JOURNAL
OF THE
ARNOLD ARBORETUM
HARVARD UNIVERSITY
EDITORIAL BoarpD
A. C. SMITH, Editor P. C. MANGELSDORF
I. W. BAILEY E. D. MERRILL
J. H. FAULL H. M. RAUP
I. M. JOHNSTON A. REHDER
C, E. KOBUSKI K. SAX
VOLUME XXVI
JAMAICA PLAIN, MASS.
1945
Reprinted with the permission of the
Arnold Arboretum of Harvard University
KRAUS REPRINT CORPORATION
rk
1968
No
No
No
No
eT as
(WV CCP), te
DATES OF ISSUE
. 1 (pp. 1-121, 5 pl.) issued January 15, 1945,
. 2 (pp. 123-228, 11 pl.) issued April 16, 1945.
. 3 (pp. 229-364, 5 pl.) issued July 16, 1945.
. 4 (pp. 365-518) issued October 15, 1945.
Printed in U.S.A.
TABLE OF CONTENTS
PLANTAE PAPUANAE mm. longis, 0.7-0.8 mm.
latis, petalis minutis albicantibus subdolabriformibus cuspidatis, ca. _ 5
mm. longis, 0.6—-0.7 mm. latis, disco annulari-pulvinato integro ca. 0.25 n
alto; staminibus pro seer floris gc brevibus, oppositisepalis, le seiee
oblongis ca. 0.7 mm. longis et 0.6 mm. latis, apiculatis, basi divaricatim
og pining filamento paar ne ca 0. 5 mm. longo; pistillodio ovoideo ca.
m. longo, apice 3- -partito, laciniis aequalibus stylum cum stigmatibus
ie sia ris @ perianthio unico sub fructu viso, putrido, saltem
annotino, ca. 3 mm. lato, breviter stipitato, columella ca. 3.5 mm. longa,
basi florum 3 Sean i glomerulis fulto.
Viti Levu: Serua: In hills, alt. about 200 m., Greenwood 1018 (A, TYPE), May,
1943 (tree 5 m. high)
This is the first record of Cleistanthus Hook. f. in the Fiji Islands. The
genus has so far been known only from certain islands of Micronesia, such
as the Pelew and Truk groups (C. carolinensis Jabl., C. insularis Kanehira,
C. angularis Kanehira, C. Morii Kanehira), and New Caledonia (C. stipi-
tatus Muell. Arg.), not to mention points farther west, from Australia to
eastern Africa, which represents its probable center of dispersal. Cleistan-
thus micranthus is most nearly allied to C. stipitatus Muell. Arg., but differs
in the spreading primaries, the less coriaceous and blunter leaf, and the
smaller, glabrous inflorescence. In the classification of Jablonszky (in
Pflanzenr. 65[IV. 147. VIII]: 35. 1915), C. micranthus apparently falls
into Sect. Australes Jabl., together with C. stipitatus (Baill.) Muell. Arg.
and C. Dallachyanus (Baill.) Benth. Thus composed, this section ranges
from Australia to New Caledonia and the Fiji Islands.
Croton Parhamii Croizat, sp. nov.
Frutex innovationibus hispido- senate sena serius glabrescentibus, tri-
chomatibus saepius brunneis pilo centrali porrecto. _ Foliis elliptico- lanceo-
latis brunnescentibus, 3-9 cm. longis, 2—4 cm. latis, tenuiter chartaceis,
apice sat obtuse acuminatis, basi rotundatis vel rotundato- cuneatis, margine
sat obscure crenato-dentatis, supra glabris, subtus trichomatibus saepissime
stellatis hispidis in lamina parcius in costa crebre indutis, venis penninerviis
arcuato-adscendentibus anastomosantibus ca. 8-12-jugis, primo jugo haud
teens. petiolo gracili aeque ac innovationibus induto 0.5—2 cm. longo,
glandulis posticis ad basim laminae 2, breviter stipitatis disciformibus.
1945] SMITH, PACIFIC ISLAND PLANTS, IV 99
Inflorescentia subspicata ad 10 cm. longa. Floribus ¢ : pedicello ca. 3-6
mm. longo, perianthio in alabastro subglobuloso, in anthesi latius cyathi-
formi (ut videtur), ca. 2 mm. longo et 3 mm. lato, petalis cum sepalis sub-
aequilongis, staminibus ca. 10. Floribus @: perianthio extus pubescente
late campanulato ca. 3 mm. longo et 5 mm. lato, pedicello 8-10 mm. longo,
perianthii lobis ad basim liberis ellipticis, ca. 1-2 mm. longis et 1.5 mm.
latis, margine integris, uno alterove interdum subanisomero, petalis glandu-
lisque ut videtur nullis, ovario late ovoideo indumenti copia brunneo-hispi-
dulo, ca. 1.5 mm. magno in stylum evidenter abeunte ; stvlorum cruribus 3,
pro ratione sat crassis, nigris, epapillosis dorso puberulis, ca. 2 mm. longis,
fere ad basim 2-partitis, columella fructu delapso gracili ad 5 mm. longa.
Vitt Levu: Tholo West: Ridge between Naloka and Naraiyawa, in forest, alt.
about 900 m., B. E. Parham 2464 (A, tyPE), July 26, 1938.
The slightly accrescent @ perianth in fruit, with evolute lobes at anthesis,
readily distinguishes this new species from C. heterotrichus Muell. Arg.,
which it suggests at first in the pubescence of the lower leaf-surface.
Croton Storckii (Muell. Arg.) Seem. has a different foliage and @ perianth,
with a much smaller ovary and scarcely accrescent lobes. Croton Ver-
reauxii Baill. is an Australian species, with a ? perianth that has characters
similar to those of C. Storckii. Croton microtiglium Burk. is known only
from Tonga and suggests the characters of C. leptopus Muell. Arg., to judge
from the description and an authentic specimen of the latter here available.
Croton Levatii Guillaumin, of the New Hebrides, is very summarily de-
scribed (in Bull. Soc. Bot. France 66: 275. 1919), but the remark “sepalis
petalisque angustioribus” rules out C. Parhamii.
ELAEOCARPACEAE
Elaeocarpus cassinoides A. Gray, Bot. U. S. Expl. Exped. 1: 204. 1854; C. Muell. in
Walp. 4: 331. 1857; Seem. FI. Vit. 29. 1865; Hemsl. in Jour. Linn. Soc. Bot.
30: 171. 1894.
Vanua Levu: Mbua: Mbua Bay, U. S. Expl. Exped. (GH, Type covt.) ; lower
Wainunu River Valley, alt. 0-200 m., Smith 1735 (GH) (tree 13 m. high, in open
forest; native name: wailoaloa). Koro: Eastern slope of main ridge, alt. 200-300
m., Smith 1007 (GH) (spreading tree 23 m. high, in dense forest ; trunk 1 m. diam
western slope, alt. 300 m., Smith 1086 (GH) (tree 14 m. high, in thickets; petals pale
pink).
Elaeocarpus cassinoides has previously been known only irom the type
collection, a fruiting specimen. Although it was originally reported by
Gray from both Fiji and Tonga, its occurrence in the latter group is ques-
tionable; Gray remarks, “Those [specimens] from the two localities, as
ticketed, are so exactly alike that they might have been taken from the
same stem, and, since the habitats are not to be verified from Dr. Picker-
ing’s notes, one or the other may be considered doubtful.” Mueller, See-
mann, and Hemsley, in the publications cited above, repeat Gray’s data
without citing additional specimens, and it seems that the species has not
been re-collected in Tonga. Since the Fijian specimens cited above are
obviously conspecific with the type, I believe that the record of this species
from Tonga is erroneous. In view of the inadequacy of the original descrip-
100 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
tion, I here redescribe the species. Of the cited specimens, Smith 1007 and
1086 are in flower and the others in fruit.
Tree, up to 23 m. high, the branchlets subterete, slender, brownish, at first
pale-puberulent, soon glabrescent; petioles puberulent and glabrescent like
the branchlets, slender, canaliculate, 5-15 mm. long; leaf-blades papyra-
ceous or chartaceous, oblong- or obovate-elliptic, 5-10 cm. long, 2—4.5 cm.
broad, subacute to attenuate at base, obtuse or obtusely cuspidate at apex,
inconspicuously serrulate especially distally (teeth 2 or 3 per centimeter),
glabrous on both surfaces or obscurely puberulent on nerves beneath when
For the time being I am unable to refer E. cassinoides to its appropriate
section. It is probably related to such New Guinean sections as § Dactyl-
osphaera Schlechter and § Fissipetalum Schlechter, but it does not entirely
agree with either of these. Another Fijian species of this general relation-
ship is E. kastensis A. C. Sm., which was originally referred to § Dicera.
However, since I have studied the New Guinean species of Flaeocarpus and
looked into the typification of § Dicera (see Jour. Arnold Arb, 25: 223.
1944), I realize that the two Fijian species here discussed should not be
placed in § Dicera. A study of all the Pacific species is advisable before
these can be properly placed. Two other Fijian species possibly of this
relationship, both very inadequately known, are E. laurifolius A. Gray and
E. pyriformis A. Gray.
Elaeocarpus (§ Dicera) pittosporoides sp. nov.
Arbor ad 6 m. alta partibus florum exceptis ubique glabra, ramulis
vel chartaceis in sicco fuscis vel olivaceis, obovato-ellipticis, 6.5-10 cm
longis, 2.5—4.5 cm. latis, basi attenuatis et in petiolum decurrentibus, apice
1945] SMITH, PACIFIC ISLAND PLANTS, IV 101
obtuse cuspidatis, margine leviter recurvatis et remote undulato-crenulatis,
u erect
acutis, extus glabris, intus minute puberulis et conspicue carinatis, margine
leviter incrassatis; petalis 5 submembranaceis, obovatis, longitudine sepalos
aequantibus, 2.5—3 mm. latis, intus basim versus paullo tomentellis ceterum
glabris, apice 3—5-lobulatis, lobis 1-2 mm. longis subacutis; disco annulari-
pulvinato circiter 0.5 mm. alto obscure 5-lobato superne sparse hispidulo;
staminibus 26 vel 27 circiter 3.5 mm. longis 1- vel 2-seriatis ubique minute
papilloso-hispidulis, filamentis subteretibus 1—1.5 mm. longis, antheris
2-2.5 mm. longis, apice subacutis erostratis; gynaecio glabro, ovario
ovoideo 2-loculari, loculis 4-ovulatis, stylo crasso subulato 2—2.5 mm. longo.
Vit1 Levu: Namosi: Hills between Navua River and Suva, alt. 200-300 m.,
Greenwood 1010 (A, TYPE), May, 1943 (upright tree 5-6 m. high; inflorescences in
leaf-axils near apices of branchlets; flower-buds yellow, somewhat dependent on very
thin pedicels).
Elacocarpus pittosporoides appears to have no close relatives among the
known Fijian species. I have recently (in Jour. Arnold Arb. 25: 223.
1944) had occasion to discuss the typification of § Dicera, to which the new
species definitely belongs. In floral characters it is very similar to E.
dentatus (J. R. & G. Forst.) Vahl, the type of the section, but in foliage
and in its slender lax inflorescences it is quite different.
TILIACEAE
Brownlowia sp.
Virt Levu: Lautoka: Mountains near Lautoka, alt. about 550 m., Greenwood
957 (A) (shrub or small tree) ; Namosi: Hills between Navua River and Suva, alt.
about 250 m., Greenwood 957A (A); vicinity of Namuamua, alt. 400 m., Gillespie 3000
(GH); Rewa: Slopes of Korombamba Mt., alt. 200 m., Gillespie 2372 (GH).
The cited specimens indicate a substantial extension of the known range
of Brownlowia; all are sterile, but their place in the genus seems certain,
and it is nearly equally certain that they represent an undescribed species.
The Fijian specimens are probably of the relationship of B. argentata Kurz,
as this is represented by material from New Guinea and the Solomon
Islands. The occurrence of the genus in the New Hebrides, not previously
reported, is indicated by Kajewski 616 (A) from Vanikoro, a specimen
probably referable to B. argentata. From this species the Fijian specimens
differ in obvious foliage characters, the leaf-blades being longer and with
more numerous lateral nerves.
Yuncker (in Bishop Mus. Bull. 178: 80. 1943) has recently reported
Brownlowia from Niue, also in sterile condition; this record appears to be
the easternmost for the genus.
ELATINACEAE
Elatine gratioloides A. Cunn. in Ann. Nat. Hist. 4: 26. 1840, Nied. in E. & P. Nat.
Pfl. ed. 2. 21: 276. 1925; Greenwood in Jour. Arnold Arb. 25: 398. 1944.
102 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
Elatine americana sensu Hook. f. Fl. Nov. Zel. 1: 27. 1853
sage americana var. australiensis Benth. FI. rena 1; 178. 1863; Cheesem. Man.
I . Fl. 73. 1906, ed. 2. 568. 1925; F. M. Bailey, Weeds & Pois. Pl. Queensl.
22. 37. [1907].
Vitt Levu: Lautoka: Mountains near Lautoka, alt. about 600 m., Greenwood
952 (GH) (entire plant light green; creeping on mud under two inches of slowly
running water, in taro plantation).
This species, until Greenwood (1. c.) mentioned it, had previously been
reported only from Australia and New Zealand. The collection of a speci-
men of Elatine in Fiji is of especial interest, since, to the best of my knowl-
edge, only one other collection of the genus has previously been cited in
literature from the entire Pacific region (other than New Zealand). The
earlier collection was Seemann 183, from the island of Taveuni, Fiji, which
was referred by Seemann (FI. Vit. 10.1865) to E. ambigua Wight. Accord-
ing to Niedenzu (in E. & P. Nat. Pfl. ed. 2. 21: 276. 1925), E. ambigua and
E. gratioloides are distinct species, the flowers being pedicellate in the
former and sessile in the latter. Niedenzu states the range of E. ambigua
to include Fiji, and I assume that his record is based upon Seemann 183.
From an examination of a duplicate of this number in the Gray Herbarium,
I am inclined to believe that it also represents E. gratioloides, since the
flowers are usually essentially sessile, only rarely being on short pedicels
about 1-2 mm. long. The value of pedicel-length as a specific character
may be questioned, since in all other features Seemann 183 agrees with
Greenwood 952 and the several New Zealand and Australian specimens in
the Gray Herbarium. It seems probable to me that the only Fijian species
of the genus is E. gratioloides, but the difference between this species and
E. ambigua should be carefully checked.
Bentham, following his original description of E. americana var. austra-
liensis, states: “This plant, whether a distinct species or a variety of the N.
American one, is found also in New Zealand and the Fiji islands, and is
very variable.” His record from Fiji was doubtless based upon Seemann
183, which had vig eae referred to E. americana by A. Gray in
Bondlandix 10: 36.
In the most nee survey of the genus in its entire range, Niedenzu (I. c.
274-276) expresses the opinion that E. gratioloides is distinct from both
E. triandra Schkuhr and E. americana (Pursh) Arn. The customary pro-
cedure by writers on the flora of Australia and New Zealand is to mention the
austral plant as E. americana var. australiensis Benth. Fassett has recently
reduced #. americana to varietal rank, as E. triandra var. americana
(Pursh) Fassett (in Rhodora 33: 72. 1931, 41: 373. 1939), but the austral
variety is not placed by him. It remains for a monographer to decide what
status £. gratioloides merits; for the time being I follow Niedenzu in con-
sidering it a specific entity. The type was collected by R. Cunningham in
1833, in a bog at Tauraki, Hokianga River, North Island, New Zealand.
FLACOURTIACEAE
Flacourtia vitiensis (Seem.) comb. n
Thacombania vitiensis Seem. FI. Vit. 426. pl. 100. 1873.
Flacourtia ovata Gillespie in Bishop Mus. Bull. a 27. f. 34 (excl. a, f, g). 1931.
1945] SMITH, PACIFIC ISLAND PLANTS, IV 103
The monotypic genus Thacombauia was originally described by Seemann
as a member of the Humiriaceae, but subsequently it was questionably
referred to the Euphorbiaceae by Durand, Pax, and Dalla Torre and Harms.
A glance at Seemann’s plate demonstrates the identity of the plant with
Flacourtia ovata, which was recently emended by me in Sargentia 1: 61.
1942. While it is regrettable that the name honoring the famous King
Thacombau thus falls into synonymy, nevertheless it is a source of satis-
faction to place the genus.
MYRTACEAE
Syzygium phaeophyllum mi & Perry, nom.
Eugenia durifolia A. C. Sm. in Bishop Mus. Bull. 141: 105. f. 56. 1936.
Syzygium durifolium Mertill . ats in Sargentia 1: 76. 1942, non in Mem. Am.
Acad. Arts & Sci. 18: 176.
Dr. E. H. Walker has eee our attention to an oversight in our use of
the specific epithet durifolium for two different species, the earlier from
Borneo and the later from Fiji. Consequently, the Fijian species is here
renamed. — E. D. Merrill and L. M. Perry
MYRSINACEAE
Tapeinosperma Greenwoodii sp. nov.
Arbor ad 5 m. alta ubique partibus juvenilibus et inflorescentiis puberulis
exceptis glabra, ramulis subteretibus rugulosis cinereis apicem versus 3—6
mm. crassis; foliis apicem ramulorum versus aggregatis, petiolis angulatis
supra complanatis crassis (1.5—2 mm. diametro) 5—8 mm. longis, Jaminis
chartaceis juventute copiose pellucido- -punctatis mox opacis in sicco fusco-
olivaceis elliptico-oblongis vel obovatis, 12-16 cm. longis, 5—8 cm. latis,
basi gradatim angustatis et in petiolum decurrentibus, apice anne vel
obtuse cuspidatis, margine saepe undulatis et paullo recurvatis, costa supra
leviter canaliculata subtus prominente, nervis lateralibus primariis utrinse-
cus 15-20 cum aliis debilioribus interspersis patentibus rectis marginem
versus anastomosantibus utrinque valde prominulis, rete venularum intri-
cato utrinque paullo prominulo; inflorescentiis axillaribus multifloris tri-
vel quadripinnatim paniculatis, 12—20 cm. longis, 6-14 cm. latis, pedunculo
4—6 cm. longo et rhachi ramulisque gracilibus dense brunneo-glanduloso-
puberulis; bracteis deltoideo-oblongis circiter 1.1 % 0.7 mm. subacutis
utrinque puberulis et margine ciliatis; pedicellis teretibus gracilibus 0.8-1
mm. longis puberulis; calyce rotato-cupuliformi circiter 2.5 mm. diametro,
basi rotundato, extus parce glanduloso- ee intus glabro, lobis 5 fere
ad basim aperts late ovatis, circiter 0.8 & 1.1 mm., margine minute
glanduloso-ciliolatis, apice emarginatis vel interdum rotundatis; corolla
(paullo ante anthesin) circiter 3 mm. longa, lobis fere ad basim liberis
ovatis 2—2.5 mm. latis, apice acutis, superne leviter punctatis; staminibus
prope basim corollae insertis subsessilibus, antheris elongato- deltoideis
circiter 1 X 0.8 mm. saepe glandulas 2 vel 3 dorso gerentibus; gynaecio sub
ae circiter 1.6 mm. longo, ovario ovoideo in stylum. gracilem circiter
1 longum attenuato, stigmate minuto subpeltato, placenta ovoidea
2- Ses
Vitrt Levu: Lautoka: Mt. Evans, alt. about 900 m., Greenwood 944 (A, TYPE),
Oct. 25, 1942 (small tree 3-5 m. high, the inflorescences lax, dependent).
104 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
Tapeinosperma Greenwoodii is most closely allied to 7. clavatum Mez,
with which it has in common short petioles, elliptic leaf-blades of moderate
size and with prominulous venation, and ciliate and usually emarginate
calyx-lobes. However, the new species has a much more ample and more
complex inflorescence, the pedicel is slender and not conspicuously swollen
distally into a conical calyx like that of 7. clavatum, and the flowers are
smaller throughout. The calycine character seems dependable, as the
several specimens of 7. clavatum examined have the calyx as described by
Gillespie (in Bishop Mus. Bull. 74: 9. f. 8. 1930). The new species may
also be compared with T. //ornez Mez, a species with long-petiolate large
leaves, a shorter, simpler, and lepidote inflorescence, and larger flowers with
acute or acuminate calyx-lobes.
RUBIACEAE
Mussaenda L.
Although it has been customary to refer the Pacific representatives of
Mussaenda to M. frondosa \.., examination of specimens and literature
indicates that this is erroneous. Students of the Pacific floras have appar-
ently been aware of this fact, but no alternative identification has been
seriously suggested for the common Pacific plant, which is, in some areas,
one ci the most abundant elements of second growth vegetation. However,
a species supposedly endemic to Raiatea, M. raiateensis J. W. Moore, was
described in 1933. A careful examination of abundant Pacific material of
the genus, including an isotype of M. raiateensis (for the loan of which I
am greatly indebted to Dr. Moore, of the University of Minnesota), in-
clines me to believe that a single species occurs from the New Hebrides to
the Society Islands, to which Moore’s binomial may be applied. A dis-
cussion of the relationships of this Pacific species and an emended descrip-
tion follow.
The Linnaean species, M. frondosa (Sp. Pl. 177. 1753), is based upon
several earlier references, among them Linnaeus’ Flora Zeylanica (p. 35.
1747) and Burman’s Thesaurus Zeylanicus (p. 165, tab. 76.1737). The
actual type is a collection of Hermann from Ceylon. The Ceylon plant
described in the above references, and further amplified by Trimen (Hand-
book FI. Ceylon 2: 323. 1894), is a scrambling shrub, with the branchlets,
leaves, and the enlarged calyx-lobe densely and closely velutinous-tomen-
tose. A representative specimen from Ceylon is J, M. de Silva 38 (A).
The actual geographical range of M. frondosa is in doubt, but its occur-
rence in the eastern Malaysian and the Pacific regions is highly question-
able. In their recent work on the genus in Papuasia (in Jour. Arnold Arb.
25: 192-196. 1944), Merrill and Perry did not find the species represented
in the available material. The true M. frondosa, as represented by speci-
mens from Ceylon and India, differs from the Pacific entity not only in its
habit and its generally more obvious pubescence, but also in its longer
normal calyx-lobes (5-10 mm. long), in its anthers being more deeply in-
serted on the corolla-tube (tips of anthers 4-9 mm. below apex of tube),
and in its fruit being elenticellate or essentially so.
1945] SMITH, PACIFIC ISLAND PLANTS, IV 105
A closer relative of the Pacific plant than M. frondosa is M. philippica
A. Rich., of the Philippine and Solomon Islands. In common with the
Pacific entity, Richard’s species differs from M. frondosa in its more highly
placed stamens and its lenticellate fruits. From M. philippica, the Pacific
plant differs in its longer and more or less subulate, rather than deltoid-
lanceolate, calyx-lobes, and in its comparatively thick pericarp, which, when
mature, remains firm and does not readily break. Another species of the
Solomon Islands, M. Kajewskii Merr. & Perry (in Jour. Arnold Arb.
25: 194, 1944), is very close to M. philippica and differs from the Pacific
plant in the same characters pertaining to comparatively short calyx-lobes.
The common Mussaenda of Micronesia has been mentioned in the litera-
ture as M. frondosa L. (Volkens in Bot. Jahrb. 31: 475. 1901; Safford in
Contr. U. S. Nat. Herb. 9: 330. 1905; Merr. in Philip. Jour. Sci. Bot.
9: 147. 1914; Fosberg in Occ. Pap. Bishop Mus. 15: 215. 1940) and M.
sericea Bl. (Valeton in Bot. Jahrb. 63: 300. 1930; Kanehira in Bot. Mag.
Tokyo 45: 351. 1931, Fl. Micrones. 369. 1933, in Jour. Dept. Agr. Kyushu
Univ. 4: 421. 1935). This Micronesian Mussaenda is certainly not M.
frondosa in the limited sense of Ceylon and Indian plants, and its identity
with M. sericea, based on a Moluccan specimen, is open to doubt. How-
ever, I see no reason to exclude the Micronesian specimens now available to
me (Kanehira 90, 1157, 1177, 1993, Herre 12 {all NY]) from M. philip-
pica A. Rich. At any rate, these Micronesian plants differ from those of
the more southerly islands (New Hebrides to Societies) in the characters
above discussed for M. philippica.
Another species of Mussaenda occurring in the Pacific region is M. cylin-
drocarpa Burck, found in the New Hebrides (Guillaumin in Jour. Arnold
Arb. 13: 4. 1932) but apparently not extending farther east into the Pacific.
This species is characterized by having its stamens deeply inserted on the
corolla-tube and its fruit cylindric, and it is not concerned in a discussion of
the identity of the species which is so abundant in Fiji, Samoa, and east-
ward.
To summarize, it seems obvious that the common Mussaenda which ex-
tends from the New Hebrides to Rarotonga is amply distinct from M,
frondosa L. and that its closest allies are more probably M. philippica A.
Rich. and M. Kajewskii Merr. & Perry. This entity does not appear
separable from M. raiateensis J. W. Moore, although it is a fairly variable
species, in which subspecific divisions may eventually seem desirable. For
the time being I cannot tie up the slight morphological variations, such as
those pertaining to the degree of pubescence, with geographical regions.
The following description is based on all material of the species now avail-
able to me.
Mussaenda raiateensis J. W. Moore in Bishop Mus. Bull. 102: 44. 1933.
Mussaenda frondosa sensu Forst. f. Fl. Ins. Austr. Prodr. 17. 1786; A. Rich. Sert.
TO
Fl. Vit. 123. 1866; Powell in in our. Bot. 6: 370. 1868; Engl. in Bot. Jahrb. 7: 477,
pro parte. 1886, Forschungsr. Gazelle Siphon. 46, pro parte. 1889; Drake, Ill. FI.
106 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
Ins. Mar. Pac. 189. 1890; Hemsl. in Jour. Linn. Soc. Bot. 30: 180. 1894; poegprore
in Bot. Jahrb. 25: 690. 1898; Burkill in Jour. Linn. Soc. Bot. 35:40. 1901;
Cheesem. in Trans. Linn. Soc. Bot. II. 6: 282. 1903; Gibbs in Jour. iva Soc.
Bot. 39: 151. 1909; Wilder in Bishop Mus. Bull. 86: 102. 1931; Guillaumin in
Jour. Arnold Arb. 13: 4. 1932; Fosberg in Bull. Torr. Bot. Club 67: 420. 1940;
non L.
on formosa ene Seem. FI. - 123, as synonym. 1866 (forsan quoad
. Icon. ined. t. 56, 57), non Jac
Pritt frondosa var. pilosissima ee Reinecke in Bot. Jahrb. 25: 690. 1898;
Rechinger in Denkschr. Akad. Wiss. Wien a 368. 1910 (repr. 3: 194) ; Setchell
n Carn. Inst. Publ. 341: 43. 1924; non Eng
Shite sp. Christoph. in Bishop Mus. Bull. os 199. 1935.
Shrub or slender tree 2-10 m. high, sometimes said to be epiphytic or
subscandent, the branchlets lightly flattened when young, densely hispidu-
lous with pale or brownish hairs 0.3—0.5 mm. long, at length terete, glabres-
cent, brownish, striate, nace eran stipules interpetiolar, 7-17 mm
long, deeply bifid, sericeous toward base, sparsely hispidulous distally, soon
caducous, each lobe subulate-lanceolate; petioles slender, subterete or shal-
lowly canaliculate, 5-25 (-40) mm. long, densely hispidulous to glabrescent
like the branchlets; leaf-blades papyraceous, olivaceous or brownish when
dried, often slightly paler beneath, ovate or oblong-elliptic, 10—-18(—20)
cm. long and 5—10(—14) cm. broad (uppermost leaves sometimes smaller),
acute to attenuate (rarely subrounded or obtuse) at base and decurrent on
the petiole, gradually acuminate at apex (acumen 5—30 mm. long), entire
or lightly undulate at margin, sparsely hispidulous (often densely so on
veins) and often soon glabrescent above, more densely hispidulous beneath
(especially on nerves and in axils, the hairs pale, spreading, 0.5-1 mm.
long) but at length often essentially glabrescent, the costa nearly plane or
raised above, prominent beneath, the secondary nerves 7—10(—12) per side,
ascending, curved or nearly straight, prominulous above, strongly elevated
beneath, the veinlet-reticulation copious, plane or faintly impressed on both
sides; inflorescence terminal, several- to many-flowered, often wide-spreading
at maturity and up to 12 cm. long and broad but usually much smaller, the
branches and pedicels hispidulous like the petioles or puberulent, essentially
glabrous in fruit, the pedicels slender, short, up to 5 mm. long or essentially
none, the bracts (up to 6 mm. Jong) and the minute bracteoles linear-
subulate and soon caducous; calyx-tube oblong-cylindric or turbinate, at
anthesis 3-5 mm. long, hispidulous or sericeous with pale or brown hairs
mm. long, the lobes 5, erect, linear-subulate, 4—6.5 mm. long,
hispidulous, glabrescent within and distally, one lobe often enlarged :
base and contracted into a slender stipe 5-20 mm. long, short-acuminate or
cuspidate at apex, sparsely hispidulous on both sides or glabrescent, several-
nerved from base, the veinlet-reticulation often obvious; disk annular,
glabrous; corolla slenderly infundibular, pale yellow proximally, yellow or
orange distally, 23-40 mm. long including lobes, the tube hispidulous or
sericeous (often inconspicuously so) without, glabrous or subglabrous
within toward base and distally (above attachment of anthers) copiously
and densely soft-strigose with pale ascending hairs about 1 mm. long, the
lobes 5, spreading, oblong or ovate-deltoid, 3—6.5 mm. long and broad,
cuspidate at apex, usually puberulent without and copiously glandular-
1945 ] SMITH, PACIFIC ISLAND PLANTS, IV 107
puberulent within; stamens 5, glabrous, inserted in throat of corolla (tip of
anthers 1-3 mm. below apex of corolla-tube), the filaments very short, the
anthers linear, 3.5—5.5 mm. long, acute at apex, sagittate at base; style
filiform, glabrous, equaling or slightly exceeding the corolla-tube, bifid for
about is mm. at apex; fruit ellipsoid or turbinate, glabrous at maturity,
green when eh brownish or blackish when dried, 10-15 mm. long, 7—11
mm. broad, truncate at apex, obtuse at base and contracted into a slender
stipe 2-4 mm. long, the calyx-lobes caducous or the enlarged one some-
times subpersistent, the pericarp firm, copiously marked with large gray
elliptic lenticels, the seeds minute, black, reticulate
DISTRIBUTION: New Hebrides, Fiji, Samoa, Tonga, Rarotonga, and Society Islands;
probably also in some of the adjacent groups. Reported from a variety of habitats,
including clearings, thickets, open forest, and occasionally rain-forest; in Fiji it is one
of the commonest plants in second growth thickets and is rarely absent from this
habitat. Reported altitudes range from sea-level to 900 m. The type is Moore 684
c
NEW HEBRIDES: a a (NY) ; eaecenr Bay, Kajewski 922
(A, NY). FIJI: Vitr Levu: Lautoka: Mountains near Lautoka, Greenwood
164 (A); Tholo North: Nanda vivati, hep ien & aes 13526 in NY), Reay 9
(A); Sovutawambu, near Nandarivatu, Bagene 14602 (A, NY); Naitasiri: Vicinity
of Nasinu, Ce 3555 (NY); Waindina River basin, MacDaniels 1053 (A). Kan-
pAvu: Namalata Isthmus region, Smith 25 (GH, NY). Vanua Levu: Thakaun-
drove: Hills south of Nakula Valley, Smith 330 (GH, NY); Valanga, Savu Savu
Bay, Degener & Ordonez 13987 (A, NY); summit of Uluinabathi ae Savu Savu Bay
region, Degener & Ordonez 13936 (A, NY); Maravu, near Salt Lake, Degener &
Ordonez 14185 (A). TaAvEuNI: Vicinity of Wairiki, aia pe (NY). OVALAU:
Vicinity of Levuka, Gillespie 4471 (NY), Degener & Ordonez 13795 (A, NY). MAakon-
DRONGA: Degener & Ordonez 13814 (A, NY). Koro: pore slope of main ridge,
Smith 1030 (NY). Oneata: U. S. Expl. Exped. (NY). FuLanca: Smith 1173
(NY). Fiji, without other locality: U.S. Expl. Exped. (GH, NY), Seemann
238 (GH), Horne (GH), Prince (GH). SAMOA: Savai: Vaipouli, Vaupel 355
(NY); Salailua, Christophersen 2999 (NY). Upotu: Near Malololelei, Christopher-
sen 323 (NY). TAv: ee trail back of Luma, Garber 610 (NY). Samoa, with-
out other locality: U. S. Expl. Exped. (GH). TONGA: Toncatasu: Along
the Hala Loto, pnd Parks 15490 (GH). Eva: Western edge of the Plateau,
Parks 16200 (A, G Y). COOK ISLANDS: Rarotonca: Parks & Parks 22209
(GH). SOCIETY Lace RatATEA: West side of highest mountain, March 24,
1927, Moore 684 (TYPE COLL., herb. Univ. Minnesota).
MON NAMES (and sources): In Fiji: snes or Mbovo (all emilee Vakath-
arendavui (Smith), Vombo (Reay); in Samoa: Uto’uto, Ma’osina (Powell), Fua i
tausaga, Laupaepae (Setchell), Alo alo vao, ae (Christophersen) in Pa inice
Kotuku (Cheeseman, Wilder)
uses: In Fiji decoctions of the leaves and bark are commonly used as a
cure for fever, chest complaints, and kidney diseases.
Airosperma
Airosperma Lauterb. & K. Schum. in K. Schum. & Lauterb. Fl. Deutsch. Schutzgeb.
Siidsee 565. 1901.
Abramsia Gillespie in Bishop Mus. Bull. 91: 27. 1932.
r. L. M. Perry has kindly pointed out to me the apparent similarity of
Aweipermea (with four species endemic to New Guinea) and Abramsia
monotypic and supposedly endemic to Fiji). Careful examination of the
literature and the available specimens indicates that these two generic con-
108 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
cepts are essentially identical. We thus have another illustration of a
genus extending from New Guinea to Fiji, presumably to be anticipated in
the intervening island groups, from which it has not yet been reported.
In their original discussion of Airosperma, Lauterbach and K. Schumann
place the genus in the “‘Albertinae,” pointing out its essential characters of
pendulous solitary ovules and contorted corolla-lobes. In the classification
of K. Schumann (in E. & P. Nat. Pfl. IV. 4: 16. 1891), this appears to be
correct, and Azrosperma may be placed in the Coffeoideae-Guettardinae-
Alberteae (op. cit. 87-89). This is the position assigned to the genus by
Krause (in E. & P. Nat. Pfl. Nachtr. 3: 328. 1908).
Gillespie, in placing his new genus Abramsia, expresses a belief that its
relationships are in the Ixoreae; the pendulous ovules, however, would seem
to exclude consideration of this or any other division of the Psychotriinae.
Gillespie emphasizes precisely the same combination of characters which
differentiates Airosperma, and indeed his illustration (op. cit. fig. 31)
shows that the fundamental characters of Abramsia are similar to those of
Airosperma (illustrated in K. Schum. & Lauterb. Fl. Deutsch. Schutzgeb.
Sudsee ¢. 27. 1901).
Although the specific epithets originally associated with Airosperma were
psychotrioides and ramuensis, and although subsequent discussions have
treated the generic name as feminine, it must be considered neuter, accord-
ing to Art. 72 (2) of the International Rules of Botanical Nomenclature
(1935).
Airosperma trichotomum (Gillespie) comb.
Abramsia trichotoma Gillespie in Bishop a. ‘Bu Il. 91: 29. fig. 31. 1932; Fosberg in
Bull. Torrey Bot. Club 67: 422. 1940, in Sargentia 1: 125, 1942
Airosperma trichotomum is known from several collections from the
Fijian islands of Viti Levu, Vanua Levu, and Taveuni, which are cited in
the listed publications. The available material shows that the species is
fairly variable as to leaf-size, but in important respects it is reasonably
consistent.
Although linear calyx-lobes characterize Airosperma psychotrioides (the
illustrated New Guinean species), other species of the genus, notably A.
grandifolium Val., have the calyx-lobes more or less deltoid. In the Fijian
species the ily lobes are still smaller, but this character is hardly of
generic value. Gillespie’s species is nee marked by its large thin leaves
and comparatively ample inflorescences.
Mastixiodendron
ag rina Melchior ¥ Bot. Jahrb. 60: 167. 1925.
ia Gillespie in Hook. Pl. 32: pl. 3190. 1933.
> Ta Te and eee were both originally described as members
of the Cornaceae allied to Mastixia Bl. Géillespie’s genus was transferred
to the Rubiaceae by the present writer in 1936 (in Bishop Mus. Bull.
141: 140) and was referred to the tribe Chiococceae, but at that time
Mastixiodendron was insufficiently known to me to be carefully considered.
More recently Merrill and Perry (in Jour. Arnold Arb. 23: 416. 1942) have
1945] SMITH, PACIFIC ISLAND PLANTS, IV 109
also removed Mastixiodendron from the Cornaceae to the Rubiaceae,! com-
menting on its close relationship to Dorisia. Further examination of the
two specific entities involved demonstrates that the differences between
them are scarcely generic in nature. Those differences pointed out by Gil-
lespie deal merely with the comparative length ae position on the fruit of
the calyx-tube and appear to be of specific value o
In order to have the reduction of Dorisia to. ee re verified,
ample material of the two species involved — Dorisia flavida (Seem.) A. C.
Sm. from Fiji and Mastixiodendron pachycladon |“pachyclados”| (K.
Schum.) Melchior from New Guinea — was submitted to Dr. Charlotte G.
Nast for study. Her report states that: “Dorisia and Mastixiodendron
could be taken as congeneric. Their nodal anatomy (trilacunar), stomata
(the usual rubiaceous type), wood, and pollen are similar. The wood has
oblique-porous vessels, septate fibers, no wood parenchyma, and fairly short
heterogeneous rays. The tricolpate pollen grain has a fairly fine but dis-
tinctly reticulate exine. Mastixia Bl. is not related to Mastixiodendron.
The wood of the two genera is taal different, that of Mastixia having
scalariform vessels with many bars, diffuse-aggregate parenchyma, and
very long heterogeneous rays. The stomata of Mastixia are not of the
rubiaceous type, and the tricolpate pollen grain is very finely reticulate,
almost granular in appearance, the pores being smaller and the grooves
larger than those of the pollen grain of Mastixiodendron.”
In view of this evidence, and because of the lack of differentiating tax-
onomic characters, it appears that Dorisia may be incorporated in the older
genus. Mastixiodendron, as thus amplified, has a range extending from
Halmahera (see Merrill and Perry in Jour. Arnold Arb. 25: 205. 1944)
through New Guinea to Fiji. Its discovery in the Solomons and New
Hebrides is to be anticipated and would complete a very natural geograph-
ical distribution.
Mastixiodendron flavidum (Seem.) comb. nov.
Canthium flavidum Seem. FI. Vit. 132. 1866.
Plectronia flavida Benth. & Hook. f. ex Drake, Fl. Ins. Mar. Pac. 194. 1890.
Dorisia rarissima Gillespie in Hook. Ic. Pl. 32: pl. 3190. 1933
Dorisia flavida A. C. Sm. in Bishop Mus. Bull. 141: 140. 1936; Fosberg in Sargentia
This species is known from the Fijian islands of Vanua Levu, Rambi, and
Viti Levu, as mentioned in the above-cited publications. Gillespie men-
tions Horne 1132, and Fosberg Parham III, as pubescent-leaved individuals
possibly worthy of nomenclatural recognition. These two specimens, both
in fruit, appear to the present writer to represent an undescribed species.
Mastixiodendron pilosum sp. nov.
Arbor, ramulis leviter angulatis demum subteretibus cinereisque, sity
versus copiose puberulis; stipulis rigidis oblongo-lanceolatis 18-25 m
longis inconspicue puberulis mox caducis; petiolis subteretibus vel eotey
canaliculatis robustis (1-2 mm. diametro) 2—4 cm. longis dense puberulis;
1 Apparently Mastixiodendron was first referred to the Rubiaceae by Danser (in
Blumea 1: 69. 1934), in his study of the Cornaceae of the Netherlands Indies.
110 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
copioso albo.
Vanua Levu: Mbua: Between Mbua and Ndevoka, Mrs. H. B. R. Parham III
(A) (tree near creek; fruits orange). Fiji, without definite locality: Horne
1132 (GH, type). The Horne collection, the better of the two listed, was probably
also obtained on Vanua Levu, from which the bulk of his collection came.
The leaves of M. flavidum (Seem.) A. C. Sm. are entirely glabrous
beneath or very minutely puberulent on the costa and principal nerves,
whereas those of the new species are densely, conspicuously, and persist-
ently soft-pilose beneath with pale spreading hairs. The young branchlets,
stipules, inflorescence-branches, fruiting pedicels, mature calyces, and per-
sistent calyx-lobes of M. pilosum are similarly pubescent, the corresponding
parts in M. flavidum being glabrous. These differences appear to be of
specific value, and I doubt if such a well-defined species as M. flavidum
should be interpreted to include the pubescent form here described as new.
Both of the Fijian species differ from the New Guinean M. pachycladon
in their less robust habit, longer and more slender petioles, narrower leaf-
blades, and more nearly completely inferior fruits.
COMPOSITAE
Erigeron sumatrensis Retz. Obs. Bot. 5: 28. 1789.
Vv EvU: Parks s. n. (Bish); Tholo North: Nandarivatu, on open slopes,
Parks 20623 (Bish).
This widespread weed, often recorded under the later binomial of F.
linifolius Willd. (1804), has been reported from only Hawaii and Faster
Island in the Pacific region.
ARNOLD ARBORETUM,
Harvarp UNIVERSITY.
1945] BOIVIN, NOTES ON THALICTRUM 111
NOTES ON SOME CHINESE AND KOREAN SPECIES
OF THALICTRUM
BERNARD BOIVIN
With one plate
RECENT collections in China made under the auspices of the Arnold Arbo-
retum have resulted in the assembling of about 200 specimens of Thalic-
trum, mostly from Yunnan; these are now deposited in the Gray Herbarium.
The purpose of the present paper is to present descriptions of certain new
species found among this material and to discuss other noteworthy species.
When this work was first undertaken, in October, 1943, there was a nearly
complete lack of material for comparison in the Harvard herbaria, and I
was forced to depend upon the original descriptions in making identifica-
tions. Since then, however, at Dr. Merrill’s request, Sir William Wright
Smith, of the Royal Botanic Garden at Edinburgh, has very kindly sent us
thirteen packets containing authentic fragments of as many of Léveillé’s
species. Each packet is fully annotated and contains fairly large frag-
ments, so carefully selected that no trouble was experienced in identifying
them, except, of course, when the plants had been badly preserved or col-
lected too early.
I am deeply indebted to Sir William Wright Smith for his generosity in
sending these fragments, and also to the authorities of the Arnold Arbo-
retum and the Gray Herbarium for the specimens and facilities placed at
my disposal. All cited specimens, unless otherwise stated, are deposited
in the Gray Herbarium.
Subgenus Thalictrum (DC.) Reichenb.
§ Homothalictrum Boivin
Thalictrum Esquirolii H. Lév. & Vaniot in Bull. Acad. Int. Géogr. Bot. 17(210-211):
ii. 1907.
Part of the type material, consisting of a complete plant and an inflores-
cence, is in the Gray Herbarium; this is very good material in full bloom.
Most of the herbarium specimens which I have seen identified as this species
have been correctly named.
Thalictrum minus L. var. elz ey ete in Bull. Soc. Bot. Belg. 24: 202. 1885.
Thalictrum amplissimum H. . & Vaniot in Bull. Acad. Int. Géogr. Bot. 11: 51.
1902.
Thalictrum minus var. amplissimum H. Lév. & Vaniot ex H. Lév. Fl. Kouy-Tchéou
339. 1915.
A fragment of a syntype of 7. amplissimum is preserved in the Gray
Herbarium. This is a side branch of the inflorescence of a plant 2 feet high,
collected in flowering condition. Whether the actual type is at Edinburgh
112 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
or at the Académie Internationale de Géographie Botanique is not known,
nor whether the original material is made up of two different collections or
a single collection.
§ Leptostigma Boivin
bial sic oe Sieb. & Zucc. var. Se aeaiaaa (Franch.) Finet &
Gagnep. in Bull. Soc. Bot. France 50: 611
The se known from Szechuan afl Cae is pubescent throughout
and its anthers are about 1 mm. long, while the typical form is entirely
glabrous and has anthers about 1.5 mm. long. Specimens of the typical
form are at hand from Japan, Korea, Kiangsi, and Chekiang.
Thalictrum Atriplex Finet & Gagnep. in Bull. Soc. Bot. France 50: 613. pl. 19, B. 1903.
This species is closely related to T. baicalense Turcz., and it is probable
that the latter should be placed in § Leptostigma rather than in § Physo-
carpum. Indeed, 7’. baicalense is a more or less heterodox species.
Thalictrum cirrhosum H. Lév. in Rep. Nov. Sp. 7: 97. 1909.
A fragment of an isotype, preserved in the Gray Herbarium, proves to be
altogether different from any other specimen at hand. The whole plant is
pubescent except upon the upper surface of the leaves. All the hairs are
simple and short except those on the lower surfaces of the leaves, these
bearing numerous short branches and having stellate tips. This is a unique
type of pubescence in Thalictrum. The specific epithet cirrhosum is un-
doubtedly wrong, for the type has no cirrhi, nor have any other specimens
of the genus. But the Edinburgh isotype is labeled ‘‘cirrosum” in Léveillé’s
hand. This could be the adjectival form of “cirrus” and could well be used
to describe a plant having the habit of 7. trichopus Franch. or T. cincinna-
tum Boivin. However, as the fragment looks much more like 7. deciterna-
tum Boivin than like any other species, one is at a loss to understand
Léveillé’s choice of a specific epithet
Thalictrum deciternatum sp. nov. Pu. I, Fics. 4-7.
