 28, 34) . LECTOTYPE SPECIES-L. uiscosa H.B.K. 

DISTRIBuTIoN.-Twelve species, northern Mexico 

to central Panama (Figures 29, 30, 33). 

15. L. nelsonii Robinson &Greenman, Amer. Jour. 

Sci. 50: 174. 1895 (Contr. Gray Herb. 9: 174. 

1895). 
NUMBER 6 49 

TYPE LocALIm.-“Six miles above Dominguillo, 

Oaxaca, altitude 4,500 to 5,000 feet, 30 October: 

1894, Nelson, 1833.” 

Stems erect, to 1.5 m long, more or less viscid 

glandular pubescent throughout, Leaves ovate to 

lanceolate, drying membranous, the larger 26 (42) 7 1 

mm long, 8( 11) 15 mm broad, mostly sessile and 

truncate or cordate at base, sometimes appearing as 

though narrowed to the base, margins variously serrate to dentate or almost crenate with 8-14 teeth on 

a side, midrib prominent below, sometimes with 2 

to several more or less obscure lateral veins. Inflorescences ascending, the pedicels 2 (4) 8 mm long. 

Calyx glandular pubescent, 8( 13) 18 mm long, 4(5) 7 

mm broad, the lobes mostly narrowly triangular to 

oblanceolate, extremely variable, 4-24 mm long, 

sometimes with irregular margins. Corolla red (to 

pinkish vermilion with lower lip pinkish purple), 

46(52)63 mm long, 5(7)9 mm broad, the upper lip 

12(19)23 mm long, little or not galeate, the lobes 

slightly cuspidate, reflexed, the lobes of the lower 

lip 2-3 mm long. Stamens dimorphic, the upper pair 

rudimentary, the lower with expanded filaments and 

shaggy, mucronate anthers. Capsules. ovoid, to 17 mm 

long, 8 mm broad. (Figure 28). Altitude: 900-2205 

m. Flowering: May to December, peak July to 

August. 

TYPE SPECIMEN.-Ne1son 1833, us 566389. 

16. L. viscosa H.B.K., Nov. Gen. 2: ed. folio 272, ed. 

quarto 338. “1817,” 1818. 

TYPE LocALITY.-“Crescit in litore Mexican0 

Oceani Pacifici prope Acapulco . . . Floret Martio.” 

TYPE SPECIMEN.-ACapukO, P (in Humboldt & 

Bonpland Herbarium and in General Herbarium). 

SYNONYMY .- 

L. cordata Schlechtendal & Chamisso, Linnaea 5: 103. 1830. 

Type locality: “Inter Tenestepeque et Quantotalapa. 

Sept.” 

L. viejensis Oersted in Bentham & Oersted, Videnskabelige 

Meddelelser fra den Naturhistoriske Forening i Kjobenhavn 1853: 28. “1854,” probably 1853. Type locality: 

“November paa Vulkanen Viejo i Nicaragua (3000’) .” 

Type specimen: Oersted 9401, C. 

L. coccinea A. Gray, Proc. Amer. Acad. 21: 404. 1886. 

Type locality: “Rocky hills near the city of Chihuahua, 

Pringle, 656.” Type specimens: Pringle 656, BR, F, G, 

GH, K, PH, RSA, US. 

Galuesia sessilis M. E. Jones, “Extracts” Contr. Western 

Bot. 18: 51. 1933. Type locality: “NO. 27410 La BarType specimen: Schiede ti? Deppe 161, W. 

FIGURE 28.-Section Hemispadon, diagrams of flowers and 

leaves. a, L. viscosa, Ernst 2442, US; a‘, Pringle 11654, 

US; b, L. nelsonii, Pringle 9334; b‘, Pringle 6331, US, elongated calyx; b“, Pringle 6777, US, capsule with calyx; c, L. 

smithii, Pringle 6705, US; d, L. rhinanthifolia, Pringle 

11064, MICH; d’, Stanford, Lauber, &’ Taylor 2392, MO; 

e, L. jalircana, McVaugh 24250, MICH. 

ranca, Guadalajara, No. 21. 1920.” Type specimens: 

Jones 27410, 21 Nov. 1930, GH, NY, date mixed. See 

Pennell in Morton, Contr. US. National Herb. 29: 91. 

1945. 

DISTRIBUTION AND NUMBER OF COLLECTIONS EXAMINED (Figure 29) .-MEXICO : State of Mexico 

(Coatepec near Villa Guerrero, Cilly, Alexander, &’ 

Hernandez 59, MO, MSC) ; Morelos (2) ; Oaxaca 

(5); Puebla (15). 

This species is very distinctive but variable and 

difficult to characterize. The apparently membranous, 

narrow, and abundantly glandular leaves frequently 

with widely spaced teeth, the short lobes of the lower 

lip of the corolla, the highly variable calyx lobes, and 

the long attenuated capsules help to identify this 

species. It probably is closest to L. viscosa. 

Plants to 3 m tall. Leaves mostly glandular pubescent, coriaceous, sessile, the larger 27 (66) 1 12 mm 

long, 10 (26) 53 mm broad, broadly or narrowly ovate 

to elliptical or lanceolate, acute or rounded apically, 

truncate or cordate basally ; serrate to doubly serrate 

to nearly dentate, the teeth often mucronulate, 14-27 

on a side, tertiary veining conspicuously reticulate and 

usually standing out below. Inflorescences ascending; 

pedicels 2(3)6 mm long. Calyx mostly glandular 

pubescent, 5(7)8 mm long, 4(5)7 mm broad, the 

lobes triangular, about as long as the cup, 2 (3) 4 mm 
50 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 29.-Distribution of section Hemispadon in Mexico and Central America. 

long. Corolla red, sometimes tinted orange, salmon, 

pink, or purple, 30(46) 60 mm long, 4(6) 12 mm 

broad, upper lip 12(18)27 mm long, mostly not 

galeate, the forward margins rounded cuspidate, 

slightly reflexed, the lobes of the lower lip 1-3 mm 

long, 1-2 mm broad. Capsules ovoid, 8-12 mm 

long, 6-7 mm wide. (Figure 28). Altitude: Mexico, 

450-2800 m; Guatemala, 600-2400 m. Flowering: 

Throughout year, more common September to 

December. 

