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UNIVERSITY OF ILLINOIS LIBRARY AT URBANA-CHAMPAIGN

L161— O-1096

FIELDIANA: BOTANY

A Continuation of the

BOTANICAL SERIES

of

FIELD MUSEUM OF NATURAL HISTORY

VOLUME 31

The Library of the

JAN 1 8 1979

/&&**&

AV^ NATURAL ^\

I* HISTORY >\

FIELD MUSEUM OF NATURAL HISTORY

CHICAGO, U. S. A.

520.5

vi 1

TABLE OF CONTENTS

PAGE

1. Two New Species of Palms from Nicaragua. By S. F. Glassman ... 1

2. Tropical American Plants, VI. By Louis O. Williams 11

3. Agriculture, Tehuacan Valley. By C. Earle Smith, Jr 49

4. Flora, Tehuacan Valley. By C. Earle Smith, Jr 101

5. Preliminary Studies in the Palm Genus Syagrus Mart, and Its Allies.

By S. F. Glassman 145

6. Tropical American Plants, VII. By Louis O. Williams 165

7. Supplement to Orchids of Guatemala. By Donovan S. Correll .... 175

8. Preliminary Notes on Scrophulariaceae of Peru. By Gabriel Edwin . . 223

9. New Species in the Palm Genus Syagrus Mart. By S. F. Glassman . . 233

10. Tropical American Plants, VIII. By Louis O. Williams 247

11. Notes on the Flora of Costa Rica, I. By William C. Burger 273

12. A New Eurystyles from Nicaragua. By Alfonso H. Heller 279

13. New Species in the Palm Genus Syagrus Mart. By S. F. Glassman . . 285

14. A Revision of the Family Geastraceae. By Patricio Ponce de Leon . . 303

15. Studies in American Plants. By Dorothy N. Gibson 353

16. Two New Nicaraguan Juglandaceae. By Antonio Molino R 357

17. Studies in the Palm Genus Syagrus Mart. By S. F. Glassman .... 363

18. Tropical American Plants, IX. By Louis O. Williams 401

PRELIMINARY NOTES ON THE

SCROPHULARIACEAE

OF PERU

GABRIEL EDWIN

NEW SPECIES IN THE PALM GENUS

SYAGRUS MART.

S. F. GLASSMAN

TROPICAL AMERICAN PLANTS, VIII

LOUIS 0. WILLIAMS

Univeistiy of Illinois

MAR 20 1968

FIELDIANA: BOTANY

VOLUME 81, NUMBERS 8, 9, 10

Published by

FIELD MUSEUM OF NATURAL HISTORY

APRIL 24, 1967

WOLOGY LIBRARV

PRELIMINARY NOTES ON THE

SCROPHULARIACEAE

OF PERU

GABRIEL EDWIN

Assistant Curator, Vascular Plants

FIELDIANA: BOTANY

VOLUME 31, NUMBER 8

Published by

FIELD MUSEUM OF NATURAL HISTORY

APRIL 24, 1967

PUBLICATION 1023

Printed with the Assistance of

The Frederick R. and Abby K. Babcock Fund

Library of Congress Catalog Card Number: 67-196^9

PRINTED IN THE UNITED STATES OF AMERICA

BY FIELD MUSEUM PRESS

Preliminary Notes on the Scrophulariaceae

of Peru

During preparation of the Scrophulariaceae for the Flora of Peru,

specimens were studied which necessitated reconsideration of the

generic limits within the tribe, Gratiolae. A number of others were

taxa not yet described. Finally, the establishment of a new tribe

appeared necessary.

This paper will concern itself with the first and third groups noted

above as well as novelties of the second, except those in the genera

Calceolaria and Bartsia.

Familial and generic limits of the "figworts" have been discussed

almost since the establishment of the family in 1789. Opinion about

these limits is far from unanimous today.

The genera occurring in Peru with the notable exception of Cal-

ceolaria (and to a lesser degree Alonsoa and Bartsia) have been elab-

orated elsewhere. None of the genera is represented in Peru by more

than six species, most having fewer than four. At least two genera,

Maurandya and Antirrhinum, are known only from cultivation.

In the course of time, evolution, natural selection and other proc-

esses have led to variation. However, in my opinion, not enough

change is apparent to warrant the current generic separation in the

Gratiolae. First, it is necessary to divide the Gratiolae into two sub-

tribes.