Planta omnino glabra, sed foliolis inferne interdum pubescentibus, caule,
ramis, petiolis petiolulisque pruinosis, 50-125 cm. alta, radicibus fibrosis,
haud stolonifera. Folia basilaria aetate florendi desunt. Folia caulinaria
10-30 cm. longa, sessilia in apice dilatationis petiolaris vel breviter petio-
lata, 6-10-ternata. Stipellae desunt. Foliola 3-10 mm. longa, bi-trilo-
bata, crassa, margine revoluto, superne dense viridia et nervis paullo
impressis, inferne glauca et nervis rugosis valde reticulatis. Foliolum
terminale basi cuneatum apice rotundo-truncatum, mucronatum. — Inflores-
centia paniculata ramosa copiosa paullulum foliosa. Pedicelli sub recepta-
culo recurvati. Sepala elliptica lutea ca. 4 mm. longa. Filamenta lutea
filiformia ca. 5 mm. longa. Antherae oblongo-lineares 3—3.5 mm. longae,
mucrone 0.1—0.3 mm. longo. Ovaria subsessilia 4-5. Stigma sessile 1—1.5
mm. longum anguste bialatum a sepalis recedit. Carpella immatura com-
pressa subsessilia, ventre basi cuneato, summo rotundo, nervo ventrali quam
dorsali paullulum convexiore. Floret junio julio et augusto
YuNNAN: Ad ee : mont. niveor. prope Lichiang, in dumetis, ad 1.25 m., alt. ca.
3200 m., junio 16, , C. Schneider 1805 (type); Lichiang Snow Range, comm
along mountain ae "August 27, 1937, T. T. Yi 15480; eastern slopes of Likiang
1945] BOIVIN, NOTES ON THALICTRUM 113
Snow Range, gulch leading to main peak, among rocks, fis. yellow, May 24, 1922, J. F.
Rock 3801; Likiang Snow Range, open pasture, plant 3 ft. high, fls. green-yellow, June
25, 1939, R. C. Ching 30272; Li-kiang ene grassy slope under forest, fls. greenish
white, alt. 3000 m., July 1935, C. W. Wang 70955; Wei-si Hsien, Yeh-chih, mountain
slope, fis. yellow, alt 3600 m., Aug. 1935, C. W. Wang 68341; Chi-ling Shan, Cheng-
rhe fls. light ae oe 1960 m., Aug. 25, 1939, H. Wang 41681; without detailed
data, T. T. Yu 9891
Thalictrum Delavayi Franch. in Bull. Soc. Bot. France 33: 367.
?Thalictrum ee dite bili H. Lév. in Bull. Acad. ee pre noe ie 297. 1902.
?Thalictrum Duclouxii H. Lév. in Rep. Nov. Sp. 7: 909.
Thalictrum grandisepalum recalls T. pee in its flowers, but it might
possibly be a different species. The type fragment at hand shows a plant
collected too early in bloom and very poorly preserved. A fragment of a
syntype of 7. Duclouxii is also at hand, but this was collected when the
flowers were still in bud. Consequently, it is impossible to identify with
certainty either of Léveillé’s species listed above.
Thalictrum Finetii sp. nov. PL. I, Fics. 1-3.
Planta 50-200 cm. alta, foliolis inferne et fructibus dense pubescentibus,
caule et petiolis puberulis. Pubescentia caulinaris fructuum et petiolorum
e pilis capitatis translucidis minutis, foliorum opacis minutis crassiusculis.
Radices fibrosae, nec planta stolonifera. Caulis et rami flexuosi. Folium
basilarium deest aetate florendi. Folia caulinaria regulariter 4-ternata et
fere omnia sessilia in apice dilatationis petiolaris. Petioluli arcuantes.
Stipellae desunt. Foliola membranacea ovata 5-15 mm. longa, trilobata
lobis crenatis. Inflorescentiae variabiles, aliae reductae, aliae amplae.
Pedicelli sub receptaculo recurvati et fere omnes ex axillis foliorum 1—3-
ternatorum. Sepala 4—5, elliptica, 3.5-4.5 mm. longa. Filamenta fili-
formia pallide lutea ca. 6 mm. longa. Antherae oblongo-lanceolatae ca.
2.5 mm, longae, acumine ca. 1 mm. longo. Ovaria 7-10 tigma 0.9—1.2
mm. longum haud alatum sed ventrale. Carpella matura BE Se ie
cata, stipite ca. 0.4 mm. longo, ventre semi-obovato ca. 4 mm, lon t 1.5
mm. lato, nervo dorsali recto, ventrali convexo et alis angustis ee
munito. Floret julio et augu
SzECHUAN: Mt. Omei, hillside, fls. white, alt. 2200 m., Aug. 1, 1938, K. N. Yin 117
(tYPE); Mt. Omei, shrubby flat, 3 ft. high, fls. white, alt. 3300 m., July-Aug. 1931,
F. T. Wang 23458; Lieng-ho-kou, grassland, fis. yellow, alt. 12000 ft., Aug. 1938, T. S.
Wen 563. YUNNAN: Mekon eye Divide, rocky places in mountains, casual,
15-20 inches high, fls. yellow, alt. , Aug. 11, 1938, T. T. Yi 22298; Chengkang,
Snow Range, common on grassy nae 2-3 ft. high, fls. white, alt. 3460 m., July 24,
tse- fa Pi-lo Shan, fis. yellow, alt. 4000 m., Aug. 18, 1934, H. T. Tsai 58014; Litiping,
between Likiang and Weihsi, in ies forests, 3-5 ft. high, Oct. 9, 1939, R. C. Ching
22072; Wei-si Hsien, Yeh-chih, common on mountain slope, alt. 3200 m., July 1935,
C. W. Wang 68033; north flank of Haba Snow Range, open pasture, 3 ft. high, fils.
white, Aug. on 1939, K. M. Feng 2100; Ta-li Hsien, on open slope, 3 ft. high, - pe
alt. 3500 m., July 30, 1933, H. T. Tsai 53943; without detailed data, T. 7. Yu
This species is dedicated to the botanist A. Finet, who ee a
collaboration with A. Gagnepain, a very good treatise on the eastern Asiatic
species of Thalictrum. The nearest relative of T. Finetii is T. platycarpum
Hook. f. & Thoms., of which a syntype is at hand. However, the new
species is much larger throughout, with longer anthers and with a narrower
114 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
fruit, which has less prominent ribs, a shorter stipe, and a well defined style.
Thalictrum Smithii sp. nov. Pt. I, Fics. 20-22.
Planta 40-150 cm. alta, plus minusve pubescens vel fere glabra, radicibus
fibrosis in locis tuberosis, tuberibus ovoideis. Folia caulinaria ca. 5-ternata.
Stipellae desunt. F ‘oliola membranacea ovata crenata, (0.5—)1(—1.5) cm.
longa. Inflorescen mpla paniculata ramosa ramis flexuosis et fere
nudis. Pedicelli pine eerie (3—)5(—15) mm. longi recti. Sepala —
dorso puberula ca. 2.5 mm. longa. Filamenta filiformia ca. 4 mm. lon
Antherae oblongo-lineares, ca. 2.5 mm. longae, apice obtusae. Ovaria 4— -
Stigma ventrale subsessile haud alatum, 0.7-1 mm. longum. Carpella
matura haud compressa costata sessilia, ventre obovoideo 1.2—1.6 mm.
longo et 0.8—1.2 mm. lato, nervis gibbosulis. Floret septembri.
S1kaNnc: Muli, Wachin, Shawan, common on margin of thickets, 1-2 bs high,
achenes black, alt. 3000 m., Oct. 11, 1937, T. T. Yui 14487 (TYPE). Yun Che.
tse-lo, in forest, fls. vallaw, alt. 3200 m., Sept. 7, 1934, H. T. Tsai Perey ig es ae
Hsien, on open slope, 5 ft. high, fls. vellowith, alt. 1500 m., Sept. 27, 1933, H. T. Tsa
54683; without detailed data, H. T. Tsai 57289.
This new species is dedicated to Sir William Wright Smith.
Thalictrum samariferum sp. nov. PL. I, Fires. 31, 32.
Planta omnino glabra 10-45 cm. alta. Radices fibrosae, interdum in
locis paullulum tuberosis. Stolones desunt. Folia ab internodii iis supe-
rantur, 5—6-ternata et sessilia in apice dilatationis petiolaris. Foliola cori-
acea, margine revoluta, minora, ca. 3 mm. longa, basi cuneata, apice trifida
vel saepius (1—)3(—5)-lobata. Stipulae adnatae integrae lanceolatae.
Stipellae desunt. Inflorescentia interdum simplex, saepius plus minusve
ramosa, semper foliosa. Pedunculi fere omnes ex axillis foliorum 2—5-
ternatorum. Pedicelli recurvati, vel, si recti, sub receptaculo valde reflexi.
Flores 5-25. Carpella matura ca. 10 in receptaculo. Stigma paullulum
1 1 ;
longus haud alatus nisi prope ventre. Venter carpelli maturi 5—7 mm.
longus et 4-5 mm. latus, obcordatus et laminaris bialatus, alis regularibus,
alia ventrali convexa, altera dorsali convexiore, et ambae 0.5—2 mm. latae
et membranaceae. Ner rvi carpelli maturi quatuor, quorum unus ventralis,
unus dorsalis et duo laterales, omnes vero paullulum rugosi. Flores mi ihi
ignotae sunt
sat A-tun-tze, mountain slope, fruit greenish white, alt. 2700 m., Sept. 1935,
C. W. Wang 70156 (tyPE) ; A-tun-tze, border of woods, fruit green, alt. 3000 m., July-
Aug. 1935, C. W. Wang 64885, 64772; without detailed data, T. T. Vii 12993. South-
eastern Tipet: Sacred mountain Kar-war-kar-boo, Tsa-wa-rung, on bare rocks at
foot of mountain, fruit light brown, alt. 3500 m. , Sept. 1 935, C. W. Wang 66468; Dier-
mai, Tsa-wa-rung, alt. 3200 m., Aug. 1935, C. W. Wang 65720; Gerda, Ree-su-la,
Tsa-wa-rung, on grassy slope, fruit nes alt. 3600 m., Aug. 1935, C. W. Wang 65913.
In habit this species closely resembles 7. elegans Wall., but it has a much
larger fruit, a usually simpler and more leafy inflorescence with longer
pedicels, and is always glabrous. Another specimen from Yunnan, 7. T.
Yu 6236, collected early in bloom, may also belong here.
Thalictrum trichopus Franch. in Bull. Soc. Bot. Rieei 33: 370. 1886.
Thalictrum Tenii H. Lévy. in Rep. Nov. Sp. 7:
A tragment of an isotype of 7. Tenii is er in the Gray Herbarium.
1945] BOIVIN, NOTES ON THALICTRUM ~ 115
Thalictrum Malai Hook. f. & Thoms. Fl. Ind. 1: 14. 1855. Pu. I, rics. 25-27.
Thalictrum virgatum var. stipitatum Franch. in Bull. Soc. tae ia aie 369. 1886.
Thalictrum verticillatum H. Lév. in Rep. Nov. Sp. 7: 97, 99.
Léveillé’s reduction of his own T. verticillatum to T. ae is correct,
judging from part of the type material of the former and an isotype of the
latter preserved in the Gray Herbarium. Franchet’s var. stipitatum is the
typical form of this species. Lecoyer’s drawing (in Bull. Soc. Bot. Belg.
24: pl. 4, fig. 10. 1885) does not represent T. virgatum.
hacia bie a nov. cai I, Fics. 23, 24.
foliolis ue pilis crassiusculis et opacis. Ovaria conspicue stipitata.
Stig _1mm. longum. Carpella matura compressa subcostata, stipite
32. pe m. longo, ventre 2.2—2.8 mm. longo et 1.5—-1.7 mm. lato, nervo
dorsali palm convexo, ventrali multo convexiore quam dorsali.
-SIKA , Kulu, rocky places in thickets, si 2-3 ft. high, achenes pale
brown, alt, ere Sept. 14, 1937, 7. T. Yu 14273 (TYPE
§ Erythrandra Boivin
Thalictrum alge BI. Bijdr. Fl. Ned. Ind. 2. 1825.
Thalictrum Argyi H. Lév. in Bull. Herb. Boiss. II. 6: 504. 1906.
A fragment of an isotype of J. Argyi is preserved in the Gray Herbarium.
Thalictrum ramosum sp. nov. Pv. I, Fics. 12-15.
Planta diffusa e basi ramosa viridula glabra debilis, 25-40 cm. alta. Folia
basilaria adsunt, vers. 20-30 cm. longa. Stipellae desunt. Foliola
membranacea, (1—)1.5(—2) cm. longa, saepius obovata et apice crenata,
nervis inferne rugosulis. Flores subcorymbosi. Biter orbicularia 2
mm. longa. Filamenta alba clavata 1.5—2.2 mm. long Antherae ellip-
soideae, ca. 0.7 mm. longae, apice rotundae. corn een parum
longiora, 8-12. Stigma ventrale haud alatum ca. 0.3 mm. longum. Stylus
cum stigmate ca. 0.5 mm. longus. Carpella matura brunnea sessilia fusi-
formia haud compressa nec costata, ventre lanceolato ca. 4.5 mm. longo et
1.2 mm. lato, nervo ventrali paullum convexiore quam dorsali. In eadem
planta nonnunquam et flores et carpella matura inveniuntur quia flores ad
anthesim veniunt in successione a mense aprili.
SzECHUAN: South of Kuan Hsien, along ditch, fls. reddish, alt. 850-950 m., Apr.
1930, F. T. Wang 20378 (tyPE); Mt. Omei, under shady rocks, fls. greenish Ghite, at
900 m., Apr. 15, 1932, T. T. Yu 274; Chengtu, Mar. 15, 1937, S. S. Chien 5880.
Thalictrum Rockii sp. nov. PL. I, Fics. 28-30.
Planta 1.5 m. alta, omnino pubescens nisi in caule inferiore e pilis uni-
seriatis translucidis aetate florendi, aetate maturandi glabra nisi in foliolis
ternata estipellata, foliolis 0.8-1.5 cm. longis, trilobatis, lobis saepius
crenatis. Inflorescentia copiosa ramosissima paniculata. Sepala ovata ca.
3 mm. longa. Filamenta clavata 5.5-7.5 mm. longa. Antherae oblongo-
lanceolatae ca. 1 mm. longae. Ovaria ca. 4, ventre compresso breviore
quam stipite, stigmate ca. 0.5 mm. longo breviore quam stylo libero. Car-
pella matura pendula ae costata laminaria, stipite tenui 3-3.5 mm. longo,
116 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
ventre semi-obovato 5—5.5 mm. longo, 2.8-3.2 mm. lato, nervo ventrali
multo convexiore quam dorsali. Floret junio julioque.
Upper Tebbu (T’ieh-pu) country, below Shihmen, Tsaluku valley, in
forests of spruces, willows, etc., fls. creamy white, alt. 11000 ft., July-Aug. 1925, J. F.
Rock 13054 (type); T’ao River basin, valley of Tayiiku, grassy slopes, fls. greenish,
alt. 8500 ft., July 1925, J. F. Rock 12835. Tstncuat (Ch’ing-hai): Ba valley, under
willows, fls. greenish yellow, alt. 9900 ft., June 23, 1926, J. F. Rock 14271
In the herbarium of the New York Botanical Garden there is a fruiting
specimen of this species, collected by W. Y. Hsia (no. 2400) at Ling-shan-
kou in Cho-lu Hsien.
Thalictrum Wangii sp. nov. PL. I, rics. 8-11.
Planta 30-40 cm. alta, omnino dense pubescens pilis capitatis translu-
cidis. Folia caulinaria 3-ternata, ea inflorescentiae 1—2-ternata. Stipulae
semi-ovatae ca. 1 mm. longae. Stipellae desunt. Foliola membrancaea
ca. 8 mm. longa, fere orbicularia, apice trilobata, lobis crenatis. Inflores-
centia 4—8-flora. Pedicelli 1-2 cm. longi, sub receptaculo plus minusve
recurvati, omnes ex axillis foliorum 1—2-ternatorum. Flores albi. Sepala
elliptica ca. 5 mm. longa. Filamenta alba apice clavata ca. 4.5 mm. longa.
Antherae oblongo-lineares, albae, ca. 1.5 mm. longae, apice rotundatae.
Ovaria fusiformia 4-6. Stylus cum stigmate ca. 1.8 mm. longus. Stigma
anguste bialatum ca. 0.6 mm. longum. Carpella matura ignota. Floret
julio,
YUNNAN: Li-kiang Hsien, in woods, fls. yellowish white, alt. 2700 m., July 1935,
C. W. Wang 71546 (type). Southeastern Trpet: Dzer-nar, Tsa-wa-rung, dry slope
on border of woods, alt. 3000 m., Sept. 1935, C. W. Wang 66523.
The closest relative of this species seems to be 7. Fargesii Franch.
§ Physocarpum DC.
Thalictrum coreanum H. Lév. in Bull. Acad. Int. Géogr. Bot. 11: 297. 1902.
In my recently published monographic study of T'halictrum, I expressed
doubt (in Rhodora 46: 368. 1944) as to whether 7. coreanum and T. ichan-
gense Lecoyer (ex Oliv. in Hook. Ic. Pl. 18: ¢. 1765. 1888) were distinct
species. At the time this original paper was prepared, we had very good
and abundant material of the latter but only one poor specimen of the
former. At present, the situation is reversed, and I have available very
good fragments of 7. coreanum but only one poor specimen of JT. ichan-
gense. From the material at hand, and from my recollection of that which
I have previously examined, these two species are undoubtedly distinct.
Thalictrum coreanum has a compact subcorymbose inflorescence and a fruit
which is sessile or nearly so, with a body about three times longer than
broad. Its leaflets are orbicular, 9 in number to each basal leaf, and very
leathery at maturity. Thalictrum ichangense has a somewhat diffuse in-
florescence with the flowers more or less racemose on the branches. Its
fruit is conspicuously stipitate, with a body about five times longer than
broad. The leaflets of its basal leaves are more or less triangular, with a
rounded base, and usually are 3 or 5, or very rarely 9, per leaf. At ma-
turity the leaflets are firm, strongly bicolored, and always dull beneath.
Thalictrum coreanum has a much stouter appearance throughout.
1945 | BOIVIN, NOTES ON THALICTRUM 117
Thalictrum tuberiferum ps in Bull. Acad. Sci. St. Pétersb. 22: ey 1876.
Thalictrum Fauriei H. Lév. in Rep. Nov. Sp. 7: 100. 1909; nec. H. Lév. & Vaniot in
Bull. Soc. Bot. France 53: 388. 1906; nec Hayate: in Jour. Coll. “Tokyo 22275
6
Thalictrum punctatum H. Lév. in Rep. Sp. Nov. 10: 376. 1912.
Fragments of the type of T. Faurier H. Lév. and of the syntype of T.
punctatum are available; both apparently belong to T. tuberiferum Maxim.
§ Tripetrium DC.
Thalictrum aquilegifolium L. Sp. Pl. 547. 1753.
Thalictrum Taqueti H. Lév. in Rep. Nov. Sp. 7: 339 (nec ite 1909.
Thalictrum Dunnianum H. Lév. in Rep. Sp. Nov. 8: 549
Fragments of type material of Léveillé’s species are eorere in the Gray
Herbarium. Thalictrum Dunnianum is merely a renaming of the T.
Taqueti based on T'aquet 508, this specific epithet having been used twice
by Léveillé
Subgenus Lecoyerium Boivin
§ Cincinneria Boivin
Thalictrum Mairei H. Lév. in Rep. Nov. Sp. 7: 339. 1909. PL. I, Fics. 16-19.
Planta robusta omnino glabra 50-150 cm. alta. Folia basilaria petiolata,
petiolo basi valde dilatato. _ Folia caulinaria 4—5-ternata petiolo brevi.
m. longa, m
ramosa et foliosa. Pedicelli robusti 1-3 cm. longi, sub receptaculo paullo
incurvati, fere omnes ex axillis foliorum 2—3-ternatorum. Sepala fere
orbicularia 4. Ovaria ca. 10. Stigma conspicue bialatum lanceolatum
sepala excedens, ca. 3 mm. longum et 0.5—-1 mm. latum. Carpella matura
costata fere haud ale valde reflexa, stipite crasso ca. 1 mm. longo,
ventre lanceolato 6-7.5 mm. longo et 1.8—-2 mm. lato. Stigma persistens
circinatum. Floret maio et julio.
SZECHUAN: Inter Oti et Ouentin, in dumetis, fl. roseo-lilacini, alt. ca. 2500 m., junio
3, 1914, C. Schneider 1170; inter Kuali et Molien, planta ad 1 m., alt. ca. 3200 m., majo
1914, C. Schneider 1382. S1KANG: Muli, Kulu, rie oe us thickets, Chea:
5 ft. high, achenes pale brown, alt. 3100 m., Sept. 22, 1937, T. Yu 14335. Yun-
Likiang, Ah-nar-koo, near Shik oo, in open aN e, fls. canary, June 1,
4
>
vA
a:
1939, R. C. Ching 30228; eastern slopes of Likiang Snow Range, Pai-shui Ho, fis. dull
oe purple, May 17, 1922, J. F. Rock 3572; without detailed data, T. T. Yu 5700,
ais species evidently belongs to the Section Cincinneria, along with T.
macrostigma Finet & Gagnep. ‘The closest relative of these two species is
T. macrocarpum Gren., from the French Pyrénées. At first I took the
species described above to be new, but, through a fragment received from
Edinburgh, I was able to ascertain its identity with 7. Mairez. However,
the above description, based on the cited specimens, does not seem super-
fluous.
118 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
EXPLANATION OF THE PLATE
All drawings are x 6 except Fig. 4, which is x 2, and Fig. 19, which is x 1.
Fics. 1-3. Thalictrum Finetii Boivin: stamen and ovary, Yin 117; fruit, Ching 22072.
sepal, stamen, ovary, and young fruit, Wang 71546. Fics. 12-15. Thalictrum
ramosum Boivin: stamen, sepal, and ovary, Wang 20378; fruit, Yu 274. Fics. 16-19
1
54683; fruit, Yai 14487. Fics. 23, 24. Thalictrum Yui Boivin: ovary and fruit, Vi
14273, Fics. 25-27. Thalictrum virgatum Hook. f. & Thoms.: stamen and ovary,
J.D. Hooker, Sikkim; fruit, Yu 14544. Fics. 28-30. Thalictrum Rockii Boivin: fruit,
Hsia 2400; stamen and ovary, Rock 13054. Fics. 31, 32. Thalictrum samariferum
Boivin: ovary and fruit, Wang 70156
Gray HERBARIUM,
Harvarp UNIVERSITY.
Jour. Arnoip Ars. VoL. XXVI
1945] LI, NOTES ON THE FLORA OF INDO-CHINA 119
FURTHER NOTES ON THE FLORA OF INDO-CHINA1!
Hut-Lin Li
Tuts brief article supplements a previous paper (Jour. Arnold Arb.
24: 362-374. 1943), and in it three new species and two new varieties are
described, one new name is proposed, and four previously described species
are for the first time recorded from Indo-China. The genera Bredia and
Stapfiophyton are new to that country. The types of the new forms herein
described are deposited in the herbarium of the Arnold Arboretum.
oe
Elaeocarpus hainanensis Oliv. in Hook. ge t. 2462, 1896
Elaeocarpus linearifolius Knuth, ee. cae 7. 49: 66. 1940, syn, nov.
ND nam, Tourane and eae J.& M.S. Clemens 3484 (isotype of
E. OT May- Tine. 1927, shrub or small tree, river-margin, flowers dull yellow.
This Indo-Chinese plant, described by Knuth as E. linearifolius, is mani-
festly the same as the Hainan species. This is one of the many species
common to both regions.
THEACEAE
Cleyera japonica Thunberg var. ronmInehsss var. nov.
A typo speciei differt foliis obovatis, 4-6 cm. longis, 2.5-3.5 cm. latis,
obtusis, fructibus longe pedicellatis, pedicellis ad 3 cm. longis.
INbDo- Cu INA: Tonkin, Ha-coi, Chuk-phai, Taai Wong Mo Shan and vicinity, W.
T. Tsang 26987 (TyPE), Oct. 16-22, 1936, 29371, July 14-31, 1939, a small tree 20 ft.
high, fairly common in thickets, fruits yellow or black.
Among the different varieties of the species as treated by Kobuski (Jour.
Arnold Arb. 18: 118-129. 1937), this new variety is nearest var. Morii
(Yamamoto) Masamune, but it may be readi'y distinguished by the smaller
leaves with obtuse apices and by the much longer fruiting pedicels.
FLACOURTIACEAE
sd ela annamica sp. nov.
Arbor 8-10 m. alta, ramulis teretibus puberulis vel subglabris; foliis
Series oblongo- ellipticis, 10-14 cm. longis, 3.5—4.5 cm. latis, longe
acuminatis (acumine 2—2.5 cm. longo), basi acutis, margine integris, supra
in sicco atro-brunneis, glabris, subtus pallidioribus parce puberulis, venis
lateralibus utrinsecus 6 vel 7, supra distinctis, subtus elevatis, valde arcu-
atim adscendentibus, rete venularum utrinque subconspicuo; petiolo 1—2
cm. longo, puberulo vel glabrescente; floribus ignotis; fructu axillari soli-
tario oblongo-ovoideo, circiter 3 cm. longo et 2 cm. lato, dense fulvo-
oe)
at
1 Prepared with partial support of a grant from the Penrose Fund, American Philo-
sophical Society, to Dr. E. D. Merrill, to assist him in working up the accumulated col-
lections of Chinese and Indo-Chinese botanical material in the herbarium of the Arnold
Arboretum
120 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
velutino, pericarpio 0.5 mm. crasso, semine solitario, ovoideo, circiter 1.5
cm. longo et 1.3 cm. lato; pedicello 7 mm. longo, crasso.
Inpo-Cu1na: Annam, 12 kilometers north of Dankia, Lang-Biang, E. Poilan
18660 (TYPE), Jan. 13, 1931, a tree 8-10 m. high, on slightly rocky argillaceous soil, in
forests, alt. 1200-1500 m.; the fruit has a strong odor.
In the absence of Abwane the species cannot be placed in the proper sec-
tion of the genus. It is strongly characterized by the one-celled, one-seeded
fruit. Prof. I. W. Bailey, who examined the structure of the wood, reports
that in all respects it is that of a Hydnocarpus.
MELASTOMATACEAE
Bredia violacea sp. nov.
Frutex parvus, ramis nigienten hirsutis; ~_ in paribus aequalibus, sub-
Frei aaah oblongo-ovatis, circiter 9-10 ¢ n. longis, 5.5—6 cm. latis, acum-
inatis, basi distincte cordatis, coe rviis, pine hirsutis, supra sparse
setosis, subtus leviter puberul is, venis venulisque supra subconspicuis,
subtus distinctis; petiolis circiter 5 cm. longis, sparse hirsutis; inflorescentiis
terminalibus cymoso-paniculatis, circiter 15 cm. longis, leviter puberulis et
sparse longe hirsutis, pilis saepissime glandulosis, pedunculis 8-9 cm.
longis, pedicellis circiter 1 cm. longis, violaceis, bracteolis minutis, vix 1
mm, longis; calycis tubo 5 mm. longo, lobis 4, late triangulari-ovatis, 3 mm.
longis, 3.5 mm. latis, acutis vel subrotundatis, leviter puberulis et sparse
longis, antheris linearibus, leviter curvatis, olivaceis, circiter 8 mm. longis,
longe acuminatis, connectivo sub theca antice tuberculato, cnet n breviter
calcarato, stylo circiter 1.1 cm. longo, olivaceo.
InpO-CHINA: nkin, Pas Ho Yung Shan and vicinity, W. T. Tsang 30751
(TYPE), Oct. 13- Rs 22, 19
In general appearance, this species closely resembles Bredia tuberculata
(Guillaumin) Diels, but it differs, among other characters, in its cymose-
paniculate instead of umbellate inflorescences, In the inflorescence char-
acter, it belongs to the group of Bredia hirsuta Blume. The new species is
characterized by its broad distinctly violet calyx-lobes and its large violet
anthers. The genus is new to Indo-China
sae eels peperomiaefolium (Oliv.) Li, Jour. Arnold ‘ere - 29. 1944.
a peperomiaefolia Oliv. in Hook. Ic. Pl. 19: t. 1814.
iat salads Stapf, Ann. Bot. 6: i a
Inpo-Cuina: Tonkin, Ha-Coi, Taai Wong Mo Shan, Chan Uk Village near
Chuk-phai, W. T. Tsang 20029, May 1-10, 1939, an i fairly common in thickets, in
dry sandy soil, fruit brown. The species has previously been known only from
gtung.
This is an acaulescent plant which closely matches the syntype (C. Ford
336, photo. in herb. New York Botanical saga except that the leaves
are somewhat broader and less acute. The Indo-Chinese specimen is in
fruit, while the type is from a flowering plant.
Blastus membranifolius sp. nov. § Desmoblastus
Frutex, ramis junioribus minute subalbide tomentellis; foliis inaequalibus
1945] LI, NOTES ON THE FLORA OF INDO-CHINA 121
vel aequalibus, tenuiter membranaceis, longe petiolatis, oblongis, 10-12
cm. longis, 3.5—6 cm. latis, longe acuminatis, basi subrotundatis vel rotund-
atis, utrinque et margins setulosis, nervis primariis 3 marginalibus duobus
gracilioribus additis, venis transversalibus supra inconspicuis, subtus prom-
inentibus; petiolo 1.5-7 cm. longo, subalbide tomentello; floribus axil-
laribus, solitariis vel 2- vel 3-fasciculatis, pedunculis circiter 7 mm. longis,
tomentellis; calycibus obconico-quadrangularibus, extus tomentellis, 1.5
mm. ae ‘lobis brevibus; petalis rhomboideis, 2-3 mm. longis; staminibus
4, filamentis 2-3 mm. ongis, antheris 1.5—2 mm. longis, apice rotundatis,
pea = theca calloso.
Inpo-Cui1na: Tonkin, Dam-ha, cee Wong Mo Shan, Lung Wan Village, W. T.
Tsang 30112 (TYPE), May 18-July 5,
The new species is allied to ae tenuifolius Diels and B. setulosus
Diels, differing from both in the leaves being setulose on both surfaces and
in the longer pedicels.
pon ee lac Ssapeih Hook. & Arn. Bot. Beechey Voy. 186. 1833; Merr. Lingnan
Sci Uae 934
INpO- ok eee Tien-yen, Ho Yung Shan and ie cried W. T. Tsang 30643,
Oct. 13-Nov. 22, 1940. Kwangtung, Hainan; new to Indo-Chin
Memecylon ligustrifolium Champ. ex Benth. in Hook. Jour. Bot. Kew Gard. Miscel.
4: 117. 1852.
eee Cutna: Tonkin, northeast of Mon-cay, Pac-si and vicinity, W. T. Tsang
950, Oct. 1-8, 1936, a shrub 10 ft. high, fairly common in thickets, in dry ay pis
tie black; Dam-ha, Sai Wong Mo Shan, Lomg Ngong Village, W. T. Tsang 30361
July 18-Sept. 9, 1940; Tien-yen, Ho Yung Shan and vicinity, W. T. Tsang 30654,
Oct. 13-Nov. 22, 1940. Kwangtung, Kwangsi; new to Indo-China.
Memecylon coacerviflorum nom. no
Memecylon confertiflorum Merr. seks Arnold Arb. 19: 58. 1938, non Cogn. 1891.
Inpo-CuHInA: Southern Annam.
VERBENACEAE
Tsoongia axillariflora Merr. var. trifoliolata var. nov.
A typo speciei differt foliis plerumque trifoliolatis.
Inpo-CuInA: Tonkin, Dam-ha, - Wong Mo Shan, oe Wan Village, W. T.
Tsang 30135 (TYPE), May 18-July 5, 1940. Kwanctunc: Shih Wan Tai Shan, Foo
Lung, H. Y. Liang 69710, July 16, oe shrubby, scandent, in open forests, flowers
yellow.
Callicarpa formosana Rolfe, Jour. Bot. ea 1882.
Inpo-Cu1na: Tonkin, Ha-coi, Chuk-phai, Taai Wong Mo Shan, Shuei Mei Vil-
lage, W. T. Tsang 29378, July 14-31, 1939, a irre 7 ft. high, fairly common, es in
aod soil among scattered shrubs, flowers lavender; Dam-ha, Sai Wong Mo Shan,
Lung Wan Village, W. T. Tsang 29947, May 18- ee 5, 1940. Formosa, the Philip-
pines, eastern and southern China; new to Indo-Chin
ARNOLD ARBORETUM,
HarvaArD UNIVERSITY.
JOURNAL
OF THE
ARNOLD ARBORETUM
VoL. XXVI APRIL, 1945 NuMBER 2
A TAXONOMIC REVIEW OF TROCHODENDRON AND
TETRACENTRON
A. C. SMITH
With one text-figure
INLRODUCTLTION
CONTINUATION of the studies of certain woody families of Ranales
undertaken by Prof. Bailey, Dr. Nast, and the writer, and published in
recent volumes of this Journal, we propose next to consider a group of
eastern Asiatic genera which have, from time to time, been associated with
one another in the same family, subfamily, or tribe. These genera are
Trochodendron Sieb. & Zucc. (1838), Euptelea Sieb. & Zucc. (1841).
Cercidiphyllum Sieb. & Zucc. (1846), Tetracentron Oliv. (1889), and
Eucommia Oliv. (1890). That these genera, all of which are small and
some of which are monotypic, have aroused a high degree of interest among
taxonomists, phylogenists, morphologists, anatomists, and horticulturists
during the last century is demonstrated by the fact that we have examined,
in connection with our studies, more than 250 separate papers discussing
one or more of them. Many of these papers are floristic or horticultural
notes, others are standard works of a taxonomic nature, but not a few are
detailed morphological and anatomical studies. Our examination of past
work has been as exhaustive as possible, in order that we might become
familiar with the many and often conflicting opinions concerning the phy-
logeny and morphology of the genera concerned. These opinions have
ranged from that of van Tieghem (26), who places each of the five genera
in a separate family, to those of Finet and Gagnepain (5), who join all the
genera in a single tribe of the Magnoliaceae, Oliver (23), who joins them
all in the Trochodendraceae, and Hallier (8), who places them all in the
Hamamelidaceae. Between these extreme views are to be found all shades
of opinion, and the classification of these five genera remains an enigma to
most students.
As a result of our study of ample material of the groups in question —
material doubtless far exceeding in quantity and quality any available to
124 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
past students, largely because of the extensive Asiatic herbarium of the
Arnold Arboretum — we are inclined to support the view of van Tieghem
and to erect separate families for the five genera. This course may appear
radical to conservative taxonomists, who are perhaps inclined to overlook
the fact that, in dealing with very ancient and primitive groups such as the
Ranales, a sound classification cannot be attained by the use of traditional
comprehensive family limits. Our criteria for family delimitation in the
Ranales, we believe, are as sound and conservative as those in current usage
by students of other orders; that the application of these criteria results in
the establishment of numerous small, or even monotypic, families in the
Ranales reflects upon the extraordinary diversity and great age of the group,
and not upon an exceptionally radical viewpoint on our part. At any rate,
in placing Trochodendron, Euptelea, Cercidiphyllum, Tetracentron, and
Eucommia in separate families we are proposing nothing new. Such recent
phvlogenists as Hutchinson (19), Wettstein (30), and Diels (4) have
arranged the genera in question in either three or four families. Our prin-
cipal disagreement with these students is the separation of Tetracentron
from the Magnoliaceae and of Euptelea from the Trochodendraceae (the
latter course also proposed by Wettstein). As a matter of fact, Cercidi-
phyllum and Eucommia, according to many modern treatments, are to be
removed from the Ranales altogether and placed as relatives of the Hama-
melidaceae and Urticaceae respectively.
The present paper is planned to redescribe the monotypic genera Trocho-
dendron and Tetracentron, to give reasonably full citations to the literature
pertaining to these genera, to list the specimens available in the larger
American herbaria, and briefly to discuss the inter-relationships of the two
groups. In the following paper in this Journal, Prof. Bailey and Dr. Nast
initiate a series designed to discuss the morphological and anatomical details
of Trochodendron and Tetracentron. In the course of this study we have
fortunately been able to examine material from the following herbaria,
hereafter designated by the parenthetical letters: Arnold Arboretum (A),
Chicago Natural History Museum (Ch), Gray Herbarium (GH), Missouri
Botanical Garden (M), New York Botanical Garden (NY), University of
California (UC), and U. S. National Herbarium (US). The privilege of
borrowing this material is greatly appreciated.
Perhaps the best way to outline the principal studies on the five genera
mentioned above is to give a brief chronological sketch of past work; those
treatments which appear to be of especial importance in an understanding
of the groups are discussed in the following paragraphs.
HISTORICAL SKETCH
Until the nineteenth century was well advanced, Japan remained an
unknown land to European botanists. For that reason many of the con-
spicuous and widely cultivated trees of that country received no botanical
names until the publication of the important Flora Japonica by Siebold and
Zuccarini (1826-1870). In this publication, in 1838, Trochodendron
aralioides was described and well illustrated, being placed in the ‘“Winter-
1945 | SMITH, TROCHODENDRON AND TETRACENTRON 125
aneae” of R. Brown. In 1840 Junghuhn (in Tijdschr. Nat. Gesch. Phys.
7: 308) independently described Gymnanthus paradoxus as a Japanese
plant of the Magnoliaceae, related to [dlicium., The identity of his plant
with Trochodendron aralioides, apparently first pointed out by Endlicher
(Gen. Pl. Suppl. 2: 73. 1842), has been obvious to all subsequent students.
Euptelea was founded in 1841 by Siebold and Zuccarini (Fl. Jap. 1:
133), with one species, E. polyandra, as a new genus of the Ulmaceae.
Cercidiphyllum, proposed without a specific epithet by the same authors in
1846 (in Abh. Bayer. Akad. Wiss. Math. Phys. Cl. 4[3]: 238), was not
related by them, their specimen having been in fruit only. The specific
epithet japonicum was added to Cercidiphyllum, and accredited to Siebold
and Zuccarini, by Hoffmann and Schultes in 1852 (in Jour. Asiatique IV.
202 262):
The nomenclators of the period preceding 1860 — namely Endlicher,
Steudel, Walpers, Meisner, Dietrich, and Lindley — agreed in referring
Trochodendron to the Magnoliaceae as a relative of J/licium, in placing
Euptelea in the Ulmaceae, and in ignoring Cercidiphyllum.
In 1858 Miers (22) published a study of the Winteraceae, pointing out
that Trochodendron is not a relative of the group but more nearly ap-
proaches the Ternstroemiaceae — a suggestion which was not taken seri-
ously by subsequent students. Bentham and Hooker, in their Genera
Plantarum (1: 17. 1862), unwisely stated that Trochodendron is“... Ara-
liacea anomala ovario subsupero,” for which they were rebuked by Seemann
in his revision of the Hederaceae (in Jour. Bot. 2: 238. 1864). For Trocho-
dendron and Euptelea, Seemann proposed the new “order” Trochodendreae,
relating it to the Wintereae, Ranunculaceae, and Magnoliaceae. Seemann
was thus the first to suggest the isolated nature of Trochodendron, although
he is not usually recognized as the author of the family Trochodendraceae
because of his spelling of the name.
Hooker and Thomson (16), in 1864, in a short but important study of
Euptelea, agreed with Seemann in removing it from the Ulmaceae. They
concluded to place the genus in the Magnoliaceae, ‘“‘section’”’ Wintereae,
noting its highly anomalous nature and stating that it might possibly rep-
resent a distinct family. Hooker and Thon.son first called attention to
the existence of this group of plants outside of Japan, by proposing the new
species Euptelea pleiosperma from northeastern India.
Also in 1864, Eichler (2) published a very important anatomical study
of Drimys and Trochodendron, concluding that the latter genus belongs in
the vicinity of the Winteraceae. The following year Eichler (3) empha-
sized his conclusions and his agreement with Seemann in a supplementary
paper.
Bentham and Hooker profited from the work of Seemann and Eichler,
and in a supplement to the first volume of Genera Plantarum (1867) they
erected the tribe Trochodendreae in the Magnoliaceae, for Trochodendron
and Euptelea. The same position was indicated for the two genera by
Baillon (Hist. Pl. 1: 191. 1868-69), under the name Magnoliaceae IV.
[series?] ‘“Eupteleeae.”
126 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
The genus Cercidiphvllum, after a long period of neglect, was again
brought into the discussion by Baillon (in Adansonia 10: 132. 1871) in
connection with his notes on the Hamamelidaceae; his noncommital ac-
count also suggested that the genus is related to the Saxifragaceae, series
“Cunoniées.””. Maximowicz (in Mél. Biol. 8: 142. 1872) was the first to
associate Cercidiphyllum with Trochodendron and Euptelea in the Mag-
noliaceae. Maximowicz also proposed a second species of Cercidiphyilum
and divided Trochodendron aralioides into two varieties, one based upon
his own inadequately published 7. longifolium. Franchet and Savatier, in
1873, in their enumeration of Japanese plants, followed Maximowicz in
referring all three genera to the Magnoliaceae.
e family Trochodendraceae, in its modern spelling, was proposed by
bea in 1888 (24) to include Cercidiphyllum, Euptelea, and Trochoden-
dron,; in a supplement to the same work in 1891 Prantl further extended
the family to include Tetracentron. The last genus was described by
Oliver in 1889 (in Hook. Ic. Pl. 19: pl. 1892), on the basis of Henry’s col-
lections from Hupeh, and was placed in the Magnoliaceae, tribe Trocho-
dendreae. In the following year, Oliver described the monotypic Eucom-
mia (in Hook. Ic. Pl. 20: pl. 1950. 1890), also based on Henry’s plants
from Hupeh, without referring it to a family.
Toward the end of the nineteenth century, some of the genera under
discussion were introduced into Europe and North America for cultivation,
especially Cercidiphyllum, a striking tree which is now to be seen in many
parks and botanical gardens. Eucommia attracted attention for the gutta-
percha content of its bark. During the following years the remaining
genera were also brought into cultivation to a more limited extent, largely
through the efforts of Sargent at the Arnold Arboretum.
By far the most important morphological discussion of these plants up to
that time was Harms’ study of the Trochodendraceae in 1897 (10). Harms
pointed out that the wood of Tetracentron is vesselless, like that of Trocho-
dendron and Drimys. He retained Tetracentron in the Magnoliaceae with
the explanation that this family, which he interpreted to include such an
anomalous genus as Drimys, is not unduly expanded by the inclusion of
Tetracentron, The remaining four genera were placed by Harms in the
Trochodendraceae — the Eupteleoideae for Cercidiphyllum, Euptelea, and
Eucommia, and the Trochodendroideae for Trochodendron. While one
cannot agree with Harms’ disposal of these genera, it should be pointed
out that his subsequent studies caused him to modify his opinions; his
morphological and anatomical work on Trochodendron and its “allies” is
of the highest quality and his several papers are indispensable for a know]l-
edge of these plants.
A work of even greater import was van Tieghem’s study of the “Homo-
xylées,” published in 1900 (26). It is unfortunate that van Tieghem’s pres-
entation was so untraditional from a taxonomic point of view, for had his
conclusions been presented in a more orthodox manner his treatment surely
would have received the attention it merited from his contemporaries. Rec-
1945 ] SMITH, TROCHODENDRON AND TETRACENTRON 127
ognizing that the three groups typified by Trochodendron, Tetracentron, and
Drimys stand apart from all other dicotyledons in their vesselless wood, van
Tieghem proposed for them the subclass Homoxylees, as opposed to the
Hétéroxylées, including all the other dicotyledons. His Homoxyleées are di-
vided into three families, the Trochodendracées, Tetracentracées, and Dri-
mytacées (Winteraceae of other authors). With some emphasis, van Tieg-
hem pointed out that these families could not be associated with the genera
Eucommia, Cercidiphyllum, and Euptelea, for which he proposed the family
names Eucommiacées, Cercidiphyllacées, and Euptéléacées respectively.
While van Tieghem’s work on these genera leaves much to be desired, his
conclusions appear to be far sounder than those of his contemporaries. An
amusing commentary of van Tieghem’s perhaps deserves to be quoted in
full (26: 261): “Il [le petit groupe des Homoxylées| se compose, pour le
moment, des trois genres Trochodendre, Drimyte et Tétracentre. Malgré
leur caractére si remarquable, qu’on vient de rappeler, ces trois genres ont
été, et sont encore aujourd’hui classés tous purement et simplement dans
la famille des Magnoliacées, ou dans son voisinage immédiat. Quand on
étudie cette famille, on se borne a dire a leur sujet: ‘Chose curieuse, il y a
tel et tel genre de Magnoliacées, ott le bois secondaire est homogene, dif-
férent de celui de toutes les autres Dicotylédones et semblable a celui de
la presque totalité des Gymnospermes.’ Et puis, c’est tout. L’idee ne vient
a Vesprit de personne, que précisément ces genres singuliers ne sont peut-
étre pas des Magnoliacées, et qu’en tout cas, il y a lieu d’y regarder de plus
res. Et si lPidée n’en vient pas, c’est sans doute parce que l’on admet
implicitement que d’avoir telle ou telle sorte de bois secondaire, c'est chose
sans aucune importance pour la Classification des plantes.”
With van Tieghem’s publication, what may perhaps be termed the ex-
ploratory phase of the study of Trochodendron and its “allies” came to a
close. In the years since 1900 many students have given attention to the
group, and those interested in the larger aspects of family relationships
have proposed their own solutions. The principal detailed studies are
here mentioned briefly.