DISTRIBUTION AND NUMBER OF COLLECTIONS EXAMINED (Figure 29) .-MEXICO : Aguascalientes 

(Hartweg 189, G, GH, W) ; Chiapas (24) ; Chihuahua (9) ; Durango (5) ; Guererro ( 11 ) ; Jalisco (24) ; 

State of Mexico (1 1) ; Michoacan (11) ; Morelos (4) ; 

Nayarit (10) ; Nuevo Le6n (near Dulces Nombres, 

2000 m, 8 Aug. 1948, Meyer &? Rogers 2879, BR, G, 

MO, PH) ; Oaxaca (24) ; Puebla (Pahuatlan, 15 

Sept. 1945, Miranda 3692, MEXU) ; San Luis Potosi 

(2) ; Sinaloa (Mesa Malqueson, Cerro Colorado, 

2500 ft, 8 Dec. 1939, Gentry 5268, DS, GH, MO, PH, 

UC) ; Sonora (10) ; Tamaulipas (4) ; Veracruz ( 13). 

GUATEMALA : Chimaltenango ( Alameda, Johnston 

315, F) ; Chiquimula (Volcan Ipala near Amatillo, 

900-1510 m, 25 Oct. 1939, Steyermark 30571, F); 

Escuintla (2) ; Guatemala (5) ; Huehuetenango (5) ; 

Jalapa (3) ; Jutiapa (3) ; QuichC (2) ; SacatepCquez 

(3) ; San Marcos (between San Marcos and Castalia, 

1600-2200 m, 16 Dec. 1963, Williams, Molina @ 

Williams 26143, F) ; Santa Rosa (near Cuilapa, 900- 

950 m, 24 Nov. 1940, Standley 78232, F) ; SololB (3) ; 

Zacapa (2). BELICE: EI Cay0 (3). EL SALVA. 

DOR : Chalatenango (Cerro Roblar, Nov. 1928, Cab 

deron 2475, F) ; Chalchuapa (2) ; Santa Ana (near 

Santa Ana, 655-900 m, Jan. 1922, Standley 20391, 

GH, US). HONDURAS: Choluteca (near San 

Marcos, 1000 m, 16 Nov. 1946, Williams @ Molina 

10886, F, GH) ; Copin (between Llave de la Puerta 

and El Salto, 900 m, 10 Jan. 1907, Pittier 1852, US) ; 

El Paraiso (2) ; Morazin (9) ; Santa BBrbara (San 
NUMBER 6 51 

Pedro Sula, 600 m, ?Thieme, US). NICARAGUA: 

Chinandega (3) ; Jinotega (near Jinotega, 1030-1300 

m, June-July 1947, Standley 9873, F) ; plus 4 uncertain. COSTA RICA: Alajuela (near San Ram6n, 

1025 m, 8 Nov. 1926, Brenes 2134, F) ; Cartago (3) ; 

Heredia (Alto de Santa Birbara, 1400 m, 21 Dec. 

1889, Pittier 1666, BR) ; Puntarenas (2) ; San JosC 

(Cerro de Pralti Escasli, 1300 m, Aug., 1935, solis 

318, F, MO) . PANAMA: Chiriqui (between Cerro 

Vaca and Hato del Loro, 850-1100 m, 27 Dec. 

1911, Pittier 5398, US; Valle de Anthn, 800 m, 8 

Sept. 1941, Allen 2235, MO, US). 

CHRoMOSOMES.-n= 14, counted by Ernst from 

D. E. Breedlove 18676, Jalisco, 16 miles northwest of 

Ameca along the road to Mascota, 4300 ft, 1 Nov. 

1970, CAS, US. 

With more than 250 separate collections, this is 

the most frequently collected and the most variable 

species, as would be expected for a taxon distributed 

from northern Mexico to central Panama. The broad, 

coarse, mostly conspicuously reticulately veined leaves 

with cordate base usually separate this from all other 

species. Sometimes the leaves are narrow and appear 

tapered to the base. This species probably is closely 

related to the uncommon L. nelsonii and to L. rhinanthifolia, which is very common in Mexico and at 

times appears to have overlapping characters sug 

gesting the possibility of hybridization. The thick 

leaves, glandular calyx, and narrow, more pointed 

corollas usually distinguish L. viscosa from L. rhinanthifolia which generally has thinner leaves, mostly 

nonglandular calyx with incised lobes, and broader, 

often more blunt corollas ; both species sometimes have 

scabrous leaves. The corollas of L. viscosa in Chiapas 

are unusually broad ; however, L. rhinanthifolia, with 

which these plants might be confused, does not cross 

the Isthmus of Tehuantepec. The habitats most frequently mentioned are grassland openings in arid oak 

and/or pine forest, sometimes on limestone, near cultivation, or in Acacia-Yucca-cactus woodland. 

17. L. rhinanthifolia H.B.K., Nov. Gen. 2: ed folio 

271, ed. quatro 337. pl. 169. "1817," 1818. 