Gratiolinae, Subtribus novum.

Capsula plerumque septicidalis, aliquando secondarius loculicidalis, findens

ad basim aut si loculicidalis nunc sepalum extimum latissimum. Placenta simplex.

Sepala plerumque discreta aut subdiscreta (Linaria spp. excepto), interdum late

valde inaequalis. Lamina et capsula plerumque glanduloso-punctatis. Inflores-

centia plerumque racemus simplex aut racimo interdum ferens flores fasciculatum

et pedicellatum. Lobi corollae anticae plerumque interium glabrum.

Capsule usually septicidal, occasionally secondarily loculicidal, splitting to the

base, or if loculicidal the outermost sepal broader than the others; placenta simple;

sepals usually distinct or almost so, sometimes very unequal in width (Bacopa);

225

226 FIELDIANA: BOTANY, VOLUME 31

leaf-blades and capsules usually glandular-punctate; inflorescence usually simply

racemose or the raceme sometimes with fascicles of pedicellate flowers; anterior

lobes of the corolla usually smooth and glabrous within.

Type species: Scoparia dulcis L.

Mimulinae, Subtribus novum.

Mimulae Benth.

Capsula loculicidalis et sepalum extimum aequans cetera in late, aut indehis-

cens, secondarius vix aut haud septicidalis. Placenta ramosa. Sepala connata

plus quam media longitudo. Lamina et capsula glabra aut pubescentia, non glan-

duloso-punctatis. Inflorescentia racemosa aut cymis axillariis unique terminans

pedunculam basalis. Lobi corollae anticae 2-porcata et interium pubescentium.

Capsule loculicidal and the outermost sepal about equal to the others in width,

or indehiscent (secondarily barely if at all septicidal); placenta branched; sepals

united over half their length; leaves and capsules glabrous or pubescent but not

glandular-punctate; inflorescence racemose or of axillary cymes one of which is

terminal to the basal peduncle; anterior corolla lobes 2-ridged and pubescent in-

ternally.

Type species: Mimulus ringens L., of North America.

As now defined the Mimulinae consists of the genera Mimulus and

Leucocarpus. The remaining genera of the Gratiolae pertain to the

Gratiolinae.

Stemodia L., Syst. Nat. ed. 10, 1118. 1759. Lendernia Minod, Bull.

Soc. GeneVe 10 : 240. 1918. Unanuea R. & P. ex Pennell, Proc. Acad.

Nat. Sci. Phila. 72: 160. 1920.

Lendernia according to Minod differed from Stemodia in the de-

hiscence of the capsule, the splitting in Lendernia almost reaching the

base of the capsule while in Stemodia the rupture is complete. This

is hardly of generic significance.

Unanuea, based on Stemodia suffruticosa HBK., proposed by Pen-

nell was said to differ from Stemodia L. in having widely spreading

corolla lobes, internally glabrous corollas, ebracteolate pedicels and

sessile or clasping leaves. Without comment on the value of such

characters, it is necessary only to note that Pennell annotated all

specimens as Stemodia suffruticosa HBK. for at least 25 years after

publication of his new genus in 1920.

Bacopa Aubl., Hist. PI. Guiane Fran. 128, t. 49. 1775 (nom. cons.).

Moniera P. Br., Hist. Jam. 269. 1756. Bramia Lam., Encyc. Meth.,

Bot. 1 : 459. 1785. Mecardonia R. & P., Fl. Peruv. et Chil. Prod. 95.

EDWIN: SCROPHULARIACEAE OF PERU 227

1794. Herpestis Gaertn. f., Fruct. et Sem. Plant. 3: 186. 1807.

Macuillamia Raf., Neogenyton 2, 1825.

In 1920 and in 1935 Pennell (1920, 1946) divided Bacopa into a

number of genera including all the above synonyms as well as Pagesia

Raf., Hydranthelium HBK., Niadothrix Pennell and a few others.

By 1946, after studying more material, especially of eastern South

America, he united most of these segregates but retained Mecar-

donia, on the basis of capsule dehiscence, corolla color and anther

position on the filament. These characters vary throughout Bacopa.

Pennell 's treatments of the Bacopa-Herpestis "complex" are in-

adequate. The taxa concerned vary randomly in calyx lobe shape,

stamen and anther characteristics, style branching, corolla color and

in other ways.