Solereder, in 1900 (25), published an important detailed morphological
study of Cercidiphyllum, pointing out that its floral characters do not seem
to be those of the Ranales; he referred the genus, and also Eucommia, to
the Hamamelidaceae. Wagner (28) presented in 1903 a study of the
morphology of the branchlets and inflorescence of Trochodendron. Also in
1903, Hallier (6) discussed the relationships of the five genera under con-
sideration, suggesting that all demonstrate affinities with the Hamameli-
daceae; he further emphasized this opinion in 1904 (7) and 1905 (8, 9) —
it is an opinion which has found no supporters and which is scarcely sub-
stantiated by the facts. Harms’ treatment in Engl. & Prantl, Nat. Pfl.
Nachtr. 3: 110-112 (1906) is important because of its concise summary
of then current opinions, including van Tieghem’s, certain of whose family
names were properly latinized. Some of these family names now came
into general usage, the Cercidiphyllaceae and Eucommiaceae being accepted
by Engler in the sixth edition of his Syllabus (1909).
128 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
From a taxonomic point of view, the most carefully documented work
on the Chinese representatives of Tetracentron, Cercidiphyllum, Euptelea,
and Eucommia has been that of Rehder and Wilson, in Sargent’s Plantae
Wilsonianae (vol. 1, 1913). In very detailed studies of the Cercidi-
phyllaceae in 1916 (11) and 1918 (12), Harms concluded that the family
is correctly placed in the Ranales. An elaborate study of the leaf-anatomy
of Cercidiphyllum was published in 1923 and 1924 by Weisse (29). The
family Eucommiaceae was treated in excellent detail by Harms in 1930
(13) and was placed in the Rosales following the Roridulaceae. The
Flora of Japan by Makino and Nemoto (Nippon-Shokubutsu-Soran, ed.
2. 1931) is perhaps the first major floral effort to recognize the family
Eupteleaceae as distinct. In 1933 McLaughlin (21), on the basis of his
examination of their wood anatomy, suggested that Euptelea and Tetra-
centron be made the types of two separate families, although the use of
such family names was avoided. In the same year, Harms (14) revised
his conclusions on the Eucommiaceae to place the family in the Urticales.
This position was given support by Tippo’s careful study of the Eucom-
miaceae in 1940 (27).
In the two preceding paragraphs I have listed only the recent major
studies pertaining to this group of plants. There remain to be consid-
ered the opinions of various recent phylogenists. The opinion of Hallier
has already been mentioned, and to a certain extent this doubtless influ-
enced that of Lotsy, who in 1911 (20) placed Tetracentron in the Mag-
noliaceae, Trochodendron and Euptelea in the Trochodendraceae, and
Eucommia and Cercidiphylilum in the Hamamelidaceae, suggesting the
derivation of the latter family from a ranalian ancestor through this series.
Bessey, in 1915 (1), placed the Trochodendraceae and Cercidiphyllaceae
in the Ranales and the Eucommiaceae in the Rosales, but unfortunately
he offered no opinion as to the position of Euptelea and Tetracentron.
Hayata (15) agreed with Bessey in the position of the three above-
mentioned families. Hutchinson, in his several discussions pertaining to
the ranalian complex (17, 18, 19), placed the Trochodendraceae (includ-
ing Trochodendron and Euptelea) and Cercidiphyllaceae in the Magnoliales
and the Eucommiaceae in the Hamamelidales; he did not account for
Tetracentron. Wettstein’s treatment in 1935 (30) has the Eucommiaceae
in the Urticales, the Cercidiphyllaceae and Eupteleaceae in the Hamame-
lidales, and the Trochodendraceae (Trochodendron only) in the Poly-
carpicae following Magnoliaceae, which includes Tetracentron. In the
most recent edition of Engler’s Syllabus, Diels (4) has placed the Trocho-
dendraceae (including Trochodendron and Euptelea) and Cercidiphy]-
laceae in the Ranales and the Eucommiaceae in the Rosales, while Tetra-
centron is not mentioned.
From these more or less current opinions, the reader may observe that
agreement as to the delimitation and phylogenetic position of these groups
has not been reached; yet it is evident that progress has been made and
that these small genera have appeared highly important to numerous stu-
1945] SMITH, TROCHODENDRON AND TETRACENTRON 129
dents. We propose to discuss the genera and to indicate the characters
that have led us to the opinion that each represents a unigeneric family.
Eucommia may be left out of consideration, for Tippo (27) has ably sum-
marized the evidence for placing this genus in the Urticales. Cercidiphyl-
lum remains highly puzzling and must still be examined in great detail:
we feel that it may be included in the Ranales, but its exact relationships
appear very questionable. The remaining genera — Trochodendron,
Tetracentron, and Euptelea — will be discussed from a taxonomic view-
point in this paper and a subsequent one.
TROCHODENDRACEAE
Trochodendraceae Prantl in E. & P. Nat. Pfl. 3(2): 21, p. P. (quoad Trochodendron).
1888; Sargent in Garden and Forest 7: 105, p. p. 1894, For. Fl. Jap. 13, p. p 894:
Oliv. in Hook. Ic. Pl. 24: pl. 2361, p. p. (quoad Trochodendron) 895; Harms in
Ber. Deutsch. Bot. Ges. 15: 359, p. p. (quoad Trochodendron). 1897, in E. & P.
Nat. Pfl. Nachtr. 1: 158, p. p. 1897; Engl. Syllabus ed. 2. 116, excl. Cercidiphyl-
lum. 1898; v. Tiegh. in Jour. de Bot. 14: 262 seq. (Troc
03, ed. 4. 124, p. p. 1904; Schneid. lll. Handb. Laub-
Y PLN
Syllabus ed. 6. 132, excl. Euptelea. 1909; Lotsy, Vortr. Bot. Stammesg. 3: 456,
in Ann. Mo. Bot. Gard. 2: 128. 1915; Engl. & Gilg,
Kew Bull. 1924: 120. 1924, Fam. Fl. Pl. Dicot. 85, excl.
Man. Cult. Trees & Shrubs 212, excl. Euptelea. 1927, ed. 2. 244, excl. Euptelea.
1940; Makino & Nemoto, Nippon-Shokubutsu-Séran (FI. Jap.) ed. 2. 307. 1931;
Wettst. Handb. Syst. Bot. ed. 4. 2: 690. 1935; Diels, Engl. Syllabus ed. 11. 205,
excl. Euptelea. 1936; Masamune, Short Fl. Formos. 63. 1936; Nemoto, Nippon-
Shokubutsu-Soran-Hoi (Fl. Jap. Suppl.) 208. 1936.
Magnoliaceae subord. Illicieae Endl. Enchir. Bot. 428, p. p. (quoad Trochodendron)
1841
1. a A. \
Magnoliaceae trib. Illicieae sensu Walp. Rep. 1: 72, p. p. (quoad Trochod
maa a
yet =D: d Trochodendron). 1843; non DC.
Magnoliaceae 11. Wintereae Lindl. Veg. Kingd. ed. 2. 419, p. p. (quoad Trochoden-
dron). 1847, ed. 3. 419, p. p. 1853.
Trochodendreae Seem. in Jour. Bot. 2: 238 (order), excl. Euptelea. 1864; Eichl. in
Flora 48:12 (familie), excl. Euptelea. 1865, in Jour. Bot. 3: 150 (order), excl.
Euptelea. 1865.
Magnoliaceae trib. Trochodendreae Benth. & Hook. f. Gen. Pl. 1: 954, excl. Euptelea.
1867; Baill. Hist. Pl. 1: 164, p. p. 1868-69; Durand, Ind. Gen. Phan. 4, p. p.
(quoad Trochodendron). 1888; Finet & Gagnep. in Bull. Soc. Bot. Fr. 52: Mem. 4:
23 (Trochodendrées), p. p. (quoad Trochodendron). 1905 (repr. Contr. Fl. As.
Or. 22.23; 1907):
Magnoliaceae II. Trochodendreae Eichl. Bliithendiagr. 2: 150, excl. Euptelea. 1878.
Trochodendraceae I1. Trochodendroideae Harms in Ber. Deutsch. Bot. Ges. 15: 359.
1897, in E. & P. Nat. Pfl. Nachtr. 1: 159. 1897.
The Trochodendraceae is in this paper interpreted as a unigeneric fam-
ily, whereas most of the above references are to a more inclusive concept.
For the most part, Euptelea has been included in the family, while many
writers have extended it to include Cercidiphyllum and even Eucommia,
except by a few writers Tetracentron has been excluded, but this genus is by
130 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
far the closest relative of Trochodendron. Of the references listed above,
the following interpret the Trochodendraceae in the limited unigeneric
sense: van Tieghem (1900), Makino & Nemoto (1931), and Wettstein
(1935). Harms’ subfamily Trochodendroideae (1897) is also limited to
the single genus Trochodendron.
Trochodendron Sieb. & Zucc. Fl. Jap. 1: 83. 1838; Endl. Gen. Pl. 839. 1839, Enchir.
28. 1841; Steudel, Nom. ed. 2. 2: 720. 1841; Endl. Gen. Pl. Suppl. 2: 73.
1847, ed. 3. 419. 1853; Miers in Ann. Mag. Nat. Hist. III. 2: 115. 1858, Contrib.
Bot. 1: 144. 1861; Benth. & Hook. f. Gen. Pl. 1:17. 1862, 954. 1867; Seem. in
Bluthendiagr. 2: 150. 1878; Durand, Gen. Ind. Phan. 4. 1888; Prantl in E. & P.
Nat. Pfl. 3(2): 23. 1888; Harms in Ber. Deutsch. Bot. Ges. 15: 350. 1897, in FE. &
P. Nat. Pfl. Nachtr. 1: 159. 1897; Solereder in Ber. Deutsch. Bot. Ges. 17: 397.
1900; v. Tiegh. in Jour. de Bot. 14: 262. 1900; Hall. f. in Ber. Deutsch. Bot. Ges.
23:89 (Trochodendrum). 1905, in New Phyt. 4:157 (Trochodendrum). 1905;
Harms in E, & P. Nat. Pfl. Nachtr. 3: 111. 1906; Lotsy, Vortr. Bot. Stammesg.
3: 456. 1911; Rehder in Bailey, Stand. Cycl. Hort. 6: 3386. 1917; Hutchinson,
Fam. Fl. Pl. Dicot. 85. 1926; Rehder, Man. Cult. Trees & Shrubs 213. 1927, ed. 2.
245. 1940; Wettst. Handb. Syst. Bot. ed. 4. 2: 689, 1935.
Gymnanthus Jungh. in Tijdschr. Nat. Gesch. Phys. 7: 308. 1840.
A monotypic genus of trees; branchlets terete, marked at the nodes with
conspicuous pseudoverticillate leaf-scars and, above these, by the narrow
scars of bud-scales, terminated by a conspicuous ovoid bud, the bud-scales
teoles on the torus or just below it and sometimes another 1—3 farther down
(these linear, bract-like); perianth none (unless the above-described toral
bracteoles are perianth-remnants) ; pedicel swollen into the subcoriaceous
obconical torus, this bearing the numerous stamens on its rugulose outer
basifixed oblong 4-sporangiate mucronulate anthers, the thecae dehiscing
by lateral clefts for their entire length: carpels fused to the torus at base,
spreading, at anthesis laterally coalescent, the ovary unilocular, with 2
rows of anatropous ovules borne near the ventral suture, abruptly narrowed
distally into a style, this (like the ovary) distinctly conduplicate and
deeply canaliculate ventrally; fruit a follicetum composed of laterally
1A detailed discussion of the development of the inflorescence is given by Wagner
(28), who considers it an interrupted “Primanpleiochasium.”
Fouiar SCLEREIDS OF TTROCHODENDRON
ie t tei 5
Vite vie
ae
A Phy Kg
a ore
t <— ¢ 2 mY ys
Puate I
Jour. Arnotp Ars. Voi. XXVI Piate II
FoLIAR SCLEREIDS OF TROCHODENDRON
Jour. Arno_p Ars. VoL. XXVI Prate III
FOLIAR SCLERE}OS OF TROCHODENDRON
Jour. Arnoip Ars. Voi. XXVI Pate 1V
FOLIAR SCLEREIDS OF ‘TROCHODENDRON
1945] MERRILL, NEW SPECIES FROM HONGKONG 163
TWO NEW SPECIES FROM THE VICINITY OF HONGKONG
E. D. MERRILL
With three text-figures
IN THE course of identifying a collection of plants made by Mr. Y. W.
Taam at Hongkong, Lantao Island, and Hongkong, New Territory, I was
rather surprised to note several apparently undescribed species from a
limited area that has been intensively explored from a botanical standpoint
for more than a century. The material studied represents approximately
575 numbers, and it was submitted to me by Prof. H. H. Bartlett of the
University of Michigan, who financed the field work. The material was
well selected and well prepared.
Two novelties are described in this paper. In addition to these, one
species previously described from outside of China is recorded for the first
time from that country. A third species illustrated is what I take to be the
little known Maackia australis (Dunn) Takeda. Incidentally, the collec-
tion contains a number of species previously described from various parts
of Kwangtung Province but which hitherto have not been recorded from
Hongkong and Lantao Islands or from Hongkong, New Territory. WNat-
urally, the collection made in what is a most important classical region
contains a great many topotypes— that is, specimens collected from
approximately the same localities as were those on which many scores of
descriptions of new species were based. Fortunately it was possible for
Mr. Taam to ship his material to the United States before the Japanese
occupation of Hongkong took place. In addition to the few records in-
cluded in this paper, his material showed clearly that the Kwangtung-
Hongkong form long passing as Hippocratea obtusifolia Roxb., the type of
which was from India, is actually rather remote from that species, and else-
where in this Journal (p. 170) Dr. A. C. Smith has described this as a new
species of Loeseneriella.
ORCHIDACEAE
Eulophia R. Brown
Eulophia macrorhiza Blume, FI. Jav. Orch. 155 (Orch. Archipel. Ind. 183). t. 63, f. 2,
t. 66B. 1858
Honcxonc: Lantao Island, near Tungchung, rare in thickets, Y. W. 2092,
June 3, 1941, det. C. Schweinfurth. A variable species extending from Java ‘a Celebs
New Guinea, and the Philippines (Luzon, Leyte) ; new to the area and to Chi
LEGUMINOSAE
Maackia Ruprecht & Maximowicz
Maackia ellipticocarpa sp. nov. Fie al:
utex circiter 2 m. altus, inflorescentiis exceptis glaber vel subglaber
(floribus ignotis); ramis teretibus levibus, atro-purpureis vel olivaceo-
purpureis, conspersissime lenticellatis, ramulis ultimis glabris, 2 mm.
164. JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
diametro; foliis 9-15 cm. longis, foliolis 7-9, ovatis vel elliptico-ovatis,
plerumque acutis vel leviter acuminatis ‘basi plerumque rotundatis, inter-
dum obtusis, subcoriaceis, 2—4.5 cm. longis, 1.2—2.5 cm. latis, olivaceis,
subopacis, subtus paullo pallidioribus, utrinque glaberrimis, nervis primariis
utrinsecus circiter 6, arcuato-anastomosantibus, subtus elevatis, distinctis,
reticulis ultimis subconfertis: petiolulis leviter incrassatis, 1 1.5 mm.
longis; inflorescentiis terminalibus, sub fructu ad 12 cm. longis, ramis
primariis 2 vel 3, racemiformibus, partibus junioribus obscure breviter
pubescentibus glabrescentibus; pedicellis 5 mm. longis, obscure breviter
adpresse pubescentibus, vetustioribus glabrescentibus; fructibus ellipticis
vel oblongo-ellipticis, compressis, 23.5 cm. longis, 1-1.5 cm. latis, valvis
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Fic. 1. — ellipticocarpa Merrill: a. a fruiting branchlet, x 4; 6. a pod with
seeds, X
glabris vel deorsum leviter adpresse pubescentibus, distincte consperseque
glandulosis, utrinque rotundatis vel basi interdum late acutis vel obtusis,
irregulariter subtransverse reticulato-venosis, ~~ enemas anguste
carinatis, haud alatis; seminibus 1-3, ad 7 mm. longis
Hon N Lantao Island, near Simca ng, Y. W. Taam nee Sept. 12, 1940,
iar on dry clay slopes near the seashore, among scattered shru
This species is clearly allied to Maackia Tashiroi ae Makino, of
southern Japan and the Liu Kiu Islands, differing in its fewer leaflets and
its distinctly glandular fruit-valves. The original illustration of Cladrastis
Tashiroi Yatabe, Bot. Mag. Tokyo 6: 345. ¢. 10. 1892, is excellent, and we
fortunately have a fine series of specimens from Oshima representing the
species, these with both flowers and fruits. The number of leaflets on this
1945] MERRILL, NEW SPECIES FROM HONGKONG 165
material varies from 9 to 15 on each leaf. Makino, Notes Bot. Gard.
Edinb, 8: 102. 1913, has explained the confusion regarding Derris chinensis
Benth., the fruiting specimen of which, from Oshima, actually represents
Maackia Tashiroi (Yatabe) Makino, while the Hongkong flowering speci-
men represents Millettia pulchra Benth.
Maackia australis (Dunn) Takeda, Notes Bot. Gard. Edinb. 8: 102. t. 27, f. 57-62.
1913. Fic. 2.
Cladrastis australis Dunn, Kew Bull. Add. Series 10: 86. 1912.
Honcxonc: Lantao Island, ¥. W. Taam 2134, June 7, 1941, a shrub 2 m. high,
abundant on damp slopes among scattered shrubs, flowers white, odorless.
Fic. 2. Maackia australis (Dunn) Takeda: a. a flowering branch, x 4; b. calyx,
ovary, and style; c. ovary and style; d. calyx, spread, showing the three small lobes and
the fourth larger one; e. keel petals; f. a wing petal; g. standard; b-g all enlarged.
When this specinren was first studied it was thought to represent a
distinct species, because of certain discrepancies between its characters and
those of Cladrastis australis Dunn as indicated in the original description
of the latter, and in Takeda’s consideration of it. In this Taam specimen
the standard is oblong or narrowly oblong-obovate, not ovate as in Dunn’s
description, and the leaflets have 7 or 8 pairs of lateral nerves rather than 5S.
Dunn described the leaflets of his species as ‘‘in apicem obtusum angustata,”’
but Takeda, on the basis of the same material that Dunn had, says that
they are mucronulate, and his figure, ¢. 27, f. 61, shows them to be very
slenderly and sharply acuminate. His figure of the standard, t. 27, f. 60,
166 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
shows it to be very narrowly oblong-obovate (not ovate as Dunn says),
the slender nerves extending to the base, the basal parts not thickened and
nerveless or nearly so as in the Taam collection. A re-examination of the
type specimen and a comparison of the Taam material with it is desirable,
as it may reasonably be expected that there are certain errors in the original
description, or possibly representatives of more than a single species were
included.
Dunn’s description was based on a collection made on the North West
River in Kwangtung Province, and a Millet collection, “Sophora from
China, Hort. Soc. Hort. Lond. 1838,” indicating that the species was culti-
vated in England at that date. It is by no means impossible that this
Millet specimen was grown from seeds collected by him on Lantao Island.
Takeda says that the Millet specimen in the Kew Herbarium bears the date
1835. Certainly at the date indicated there was no possibility of the Millet
collection having been made on the North West River in Kwangtung
Province.
EBENACEAE
Diospyros Linnaeus
Diospyros Taamii sp. nov. Fic. 3
Arbor parva, circiter 7 m. alta, partibus junioribus fructibusque exceptis
glabra; ramis teretibus, glabris, in sicco longitudinaliter subrugosis, ramulis
M. SUTT/S
Fic. 3. Diospyros Taamii Merrill: a. a fruiting branch, x 3; b. a leaf, with a por-
tion enlarged to show the ultimate reticulations; c. an immature . x 14; d. cross-
section of an immature fruit, showing the persistent sepals; e. an Individual sepal,
dorsal view; f. dorsal view of a fruiting calyx; all somewhat enlarged.
1945] MERRILL, NEW SPECIES FROM HONGKONG 167
ultimis gracilibus, glabris, 1 mm. diametro; foliis firmiter chartaceis vel
subcoriaceis, utrinque glabris, in sicco pallidis, oblongis vel oblongo-ellip-
ticis, 6-10 cm. longis, 2—4 cm. latis, sursum angustatis, distincte ae oe
acuminatis, basi subrotundatis vel late acutis, nervis primariis utrinsecus
5 vel 6, supra obscuris, subtus distinctis, elevatis, curvato-adscendentibus,
arcuato-anastomosantibus, utrinque sat dense reticulatis sed vix foveolatis;
petiolo 6—8 mm. longo, glabro; floribus ignotis; fructibus in ramulis ultimis
solitariis vel 2 vel 3 in ramulis specialibus racemosim dispositis, his plus
minusve subadpresse hirsutis et aliquando foliiferis, fructibus globosis,
immaturis circiter 1 cm. diametro, 8-locellatis, densissime adpresse breviter
pallide pubescentibus; sepalis persistentibus coriaceis, ovatis, circiter 1 cm.
longis et 8 mm. latis, sursum leviter angustatis, rotundatis, utrinque dense
subreticulato-venosis, venis venulisque subelevatis, extus consperse hirsutis,
pilis longis, adpressis, pallidis.
Honckonc: Lantao Island, Tai Shan, Y. W. Taam 2169, June 12, 1941, abundant
on moist slopes in thickets
With fruiting mena alone available it is difficult to place this rather
strongly marked species in its proper group. It seems to be allied to
Diospyros Tutcheri Dunn, but in that species the fruits are described as
glabrous.
ARNOLD ARBORETUM,
HARVARD UNIVERSITY.
168 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
DAVID DON’S “PRODROMUS FLORAE NEPALENSIS”
WILLIAM T. STEARN
Tue ‘“Prodromus Florae Nepalensis, sive Enumeratio Vegetabilium quas
in Itinere per Nepaliam proprie dictam et Regiones conterminas, ann.
1802-1803, detexit atque legit D. D. Franciscus Hamilton, (olim
Buchanan)” by David Don (1799-1841) is an unpretentious little book of
considerable importance to students of the Himalayan flora,! for many
species were first described in its 256 pages. It evidently took a long time
to prepare. A letter to Wallich from Buchanan-Hamilton dated 16 October
1821 states that “A Mr. Don, however, who lives with Mr. Lambert, to
whom I gave duplicates of the collection presented to Sir J. E. Smith, is
engaged in publishing an account of them together with those which you
have sent, and I believe he has both abilities and industry to produce a
very valuable work.” (Ann. Roy. Bot. Gard. Calcutta 10[2]: xxxi. 1905),
Don’s preface is dated “‘Kalendis Octobris, Ann. 1824” and in Trans. Linn.
Soc. London 18: 518 (1841) he states that it was “completed and some
copies of the work distributed before the close of 1824.” As the title-page
is dated “MDCCCXXV,” it was evidently not intended or expected to be
available to the public until 1825, and the Monthly Literary Advertiser,
no. 240, p. 28, 32 (9 April 1825) lists it among the new books published
in March 1825. 0.6-0.7 mm.; ovario in disco
immerso trigono, ovulis in quoque loculo i oblique adscendentibus, stvlo
conico-subulato 0.8—1 mm. longo truncato; capsulis divergentibus obovato-
1945] SMITH, NOTES ON HIPPOCRATEACEAE 171
ellipticis, maturitate 3.5-6 cm. longis et 1.5—3.5 cm. latis, basi obtusis, apice
emarginatis, pericarpio venis longitudinalibus anastomosantibus prominulis
copiose striato; seminibus plerumque 4, ala basali submembranacea vel
papyracea ellip tica maturitate ad 3 2 cm. basi obtusa, parte embrvoni-
fera fies Slipcides ad 15 mm. mon
Fic. 1. Loeseneriella concinna; a. flowering branchlet, x 3; 6. detail of inflorescence.
x 4; c. flower slightly before full eae x 3; d. flower with two petals removed,
3; e. fruit with one capsule aborted, ; f. seed, x 4%. Figs. a-d drawn from the
type, e from Tsang 21743, f from Tsang ee
Cuina: Hongkong: Ford (A); Lantao Island, Shantao, Tungchung and
vicinity, Y. W. Taam 2105 (A, type), June 4, 1941 (fairly common on sandy stony
slope among scattered shrubs; flowers yellow) ; Lantao Island, Taai Ue Shan, Lingnan
Univ. 16674 (Tsang) (A, NY) (in a ravine; fruit yellow; native pe Ch’ing Heung
Tsai Shue); Kwangtung: Tung Koo Shaw Tapu District, W. T. Tsang 21743 (A,
NY) (fairly common on dry steep iit in are sandy rocky soil, among scattered
shrubs [fruit]); Kwangsi: Shap Man ai Shan, near Iu Shan village, Shang-sze
District, Kwangtung border, W. T. Res 20444 (A) (fairly common in thickets in
silty clayey rocky soil; owe yellow).
This species of southern China, which has been passing in herbaria as
Hippocratea obtusifolia Roxb., may be readily distinguished from that
Indian species, discussed above, as follows:
Petioles 4-10 mm. long; leaf-blades elliptic, often broadly so, 4-11 x 2.5— 7 cm.; inflo-
rescence ape. often as long as the leaves, many-flowered; pedicels 1.5 -3 mm.
long at ae with the sepals brown-puberulent; style 1.2-1.6 mm. long; ovules
gs a OE nn TPE YP! etek meee ee eT On L. obtusifolia.
172 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
Petioles 2-4 mm. long; leaf-blades oblong-elliptic, usually 4-7 x 1.5-3.5 cm.; inflo-
rescence compact, shorter than the leaves, few-flowered; pedicels 5-15 mm. long
at anthesis, with the sepals essentially glabrous; style 0.8-1 mm. long; ovules 4
ge | erry ey rere rr er er err ee eer res Se ree L. concinna.
Loeseneriella Merrilliana sp. no Fic
— obtusifolia sensu Merr. in ak Sci. Jour. 6: 328. 1928 [1930]; non
mri uotabilis ramulis divaricatis glabris fusco-cinereis saepe copiose
scabrido- lenticellatis: foliis oppositis glabris, petiolis robustis canaliculatis
5-8 mm. longis, laminis chartaceo-coriaceis in sicco fusco- sehr? sub-
nitentibus oblongo-ellipticis, (5-) 6-10.5 cm. longis, (2—) 2 cm.
latis, basi obtusis vel anguste rotundatis et in petiolum Fh ot Rk i
¢ Suttrcs
Fr oo epee a. flowering branchlet, x 4; b. flower with two
ae saaheie’ x 3; c. flower, ; d. fruit with one persistent capsule, x 4; e. seed,
. Figs. a-c en from the ae d and e from How 73743.
in acuminem obtusum 5-10 mm. longum abrupte cuspidatis, margine incon-
spicue crenato-serrulatis (dentibus circiter 2 per centimetrum obscure
oe costa utrinque subprominente, nervis secundariis utrinsecus
4-6 adscendentibus utrinque paullo elevatis, rete venularum intricato
utrinque priminulo vel supra subimmerso; inflorescentiis cymoso-panicu-
latis apicem ramulorum versus axillaribus 2. ‘5-6 cm. longis saepe PABST,
ramulis pedicellisque dense sed minute farinoso-puberulis, pedunculo com
muni plerumque 1—1.5 cm. longo, bracteis bracteolisque deltoideis acutis
is; floribus ultimis binis vel ternatis, pedicellis
gracilibus sub anthesi in dichotomiis 3-8 mm. longis alteris circiter 2 mm.
]
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1945] SMITH, NOTES ON HIPPOCRATEACEAE 173
longis; sepalis ovato-deltoideis, 1-1.2 mm. longis, 1.5—1.8 mm. latis, apice
obtusis, margine ciliolatis; petalis tenuiter carnosis oblongo- lanceolatis,
4-5 mm. longis, 1.7-2.5 mm. latis, apice dorso mucronulato-calcaratis,
versus obscure constricto; filamentis ligulato- deltoideis 1-1.7 mm. longis,
basi es 1.5 mm. latis, superne angustatis, antheris 0.4-0.5 x 0.6— 0.8
mm.; ovario in disco semi-immerso trigono, ovulis in quoque loculo 8, stylo
conico- subulato 1-1.5 mm. longo truncato; capsulis immaturis obovato-
ellipticis vel anguste ellipticis, ad 6 cm. longis et 3.2 cm. latis, basi obtusis
et minute stipitatis, apice rotundatis vel emarginatis, pericarpio venis in-
conspicue striato, seminibus abortu paucis, ala basali membranacea late
si ae
CH : Sin Woh, Taam Chau District, W. T. Tsang 381 [Lingnan
Univ. pair (A, Type; NY), May 19, 1928 (growing on the plain of a stream; flowers
white, fragrant) ; Po-ting, alt. 350-550 m., F. C. How 72209 (A), 73743 (A) (twining
plants in forested ravines; flowers [72209] pale green; fruits [73743] lustrous green) ;
Yeung Lam Shan, near Yeung Lam village, Yai Hsien, S. K. Lau 6337 (A) (rare woody
climber on dry steep slop sandy soil in forest; flowers white); Yaichow, H. Y.
Liang 62274 (NY) Cees in forested ravine: immature fruits green).
This new species from Hainan differs from the plant of Hongkong,
Kwangtung, and Kwangsi above described as L. concinna in several obvious
characters, namely the longer petioles, the more coriaceous and larger
leaf-blades with a more obvious acumen, the usually more ample inflo-
rescence with shorter pedicels and larger sepals, the farinose-puberulent
character of its inflorescence-branches and petals, and in having 8 rather
than 4 ovules per locule. From the Indian L. obtusifolia (Roxb.) A. C.
Sm. the new species differs in the proportionately narrower leaf-blades,
which are more coriaceous in texture and have a more obvious acumen, in
the longer pedicels and slightly larger petals, and in having 8 rather than 6
ovules per locule.
A closer relative of the new species is apparently the Indo-Chinese Loe-
seneriella dinhensis (Pierre) comb. nov. (Hippocratea dinhensis Pierre,
Fl, For. Cochinch, 4: pl. 301A. 1893). The Hainan plant differs from this,
however, in its fewer secondary nerves, its more ample inflorescence, and
its less highly elevated disk. It should be noted that the Indo-Chinese
specimen Pételot 2119 (or 2219) cited by Merrill (in Jour. Arnold Arb.
21: 374. 1940) as Hippocratea obtusifolia Roxb. is very close to the new
species and possibly identical with it. However, Pételot 2119 has the leaf-
blades proportionately a little broader than our species, the pedicels shorter,
and the ovules only 6 per locule. The Pételot plant is somewhat more
suggestive of the true L. obtusifolia than the other specimens here discussed,
but I doubt if it can be referred to the Indian species.
Hippocratea obtusifolia and H. dinhensis are placed by Loesener (in
Nat. Pfl. ed. 2. 20b: 213. 1942) in Hippocratea subgen. Euhippocratea
sect. Barbatae, the type-including section of Hippocratea, which in the
writer’s opinion (see Brittonia 3: 356-367. 1940) is a monotypic American
genus.
174 JOURNAL OF THE ARNOLD ARBORETUM (VOL. XXVI
Loeseneriella yunnanensis (Hu) comb. nov.
Hippocratea yunnanensis Hu in Bull. Fan Mem. Inst. Biol. 10: 152. 1940.
Viinnan: Pu-Erh Hsien, Po-Pien-Kiang, alt. 1100 m., C. W. Wang
§1219 (A, TYPE COLL.) (on open dry slope along river-bank; flowers greenish) ; Chu-
yiian, A. Henry 10865 (ANY) (large climber; flowers green) ; Shih-Ling, alt. 1200 m.,
A. Henry 13274 (A,NY) (climbing shrub).
This is another species of the general alliance of L. obtusifolia (Roxb.)
A. C. Sm., to which it is very similar in foliage, differing perhaps in having
slightly shorter petioles and thicker leaf-blades. The flowers of L. yun-
nanensis are noticeably larger (sepals about 1.5 2 mm.; petals 5-8 x
2.5-4 mm.) than those of L. obtusifolia (sepals 0.7-0.8 0.6-1.2 mm.;
petals 3.5-4.5 1.5-2 mm.). The disk is less highly elevated, being only
1-1.5 mm. high but widening to a base 3.5-4 mm. across; the disk of the
Indian species is 1.7—2.3 mm. in diameter but proportionately higher. The
filaments of the Yiinnan plant are longer (2—2.5 mm. rather than 1—1.5
mm.), and the ovules are 8-10 per locule rather than 6. The available
fruits of the two species show no consequential differences. Hu compared
his new species with Hippocratea puberula Craib, but that species has much
larger and longer-petioled leaves and smaller flowers.
Loeseneriella Arnottiana (Wight) comb.
ye sag! Arnottiana Wight, Ill. “Ind on 1: 133. pl. 46, 47A. [1839]; Lawson
Fl. Brit. Ind. 1: 624.
INDIA: eae Wight 463 — con 2445 (NY) (probably parts of TYPE
COLL.) ; Kodaikanal Region, Palni Hills, Madura District, Anglade (A).
The Wight specimens cited above agree well with the original description
and illustrations and are very probably a part of the type collection, cited
without number; nos. 463 and 2445 are so similar that it seems likely that
they are parts of a single original collection, subsequently re-numbered.
The original locality is mentioned as ‘Malabar’ by Wight, and as Quilon
[Travancore State, Madras] by Lawson. The Anglade collection, from
the same part of India, agrees precisely with Wight 463 and 2445. The
species is very distinct in its large flowers (about 8 mm. in diameter at
anthesis), spreading spatulate clawed petals, and papillose disk; the ovules
are 8 or 10 per locule.
Loesener (in Nat. Pfl. ed. 2. 20b: 214. 1942) places Hippocratea Arnot-
tiana in Hippocratea subgen. Euhippocratea sect. Scutellatae, a synonym
of Prionostemma Miers, a very different American genus in the writer’s
understanding (see Brittonia 3: 391-396. 1940).
Loeseneriella serrata (Griffith) comb. no
Hippocratea serrata Griffith, Not. Pl. es 4: 473. 1854, Ic. Pl. As. 4: pl. 382. 1854.
Griffith’s species is difficult to interpret, due to the inadequacy of the
original description and plate, but the latter is sufficiently clear to suggest
with reasonable certainty that a species of Loeseneriella is represented,
characterized by narrowly oblong-elliptic serrate leaf-blades and fairly short
inflorescences. The original locality is given as: ‘Journey from Assam to
Ava. Tsakan Delvi,” probably in central Burma.
A specimen which agrees well with Griffith’s plate is Helfer 905 (GH),
from ‘““Tenasserim and Andamans.”’ The leaves of this specimen are essen-
1945] SMITH, NOTES ON HIPPOCRATEACEAE 175
tially identical with those of the plate and agree with the description as to
“costa venisque lutescentibus, subtus pallidioribus, ...” The flowers of
the Helfer specimen have the pedicels and sepals glandular-puberulent, the
petals lanceolate-oblong, 3.5-4 mm. long, the disk obvious, the filaments
and style about 1 mm. long, and the ovules 6 or 8 per locule.
Pristimera Miers
Pristimera indica (Willd.) A. C. Sm. in Am. a Bot. 28: 440. 1941.
shee indica Willd. Sp. Pl. 1: 193. 179
Cu Hainan: ie Chik Shan and vicinity, Ch’ang-kiang District, S. A. Lau
1675 (NY) ; Ue Lung Shan, Ch’ang-kiang District, S. K. Lau 3731 (A); Pak Shik Ling
and vicinity, Ching Mai earns G21: Let 847 (NY); Yaichow, H. Y. Liang 62979
a
The above specimens are listed because the species appears not to have
been otherwise recorded from China. Although, as indicated by herbarium
records, Pristimera indica is a very widespread species, it is fairly variable
as currently interpreted and its range cannot be stated without detailed
study of the genus. The Hainan material, however, appears quite identical
with that from India and Ceylon. Willdenow gives the original locality as
‘in India orientali.”
Pristimera setulosa sp. nov. Fice3.
Frutex scandens, ramulis hornotinis acute quadrangularibus copiose setu-
losis (pilis 0.1-0.15 mm. longis, glandulosis [?]), annotinis glabrescentibus,
vetustioribus teretibus cinereis; foliis oppositis glabris, petiolis gracilibus
leviter canaliculatis 3-5 mm. ongis, laminis chartaceis in sicco fusco-viridi-
bus eects 4—6.5 cm. longis, 2—3. 5c m. latis, basi obtusis vel subacutis et
in petiolum decurrentibus, apice abtusis vel breviter et obtuse cuspidatis,
cm. longis et latis, pedunculo ad 17 m ig eee ramulisque acest teri
et copiose setulosis, bracteis oe glabris deltoideo-oblongis 0.5—0.8
mm. longis acutis, bracteolis similibus minoribus; floribus in ramulis ultimis
binis pedicello gracili 0.6-0.8 mm. longo obscure setuloso excepto glabris:
sepalis papyraceis deltoideo-oblongis, 0.4—0.5 mm. longis, circiter 0.3 m
latis, obtusis, margine ee petalis tenuiter carnosis elliptico- oblongis,
circiter 1 mm. longis et 0.5 mm. latis, apice obtusis, integris, utrinque
obscure Sara “abr disco obscuro minutissime annulari; staminibus
3 minutis, filam s ligulatis circiter 0.15 mm. longis, antheris Snag
(OE 0.15 mm.; ovario depresso- subgloboso sub anthesi circiter 0.4 n
diametro 3- apis ovulis in quoque loculo 2 collateralibus, stylo incon-
spicuo circiter 0.15 mm. longo truncato.
INA: -hao [on Yang Chiang (Red River) near Indo-Chinese
boundary], A. Henry 9612 (NY, Type) (large climber with yellow flowers, coll. June
19 [year?]).
The new species differs from the common P. indica (Willd.) A. C. Sm.,
which apparently does not occur in interior China, in its densely setulose
176 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
young branchlets and inflorescence-branches. The hairs are very abun-
dant, stand out stiffly, and are glossy when expanded, as though glandular
in nature. The young branchlets and inflorescence-branches are sharply
quadrangular, whereas these parts in P. indica are usually subterete.
Marthe
Suttis
Fic. 3. Pristimera setulosa; a. flowering branchlet, x 4; 6. detail of inflorescence,
x 3; c. young flowers, x 5; d. flower with two petals removed. x 20. Drawn from
the type.
Pristimera arborea (Roxb.) comb. nov.
Hippocratea arborea Roxb. Hort. Beng. 5, nomen. 1814, Pl. Coast Corom. 3: 3. pl.
205. 1819, Fl. Ind. 1: 171. 1820.
INDIA: Cult. Calcutta Botanic Garden, Wallich 4212C (NY), Collector?, Feb.
1901 (A); ‘East ee rile 910 (GH); “W. Himalaya,” J. F. Duthie ve
BurMa: (Without data), . Prazer 23 (NY). Cuina: Yiinnan: Puerh cliffs,
alt. 1800 m., A. Henry see at (large climber) ; Sheau-meng-yeang, Che-li Hsien,
alt. 960-1000 m., C. W. Wang 75610 (A), 79608 (A) (vines, in woods on mountain-
slopes ; ae penn
The Wallich specimen cited above is listed under Hippocratea arborea
Roxb. by Wallich (Cat. no, 4212C. [1830]) as ‘“HBC.,” thus indicating
that it was taken from a plant cultivated in the Calcutta Botanic Garden.
In habit this Wallich specimen agrees precisely with Roxburgh’s descrip-
tions and illustration; in floral details these descriptions and the illustration
are highly inaccurate, which is not surprising in view of the fact that at
anthesis the flower is little more than 1 mm. in diameter. However, there
1945] SMITH, NOTES ON HIPPOCRATEACEAE 177
seems no doubt that Roxburgh had at hand a species of Pristimera closely
allied to P. indica (Willd.) A. C. Sm. The original collection is said to
have come from the “interior parts of India,’ and Roxburgh (PI. Coast
Corom. 3: 4. 1819) states that the species was in cultivation at Calcutta.
It seems very likely, therefore, oe the Wallich specimen was taken from
the type plant or a descendant of
It is also possible that the tear specimen cited above (Collector?, Feb.
22, 1901) came from a descendant of Roxburgh’s original plant, although
its leaves are slightly thicker and less obviously serrate than those of Wal-
lich 4212C. In its fruit, the collection of 1901 seems to agree well with
Roxburgh’s concept.
Of the other Indian specimens cited above, Griffith 910 has inflorescences
identical with those of Wallich 4212C, although its leaves are somewhat
more coarsely serrate; I believe that the Griffith specimen can be referred
here with reasonable certainty. It is probably part of the same collection
which was questionably referred to Hippocratea arborea by Lawson (in
Hook. f. Fl. Brit. Ind. 1: 625.- 1875), as from “Bhotan and the Khasia
Mts.” The Duthie specimen consists of young leaves and inflorescences
and is accompanied by mature fruits; it is referred to the species with con-
fidence. Lawson (I. c.) states that this species differs from Hippocratea
indica “apparently in nothing but size.” The two species are indeed very
similar in inflorescence characters, but the much larger leaves and fruits of
Pristimera arborea make it readily recognizable.
The cited Burmese specimen is remarkably similar to Wallich 4212C in
both foliage and inflorescences, while the cited Chinese specimens, all in
fruit, seem undoubtedly to belong here. The range of the species is thus
from northeastern India across [northern?| Burma to southern Yunnan.
Apparently it has not previously been recorded outside of India.
Pristimera cambodiana (Pierre) comb. n
Hippocratea cambodiana Pierre, FI. For. aaa 4: pl. 302B. 1893; Pitard in
Lecomte, Fl. Gén. Indo-Chine 1: 896. 19
Inpo-CuH1na: Cambodge: Samrong- hee Fe Pierre 869 (CoTYPE COLL., A, NY).
Yiinnan: Lan-Tsang Hsien, alt. 890-1100 m., C. W. Wang 73135 ar
76712 (a) eee vines, on mountain slope or on outcrop on river-bank). Upper
BuRM J.C. Prazer in 1894 (A, 2 sheets), 39 (NY).
A cael flower associated with Wang 73135 agrees precisely with the
flowers described and well figured by Pierre. The species is marked by its
small flowers, involute petals, inconspicuous disk, and 6-ovulate locules.
In foliage, the cited Chinese specimens agree very well with the available
cotype collection. Wang 76712 bears juvenile fruits, the capsules of which
are much shorter than those of Pierre 869 but similar in texture. The cited
Prazer specimens from Burma agree precisely with Pierre 869 in foliage,
and the available capsules are identical in shape with those of the Pierre
specimen but are shorter, perhaps due to immaturity.
These collections extend the range of the species to southern Yunnan and
northern Burma; otherwise it has been recorded only from the southern
part of Indo-China (Cochinchine and Cambodge).
178 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
Pristimera cambodiana differs from the widespread P. indica (Willd.)
A. C. Sm. in its much larger leaves, flowers, and fruits, its 6- rather than
2-ovulate locules, and in other obvious characters. Nevertheless the floral
characters of the two species are fundamentally similar, and I believe that
Pierre’s species is safely referable to Pristimera, as emended by the writer
in Brittonia 3: 367-383. 1940, and in Am. Jour. Bot. 28: 440. 1941.
A closer relative of P. cambodiana is Pristimera Grahamii (Wight)
comb. nov. (Hippocratea eta Wight, Ill. Ind. Bot. 1: 134. [1839],
Ic. Pl. Ind. Or. 2: pl. 380. 1840). From this Indian species, P. cambodiana
differs in its somewhat fae leaves, shorter pedicels, glabrous rather than
faintly puberulent disk, and longer filaments and_ style. i.
Grahamit is represented by two specimens collected by Law (GH) in th
Kanara District, Bombay; the type (coll. Graham) is said to Ae come
from Bombay.
It should be noted that Wang 73135 was cited by Hu as the type of
Mangifera austro-yunnanensis (in Bull. Fan Mem. Inst. Biol. 10: 160.
1940). This circumstance is doubtless due to a mixture of numbers, as the
data on the field label of our specimen are different, while Hu’s description
of the Mangifera obviously does not pertain to our plant.
Both Hippocratea cambodiana and H. Grahamii are placed by Loesener
(in Nat. Pfl. ed. 2. 20b: 216. 1942) in Hippocratea subgen. Euhippocratea
sect. Thyrsiflorae, whereas Hippocratea indica and H. arborea are placed
by him in Hippocratea subgen. Euhippocratea sect. Micranthae (op. cit.
212)
SUMMARY
Three new Chinese species of Hippocrateaceae are above described and
seven new combinations are proposed in Loeseneriella and Pristimera for
species of southeastern Asia. The seven species of capsular-fruited Hip-
pocrateaceae now known from China may be keyed as follows:
Flowers comparatively large, the petals at least 4 mm. long, often spreading at anthesis,
isk conspicuous, annular-pulvinate, 1-1.5 mm. high, slightly constricted near
the middle; capsules usually ecu: -elliptic Laateneriatlas
Inflorescence-branches and cg essentially glabrous, the hoe 5-15 mm. lon
esis; OV pong 4 per locule; petioles 2-4 mm. long; leaf-blades usually
4-7 x 1.5-3.5 ¢ gf tet ea Kwangtung, Kwangsi). g reseiennn concinna.
Inflorescence- icin and — puberulent at anthesis, the pedicels 2-8 mm. long
nthesis; ovules 8-10 per locule; leaf-blades 5—10.5 6 cm.
PET NON oblong-elliptic, = acumen 5-10 mm. long; petioles 5-8 mm. long;
petals 4-5 x 1.7-2.5 mm.; filaments 1-1.7 mm. long (Hainan)............
pee piSens thee g eee eeee esa parses eeeaus star e4e Poe Merrilliana.
Leaf- aie eae wei or short-cuspidate; petioles 2-5 mm. long; i
mm.; filaments 2—2.5 mm. long (southern Pane
CMEC ASERCL AAD Rh ERGO ERS Se OE Sag 8a eS Loeseneriella yunnanensis.
Flowers small, the petals less than 3 mm. long, suberect at anthesis, the disk obscure,
arcely apparent as a pulvinate thickening at base of stamens; capsules narrowly
oblong (Pristimera).