TYPE LocALITY.-"Crescit locis temperatis Novae 

Hispaniae prope pagos Magdalena, Actopan et Sancta 

Rosa de la Sierra, alt. 1040-1300 hex . . . Floret 

Julio-Septembri . . . Descriptio speciminis propter 

pagum Magdalena lecta . . . ." 

TYPE SPECIMEN.-^. Rosa, n. 4274, P (in Humboldt & Bonpland Herbarium). 

Stems erect to 1 (3) m tall. Leaves membranous 

and pubescent to coriaceus and scabrous, the larger 

16(34) 52 mm long, 11 (17) 28 mm broad, sessile, 

mostly ovate or elliptical, rounded or obtuse apically, 

truncated to cordate basally, serrate to doubly serrate 

or nearly dentate, the teeth 8-21 on a side, sometimes 

mucronulate; midrib standhg out below with a few 

laterals. Inflorescence ascending; pedicels l(3) 5 mm 

long. Calyx pubescent, 7 (9) 12 mm long, 5 (8) 11 mm 

broad, the lobes 3 (4) 5 mm long, broadly triangular 

and usually deeply serrate, sometimes entire. Corolla 

variable red (tinted pink, orange, yellow, purple), 

47(57)68 mm long, 8(10)13 mm broad, often subventricose toward calyx, upper lip 15 (22) 26 mm long, 

little or not galeate, forward margins mostly rounded. 

reflexed; the lobes of lower lip to 4 mm long. Stamens 

dimorphic, the upper pair of anthers vestigial, lower 

pair shaggy, the filaments expanded, and the lobes of 

the anthers short mucronate. Capsules ovoid, to 13 

mm long, 8 mm broad. (Figure 28). Altitude: 793- 

3110 m. Flowering: May to October, peak June to 

DISTRIBUTION AND NUMBER OF COLLECTIONS EXAMINED (Figure 30) .-MEXICO : Aguascalientes 

(Hartweg, BR) ; Coahuila (La Casita, Kenoyer €3 

Crum 3118, A, MICH) ; Distrito Federal (13) ; Durango (13) ; Guanajuato (3) ; Hidalgo (5) ; Jalisco 

(near Hacienda Chinampas, McVaugh 17027, 

MICH) ; State of Mexico (27) ; Michoach (near 

Coalcomin, Hinton 12356, DS, GH, MEXU, MICH, 

US) ; Nuevo Le6n ( 10) ; Oaxaca (19) ; Puebla (8) ; 

San Luis Potosi (8) ; Sinaloa (2) ; Tamaulipas (4) ; 

Veracruz (2) ; Zacatecas (2). 

After L. viscosa and L. multifida, the third most 

common species is L. rhinanthifolia. The broad, ovate, 

usually membranous leaves with relatively close, deep, 

and plentiful teeth, with conspicuous venation below; 

the broad, serrate sepals; and the relatively broad 

corollas identify this species. The thick flower tubes 

recall those of L. tenuifolia but these species otherwise are only remotely similar. This species probably 

is closest to L. viscosa but is distinguished by its 

usually smaller and more membranous leaves, serrate 

calyx lobes, and broader corollas; however, the two 

sometimes are confusingly similar. The relationship 

to L. smithii probably also is very close. Limestone 

substrate is mentioned for Nuevo Le6n, San Luis 

Potosi, and Tamaulipas. 

i\ugust. 
52 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 30.Distribution of section Hemispadon in Mexico and Central America. 

18. L. smithii Robinson & Greenman, her. Jour. 

Sci. 50: 172. 1895 (Contr. Gray Herb. 9: 172. 

1895). 

TYPE LOCALITY.-“Mountains of Jayacatlan, Oaxaca, altitude 5,000 feet, 13 August, 1894, L. C. 

Smith, 155.” 

TYPE SPECIMEN.Cmith 155, GH. 

Stems erect, probably 1 m tall. Leaves pubescent, 

membranous, ovate, 50 (65) 72 mm long, 29 (35) 38 

mm broad, sessile and conspicuously cordate basally; 

margins dentate to serrate, sometimes doubly so, the 

projections 12-27 on a side, 2-4(6) mm long, often 

mucronulate ; midrib conspicuous above and standing out below with 4-6 laterals. Inflorescences ascending; pedicels 3-5 mm long. Calyx pubescent, 8( 10) 11 

mm long, 5-6 mm broad, the lobes 4-6 mm long, 

narrowly triangular. Corolla presumed red (or orange), 44(49) 54 mm long, 8(9) 10 mm broad, the 

upper lip 17 (19) 22 mm long, little or not galeate, 

the forward margins rounded or slightly cuspidate, 

slightly reflexed; the lobes of the lower lip about 

2 mm long. Stamens dimorphic, the upper pair of 

anthers vestigial, the lower pair shaggy, the lobes 

mucronate at base, the filaments expanded (Figure 

28). 

DISTRIBUTION AND COLLECTIONS EXAMINED (Figure 

30) .-MEXICO: Oaxaca (Smith 155, see type; 

Tomellin Canyon, 5000 ft, 17 July 1897, Pringle 

6705, BR, F, G, GH, IPN, MBG, MEXU, MSC, PH, 

UC, US, W; Cuicotlan, 580 m, 3 Aug. 1872, Urbina 

sn, MEXU). 