At present, there probably is no truly satisfactory way to arrange

the species. However, when treated as one genus, they are quite

readily separable from the remainder of the Gratiolae on the basis of

the structure of the calyx lobes, the basis used by Wettstein in 1891.

While one character is usually considered insufficient for generic dis-

tinction, it should be kept in mind that in this case it is the best of

poor choices. Uniting the taxa in one genus allows for ready iden-

tification and naming of the taxa concerned1 and for the combining

of random and overlapping variations.

Ouriseae, Tribus novem.

Antice lobi corollarum extimae et superpostae alabastrae. Stigmatis lobi con-

nata; antherae divaricatae.

Anterior lobes of the corolla external and overlapping in bud; stigma lobes

united; anthers divaricate.

Type species: Ourisia chamydrifolia Benth.

A unigeneric tribe, as circumscribed, which lies between the Digi-

taleae and the Veroniceae in the Rhinanthoideae. The Ouriseae has

the united stigma lobes of the Digitaleae and the divaricate anthers

of the Veroniceae.

Species of Ourisia are almost always acaulescent herbs; the five

calyx lobes are united at least part way; they have tubular corollas

and didynamous and included stamens. Except for their habit, the

Ouriseae resemble the Digitaleae more than the Veroniceae.

1 It is useful to note that the species Bacopa acuminata (Walt.) Robinson has

been interpreted by various workers to belong to five of the segregates of Bacopa

discussed above as well as to five other genera, one in the other subfamily!

228 FIELDIANA: BOTANY, VOLUME 31

A great number of new species and novelties below the rank of

species could be presented with some justification. However, those

few appearing are proposed on a most conservative basis. A very

broad view of the species is taken: it must be distinguished easily

from its nearest (morphological) relatives. No other criteria have

been employed, largely since so little of such data (pollen analysis,

chromosome number, etc.) are available from Peru. The same cri-

terion applies to subspecific entities.

Sibthorpia rotundifolia (R. & P.) Edwin, comb. nov. Veronica

rotundifolia R. & P., Fl. Peruv. 1 : 6. 1798. Sibthorpia pichinchensis

HBK., Nova Gen. & Sp. PI. 2: 390, 1. 176. 1818.

The Veronica of R. & P. is unquestionably the Sibthorpia of HBK.

Sibthorpia rotundifolia (R. & P.) Edwin f. alba Edwin, f. nov.

Differt a typica corollis viridi-albus, capsulis pubescentiis et pilis rigidus.

Differing from the typical in having greenish-white (instead of rose-purple)

corollas and bearing stiff hairs on the capsules.

Cusco: Rio Yanamayo, below Pellahuata. Repent herb. Wet

mossy rocks by cascade at 2100-2300 m. alt., 4-5 May 1925, F. W.

Pennell 13935 (F, Type; GH; NY; PH).

The several synonyms not pertaining to Peru, and therefore un-

listed, only begin to indicate the taxonomic and nomenclatural prob-

lems remaining in Sibthorpia. Two Central American species and at

least one species each in Africa, Europe and South America require

further study.

It is unfortunate that Pennell's (1921) excellent paper concerning

Veronica and its allies in North and South America did not treat Sib-

thorpia. The generic lines as drawn in this work appear to be as valid

now as when published. The exclusion of Hebe (and Veronicastrum)

from Veronica permits a treatment of the taxa occurring in Peru that

is both natural and direct, in contradiction to the species limit prob-

lems of Sibthorpia.

Alonsoa integrifolia Edwin, sp. nov.

Frutex ramosus, caulis teres, longitudiner striatus, supra glanduloso-pubescen-

tia. Folia lineare-lanceolata usque ad ellipticolanceolata interdum linearia, ad

2 cm. longa, sessilia, angusta ad bases et apices margines integri. Inflorescentiae

racemi. Calycis lobi lanceolati, glandulosa, pubescentes, 7-8 mm. longa, plus

quam quarta longi capsula maturata. Corollae rosae. Antherae longiores quam

filamenta. Ovaria ovoidea, stigmatis-lobi connati, subpatelliformes.