Petals about 1 mm. long at anthesis; ovules 2 per locule; capsules 2-seeded.
1945] SMITH, NOTES ON HIPPOCRATEACEAE 179
Leaf- Pas a pene Daan 3.5-10 x 2-5 cm.; capsules at maturity (not
kno or P. se sa) 3-4.5 X 1-1.5 ¢
inflorescence- ee glabrous, usually subterete
ainan; also India, Indo-China, Malaysia, etc.)...... Pristimera indica
and inflorescence-branches ane setulose with eile
ndular hairs, sharply quadrangular (Yu ) .Pristimera setulosa.
Leaf-blades chartaceous to subcoriaceous, usually 10-17 x 5-9 cm.; capsules a
x 2.5-3.5 cm. (southern Yiinnan; also noiihedstcen India
istimera arborea.
Santee eee os
(H
Young branchlets
gla
ATIC SUTIN A) ese ee eet toa. sb. a ae cis anaes encmeneeeneya a eeerans
Petals 2.5-3 mm. long at anthesis; ovules 6 per locule; capsules 7-10.5 x 2-3 ¢
usually 6-seeded; leaf-blades coriaceous, usually 12-15 x 5-9 c Gaee
Yiinnan; also Indo-China and Burma)............... Pristimera eee:
In addition to these seven species, another apparently undescribed species
of Loeseneriella from China is represented by R. C. Ching 6573 (NY),
from Chang Tung, E. Tan Shan, Kwangsi. This is a distinct species with
copiously pubescent petals, but the collection seems inadequate to be the
sole basis of a new species.
ARNOLD ARBORETUM,
HARVARD UNIVERSITY.
180 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
A NEW SPECIES OF ISOETES FROM NEW GUINEA*
A. H. G. ALston
Isoétes habbemensis sp. nov.
Rhizoma bilobatum(?), circa 3.5 & 1.5 cm., | cm. altum, appresse semi-
globosum, sursum foliorum fasciculum gerens, subtus radicibus nigro-
pallide rufo-fuscescente, basin versus late alata usque ad 1 cm. lata.
Stomata nulla. Ligula deltoidea. Velum nullum. Sporangia obovato- oblonga,
circa 1 cm. longa, 4 mm. lata, multilocularia. Megasporae circa 575 u in
diametro, sublaeves, valde tricarinatae, siccitate pallide griseo-albidae.
Microsporae dense echinulatae, circa 43 uw in diametro, siccitate fuscescentes.
NETHERLANDS NEw GuineEA: Lake Habbema, alt. 3225 m., Brass 9440 (TYPE in
herb. Brit. Mus.), 9441 (form with short recurved leaves), very abundant in marginal
shallows; 4 km. N. E. of Wilhelmina-top, alt. 3660 m., Brass & hel oS 9974 (form
with a leaves), common in stony shallows of a lake. The cited specimens are
on eel in the herbarium of the British Museum, and iva are in the Gra
Her
This species is separated from /. neoguineensis Bak., the only other
species recorded from New Guinea, by its nearly smooth megaspores, which
are very slightly rugose on the back. These megaspores are greyish white
when dry and light brown when wet. Jsoétes neoguineensis was first col-
lected by A. Giuliarietti on Mount Scratchley at 10000-13000 ft., and re-
cently by Brass (no. 4366) in the shallows of an alpine lake at 3680 m. on
Mount Albert Edward. The megaspores were incorrectly described as
“laeves” by Baker; they are strongly tuberculate. J/soétes philippinensis
Merrill & Perry, from a stream at 400-500 m. near Momungan, in Lanao
Province of Mindanao, is separated by the reticulate sculpturing of its
megaspores. The megaspores of I. sinensis Palmer, which was found in a
pond near the Ming tomb in Spirit Valley (Nanking), are conspicuously
crested. The other Chinese species, J. hypsophila Hand.-Maz., from moor-
land pools at 3600 m. in Yunnan, has smooth mega- and microspores, while
the Indian /. coromandeliana L. f. has tuberculate megaspores
British Museum (Natura History),
Lonpon
*Botanical Results of the Richard Archbold Expeditions.
1945] CROIZAT, EUPHORBIACEAE FROM THE AMERICAS 181
NEW OR CRITICAL EUPHORBIACEAE FROM THE AMERICAS
L. CroizAt
With one plate
Tus PAPER consists of the descriptions of various new species and va-
rieties, a new genus Moacroton from Cuba, and critical notes and records,
reductions, and transfers. Unless otherwise stated, the types of the forms
herein described are preserved in the herbarium of the Arnold Arboretum.
Phyllanthus Linnaeus
Phyllanthus vichadensis sp. nov.
Fruticulus caules strictos plures erectos e rhizomate perenni edens, ramis
ee exalatis subexalatisve. Foliis ellipticis carnosulis enerviis 5—7
m. longis, 2 mm. latis, utrinque aequo iure acuminatis petiolulo ca. 1 mm.
fore: stipalis subsetaceis integris ca. 2 mm. longis. Floribus secus apicem
ramulorum dispositis, ¢ ad 3 capitulatim aggregatis subsessilibus, ? lon-
gius pedicellatis saepissime singulis. Perianthio ¢ ca. 2 mm. longo os
mm. lato 5-lobo, antheris 3 in columna ca. 1.5 mm, alta coalitis longitudi-
naliter dehiscentibus, columna basi incrassata revera haud glandulosa.
Perianthio @ magnitudine fere g 5-lobo, lobis costatis rotundato- ellipticis,
disco apes Eanes ovario globuloso depresso ca. 1 mm. in anthesi
magno, stylis 3 ca. 1 mm. longis e basi liberis subliberisve apice bilobato
partitis, aera sub fructu nonnihil apice incrassato ad 4 mm. longo.
CotomsBiA: Comisaria El Vichada: About 70 km. southeast of Orocué,
Haught 2779.
This new species is described as an inconspicuous perennial of the open
llanos. Its habit is reminiscent of P. diffusus Kl., an annual weed with
which it was originally confused in the herbarium. Fragments of Le-
prieur 319 and Schomburgk 529 in our herbarium show that it definitely
differs from P. hyssopifolioides H. B. K. (well represented by Wiliams
15945a; Venezuela, Amazonas, Puerto Ayacucho) and P. guianensis Kl.
in foliage as well as in floral characters. It may be remarked that the habit
and gross morphology of the latter is reminiscent of the species in the
vicinity of P. caroliniensis Walt.
Croton Linnaeus
Croton heen ap: nov
5-8 cm. longis, 2-5 cm. resi olivaceo-brunneis vel ee Anal
adscendentibus ca. 7—9-jugis, primo jugo ramoso, glandulis stipulisque
182 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
nullis subnullisve, petiolo tomentello gracili, 0.5-2.5 cm. longo. Inflores-
centia spicata brevi ad 4 cm. longa. Floribus @ gracilius pedicellatis, pedi-
cello ca. 3 mm. longo, perianthio totidem lato longoque, petalis sepalisque
subaequalibus, staminibus ad 15. Floribus 9: perianthio ca. 3 mm. longo,
totidemque lato, apetalo, lobis 5 fere ad basem liberis, costulatis, caeterum
glabratis tenuibusque margine integris, ovario subglobuloso pallide ochraceo
hispido ad 3 mm. magno, stylis 3, cruribus fere ad basem Sacro 3.2 mm.
_ longis, glandulis 5 discretis, ‘pedicello crasso ca. 1 mm. longo, semine
. 7 mm, longo, 3—3.5 mm. lato, pallido, valde nitido, en facie anteriori
ne obscurissime grosseque rugulosa, caruncula subquadrangula
parva, columella fructu delapso ad 7 mm. longa, coccis tomento hispido
indutis, epicarpio verruculoso.
Mexico: Baja tag tle Todos Santos, Cape Region, 7. S. Brandegee s. n.,
Oct. 4, 1899 (typE, Herb. Univ. Calif.) ; San José del Cabo, T. S. Brandegee s. n., Sept.
11, 1891; Corral Piedras, Rdkceine Brandegee s.n., Sept. 16, 1893.
My first impulse on examining this peculiar plant was to treat it as a
subspecies of C. fragilis H. B. K., for it clearly belongs in the affinity of
that species. However, a careful study of the group of C. fragilis H. B. K.
as a whole indicated that if this form be placed under that species it was
necessary to merge under this binomial numerous other allied forms that
have uncontroversially been named, characterized and accepted as distinct
entities. In addition, the apparently substrigose indumentum of the upper
surface of the leaf, the large and shiny seeds, and the very hispid indumen-
tum of the young capsules are characters which separate C. caboensis
Croiz. from all other allies of C. fragilis H. B. K. at a glance. Baja Cali-
fornia, lastly, is well known as a region of much endemism. Croton flaves-
cens Greenm. has a quite different indumentum, and sharp keels on its
capsule valves.
Croton culiacanensis sp. nov.
Frutex 1-2 m. altus, innovationibus tenuiter puberulis griseis, raion
Penns Foliis lanceolato- ovatis vel lanceolatis, 3-7 cm. longis,
latis, subtus griseo-tomentulosis, supra pallide viridibus, apice bon
acuminatis, basi rotundatis ve cuneato-rotundatis, eglandulosis sed lami-
bus @: perianthio vix 2 mm. magno albicante, lobis (fructu ineunte) ca.
3.5 mm. longis 2 mm. latis, margine subreduplicativis subintegris, costatis,
petalis nullis, glandulis in disculum connatis latis at haud conspicuis, ovario
ca. 2 mm. longo latoque albicante, hispidulo, obvie 3-cocco, stylis 3, quove
iteratim partito, ca. 2.5 mm. go.
ME Sinaloa: Culiacan, E. Palmer 1507 (type); Ymala, E. Palmer 1446;
Culiacan, Brandegee s. n., 1904; Mazatlan, Eyerdam & Beetle 8660.
Currently this has been identified as representing C. Watsonii Standl.,
or as C. Ortegae Stand]. It is close to both in its affinities, but cannot
easily be confused with either. Croton Watsonii Standl. is endemic to the
eastern coast of Mexico, and has much larger 9 flowers, with a definitely
1945] CROIZAT, EUPHORBIACEAE FROM THE AMERICAS 183
silvery indumentum; the undersurface of the leaf is silvery lepidote, not
puberulous or tomentose. Croton Ortegae Standl. which is known from
the western coast of Mexico, also has much larger ? perianths, styles cleft
but once, smaller ovaries, and, above all, conspicuous tubular glands at the
apex of the petiole. In C. culiacanensis Croiz. no glands appear, and the
base of the blade is somewhat corrugated, and subauriculate in a peculiar
manner.
Croton gynopetalus sp. nov.
capes innovationibus griseo-tomentosis tardius glabratis. Foliis ovatis,
—5 cm. longis, 0.5-2.5 cm. latis, utrinque sed subtus magis conferte
Cee supra olivaceis subtus cinereis vel cinereo-roseatis apice latius
acuminatis basi rotundato-cuneatis vel rotundatis, margine eae vel
plus minusve grosse dentato-s erratis, penninerviis, nervis ca. 5—8-jugis
adscendentibus, glandulis eis patelliformibus ceraceis sivas -
petiolo 1-1.5 cm. longo, stipulis fere nullis. ee spicata ca. 3—
cm. longa. Floribus g : perianthio in alabastro ca. 2 mm. magno, poet
bus, ut videtur, 10-12, pedicello brevissimo. F (onus oe perianthio ca.
3 mm. longo, 2 mm. lato, lobis 5 erectis, postico 1 minore, caeteris subae-
qualibus obovato- ligulatis, integerrimis, petalis setaceis, ovario ellipsoideo
ongo tomentello griseo, stylis 3 ad basem partitis ca. 2 mm.
longis, disco sat evoluto.
Mexico: Oaxaca: San Geronimo, Purpus 7159 (type, Herb. Chicago Nat. Hist.
Museum).
The position of this new species is doubtful. The material now avail-
able clearly shows that a complex of more or less closely related forms
ranges between Guatemala and Vera Cruz, speciating as follows: (1) by
more or less extensive reductions in the size of the lobes of the @ perianth
which are nearest the axis of the inflorescence, and by an increase in the
number of the lobes so reduced; (2) by a tendency in the foliage to be-
come stabilized either as entire or as serrate, the difference between ex-
tremes being of specific order, all floral characters left aside; (3) by varia-
tions in the size of the @ perianth and capsule.
To the complex just mentioned belong: (a) C. jutiapensis Croiz., Gua-
temala; (b) C. itzaeus Lund., Yucatan; (c) C. gynopetalus Croiz., Oaxaca;
(die. ‘ramillatus Croiz. and its var. insignilobus Croiz., Chiapas ‘and Vera
Cruz. Of these five entities, two, C. ramillatus Croiz. and C. jutiapensts
Croiz. have plainly anisomerous ? perianths, the remaining three showing
also some irregularity in the size of the lobes of the @ flower, one or more
of which tend to be smaller than the rest. This group, consequently,
varies in the 2 perianth in a manner strongly suggesting the modifications
taking place in this organ in C. capitatus Michx., C. Palmeri S. Wats., C.
leucophyllus Muell.-Arg. of northeastern Mexico and Texas, and in C. men-
tiens (S. Moore) Pax and its allies of Brazil.
Croton gynopetalus Croiz. has foliage intermediate between that of
C. ramillatus Croiz. and C. itzaeus Lund., that is, more or less entire to
dentate-serrate. It differs from both in the presence of petals in the @
perianth; from the former also in the much larger perianth-lobes, only
184 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
one lobe being reduced; and from the latter in the much less pubescent
and grayish, more definitely ellipsoid ovary. The position of C. ramillatus
Croiz. var. insignilobus Croiz. is still doubtful as between C. ramillatus
Croiz. and C. gynopetalus Croiz., and will remain so until the full signifi-
cance of the petals in the @ flower of single known collection of the latter
is ascertained.
Croton ramillatus sp. nov,
Fruticulus lignosus, videtur dioecius, plus minusve intricatim ramosus,
innovationibus trichomatibus stellatis haud delicatis pallide luteis vel sub-
cinereis vestitis, citius glabratis. Foliis ellipticis vel ovato-lanceolatis, utrin-
que fere aequo jure attenuatis tomentosisque, supra brunneis vel olivaceis
subtus sordide cinereis, margine integerrimis, nervis adscendentibus ca.
5—7-jugis penninerviis, ‘glandulis tubulosis minimis vel saepius subnullis,
petiolo sat gracili 3-5 mm. longo. eae aciapgge exacte spicatis, plerum-
que 7—10 cm. ane Floribus é : staminibus ca. 10 in perianthio ca. 2 mm.
longo latoque, pedicello vix 1.5—-2 mm. longo. Floribus 9 : perianthio habitu
erecto, more proprio anisolobo, scilicet lobis posticis ca. 4 valde diminutis
subsetaceis ad 1 mm. longis, anticis ca. 3 ligulatis rotundatis margine inte-
gerrimis ca. 2 mm. longis, disco valde adpresso subnullove, ovario rotundato
pallide hispidulo ca. 1.5 mm, longo latoque stylis 3, quove ad basem partito
m. longo, capsula sub fructu glabrata, coccis subtrigonis vel potius
oe ta
XICO: peel Cruz: Barranca de Panoya, Purpus 8450 (type, Herb. Chicago
Nat. Hist. Museum) ; Zacuapan, Purpus 2114, 2415; Mata Zarza, Purpus 9324. Three
$ collections, all from Vera Cruz, Purpus 14045, 14145, 16369, may also belong here.
This new species 7 Lge confused with C. rhamnifolius H. B. K. and
C. sphaerocarpus H. B. K. (C. morifolius Willd. sensu Muell.-Arg.), which
it hardly resembles - gt In its vegetative characters it strongly sug-
gests C. Sutup Lund., but the erect 2 flower with uneven lobes up to 8 in
number, and the simple cleft styles set it apart from that species in a
decisive manner.
Croton ramillatus var. insignilobus var. nov.
A var. typico lobis perianthii ad 5, anticis oe postici minus dimi-
nutis, capsula — ut videtur — paullo majore discedit
Mexico: Chiapas: East of Monserrate, Purpus 10066 (Herb. Univ. Calif.).
This variety is discussed under C. gynopetalus Croizat.
Croton Shreveanus sp. nov.
Frutex ca. 2-metralis, innovationibus apice hispido-lanulosis citius gla-
bratis. Foliis griseo- viridibus longe ovatis vel elliptico-ovatis, 5-12 cm.
longis, 2.5—6.5 cm. latis, stipellis anticis 2—4 (neque glandulis more gene-
perianthio ca. 3 mm, lato, 2.5-3 mm. longo, petalis sepalisque subaequi-
longis, staminibus ca. 15, filamentis basi valde lanulosis. Floribus ¢:
1945] CROIZAT, EUPHORBIACEAE FROM THE AMERICAS 185
perianthio minuto, vix 1.5—2 mm. lato, 1-1.5 mm. longo, lobis valde acutis
triangularibus integris vix accrescentibus, petalis minimis ligulatis, ovario
globuloso ca. 1.5 mm. magno, albicante, hispidulo, stylis 3 ca. 5 mm.
longis, cruribus ad tertium inferum partitis.
Mexico: Sinaloa: Capadero, Sierra Tacuichamona, 3000 feet above sea-level,
“slender spreading shrub 2 m. high,” Gentry 5580 (Herb. Univ. Calif.; duplicate in
Herb. F. Shreve).
This shrubby species belongs in the affinity of C. adspersus Benth., which
also occurs in western Mexico but ranges southward. It is a very distinct
entity, easily recognized by the peculiar stipellae of the limb, reminiscent
of the appendages of certain species of Alchornea Sw., but very seldom
found in Croton. It is not C. Roxanae Croiz. from Nayarit, Sinaloa and,
possibly, Guerrero, which is directly allied with C. sphaerocarpus H. B. K.,
and C. fragilis H. B. K.
Croton sancti-lazari sp. nov.
Fruticulus dioecius intricatus, innovationibus tomento stellato hispido
pallide citrino vel aurantiaco parcius indutis, serius glabratis. Foliis ellipti-
cis vel sublanceolato-ellipticis 1-2 cm. longis, 0.3—0.9 cm. latis, integerrimis,
apice plus minusve late rotundatis, basi rotundato-cuneatis, subtus cinereis
vel pallide luteis hispido-tomentosis, supra viridibus tomento stellato dis-
sito glabratis, tenellis toto hispido aurantiacis vel fulvescentibus, haud
lepidotis, nervis adscendentibus ca. 5—7-jugis, petiolo brevissimo 3 mm.
longo vel minore, glandulis nullis, stipulis subcarnosis, triangularibus. Inflo-
rescentiis spicato-capitatis aurantiacis vel lutescentibus, pro more oligan-
this. Floribus ¢: perianthio hispido-tomentoso ca. 2 mm. lato, 1.5 mm
longo, lobis triangularibus, petalis glabris subaequilongis, staminibus ca. 10.
Floribus 9 : perianthio vix 2 mm. longo latoque, lobis acutis triangularibus,
petalis (videtur) nullis, ovario trigono subglobuloso dilute aurantiaco
grossius tomentoso ca. 2 mm. longo, 1.5 mm. lato, stylis 3 carnosullis, ca.
22.5 mm. longis, quove fere ad basem partito, semine ovato apice sat
acuminato ca. 3 mm. longo, 2 mm. lato.
Mexico: Coahuila: South of Castafios, rocky slopes of El Puerto de San
Lazaro, Wynd & Mueller 155 (type 6, Herb. Arnold Arb.; @ Herb. F. Shreve) ;
Coahuila-Chihuahua boundary: Sierra Almagre, “bushy 1-2 feet tall, on floor
of open canyons,” Johnston & Mueller 1141 (Herb. Gray).
A very strong species, related, but not closely, to C. hypoleucus Schlecht.
and its group. It suggests in certain of its vegetative parts dwarfed states
of C. fruticulosus Torr. The absence of lepidote hairs throughout well
separates C. sancti-lazari Croiz. from somewhat similar early spring growth
forms of C. neomexicanus Muell.-Arg., as represented by M. £. Jones
29177 collected near Laredo, Texas.
Croton lotorius sp. nov.
186 JOURNAL OF THE ARNOLD ARBORETUM (VOL. XXVI
tiis gracilibus spicatis 2-sexualibus ca. 10 cm. longis. Floribus ¢ : perian-
thio ca. 1.5-2 mm. lato, staminibus paucis, pedicello ca. 1 mm. longo.
Floribus @: perianthio inter generis minimo vix 1.5—-2 mm. longo sub-
campanulato, pedicello ca. 0.5 mm. longo; calyce hispidulo, fere ad basim
partito, lobis integris apice virescentibus, ovario globuloso hispidulo albi-
cante, stylis gracilibus ca. 2 mm. longis partitis. Caetera desunt.
GuateMara: Huehuetenango: Between Santa Ana Huista and Rancho Lucas,
Sierra de los Cuchumatanes alt. 800-900 m., Steyermark 51332.
This plant is used in baths and is locally known as “‘sanalotodo,” a name
frequently given to herbs supposed to be panaceas. It suggests C. trini-
tatis Millsp., but that is a strictly herbaceous plant, while C. lotorius Croiz.
is manifestly woody, and differs further in the characters of its indumentum.
Croton pyriticus sp. nov.
triangulari-ovatis 4—9 cm, latis, 4-11 cm. longis, apice abrupte breviterque
ca. 6-jugis, primo jugo majore ramoso, glandulis nullis, petiolo 1.5—6 cm.
longo, stipulis nullis. Inflorescentiis apicalibus lateralibusque spicatis ad
10 cm. longis, interdum 1-sexualibus paniculato-ramosis. Floribus ¢ ca.
4 mm. longis, 7 mm. latis, lobis late ovatis petalis subaequilongis, stamini-
bus ca. 15, Floribus @ ca. 7 mm. longis, fere totidem latis, perianthio ad
basem partito, lobis late ovatis abrupte acuminatis ca. 2.5 mm. latis margine
indumento molli sulphureo tectis dorso leproso-lepidotis, petalis obovato-
ligulatis ad 2 mm. longis, aeque ac lobis margine indutis, ovario globuloso
ca. 2 mm. magno, subargenteo-lepidoto, evoluto sub lente gravi nonnihil
verrucoso, stylis pluri-partitis carnosulis ad 4 mm. longis, semine (e David-
son 865, in capsula ad specimen tantum solutum) 15 mm. longo, 10 mm.
lato, antice ad latera compresso lineolis 3—4 e caruncula ortis obliquis
utrinque exarato, rubro-brunneo maculoso-striato, postice nonnihil carinato,
cocco (delapso) ad 19 mm. longo, epicarpio dissite verrucoso.
Costa Rica: Vicinity of El Alto R. R. Station on road to Cartago, alt. 1550 m.,
Allen 661 (TyPE); slope of Cerro Carpintera above La Unidén de Tres Rios, alt.
1350-1500 m., Dodge & Goerger s.n.; Alajuela: San Juanillo, alt. 1525 m., A. Smith
2756; same locality, alt. 1400 m., “Pacific tropical,” A. Smith 10008. PANAMA:
Chiriqui: Boqueli, alt. ca. 1300 m., Davidson 865.
In a previous paper, in Field Mus. Publ. Bot, 22: 448. 1942, I provision-
ally referred this to C. eluterioides Lotsy. A careful study of all the species
of this group so far known from Alabama to Costa Rica, undertaken on
the basis of capsules and seeds, has eventually shown that Lotsy’s species is
not represented.
The forms in this affinity belong to Sect. Eluteria Griseb., and are char-
acterized by a more or less densely lepidote foliage, and petaliferous @
flowers. Mueller-Arg., Schlechtendal, Baker and other authors have treated
this section in a very unsatisfactory manner, which is understandable,
considering that with few exceptions the species under it cannot be identi-
fied from characters other than those of the capsule and seed.
1945] CROIZAT, EUPHORBIACEAE FROM THE AMERICAS 187
Croton argyranthemus Michx., extending from Georgia to Texas, sug-
gests Sect. Eluteria Griseb. in some respects, and might ultimately prove
to be an extreme depauperate form of this group. The species of Sect.
Eluteria Griseb. which are all woody shrubs or trees, more or less densely
lepidote and silvery, their @ flower petaliferous, styles usually several times
dichotomous, and are natives of the region extending from Alabama to
Panama may be separated as follows:
Seed 11 mm. long, 6 mm. broad, or larger.
Epicarp at maturity densely muricate with su
7 mm. broad, caruncle apical, conspicuous, testa ungroo
Tila aa Y UGA CAM = COIADAS sce cis eceser cig: ars oiud oa Suarulene oy ode aeeueeae
Epicarp at maturity with scattered subgranular warts; seed 14 mm. long, 9-10 mm.
b aruncle frontal, inconspicuous, testa with 3-4 shallow grooves radiating
down ward from the caruncle: Panama to peers Rica, possibly northward to
C. pyriticus Croizat.
ubwoody processes; seed 11 mm. long.
oved: Eastern Guate-
C. eluterioides Lotsy.
Seed not over 10 m ong, 5 mm. broad.
Pedicel af ak eae beareely or not at all woody, (1.5)—2 to 3.5 cm. long.
aves definitely elliptic, earns with 8-14 or more pairs of subparallel
very broadly spreading veins; seed 9-10 mm. long, 4-5 mm. broad; epicarp
at maturity with few ed warts: Vera Cruz and Chiapas, Mexico, to
Venezuela, Ecuador, and the West Indies generally............ C. nitens Sw.
— hear tied ovate to lanceolate-ovate, with no more than 6-7 pairs of
easy veins ; seed about 7 mm. long, 4-5 mm. broad; epicarp at maturity
as in C. nitens Sw.; Central and sae ess Guatemala.C. eeaee Lotsy.
Pedicel of fruit ae to sibpodds almost always stiff, less than 1.5(-2 . long.
Leaves ee obtuse to short-acuminate, veins conspicuous, blade a
oe y silvery o ashen; seed ellipsoid about 6 mm. long, 4.5 mm. broad:
A SORTER Oe ee er re C. alabamensis A. E. Smith.
Leaves cere te to lanceolate-elliptic, seldom ovate-cordate, veins weak or
in ;
spicuous, blade underneath as in C. pseudochina Schlecht. & Cham.; seed
scaraboid to sublenticular about 6 mm. long, 5—5.5 mm. broad: ge
Guerrero: tO) SONOta. = see cet ee ea seen C. reflexifolius H. B.
The specific limits in this group are not all evident in the same ne
Careful field studies and experimental cultivation are needed to elucidate
the status of puzzling forms which gravitate around C. pseudochina
Schlecht. & Cham., and C. reflexifolius H. B. K
Croton flavescens Greenm. var. Brandegeanus var. nov.
A typo seminibus 4.5—5 mm. longis, neque 7-8 mm. longis recedit.
Sonora: Between Mazatan and Matapé, 4 miles north of Macori,
Wiggins & Rollins 391.
Croton flavescens Greenm. in Proc. Amer. Acad. 39: 81. 1903, and its
forms constitute a complex, ranging from Michoacan, possibly farther
south, to central Sonora. All these forms are more or less closely related
to C. fragilis H. B. K., and C. limnocharis Croiz., but can be distinguished
from these two species by the keel of the ovary and mature capsule, which
is narrowed and prominent in C. flavescens Greenm., rounded off and quite
188 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
obscure in the other species. The entire group is most controversial, and
should be subjected to experimental cultivation.
Here also belong: Brandegee s. n., Culiacan, Sonora; Rose, Painter &
Rose 9413, Guerreo, Iguala, and ome & Mexia 5164, Sinaloa, Labradas.
The type- -collection of C. flavescens Greenm. is Pringle 8667, Michoacan:
Volcanic hills, Monte Leén Station, 5000 feet above sea-level.
Croton glandulosus L. var. septentrionalis Muell.-Arg. in DC. Prodr. 15(2): 686.
: Ferg. in Missouri Bot. Gard. Rept. 12:51. pl. 17. 1901, p.p. descr. emend.
Croton glandulosus L. var. angustifolius Muell.-Arg. loc. cit.; Ferg. op. cit. 53. pl.
16. f. 2. syn. nov.
Semine ca. 3.25 mm. longo, 2.75—3 mm. lato (inde primo intuitu scara-
boideo neque ellipsoideo), caruncula in speciminibus siccis subventrali
(i. e., seminis a dorso spectata haud manifesta), arillo in semine maturo
saepius brunneo badiove. Foliis saepius (saltem ad ramorum floriferorum
radicem) acuminatis.
Lectotypus: Arkansas, Little Rock, Engelmann 6, 1835 (in Herb. Missouri Bot.
rd.)
Engelmann 6 is the lone specimen out of the six cited by Mueller-Arg. at
publication which perpetuates the epithet septentrionalis in connection with
the form currently identified as such in the literature. I cannot find the
slightest differences by which to separate Engelmann 6 from Lindheimer
172, Ferguson’s presumed lectotype of C. glandulosus L. var. angustifolius
Muell.-Arg.
Croton glandulosus L. var. parviseminus var. nov.
Semine ca. 3.25 mm. longo, 2.25 mm. lato (inde primo intuitu ellipsoideo
neque scaraboideo), caruncula in speciminibus siccis subapicali (1. e., semi-
nis a dorso spectata manifesta), arillo pallido maculis obscurioribus parcius
notato
aici Lindheimer 691b (Herb. Gray).
This variety, which better specimens and experimental cultivation may
yet suggest to be better treated as a form, appears to be delicate, sparingly
branched and subumbellate in habit. It does not seem to be frequent, for
of it I have seen only three specimens, Reverchon s. n., Buzzard Springs,
Texas; Sister Mary Clare s. n., Applewhite Rd. 18 miles South of San
Antonio, Texas; Sister Mary Clare 291, Somerset Rd. 13 miles southwest
of San Antonio, Texas. The type-collection was originally included by
Mueller-Arg., in DC. Prodr. 15(2): 686. 1866, among the specimens cited
as representing C. glandulosus L. var. septentrionalis Muell.-Arg.
Croton glandulosus L. var. pubentissimus var. nov.
Perennans, e loco natali a basi conferte ramosus subdumosus ca. 30 cm.
altus, 50-60 cm. latus, cultus laxior at ramorum dispositione idem. Indu-
mento subhispido copioso, foliorum lamina utrinque bene induta, pilis inter-
dum adpressis habitu subsimplicibus. Perianthii lobis sub fructu quam
capsula longioribus, semine ovoideo ca. 4 mm. longo, totidemque lato.
Texas: Kleberg Country, Padre Island, Cory s. n., 1940.
This is a strong variety, quite unlike C. glandulosus L. var. hirtus
(L’Hér.) Muell.-Arg., which it somewhat suggests in the abundant pubes-
1945] CROIZAT, EUPHORBIACEAE FROM THE AMERICAS 189
cence. It certainly differs from C. glandulosus L. var. Lindheimeri Muell.-
Arg., which is a more delicate plant, with short indumentum, well repre-
sented by the material of Martindale and Parker, collected at the Phila-
delphia Navy Yard, and erroneously identified by Ferguson (in Missouri
Bot. Gard. Rept. 12:52. 1901) as C. glandulosus L. var. scordioides
(Lamck.) Muell.-Arg. Croton glandulosus L. var. Simpsonii Ferg. which
may not differ from C. arenicola Small, has none of the characters of this
new variety.
Croton yavitensis sp. nov.
Arbuscula ad 7 m. alta innovationibus grossius stellato-tomentosis pal-
lide luteis ochraceisve. Foliis fere exacte ellipticis 6-15 cm. longis 2—6 cm.
mote obscureque dentato-serratis, petiolo ca. 5 cm. longo, stipulis subnullis.
Floribus ¢ ignotis. Floribus 9? ‘sub fructu tantum visis sat mancis: peri-
anthio 1 cm. minore lobis 5 integris triangularibus haud accrescentibus,
petalis abortivis nigricantibus vix 1 mm. longis; capsula sub-3-dyma
cylindrico-truncata ca. 1.5 mm. magna indumento pallide luteo fere
eradiato scabrida, stylis Sapa iterum partitis involutis, semine ellipso-
ideo ca. 1 cm. longo 0.8 cm. crasso, brunneo haud nitido, caruncula mani-
festa antica nempe crinito- ane
VENEZUELA: Amazonas: Yavita, Williams 14029.
This is a strongly marked species of uncertain affinities.
Moacroton Croizat
Moacroton gen. nov
Frutices vel arbors humiles e Cuba orientali, foliis coriaceis supra nitidis,
subtus lepidibus plus minusve confertis adpressis tectis. Flore ¢ : perian-
thio in serie duplici 3—(5)-mero, petalis sepalis subsimilibus, pro ratione
affinitatis valde diminuto: antheris 3—6 subsessilibus, scilicet in apice fila-
menti brevissimi potius pro glandula salutandi impositis, in anthesi sub-
Porivontalibus. staminodiis nullis. Flore @ : perianthio 4—(5)-mero laciniis
integris erectis vel suberectis, glandulis petalisve nullis: capsula coccorum
dorso acutata: stylorum cruribus more proprio abbreviatis, integris vel
3—4-lobulatis: semine fere Crotonis L. in affinitate C. reflexifolii Kunthiani.
Genus optime a Crotone L. discedit: (1) Habitu proprio; (2) Natura
androecii a genere Linneano omnino aliena, flore ¢ caeterum minimo;
(c) Stylorum fabrica in Crotone L. haud obvia. Genus sistit relictum in
regione Cubana formis peculiaribus celebrata. In systemate ad Julocrotonem
Mart. inserendum,
Species typica: Moacroton Leonis Croiz.
This new genus keys out as follows:
A) ¢ flower saereniay
a) Filaments filifor at least elongate and cylindric; anthers introrse in anthesis,
parallel with ae center-line of the flower; stamens always more than 6.......
b) Filaments very short, subglandular club-shaped; anthers almost horizontal in
anthesis, perpendicular with the center-line of the flower; stamens 3-6........
Moacroton Croiz.
190 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
B) é flower apetalous.
a’) Stamens free... ..... ce cece eee eter cere terete eee reee Lasiocroton Griseb.
b’) Stamens in a column........... eee eee eee eee eens Leucocroton Griseb.
This assemblage is remarkable on account of its genera differing mainly.
if indeed not wholly, in the characters of the ¢ flower. Its species, how-
ever, can be separated only from the characters of the ? perianths. Great
caution is therefore to be used, lest the ¢ flowers be overlooked as having
slight significance.
Moacroton Leonis sp. nov. PL. I, Fics. 5, 6.
Frutex videtur, innovationibus brunneo-furfuraceis. Foliis primo intuitu
supra nitidis, glaberrimis pico olivaceis valde coriaceis, margine bene revo-
lutis, costa medio supra impressa, subtus prominente, nervis lateralibus
nullis. Lamina longe elliptica, interdum vix ad medium parum dilatata, ad
6 cm. longa, 0.7 cm. lata vel minore, indumento supra nullo subtus lepidi-
bus minimis valde sparsis, petiolo sat crasso ca. 0.5 cm. longo, glandulis ad
basem limbi ipsissimam 2 pro more lente caute inquirendis, stipulis ari
obsoletis. Inflorescentiis mihi adhuc obviis 1-sexualibus spicatis, ad 4—
cm. longis, more proprio simul cum innovationibus prorumpentibus, ae.
propter aegre pro apicalibus lateralibusve salutandis, revera exquisite inter-
calaribus (vide Croizat in Bull. Torrey Club 70: 496- 509. 1943), rachide
crasso, floribus 6 omen paucis in axilla squamulae valde ciliatae.
Flore ¢ minimo qua mm. minore pedicello gracili ca. 3 mm. longo:
sepalis 4, petalis 3 ae re pellucido-puncticulatis: androecio 3-mero
antheris in apice filamenti clavati incrassati brevissimi. Flore @ in situ
haud viso (adest fructus juvenilis tantum solutus, qui diligenter cum
M. trigonocarpi (Griseb.) Croiz. comparatus, huic valde similis evadit) :
capsula 8 mm. longa, laevi, ca. 6 mm. lata, coccorum dorso carinato, hinc
inde lepidibus sparsis ornata: lg more generis brevissimo, vix 1 mm,
longo, cruribus apice 1—3-lobulat
Oriente: Region of <— ichiaawa 400 m. high between the Rio Cabanas
and the Rio Moa, growing on limonite, Marie-Victorin & Clément 21735 (TyPE) ; same
region, “charrascal” on serpentine at Playa de la Vaca, Marie-Victorin & Clément
21774
It proves impossible under the present conditions to study the types of
the species of Urban. However, the descriptions suggest that my new
species is neither M. Ekmanii (Urb.) Croiz. nor M. cristalensis (Urb.)
Croiz., which do not have narrowly elliptic leaves rounded or truncate at
the tip. Nothing like my new species, likewise is to be found in Carabia’s
monographic study (Carib. Forest. 3: 114-135. 1942). The specific name
honors Hermano Leén (Dr. Joseph Sylvestre Sauget y Barbier) of the
Colegio de la Salle, Vedado, Havana, Cuba.
The inflorescence here described, if constant (which might well be the
case), would constitute one more character further to differentiate Moacro-
ton Croiz. from Croton
Moacroton trigonocarpus (Griseb.) comb.
Croton trigonocarpus Wright ex Griseb. i in ee 7 gone i. 1865, No. 7:
17 March 1Sth. 1865; Muell.-Arg. in DC. Prodr. 15(2): 576. 1866; Carab. in
Carib. Forest. 3: 125. 1942
1945] CROIZAT, EUPHORBIACEAE FROM THE AMERICAS 191
Moacroton Ekmanii (Urb.) comb. no
Croton Ekmani Urb. Symb. Antil. ‘9: 194. 1924; Carab. in Carib. Forest. 3: 125.
1942, non Leonard 1927.
Moacroton cristalensis (Urb.) comb. n
Croton cristalensis Urb. Symb. pecs io: 197. 1924; Carab. in Carib. Forest. 3: 125.
1942.
Urban appears to have overlooked Croton trigonocarpus Griseb., and it
is not impossible that Croton Ekmanii Urb. and C. cristalensis Urb. may
prove to be other than forms or varieties of Grisebach’s much earlier species.
I have seen authentic material of the latter, which patently differs from
M. Leonis Croiz., but none of Urban’s specimens. ‘The transfers to Moa-
croton of all these entities is effected, consequently, as a matter of course,
pending a final revision of the entire group.
Argythamnia P. Browne
I see no reason to maintain Ditaxis Vahl ex Juss. as a genus distinct from
Argythamnia. Accordingly, I treat the latter as Argythamnia P. Browne
subg. Ditaxis (Vahl ex Juss.) comb. nov.
Argythamnia coatepensis (Brand.) comb. n
Croton coatepensis Brand. in Zoe 5: 249. cae
The holotype, Purpus 2827 (in herb. Univ. Calif.) has a 3 flower with
the technical characters of Argythamnia, that is, the staminal column is
single-whorled with no more than 5 stamens. In all other characters it is
very difficult to separate this plant from the numerous polymorphic states
of Argythamnia guatemalensis Muell.-Arg. (A. tinctoria Millsp. in Field
Mus. Bot. 1: 303. 1896, syn. nov.) which ranges from Central America to
eastern and western Mexico and Venezuela (Potter 5145, Guanta, roadside
on the outskirts of the city, Herb. Gray)
Argythamnia micrandra sp. nov.
Fruticulus vix pedalis, radice videtur annua, pilis malphighiaceis sat
longis ad caules indutus, parce ramosus.: Foliis ellipticis vel obovato-
ellipticis 2—3.5 cm. longis, 1-2 cm. latis apice breviter acuminatis basi
longe cuneatis, ultra medium dentato-serratis subtrinerviis, aeque ac
caulibus indutis, petiolo ca. 1 cm. longo, stipulis lanceolatis ad 3-4 mm
longis. Perianthio ¢ vix 2 mm. lato, sepalis 4, petalis setaceis minimis,
columna staminali 9-andra vix 1 mm. alta. Perianthio @ ad 5—6 mm. sub
fructu vix maturo lato, sepalis petalisque 5, petalis glandulosis minimis
subnullisve, ovario in anthesi vix 0.75 mm. magno pilis erectis longe hispido,
stylis minutis bipartitis.
Mexico: Guerrero: Coyuca, Cutzamala, Hinton 6307 (Herb. Gray).
A very distinct, apparently weedy species. The 2-whorled minute
staminal column, with about 9 stamens, is the technical character that
places this entity under Argythamnia subg. Ditaxis (Vahl ex Juss.) Croiz.
Alchornea Swartz
Alchornea orinocensis sp. nov.
Arbuscula ad 10 m. alta, innovationibus rachidibusque parcius puberulis
glabratisve. Foliis penninerviis basi antice optime stipellatis, elliptico-ovatis
192 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
6-21 cm. longis 4-11 cm. latis apice breviter acuminatis basi rotundato-
cuneatis, margine dentato-serratis serraturis apice incurvis glandulosisque
ad 3 per cm., nervis ca. 7-jugis, trabeculis evidentibus, glandulis maculosis
secus s costam. in axillis nervorum positis, petiolo sat gracili ca. 10 cm. longo,
stipulis subnullis. Floribus ignotis. Fructu capsulari epicarpio tenui sublaevi
vix granuloso, coccis delapsis ad 8 mm. longis, columella ca. 5 mm. longa,
semine fere exacte ellipsoideo ad 6 mm. longo, 4.5 mm. crasso subcylindrico,
ad chalazam fere truncato ad micropylem abrupte acuminato, arillo striato-
rimoso brunneo subnitido.
VENEZUELA: Amazonas: Alto Orinoco, Tamatama, Williams 15833 (TYPE);
same locality, Williams 15087.
The strictly penninerved leaves, and the well-marked stipellae at the
anterior face of the blade near the insertion of the petiole, as well as the
cicatricose glands along the midrib are characteristic.
Gitara Pax & Hoffmann
Gitara panamensis sp. nov.
Frutex vel arbuscula videtur, innovationibus hispidulis, Saehage _—
= rimoso, habitu toto rigido. Foliis obovato-lanceolat 15 cm
longis, 2.5—6 cm. latis tenuiter chartaceis, glabrescentibus, penninerviis,
8 nervis oe adscendentibus ca. 8-jugis, basi cuneatis vix vel haud
truncatis, margine ad tertium inferum mediumve bene dentato-serratis
(dentibus ad 7-8), apice abrupte sat longe caudatis, petiolo hispidulo ca.
0.5 cm. longo, stipulis membranaceo-scariosis = incrassatis, triangulari-
acuminatis, margine integris, adpressis ad 5 mm. longis. Inflorescentiis
(videtur) lateralibus spicatis 1-4 cm. longis, mono- -dioecis, oe 2
apicalibus terminalibusve. Perianthio ¢ simplici delicato ca. 3 mm. lato
2 mm. longo, ca. 5-lobo, staminibus ca. 30, 1.5 mm. longis, cio
penicillato, staminodiis glandulisve nullis. Perianthio @ 6-mero lobis
lanceolatis acuminatis integris anthesi peracta ca. 4-5 mm. lato, ovario
gap eon e ca. 5-6 mm. lato hispido, stylis 3 basi in ric mc coalitis
. 5-10 mm. longis subplumosis, een nullis.
aig Hills between Pinogana and Yavisa, Pittier 6543, 1914.
Gitara Pax & Hoffm. is a segregate from Tragia L., which latter consists
of a multitude of forms as yet scarcely understood. Its technical char-
acters strongly suggest Angostylis Benth., to judge from the descriptions
and illustrations I have so far seen, but it is otherwise easily mistaken for
Argythamnia P. Browne, having been originally identified, distributed as
representing the latter genus. I retain Gitara Pax & Hoffm. as a genus for
the present
Gitara venezolana Pax & Hoffm. has ecaudate leaves much narrowed at
the tip, with truncate bases, the venation appearing as if markedly 3-nerved.
In these characters, and, possibly, the smaller @ flower, Pax & Hoffmann’s
species does not agree with mine.
Cunuria Baillon
Cunuria (?) casiquiarensis sp. nov.
Arbor ad 6 m. alta. Foliis firme coriaceis ellipticis 11-18 cm. longis, 7—10
cm. latis apice obtuse rotundatis, basi rotundato-cuneatis, margine inte-
1945] CROIZAT, EUPHORBIACEAE FROM THE AMERICAS 193
5 mm. longa, apice abrupte incrassata.
VENEZUELA: Amazonas: Alto Casiquiare, Capihuara, Williams 15690.
The genera in this assemblage are ill defined, and it is difficult to make
identifications unless one has complete material. Cunuria is suggested by
the intangibles of habit, and the characters of the foliage, but the inflores-
cence is somewhat unconventional, when compared with that of C. Spru-
ceana Baill., the standard-species. Conceveiba Kl. and Concevetbastrum
(Muell.-Arg.) Pax & Hoffm. appear to ruled out of consideration by their
foliage, and the manifest glands at the base of the @ perianth.
Sebastiania Sprengel
Sebastiania pusilla sp. nov.
Fruticulus glaberrimus parte epigea, ut videtur, vix ultra palmari, ramis
emortuis persistentibus intricato-ramosus. Foliis statu juvenili tantum visis
carnosulis penninerviis pro more obovatis ca. 1 cm. longis totidemque latis
margine incrassatis sat obscure denticulato-serratis omnino exaristatis,
nervis 3—4-jugis, limbo toto eglanduloso apice obtuse acuminato vel retuso-
truncato basi longius cuneato petiolo glanduloso canaliculato apice saepius
laeviter papilloso ca. 1 cm. longo, stipulis papillosis triangulari-lanceolatis
ad 0.5 cm. longis. Inflorescentiis terminalibus, basi flores 1-2 @ gerentibus
nec ultra 2—2.5 cm. longis simplicibus spicatisque. Flore ¢ vix 1.25 mm.
longo latoque in axilla squamulae basi glandulis 2 baculiformibus ornatae
singulo: perianthio subintegro cupulari, staminibus subsessilibus 3. Flore @
vix 1.5 mm. longo, totidemque lato: perianthii lobis 3 late ovatis margine
erosulis, glandulis obovoideis simplicibus subsimplicibusve cum lobis alter-
nis, ovario in carinis corniculato, stylo fere ad basem 3-partito, cruribus vix
papillosis simplicibus recurvis, semine ca. 1.75 mm. longo, 1.5 mm. lato,
carunculato, pallide ochraceo-brunneo laevissime reticulato.