This species is distinguished by its membranous, 

large, cordate, sessile, ovate leaves with their conspicuous, widely spaced venation, and the restricted 

distribution in Oaxaca. This species seems related to 

L. viscosa, but is especially close to L. rhinanthifolia. 

The leaf size and shape resemble the former; the 

texture and venation of the leaves and the broadness 

of the flowers resembles the latter. Little variation is 

shown by the few collections available; however, it 

would seem likely that if this species were variable 

it would blend into L. rhinanthifolia. 
NUMBER 6 53 

19. L. jaZiscana W. R. Ernst & M. F. Baad, Ma- jada; elevation 1800-2250 m, 19-21 November 1952. 

HoLoTYPE.-Rogers McVaugh 14250 (& J. Sooby, 

Jr.), MICH (Figure 31) and Isotype. 

Stems erect to 1.5 m tall. Leaves possibly scabrous 

above, soft pubescent below, narrowly lanceolate, 50- 

droiio 20: 344. “1970,” 17 March 1971. 

TYPE LocALITY.-JalisCo: Sierra de Caule, southwest of Talpa de Allende; southwest of Piedra RaFIGURES 3 1 .-Section Hemispadon. L. jaliscana, holotype, McVaugh 14250, MICH. 
54 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

80 mm long, 6-10 mm broad, attenuate apically and 

basally, revolute, serrate with 12-22 coarse teeth on 

a side. Inflorescences ascending; the pedicels 5-8 mm 

long. Calyx glandular pubescent, 14-16 mm long, 

5 mm broad, the lobes narrowly subulate, 10-12 mm 

long. Corolla red, cylindrical, 56-63 mm long, 9-10 

mm broad, the upper lip 16-25 mm long, little or 

not galeate, the lobes broadly attenuate, 5-7 mm long, 

reflexed; the lobes of the lower lip narrow, 5-8 mm 

long. Upper pair of stamens half as long as corolla, 

the anthers vestigial and glabrous; lower pair of stamens nearly as long as corolla, the filaments dilated 

distally, the anthers shaggy pubescent, the lobes 

acuminate at base. Style with spreading hairs. Capsules ovoid, 12-13 mm long, 7-8 mm broad. (Figure 

28). 

DISTRIBUTION AND COLLECTIONS EXAMINED (Figure 

33) .-MEXICO: Jalisco (see type; 11-12 miles 

south of Talpa de Allende, headwaters of west branch 

of No de Talpa, elevation 1200-1700 m, 23-25 November 1960, McL‘augh 21325 (& C. Feddema, R. 

Pippen), MICH) . This species, known from only two collections, is 

distinguished by its large leaves, subulate glandular 

calyx lobes, and large corolla. The lower lip of the 

corolla arches downward with three elongated lobes 

resembling section Lamourouxia, and the subulate 

calyx lobes resemble the similar condition in L. longipora and L. multifida. The dimorphic stamens with 

only one pair of fertile anthers, however, establishes 

this species in section Hemispadon in a more or less 

central position between extremes. The pubescent 

leaves and glandular calyx allies L. jaliscana to the 

L. viscosa complex; however, the narrowly elongated 

leaves with their tapered or petiolelike bases resembles 

the similar condition among the glabrous-glabrate 

species such as L. lanceolata and L. tenuifolia (Fig 

ure 32). 

The position of L. jaliscana is enigmatic. The broad 

corolla, its dangling lower lip with long, spreading 

lobes and the more or less open mouth clearly suggest 

an alliance with section Lamourouxia. The heavy, 

almost scabrous pubescence of the leaves, the glandular condition of the calyx, and the strongly dimorphic 

stamens with only two fertile anthers, on the other 

hand, fixes this species in section Hemispadon near 

the L. viscosa-L. rhinanthifolia complex. Although the 

leaves are pubescent and the calyx glandular, the 

lanceolate, petiolate condition of the leaves recalls the 

similar characters among the glabrous-glabrate species 

of Hemispadon. The position of this species in Hemispadon parallels that of L. dispar in section Lamourouxia, inasmuch as either the corollas or the stamens 

in these two species appear to have become associated 

with the wrong plants. 

20. L. gracilis Robinson & Greenman, Amer. Jour. 

Sci. 50: 174. 1895 (Contr. Gray Herb. 9: 174. 

1895). 

TYPE LocALITY.--“Summit of Sierra Madre near 

Chilpancingo, Guerrero, altitude 9,000 to 12,000 feet, 

24 December, 1894, Nelson, 2234.” 

TYPE SPECIMENS.-NelSOn 2234, GH, us. 

Stems erect 25-60 cm tall. Leaves membranous, 

elliptical to obovate, 21 (24) 28 mm long, 5 (7) 10 mm 

broad, basally attenuate, glabrous or with a few appressed hairs ; margins sometimes revolute, serrate or 

almost crenate, with 5-9 teeth on a side, the projections to 1 mm long; midrib standing out below, 

usually with 3-4 laterals. Inflorescences ascending 

with few flowers; pedicels 6-10 mm long. Calyx 

5 ( 7) 9 mm long, 3 (4) 5 mm broad, glabrate, the lobes 

broad, oblanceolate, spreading, about 4 mm long, 2 

mm broad, sometimes with 2-3 shallow indentations. 

Corolla “red with yellowish throat,” 45 (46) 49 mm 

long, 8(9) 10 mm broad, upper lip 14( 15) 17 mm 

long, rounded, slightly galeate, the short lobes reflexed; lobes of the lower lip 2-3 mm long. Stamens 

dimorphic, the upper pair rudimentary, the lower pair 

with thickened filaments and shaggy anthers, their 

lobes mucronulate below. Capsule ovoid, 10 mm long, 

6 mm broad. (Figure 34.) Flowering: September to 

December. 