EDWIN: SCROPHULARIACEAE OF PERU 229

Branched shrub drying dark brown to black, stems terete, longitudinally stri-

ate, new wood dark grey, glandular-pubescent at least on the upper part; leaf-

blades sessile, linear-lanceolate to elliptic-lanceolate, 0.5-2.0 cm. long and 0.1-

0.3 cm. wide, entire, narrowed at apex and somewhat to base, decreasing in size

from base to apex of stems, punctate beneath, lower blades glabrous, upper some-

times puberulent or sparsely glandular-puberulent beneath, costa little impressed

above, flattened and elevated beneath, lateral veins obsolete above and usually

beneath, when present little elevated; inflorescences racemose, the internodes of

the stem little if at all reduced in length, pedicels 10-12 mm. long, glandular-

puberulent, usually 2 times longer than the subtending bractlike leaf; calyx tube

very short, sparsely glandular-puberulent, calyx lobes lanceolate, 7-8 mm. long,

more than one-fourth as long as the mature capsule; corolla red; stamens 4, the

confluent anthers longer than the black, flattened filaments and dehiscing longi-

tudinally across the septum; ovary ovoid or occasionally lance-ovoid; style little

longer to little shorter than the ovary; stigma lobes united, subpatelliform; cap-

sules lance-ovoid or ovoid, ca. 12 mm. long, about 6 mm. wide at the base, narrow-

ing regularly to about 1-2 mm. wide at the apex; seeds numerous, black, longitudi-

nally furrowed, usually blunt, occasionally acute, at one or both ends. Mature

corollas few, poorly pressed, red or pale red.

CAJAMARCA: Michiquillo; shrub. On calcareous slope at 3000 m.,

7 April 1948, Pennell & Reichlin 15033 (BM, type; PH).

Alonsoa minor Edwin, sp. nov.

Frutex ramosus, in sicco brunneus vel atrobrunneus, apices ramorum et ramu-

lorum glandulosa-pubescentia. Folia angustissime lanceolata, usque ad 1.9 cm.

longa, 0.6 cm. lata, subsessilis, margines pauciserrata, dentibus angustissimis et

durissimis, attenuta ad bases et apices, plerumque minore subta paucipunctata,

puncti nigra. Inflorescentiae racemi. Pedunculi nulli, pedicelli glandulosa-pubes-

centia quam flores longiore. Calyces praefloratione hypocrateriforma, maturitati

tubi brevissimi, usque ad 1.5 mm. longa. Corollae atrorubrae, lobi quam tubo

breviore. Stamina ex tubum corollae exserta, antherae dolabriformi dense puberuli

quam filaments breviores. Ovaria anguste ovoidea, quam stylos longiore vel brevi-

ore. Capsula ovoides, apices obtusi, colli incrassati. Semina juvenalia nigra.

Branched shrub drying brown or dark brown, apices of branchlets glandular-

pubescent; leaf-blades very narrowly lanceolate, subsessile, up to 1.9 cm. long and

0.6 cm. wide, margins few-serrate, serrations very narrow and very hard, blades

attenuate to bases and apices, sparsely black punctate beneath; inflorescences

racemose; peduncles wanting; pedicels glandular-pubescent, up to 17 mm. long,

longer than the flower, fruit and subtending bracetolate leaf; calyces salverform in

bud, when mature the tube very short, up to 1.5 mm. long, calyx lobe 2.5-3.5 mm.

long, elliptical or narrowly elliptical, acuminate; corolla red to more often dark red,

lobes shorter than the tube; stamens exceeding the tube, exserted beyond the

mouth of the corolla, but shorter than the lobes, the axe-shaped, densely pubescent

anthers shorter than the black filaments; ovary narrowly ovoid, little longer or

shorter than the style; capsule (not fully mature) 9-10 mm. long and 4-5 mm.

wide, ovoid or occasionally narrowly ovoid, widest about one-third the way from

the base, the neck thickened and a little narrower than the obtuse to somewhat

rounded apex; seed (immature) black.

230 FIELDIANA: BOTANY, VOLUME 31

Good corollas are lacking. Most closely related to A. pallida (see

below) from which it differs in having smaller marginal teeth, blades

narrower in relation to length, shorter styles in relation to the ovaries

and shorter pedicels.

AMAZONAS: Above Colcamar. Shrub with red corolla, in thin soil

over limestone rocks, alt. 2600-2900 m., 24-26 June 1948, F. W. Pen-

mil 15609 (F, type; PH).