Urucuay: Paysandu, Chapicuy, banks of the Rio Uruguay, Sta. Sofia, Rosengurtt
B-4169.
This is a remarkable species agreeing with none of the sections in the
genus, and certain to become the type of a new one in the future. The
persistent large petaloid appendages of the @ flower, the corniculate ovary,
the subsessile stamens borne in a subentire perianth are characteristic.
Sapium P. Browne
Sapium contortum sp. nov.
Arbuscula ad 60 cm. alta contorta, ramulis nigricantibus glaberrimis
Foliis subchartaceis ellipticis vel ovato-ellipticis, 3-8 cm. longis 1-3 cm
latis, basi latius rotundatis, apice abrupte acuminatis, margine obscure
.
.
194 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
serrato-denticulatis hinc inde glandulis majoribus auctis, nervis latius
adscendentibus ca. 8—12-jugis, petiolo gracili ca. 1 cm. longo, glandulis
petiolaribus plus minusve elongatis retroflexis, stipulis rotundatis margine
profunde scariosis alabastra vegetativa apice in exo tegentibus ca. 2 mm
longis latisque. Inflorescentiis ¢ tantum visis apicalibus ad 8 cm. longis,
floribus glomerulatis 7-13, glomerulis basi glandulis elongatis 2—3 fultis,
perianthio generis, vix 1.5— ‘2: mm. long
VENEZUELA: Amazonas: Puerto rena Williams 15891,
This species thrives on exposed rocky ledges, and is unusually low-
growing for the genus. It might prove to be a local form of S. Aubletianum
(Muell.-Arg.) Huber, the leaf of which, according to Huber’s figure (in
Bull. Herb. Boiss., sér. 2, 6: 363. f. 23. 1906) is also obscurely serrate and
bears scattered large glands. However, the foliage of S. Aubletianum
(Muell.-Arg.) Hub. should in no case be shorter than 10 cm., and the
character of the leaf margin in this group is not primarily a specific one.
have no material at present representing Sapium guaricense Pitt., S.
naiguatense Pitt., and S. paucistamineum Pitt., but find nothing in the
descriptions to indicate that they may be dangerously close to my new
species. To judge from fragments of Spruce 511, originally determined in
the Paris herbarium as representing S. prunifolium Kl., it seems probable
that Klotzsch’s species is the same as S. Aubletianum (Muell.-Arg.) Huber.
Euphorbia Linnaeus
Euphorbia zerioides Boiss. in DC. Prodr. 15(2):
Euphorbia chiapensis Brandeg. in Univ. Calif. Sy a 6: 54. 1914. syn. nov.
As revealed by a comparison of Purpus 6895: Chiapas, Sierra de Tonala,
and Galeotti 3741, the two species are absolutely synonymous.
Euphorbia segoviensis Boiss. in DC. Prodr. 15(2): 58. 18
Euphorbia Sloanei L. C. Wheel. in Cact. Succ. Jour. 11: fig. 1939. syn. nov.
No difference exists between Andrieux 105 and Palmer 139 which may be
said to be of significance, the two specimens being distinguished only by
details of the pubescence, and the length of the petaloid appendages.
Euphorbia radians Benth. PI. Hartweg. 8. 1839; Boiss. in apie Prodr. ae 74. 1862,
Euphorbia Stormiae Croiz. in Rev. Sudamer. Bot. 6: 13. 1939. syn.
The differences that separate Bentham’s species area my own are seem-
ingly of little account. This species is treated by Boissier, op. cit. 71, 74,
under Sect. Poinsettia, and by Wheeler, in Amer. Midl. Nat. 30: 482. 1943,
under Subg. Poinsettia. A glance at material representing EF. lancifolia
Schlecht. and its allies might suggest that E. radians Benth. probably falls,
on the contrary, under Boissier’s Sect. Dichilium, that is, under Wheeler’s
Subg. Agaloma.
Chamaesyce S. F. Gray emend. Croizat
Chamaesyce arequipensis sp. nov.
Multiceps humilis erecta tota patule hispida, internodiis 1— A cm. longis
vel ultra. Foliis carnosulis ovatis vel rotundato-ovatis 1-1.5 cm. longis, 1
cm. latis sub lente subeveniis 3-nervis, margine subintegris apice tantum
1945] CROIZAT, EUPHORBIACEAE FROM THE AMERICAS 195
obscure lateque dentatis, petiolo 1-2 mm. longo, stipulis obscuris. Cyathiis
in ae soot pedice ello ad 4 mm. longo as late campanulatis ca.
s 1.3 mm. latis glandulis bl nspicue append
latis Bapentice albicante inciso-dentata, ‘laciniis ca. ah ‘ovario ita.
trigono hispidulo albicante, stylorum cruribus 3 fere ad medium partitis,
basi in columnam brevissime coalitis, stigmatibus capitato-globosis; semini-
bus, ats ellipsoideis acuminatis, albicantibus, testa videtur sublaevi.
PER 4 km. from Caraveli on the road to Atico,
alt. fsoo2iono Metcalf 30341. (U. S, a Herb.).
The habit of this new species is reminiscent of that of C. oranenstis Croiz.
of Argentina, and C. Barberiana Croiz. of Paraguay.
Chamaesyce rochaensis sp. nov.
Annua vel pérennans repens tota pilis albidis intricatis sat grosse lanu-
losa caulibus prostratis crassitie fili emporetici minoris vel gracilioribus
internodiis 1-2.5 cm. longis, stipulis interpetiolaribus profunde partito-
ee Foliis bene anisomeris ellipticis rotundatisve emaculatis cum petiolulo
x 1-2 mm. longo ad 1 cm. longis vel brevioribus margine (apice rotundato
pee atini sub lente grossius serratis fabrica haud coriacea. Cyathiis
subsolitariis ob ape abbreviata sub apices sat congestis campanulatis
vix 1 mm. longis pedicello ca. 1-1.25 mm. longo, glandulis appendice peta-
loidea minima praeditis, capsula rotundato-trigona tota lanulosa ad 2 mm.
longa lataque, semine carinato-ellipsoideo maturo pallide brunneo haud
acuminato sublaevi, ae rugis subnullis lente acri caute inquirendis, ad
1.3 mm. longo, 0. 65m ato
RUGU For ie ae Theresa, “rara in uliginosis,” Rosengurtt B-2645
(TYPE) ; same ce Lombardo 2
This is one of the forms ee placed in the herbarium under
Euphorbia thymifolia. It differs, however, from Indian material, such as
Metz (Hoehnacker) 67, cited by Boissier in DC. Prodr. 15(2): 47. 1852,
in its much larger capsules, its more robust habit, heavier pubescence and
larger cyathia. It is not the same as C. oranensis Croiz. and C. portu-
casadiana Croiz., both of which are stouter plants with a different pubes-
cence and other floral characters.
Chamaesyce lutulenta sp. nov.
Annua vel perennans radice recta subsimplici descendente quam par
epigea a Mie majore insignis, cauliculis fili crassitie vel see erate
ad 3-5, vix 2—4-ies partitis, totis, ut visis, nec ultra 5 cm. longis indumento
lanoso ficeene hispidulis. Foliis sat laxe lanosis suborbicularibus aniso-
meris ca. 0.5—0.7 cm. longis totidemque latis parte supera praesertim sub
lente grosse denticulato- serratis emaculosis, petiolulo vix 1 mm. longo
vel minore, stipulis interpetiolaribus late triangularibus denticulato-fissis.
Cyathiis subsingulis purpurascentibus glabratis glabrisve arcte campanu-
latis vix ultra 1 mm. longis, glandulis rotundatis exappendiculatis (interdum
margine albicanti-incrassatis), ovario rotundato-trigono glabro vix 1 mm.
magno, semine ellipsoideo haud acutato nigricante sublaevi.
Florida, a Paes. Estancia Sta. Clara del Dr. Gallinal, rare in moist
localities, Rosengurtt B- 16
196 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
This diminutive new species is strongly reminiscent on the whole of the
North American Chamaesyce Fendleri (Torr. & Gray) Small, but is laxly
hispid-lanose and has rounder dentate-serrate leaves.
EXPLANATION OF PLATE I
Fic. 1. Croton poecilanthus Urb. Section through ¢ ee in bud, showing posi-
tion of rire rs in vernation characteristic of Croton L. Fic. Croton corylifolius
Lam. Section through ¢ flower at anthesis, showing nature = position of the
stamens (only few left) characteristic of Croton L. Fic. 3. Moacroton trigonocarpus
(Griseb.) Croiz., 2 flower. Fic. 4. Moacroton ——— (Griseb.) Croiz —
and stigmas at anthesis seen from above. Fic. S. Moacr n Leonis Croiz., 3 flow
at anthesis seen from above. Fic. 6. Moacroton Leon a ed Stamen seen from oe
side (in slight foreshortening). Fic. 7. Croton seatesha Urb., @ flower; left, a
style. Fic. 8. Croton corylifolius Lam., 2 flower; right, a style
ARNOLD ARBORETUM,
Harvarp UNIVERSITY.
Puiate I
Jour. Arnotp Ars. VoL. XXVI
uP .
OX
y
2.
0
yA
>
EUPHORBIACEAE FROM THE AMERICAS
1945 | METCALF, FLORA OF KUNG PING SHAN, KWANGTUNG 197
NOTES ON THE FLORA OF KUNG PING SHAN, KWANGTUNG
F. P. MetTcaLF*
RECENT expeditions to Indo-China during the years 1936, 1939, and 1940,
undertaken jointly by the Botanical Survey, Lingnan University, and the
Arnold Arboretum, Harvard University, resulted in the collection of ap-
proximately 2800 numbers, each number with abundant duplicates. Of
these, about three hundred and fifty were actually collected in Kwangtung
Province. Kung Ping Shan is located in the extreme western part of
Kwangtung Province, a few miles north of the Tonkin border.
This collection was made by Tsang Wai-Tak during July, August, and
September, 1936, from two localities, Na Leung and Kung Ping Shan, Fang
Cheng District, located on the north side of the river that separates Kwang-
tung from Tonkin, northeast of the town of Tung Hing, Kwangtung, and
about equidistant between the sea-coast of Kwangtung at the Tonkin
border and the Kwangsi boundary.
This region, so close to both the Tonkin (Indo-China) and the Kwangsi
borders, would be expected to have a flora somewhat similar to both these
regions, but an unusual aspect of the problem is the fact that the flora is
apparently more closely related to that of Hainan than to those of central,
eastern, or northern Kwangtung. Of the species recorded in this paper as
new to Kwangtung, about ten were formerly known only from Hainan,
eight were previously known from both Hainan and Tonkin, and ten were
known only from Tonkin.
In this paper notes are given on 32 species that are here first reported
from Kwangtung, representing the following genera: Tacca, Hedvosmum,
Ficus, Artocarpus, Boehmeria, Helicia, Talauma, Fissistigma, Desmodium,
Walsura, Dichapetalum, Elaeocarpus, Flacourtia, Homalium, Clethra,
Diospyros, Anodendron, Xylinabariopsis, Heterostemma, Tabernaemon-
tana, Erycibe, Premna, Gomphostemma, Rhynchotechum, Hedyotis, Lasi-
anthus, Nauclea, Paederia, Uncaria, Wendlandia, and Xanthophytum. Of
these 32 species, about 20 are actually new to China proper. The genera
Talauma and Xylinabariopsis are here first recorded from Kwangtung and
China. In addition, distributional or technical notes are included on
species of Jtea, Prunus, Turpinia, Schizomussaenda, Urophyllum, and Pen-
taphragma.
These notes constitute only a preliminary report, as the novelties, of
which there are about ten, need further study. In addition, some special
groups, such as the Euphorbiaceae, Lauraceae, Caprifoliaceae, and Thea-
*This paper was prepared by Dr. Metcalf on the basis of a Milton-Clark Fund
Grant, Harvard University, to Dr. E. D. Merrill, Director of the Arnold Arboretum,
for use in assisting him in working up the large accumulated collections of Chinese
material at that institution. Other papers will follow.
198 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
ceae are being taken care of by specialists, while the species of Ormosia and
Sabia are reported in Sargentia 3: 1-120. 1943. It is probable that in this
small collection from Kwangtung at least twenty per cent of the species will
represent new records for the province.
TACCACEAE
Tacca Esquirolii (H. Lév.) Rehder, sg? Arnold Arb. 17: 64. 1936
Clerode see Esquirolii H. Lév. Rep. Sp. Nov. 11: 298, 302. 1912; Fl. Kouy-Tchéou
Tacca Pasian Limpr. Pflanzenr. 92(1V. 42): 16. 1928; Pei, Mem. Sci. Soc. China 1:
162. 1932.
arian Kung Ping Shan, Fang Cheng District, along the Kwangtung-Tonkin
border, W. T. Tsang 26661, Aug. 15-24, 1936. New to Kwangtung. Previously re-
corded from Kweichow, Funai, and Tonkin.
CHLORANTHACEAE
Hedyosmum orientale Merr. & Chun, Sunyatsenia 3:36. 1940.
Hedyosmum nutans sensu ap aaa Sci. Jour. 5: 59. 1928; Groff, Lingnan Sci.
Jour. 11: 87. 1930; non Sw
Kwanctunc: Kung Ping . om Cheng District, along the Kwangtung-Tonkin
border, W. T. Tsang 26692, Aug. 15-24, 1936. The first record for Kwangtung and
China proper. Formerly known only from Hainan.
MORACEAE
Ficus siren Reinw. in Blume, wie 470. 1825; King, a Bot. Gard. Calcutta 1:
150, 151, 1888; Hook. f. Fl. Brit. Ind. 5: 525. 1888.
ih TUNG: Kung Ping Shan, a, Cheng District, ue the Kwangtung-Tonkin
border, W. T. Tsang 26829, Sept. 10-18, 1936. The first record for Kwangtung and
China. Formerly known from India to Burma, Indo- China, and the Malay Archi-
pelago.
Ficus sikkimensis Miquel, Ann. Mus. Lugd.-Bat. 3: 225, 292. 1867; ian Ann. Bot.
Gard. Calcutta 1: 89. t. 113. 1888; Hook. f. Fl. Brit. Ind. 5: 521.
egy ia Kung Ping Shan, Fang Cheng District, along the stone Tonkin
border, W. T. Tsang 206856, Sept. 10-18, 1936. The first record si Kwangtung.
we known from Sikkim to the Khasia Hills in India, and Yunna
ado tonkinensis A. Chev. ex Gagnep. Bull. Soc. by France 73:90. 1926;
a Sei. Jour. 6: 275. 1930, op. cit. 7: 303.
ae g Ping Shan, Fang ig — ei the Kwangtung-Tonkin
border, W. T. os peas Aug. 25-30, 1936. Yun Szemao, Henry 13015. The
first record for Yunnan, and the first as a wild Reaier in ius cue Formerly known
from Tonkin and Hainan; cultivated in Kwangtung.
This species seems to be very close to Artocarpus Petelotii Gagnep., which
was described by him at the same time as 4. tonkinensis. Specimens from
Indo-China, determined as representing 4. Petelotit, have very similar but
somewhat more pubescent leaves.
URTICACEAE
re macrophylla D. Don, Prodr. Fl. Nepal. > 1825; Wedd. in DC. Prodr.
6(1): 209. 1869; Hook. f. Fl. Brit. Ind. 5: 577. 1888.
1945] METCALF, FLORA OF KUNG PING SHAN, KWANGTUNG 199
Kwanctunc: Kung Ping Shan, Fang Cheng District, along the Kwangtung-Tonkin
border, W. T. Tsang 26844, Sept. 10-18, 1936. The first record for Kwangtung.
As usually interpreted, this species is reported from India, Indo-China,
and Yunnan. The Yunnan and Indo-China material undoubtedly repre-
sents the same form as this Kwangtung specimen. There is, however, some
question as to whether or not this represents the same species as that de-
scribed by D. Don from Nepal; unfortunately we have seen no material
from the classical locality.
PROTEACEAE
Helicia silvicola W. W. Smith, Notes Bot. Gard. Edinb. 10: 181. 1918.
Kwanctunc: Kung Ping Shan, Fang Cheng District, along the Kwangtung-Tonkin
border, W. T. Tsang 26665, Aug. 15-24, 1936.
This species was previously known only from the type locality, Szemao,
Yunnan. Two specimens of Henry 13075, the type collection, are avail-
able for study; these are in flower. The Kwangtung specimen has mature
fruits, of which a description follows:
Fruit ovoid to ellipsoid, 6-8 mm. long and 4-6 mm. wide, minutely and
closely rufous-puberulent, with persistent style 15-18 mm. long, and short,
rather stout pedicels 4-6 mm. long.
MAGNOLIACEAE
Talauma Candollii Blume, Bidjr. 1: 9. 1825; Hook. Bot. Mag. 72: t. 4251. 1846.
Kwanctunc: Kung Ping Shan, Fang Cheng District, along the Kwangtung-Tonkin
border, W. T. Tsang 26656, Aug. 15-24, 1936.
This represents the first record for the genus from China and from
Kwangtung. The species ranges from Java, Sumatra, and Borneo to the
Malay Peninsula, Siam, and Indo-China. The cited specimen closely
matches a specimen collected by Sargent in Java and Pierre 742 from Cam-
bodia, Indo-China.
ANNONACEAE
Fissistigma polyanthum (Wall.) Merr. Philip. Jour. Sci. 15: 135. 1919; Merr. & Chun,
nyatsenia 5: 59. 1940.
Uvaria polyantha Wall. List No. 6467. 1832, nomen nudum.
Melodorum polyanthum Hook. f. & Thoms. Fl. Ind. 121. 1851; King, Ann. Bot.
Gard. Calcutta 4: 131. t. 172A. 1893.
SAXIFRAGACEAE
Itea amoena Chun, Sunyatsenia 1: 258. 1934, Ic. Pl. Sin. 5:19. ¢. 219. 1937.
Kwanctunc: Na Leung and Kung Ping Shan, Fang Cheng District, along the
Kwangtung-Tonkin border, W. T. Tsang 26510, 26675, July-August, 1936.
This represents the second collection for this province, the type being
from Shap-Man-Tai Shan, a mountain on the Kwangtung-Kwangsi border.
The species is to be expected in Tonkin, Indo-China, as the present collec-
200 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
tion was made only a few miles north of the border. Also known from
Kwangsi (Ching 8059).
ROSACEAE
Prunus Fordiana Dunn, Jour. Bot. 45: 402. 1907; Dunn & Tutcher, Kew. Bull. Add.
Ser. 10: 93. 1912; Groff, Lingnaam Agr. Rev. 2:12. 1924; Merr. Lingnan Sci.
Jour. 5: 87. 1930, in nota, sub Prunus phaeosticta (Hance) Maxim
Kwanctunc: Na Leung and Kung Ping Shan, along the nee Tonkin
border, W. T. Tsang 26588, 26590, 26627, 26741, August, 1936. HAINAN: U
CCC 7742 (McClure) ; Paak Shek Shan, LU 17444 (Tsang) ; Chim Shan, Fung 20182;
Manning and Poting, How 71520, 71653, 72723.
This species was based on a specimen collected by Dunn’s collector at
Sanning, southern Kwangtung. Apparently this is its second collection in
Kwangtung, though the species has been credited to Hainan. It has been
confused with Prunus phaeosticta (Hance) Maxim., both having black-
punctate leaves, but in P. Fordiana Dunn the apex is acuminate, not
caudate-acuminate, the veins are obscure, and the fruit is larger, 8 mm.
long, and ellipsoid rather than globose.
LEGUMINOSAE
asa orn longipes ees) Kew Bull. 1910: 20. 1910; Gagnep. in Lecomte, FI. Gen.
o-Chine 2: 570.
ae as sensu Williams, Bull. Herb. Boiss. II. 5: 20. 1904, non Bak.
(fide Schindler
Desmodium hinhinehse Schindler, Bot. pestis 54: 53. 1917.
Phyllodium longipes Schindler, Rep. Sp. Nov. 20: 270. 1924.
Kwanctunc: Na Leung and Kung Ping Shan, Fang Cheng District, along the
Kwangtung-Tonkin border, W. 7. Tsang 26511, 26702, July-August, 1936; also Sie-
lung, Loting, Y. Tstang 1141, Sept. 14, 1928.
This is the first record for this species from Kwangtung and China.
Formerly known from Siam and from Cambodia, Laos, Cochinchina,
Annam, and Tonkin, in Indo-China. The species has been confused with
both Desmodium pulchellum (Linn.) Benth. (PAvillodium pulchellum
Desv.) and Desmodium elegans (Lour.) Benth. (Phyllodium elegans
Desv.), but it is easily distinguished from both of these by the much larger
bracts (3.5 cm., not 1-1.5 cm.) and the larger leaves. In the size of the
bracts it resembles Desmodium Kurzii Craib, a species of Siam, Burma, and
China (Kwangsi), but it is readily separated from that species by the fact
that the trifoliolate leaves have leaflets that are very variable in size: in
Desmodium longipes Craib the terminal leaflet is very much larger than
the two lateral leaflets, while in D. Kurzii Craib the three leaflets are all
approximately of the same size.
MELIACEAE
Walsura inser Roxb. Hort. Beng. 32. 1814, nomen nudum, FI. Ind. ed. 2,2: 386. 1832:
Pelleg viata Fl. Gén. Indo-Chine 1: 785. 1911; Merr. Lingnan Sci. Jour.
14: a.
KWANGTUNG: ‘ans Ping Shan, Fang Cheng District, along the Kwangtung-Tonkin
border, W. T. Tsang 26690, August 12-24, 1936. The first record 7. Kwangtung.
Formerly known from India to Burma, Indo-China, Yunnan, and Haina
1945 ] METCALF, FLORA OF KUNG PING SHAN, KWANGTUNG 201
DICHAPETALACEAE
Dichapetalum hainanense Nees e) se ee te be Prant!, Nat. Pflanzenfam. 3(4):
348. 1896; Merr. Lingnan Sci. Jour. 5: 105.
Chailletia hainanensis Hance, Jour. Bot. es a on
Kwanctunc: Kung Ping Shan, Fang Cheng District, along Kwangtung-Tonkin
border, W. T. Tsang 26818, Sept. 10-18, 1936
This represents the first record af this species for Kwangtung and China
proper; formerly known only from Hainan. The species is rather close to
Dichapetalum tonkinense Engl., recorded from Hainan and from Tonkin,
Indo-China. It can usually be separated from that species by the fact
that the lower surfaces of the leaves are rather uniformly pubescent, the
pubescence not being confined to the midrib and secondary nerves as in
D. tonkinense Engl. The leaves also average slightly smaller.
ELAEOCARPACEAE
Elaeocarpus hainanensis Oliver in Hook. Ic. Pl. 25: t. 2462. 1896; Merr. Lingnan Sci.
Jour. 1928.
Kwanctunc: Na Leung, Fang Cheng District, along the Kwangtung- ba
border, W. T. Tsang 26636, Aug. 1-10, 1936. Formerly known only from Haina
STAPHYLEACEAE
Turpinia glaberrima Merr. var. stenophylla Merr. & Perry, Jour. Arnold Arb. 22:
552. 1941.
Kwanctunc: Kung Ping Shan, Fang Cheng District, along the Kwangtung-Tonkin
border, W. T. Tsang 26739, Aug. 25-30, 1936. This number is the type collection of
the variety, which also occurs in Kwangsi. The species occurs in Kwangtung, Kwangsi,
Hainan, and Indo-China.
FLACOURTIACEAE
Flacourtia pie Zoll. & Mor. Syst. Verz. 33. 1854; eee Jour. As. Soc. Beng. 59(2):
il WY) ; Merr. Philip. Jour. Sci. 10: Bot. 329. 1915, op. cit. 29: 401. 1926, op.
cit. an: a . 1926, Lingnan Sci. Jour. 5: 132. 46
Kwanctunc: Kung Ping Shan, Fang Cheng District, along the Kwangtung-Tonkin
border, W. T. Tsang 26837, Sept. 10-18, 1936. Indo-China to neat Polynesia,
the Philippines, and Yunnan and Hainan in China. New to Kwangtung.
Homalium Petelotii Merr. Jour. Arnold Arb. 21: 377. 1940.
KWANGTUNG: Na a eung, Fang Cheng District, along the Kwangtung-Tonkin
er, W. T. Tsang 26568, Aug. 1-10, 1936. Formerly known only from the type
collection, Pételot 5983, from Tonkin, Indo-China. New to China and to Kwangtung.
CLETHRACEAE
Clethra annamensis F. Dop, eee Soc. Bot. France 75: 732. 1928, et in Lecomte, FI.
Gén. Indo-Chine 3: 717.
KWANGTUNG: Kung Ping ae Fang Cheng District, along the Kwangtung-Tonkin
border, W. T. Tsang 26761, Sept. 1-9, 1936. Formerly known only from Annam and
Gane tri, Indo-China. Nee to China and to Kwangtung
EBENACEAE
Diospyros potingensis Merr. & Chun, Sunyatsenia 5: 164. 1940.
Kwanctunc: Kung Ping Shan, Fang Cheng District, along the Kwangtung-Tonkin
border, W. T. Tsang 26705, Aug. 25-30, 1936.
The cited specimen apparently represents this species. It differs slightly
202 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
from the type in having proportionately longer and narrower leaves of a
slightly different color, but the color difference may be due to the method
of drying. If it is correctly named, this is the first record of the species
for Kwangtung. Formerly known from Hainan and Hupeh.
APOCYNACEAE
Anodendron punctatum Tsiang, Sunyatsenia 2: 129. 1934, op. cit. 3: 140. 1936.
Kwancrunc: Kung Ping Shan, Fang eg District, along the Kwangtung- -Tonkin
border, W. T. Tsang 26724, Aug. 25-30, 1936. Formerly known only from three col-
lections from Hainan. A species condlily ais by its punctate leaves.
Xylinabariopsis napeensis (Quintaret) comb.
Micrechites napeensis Quintaret, Compt. Rend, yey Sci. 134: hese feet du Jundi
17 Février; received Harvard College eased March 21).
Xylinabaria Reynaudi Jumelle, Rev. Cult. Colon. 11: 228. on
Ecdysanthera napeensis Pierre, Rev. Cult. Colon. 1]: 228.
Parabarium napeense Jumelle, in Spire, Caoutch. Indo- pai 33
sncyaas i Reynaudi Pitard in Lecomte, Fl. Gén. Indo- a. . ae 1933.
Kw Na Leung and Kung Ping Shan, Fang Cheng District, along the
mercer Tonkin border, W. T. Tsang 26523, 26643, July-August, 1936
It is difficult to understand why Pitard used the specific epithet Rey-
naudi, when both Pierre and Jumelle had already accepted the earlier
epithet napeense. Formerly known from Annam, Laos, and Tonkin (Indo-
China), and hence new to both China and Kwangtung.
ASCLEPIADACEAE
Heterostemma oblongifolium Constantin - Lecomte, Fl. Gén. Indo-Chine 4: 120.
1912; Merr. oe Sci. Jour. 5: 153
Kwanctunc: Kung Ping Shan, Fang iin District, along the Kwangtung-Tonkin
border, W. T. Tsang ae Sept. 10-15, 1936. Formerly known only from Laos, Indo-
China, and Hainan, and a new to both the Chinese mainland and Kwangtung.
Tabernaemontana bufalina Lour. Fl. Cochinch. 117. 1790; Pitard in Lecomte, Fl. Gén.
Indo-Chine 3: 1160. 1933; Merr. Trans. Am. Philos. Soc. II. 24(2): 312. 1935
(Comment. Lour. Fl. Cochinch.).
Kwanctunc: Na Leung and Kung Ping Shan, Fang Cheng District, along the
Kwangtung-Tonkin border, W. T. Tsang 26547, 26654, July-August, 1936. Previously
known from Indo-China and Hainan, and hence new to the Chinese mainland and to
Kwangtung
CONVOLVULACEAE
sialase eee Merr. Philip. Jour. Sci. 21: 353. 1922, Lingnan Sci. Jour. 5: 153.
Kwanctunc: Kung Ping Shan, Fang Cheng District, along the Kwangtung-Tonkin
border, W. T. Tsang 26640, Aug. 15-24, 1936. Formerly known only from Hainan
VERBENACEAE
Premna Maclurei Merr. Lingnan Sci. Jour. 6: 330. 1930.
Kwanctunc: Kung Ping Shan, Fang Cheng District, along the Kwangtung-Tonkin
border, W. T. Tsang 26657, Aug. 15-24, 1936. Formerly known only from Hainan
LABIATAE
a re er leptodon Dunn, Notes Bot. Gard. Edinb. 8: gee oo op. cit. 6: 190.
; T. Doan in Lecomte, Fl. Gén. Indo-Chine 4: 1036.
1945 | METCALF, FLORA OF KUNG PING SHAN, KWANGTUNG 203
Kwanctunc: Kung Ping Shan, Fang Cheng District, along the Kwangtung-Tonkin
border, W. T. Tsang 26790, 26825, Sept., 1936. Formerly known only from Kwangsi
(Morse 257) and from Tonkin, Indo-China; new to Kwangtun
GESNERIACEAE
ee ellipticum (Wall.) A. DC. in DC. Prodr. 9: 285. 1845, in nota; C. B
, Commel. Cyrtandr. Bengal. t. 91. 1874 (Rhyncotechum), in DC. Monogr.
me ey 198. 1893; Hook. f. Fl. Brit. Ind. 4: 373. 1884; Merr. Lingnan Sci. Jour.
13:71. 1934
Kwanctunc: Kung Ping Shan, Fang Cheng ere along the Kwangtung-Tonkin
border, W. T. Tsang or Sept. 10-18, 1936. rmerly ot from India and
Hainan; the first record for the Chinese mainland a peeee
RUBIACEAE
Hedyotis obliquinervis Merr. Lingnan Sci. Jour. 14: 56. 1935.
Kwanctunc: Na Leung and Kung Ping Shan, Fang Cheng District, along the
Kwangtung-Tonkin border, W. T. Tsang 26518, 26644, July-August, 1936. Previously
known only from the type collection, Lau 452, from Fung Leng, Ngai District, Hainan,
Additional collections of this species from Hainan are: Lau 1265, Tai Tin Shan, Chang-
kiang District, Liang 62029, Yaichow, Gressitt 960, Ta Hau, and without definite
locality, Liang 64974 and 66230 and Wang 32842. The first record for the Chinese
mainland and for Kwangtung
Lasianthus Koi Merr. & Chun, Sunyatsenia 2: 47. 1934.
Kwanctunc: Na Leung and vicinity, Fang Cheng District, along the Kwangtung-
Tonkin border, W. T. Tsang 26558, July 24-31, 1936. Previously known only from
the type collection, Ko 52243, from Tingnan, Hainan. Liang 54368 and 65280 and
Wang 35807, from Hainan, without definite locality, also represent this species.
The original description was based on a flowering specimen. The follow-
ing description of the fruit is based on Tsang 26558.
Fruit sessile, subglobose, black, subglabrous, 5 mm. long and about 4 mm.
wide, capped by persistent calyx-lobes; lobes of calyx PAeeiate glabrous
toward base, but covered with numerous, prominent, whitish, jointed hairs
above the middle.
Schizomussaenda dehiscens (Craib) Li, Jour. Arnold Arb. 24: 100. 1943.
Mussaenda dehiscens Craib, Kew Bull. 1916: 263. 1916; Pitard in ne Fl. Gén.
Indo-Chine 3: 174. fig. 12, 2-3. 1923; Chun, Sunyatsenia 1: 306.
Kw ung and Kung Ping Shan, Fang Cheng Bini along the
fe che Perkin es ih W.T. Tsang 26513, 26575, 26668, Sale Annee 1936.
While this note was originally prepared to record the species as Mus-
saenda dehiscens Craib as occurring in Kwangtung, Dr. Li’s findings have
been accepted. The species extends from Burma, Siam, and Indo-China
to Yunnan, Kwangsi, and Kwangtung, having first been recorded from the
latter province by Chun, without any citation of specimens, and by Li, who
cites Liang 69546 from Shih Wan Tai Shan. Schizophragma macro-
sepalum Hu (1930) and Emmenopterys Rehderi Metcalf (1932) are
synonyms, according to Dr. Li, both of these genera belonging to the Saxi-
fragaceae.
Nauclea officinalis Pierre ex Pitard in Lecomte, Fl. Gén. Indo-Chine 3: 26. 1922, in
yn.; Merr. & Chun, Sunyatsenia 5: 188. 1940.
Sarcocephalus Er teat Pierre ex Pitard, 1. c.
204 JOURNAL OF THE ARNOLD ARBORETUM (VOL. XXVI
Kwanctunc: Kung Ping Shan, Fang Cheng District, along the Kwangtung-Tonkin
— W. T. Tsang 26867, Sept. 10-18, 1936. Formerly known from Indo-China and
Haina
— gre Blume, mg 968. 1825; Merr. Bibl. Enum. Born. Pl. 580. 1921,
num. Philip. FI. Pl. ies 1923
KWANGTUNG: ee ng Ping “1 ies Cheng District, along the Kwangtung-Tonkin
border, W. T. Tsang 967 51, 26787, 26874, Aug.—Sept., 1936. A widespread species,
known from the ee penton to Java and the Philippines. The first record tor
China and for Kwangtu
Unearia een pa Hutch. in Sargent, Pl. Wils. 3: 406. 1916; Merr. Lingnan Sci.
Jour. 11: 59. 1930; Rehder, Jour. Arnold Arb. a - 1935
pn scandens Sm. in Rees, Cyclop. 39: no. 9.
Uncaria pilosa Roxb. Fl. Ind. ed. 2. 1: 520. 1832; Hook {. Fl. Brit. Ind. 3: 32. 1880;
ard in Lecomte, Fl. Gén. Indo-Chine 3: 47. 19
Kwanxctunc: Kung Ping Shan, Fang Cheng sea along the Kwangtung-Tonkin
border, W. T. Tsang 26833, Sept. 10-18, 1936. Also known from India and Indo-
— and from Yunnan, Kweichow, and Hainan, China. The first record for Kwang-
ng.
Cepia chinense Merr, & Chun, Sunyatsenia 2: 19. 1934.
Na Leung _ Kung Ping Shan, Fang Cheng District, along the
Mccasncesiy Tonkin border, W. T. Tsang 26512, 26770, July and Sept., 1936
This species is the only representative of this genus known from China
proper, and it was formerly known only from the type collection, T's0 23555,
from Sup Man Ta Shan, along the Kwangtung-Kwangsi border. One of
the two collections cited above is in bud; the original description was based
on a fruiting specimen only. A description of the young flowers is here
added.
Buds ovoid, noticeably constricted at the base, glabrous; calyx saucer-
shaped or broadly and shallowly campanulate, 5- ‘lobed, the lobes broadly
ovate- ermine’ with glandular margins; pedicels filiform, about 6 mm.
long, puberulen
re tinctoria DC. ee orientalis Cowan, Notes Bot. Gard. Edinb. 16: 268.
, op. cit. 18: 184.
Meee glabrata ek in part, non DC.
Kwanctunc: Na Leung, Fang Cheng District, along the Kwangtung-Tonkin
border, W. T. Tsang 26542, July 24-31, 1936. Formerly known from India, Siam,
Indo-China, and from Yunnan and Kwangsi in China; new to Kwangtung.
— ferrugineum (DC.) Merr. es Inst. Bot. Hamb. 7: 270. 1937.
etabolos ferrugineus DC. Prodr. 4: 436. 1830.
pemeei tum Villari Vidal, Rev. PI. co Filip. 150
Xanthophytum anaes sensu oe ae Jour. ae ¥ Bot. 328. 1909, Enum.
Philip. Fl. Pl. 3: 492. 1923, non Rei
KWANGTUNG: aa Ping Shan, Fang an District, along the Kwangtung-Tonkin
border, W. T. Tsang 28721, Aug. 25-30, 1936. Kwancsi: Shap Man Taai Shan, near
Iu Shan village, Siangsze Disteiet, along the Kwangtung border, W. T. Tsang 22306,
May 16, 1933. New records for both Kwangtung and Kwangsi, ‘ond also for China
These specimens are associated with the Philippine species, which as
only recently been segregated from the form of Java, Borneo. and Sumatra,
on account of the differences in the inflorescences. The specimens seems to
agree fairly well with the Philippine plant, but when flowering material
1945] METCALF, FLORA OF KUNG PING SHAN. KWANGTUNG 205
becomes available, some other disposition of them may prove to be desira-
ble
CAMPANULACEAE
Ce spicatum Merr. Philip. Jour. Sci. 21: 511. 1922, Lingnan Sci. Jour. 5:
1930.
ies UNG: g Ping Shan, Fang — District, along the Kwangtung-Tonkin
border, W. T. Nene ee Sept. 10-18,
This species was originally based on two specimens, one from Tung Sing,
Kwangtung (K. K. Tsoong 1907), and the other from Ng Chi Leng,
Hainan (CCC 8675 [McClure|). This is apparently the second record of
the species for Kwangtung.
ARNOLD ARBORETUM,
Harvarp UNIvERSITY.
206 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
AN APPARATUS TO MAINTAIN A SURFACE FILM OF WATER
FOR USE IN VEGETATIVE PROPAGATION
Kart A. GROSSENBACHER*
With three text-figures
VEGETATIVE propagation has long been of practical importance to nurs-
erymen and others engaged in work with plants. The development of the
theory of the hormone mechanism of root formation and availability of
synthetic chemicals for treatment of cuttings has greatly increased the
amount of work being done with vegetative propagation. Plant physiolo-
gists are interested in the phenomena of root formation as well as in the
use of cuttings as a source of uniform experimental material. In the study
of genetics of trees vegetative propagation becomes a valuable tool for the
more accurate estimate of individual clones.
Doran (1) presents a summary of the work with trees and shrubs as well
as an extensive bibliography. The standard practice has been to shade or
cover propagating beds and to sprinkle the cuttings to prevent them from
drying out during the period of root formation. Where leafless stems can
be used, there is less evaporating surface and maintaining cuttings is a much
simpler problem. For many species, however, it is necessary to use leafy
cuttings, and their maintenance presents many problems. The water con-
tent must be kept up. They must have sufficient light to function prop-
erly and fungi must be controlled.
Raines (3) reports on a spray chamber technique for handling plant
materials which promises to lead to new developments. Mitchell et al. (2)
developed an out-of-door modification used to maintain cuttings in full
sunlight. They found the most favorable environment for the propaga-
tion of slash pine (Pinus caribaea Morel.) to be “well drained sand. . .
75° —90° F.... fine spray of water ...on 5 minutes out of each 10-minute
cycle for 10 to 12 hours each day.” " Others have pointed out that spray
humidification in greenhouses has extended the limits of propagation of
plants which are difficult to root (5).
Preliminary work in the greenhouses of the Harvard Biological Labora-
tories indicated that cuttings of several forest trees responded better in
fuller exposure to sunlight than in the more conventional shaded beds.
Tests were made with mechanical humidifiers as well as with intermittent
ray. During the summer of 1944 an apparatus was developed and oper-
ated at the Harvard Forest. Taking advantage of full sunlight, out-of-
*This work has been financed by the Maria Moors Cabot Foundation for Botanical
Research of Harvard University and ae carried out at the Harvard Forest. The
author wishes to thank C. T. Brown, Jr., S. H. Spurr, and P. R. Gast for their aid in
the development and construction of this fear
1945] GROSSENBACHER, APPARATUS TO MAINTAIN FILM OF WATER = 207
door propagating beds were equipped with a sprinkler system and an auto-
matic control mechanism. This apparatus was designed to give maximum
surface wetting with a minimum amount of water and was actually con-
trolled by the evaporation of water from a surface film.
Cuttings of various forest trees were maintained in excellent condition.
Satisfactory rooting was obtained with cuttings of red maple (Acer rub-
rum), sugar maple (A. saccharum), and paper birch (Betula papyrifera).
No satisfactory rooting was obtained during the relatively short 6-8 week
period with cuttings of hemlock (Tsuga canadensis), red pine (Pinus
resinosa), white pine (P. Strobus), or white spruce (Picea glauca).
APPARATUS
Propagation beds, as illustrated in Fig. 1, were built with gravel drainage
and heating coils, allowing a 6-inch depth of builders’ sand for planting
cuttings. Sprinkler pipes were placed 10 ft. apart (every two beds) with
flat misting nozzles (#.031F, Spray Engineering Co., Somerville, Mass.)
every 3 ft. They are nected by a motor-driven crank, connected by
rods to lever arms clamped directly on the pipes. The oscillating pipes are
connected by loops of garden hose. The water supply (40 Ib. pressure ) is
controlled by a single solenoid valve. An emergency by-pass with a nor-
mally open solenoid valve has been installed to supply constant spray in
the event of any power failures.
| |
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Fic. 1. Diagram of propagating beds.
The regulator, shown in Fig. 2, consists of a Livingston black bulb
atmometer (B. E. Livingston, Riderwood, Md.) connected to a water-oil-
mercury column. When a small amount of water is lost from the evaporat-
ing surface, the mercury moves up, making contact with the platinum wire
activating the relay. This starts the oscillating motor and opens the sole-
noid valve. The regulator is so placed that water from the sprinklers falls
on the evaporating surface, thereby replacing the water lost and opening
the circuit. A distilled water reservoir is connected into the water column
through a mercury safety valve to prevent undue water tension from de-
veloping in the regulator in case of mishap. By applying air pressure
gently to the distilled water, the walls of the bulb can be conveniently
flushed.
In order to reduce electrical action in the mercury-oil-platinum switch as
208 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
EVAPORATING
SURFACE
APPLY MOUTH
FLUSHING
ATMOMETER
as =>
EN
a\ ane
OIL COLUMN ss
DISTILLED WATER
== RESERVOIR ==
MERCURY
SWITCH
MERCURY =
SAFETY VALVE =
(Co
COTTON PLUG
cy 1944
Fic. 2. Detail of regulator.
1945] GROSSENBACHER, APPARATUS TO MAINTAIN FILM OF WATER — 209
well as to assure uniformity of behavior, a two-coil, locking relay is used
(Cenco #99740). The activator current is drawn directly from the 110-
volt A.C. line with the load, relay coil, motor, and solenoid valve, in series.
A single low amperage impulse (1/120 sec.) is enough to lock the relay in
the on-position. A separate coil and circuit is used to open the relay, and
the necessary impulse is drawn in a similar manner from the A.C. line.
The circuit is closed by a contact point on a wooden idler wheel, which is
driven by the crank shaft of the oscillating mechanism. While the idler
wheel makes one revolution in 30 seconds, the crank shaft makes 5, thereby
giving five sweeps of the sprinklers per 30-second period. The momentum
of the apparatus is sufficient to carry the contact points past the make-
position, opening the off-circuit and clearing the relay for the next starting
impulse. A small amount of chattering occurs if water from the sprinkler
has not opened the starter circuit during the 30-second on-period. With
the regulator placed midway between pipes and well within the range of
several nozzles, this can only happen occasionally in dry windy weather.
This regulator mechanism has two fundamentally weak aspects. The
evaporating surface and porous walls of the atmometer tend to accumulate
deposits of materials dissolved in the spray water. Also, the electric cur-
rent causes sludge to form at the junction of the oil and mercury. Though
satisfactory contact is made when relatively large amounts of sludge have
accumulated, the sludge can be expected eventually to cause trouble. The
present apparatus has been operated for more than three months without
interruption.
PERFORMANCE
The oscillation of the sprinkler pipes can be adjusted to give satisfactory
cover of the beds by changing the lever arms and connecting rods, etc. The
two edge sprinkler pipes are set to supply a single bed with a short arc, thus
giving a heavy watering to compensate for the effects of the wind. The
center pipe swings through a larger arc, watering a bed on each side.
The regulator can be adjusted to give a wide range of performance. In
order to maintain a film of surface water, the mercury in the outer arm of
the mercury switch is put enough higher than the contact point nearly to
balance the weight of water in the column supplying the bulb, thereby main-
taining almost a free water surface on the bulb. Excess water drains off,
and only a slight evaporation is necessary to start the sprinklers.
With this arrangement, the sprinklers turn on every 2 to 2} minutes in
bright dry weather, less frequently in cloudy weather, and only occasionally
during the night. This means that the sprinklers are turned on about 240
times a day, or they run for about 2 hours out of the day, delivering 500
gallons over an area of about 500 sq. ft. (allowing for edge effects). This
is equivalent to a rainfall of about 1.6 inches. However, a large portion
of this water is actually lost by evaporation or blown away by the wind.
Records of sprinkler frequency were obtained with an independent cir-
cuit operating a chronograph. The drainage of water through the propa-
gating bed was measured from three sampling buckets, as shown in Fig. 1.
210 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
Atmospheric temperature and humidity records were obtained in a stand-
ard weather shelter some 75 yards from the beds. Sample data for August
28, 1944, are presented in Fig. 3. The low temperature and high humidity
of the preceding night kept the sprinkler off, and the drainage rate was
therefore very low. As the day advanced, and the temperature rose and
the humidity fell, the sprinkler came on more often, reaching by nine
o'clock a fairly constant rate of 25 to 30 times per hour (highest recorded
rate: 36 times per hour). The drainage for the 9 to 5 period reached the
large amount of .326 cc per cm? per hour, being one of the highest drainage
rates observed. As night came on and the humidity increased, the sprinkler
j ———— 80 90
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Arboretum. The codperating institutions made available the ae facilities and
covered all expenditures of the Committee
~The authors bow to acknowledge the codperation of the United States Plant Intro-
duction Garde en at Coconut Grove, Florida, and that of camoufla age officers in the
Corps of suite and other branches of the United States Army. Among = mem-
bers of the Harvard Camouflage een I. W. Bailey, Ernest Ball, P. R. Gast,
R. J. Lutz, E. D. Merrill (Chairman), K. V. Thimann, and R. H. Wetmore wate freely
of their time and energy to those phases pie by the present report
*Harvard Camouflage Committee. Using cut foliage for santas. 2 manuals,
17 pp. and 19 pp. Illus. Harvard Forest, Petersham, Mass. 1943,
1945] GROSSENBACHER, SPURR, & VLAMIS, CUT FOLIAGE 215
tropical plants tested were lowland species, particularly those native to
coastal regions.