COLLECTIONS EXAMINED (Figure 33) .-MEXICO: 

Guerrero (see type) ; Morelos (near Cuernavaca, 

Pringle 6510, BR, CAS, ENCB, F, G, GH, K, MEXU, 

MO, MSC, PH, UC, US, W; near Cuernavaca, 

Pringle sn, 19 Oct. 1900, C) , 

This species is distinguished by its small, but elongated, petiolate, glabrate, membranous leaves, broad, 

reflexed, calyx lobes, and broad corolla. It appears to 

be closest to L. microphylla. 

21. L. microfihylla Marteqs & Galeotti, Bull. Acad. 

Sci. Bruxelles 12(2) :31. 1845. 

TYPE LocALITY.-“Dans les bois humides de Yolotepeque et Sola (cordillere au sud d’oaxaca), de 
NUMBER 6 55 

FIGURE JZ.--Section Hemispadon. L. tenuijolia (a syntype specimen of L. conzatti, Gonzales 

&? Conzatti 760, GH). 

6,500 a 7,500 pieds . . . Septembre . . . H. Gal. No 

1005.” 

TYPE SPECIMENS.-Gu~eOtti 1005, mixed data representing tlvo or three collections, BR (two specimens) , G, K (tlvo specimens) W. 

(v. in herb. Hook.).” Type specimens: Galeotti 2005, 

BR (two specimens), G, K (two specimens), W. 

Stems erect to 1 m tall. Leaves glabrous, probably 

coriaceous, 610 mm long, 2-3 mm broad, elliptical 

to narrowly obovate, attenuate basally and appearing 

to have a petiole to 1 mm long; margins revolute, 

serrate or sinuate, with 3-4 lobes on a side, midrib 

standing out below, sometimes with 2-3 faint laterals. 

SYNONYMY.- 

L. parvifolia Bentham in DC. prod. 10: 542. 1846. T~~~ 

locality: “In sylvis prov. Oaxaca (Galeotti. n. 1005) , . . 
56 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 33.-Distribution of section Hemispadon in Mexico and Guatemala. Localities in 

Guatemala of specimens appearing to blend characters of L. integerrima and L. lanceolata 

shown by open triangles. 

Inflorescences ascending, frequently branched, pedicels 

about 4 mm long. Calyx glabrous, 5-6 mm long, 

3-4 mm broad, the lobes triangular, to 1 mm long. 

Corolla red, 37-41 mm long, 6 mm broad, the upper 

lip 16-18 mm long, little or not galeate, the forward 

margins reflexed, the lobes of the lower lip 2-3 mm 

long. Stamens dimorphic, the upper pair of anthers 

rudimentary, the lower pair shaggy, the lobes acuminate below, the filaments expanded. Capsules subspherical, 6 mm long, 5 mm broad. (Figure 34). 

DISTRIBUTION AND COLLECTIONS EXAMINED (Figure 

33) .-MEXICO: Oaxaca (Galeotti 1005, see type; 

Cuesta de S. Pedro el Alto, 10/42, Liebmann 9394, 

A, C (3), F, NY, US; granitic hills, Sierra de Clavellinas, 6000 ft, 19 October 1894, Pringle 6000, BR, 

BM, ENCB, G, GH, K, MEXU, MO, MSC, NY, P, 

PH, RSA, UC, US, W, WIS. Oak-pine woods, 6900 

ft, Sierra Madre del Sur south of Miahuatlhn, 14 

Nov. 1966, Ripley tY Barneby 14640, CAS) . 

This species is distinguished by its very small, 

glabrous leaves, glabrous calyx with very short lobes, 

a 

FIGURE 34.-Section Hemispadon, diagrams of flowers and 

leaves. a, L. tenuifolia, Halberg 939, MICH; b, L. integerrima, Steyermark 43293, F; G, L. lanceolata, Tucker 658, 

IJS; d, L. colimae, McVaugh 18077, MICH; e, L. gracilis, 

Pringle 6510, US; f, L. microphylla, Pringle 6000, US & 

MEXU; g, L. gutierrecii, Allen 1496, US. 
NUMBER 6 57 

and narrow corollas. There are too few collections for 

a sense of variation. This species seems most closely 

allied to L. gracilis. 

22. L. tenuifolia Martens & Galeotti, Bull. Acad. 

Sci. Bruxelles 12 (2) : 32. 1845. 

TYPE LOCALITY.-“DanS les bois de r6gions tempCrCes froides de la cordielh orientale d‘Oaxaca (San 

Andreas, Roayaga, etc.), ai 6,000 pieds . . . Juin . . . H. Gal. No 922.” 

SYNONYMY.- 

TYPE SPECIMENS.-GUleOtti 992, BR, G, w. 

L. conzattii Greenman, Proc. Amer. Acad. 35: 312. 1900 

(Contr. Gray Herb. 18: 312. 1900). Type locality: “Collected by V. Gonzales and C. Conzatti on the Cerro de 

Pbpalo, camino de Cheve, Cuicatlan, Oaxaca, alt. 2,500 

m., 16-22 June 1898, no. 760, and C. Conzatti on the 

Cerro de Yalina, District of Villa Alta, Oaxaca, altitude 

1,500 m., June 1899, no. 965.” Syntype specimens: Gonzales d Conratti 760, GH, US; Conratti 965, GH. 

Stems erect, probably 1 m or more tall. Leaves 

glabrous or glabrate, very narrowly lanceolate or 

subulate, attenuate at either end, the larger 21 (28) 32 

mm long, 3 mm broad, margins revolute, sinuate to 

serrate, the teeth 6-10 on a side, usually less than 1 

mm long, midrib standing out below or with 3-4 faint 

laterals. Inflorescences ascending; pedicels 4 (5) 7 mm 

long. Calyx glabrous or minutely papillate, 7 (9) 10 mm 

long, 3 (4) 6 mm broad, the major veins usually standing out, the lobes triangular, 4-5 mm long. Corolla 

red, 45(51)55 mm long, 9(10)13 mm broad, the 

upper lip 11 (16) 19 mm long, little or not galeate, 

margins reflexed, the lobes of the lower lip 2-3 mm 

long. Stamens dimorphic, the upper pair rudimentary, 

the lower pair with thickened filaments and shaggy 

anthers, mucronate below. Capsules 10-12 mm long, 

6-8 mm broad. (Figure 34). Altitude: 1372-2500 m. 