Alonsoa pallida Edwin, sp. nov.

Glabra herba erecta, in sicco pallidibrunneus, ramosa, rami et ramuli oppositi,

ramuli quadrati. Folia elliptica vel subovata elliptica, ca. 1 cm. longa X ca. 0.6 cm.

lata, brevipetiolata, margines paucidentati, serrati ad profundeserrati usque ad

sublaciniatos, interdum prope basim dimidiati, nervi subter elevatis, supra juve-

nalis puberulentis. Inflorescentiae racemi. Flores maturi ignoti, pedicelli glabri,

usque ad 9 mm. longos, plerumque capsulae non aequans. Bracteae minutae.

lanceolatae vel anguste-elliptici, acuminati, 3-nervati, ca. 3 mm. longi, calyx tubi

nulli. Corollae (fide Pennell) albae. Stamina in tubum corollae inclusa, antherae

breviores quam filamenta, sagittatae. Capsulae 7-10 mm. longae X 3-5 mm. latae,

ovoideae. Semina anthracina.

Glabrous, erect, branched herb, branches and branchlets opposite, branchlets

square, drying pale brown; leaf-blades elliptical or subovate-elliptical, up to 1 cm.

long and 0.6 cm. wide, short petiolate, margins few-toothed, serrate to deeply ser-

rate or laciniate blades sometimes dimidiate at base, young blades sometimes

puberulent above, nerves elevated beneath; petioles up to 2 mm. long, flattened;

inflorescences racemes; bracts minute, lanceolate or narrowly elliptic, margins en-

tire, apices obtuse, bases obtuse to occasionally subacute; pedicels glabrous, up to

9 mm. long, shorter than the capsules; mature flowers lacking; calyx lobes about

3 mm. long, lanceolate or narrowly elliptic, acuminate, 3-nerved; calyx tube lack-

ing; corolla white (fide Pennell); stamens included in the corolla tube, the sagittate

anthers shorter than the filaments; capsule 7-10 mm. long and 3-5 mm. wide,

ovoid: seeds shining black.

Mature flowers are lacking but this taxon is quite easily separated

from A. minor (see above) and from A. caulialata R. & P. by its

much smaller leaf -blades with fewer teeth, white corolla, and shorter

petioles and pedicels. In addition, A. caulialata has pedicels most

often longer than the capsules.

CAJAMARCA: Below Llama. Herb with whitish corolla. Banks,

1900-2100 m. alt., 17 July 1948, F, W. Pennell 15918 (BM, type; PH).

Alonsoa linearis (Jacq.) R. & P. var. glaberrima Edwin, var. nov.

Differt a typica antheris brevioribus quam filamentis et laminis et pedicellis

et calycibus glabrous.

Differs from the typical variety in having anthers shorter than the filaments

and the herbage, pedicels and calyces glabrous.

EDWIN: SCROPHULARIACEAE OF PERU 231

LIMA: Canta; shrubby, corolla "scarlet red." On open rocky

slopes, alt. 2700-2900 m., 11-19 June 1925. F. W. Pennell 14330,

(F, type; GH; NY; PH). Lima: San Buenaventura; anthers yellow.

On open rocky slopes, F. W. Pennell 1^556 (F).

REFERENCES

BENTHAM, G.

1846. In A. de Candolle, Prodromus 10: 186-586 (passim).

PENNELL, F. W.

1920. Scrophulariaceae of Colombia. I, Proc. Acad. Nat. Sci. Phila. 72: 136-188.

1920. In Britton, Descriptions of Cuban Plants New to Science, Mem. Torr.

Bot. Club 16: 103-106.

1921. "Veronica" in North and South America, Rhodora 23: 1-41.

1935. Monogr. Acad. Nat. Sci. Phila. 1: 49-69.

1946. Reconsideration of the Bacopa-Herpestis Problem of the Scrophularia-

ceae, Proc. Acad. Nat. Sci. Phila. 98: 83-98.

THIERET, J. W.

1954. The Tribes and Genera of Central American Scrophulariaceae. Ceiba 4:

164-184.

WETTSTEIN, R. v.

1891. Die Natur. Pflanzenf. 4(3b): 76.

Publications 1023, 1024 and 1025

UNIVERSITY OF ILLINOIS-URBANA