Sprays of cut foliage of a uniform size (6 to 8 feet in Petersham and 3
to 5 feet elsewhere) were exposed on stakes or racks to full sunlight and
all weather conditions. Careful notes of the condition of the plants were
taken several times a day during the critical stages of drying, and daily at
other times until the branches were completely withered. Each species was
tested with at least two treatments: (1) base of branch suspended in air,
and (2) base of branch immersed in water. Every treatment was repli-
cated (five times in Petersham and three times elsewhere) and was repeated
until reasonably consistent results were obtained.
The greatest difficulty encountered in these species surveys was that of
developing a system of rating foliage condition that would give comparable
results throughout the great variety of plants tested. Cut foliage reacts in
many ways to drying out and death. Many plants brown; others, like
Viburnum, blacken; still others, like Eucalyptus and some palms, whiten;
and others, like pin cherry, develop autumn coloration. Some crisp, while
others merely wilt. After initial trials, the following classification was set
up and successfully used in dividing all plants into five broad classes of
Crass I. Normal. No evidence of wilting or injury.
Cuass II. Slight wilt. Essentially normal in appearance but evidence of drying
such as (1) slight drooping of leaves, (2) marked wilt of young growing tips only, or
(3) slight discoloration, usually on undersides of leaves only.
Crass III. General wilt. Foliage wilted, but little or no (less than ten percent)
crisping. Obviously abnormal in appearance, but foliage still essentially green and
soft. Browning largely confined to undersides of leaves.
Crass IV. Partially crisp. Marked crisping and discoloration of foliage, but more
than fifty percent of leaves essentially green and soft
Crass V. Crisp. More than fifty percent of leaves crisp and discolored. Dis-
coloration may consist of browning, blackening, whitening, autumn coloration, or
merely marked fading.
During a given test, all foliage passed from Class I to Class V. At the
time of each observation, a single rating was given for each species unless
the variation was so great that single branches of the same species fell into
different classes. The time necessary to pass through various classes was
different for different plants. This difference might be used to explore
certain physiological properties of species. The dividing line between Class
II and III was taken as the limit of usefulness of plant material to be recom-
mended for camouflage purposes. Selecting of any other point would
result in a different ranking of species.
As a result of these surveys, it was possible to rate the relative lasting
qualities of the different species, both with and without water (Tables 1
and 2). The values given for each plant cannot be considered as exact,
since the lasting qualities of any plant are affected by many factors. The
relative ranking of the different species, however, is reasonably accurate,
and the values presented are a good indication of the number of days each
species will last if exposed to full sunlight in the middle of the growing
216 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
TABLE 1.
LASTING OF CUT TEMPERATE FOLIAGE
Days in sunlight Days in sunlight
without water Species with water
120rmore Picea Abies (Norway spruce) 14-21
Pinus resinosa (red pine) 14-21
Pinus silvestris (Scotch pine) 12-16
Pinus Strobus (white pine) 10-14
6-12 Picea glauca ular spruce) 14-21
Juniperus communis (common juniper) 14-21
2-6 Tsuga canadensis rane 4-10
Larix decidua (European larch) 4-8
Larix Kaempferi (Japanese larch) 4-8
1-2 Malus pumila (apple) 6-10
Ligustrum vulgare (common privet) 6-10
Ligustrum ovalifolium (California privet) 6-10
Vacciniu p. (blueberry) 4-8
Quercus velutina (black oak) 3-6
uercus borealis (red oak) 3-6
Crataegus ean (English hawthorn) 2-4
Acer saccharum (sugar maple 2-3
Acer rubrum (red ple 2-3
Acer campestre (European maple) 2-3
3-1 Syringa vulgaris (lilac) 6-10
Fagus s tica (European beec 5-8
Fagus grandifolia (American beech) 3-6
Quercus robur (Englis 3-6
Quercus petraea (durmast oak) 3-6
Quercus alba (whi — 24
Castanea dentata (c ut) 2-4
Tilia platyphyllos ea scan linden) 2-4
Tilia cordata (small-leaved linden) 2-4
Acer pseudoplatanus (sycamore maple) 2-3
4-3 Prunus pennsylvanica (pin cherry) 3-6
Acer pennsylvanicum (moosewood) 2-3
Ginkgo biloba (gin 2-3
Platanus acerifolia (London plane) 1-3
Fraxinus excelsior (European ash) 1-3
Prunus serotina (black cherry) 1-2
Prunus Padus (European bird cherry) 1-2
Alnus rugosa (smooth al 1-2
Alnus glutinosa (black alder) 1-2
Betula lutea (yellow birch) 1-2
Betula papyrifera (paper birch) 1-2
Betula populifolia (gray ) 1-2
Liriodendron tulipifera (tulip tree) 1-2
Populus nigra (black poplar 3-2
Ulmus americana (American elm) 3-1
Ulmus procera (Engli ) 4-1
Carya ovalis (small pignut hickory) 3-1
1945] GROSSENBACHER, SPURR, & VLAMIS, CUT FOLIAGE
TABLE 1. (Continued)
Days in sunlight Days in sunlight
without water Species with water
0-4 Rhus typhina (staghorn sumac) 3-6
Viburnum cassinoides (withe-rod 3-2
Populus grandidentata (big-toothed aspen) 3-2
Populus tremuloides (trembling aspen 0-2
Fraxinus americana (white ash) 0-3
Juglans cinerea (butternut) 0-2
Robinia pseudoacacia (black locust) 0-2
Salix nigra (black willow 0-2
Salix fragilis (crack willow) 0-3
Salix pentandra (bay willow) 0-3
Sambucus canadensis (elder) 0-4
Ailanthus spp 0-3
season. ‘These species surveys, covering more than one hundred different
kinds of plants of the north temperate and tropic zones, reveal a number of
phenomena, some commonplace and expected; others, rather surprising
Tropical species reacted much the same as temperate ones. The range
in number of days that plants would retain a normal appearance in sunlight
is about the same, both in trials with water and in trials without. Plants
from the tropics, as a group, seemed to respond to water about the same
as temperate plants, a few profiting immensely, most lasting two or three
times as long with water as without, and some incapable of taking up
sufficient water to prolong the fresh appearance. The amount of water
required to keep a cut branch alive is considerably less than the amount
transpired by the same branch before being cut. A cut six-foot branch will
ordinarily require up to a quart or more of water the first day, and decreas-
ing quantities thereafter until the branch is dead. Coniferous foliage
ordinarily requires less water than hardwood foliage, probably because of
its better protection against water loss.
Temperate Zone Foliage
The most obvious generality that may be stated upon the basis of tem-
perate zone tests is the common observation that conifers of all types retain
their normal appearance far longer than hardwoods. The poorest conifer
tested (larch) lasted longer than the best hardwood (apple)
The other obvious tendency is that plants of the same genus tend to have
similar lasting qualities regardless of species, locality, or origin. Certain
genera were particularly consistent in this regard, among them being
Larix, Ligustrum, Alnus, Betula, Ulmus, and Salix. In all cases, variation
within the genus was far less than variation between the gene
Species of several genera were more or less consistent, but yet reacted
differently in various degrees. In some genera, species characteristic of
drier sites — plants whose structure was better adapted to retaining mois-
218 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
ture — apparently remained fresh the longest. Thus, red pine was some-
what better than Scotch pine, which, in turn, was definitely superior to
white pine. A number of European species seemed to be somewhat better
than their American counterparts, possibly because they had developed in
a drier climate. Among these were European beech, European bird cherry,
European ash, and European black poplar. This was not universally true,
however, for the American maples lasted as well as the European hedge
maple, and considerably longer than the European sycamore maple with its
large thin leaves. The foliage of the latter, though, lasted longer than did
the similar foliage of the American moosewood. When supplied with
water, all five maples tested remained fresh for about the same period.
Among the oaks, the northern red oak was superior to the white oaks,
both European and American, when tested in the absence of water. That
the red oak group as a whole has greater lasting powers than the white oak
group was substantiated by repeated tests not indicated in Table 1. With
water supplied, little difference in reaction was noted.
Among the temperate species, the response to added water was relatively
consistent. Practically all species benefited markedly from water except
for a few plants which wilted and withered nearly as quickly with water as
without. The response of staghorn sumac to a supply of water was excep-
tional. Cut sumac foliage would wilt the first day after being placed in
water. Then, surprisingly enough, it would begin to recover. For several
days thereafter, the leaflets would remain turgid, the plant being appar-
ently normal except for a droop of the rachis. No other species tested
showed this recovery from leaf-droop and wilt.
Tropical Zone Foliage
Tropical plants varied a great deal in their capacity to absorb water and
in their lasting properties. Certain genera, such as Casuarina, Ixora,
Eugenia, and Eucalyptus, lasted as much as five to fifteen times as long
with water as without. Many others tested apparently were totally unable
to utilize water (Table 2).
In one respect, tropical species behaved rather differently from temperate
plants. Because of the large size and weight of many of the leaves, abnor-
mal appearance was often first manifested by marked wilting of the petioles
and consequent leaf droop, rather than by a change in the leaf itself. For
instance, the leaves of the India rubber tree remained normal for two to
four days after cutting, although petiole wilting rendered the plant abnor-
mal in appearance in half that time. In the case of Pandanus, the leaves
remained normal for six to twelve days, although severe stem droop occurred
in half that period.
Among the palms, the coconut was among the several that lasted well.
Others, however, whitened or drooped very soon after being cut. Foliage
that was thick, heavy, shiny, and leathery, such as that of Ficus, Rhizo-
phora, Hernandia, and Barringtonia, was moderately long-lived. Branches
of these species were among those that drooped badly long before the leaves
had begun to dry noticeably.
1945]
GROSSENBACHER, SPURR, & VLAMIS, CUT FOLIAGE
TABLE 2.
LASTING OF CUT TROPICAL FOLIAGE
219
Days in sunlight
Days in sunlight
without water Species with water
3-6 Calophyllum inophyllum (palo maria) 8-16
Aralia s 8-16
Cocos nucifera (coconut )—entire tree 8-16
Cocos nucifera (coconut )—separate fro 4-8
Caryota spp. (fishtail palms)—separate fronds 4-8
Ptychosperma spp. (MacArthur palm)—separate fronds 4-8
Pandanus tectorius (screw pine) 3-6
1-3 Casuarina equisetifolia (Australian pine) 8-16
Ixora 8-12
Mangifera indica (mango) 2-4
Pongamia pinnata 2-4
Acacia spp. 2-4
Musa sapientum (banana) 1-3
Ficus bengalensis (banyan) 1-3
2-1 Eugenia desu (rose apple) 8-16
Barringtonia asiatica 2-4
Livistona spp. “(fan palms)—separate fronds 2-4
Hernandia peltata 2-4
Melaleuca Jeucodendron 2-4
Morinda citrifolia 2-4
Ficus elastica (India rubber tree) 1-2
Guettarda spp. 4-1
Rhizophora Mangle (mangrove) 3-1
Sterculia foetida 3-1
4-3 Eucalyptus algeriensis 4-8
Corypha elata (giant fan Je all fronds 1-2
Hevea brasiliensis (rubber 1-2
Aleurites moluccana (can ; 3-1
Terminalia Catappa (Indian almond) 3-1
Thespesia populnea 3-1
0-4 Dodonaea viscosa 1-2
Bambusa spp. (bamboo) 3-1]
Hibiscus tiliaceus 4-3
Styloma pacifica (palm)—separate fronds 4-3
spp. 0-4
Erythrina s
Few generalities can be drawn concerning the lasting of tropical foliage.
This is due to the immense number of species in this zone and to the great
difference in their size, structure, and growth habits.
The fifty or sixty
genera actually tested are sufficient only to tell us something of the range
of lasting properties, of the reaction of some of the more important species,
and of techniques useful in prolonging the lasting qualities of plants similar
to those tested.
SELECTION AND HANDLING OF CUT FOLIAGE
The length of time that foliage of a given species will last is affected by
220 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
(1) the size of the branch, (2) the part of the tree from which it is cut, (3)
the time of day, and (4) the time of year the branch is cut. The impor-
tance of these factors was repeatedly demonstrated.
The larger the branch, the longer it will last. This applies both to
branches with and without water. For instance, a twenty-foot red oak
tree, cut at the base and guyed upright in a canvas-lined hollow filled with
water, lasted about ten days, while few red oak branches lasted more than
six days. Small sprigs of red oak foliage ordinarily drooped in two or three
days. Similar results were obtained with hemlock in many repetitions. In
Florida, entire coconut trees cut at the base and placed in water lasted
about twice as long as separate fronds. The superior lasting qualities of
entire trees and large branches as compared to small branches is probably
due to the water supply available in the reservoir of the trunk and large
limbs. This water is drawn upon by the foliage long after the water sup-
plied is no longer taken up through the base of the cut stem.
That the part of the tree whence the branch came affects the lasting of
the foliage is easily demonstrable, but the results obtained are difficult to
explain. In the case of temperate zone species tested, foliage grown in
full sunlight lasts longer when cut than foliage grown in partial shade.
Sprout foliage of red oak, red maple, and yellow birch grown in full sunlight
lasted longer than shade-grown branches of the same species. This phe-
nomenon is to be expected, as the leaves of the former type are better pro-
tected against water loss, having thicker blades, thicker cuticle, and fewer
stomata. Yet tests in Florida showed that the upper sun-grown foliage of
Pongamia, Sterculia, and the India rubber tree definitely did not last as
long as the low shade-grown branches from the same trees.
It has long been known that foliage cut in bright sunlight and supplied
with water withers more quickly than foliage cut during cloudy periods or
at night and similarly treated. This fact was confirmed by tests with six
northeastern hardwoods. In all cases, foliage cut at 5 A.M. lasted longer
than that cut at 11 a.m. The greatest difference occurred in red oak.
Branches of this tree cut at 5 A.M. lasted six days; at 7 a.M., four days;
and at 11 a.m., two days. This effect was observed only when branches
were supplied with water, and can be satisfactorily explained by air lock, a
phenomenon discussed below.
As would be expected, immature foliage is poorer in lasting quality than
hardened mature foliage. It follows that branches cut in the spring will
not last as long as those cut later in the season. Hardwood branches cut
in the fall, however, are inferior in their lasting qualities to those cut in
mid-summer. This is because cutting the branch is apt to hasten autumn
coloration, as in cherry and sumac; and, more important, is apt to hasten
leaf abscission (many species). The rapid drop of leaves in autumn-cut
branches more than compensates for the low loss of water from the mature
foliage in cool weather.
Experiments with the use of synthetic plant hormones (auxins) indicated
that, in certain instances, these substances had beneficial effects on the
1945] GROSSENBACHER, SPURR, & VLAMIS, CUT FOLIAGE 221
lasting qualities of cut foliage. It has been well established that auxin
treatment retards the formation of the abscission layer, a principle utilized
commercially by apple growers, and this retardation apparently may
lengthen the life of branches cut in the fall. Also, auxin spray seems to
maintain higher water content in leaves. It is further possible that auxin
treatment influences metabolism, but it is difficult to determine whether
the influence is direct or whether auxin influences metabolism by increasing
water content by retarding abscission layer formation. The effect of the
auxin treatment was observed in certain instances when branches were
sprayed with water solutions (ca. 0.01%), but was more pronounced when
applied in similar concentrations in combination with a wax emulsion spray
(see below).
SUPPLY OF WATER
Cut foliage can be kept alive for extended periods as long as its moisture
content can be maintained. The problem of supplying water is the limiting
factor in the preservation of cut foliage. Once water no longer reaches the
leaves, wilting and crisping quickly occur.
Plugging
Simply supplying water to the cut bases of branches does not necessarily
keep the foliage fresh, as most plants have a wound reaction which plugs
the cut stem and prevents water from reaching the leaves. The nature of
this plugging is variable. It may be structural plugging, such as the de-
velopment of tyloses and gum-like plugs in the vessels; it may be due to
secretions such as latex and resin; it may be due to external agents such as
bacteria and fungi; or it may be due to bubbles of gas in the vessels.
That plugging of vessels with tyloses and gum-like deposits is correlated
with the cessation of water uptake in certain temperate plants was deter-
mined morphologically by Prof. R. H. Wetmore and Prof. I. W. Bailey, of
Harvard University. In red oak, heavy tylosis formation takes place
d
tvloses and ‘‘amber” plugs were formed very rapidly in willow.
In resinous plants, accumulations of exuded resin tend to seal off cut
faces exposed to air. When the cut face is immediately placed in water,
however, resin does not appear to interfere seriously with water uptake.
Secretions of various tropical plants were much more troublesome. In par-
ticular, the latex of Ficus, Hevea, the mucilaginous sap of Musa, and similar
secretions in representatives of other genera, tend to become suspended in
the water, and in turn are drawn into vessels, where they form plugs at the
first cross-wall.
It was further observed that films caused by bacterial accumulations
occurred on the cut surfaces of stems containing no toxic substances such
as tannin. These might well hinder water uptake, as would also extensive
fungal activity.
In cut branches, air bubbles enter the vessels, forming “air locks,” which
presumably plug the stem and reduce water uptake. When the stem is
222 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
cut, transpiration immediately draws air up through the cut surface into the
stem. If the stem is then placed in water, the air is confined within and
will form bubbles at the first effective cross-wall. This explains why
foliage cut before sunrise lasts longer than that cut at mid-day. In full
sunlight, the heavy transpiration of foliage draws air into the stem as soon
as the stem is cut. Consequently, regardless of how quickly the stem is
placed in water, enough air has entered the stem to form an air lock. Early
in the morning, or at any other time that the transpiration rate is low, little
or no air enters the stem if the cut base is promptly immersed in water.
Recutting
It is possible to prevent the formation of an air lock by cutting the
branch under water. When bent through a pail of water and severed from
the tree by being cut under water, aspen and white ash branches took up
two to three times as much water as controls cut in the air. This enables
them to last longer.
Such a procedure, though sound, is not very practicable. Once a branch
cut in air has been placed in water, however, it can be recut under water
advantageously. This will remove any plugging in the basal portion of the
stem, whether due to tyloses, secretions, bacteria, or air locks.
The efficacy of the recutting treatment depends upon the species involved.
It materially lengthens the life of those in which plugging occurs largely
at the base, and is naturally of little value when plugging occurs throughout
the stem. In trials at Petersham, the life of cut branches of sugar maple,
red maple, beech, yellow birch, and hemlock was materially lengthened by
recutting under water. Red oak, chestnut, sumac, and pin cherry were not
materially aided. The results obtainable by recutting under water are
occasionally spectacular. In one instance, a sugar maple branch taken in
midwinter was kept alive for three months. By a combination of recutting
and the basal injection of water with pressures up to 15 lbs. per square
inch, a full crop of leaves was grown and maintained for many weeks.
Apparently species with a long functional vessel length are not helped by
recutting, as the air locks and other plugs are not confined to the base of
the stem. By testing cut segments of stems with eight to fifteen pounds
air pressure applied to one end and collecting the air bubbles under water
at the other, it was possible to calculate the approximate functional length
of open vessels. This length was several feet or more for red oak, Lom-
bardy poplar, white ash, elm, cherry, and ailanthus; several inches for red
maple, sugar maple, yellow birch, and black walnut. In conifers, the
tracheids are so short that such measurements were not made. The longest
functional vessel lengths observed were 35 feet for red oak and 20 feet for
Lombardy poplar.
Recutting a cut stem in air proved better than no treatment but inferior
to recutting under water. This is to be expected, as the structural basal
plugs are removed by the treatment, but air locks are not prevented from
being reformed.
The lasting properties of lactiferous plants can be materially lengthened
1945] GROSSENBACHER, SPURR, & VLAMIS, CUT FOLIAGE 223
by washing the latex off the wound until it stops running, and also, to a
lesser extent, by charring the base of the cut branch to coagulate the latex
in the latex tubes, thereby preventing it from plugging the vessels.
Purity of Water
Water purity greatly influences the lasting of foliage, because impurities
tend to clog up cut stems, thus reducing the rate of the water supply. This
was demonstrated by trials at Petersham with sugar maple, red oak, beech,
and hemlock. The reaction of sugar maple was typical. It lasted six days
when supplied with water from a deep driven well; four days with clear,
swift-flowing river water; three days with sphagnum swamp water; and
two days with water from a stagnant pool. Water uptake for all species
was closely correlated with lasting qualities.
In Florida, tests were carried out to see whether certain coastal species
could be maintained with sea water. Pongamia and Hernandia were in-
jured by salt water, and separate fronds of coconut quickly browned.
Entire cut coconut trees and Casuarina were successfully maintained. In
the case of coconut trees, the stem apparently filtered out the salt before
the salt reached the foliage. Mangrove was successfully maintained in salt
water as well as in fresh water only if the bases were cut under water and
maintained there.
Extensive exploratory tests carried out by Dr. P. R. Gast and by the
authors failed to reveal any chemical or combination of chemicals which
when added to the water supply was much more effective in prolonging the
fresh appearance of cut foliage than the use of water alone.
Method of Supply
The most obvious and most practicable means of supplying water to cut
foliage is through cut bases. Water may also be made available to the
plant in other ways.
Merely placing the cut bases in moist soil is sufficient to prolong the life
of coniferous branches. A certain amount of water is supplied by capillary
action, although this method is much inferior to standing the cut base in
clean water. The value of placing coniferous stems in the ground was
demonstrated by a number of tests, both indoors and outdoors, one of
which is detailed in Table 3. Hardwoods require so much water that they
are not helped by this treatment.
Water was also supplied through incisions in the stem, either made by a
sharp blade or by an auger, and through the stubs of cut laterals. In the
experiment summarized in Table 3, the amount of water taken up by hem-
lock foliage through cut bases and cut laterals was roughly proportional to
the surface area of the sapwood exposed in the cuts. The most successful
treatment, exclusive of wax treatments, was that in which water was sup-
plied both through the cut base and a cut Jateral.
Cut foliage can be maintained for long periods of time by placing the
bases in sand and preventing the tops from drying out with an intermittent
spray. In sucha case, the rate of water loss is reduced to a very low level,
TABLE 3
WATER SUPPLY AND WAX TREATMENT OF CUT HEMLOCK TREES!
Treatment Water content? Water uptake in 17 days Leaf fall
(percent) (cc. per gr. total dry wt.) (percent )
Base in 13 days 27 40 &0 40 days 80
Air 34 12 7 3 100 100
37 14 8 3 100 100
Soil 49 36 24 3 10 100
49 39 19 11 30 100
Air—one lateral 42 17 7 3 047 100 100
with water 42 19 8 3 051 100 100
Air—two laterals 44 31 14 3 057 + .064 96 100
with water 41 2 9 3 031 + .057 100 100
Water 54 55 40 3 854 6) 100
55 55 48 3 717 0) 100
Water—one lateral 54 55 55 38 607 + .059 0) 0
with water 56 56 56 8 738 + .076 0 70
A‘ir—sprayed 48 31 18 3 4 100
with Dowax? 41 34 ) 3 100 100
Air—sprayed 48 45 40 7 0) 95
with wax solution! 46 44 44 5 0 30
iin ed S4 54 49 4 582 0 95
with Dow oP] 53 48 5 404 0 95
iiseuaersd 54 56 a7 40 259 0 3
with wax ihinlanita 55 aan 56 43 393 0 6
Six : to eight feet tall placed in ‘large 1 room on n December 16, 1942. Average daily 1 range in : temperature, 45 to 60° a "Average
relative humidity 50%.
“Original water content, average ee ™ trees, 54%. All values below 35% represent fallen needles.
“One pa Dowax to four parts w
4One part paraffin to one part cuclean to eight parts kerosene.
b2S
WILAYOdUV GIONYV AHL AO TYNYNOFL
IAXX “TOA]
1945] GROSSENBACHER, SPURR, & VLAMIS, CUT FOLIAGE 225
and enough water is taken up by the leaves to make up any deficit which
may occur. The stems may be heavily plugged, but the development of
new xylem will re-establish translocation after roots are formed. An auto-
matic apparatus to accomplish this was constructed at the Harvard Forest,
which maintained leafy cuttings of various species for the entire growing
season.+ Maintenance of this type is of no value in camouflage, but is a
promising tool in the rooting of leafy cuttings.
REDUCTION OF WATER LOSS
Inasmuch as loss of water is the primary cause of the death of cut foliage,
it follows that any method which reduces water loss without otherwise
affecting the plant will increase the period of normal appearance. The
principal way of accomplishing this is to coat the foliage with a non-toxic
substance, usually a wax or a mixture containing wax.
In the course of the present investigations, a great deal of exploratory
work was carried on to investigate the possibilities of this type of treatment.
A large number of substances were tried in many combinations under a
variety of conditions. Although no material was found which could be
highly recommended, enough was learned to reveal the limitations and
possibilities of this means of reducing water loss.
A satisfactory coating must have two properties: it must form a thin
pliable inconspicuous film capable of markedly reducing water loss, and it
must be non-toxic to the living tissue of the plant. Unfortunately, these
two characteristics rarely occur together.
The effectiveness of a film in reducing water loss may be determined by
observations of the lasting quality of branches, by measuring water uptake,
by measuring changes in total plant weight, and by determining moisture
content. Observation suffices only to distinguish living foliage from dying
and dead foliage. Water uptake may be used as an index of water loss,
especially for longer time periods. Thus, in Table 3, the hemlocks sprayed
with wax solution required less water and lasted better than trees sprayed
with Dowax. Moisture content, whether determined directly from samples,
or indirectly by measuring the loss of weight of a drying cut branch, is an
index of the ability of the coating to reduce water loss. This, too, is illus-
trated in Table 3
Death of living tissue is indicated by color and structural changes which
markedly alter the normal appearance of the plant. Any substance caus-
ing the death of tissue, therefore, is undesirable as a protective coating.
Toxicity is governed i a large embeE of factors, such as temperature,
method of application, and chemical composition of the coating. Chem-
icals toxic to foliage even in a low concentration are not necessarily toxic
in combination, as in the case of ammonium hydroxide in Emulsion B,
Table 4. Most of the coatings tried in the present investigation proved
to be toxic in one way or another. Conifers are much more resistant to
4Grossenbacher, Karl A. An apparatus to maintain a surface film of water for use
in vegetative propagation. Jour. Arnold Arb. 26: 206-211. 1945.
226 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
TABLE 4.
TESTS IN FULL SUNLIGHT WITH WAX EMULSION DIPS
ON BRANCHES SUPPLIED WITH WATER
Species Control Dowax! Emulsion A? Emulsion B#
Average number of days before reaching Class II
Red oak 4 6 8 5
White oak 3 5 8 6
— 4 2 6 5
ech 4 3 6 Ri
Ge birch 2 12 3 2
Elm 1 1 1 1
Aspen 0) 0 1 0
10ne part Dowax to three parts water.
“Paraffin, 7.7g.; Duponol WS, 2.3g.; linseed oil, 3.5ml.; glue, 1.0g.; water, 200ml.
38Candellila wax, 5.0g.; Neomerpin, 1.0g.; Ammonium hydroxide, 4.0g.; kerosene,
4.2ml.; water, 6.3ml. Concentrate diluted, one part to three parts water
toxic effects than hardwoods. The development of a non-toxic coating is
the principal difficulty in solving the problem of how effectively to reduce
water loss and maintain living foliage.°
Coatings fall into two general classes: solutions and emulsions. Solu-
tions of various waxes in organic solvents are highly effective in preventing
water loss. Most of the solvents, however, are highly toxic to living tissue.
The most promising solution used was a mixture of one part paraffin and
one part petrolatum in eight parts of kerosene. This proved extremely
effective in preventing water loss (Table 3), and of low toxicity when
sprayed on foliage at such a temperature that the film solidified as soon as
it was formed. It was very toxic, however, when used at higher tempera-
tures, or when applied as a dip. In these cases, greater penetration
occurred, and penetration of any coating was found to be conducive to
toxicity.
Emulsions have a basic advantage over solutions in that they may be
applied in a non-toxic medium — water. Among the emulsions tested, a
prolonging the life of conifers but not of many hardwoods (Tables 3 and
4). It was toxic in varying degrees with several of the plants tried. Ex-
periments made by Dr. P. R. Gast in Florida indicated that its use mate-
rially prolonged the life of fronds of the cabbage palm (Sabal palmetto)
and the leafy foliage of blackjack oak (Quercus marilandica). Auxin
treatment, as mentioned above, apparently lengthened the life of cut foliage
somewhat when used with Dowax on branches supplied with water. This
5Comar, C. L., and Barr, C. G. Evaluation of foliage injury and water loss in con-
nection at use of wax and oil emulsions. Plant Physiology 19: 90-104. 1944.
1945 ] GROSSENBACHER, SPURR, & VLAMIS, CUT FOLIAGE 227
emulsion was developed by E. J. Miller® and contains a wax, an ammonium
salt of a drying acid, and a colloidal earth.
Emulsion A (Table 4, footnote 2) was the most promising developed in
the present investigations. It is a non-ammonical mixture but inconvenient
in that it cannot be prepared in a concentrated form. For hardwood foliage
it was of low toxicity and highly effective in reducing water loss. This
emulsion was developed by Dr. Ernest Ball, of Harvard University, who
participated actively in the development of satisfactory coatings.
Emulsion B (Table 4, footnote 3), an ammonical mixture which could be
prepared in a concentrated form, was also effective in reducing water loss.
The emulsifiers used in this and other emulsions, as well as certain other
chemicals, were provided by the E. I. Dupont de Nemours Company.
All the coatings mentioned above are effective only when water is supplied
to the treated foliage. Without water, improvement was noted only in
certain cases. The lasting of conifers could usually be lengthened some-
what. Emulsions A and B also helped certain hardwoods, such as red oak
and paper birch. This limited effect in the absence of water is to be
expected, inasmuch as a protective coating is of value only in reducing
water loss to a rate comparable with that of water uptake, thus keeping
the plant tissue alive. If water is not supplied, a protective coating can
lengthen life only a short time by retarding the loss of water already
present in the plant.
SUMMARY
Extensive investigations were carried out to determine how long cut
foliage of different species will retain its normal appearance under various
conditions, and how the life of cut foliage can be maintained.
More than one hundred representative species of the northeastern United
States, western Europe, Asia, and the tropics were tested and classified as
to their lasting qualities, both with and without a supply of water.
qualities. Plants of the same genus tended to have similar lasting qualities
regardless of their geographic origin.
The range in lasting qualities of tropical species was much the same as
that of temperate plants. In the tropics, however, because of the large
size and weight of many of the leaves, abnormal appearance of drying
foliage often was first manifested by marked wilting of the petioles and
consequent leaf drooping, rather than by a change in the leaf itself.
A number of factors involving the selection and handling of cut foliage
affect their lasting qualities. The larger the branch, the longer it will last.
Foliage grown in full sunlight generally lasts longer than shade-grown
foliage, although upper sun-grown branches of certain tropical species were
found to be inferior to lower limbs grown in the shade. Mature foliage
6Miller, E. J., Meilson, J. A., and Bandemer, Selma Wax emulsions for ee
nursery oe and other plant materials. Michigan a Exp. Sta. Spec. Bull. ;
7U. S. Patent no. 2,013,063.
228 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXvI
lasts better than immature, but late season foliage does not last as well
because of abscission layer formation. In certain instances, auxin treat-
ment has beneficial effects on the lasting qualities of cut foliage, both in
retarding abscission and in maintaining water content.
Supplying water generally increases the life of foliage two to four times.
Water ceases to reach the leaves, however, in a few days, due to plugging
of the stem. This plugging may be due to tyloses or gum-like deposits
in the vessels, to secretions, such as latex or resin, to bacteria or fungi, or to
air locks—bubbles of air in the vessels. By cutting foliage when the
transpiration rate is low (during rains, early in the morning, etc.), plug-
ging from initial air locks can be minimized.
In plants with functionally short vessels, where plugging occurs mainly
in the basal portion of the stem, recutting the stem under water will remove
the plug and materially increase the life of the foliage.
Cut foliage lasts best in pure water, as solid particles clog the vessels and
chemical impurities may injure the living tissues. Salt water (sea water),
however, successfully maintained cut branches of a few tropical coastal
plants.
No chemical or combination of chemicals was found which would in-
crease the lasting of cut foliage.
Water can be successfully applied to cut branches through the base,
through cut laterals, and through stem incisions.
The reduction of water loss by means of a non-toxic coating will mate-
rially increase the life of cut foliage. Despite extensive experiments, no
highly successful coating was developed. Several coatings however, were
satisfactory under certain conditions. Among these coatings were one solu-
tion of paraffin and petrolatum and two emulsions: a non-ammonical one
with paraffin, and an ammonical one with candellila wax. Waxy coatings
are particularly effective when the treated foliage is supplied with water.
The most satisfactory coatings approximately double the life of many
species.
HaArvaArD Forest,
PETERSHAM, Mass.
JOURNAL
OF THE
ARNOLD ARBORETUM
VoL. XXVI JULY, 1945 NuMBER 3
PLANTAE PAPUANAE ARCHBOLDIANAE, XVI*
E. D. MERRILL AND L. M. PERRY
With eleven text-figures
THE FOLLOWING genera are considered in this article: Urophyllum,
Pachystylus, Canthium, Antirhea, Timonius, Mastixiodendron, Coffea,
Ixora, Versteegia, Coprosma, Coelospermum, Morinda, and Galium. We
still have Psychotria and its closely related genera to be included in another
paper, and that, we hope, will complete the Rubiaceae until such time as
certain specialists can give the collections their attention.
RUBIACEAE (in part)
Urophyllum Wallich
Urophyllum umbelliferum Val. Nova Guin. Bot. 14: 262, t. 29A. 1925.
British New Guinea: Fly River, 528 mile Camp, Brass 6646, May 1936, alt. 80 m.,
plentiful in undergrowth on ridges (tre ee 4 m. high; flowers green; ripe fruit black,
fleshy, about 1 cm. long).
Described from Netherlands New Guinea. This collection is not so
pubescent as the original, and the petioles are slightly longer than those
shown in the plate, yet these are only minor variations. The calyx-lobes
vary in length, being 2-4 mm.; in the open flower the corolla-tube is 4 mm.
long and sriecmal glabrous, and the lobes are 3—3.5 mm. long and pilose
on the outside
Pachystylus K. Schumann
Pachystylus Guelcherianus K. Schum. in Schumann & Hollrung, Fl. Kaiser Wilhelms
889; Schumann & Lauterb. Fl. Deutsch. Se Stidsee 574. 1900.
Tarenna Guelcheriana (K. Schum.) Val. Bot. Jahrb. 60: 8
NETHERLANDS NEw GuINEA: Bernhard Camp, seen . er, Brass 13886, Apr.
1939, alt. 120 m., rain-forest of lower mountain slopes (undergrowth tree 2.5 m. high).
British New Guinea: Oroville Camp, Fly River, Brass 7401, a 1936, plentiful
in rain-forest undergrowth (tree 2-3 m. high; flowers cream-colore
Described from Northeast New Guinea, and Beene reported from
* aan Results of the Richard Archbold Expeditions. See Jour. Arnold Arb.
26: 1-36. 1945.
230 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXVI
Netherlands New Guinea. Although the genus was reduced to Tarenna
Gaertner by Valeton, Bremekamp believes it should be maintained as a
separate genus, and from our little available material we agree with his
point of view
Canthium Lamarck
Canthium suborbiculare (White) comb.
Plectronia suborbicularis a ia Linn. Soc. N. S. Wales 51: 296, t. 17. 1926,
Jour. Arnold Arb. 10: 268
Known only from Papin,
Canthium cymigerum (Val.) B. L. Burtt, re Bull. 1936: 463. 1936.
Plectronia cymigera Val. Bot. Jahrb. 61:
NortTHEAST NEW GUINEA: evra ese 19513, May 20, 1909, in forest.
Sotomon Istanps: Bougainville: Kugimaru, Buin, Kajewski 1805, June 1930,
alt. 150 m., rain-forest (tree up to 20 m. high; leaves dark glossy green; fruit 9 mm.
long, 1.2 cm. broad, 6 mm. thick) ; Koniguru, Buin, ci ey 2111, Aug. eae alt. 900
rain-forest (tree up to 10 m. high; fruit 6 mm. long, 1 cm. broad); Ysabel:
Tasia, Brass 3286, Dec. 1932, coastal rain-forests (small ee with drooping branches;
leaves shining, the lower surface very pale; flowers white).
In the original publication of this species the type-number is given as
17513, so that either the number on our duplicate was copied wrong or
there was an error in publication. It does not seem possible that both
Schlechter’s numbers could have been collected in the same place in the
same month, unless an exceptionally large collection was made. There are
two unbroken leaf-tips in the Schlechter specimen cited which tend to be
broadly and obtusely acuminate. In the Solomon Islands material the
leaves are ovate-elliptic, up to 10 mm. broad, more abruptly rounded at
the base, and rather abruptly obtusely acuminate.
Canthium odoratum (Forst. f.) Seem. Fl. Vit. 132. 1866.
Coffea odorata Forst. f. Prodr. 16(no. 94). 1786.
British New Guinea: Mabaduan, Brass 6539, April 1936, common in small
scrubby rain-forest patches in savanna-forests (tree 5-6 m. high).
This fruiting collection shows a very close resemblance to the Polynesian
material. The secondary venation of the leaves is more obvious in the
material above cited than in the other specimens at hand, the fruit is more
strongly rugose on the outside, and the semicircular pyrenes have a very
deep indentation on the inner surface.
3
Canthium graciliflorum sp. nov.
Arbuscula 4—5 m. alta; ramulis ultimis 4-angulatis demum subrotundatis,
minute puberulis, internodiis 1.5—-4 cm. longis; stipulis valde inaequalibus,
altera 2 mm. longa, altera 4 mm. rap basi latis, ensiformibus; foliis
lanceolatis utrinque angustatis, 3-4.7 cm. longis, 1—1.8 cm. latis, apice
anguste obtusis, basi in petiolo 2-3 mm. longo attenuatis, subcoriaceis,
supra subnitidis, olivaceis, subtus opacis, interdu 0 sus basim
minute puberulis, ceterum glabris, nervis lateralibus utrinsecus + 4 oblique
adscendentibus marginem versus arcuatim peer ae utrinque incon-
spicuis; corymbis axillaribus bifurcatis vel trifurcatis, 1.5-1.7 cm. longis
latisque; floribus plerumque breviter pedicellatis; ovario subgloboso glabro
1945] MERRILL & PERRY, PLANTAE ARCHBOLDIANAE, XVI 231
vix 1 mm. longo; calycis limbo brevissimo, minute 5-dentato; corollae tubo
1 mm. longo, fauce pilis reflexis barbato, lobis 2.5 mm. longis, ey
patentibus vel reflexis; staminibus fauce insertis, filamentis 1.5 mm. longis,
antheris sagittatis, 1. 5 mm. lo ongis, apice apiculatis; stylo See 4 mm.
longo; stigmate mitriforme, apice bilobato; fructibus non visis.
Britiso New Gunes: Tarara, Wassi Kussa River, Brass 8596 (TYPE), Dec. 1936,
common and conspicuous in brushy rain-forest fringing river (slender tree 4-5 m. tall;
branches semi-erect ; leaves smooth and shining; very numerous cream-colored flowers).
This species is related to Canthium odoratum (Forst. f.) Seem., but the
leaves are narrower and without domatia; the inflorescence is much shorter
and the flowers are slightly smaller in all the parts. In some ways, partic-
ularly in the outline of the leaves and their tendency to shine, the plant sug-
gests one passing in Australia as Canthium lucidum Hook. & Arn., which
perhaps more correctly should be known as C. lamprophylium F. v. Muell.
Here again in comparison the Papuan material is less coarse in every way
than the Queensland collections. In the Gray Herbarium is a small frag-
ment from Hooker’s herbarium collected on Gambier Island by Beechey
labelled C. lucidum Hook. & Arn. The fragment is probably from the
type or an isotype and there is no question in our minds that it is identical
with C. odoratum (Forst. f.) Seem., but it is scarcely the same as the
Australian material so-named.
Canthium brevipes sp. nov.
Arbuscula 3—5 m. alta; ramulis in sicco atro-cinereis vel brunnescentibus,
novellis nigrescentibus, glabris, internodiis 2-4 cm. longis; stipulis brevibus,
1-2 mm. longis, caducis; foliis ellipticis vel oblongo- -ellipticis, 7-12 cm
longis, 2.2-4.5 cm. latis, utrinque aequaliter ee apice obtusis vel
obtuse acuminatis, basi cuneatis, chartaceis, in sicco supra atro-fuscis,
subtus olivaceis, glabris, nervis lateralibus ieee 6-8 utrinque mani-
festis non prominulis in axillis domatia minuta foventibus; floribus in
tuberculis axillaribus 1—9-umbellatis; pedicellis 2-3 (—6 in fructu) mm.
longis; ovario subgloboso, cum calyce + 1.5 mm, ee calycis tubo incon-
spicuo, margine minute dentato ciliato; corollae tubo 3-4 mm. longo intus
retrorse pubescente, fauce et lobis basi villosis, lobis 2.2—2.5 mm. longis,
lanceolatis acutis; antheris circiter 1 mm. longis, sessilibus, in fauce insertis,
facie fructum Guioae admonentibus; pyrenis extus rugulosis, oblique semi-
rotundis latere ventrali incisis; seminibus valde curvatis.
British New Guinea: Lake Daviumbu, Middle Fly River, eee che Aug. 1936,
rain-forest, common in lakeshore undergrowth (tree 3-5 m. high; fruit red) ; Penzara,
between Morehead and Wassi Kussa Rivers, Brass 8443 (TYPE), rain- ibe (small tree
fringing a permanent waterhole; flowers yellow)
Canthium brevipes is closely related to C. Valetonianum S. Moore, if we
have rightly interpreted the latter species. In the former the veins of the
leaves are closer together; the fruit is broader and at the apex has a broad
shallow depression not characteristic of Moore’s species. At a ae the
fruit somewhat suggests that of certain species of Guioa in contour; in
fruit the pedicel is only 6 mm. long, whereas in the other species aha we
232 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XxvI
have seen the pedicel has elongated to 1 cm. or more in length. This group
of species with umbellate flowers needs a revision from a specialist’s point
of view.
Canthium korrense (Val.) Kanehira, Bot. Mag. Tokyo 46: 671. 1932.
Plectronia korrensis Val. Bot. Jahrb. 63: 311. 1930
Sotomon Isianps: Florida (N’Gela): North end of the island, Brass 3510, Jan.
1933, alt. 75 m., hill rain-forests (tree 5 m. tall, with close gray bark; flowers white or
pale yellow; fruit smooth, pale red); Guadalcanal: Mamassa, Konga, Kajewski
2479, Feb. 1931, alt. 400 m., rain-forest (tree up to 18 m. high, on banks of freshwater
creeks; fruit dull red when ripe, 1.2 cm. long, 1.4 cm. broad).
This material seems to be a very good match for the description and
material collected at Ponape. ‘The flowers are perhaps nearer 6 mm. long
rather than 4 mm. as given in the =_— description.
Canthium longiflorum (Val.) comb.
Plectronia longiflora Val. Bot. Jabrb. "61: 56. 1927.
NeTtTHERLANDS New Guinea: Bernhard Camp, Idenburg River, Brass 13971, Apr.
1939, alt. SO m., rain-forest subject to occasional flooding (undergrowth tree 6 m. tall,
flowers white). British New Gurnea: Tarara, Brass 8509, Dec. 1936, common in
rain-forest undergrowth (shrub 2 m. high; fruit nike’,
Described from Northeast New Guinea. The second specimen cited is
in fruit only but seems to fit better here than elsewhere at present.
Canthium Schlechterianum nom. no
Plectronia nitens Val. Bot. Jahrb. 61: 57. 1927.
Nortueast New Guinea: Yoangen, Clemens 6492, June 1937, alt. about 1250 m.
This collection agrees with Valeton’s description except that the corolla-
lobes are about 12 mm. long rather than 5mm. _ The specific epithet mtens
is pre-empted in the genus Canthium. The species is known only from
New Guinea, and only in the flowering stage.
? Canthium megistocarpum sp. nov
rbor parva; ramulis hegimeieansbun: novellis in sicco atro-olivaceis,
glabris; internodiis 3—7.5 cm. longis; stipulis inaequalibus, 5-10 mm.
longis, pie latis, ensiformibus; _ oblongo- lanceolatis, 18-27 cm. longis,
5.5-7.5 Cc latis, apice longe acuminatis, acumine 12 cm. longo, basi
cuneatis, pen glabris, nervis lateralibus utrinsecus 8 vel 9 supra
leviter impressis, subtus prominulis, venis inconspicue manifestis, laxis;
petiolo page 9 mm. longo; floribus non visis; fructibus pyriformi us, in
sicco 7 cm. longis, 4, 5 cm. diametro; pyrenis 2, endocarpio osseo extus
irregulariter tuberculato; semine pendulo ad tertium superum ovarii longi-
tudinis affixo
a New Guinea: 2 km. southwest of Bernhard Camp, Idenburg River,
Brass 13476 (type), March 1939, alt. 800 m., small tree of rain-forest substage (large
sis soa green pithy fruit up to 10 cm. long and 7.5 cm. in diameter).