Flowering: April and June to August. 

DISTRIBUTION AND NUMBER OF COLLECTIONS EXAMINED (Figure 33) .-MEXICO : Oaxaca (near 

Ixtlan, July 1906, Conratti sn, A, ENCB, G, UC, US, 

W; Gonzales B Conratti 760, Conzatti 965, see syntypes for L. conzattii; Sierra de Ixtlbn, 2200 m, July 

1906, Conzatti 1379, ENCB, GH, MEXU; Ixtlbn, 

July 1906, Contatti (or ?Pringle) 10371, F, G, K, 

MO, MSC, PH; Galeotti 992, see type; vicinity of 

Cerro Zempoaltepec, 2400 m, 13 Aug. 1950, Hallberg 

939, MICH; Lachopa, 7500-8000 ft, 6/1842, Liebmann 9393, C, US, W; Villa Alta, April 1949, Paray 

122, MEXU). 

The very narrow, glabrous leaves, glabrous sepals, 

and relatively broad corollas distinguish this species, 

which does not seem to be very closely allied to any 

of the others. The name L. tenuifolia has been misapplied to many collections of L. dasyantha, a very 

different species in section Adelphidion, which, coincidentally, usually has narrow leaves. The type collection for L. tenuifolia var. micrantha Greenman is 

L. dasyantha. 

23. L. integerrima J. D. Smith, Bot. Gaz. 13: 189. 

1888. 

TYPE LocALITY.-Guatemala, “Pansamala, Alt. 

4,000 feet, January, 1887. (Ex P1. cit. 11 12.) .” 

TYPE sPEcIMENs.-~on Tuerckheim 11 12, G, GH, 

PH, US (3 sheets) (Figure 35). 

Stems to 2.5 m long. Leaves membranous, glabrous 

or glabrate, lanceolate to narrowly elliptical, attenuate 

to base, 20 (30) 42 mm long, 3 (5) 7 mm broad, margins entire or.sinuate or slightly serrate, midrib and 

a few lateral veins standing out below. Inflorescences 

ascending; pedicels 4 (6) 8 mm long. Calyx glabrous or 

glabrate 7 (8) 10 mm long, 3 (4) 5 mm broad, the lobes 

2(4) 6 mm long. Corolla red, 43(47)51 mm long, 

8(9) 11 mm broad, the upper lip 14( 17) 19 mm long, 

slightly or not galeate, the lobes reflexed; the lobes 

of the lower lip to 4 mm long. Stamens dimorphic, 

the upper pair rudimentary, the lower pair with 

expanded filaments and shaggy anthers, the lobes 

acuminate basally. Capsules ovoid, 10-14 mm long, 

7-8 mm wide. (Figure 34). Altitude: 1220-2865 m. 

Flowering: December to February and June to 

October. 

DISTRIBUTION AND MORE OR LESS TYPICAL COLLECTIONS EXAMINED (Figure 33) .-GUATEMALA: Alta 

Verapaz (see type) ; El Progreso (between Finca Piamonte and summit of Volch Santa Luisa, 2400-3333 

m, Steyermark 435557, F) ; Huehuetenango (4 mi E 

of San Mateo Ixtath, 8500 ft, Breedlove 8768, DS, 

F, MICH, US; Cerro CananL near Nucapoxlac, 

2500-2800 m, Steyermark 49083, F) ; QuichC (Nebaj, 

7600 ft, Skutch 1897, A, GH, F, PH, US) ; Zacapa 

(between Lomas El Picacho and Cerro de Monos, 

2000-2600 m, Steyermark 42835; Volciin Gemelos, 

2100-3200 m, Steyermark 43923, F, PH). 

COLLECTIONS PRESUMED CLOSE TO L. integerrima, 

RUT INCOMPLETE OR WITH VERY NARROW AND/OR 
58 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURES 35.--Section Hemispadon. L. integerrima, presumed holotype, Vcn Tuerckheim 1 112, US. 

SERRATE LEAVES, AND OFTEN WITH LONGER SEPALS.- 

GUATEMALA: Alta Verapaz (trail to SenahG, G. P. 

Go11 116, US) ; El Progreso (between Finca Piamonte 

and summit of Volchn Santa Luisa, 2400-3333 m, 

Steyermark 43475, F. US) ; Huehuetenango (Cerro 

Huitz between Barillas and Mimanhuitz, 1600-2600 

m, Steyermark 48534, F) ; Quezaltenango (Volchn 

Zunil, 2500-3800 m, Steyermark 34588, F; same locality, Steyermark 34589, F, PH) ; Solola (between 

San Pedro and Volan Santa Clara, 1900-2100 m, 

Steyermark 4711 I, F) ; Suchitepequez (Volah Atitlan, 8000 ft, Skutch 1519, A, F, PH, US). 

The broad, entire, lanceolate leaves, the ascending 

inflorescence tips, the short, broad calyx lobes, and 

the broad corolla distinguish this species. Collections 

other than the type, however, tend to have narrower 
NUMBER 6 59 

and often serrate leaves as well as longer calyx lobes. 