This species may be related to Canthium Schlechterianum Merr. & Perry
(Plectronia nitens Val.), but up to the present the only comparable parts
are the foliar characters, one having very large flowers, the other very large
fruits. Valeton’s species, if we have correctly interpreted it, has equal
stipules and firmer, broader, and shorter leaves with shorter petioles. Sup-
plementary material of both species is greatly needed.
1945] MERRILL & PERRY, PLANTAE ARCHBOLDIANAE, XVI 233
? Canthium aurantiacum sp. nov.
Arbor parva, usque 5 m. alta; ramulis novellis atro-fuscis, glabris; inter-
nodiis 1.5—3 cm. longis; stipulis subaequalibus, 5-10 mm. longis, basi latis,
ape elongato- ppg foliis oblongo-lanceolatis, 12.5—-14.5 cm. longis,
S apice breviter acuminatis, acumine obtusiusculo, basi
anguste Sens a acutis, glabris, coriaceis, nervis lateralibus utrinsecus
6-8 oblique adscendentibus, utrinque manifestis non prominulis, venis
obscuris; petiolo circiter 8 mm. longo; floribus non visis; tuberculis in
axillis foliorum 2- vel 3-cicatricosis; fructibus in sicco obovoideis 4 cm.
longis, 2.5 cm. diametro, gaan rugulosis; pyrenis 2, endocarpio osseo extus
irregulariter tubercu ato: emine pendulo ad tertium superum ovarii longi-
tudinis affixo; embryone ne cm, longo, teretiusculo, recto, cotyledonibus
brevibus, radicula supera.
British New Guinea: Kubuna, Brass 5580 (type), Nov. 1933, alt. 100 m., ridge-
forest substage (small tree 5 m. high; leaves thick, smooth; fruit solitary in axils,
orange-vellow, about 4.5 cm. long, 3 cm. or more in diameter).
The leaves of this species are smooth and thicker than are those of most
species of Canthium from New Guinea; the tubercles in the axils of the
leaves are very short and have 2 or 3 scars on the ends, probably indicating
the number of flowers or fruits borne there. The fruit is very large com-
pared with that of the other New Guinean species except one, but C.
glabrum Bl. of Malaysia has fruit as large or larger. In the latter species,
however, the leaves are larger, thinner, and broader than in the New
Guinean material.
Antirhea Commerson
S. Moore, Jour. Bot. 65: 266. 1927, and Fosberg, Sargentia 1: 121. 1942,
include within their concept of the genus Timonius DC. plants with fruits
characterized by a 4—10-loculed putamen. The latter character, coupled
with a persistent calyx and a corolla with imbricate lobes, belongs to the
concept of the genus Antirhea Commers. As far as we may judge from
the material at hand the genus is a valid one. Valeton, Bull. Dép. Agr.
Ind. Néerl. 26: 12. 1909, indicates the similarity of the inflorescences in
the entire tribe Guettardeae, in which case one would naturally look for
generic differences in the fruits. S. Moore simply indicates that Brass 946
is apparently a new species of Tzmonius, suggesting its resemblance to T.
subcoriaceus Val. but noting that the center of the fruit is occupied by a
star-shaped woody mass containing 9 pyrenes. We believe that another
genus is represented when the fruit contains a putamen (a hardened con-
crete mass containing the seeds) or, as Fosberg calls it, a fused stone. In
the case of Timonius Kajewskii (Guill.) Fosb., at least as to the fruiting
specimen cited by Fosberg, the pyrenes cohere very closely, but the walls
of the adjoining pyrenes maintain their identity; this is apparent in a cross-
section of the fruit which shows a definite line between adjacent pyrenes.
In Timonius Smithii Fosb., from Fiji, the fruit does not contain separate
pyrenes as such, but a putamen with 10 locules. Since the latter is a char-
acter of Antirhea Commers. rather than of Timonius DC., the Fijian plant
should be called Antirhea Smithii (Fosb.) comb. nov. (Timonius Smithii
Fosb. l.c.).
234 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXVI
Antirhea tenuiflora Benth. Fl. Austr. 3: 418. 1867; F. M. Bail. Queensl. Fl. 3
1900
British New Guinea: Tarara, Wassi Kussa River, Brass 8525, 8585, Dec. 1936,
comenon in undergrowth of rain-forest (shrub of weak habit 2—5 m. high; leaf-nerves
pale; flowers white; fruit red, ovoid).
These collections are intermediate between two Queensland specimens
in our herbarium and agree reasonably well with the description of this
species. This seems to be the first record of the presence of the genus in
New Guinea.
A. — costata Merr. & Perry: a. habit, «x 3; b. leaf, x 3. B. Antirhea
mega suede Me r. & Perry: a. leaf and terminal bud, , Geena ee ere Oe Aces ee SSii.
oe cae stipules dentate, the teeth apparently being glands alls the
Stipules se aheceat outside; midrib of leaves sparsely hirtellous beneath.
Leaves smaller, 9-12 XK 3-3.5 mm.......... 2c eee e eee eee amiana.
Leaves larger, 16-35 & 3-8 mM............000eeee eens GC habbemensis.
Stipules glabrous with a thick margin of hairs; leaves glabrous............
ST ERCE OTIC RORIC TEC TLE SORE aa eT rere et ee C. novoguineensis.
Coprosma divergens Oliver, Records Dominion Museum [N. Zeal.] 1: 44. 1942.
Brass collected two numbers (4215 and 4216) of this species, one ¢ , the
other 2 ; it would seem that the numbers were mixed when duplicates were
diinbuied. as mo. 4216 in the Arnold Arboretum set is froma ¢
However, the two are so much alike that the only difference we can see is
that one has stamens where the other has styles, the calyx and corolla
being alike in the two plants.
Fic. 11. A. Coprosma meaning Merr. & Perry: a. habit, x 3; 6. ¢ flower, Xx
28° GO ae x 28. . Coprosma Brassii Merr. & Perry: a. habit, x 3; b. 3
flower, X 23; c. 2 Pie x 23; d. fruit 3.
Coprosma Archboldiana sp. nov. Fic. 11,A;
Frutex ‘dune caulibus minute hirtellis vel glabris, ramosis; inter-
nodiis 4-10 m ngis; foliis oppositis, lineari-lanceolatis, 0.6—1 cm. longis,
2.5-3 mm. latis, apice acutis, basi in petiolo marginato 1-3 mm. longo fere
1 mm. lato angustatis, margine consperse ciliatis, pilis brevibus, utrinque
glabris minute rugulosis, costa tantum manifesta; stipulis cum petiolorum
basibus connatis, vaginan ntibus, margine minute dentatis, ciliatis; floribus
sessilibus terminali bus; ¢ : calyce ane inaequaliter 4- dentato, minute
ciliato; corolla tubiforme, fhe 7 mm. longo, lobis 2 mm. longis, lanceolatis:
- longis
reliquis brevioribus, ‘sparsim ciliatis; corolla tubiforme, tubo 7 mm. longo,
260 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
lobis eT 2 mm. longis, costa et apice sparsim ciliatis; staminibus
nullis; stylis 2, 1.5—2 cm. longis, minute papillosis; baccis globosis, 6 mm.
diame tro
acti stare New Guinea: 5 km. east of Wilhelmina-top, Brass 9411, Aug. 1938,
alt. 3650 m., matted on exposed rock-faces; 11 km. northeast of Wilhelmina-top, Brass
& Myer-Drees 9817, Sept. 1938, alt. 3400 m., alpine grassland, dwarf shrub prostrate
9831 (TYPE), Sept. 1938, alt. 3560 m., pendent in great cushion-like masses on moist
semi-shaded cliffs (fruit red, globose, soft, fleshy).
Coprosma Archboldiana appears to be related to Coprosma ernodeoides
A. Gray and to the group of Coprosma pumila (C. pumila Hoo :
nivalis Oliver, C. Petriei Cheesem.). Coprosma Archboldiana differs from
all of these in the definitely lobed calyx of the 9° flower. The stipules are
very short and some of the younger ones under a lens seem to be minutely
denticulate. The calyx of the ¢ flower appears to have four teeth of
unequal length, but in between these are minute denticulations which might
possibly suggest that the calyx is a transitional form. he corollas of
both ¢ and @Q flowers are alike in size and shape in the New Guinean
material. In the species mentioned above the corolla of the @ flower is
always shown as the smaller in Oliver’s monograph. Nevertheless it must
be noted that the habit, the relative size of the leaves, the solitary terminal
flowers, and the trumpet-shaped corolla all indicate a close relationship
among these species.
Coprosma Brassii sp. nov. Fic. 11, B.
Frutex 1,.5—2.5 m. altus, ramosus; ramulis ultimis plerumque ternatim
imac obtuse 3- angulatis, patenti- hirtellis, pilis brevibus, internodiis
3-10 m m. longis, partim stipulis obtectis; foliis ternatim verticillatis, 1-1.7
margine minute papillosis vel erosulis; costa tantum manifesta, supra im-
pressa subtus prominente; stipulis firmis extus glabris, circiter 2 mm. longis,
in axillis supra folia breviter connatis, margine fimbriatis, processibus
inaequalibus centrali longiore (2 mm. longo, 0.2 mm. lato), margine pubes-
centibus, pilis + densis 0.2 mm. longis: floribus sessilibus solitariis vel 4
ad apicem ramulorum; ¢ : calyce + turbinato, 4—6-dentato, ——
circiter 2 mm. longis, acutis, minute ciliatis vel glabris: ; corolla campanu-
lata, tubo 3-4 mm. longo, lobis ovatis aequilongis, recurvis vel Seta
staminibus 4-6, exsertis, antheris 4 mm. longis, apiculatis; ? : calyce +
lata, tubo 2~3 mm. longo, lobis circiter 3 mm. longis; staminibus partim
inclusis vel inclusis, ate stylis 2, + 1.5 cm. longis, minute papillosis;
baccis ellipsoideis, circiter 6 mm. longis (calyce persistente inclusis), 4 mm,
diametro.
NETHERLANDS NEw GuINnEA: 11 km. northeast of Wilhelmina-top, Brass & Myer-
Drees 9807, 9809, Sept. 1938, alt. 3400 m., occasional in forest-edges (robust shrub 2 m.
high; fruit red); 7 km. northeast of Wilhelmina-top, Brass & Mvyer-Drees 9842, 9843
(TYPE), Sept. 1938, alt. 3560 m., very abundant on forest-borders (large weak shrub
of brownish appearance, 1.5—2.5 m. tall) ; 2 km. east of Wilhelmina-top, Brass & Mvyer-
Drees 10221, 10304, Sept. 1938, alt. 3800 m., common in glades and forest-borders, sub-
alpine forest (lax shrub 1.5-2 m. high).
1945] MERRILL & PERRY, PLANTAE ARCHBOLDIANAE, XVI 261
This species seems to belong to the group of Coprosma sundana, accord-
ing to Oliver’s monograph. In nos, 9807 and 9809 the branchlets are
nearly glabrous, but the collections do not seem to differ from the others in
any other character. The dominant arrangement of both leaves and
branchlets is ternate, although we have noted one node with only opposite
leaves, also in another plant there is one verticil containing four rather than
three branches. One flower of the type has three styles, but all others
examined have two. The flowers are terminal and solitary or surrounded
by three younger buds probably axillary at the next node, the internode
between being so short as to give the appearance of four flowers clustered
together. At the base of each flower is a whorl of reduced leaves and their
stipules. The distinctive characters of the species are: the ternate arrange-
ment of the leaves, their glabrousness, and under a lens the minutely eroded
margin which appears as if it might have been papillate (possibly pubes-
cent) or glandular; and the fimbriately margined stipules forming a very
short sheath, the processes glabrous on the outside but the margin hairy
(scarcely in a single line as cilia), the middle one being the longest of five
or seven which extend from the margin of a single stipule.
Coprosma habbemensis sp. nov.
Frutex 3-5 m. altus, ramosus; ramulis minute hirtellis; internodiis
@: calyce ut in 3 ; corolla campanulata, tubo + 2 mm. longo, lobis 3-4
mm. longis; staminibus parvis, filamentis brevissimis, antheris inclusis;
stylis 2, circiter 1.5 cm. longis, minute papillosis; baccis in sicco ellipsoideis,
circiter 7 mm. longis et 5 mm. diametro.
NETHERLANDS New Guinea: Lake Habbema, Brass 9367, Aug. 1938, alt. 3225 m.,
common in open parts of subalpine forest (tree or shrub 3-4 m. high) ; 9 km. northeast
of Lake Habbema, Brass 10587 (TYPE), 10588, 10929, Oct. 1938, alt. 2800 m., abundant
in forest second growths on landslips, native clearings, and openings (scrambling shrub
2-5 m. high; fruits orange in no. 10587, red in no. 10929).
This species is closely related to Coprosma W ollastonii Wernh. from the
Carstensz Mountains; in fact, we have hesitated for some time whether to
consider this a more pubescent form of C. Wollastonii or a distinct species.
The peculiar combination of characters of Wernham’s species (leaves
scabrous above, midrib beneath clothed with scattered stiff hairs, hispid
margin, glabrous stipules, glabrous branchlets), as well as the characters
found in C. novoguineensis, have finally influenced us to believe that this is
a distinct species. Apart from the size of the leaves, the longer hairs on the
262 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXVI
midrib beneath, the more conspicuous stipules, and the much more open
habit, C. habbemensis is very similar to C. Lamiana Oliver.
Coprosma novoguineensis sp. nov.
Frutex plerumque 1—1.5 m. altus, ramosissimus; ramulis ultimis brevi-
bus, pubescentibus; internodiis brevissimis saepissime stipulis fere obtectis;
foliis arena in sicco rigide subcoriaceis, confertis, lanceolatis vel ova-
tis, 0.6-1.5(-2.2) cm. longis, 0.2—-0.6 cm. latis (in geese typico
0.6-0.8 X 0.2- 0.3 cm. ), apice cCuspidatis, cuspide 1—-1.8 mm. longa, basi
ramulorum:; $: ca yce 4- vel 5- dentato, ‘dentibus circiter 1 mm. longis,
glabris vel in margine aieniee et inter dentes minute ciliatis; corolla cam-
panulata, tubo 3 mm. longo, lobis 4 vel 5, anguste ovatis aequilongis;
nulatis; 2: calyce et corolla ut in ¢ ; nibus inclusis parvis; stylis 2,
circiter 9 mm. longis, minute anlage: pen ellipsoideis, circiter 5 mm,
longis et 3 mm. diametro.
NETHERLANDS NEW GUINEA: 11 km, —— of Wilhelmina-top, Brass & Myer-
Drees 9705, 9706, 9707, Sept. 1938, alt. 3400 m., grassy place (shrub 1.5 m. high ;
flowers brownish, the anthers somewhat nine ripe fruits orange) ; 7 km. northeast of
Wilhelmina-top, Brass & Myer-Drees 9933, Sept. 1938, alt. 3740 m., in edge of forest,
not common (very slender sparsely foliaged tree 3.5 m. high) ; same locality, Brass &
Myer-Drees 9939 (type), 9940, Sept. 1938, alt. 3900 m., plentiful on grassy summits
10378, 10380, alt. 3700 and 3750 m., in marginal shrubberies of pana forest : Lake
Habbema, Brass 9028, 9063, 9144, 9145, 13736A, Aug, 1938, alt. ., abundant in
shrubberies bordering forest (loose straggling ashy 1.5-—2.5 m. ae
The collections from Mount Wilhelmina are fairly uniform except for
nos. 9933 and 9706. The first has leaves 1.5 0.5 cm., the second 1
0.6 cm.; apart from these two collections, the rest have leaves 0.6-0.9
0.2—0.4 cm. tending to dry very loosely folded along the midrib. All have
very short internodes and are profusely branched, thus giving the impres-
sion of a compact habit. In the collections cited from Lake Habbema we
can find no real differences. The glands or teeth along the margins of the
stipules do not seem to have shrunk as much as in the Mount Wilhelmina
material, and the specimens show both a compact and a loose habit: there
is a wide variation in leaf-size, the two extremes being no. 9145 with leaves
1.3-2.2 0.2-0.5 cm., and no. 9028 with leaves 0.7 0.4 cm., the latter
having very crowded nodes and the stipules also being short; in no, 9145,
on the other hand, the nodes are 5-10 mm. long, and the stipules are also
long. In practically all the collections the yellowish stipules are con-
spicuous. Coprosma Lamiana Oliver, collected at the base of Doorman-
top, differs only in the longer pubescence of the branchlets, the dense
pubescence of the stipules (on account of which there is practically no line
of demarcation between stipules and branchlet at the base of the stipules) ,
and the few short hairs on the midrib beneath.
1945] MERRILL & PERRY, PLANTAE ARCHBOI DIANAE, XVI 263
Coelospermum Blume
Coelospermum reticulatum (F. v. Muell.) Benth. Fl. Austr. 3: 425. 1867; F. M. Bailey,
Queensl. Fl. 3: 769. 1900.
Pogonolobus reticulatus F. v. Muell. Fragm. Phytogr. Austr. 1: 56. 1859.
British New Guinea: Tarara, Wassi Kussa River, Brass 8565, December 1936,
more or less gregarious on savanna-forest ridges (stiff erect shrub growing to 30-100
-from a somewhat fleshy rootstock; flowers white; leaves opposite and in 3’s).
Previously reported from Queensland and North Australia.
Morinda Linnaeus
cm. diametro), crassiuscula, irregulariter 4- vel 5-angulata, longitudinaliter
sulcata, pyrenis 4 subosseis, oblongis vel obovatis, circiter 8 mm. longis,
complanatis.
NETHERLANDS NEw GUINEA: 6 km. southwest of Bernhard Camp, Idenburg River,
Brass 13012 (type), Feb. 1939, alt. 1200 m., frequent in rain-forest (large canopy liane ;
fruit red, costate).
Morinda costata seems to be most like M. Grayi Seem., from Fiji, but the
leaves have shorter petioles, and the syncarps are much larger and sessile
in the New Guinean material. Among the New Guinean species M. costata
is somewhat like M. glomerata (Blume) Mig., but in the latter the syncarps
are short-pedunculate, the berries are smaller, and there is practically no
thin calyx-margin, i. e. the entire wall is thickened around a cup-shaped
opening about 5 mm. in diameter; in M. costata, on the other hand, there
is a thin calyx-margin about 2 mm. wide projecting beyond the fleshy free
costate part of the berry, leaving an opening at the apex about 3 mm. in
diameter.
Morinda glomerata (Blume) Migq. Fl. Ind. Bat. 2: 247. 1857; Val. Nova Guin. Bot. 8:
515, 1911
Sphaerophora glomerata Blume, Mus. Bot. Lugd.-Bat. 1: 179. 1850.
NETHERLANDS New Gurnea: Hollandia, Brass 9003, July 1938, large rain-forest
liane. British New Guinea: Budatobara, Brass 772, Dec. 1925, alt. about 90 m.,
liane in rain-forest. Soromon Istanps: Ysabel: Tasia, Brass 3273, Dec. 1932,
large liane in lowland rain-forest.
These collections are all in fruit but seem to match the description fairly
well: the species has been reported previously only from Netherlands New
Guinea.
264 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXxVI
Morinda jasminoides A. Cunn. in Hook. Bot. Mag. 61: t. 3351. 1834; F. M. Bailey,
Queensl. Fl. 3: 768. 1900.
NETHERLANDS New Guinea: Angi, Arfak Mountains, Kanehira & Hatusima 13798,
April 1940, alt. 1900 m., scandent in thicket by Iray, Lake Giji. Brirish New GUINEA:
Bella Vista, Central Divider, Brass 5443, Nov. 1933, alt. 1450 m., in forest (small
climber with cream-colored flowers on a globose receptacle; fruit rani -yellow, 1.2-1.3
cm. diameter).
The above cited material seems to be a very good match for the Aus-
tralian collections of this species at hand.
Morinda salomoniensis Engl. Bot. Jahrb. 7: 478. 1886; Val. Bot. Jahrb. 61: 154. 1927.
Nortueast NEw Guinea: Ogeramnang, Clemens 4835, Jan. 1937, alt. about 1750
m., forest-hills (vine; fruit red) ; Sarawaket, Camp Kilanda, Clemens 5236, alt. 2100 m
British New Guinea: Matulu, Brass 5187, Oct. 1933, alt. 1250 m., in tall Fisinet
(large liane with rigid branchlets; leaves very dark green; fruit red).
These collections all seem to agree with the description of Morinda salo-
moniensis Engl.; there is only one character shown here which is not men-
tioned in the original description: the corolla-tube as well as the throat is
hairy within.
Morinda hirtella sp. nov.
Planta scandens; ramis hirtellis, brunnescentibus vel viridescentibus:
internodiis novellis compressis, + angulatis; stipulis hirtellis, novellis
probabiliter tubulatis cito uno latere fissis, 3-4 mm. longis, truncatis; foliis
latis, ll X 3.4 cm., tenuiter coriaceis, sei plerumque abrupte acuminatis,
interdum ee cum acumine 3—10 mm. longo acuto, basi rotundatis vel
novellis cuneatis, supra sparsim et aa: hirtellis, subtus (costa et nervis
densius) non dense hirtellis, nervis lateralibus utrinsecus 7—9 oblique ascen-
+ 9- em pects pedicellis 1 -20, gracilibus, rs 1.5 cm. longis,
hirtellis; floribus ut videtur unisexualibus: calyce cupuliformi, integro
glabro; corolla nee puberula; tubo 1 mm. longo, lobis 3-5, circiter 4 mm.
longis, oblongis, intus dense longe albo- sea apice glabris: staminibus
3-5 tubo inter lobos insertis, filamentis brevibus, antheris 2.5 mm. longis
sai ad basim affixis; stylo non viso; disco prominente; capitulis fructiferis
n sicco circiter 10 mm. diametro, 6-8 mm ._ altis; baccis in parte superiore
vix liberis, pyrenis 3 vel 4 obovatis, dorso convexis, osseis.
NortHeast New Gutnea: Above Heldsbach, oo 1957, March 1936, alt. +
00 m., vine on roadside (fruit avid orange). TISH New GuINEA: Lower FI
River, east bank opposite Sturt Island, Brass me pe PE), rain-forest, large canopy
liane of the ridges (branches corrugated; flowers green).
This is another of the species with umbellate inflorescences, in some ways
suggesting Morinda mollis A. Gray, of Polynesia, but with many more
pedicels to the umbel and with glabrous calyces. With so little material we
are unable to say whether the species always has unisexual flowers or not,
but in the six flowers (taken from two heads in different umbels) examined
we did not find even a rudimentary style; the corolla is mostly 4- or 5-lobed,
but occasionally 3-lobed ones were observed.
1945] MERRILL & PERRY, PLANTAE ARCHBOLDIANAE, XVI 265
Morinda micrantha Val. Bot. Jahrb. 61: 153. 1927.
British NEw Guinea: Lake Daviumbu, Middle Fly River, Brass 7756, Sept. 1936
rain-forest substage liane (fruit-heads orange-yellow, less than 1 cm. diameter) ; Tarara,
Wassi Kussa River, Brass 8505, 8675, Dec. 1936, Jan. 1937, large liane, common in rain-
forests (flowers green; fruits orange-colored, 5-8 mm. diameter)
Described from Northeast New Guinea. The three cited collections
indicate the variation in the species; the leaves vary from 5 to II cm. in
length and 2-5.7 cm. in breadth, the leaf-tips in some instances with an
acumen | cm. long, in others almost rounded with a mucro 2 mm. long; the
umbels have 10-20 pedicels each bearing a 5—9-flowered head; the corolla-
tube is about 1 mm., the lobes 2.5 mm. long.
Morinda oligocephala sp. nov.
Frutex scandens glaber; ramulis subteretibus; internodiis 2.5—5 cm.
longis, novellis gracilibus; dd parvis, late ovatis, obtusis, caducis;
foliis 5-10 cm. longis, 2—5 cm. latis, anguste ellipticis vel ovato-ellipticis,
apice breviter acuminatis vel qe, basi rotundato-cuneatis vel late obtusis,
tenuiter coriaceis vel rigide chartaceis, nervis lateralibus utrinsecus 6-8
utrinque perspicuis patenti-adscendentibus, versus marginem arcuatis et
confluentibus, subtus in axillis domatiiferis, reticulo laxo utrinque promi-
nulo; petiolo 5-10 mm. longo, canaliculato; inflorescentiis umbellatis,
Saree et interdum in axillis foliorum superiorum dispositis; capitulis
- vel 3-floris, pedicellatis, pedicellis 4, circiter 5 mm. longis; calyce vix
i 5 mm. longo, minute 4- vel 5-dentato; corollae tubo 4 mm. longo, extus
glabro, intus pubescente, lobis 4 vel 5, oblongis, 5 mm. longis, extus glabris,
intus dense albo-barbatis; staminibus 4 vel 5 tubo inter lobos insertis, fila-
mentis brevissimis, antheris subsessilibus, 2.5 mm. longis; stylo 7 mm.
longo, pues 2, 3 mm. longis; vel filamentis 4 mm. longis, antheris
2.5 mm. longis, stylo 4 mm. longo, stigmatibus 3 mm. longis; capitulis
racers non visis.
H New GurInea: Tarara, Wassi Kussa River, Brass 8490 (Type), Dec. 1936,
parporses of light rain-forest (ale shrub; flowers cream-colored
This species is readily distinguished by the rather Brorainentl reticulate
leaves, the predominantly 4-pedicelled umbels, and the few-flowered heads.
The umbels are sessile, subtended by two leaves, but these sometimes have
fallen, giving the impression at first glance of a pedunculate umbel.
Galium Linnaeus
Galium asperifolium Wall. Fl. Ind. 1: 381. 1820; van Steenis, Bull. Jard. Bot. Buitenz.
Ill. :
NETHERLANDS NEw GuINnEA: Angi, Arfak Mountains, Kanehira & Hatusima 136066,
April 1940, alt. 1900 m., in open wet grassy field on inundation-area of Iray River
running to Lake Giji.
According to van Steenis the species is known from SE. Asia, Sumatra,
Java, and the Philippines.
Galium innocuum Migq ae Nederl. Ind. 2: 341. 1857; van Steenis, Bull. Jard. Bot.
Buitenz. III. 13: a 934
ERLANDS NEW aaa Lake geen Brass 9286, Aug. 1938, alt. 3225 m.,
Pease to 50 cm. on tall marsh grass, commo
Van Steenis gives the range of this species as the Philippines, the Moluc-
266 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXVI
cas, Java, and Sumatra. This seems to be the first record of it from New
Guinea.
Galium ee ee nie i Jahrb. 62: 493. 1929; van Steenis, Bull. Jard. Bot.
B _ III. 13: 247.
serait New GUuINEA: “Sees Habbema, Brass 9511, 9548, Aug. 1938, alt. 3225
m., occasional in moist forest-edges ; 7 km. northeast of Wilhelmin na-top, Brass & Myer-
Drees 9859, Sept. 1938, alt. 3560 m., pendent on mossy trees in forest-edges, common.
Previously collected only in Northeast New Guinea (Sarawaket).
— oe DC. Prodr. - 608. 1830; Benth. Fl. Austr. 3: 446. 1867; F. M. Bailey,
nsl. Fl. 3: 782. 190
British New GuINEA: ries rray Pass, Wharton Range, Brass 4730, Aug. 1933, alt.
840 m., common on wet banks of a creek flowing through grassland. Australia
Galium rotundifolium Linn. Sp. Pl. 108. 1753, sensu lato.
Netuertanpns New Guinea: 9 km. northeast of Lake Habbema, Brass 10731, Oct.
for
1933, alt. 2840 m., plentiful amongst coarse grass on banks of a grassland creek, also a
few plants in a fern-brake on the edge of the forest
Probably Brass 9321 from Lake Habbema is also a depauperate form.
This is perhaps the same species which F. v. Mueller designated as Galium
javanicum Mig. The whole complex is in need of careful study by a spe-
cialist of the group.
Galium subtrifidum Reinw. in Blume, Bijdr. 944. 1826; DC. Prodr. 4: 594. 1830.
NETHERLANDS New Gurnea: Bele River, 18 km. northeast of Lake Habbema,
Brass 11423, Nov. 1938, alt. 2200 m., trailing and pendent to a meter [in length] on a
dry face of limestone in the forest; Angi, Arfak Mountains, Kanehira & Hatusima
forest at Iray. British NEw GUIN Murray Pass, Wharton Range, Brass 4946,
Aug. 1933, alt. 2840 m., bank of an open eum gully, rare.
The collections seems to be a good match for a Javanese collection so-
named by van Steenis.
Galium bryoides sp. nov.
Planta perennis glabra; caulibus numerosis, ramosis, caespitosis, decum-
bentibus vel humifusis, probabiliter 7-10 cm. lo ongis; ramis quadrangulatis,
internodiis 1-5 mm. longis: foliis quaternis, sessilibus, lineari-lanceolatis
5-4 mm. longis, 0.4—0.8 mm. latis, apice attenuato-acuminatis et mucro-
nulatis, basi leviter angustatis margine planis, patentibus vel recurvatis,
supra enerviis, subtus 1-nerviis, luce permeante laxe reticulato-venosis;
floribus axillaribus solitariis, pedunculis sub anthesin subnullis, gee
usque 0.5 mm. longis; corolla rosea, 4-partita, lobis vix 1 mm. longis,
ovatis; staminibus quam lobis brevioribus; stylis 2 distinctis, stigmate
capitato; fructibus circiter 1 mm. longis, glabris et sub lente minute papil-
latis; embryone tantum leviter curvo.
British New Gutnea: Mount Albert Edward, southwestern slope, Brass 4416
(tyPE), July 1933, alt. 3680 m., several plants in a wet grassland hollow, rare (flowers
pink )
This is a very distinct plant readily recognized by the glabrous char-
acter, the small and slender leaves, the matted habit, the very short pedun-
cles, and the glabrous fruits.
ARNOLD ARBORETUM,
ARVARD UNIVERSITY.
1945] NAST & BAILEY, TROCHODENDRON AND TETRACENTRON, II 267
MORPHOLOGY AND RELATIONSHIPS OF TROCHODENDRON
AND TETRACENTRON
Il. INFLORESCENCE, FLOWER, AND FRUIT
CHARLOTTE G. NAST AND I. W. BAILEY
With five plates
INTRODUCTION
THE FIRsT paper (4) of this series dealt with the morphology of the stem,
root, and leaf of Trochodendron and Tetracentron and with significant
palaeobotanical data. The present paper will deal with the morphology
of the inflorescence, the floral organs, and the fruit, and will conclude the
series with a general discussion of the relationships of these remarkable
dicotyledons. In so doing, we shall present evidence in support of Dr.
Smith’s (10) conclusion regarding the close affinities between the two
genera.
INFLORESCENCE
TROCHODENDRON. The buds of this genus have numerous scales. When
a vegetative bud expands, the smaller outer scales soon drop off, but the
inner larger cataphylls are persistent for a season and become separated by
internodal elongation. A pseudoverticillate cluster of leaves of varying
sizes develops above these cataphylls, and the axis terminates in a maturing
bud. The growth of such a vegetative shoot obviously is monopodial.
When a flower bud of Trochodendron expands, the numerous cataphylls
are early deciduous, and the stem is extended as the primary axis of an
inflorescence, Fig. 1. This rachis usually, but not invariably, terminates
in a flower. It bears numerous elongated bracts, in the axils of most of
which secondary flower-bearing axes are formed. In certain cases, the
lowermost of these structures are branched, thus giving rise to tertiary floral
axes, t. fl., Fig. 1. Even when the rachis terminates in a flower, the upper-
most bracts tend to be reduced in size and have no flowers in their axils.
Elongated bracteoles do not develop on secondary axes except where these
axes are branched. A vegetative bud occurs in the axil of the last sterile
scale at the base of the inflorescence, ax. 6., Fig. 1. This bud extends the
vegetative part of the shoot, which, therefore, is of a sympodial type.
It is evident, accordingly, that the inflorescence of Trochodendron is ter-
minal and that it is a raceme-like pleiochasium with occasional tertiary
floral axes. The occurrence of such tertiary axes and the not infrequent
absence of a terminal flower render difficult the acceptance of Wagner’s
(12) interpretation of the inflorescence as an interrupted ‘“‘primanpleio-
chasium.”’
TETRACENTRON. The vegetative buds of the long shoots of Tetracentron
have two elongated scales, occasionally reduced to one in the axillary bud
268 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
closest to the apex of the shoot. Within the bud scales there is a tiny leaf,
having a terminal bud primordium enclosed within its stipular flange. The
extension of the shoots is by monopodial development.
The flower buds of Tetracentron are borne on short shoots, and at first
are enclosed within the stipular flanges of a leaf (0. /f.) as illustrated in
Fig. 3. Each matured flower bud has two scales, within which are an
incipient inflorescence (imfl. ax.) and a short axis bearing a small leaf (//.)
enclosed within an enveloping scale (br. 3). The subsequent development
of the short axis, Fig. 2, crowds the inflorescence into an apparently lateral
position. That the inflorescence actually is terminal and the leaf-bearing
shoot axillary is indicated, however, by topographical relationships within
the flower bud, Fig. 3. The occurrence of a bud (ax. b.) in the axil of the
leaf (o. /f.) demonstrates that the flower bud is terminal. Similarly, the
occasional formation of a rudimentary bud in the axil of the lower bud
scale (br. 1) indicates that the inflorescence is terminal and that the short
leaf-bearing axis is lateral, i.e. develops from a bud in the axil of the inner
scale (br. 2) of the flower bud. Such an interpretation of the morphology
of the short shoot is supported by the orientation of the scale (br. 3) of the
inner axillary bud (¢. m.). This scale is set at right angles to the scales
(br. 1 and 2) of the flower bud, and its orientation in relation to the axis
of the inflorescence is the same as that of the scales of axillary buds to the
axis of long vegetative shoots.
The apical primordium (t.m.) of the leaf-bearing axillary shoots forms
the flower bud of the next year. It is evident, accordingly, that the inflo-
rescences of Tetracentron are terminal and that the development of the
short shoots is sympodial. The inflorescence is a spike with numerous
sessile flowers, each subtended by a minute bract, Fig. 4, br. The flowers
tend to be arranged in false whorls. Four decussate bracts are usually
yi at the apex of the axis, but occasionally the rachis may terminate
ina er.
FLOWERS
PERIANTH. ‘The sessile, tetramerous flowers of Tetracentron, Fig. 4,
have four sepals oriented opposite to the stamens. The flowers of Trocho-
dendron generally are considered to be without a perianth, but Hutchinson
(7), in defining the Trochodendraceae (Euptelea and Trochodendron),
states that sepals are absent or very minute. Most of the axillary flowers
of the inflorescence have a pair of tiny, unvascularized! scales, br., Figs. 6
and 7, that commonly are attached to the base of the torus or less frequently
at a somewhat lower level. The terminal flower of the rachis may have
three or four of these rudimentary appendages. The higher levels of attach-
ment suggest that the scales may be vestiges of a calyx. It should be noted
in this connection, however, that the broad bases of the scales are decurrent
and embossed, Fig. 7, as are those of the bracts on the rachis and of elon-
1In contrast to the normal elongated bracts and bracteoles of the inflorescence,
which — a single vascular strand. The minute bracts of Tetracentron also have a
single tra
1945] NAST & BAILEY, TROCHODENDRON AND TETRACENTRON, II = 269
gated bracteoles on branched secondary, flower-bearing axes. Further-
more, the androecium is perigynous and the stamens appear to be attached
to the dorsal surfaces of the carpels. The subtending scales may, therefore,
be rudimentary bracteoles that have been upwardly displaced onto the sur-
face of the receptacle. It is unfortunate that there are no vascular strands
in these structures to aid in their interpretation.
ANDROECIUM. The numerous stamens of Trochodendron are borne indi-
vidually upon cushions that tend to be linearly arranged on more or less
embossed and decurrent ridges of the dorsal surface of the carpels, Figs. 6
and 7. The four stamens of Tetracentron are borne alternately to the
carpels and do not exhibit external evidences of perigyny.
The stamens of the two genera are, however, of a fundamentally similar
morphological type. The filaments terminate in a relatively massive in-
stead of a much constricted connective as in Drimys (Bailey and Nast, 2),
and the four sporangia are not conspicuously protuberant as in many dicoty-
ledons. The endothecium is neither as restricted as in Degeneria and
Himantandra (1) nor as extensive as in the Winteraceae, where it com-
pletely jackets the sporangia. In Tetracentron and Trochodendron, it isa
continuous subepidermal layer, not only external to the sporangia, but also
extending across the adaxial and abaxial sides of the connective, Figs. 22
and 23. It differentiates first in the region of the sporangia and subse-
quently in the connective. Its maturation in Trochodendron is completed
before dehiscence of the pollen, whereas its differentiation in the connective
of Tetracentron may be markedly retarded. Dehiscence is longitudinal-
lateral, giving to the anther a bivalvular appearance.
PoLLEN. Van Tieghem (11) described the pollen of Trochodendron as
simple, spherical grains with three pores, the pollen of Tetracentron as
simple, somewhat flattened pentagonal grains with smooth surfaces and
having five pores. He provided no illustrations, and his descriptions are
inadequate and quite inaccurate in the case of Teiracentron.
As indicated in Figs. 24 and 25, the pollen of both genera are tricolpate
with papillate, crest-like thickenings on the floor of each furrow. The
grains have a conspicuous reticulately thickened outer surface, the appear-
ance of which changes at different focal levels. If the outermost surface
of the reticulation is brought into focus, the ridges are rather broad and
the pits are small. At a lower focal level, the ridges appear quite narrow
and the pits considerably wider. This difference in surface views is due
to the conformation of the ridges and can be most convincingly demon-
strated in thin sections of pollen grains cut at right angles to their surface.
Such a section, one micron in thickness, is illustrated in Fig. 26. The basal
part of the ridges is very narrow, whereas the external part is much thicker,
appearing almost bulbous in sectional view. The ridges project outward
from a thick relatively homogeneous laver of the exine, within which is a
thin, very hyaline layer, presumably the intine. It is the latter layer that
forms the floor of the furrows and supports the crest-like thickenings of
papillate exine.
270 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
Although the pollen grains of Tetracentron and Trochodendron are en-
tirely unlike those of the Magnoliaceae and other ranalian families having
monocolpate and derived types of pollen, they are of a general morpho-
logical type that occurs at times in other dicotyledonous families, e.g. in
Disanthus (Fig. 27) of the Hamamelidaceae.
GyNoeciuM. The carpels of Tetracentron and Trochodendron are later-
ally concrescent, with free styles, Figs. 4 and 7. As in Degeneria, the sec-
tion Tasmannia of Drimys, and Himantandra, the megasporophylls are
adaxially folded or conduplicate with their ventral (upper) surfaces closely
approximated, Figs. 9 and 72, thus leaving a cleft-like opening into the
locule. The margins of the carpels are free except at the base of the svn-
carpous gynoecium, level A, Fig. 7. The conduplicate form of the carpel
is not confined to the region of the locule but occurs throughout the style,
Figs. 10 and 13.
The carpels have a conspicuous dorsal bulge which is greatly accentuated
in Tetracentron, d. blg., Figs. 4, 12 and 13, and is nectariferous according
to van Tieghem (11) in Trochodendron, Fig. 7. Insects evidently are
attracted to this succulent parenchymatous region, for the dorsal bulge has
frequently been chewed away in herbarium specimens of Trochodendron,
Fig. 6
In Tetracentron, where abaxial deformation of the carpels does not occur
until after anthesis, the placentae are oriented parallel to the vertical axis
of the flower and the ovules are set at right angles to this axis. In Trocho-
dendron, where more or less abaxial deformation occurs prior to anthesis,
the numerous ovules are set at right angles to the placentae, which are
oriented in various diagonal positions in relation to the vertical axis of the
flower, Fig. 7. The placentae are situated in the upper part of the locule,
and particularly in the case of Trochodendron far back from the free
margins of the conduplicate megasporophyll, Fig. 9.
Ovutes. The ovules of Trochodendron are considered by van Tieghem
(11) to have a unique development and morphological form, unlike that of
any other known representative of the angiosperms. Each ovule has a
pronounced sub-chalazal extension, Fig. 15, in which the vascular strand
(or the procambium) makes a “hair-pin” turn. The nucellus and integu-
ments develop laterally instead of terminally on the ovulary lobe, Fig. 14.
The sub-chalazal projection, therefore, is actually an extension of the apex
of the ovulary lobe and not to be confused with the unvascularized sub-
chalazal emergences of Bilbergia and other genera.
Our observations, based upon somewhat inadequate herbarium material,
tend to support van Tieghem’s generalizations regarding the ovules of
Trochodendron, and indicate that homologous, but less accentuated, char-
acters occur in the ovules of Tetracentron. The sub-chalazal extension is
present in ovules at an early stage of development where no procambium is
discernible, Fig. 74, and also in younger ovules with incipient integuments
and no evidences of a megaspore.
The matured ovules have two integuments. The outer one consists of
1945] NAST & BAILEY, TROCHODENDRON AND TETRACENTRON, II 271
three layers of cells in both Tetracentron and Trochodendron, and is more
massively developed at the micropylar end of the ovules. The inner in-
tegument is composed of two layers of cells.
VASCULARIZATION OF THE FLOWER. The elongated pedicel of Trocho-
dendron contains a eustele composed of a varying number of vascular
bundles. These bundles tend to increase in number toward the base of the
flower. They branch and anastomose, forming new interfascicular regions
as in the pedicel of Drimys (Nast, 8). Certain of them become cortical
bundles”, cor. b., Fig. 7, at the base of the receptacle. They are variable
in number, commonly about as numerous as the stamen-bearing ridges on
the dorsal surface of the carpels. Each of the cortical bundles tends to give
rise to three or four traces, one for each of the stamens of a particular ridge.
Occasionally the cortical bundles branch and form twice as many staminal
traces. Rarely does the individual trace of a stamen arise directly from
the eustele.
The dorsal traces of the carpels diverge from the eustele at a level slightly
below that of the base of the locules, d.t., Fig. 7. They usually are discrete
strands, although occasionally one may arise from a stelar bundle common
to it and either a ventral or cortical bundle. The dorsal trace forms, level
B, Fig. 7, two lateral branches, /.d.t., Figs. 7 and 8. Toward the upper part
of the carpel, the median and lateral dorsal strands divide and anastomose,
forming a network of strands (Fig. 7 shows the network of half of one
carpel). A number of tiny branches of the dorsal strands extend a short
distance into the parenchymatous dorsal bulge of the carpels and end
blindly, a, Figs. 7 and 9.
The bundles which ultimately form the ventral traces of the carpels
usually arise from stelar bundles that are common to them and to the
cortical bundles which form the traces of the stamens, Fig. 7, right. At the
level of the bases of the carpels, there is one of these bundles in each of the
septa formed by the lateral concrescence of carpels, v.b., Fig. 7. Ata
higher level, evel B, Figs. 7 and 8, these bundles divide, their halves be-
coming ventral traces of the adjacent carpels, v.t., Fig. 7. Most of the
ovules are vascularized by branches of the two ventral traces. However, a
dorsal strand anastomoses with each ventral bundle and the uppermost
ovules are, therefore, vascularized by these fused strands, as in many of the
Winteraceae (Bailey and Nast, 3).
The bundles of the eustele at the base of the sessile flowers of Tetra-
centron become abruptly and almost simultaneously dissociated into traces.
The four large and very short outermost bundles, which are basically cor-
tical, cor. b., Fig. 11, produce four calycine traces, one to each sepal, and
four staminal traces, one to each stamen. Occasionally a sepal may have
an extra trace, Fig. 12. The vascularization of the carpels is fundamentally
similar to that of the carpels of Trochodendron. Ordinarily there is a large
dorsal trace, d.t., Fig. 11, in the base of each carpel. This trace forms three
strands, Fig. 11, carpel at right, which divide, Fig. 11, anastomose, Fig. 12,
2 For definition of term refer to Nast (8).
272 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XxvI
and may redivide, Fig. 13. There are, in addition, four bundles which form
ventral traces. At the base of the carpels, one of these bundles occurs in
each of the septa of the laterally concrescent carpels. As in Trochoden-
dron, these bundles divide to form ventral traces of adjacent carpels, Figs.
77 and 12. Both the ventral and the dorsal strands extend upward to the
apex of the style. There is no fusion of ventral and dorsal strands in the
vascularization of ovules, as in Trochodendron, possibly owing to the reduc-
tion in size of the carpels and in the number of ovules.
FRUIT AND SEED
The fruit of Trochodendron is a follicetum with ventral, loculicidal de-
hiscence. It is somewhat flattened at the apex, Fig. 6, with the styles of the
follicles bent outwardly, forming spur-like appendages. In the develop-
ment of the fruit the conduplicate ventral side of the sporophyll enlarges
disproportionately to the dorsal side, thus displacing the styles and causing
the fruit to become flattened at the distal end. The placentae, which are
in a diagonal position in the flower, Fig. 7, assume in the fruit an apical or
horizontal orientation with the seeds pendent in the locules. The fertile
and interspersed sterile seeds are in two rows, one row on each placenta.
The raphes or dorsal sides of the seeds are arranged back to back. This
orientation of the seeds is the same as that of the ovules, which are hetero-
tropous according to Agardh’s terminology (Gray, 6, p. 282).
The fruits of Tetracentron, although superficially very different, are
essentially similar in structure. The fruit of Tetracentron also dehisces
loculicidally. However, the abaxial deformation of the megasporophyll,
due to the extreme development of the conduplicate ventral side, is much
more exaggerated in Tetracentron than in Trochodendron. The dorsal side
or bulge, d. blg., Figs. 4 and 5, retains approximately its original size. The
ventral side, however, increases to such an extent that the stvles, which are
upright in the flower, assume a basal position, Figs. 4 and 5. There is also
greater growth on the ventral or grooved sides of the styles than on the
dorsal sides, so that the stvles also curve abaxially, thus producing four
hooked spurs in the fruit. With the over-extention of the ventral sides of
the sporophylls, the placentae are brought from a_ vertical position at
anthesis to a horizontal and apical position in the fruit. The seeds are thus
pendent, with their dorsal sides or raphes back to back, as in Trochoden-
ron.