The habit varies and is sometimes noted as shrubby 

or “drooping.” The type collection with its unusual 

leaves appears to be an extreme condition. The first 

collections cited above are more or less similar to the 

type, but the second group is of uncertain relationship, perhaps representing a new taxon or intermediates with L. lanceolata. This taxonomic problem 

is restricted to Guatemala. 

24. L. lanceolata Bentham in DC. Prod. 10: 542. 

1846. 

TYPE LocALITY.-“In Mexici prov. Chiapas ad 

Amatenango (Linden! n. 198) . . . (v. in herb. 

Hook.) ,” 

TYPE SPECIMENS.-Linden 198, G, K (Herb. 

Hook). 

Plants to 3 m tall, stems frequently branching 

at nearly right angles. Leaves glabrous or with a few 

hairs along major veins, mostly narrowly lanceolate, 

rarely ovate, usually attenuate at either end, occasionally somewhat falcate, the larger leaves 19 (42) 57 

mm long, 4( 7) 14 broad, 4-1 1 serrate on a side, midrib and a few laterals standing out below. Inflorescences depressed; pedicels 1 (7) 14 mm long; flowers 

inverted. Calyx glabrous or glabrate, 10( 14) 18 mm 

long, 4(5)6 mm broad, major veins standing out, 

the lobes oblanceolate or narrowly triangular, frequently spreading, 6(9) 15 mm long, rarely serrate. 

Corolla red or red-orange, 32 (41) 47 mm long, 5 (6) 8 

mm broad, the upper lip 8( 16) 20 mm long, little or 

not galeate, the lobes reflexed; lobes of the lower lip 

2-3 mm long. Stamens dimorphic, the upper pair 

qhort and rudimentary, the lower pair with expanded 

filaments and shaggy anthers, the lobes short mucronate. Capsules ovoid to 12 mm long, to 8 mm 

broad. (Figure 34). Altitude: 1000-2800 m. Flowering: December to April, peak December to 

February. 

DISTRIBUTION AND NUMBER OF COLLECTIONS EX 

AMINED (Figure 30) .-MEXICO: Chiapas (13) ; 

TMichoacAn (doubtful-Dos Teteras, Arsene, I1 1909, 

PH) ; Oaxaca (between Suchixtepec and El Guajolote, Ernst 3 Delgadillo 2628, US). GUATEMALA: Alta Verapaz (1 1) ; El Progreso (between 

El Jute de Cobana and Finca Piamonte, Steyermark 

43368, F, PH) ; Guatemala (Montaiia of Las Nubes 

west of San JosC de Pinula, Williams @ Molina 

15272, F, GH) ; Quezaltenango (5) ; QuichC (San 

Miguel Uspatan, Heyde €3 Lux, Apr. 1892, GH, US) ; 

SacatepCquez (5) ; San Marcos (Volah Tajumulco, 

Steyermark 36561, F) ; Solo1a (2) ; Zacapa (2). EL 

SALVADOR : Chalatenango (Los Esimiles, Tucker 

1007, F, G, MICH, PH, UC, US) ; Morazan (Montes 

de Cacaguatique, Tucker 658, F, G, MICH, PH, UC, 

US) ; San Salvador (2). COSTA RICA: Cartago 

(4) ; Puntarenas (Ujarras, Oersted 9400, A, C, 

MICH, US, W) ; San JosC (1 1). 

Floral as well as vegetative characteristics of L. 

lanceolata link it to other species in Hemispadon. It 

is distinguished from all other species in Hemispadon, 

however, by its growth habit which tends to be deliquescent, the branching frequently at right angles, 

the inflorescence tips depressed, and the flower inverted. This peculiar syndrome has been discussed 

previously for L. xalapensis and its closest allies in 

section Lamourouxia. 

The plant in Oaxaca (Ernst €3 Delgadillo) was a 

low, intricately branched shrub; some other herbarium specimens look as though there were longer shoots 

arching over other vegetation. The elongated leaves, 

depressed inflorescences, and reflexed calyx lobes distinguish this species. The peculiar inflorescence characters are shared with L. dependens, L. ovata, and 

L. xalapensis of section Lamourouxia. A taxonomic 

problem exists with L. integerrima in Guatemala 

partly because of variation and partly because of 

poorly preserved specimens. 

25. L. colimae W. R. Ernst & M. F. Baad, Madroiio 20: 342. “1970,” 17 March 1971. 

TYPE LocALITY.-Colima : Mountain summits near 

pass ca. 11 miles south-southwest of Colima on Manzanillo road; elevation 500 m. 21 September 1958. 

HoLoTYPE.-Rogers McVaugh 18077 (& H. F. 

Loveland, R. W. Pippen), MICH (Figure 36). 

Stems erect to 1 m tall. Leaves glabrous or with 

very few hairs, elliptical or ovate, 18-26 mm long, 

6-10 mm broad, attenuate basally, mostly with a very 

short petiole or sessile, margins revolute, 3-6 serrate 

or dentate on a side, veining standing out below, 

midrib and 3-6 laterals recessed above. Inflorescence 

erect ; pedicels 3-4 mm long. Calyx glabrous or microscopically papillate, 6-7 mm long, 4 mm broad, the 

lobes 4-6 mm long, broadly triangular, spreading. 

Corolla scarlet, 30-35 mm long, 6-7 mm broad, the 
60 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 36.--Section Hemispadon. L. colimae, holotype. McVaugh 18077, MICH. 

upper lip 12-13 mm long, the lobes shallow, rounded, 

slightly cuspidate, little or not reflexed; the lobes of 

the lower lip about 1 mm long. Upper pair of stamens 

one-half or less the length of the corolla, their anthers 

vestigial and glabrous; lower pair of stamens as long 

as the corolla, the filaments expanded distally, the 

anthers shaggy pubescent, their lobes short acuminate 

at base. Style with a few short hairs. (Figure 34). 

DISTRIBUTION.-MEXICO : Colima (see type; and 

also see Figure 30). 

This species is known only from the type collection. 

It seems close to L. lanceolata, although the inflores- 
NUMBER 6 61 

cence is erect, somewhat similar to L. gracilis although the corollas are narrower, and recalls L. 

gutierrezii although the leaves are glabrous. The 

low altitude seems notable. 

26. L. gutierrezii Oersted in Bentham 81 Oersted, 

Videnskabelige Meddelelser fra den Naturhistoriske 

Forening i Kjobenhavn for Aaret 1853 : 29. “1854,” 

but the first part, including p. 29, probably distributed in 1853. 

TYPE LocALITY.-“Denne smukke Art voxer i den 

yppige Skov, som beklaeder Bjergene mellem Cartago 

og Candelaria i en Hoide af 6-7000 Fod, hvor jeg 

traf den med Blomst i Februar.” 

TYPE sPEcIMENs.-Costa Ria, “Inter Cartago et 

Candelaria, 8000’, 2/47,” Oersted 9398, C; ? “No. 

10, In monte Candelaria,” K (Herb. Benth.). 

SYNONYMY.- 

L. scabra Bentham ex Seemann in Bot. Voy. H.M.S. Herald, 

p. 177. pl. 33. 1854. Type locality: “Volcano of Chiriqui, Veraguas.” Type specimens : Panama, Seemann, Bot. 

Herald, tab. 33, “to be figured,” BM; Seemann, Boquete, 

Mar. 1848, K (Herb. Benth.). 

Stems erect to 2 m long. Leaves scabrous pubescent, 

elliptical to ovate, 8( 15) 25 mm long, 4(6) 10 mm 

broad, usually tapered below, petiole to 2 mm long, 

margins serrate to crenate with 4 to 10 teeth on a 

side; veining standing out below with 1 or 3 principal 

veins, usually recessed above. Inflorescences ascending; pedicels 2-3(4) mm long. Calyx pubescent, 

6 (8) 10 mm long, 4-5 mm broad, the lobes 2 (3) 4 mm 

long, broadly triangular, occasionally incised. Corolla 

red (orange or ?yellow), 32(37)44 mm long, 4(6)8 

mm broad, the upper lip 12 ( 15) 18 mm long, slightly 

or not at all galeate, the lobes reflexed; the lobes of 

the lower lip 1-2 mm long. Stamens dimorphic, the 

upper pair rudimentary, the lower pair with thickened 

filaments and shaggy anthers with mucronate lobes. 

Capsules ovoid to spheroidal, 9-13 rnm long, 7-8 mm 

broad. (Figure 34). Altitude : 945-1 750 (2000) m. 

Flowering: October to April and June to July, peak 

December to January and July. 

DISTRIBUTION AND NUMBER OF COLLECTIONS EXAMINED (Figure 30) .-COSTA RICA: Alajuela (5) ; 

Cartago (3) ; Heredia (2) ; Puntarenas (2) ; San JosC 

(9). PANAMA: Chiriqui (7). 

This species is distinguished by its far southern 

distribution in Central America, being the only one 

restricted to Costa Rica and Panama. In a general 

way, it seems close to L. colimae, L. gracilis, and L. 

lanceolata but is very pubescent and scabrous, thus 

resembling the L. uiscosa complex; the species differs, 

however, in being without glands and the leaves are 

tapered to the base. 

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62 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

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Index to Species 

A, section Adelphidion ; H, section Hemispadon ; L, section Lamourouxia 

6. betonicaefolia 

11. BRACHYANTHA (A) 

13. breviflora 

16. coccinea 

25. COLIMAE (H) 

22. conzattii 

16. cordata 

10. exserta 

20. GRACILIS (H) 

26. GUTIERREZII (H) 

23. INTEGERRIMA (H) 

19. JALISCANA (H) 

24. LANCEOLATA (H) 

12. linearis 

2. LONGIFLORA (L) 

14. loxensis 

6. MACRANTHA (L) 

21. MICROPHYLLA (H) 

9. DEPENDENS (L) 

5. DISPAR (L) 

1. gxandiflora 

2. hyssopifolia 

1. laciniata 

= MACRANTHA 

= SYLVATICA 

=VISCOSA 

= TENUIFOLIA 

= VISCOSA 

= XALAPENSIS 

= MULTIFIDA 

= LONGIFLORA 

= MULTIFIDA 

=DASYANTHA 

= VIRGATA 

9. montana =DEPENDENS 

1. MULTIFIDA (L) 

8. OVATA (L) 

3. PARAYANA (L) 

15. NELSON11 (H) 

10. parviflora Hemsley = XALAPENSIS 

21. parvifolia Bentham = MICROPHYLLA 

17. RHINANTHIFOLIA (H) 

26. scabra = GUTIERREZII 

14. serratifolia = VIRGATA 

18. SMITH11 (H) 

10. spathaceae =XALAPENSIS 

13. subincisa = SYLVATICA 

13. SYLVATICA (A) 

22. TENUIFOLIA (H) 

12. tenuifolia micrantha =DASYANTHA 

1. tepicana = MULTIFIDA 

16. viejensis = VISCOSA 

14. VIRGATA (A) 

16. VISCOSA (H) 

10. XALAPENSIS (A) 

4. PRINGLEI (L) 

7. STENOGLOSSA (L) 

* U.S. GOVERNMENT PRlNTlNQ OFFICE: 1972 0--4L14--913/9 

63