The seeds of the two genera are strikingly similar in their anatomy. In
both Trochodendron and Tetracentron, the raphe forms an embossed ridge
throughout the length of the seed. Within this ridge is the raphe-bundle,
which is surrounded by thick-walled cells, Figs. 78 and 19. The sub-
chalazal ends of the seeds are extended into a projection which is vascu-
larized, Figs. 16 and 17. The funiculus is fairly short, with the micropylar
end of the outer integument greatly enlarged and encroached upon it. The
average size of the seed of Tetracentron is slightly smaller than that of
Trochodendron.
1945] NAST & BAILEY, TROCHODENDRON AND TETRACENTRON, II = 273
Van Tieghem (11) and Netolitzsky? (9) report two seed coats in Trocho-
dendron. According to van Tieghem, the internal coat arises from the
inner integument of the ovule and is composed of two layers of thin-walled,
colorless cells. The outer seed coat is brown and has three layers, of which
the medium is sclerous and arises from the outer integument of the ovule.
In the case of Tetracentron, he merely mentions that the seed has a thick
external integument, lacunous and soft.
ur observations on the seed coats do not agree with van Tieghem’s de-
scriptions. The three-layered outer integument of the ovule matures into
an outer seed coat of thin-walled cells in both Trochodendron and Tetra-
centron, Figs. 18, 19, and 21a. In Trochodendron, the cells of the inner-
most layer of the external seed coat have thickened (cutinized?) anticlinal
walls and a thickened external (i.e. adjacent to the inner seed coat) peri-
clinal wall. It has, therefore, typical epidermal characteristics. The cells
of the other two layers become flattened radially, Fig. 19, but are greatly
extended longitudinally, giving a striated appearance to the body of the
matured seed when cleared in NaOH. In Tetracentron, this inner layer is
only slightly thickened, but the outer seed coat forms ridges by increase in
size and number of its cells. These cells grow and expand into any avail-
able space between the seeds, which are tightly packed in the fruit. The
resulting ridges give a winged appearance to the seeds of Tetracentron,
Figs. 16 and 18, opposite raphe.
The external layer of the inner seed coat of both genera is sclerenchy-
matous. The cells are small in cross sectional diameter, Figs. 18, 19, and
21a, but greatly elongated, Fig. 20. This layer or sclerotesta also appears
striated, due to its elongated cells, and is readily visible in cleared seeds,
Figs. 16 and 17. Internal to the sclerotesta are thin-walled cells which are
derived from the inner layer of the internal integument and the nucellus of
the ovule.
The endosperm, which constitutes the greatest volume of the seed, is
composed of thin-walled, isodiametric cells. The tiny embryo, embedded
in the endosperm at the micropylar end of the seed, is either undifferentiated
or has incipient cotyledonary development, Fig. 20.
In the sterile seeds, the endosperm is usually absent (a small amount is
present in occasional sterile seeds of Tetracentron). Often the sclerenchy-
matous layer is all that remains of the inner seed coat. The inner layers of
the outer seed coat, especially of Trochodendron, become densely scleren-
chymatous. The numerous sterile seeds of Trochodendron are darkly and
conspicuously castaneous.
RELATIONSHIPS
Although Tetracentron and Trochodendron differ conspicuously in their
habit of growth, in the form and vascularization of their leaves, and in the
grosser characters of their inflorescences and flowers, they exhibit many
peculiarities of internal structure, organization, and development that are
3 Netolitzsky’s description is apparently taken from van Tieghem.
274 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
fundamentally similar and that serve to differentiate these plants from other
woody ranalian genera.
As emphasized in the first paper (4) of this series, the woods of Tetra-
centron and Trochodendron are remarkably similar and differ from those of
other known angiosperms, including the vesselless Winteraceae. Particu-
larly significant in this connection are transitions from typical transverse to
diagonal and longitudinal planes of cell division in the maturation of wood
parenchyma strands. his aberrant trend of cytological specialization is
much exaggerated in Tetracentron.
The secondary phloems of the two genera are of the same morphological
type. They do not have the precocious flaring of the multiseriate rays and
the early stratification into narrow, alternating arcs of soft bast and fibers
that occur so characteristically in the Magnoliaceae (sensu stricto), De-
generiaceae, and Annonaceae. Nor is there a sclerification of the multi-
seriate rays Close to the cambium, as in Euptelea and many Winteraceae.
Although typical leaves of adult Trochodendron are exstipulate with
multilacunar nodal attachments and commonly without buds in their axils,
the earlier leaves formed by seedlings resemble the leaves of Tetracentron
in having trilacunar nodal attachments and sheathing leaf bases that en-
close axillary buds. Furthermore, the stomata of the two genera are of a
peculiar and similar structural type.
There are similar tendencies toward reduction of internodal elongation
both in the vegetative axes and the inflorescences, leading in Trochodendron
to the periodic formation of pseudoverticils of leaves and to the clustering
of axillary flowers on the upper part of the rachis, and in Tetracentron to
the production of short shoots and of false whorls of sessile flowers on the
rachis.
The complex patterns of vascularization of the flowers are essentially
similar in both genera. The form of the open, conduplicate, laterally con-
crescent carpels is fundamentally the same. There is a similar abaxial de-
formation of the follicles which is merely more extensive in Tetracentron
than in Trochodendron. The placentation, stamens, pollen, ovules, and
seeds exhibit minor differences only. The vascularized, sub-chalazal pro-
jections of the ovules and seeds appear to be as unique among angiosperms
as is the structure of the vesselless xylem.
The totality of the developmental, anatomical, and other morphological
similarities between Trochodendron and Tetracentron is so large as to leave
no doubt regarding the close affinities of the two genera. Whether the
plants should be placed in a single family or in two separate but closely
related monotypic families depends upon the taxonomic emphasis that is
placed upon obvious differences in their foliage, inflorescences, and flowers.
As will be shown in subsequent papers by Dr. Smith and ourselves, the
genus Euptelea exhibits no significant similarities to Tetracentron and
Trochodendron in any of its vegetative or reproductive parts and, therefore,
cannot be associated with Trochodendron in the Trochodendraceae.
It should be emphasized in conclusion that there are no cogent evidences
19451 NAST & BAILEY, TROCHODENDRON AND TETRACENTRON, II ZS
of close relationship between either Tetracentron or Trochodendron and the
Magnoliaceae (sensu stricto), Degeneriaceae, Himantandraceae, Wintera-
ceae, Schisandraceae, Cercidiphyllaceae, or Eucommiaceae.
LITERATURE CITED
_
Baitey, I. W., CuHartotteE G. Nast, and A. C. SmitH. The family Himantan-
draceae. Jour. Arnold Arb. 24: 190-206. pl. 1-6. 1943
2, ———— and Cuartotte G. Nast. The comparative morphology of the Win-
teraceae, I. Pollen and stamens. Jour. Arnold Arb. 24: 340-346. pl. 1-3. 1943.
3. ————— and —————.. III. _Carpels. Jour. Arnold Arb. 24: 472-481. pl. 1-0.
1943.
4, ————— and —————. Morphology and relationships of Trochodendron and
Tetracentron, I. Stem, ‘root, and leaf. Jour. Arnold Arb. 26: 143-154. pl. 1-6
1945.
mn
Foster, agri S. The se a of ee aralioides Sieb. &
Zucc. Jour. Arnold Arb. 26: 155-162
Gray, = The botanical a if ce botany. (Sixth edition.)
1879.
&
~
HUvuTCHINSON, : The families of flowering plants, I. Dicotyledons. 1926.
co
. Nast, CHARLOTTE G. The mparative morphology of the Winteraceae, VI.
erga pared of the eee shoot. Jour. Arnold Arb. 25: 454-466. pl. 1-4
1944.
\o
. NETOLITZSKY, Fritz. Anatomie a Pega eas Linsbauer’s Hand-
buch der Pflanzenanatomie, Band X
10. SmirH, A. C. A Rarer review of adios and Tetracentron. Jour.
Arnold Arb. 26: 123-142.
11. TrecHEM, P. van. Sur ks crabes du groupe des Homoxylées. Jour. de
Bot. 4: 259- 297, 330-36 900.
12. WaGNER, RUDOLPH. i zur Kenntnis der Gattung Lecco Sieb. et
Zucc. Ann. RenchRe Hofmus. Wien 18: 409-422. f. 7, 2. 1903.
276 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
EXPLANATION OF PLATES
PraTeE I
Ax. b., axillary bud; br., bract; br. sc., bract scar; d. blg., dorsal bulge; f. w., floral
whorl; jail. ax., ‘nflmescante de erry sc., inflorescence scar; /f., leaf; Lf. sc., lea scar;
n. br., naked bract; o. If., old leaf; pd., peduncle; st. pd., stamen pad; st. sc., stamen
scar; sty., style; t. m., terminal meristem; ¢. fl., tertiary flower; v. sw., ventral suture.
Fic. 1. Trochodendron. Diagrammatic drawing of an inflorescence. Fic. 2. Te
centron. Diagrammatic drawing of an opening floral bud on a short shoot; internodes
rion Fic. 3. Tetracentron. Diagram of floral bud. Fic. 4. Faivaianicon, Flower,
approx. xX 26. Fic. 5. Tetracentron. Fruit, approx. x 29. Fic. 6. Trochodendron.
Fruit, approx. x 7.
Prate II
A,, vascular strands in dorsal bulge; br., bract; cor. b., cortical bundle; d.b., dorsal
bundle; d.t., dorsal trace; fil., filament of ‘et en; Ld.b., lateral- ee bundle; ov. t.,
ovulary trace; st. cu., staminal cushion; st. sc., sae inal scar; st. t., staminal trace;
v.0:; aren bundle; v. t., ventral baa
Fic. 7. Trochodendron. bee two carpels removed at Jevel A and two removed at
level Bt to show cross sectio view; perpendicular shading lines indicate cut surfaces;
epidermal surface represented rea sHipolina: only half of dorsal vascular system shown
in one carpel; approx. xX 18
Pate III
A., vascular strands in dorsal bulge; cor. b., cortical bundle; d.b., dorsal bundle; d.
blg., dorsal bulge; d.l.b., dorsal-lateral bundle; d.l.t., dorsal-lateral trace; d.t., dorsal
trace; /. se. b., lateral sana trace; ov. t., nvulats trace; se., sepal; se. t., sepal trace;
st., stamen; st. t., stamen trace; v. ¢t., ventral trace
Fic. 8. Trochodendron. Transverse section of flower from region comparable to level
B in Fig. ue x 12. Fic. 9. The same. Transverse eer rag placentae, x 1
Fic. 10. The e. Transverse section through styles, 12. Fics. 11-13. Fatvaceuinan:
Transverse ates through flower comparable to sortie in Figs 8-10, approx. x 31.
Priate IV
Em., embryo; end., endosperm; fu., funiculus; 7. int., inner integument; mi.,
pyle; o. int., outer integument; ov. b., ovulary bundle; ra., raphe; sc. 1. nS et
tous lay
Fic. 14. Tetracentron. Young ovule, approx. X 114. Fic. 15. Trochodendron. Young
ovule, approx. x 76. Fic. 16. Tetracentron. Fertile seed cleared in NaOH, approx.
x 24. Fr
17. Trochodendron. Fertile seed cleared in N approx 24 1G. 18
Tetracentron. Transverse section through middle of fertile seed, app x 84. Fic. 19
Trochodendron. Transverse section through middle of fertile seed, appr x 84 I
83. Fic. 21a. Trochodendron. Senipeouieionian section through outer seed coat and scle-
renchymatous layer of fertile seed, approx. x 764.
Pate V
Fic. 22. Trochodendron. Transverse section through anther, x 126. Fic. 23. Tet
centron. Transverse section through anther, x 103. Fic. 24. Trochodendron. Pollen
grain, pap., papillae; bg a of seeing ion, X 1390. Fic. 25. Tetracentron. Pollen
grain, X 1390. Fic ochodendron. Section of pollen grain showing structure of
wall, approx. x ve eae 27: Disanthus cercidifolius Maxim. Pollen grain, x 1390.
BIOLOGICAL LABORATORIES,
Harvarp UNIVERSITY.
Jour. Arnotp Ars. Voi. XXVI PiateE I
MorpuHOLoGy OF TROCHODENDRON AND ‘TETRACENTRON
Jour. Arnoip Ars. VoL. XXVI PLATE I]
MorPHOLOGY OF TROCHODENDRON AND TETRACENTRON
Jour. Arnotp Ars. Voit. XXVI Prate III
MorpHOLOGY OF TROCHODENDRON AND TETRACENTRON
Jour. Arnotp Ars. Voit. XXVI Pirate IV
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1945 ] REHDER, NOMINA CONSERVANDA 277
MORACEAE, HIPPOCASTANACEAE ET VITACEAE,
NOMINA CONSERVANDA
ALFRED REHDER
COMPARATIVELY little attention has been paid to names of families and
their validity under the International Rules of Botanical Nomenclature.
The only proposal for the conservation of names of families is the list of
186 names proposed by J. Lanjouw and T. A. Sprague on pp. 64—65 of the
Synopsis of Proposals concerning Nomenclature prepared by T. A. Sprague
and submitted to the Sixth International Botanical Congress at Amsterdam
in 1935. This list was voted upon at the Congress and adopted without
change (see Proc. Sixth Intern. Bot. Congr. 1: 358. 1936). The list,
however, contains only names employed by Bentham and Hooker f., Gene
Plantarum, and also some by Engler & Gilg, Syllabus der Pflanzenfamilien
(ed. 9 & 10), formed according to Art. 23 of the Rules and the few excep-
tions conserved under Art. 23, Exceptions (1) and (3). The conservation
of additional names was left by the last Botanical Congress to the Special
Committee for Phanerogamae and Pteridophyta (see op. cit. 358, 381).
The three names proposed here for conservation are not included in the
list referred to above, as they were not used by Bentham and Hooker, but
they have been generally accepted by most recent authors, including Engler
& Gilg, and they would be the correct names for the three families if each of
them were not antedated by an older validly published and correctly formed
name. Like these three, a number of other family names proposed by
Horaninov have been entirely overlooked and have been credited to later
authors, as Myricaceae, Juglandaceae, Calycanthaceae, Loganiaceae, Scro-
phulariaceae, and others
ee eer ey, Veg. Kingd. 266 (1846), sensu stricto. — Bureau in De Candolle,
17: 211 (1873), sensu stricto. — Engler in Nat. Pflanzenfam. III. 1: 66
nad — Nakai, Fl. Sylv. Kor. 19: 90 (1933), with many synonyms. — Nomen
conservandum
versus
Artocarpaceae Horaninov, Prim. Lin. Syst. Nat. 62 (1834).— Lindley, Veg. Kingd. 269
6), sensu stricto. — Bureau in De Candolle, Prodr. 17: 280 (1873), sensu
stricto
doccing ay ul. Artocarpeae De Candolle in Lamarck & De Candolle, Fl. Frang. ed.
8 (1805).
Peiicane pars Aphor. Bot. 203 (1825), p. p.
Moreae Endlicher, Prodr. Fl. Norfolk. 40 (1833), nom.; Gen 7 (1836).
Urticaceae 1. Artocarpeae Reichenbach, Handb. Nat. eal oe (1837), nom.
Urticaceae subord. Moreae Gray, Bot. Text-book, 356 (1842).
Urticaceae subfam. Ficeae Presl, WSeob. Rostl. 2: 1365 (1846).
Moraceae subfam. Moroideae et Artocarpoideae A. Braun in Ascherson, FI. Prov.
Brandenb. 1: 57 (1864).
278 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
As the name Moraceae has been used by all authors who considered the
family distinct from Urticaceae, it seems advisable to conserve it for that
important family, which includes a number of genera of great economic
importance. The name Artocarpaceae Horan. (emend.) would then be
valid only as the name of a separate family distinct from Moraceae, as used
by Lindley in 1846 and by Bureau in 1873; in his original concept Horani-
nov used the name in a wider sense, panes Morus and related genera.
Hippocastanaceae Torrey : Gray, Fl. . 1: 250 (1838).— Pax in Nat. Pflanzen-
-_ III. 5: 273 (1895) aus Mian panera
versu
Poe Horaninov, Prim. Lin. Syst. Nat. 100 (1834).
Sapindaceae Jussieu in Ann. Mus. Hist. Nat. Paris, 18: 476 (1811), p. p
sai ari ip De Candolle[Théor. Elém. Bot., ed. 2, 44 (1819) ‘“Hippocastanées” | ;
Prodr. 1: 597 (1824).
— Link, Enum. PI. Hort. Berol. 1: 354 (1821), nom. subnud.;
rently based on ae ie though not in the sense of Miller; it is ae cited
y Link as one of the constituent genera.
oo 3. Sapindeae a. Hippocastaneae Reichenbach, Handb. Nat. Pflanzensyst.
285 (183
Aesculeae Pres] (1820) ex Presl, WSeob. Rostl. 1: 217 (1846).
Sapindaceae trib. Paviariae ee v, Char. Ess. Fam. Reg. Veg. 182 (1847).
Sapindaceae subord. Sapindeae Bentham & Hooker f., en. Pl. 1: 389 (1862), p.
Sapindaceae subfam. san eniR RDS A. Braun in ‘hig. Fl. Prov. oe.
253 (1864).
Sa pindncene 2. Hippocastanae Drude in A. Schenk, Handb. Bot. 3,2: 390 (1887).
As the name Hippocastanaceae has been used since 1838 by all authors
who considered the family distinct from Sapindaceae, and as the name
Paviaceae, though validly published by Horaninov in 1834, has never been
taken up, as far as I know, by any author, it seems savin to conserve
the name Hippocastanaceae for this well known, though small family, of
ornamental trees and shrubs lana planted throughout the temperate zone.
Vitaceae rare Nat. ed. 2, 30 ean — Gilg in Nat. Pflanzenfam, III.
896). — ioe se nee
versus
Leeaceae Horaninoy, Prim. Lin. Syst. Nat. 95 (1834).
Vites Jussieu, Gen. Pl. 267 (1798). — Lindley, Introd. Nat. Syst. Bot. 119 (1830).
1799).
Viniferae Jussieu in Mém. Mus. Hist. Nat. Paris, 3: 444 (1817).
Ampelideae Humboldt, Bonpland & Kunth, Nov. Gen. 5: 223 (fol. ed. p. 172)
(1821).— De Candolle, Prodr. 1: 627 (1824). — Bentham & Hooker f., Gen. Pl.
1: 386 (1862).
Celastraceae ... Cisseae Horaninov, Tetractys Nat. 32 (1843), nom.
Cissaceae Horaninov, Char. Ess. Fam. Reg. ies 184 (1847).
Ampelidaceae Lowe, Man. FI. Were. 80 (186
As Vitaceae is the only name formed cies to the Rules which has
been generally accepted. while Leeaceae has not been taken up by any
author and moreover is based on a genus not typical of the family, it seems
advisable to conserve the name Vitaceae. Horaninov himself abandoned
1945] REHDER, NOMINA CONSERVANDA 279
the name in his two later works, making the group in 1843 under the name
Cisseae a subdivision of Celastraceae, and in 1847 raising it as Cissaceae
again to the rank of a family.
If one were to follow the proposal made by Sprague in 1922 (in Jour. Bot.
60: 69-73), the two last family names discussed would be valid without
conservation as Hippocastanaceae (DC.) Torr. & Gray and as Vitaceae
(Juss.) Lindl. Sprague brings forward the argument that the priority of
a name of a family dates from the first valid publication of a name with
any plural ending based on a generic name, even if it consists of the plural
form of that generic name. In such cases the change to the ending -aceae
is supposed to be an orthographic correction and the original author is to
be cited in parentheses. Sprague’s proposal, however, conflicts with long-
established usage and the very spirit of the Rules. The use of parentheti-
cal authors is specifically restricted by Art. 49 to changes of rank of genera
and groups below genera, and of transfers of groups below the genus with
or without change of rank, but with no alteration in the name or epithet
itself. According to that article, a citation like Hippocastanaceae ( DC.)
Torr. & Gray could mean only that Torrey & Gray transferred a family
named by De Candolle as Hippocastanaceae from one higher group (ordo
of Art. 12 and of Recomm. VIII and IX) to another. Hippocastaneae
DC. and also Vites Juss. are not valid family names according to Art. 23;
they have no standing, and if transferred could have no validity. Ortho-
graphic corrections or changes in the spelling of a name do not call for the
citation in parentheses of the original author; they should be indicated by
citing the original spelling in quotation marks after the citation of the cor-
rect name, or in a note beneath it.
In regard to transfers of names of groups above genera (see note on pp.
68 and 78), it would seem to be in the spirit of the Rules to make the cita-
tion of the parenthetical author obligatory for transfers of the correct names
of subdivisions of families. This could be done by a proposal to insert in
the second paragraph of Art. 49 the word “family” before genus, so that
the paragraph would read: ‘“‘The same holds when a subdivision of a family,
a genus, a species, or a group of lower rank is transferred to another
family, genus or species with or without alteration of rank.”
ARNOLD ARBORETUM,
HARVARD UNIVERSITY.
280, JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
STUDIES IN THE LAURACEAE, VI*
PRELIMINARY SURVEY OF THE MEXICAN AND
CENTRAL AMERICAN SPECIES
CAROLINE K. ALLEN
THE PRESENT paper was undertaken as groundwork for the presentation
of the Lauraceae in the Flora of Panama, which is being published in fasci-
cles by Dr. Robert E. Woodson, Jr., in the Annals of the Missouri Botanical
Garden. Dr. A. J. G. H. Kostermans has published in detail on the smaller
genera of Lauraceae of this hemisphere. Consequently the present treat-
ment of these genera will not be as detailed as those on which he has not
worked. Unfortunately, at the outbreak of World War II Dr. Kostermans
had on loan in Utrecht a large number of specimens of Lauraceae from
various American herbaria. From the material at hand it is apparent that
a great deal of field work on the group is necessary. More complete and
better collections from regions heretofore little collected are important in
order to supplement data already available and to aid in the solution of
distributional problems. In passing, mention should be made of the im-
perative need of corroborative morphological studies in the group.
In each of the four major genera of the Lauraceae represented in our
region, namely Nectandra, Ocotea, Persea, and Phoebe, the majority of the
species clearly belong to the genera under which they are treated. There
are, however, a few species in each genus of which the characters appear to
overlap. It is not my belief that these represent new genera, but that the
lines between the genera are not as yet very clearly defined. An attempt
has been made to evaluate the characters used in delimiting the genera and,
where these characters overlap, to place the entity in question in that genus
with which it seems to have the greatest number of characters in common,
The majority of instances of overlapping occurs among the four genera
mentioned. One species of Ocotea, however, has double-margined cupules
that are typical of Licaria, but the flowers are definitely those of Ocotea.
In the main, the species themselves are clear-cut. On the other hand, there
is in each genus at least one entity which appears to be widespread and
variable throughout Mexico and Central America. The variations are
manifest in the size of the leaves, inflorescences, and flower-parts. Ex-
tremes of variation occur seemingly as a result of change in geographical
or ecological influences. It should be kept in mind that to date our collec-
tions from Mexico and Central America are too scattered to admit of a
sweeping reduction of species described from these different areas. In
Litsea glaucescens, more completely collected than any of the other mem-
bers of the family, one sees the trend toward a single variable species. The
* For No. V in this series see Jour. Arnold Arb. 23: 444-463. 1942.
1945 | ALLEN, STUDIES IN THE LAURACEAE, VI 281
situation in Litsea nevertheless is hardly typical for other genera in this
region. Jitsea is the only genus which occurs in the arid northern areas
of Mexico, south through Central America, and which has no representa-
tives in South America. The genus Persea extends north in the same
region, in Mexico, but is found in South America. The central portion of
Costa Rica seems to have given rise to the largest number of endemic
species. Until such time as Mexico and Central America are well repre-
sented in herbaria, the presence of few widespread variable species as op-
posed to several very closely related but separate species is a matter of
conjecture. There has been heretofore an almost wholesale linkage of
Central American and Colombian entities. Very probably there is a de-
finite and logical connection, but very possibly this connection has been
over-worked and is not as common an occurrence as has been supposed.
This may be illustrated by the South American species of Persea caerulea
and Nectandra Pichurim, neither of which occurs in Central America, but
both of which have been reported as common to both areas. Again, it is
necessary to have available more material than is at hand at present before
disposing of these distributional problems with any degree of accuracy. In
the course of this study many specimens which are too fragmentary or in-
complete for satisfactory description have been set aside until such time as
more complete material will be available.
An effort has been made to locate on modern maps as many as possible
of the localities cited, and the present spelling has usually been used.
Numerous labels of Purpus’ plants mention Zacuapan. This is on recent
maps clearly Axouacapan. From the latter locality to Cordoba numerous
small localities given by Purpus have been located. The collector Austin
Smith mentions various zones on his labels. This zoning is not botanical,
but ornithological, being determined by the presence or absence of certain
key birds. Mention should be made of the term “‘apparent petiole’, which
is used to describe the basal part of a leaf-blade which is narrowly decur-
rent and recurved for so long a distance at its base that the lower portion
of the blade seems to form with the midrib an extremely long petiole. The
apex of certain fruits frequently dries smooth and is not wrinkled as is the
remainder of the surface. This smooth unwrinkled portion sometimes dries
in such a way as to split the outer layer of the exocarp, forming a star-
shaped pattern. The measurements of the flower-parts were made with
the aid of a micrometer device, and some variation in respect to measure-
ment is to be anticipated.
I wish to thank the directors and curators of the following herbaria for
their courtesy in lending material for study: Arnold Arboretum (A), Chi-
cago Natural History Museum (Ch), Gray Herbarium (GH), University
of Michigan (Mich), Missouri Botanical Garden (Mo), New York Botani-
cal Garden (NY), University of Texas (Tex), United States National
Herbarium (US), Yale School of Forestry (Y). The parenthetical letters
indicate the place of deposit of the cited specimens. I am particularly
indebted to Dr. P. C. Standley, of the Chicago Natural History Museum
for his helpful and careful selection of interesting material from Guatemala.
282 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
At this time I should especially like to acknowledge my gratitude to the
late Dean S. J. Record, of the Yale School of Forestry, for his kind interest
in this study.
KEY TO THE MEXICAN AND CENTRAL AMERICAN GENERA
OF LAURACEA
A. Foliose trees or shrubs, not parasitic.
B. Anthers four-celled.
C. Inflorescences variously paniculate, with no involucre.
D. Staminodia large, cordate, stipitate.
E. Perianth-lobes usually unequal or at most subequal; usually a
cells of the anther touching upper at their sides...........
E. i lobes equal; usually lower cells of the anther pa
upper their -DAsGh..i40 coca ecd Meare bauereseiawent as 2. Phoebe.
D. Staminodia gor inconspicuous or occasionally lackin
E. Flowers perfect or dioecious; perianth-lobes not ae at anthesis,
thin in texture; cells of the anthers arranged in two planes, one
3. Oc
above the ot ina cas Wes cha fs pans 4 oa a eee pein dee wasnt ee otea.
E. Flowers always perfect; perianth-lobes usually fleshy, papillose and
reflexed at pets cells of the anthers arranged in an arc........
ite.
D. Flowers with three fertile stamens; fruits subtended by —s with
double or triple margins...........0. 0.00 eee eee e eee nee . Licaria
or by cupules with single margin
E. Staminodia very well developed: perianth-lobes deciduous entirely ;
dak ead tae pa citnneechee sad sasine maine oy acne ryptocarya.
E. Staminodia inconspicuous or absent; perianth ae or in part
persistent; fruits subtended by cupules with single margins.
F. Stigma well developed, peltate; leaf-blades erage on wish
ing; fruits subtended by fleshy cupule and pedicel..... 7. Aiouea
F. Stigma minute, hardly conspicuous; leaf- blades. nt . -green
on drying; fruits subtended by woody cupule and pedicel.......
-
1. Persea Miller
Persea Miller, Gard. Dict. ed. 8. 1768; Gaertner f., Fruct. & Sem. 3: 222. 1805; H.B.K
Nov. Gen. & Sp. 2:125 [157]. 1817; pear in DC. Prodr. 151: 43. 1864;
Bentham in are & Hooker f. Gen. Pl. 3: 156. 1880; Hemsley, Biol. Centr.
Am. Bot. 3: 71. 1882; Mez in Jahrb. ae nd Berlin 5: 134. 1889; Kostermans
in Meded. Bot. Sieh "Utrecht 25: 12.
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 283
DistrisuTIoN: A tropical American genus, with the exception of a single species
from the Canary Islands. A few species are native to the United States, from Virginia
and the Carolinas south to Florida and Texas, but these species do not occur in Mexico.
The West Indian species seem to have nothing in common with other tropical species
of = soca rach link which binds all tropical localities is the avocado pear, Persea
amer: whic s the species proper or in one of its many forms, is to be found
cultivated ae iiceatout the Americas.
Persea is comprised of trees or shrubs with evergreen penninerved leaves
which vary in texture from chartaceous to heavily coriaceous and are
usually pubescent to some degree. The axillary commonly paniculate
inflorescences bear perfect flowers up to 9 mm. long, the customary length
for the genus being 4-5 mm. The 3 outer perianth-lobes are generally
shorter than the inner and almost always persist in fruit. There are four
staminal whorls, the first three of which are usually fertile. The four-
celled anthers are borne on slender hairy filaments two or three times as
long as the anthers. The cells of the anthers of the first series are introrse,
whereas those of the third are extrorse, or the two upper cells may be
lateral and the lower extrorse, the Canc being distinctly biglandular
near the base. The four cells of the anthers are in two planes, the bases of
the two upper cells laterally tangential to the apices of the two lower cells.
The staminodia of the fourth cycle are conspicuous, cordate-sagittate, and
stipitate, often pubescent throughout, the stipes generally densely so. The
ovary is subglobose, pubescent or glabrous, topped by a slender usually
pubescent style that is equal to or more than the length of the ovary. The
fruit is small and globose or, in the case of the avocado-type, large, fleshy,
and obovoid, borne for the most part on the spreading perianth-lobes and
pedicels, generally not enlarged as compared with those of the other genera
of the Lauraceae. Only occasionally are the perianth-lobes deciduous.
KEY TO THE SPECIES OF PERSEA
A. Leaf-blades not more than 12 cm. long
Lower leaf-surface completely glabrou
C. Lower leaf-surface glaucescent ; ee nerves diverging from a costa at
AN ANGIE Olt S590 a sa aiete Mhastitass & ac 4! 5c @ asete Gt maaan . P. longipes.
C. Lower leaf-surface not ee lateral nerves diverging from the costa
at an angle of 90° or sia oak LESS apa anigavaauefiiem havoaiaavaeaiee 2. P. Steyermarkiti.
B. Lower leaf. surface not glabro
C. Leaf-blades ochraceous-sericeous beneath, not more than 7.5 cm. long;
lateral nerves diverging from the costa at an angle of about 90°..........
RE a Pe eT Nee treet we NIA OS ene eer . P. Brenesii.
C. ee blades ferruginous-tomentose beneath, up to 9 cm. long; lateral nerves
erging from the costa at an angle of 20-45°........... 4. P. pachypoda.
A. Leaf- ae not less than 12.5 cm
B. Leaf-blades never sessile and ate
C. Ovary pubescent
. Leaves anise- -scented with long petioles one-half the length of the thin
elliptic blades; perianth persistent or subpersistent..............-.-+-
ssa Te sch F Sa denh aide 1a eH aa Ree eee es . P. americana var. drymifolia.
D. Leaves not anise-scented, with petioles not more than one-quarter the
length of the chartaceous or coriaceous blades, varying from elliptic to
JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
E. Inflorescence, herd bani and lower leaf-surface not densely
golden-brown-serice
F. Branchlets an ‘inieidiice becoming glabrous or subgla-
brous; lower leaf-surface subglaucous; lateral nerves 4-14 pairs;
blades variable, usually elliptic, up to 10.5 cm. broad, acuminate;
fruits variable in shape...................004. . P. americana.
. Branchlets and lower leaf-surface densely brown-tomentose; lat-
_
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Ln J
ry
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pairs; blades ovate, acuminate, up to 7.5 (-8) cm. broad; fruits
depressed-globose(?).......... cece cece eee ee nens 7. P. floccosa.
E. ae ta young branchlets, and lower leaf-surface densely
den- aaa REVICCOUS:. 0445 bok Rad eee a ees 8. P. flavifolia.
C. Ovary cana
“if eared loosely paniculate or loosely subcorymbos'
-blades sometimes obovate or elliptic or comedy round-elliptic.
F. Inflorescence not less than 8 (usually 10-12) cm. long; largest
leaf-blades usually at least 10, menenaig: 15. 0M. DYOAG io okie
ee ee ere ee er rere ere oT ee onnell-Smit hit.
F. Inflorescence not more than 7 cm. long; ale pais blades not
more than 8 cm. bro
G. Leaf-blades 9- hy x 3.2-7 cm.; lateral nerves up to 8 pairs;
TiS DIACK ss 36 ccer cea veers bese salonees 0 hamissonis
G. Leaf- blades 10-20 x 5-8 cm.; lateral nerves 14-16 pairs;
11. P. cinerascens.
e.
*, oe lower leaf-surface, and inflorescence densely ferru-
nous-tomentose............. 02000 e eee 9. P. Donnell-Smit hii.
F. Bechet, veld leaf-surface, and inflorescence not densely
paren -tom
G. Leaf-blades rigidly coriaceous or chartaceous; inflorescences
numerous and more or less corymbose, densely pubescent.
H. Pesanth lobes decidedly unequal.
r leaf-surface and inflorescence subsericeous; peti-
ila up to 4.5 cm. net leaf-blades not more than 10.5
cm. broad; fruits 7 mm. diam., subtended by remnants
of scarcely thickened Eee job es..12. P. Liebmann
I. Lower leaf-surface and inflorescence shortly tomentel-
lous; petioles up to 2 cm. long; leaf-blades not more
than 6.5 cm. broad; fruits at least up to 3.5 cm. diam.,
subtended by a thickened a cupule.....
. P. vesticula.
H. Perianth-lobes equal or subequal.
I. Largest leaf-blades not more than 15 cm. long, glau-
cescent beneath; petioles up to 2 cm. long; branchlets
yellowish-pubescent to glabrous; eatasrren equal to
or slightly shorter than the leaves....... 14. P. pallida.
I. Largest leaf-blades up to 33 cm. long, not glaucescent
beneath; petioles not more than 1.5 cm. long; branch-
lets glabrous; inflorescence much shorter than leaves. .
Y Sd eg atéteadh ie og a we eR EO OE TORS R EN 15. P. rigens.
G. Leaves pergamentaceous; inflorescences not numerous and
corymbose but spreading-paniculate, fulvo- yerar Me aie bess
16. P. Skutchit.
eee ea ee eee ee ee ee ee ee ee ee ee ee ee ee
1945 | ALLEN, STUDIES IN THE LAURACEAE, VI 285
D. Inflorescence with compact shortly branched panicles more or less com-
pressed into a subpyramidal or subcapitate structure at the tips of the
peduncles.
. Lower leaf-surface densely Leesa blades of largest leaves not
less than 6, usually
F. Lower leaf-surface sie -sericeous; petioles to 4.5 cm. long; in-
aus flavo-sericeous, not more than 6 cm. long (rarely 7
TMV) rere ee eo a Gd wins nea meee alerts aie JP:
F. ee leaf-surface whitish or flavo-tomentose, not sericeous;
petioles to 3.5 cm. long; inflorescence whitish- or flavo-tomentose,
Up CO210 Cm LOM Goo a: s so sis, 5's se spore eigiotermesercteceee 17. P. Hintoniz.
E. Lower leaf-surface glabrous or glabrescent (or sericeous to gla-
brescent), eaucescent beneath; blades of largest leaves not more
oo
Rg
°e
s.
bo
i=)
=
a)
-than 5.5 cm. b
F. 2 blades elliptic, a aple acuminate or acute, not more than
5 (occasionally 8) cm.; lateral nerves 6—10
a est blades glabrous, roy margin aia the oe phen
subglaucous; lateral nerves 6-8 p : : Inflorescence not mo
than 6 cm. long; fruits blackish, pee on = remnants of
partially deciduous perianth-lobes............ . P. longipes.
G. Leaf-blades sericeous to glabrescent, eee penean
lateral nerves 8-10 pairs; inflorescence up to
°
eee tlie elu bloom, seated on the patent raster
18. P. veraguasensis.
F. Leaf- Hades lanceolate- elliptic, subobtuse, subacute to acumin ate,
not more than 4.5 cm. broad; lateral nerves 10-14 pai
G. See not more than 2 cm. long; lateral nerves a to 14
pairs, usually obscure beneath; ces care at if present very
HICONSPIGUOUS so cnsahsa:siacate eoeuatelegeccieteters erento 19. P. podadenia.
G. Petioles up to 3.5 cm. long; lateral nerves 10-12 pairs and
nce conspicuous above, very much so beneath; reticulation
picuous, poe beneath........... 20. P. Standleyi.
B. Leaf-blades sessile aa SUDGOLC ALE a ersexs sree crete shore onic 01 S641 oslo Se 21. P. sessilis.
1. Persea anes Seg narige ued heed in DC. Prodr. 151: 55. 1864; Standley in
Contr. U. erb. 23: 922.
Laurus Dasus eeieciendil in eras 7: 390. 1832.
DisTRIBUTION: Known only from Vera Cruz, Mexico.
Mexico: Vera Cruz(?): Hacienda de la Laguna, Schiede 59 (fr., photo. of TYPE,
Ch, NY). Vera Cruz: Mirador, 3/1842, Liebmann Lauraceae 72 (fl., Ch), 756 ?
Ch).
The Liebmann numbers last cited bear the determination P. brevipes,
but since the number 72 is cited by Mez in his monograph under P. longipes
and agrees with the description, it is safe to assume that the name on the
label is erroneous. The flowering specimens cited show densely hang
leafy branchlets blackish brown and ee with long (up to 3.3 ¢
gracefully petioled leaves (2.5-10 [-13] cm. & 1.5—5.5 cm.) eas
and glabrous, the midrib impressed above er the lateral nerves obscure,
the surface somewhat areolate. The midrib and nerves are slightly more
prominent beneath, being brownish against the subglaucescent lower leaf-
surface. The inconspicuous lateral nerves, 6-8 pairs, usually diverge at
an angle of 45°, but vary from 35-55°. The short rather few-flowere
panicles (6 cm. long) are narrow, subtended by glabrous dark brown
slender peduncles up to 4 cm. lon The flowers are ferruginous-sericeous
with the inner lobes of the perianth twice as long as the outer. The
286 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXxvI
stamens of the outer series are + 2.4 mm. long, the pubescent filaments
equalling in length the ovate rounded anthers, with the two upper cells
smaller than the lower. Those of the inner series are + 2.6 mm. long,
the filaments biglandular, the anthers narrower. The staminodia are
+ 1.25 mm. long, broadly cordate, borne on pubescent stipes one-half the
entire length. The glabrous gynaecium is up to 3 mm. long, and the sub-
globose ovary is about one-third the length of the entire — The
slender style is topped by a small inconspicuous discoid stigm
The nearest relatives of this species seem to be the Mexican P. sididonta,
the palish leaf-blades of which are more regularly elliptic or oblong-elliptic,
6-15 < 2-4 cm., with petiole up to 2 cm. long and with 14 pairs of lateral
nerves, and the Costa Rican and Panamanian species P. veraguasensis.
From the former it is readily separated by the broadly and irregularly
elliptic leaf-blades with undulate margins, 2.5-10 (-13) & 1.5-6.5 cm.,
with petiole up to 3.3 cm. long and with 6-8 pairs of lateral nerves. From
the latter species it is separated by the leaf-blades of more uniform shape,
not undulate at the margin, very glaucous and lightly pubescent on the
lower leaf-surface, with usually more than 8 pairs of lateral nerves, the
dark venation being very conspicuous. ‘The branchlets are covered with a
slight puberulence, and the inflorescence is densely clothed with a sub-
ferruginous somewhat sericeous pubescence. The blackish (fide Meissner )
fruits of P. longipes, to judge from the photograph of the type, are sub-
tended by the remnants of the perianth-lobes, the tips of which have broken
off, rather roughly and irregularly, showing only the line of decurrence on
the tube —a characteristic in common with P. podadenia. The bluish
glaucous fruits of P. veraguasensis, on the other hand, are subtended by the
somewhat enlarged remains of the persistent pubescent perianth-lobes,
which are always patent and often slightly reflexed.
2. Persea Steyermarkii, sp. nov.
Arbor parva ad 10.5 m. alta, ramulis apice dense foliosis glabris atro-
rubescentibus striatis mox griseis rugosulis vadose sulcatis. Folia alter-
nata, petiolis atro-rubescentibus glabris striatis canaliculatis ad 2 cm.
longis et 1 mm. vel minus latis, laminis utrinque glabris coriaceis in sicco
supra viridescenti-brunneis, subtus pallidis (caeruleo-argenteis fide coll.),
lanceolato-ellipticis vel oblon ngo- TTT 6.2-10.5 cm. longis et 2.5—4 cm.
latis, basi obtusis, apice rotundatis vel acutis saepe emarginatis, penniner-
viis, costa supra satis impressa subtus valde elevata, nervis 6- vel 7-paribus
nonnihil obscuris pallidis supra leviter impressis subtus leviter elevatis
angulo ad 80° divergentibus, rete venularum utrinque conspicuo. Inflores-
centia axillaris (foliis deciduis) foliis brevior subverticillata, paniculata,
3-4(—S) cm. longa, pauciflora, pedunculo rubescente glabrescente striato
gracili ad 3 cm. longo. Flores ad 5.5-8 mm. longi, pedicellis minute
gracilibus adpresse pubescentibus ad 7-10 mm. longis, perianthio campanu-
lato flavo-viridescente fragrante fide coll., lobis reflexis nonnihil crassis
leviter papillosis exterioribus + 4.5 mm. longis interioribus 6 mm.
longis; staminibus loculis superioribus quam inferioribus brevioribus
4.25 mm. longis, antheris truncato-ovatis longitudine subdimidio sta-
minium aequantibus filamentis pubescentibus; staminibus ser. ITI antheris
emarginatis, biglandulosis, glandulis conspicuis reniformibus_ stipitatis,
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 287
antheris subaequalibus; staminodiis conspicuis, + 3.5 mm. longis cordatis
ere eae pubescente crasso; gynaecio pubescente + 4.25 mm. longo,
eo vel subgloboso, stvlo leviter quam ovario longiore,
State eunenets conspicuo. Fructus ignotus.
DISTRIBUTION: Known only from type-locality.
GUATEMALA: San Marcos: Trail between Finca E] Porvenir and San Sebastian,
upper slopes of Volcan Tajumulco, alt. 13C0-1400 m., an 1, 1940, Stevermark 37061
(fl., Type, Ch) (small tree 9-10.5 m.; leaves ascending, erect, coriaceous, pale or olive-
green above, blue-silvery-green beneath with pee between lateral veins, the
margins revolute; flowers sweet-scented, the petals yellow-greenish, the pedicels pale
green, the anthers deep yello
This species, with ie oniete densely foliose at the apex, seems to have
no particular affinity. Its floral characters seem to be near those of P.
americana and its relatives, but the general habit indicates perhaps a rela-
tionship to the species P. longipes and P. Standleyt.
3. Persea Brenesii Standley in Field Mus. Publ. Bot. 18: 458. 1937.
DistRIBUTION: Forests or pastures of Costa Rica, known only from the type-
locality
Costa Rica: Alajuela: La Palma de San Ramon, alt. 1150 m., Brenes 4451 (fl,
TYPE not seen)
This species is one of the smallest-leaved species of Persea. The most
salient features seem to be the dense ochraceous-sericeous pubescence found
on the leafy thick angled branchlets, the petioles 10-14 mm. long, and the
lower surface of the small (5.5—7.5 1.5—2.5 cm.) obtuse or acutish oblong
leaf-blades glabrous above with the costa and nerves slightly impressed
and elevated beneath. The lateral nerves are about 6 pairs diverging from
the costa at right angles or more narrowly ascending. The inflorescences
are axillary few-flowered corymbs slightly longer than the leaves and
densely ochraceous-sericeous, with flowers sessile or shortly and thickly
in
apparently immature, are globose, glabrous, and measure | cm. in diameter.
The description seems to place the species in the vicinity of Persea
pachypoda, from Mexico, if it is not actually conspecific with it. The type
cannot be located at present.
4, Persea pachypoda Nees in Linnaea 21: 490. 1848
Oreodaphne Benthamiana Nees in Linnaea 21: 521. 1848.
Phoebe Hartwegii Meissner in DC. Prodr. 151: 30. 1864.
Persea Hartwegii Hemsley, Biol. Centr. Am. Bot. 3: 72. ae
Phoebe Benthamiana Mez in Jahrb. Bot. Gart. Berlin 5: 195.
Emote pachypoda Mez in Jahrb. Bot. Gart. Berlin 5: 196. ce ana in Contr.
S. Nat. Herb. 23: 294. 1922
DistrIBUTION: Mexico.
Mexico: ? “Bei el bruco, Ehrenberg 942, Arbor, fructu eduli, Apugato cimaron
incolis.” ({r., fragm. of TyPE of Persea pechvncdae Ch). uanajuato: AHartweg 84
(fl., fragm. of type a Oreodaphne Benthamiana, and syntype of Phoebe Hartwegit,
NY). San Luis Potosi: Minas de San Rafael, Bagre, June 1911, Purpus 5338,
5457 (fl., Ch).
NATIVE NAMES: “Aguacate cimarron,” “Apugato cimarron” (Mexico).
This small-leaved species has apical branchlets covered with a dense
288 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXVI
pale ferruginous tomentum, later becoming fuscous and black. The oblong
r oblong-lanceolate acutish or round-apiculate leaf-blades are coriaceous,
cuneate at the base, and measure 9.5 3(-5) cm. They are densely
ferruginous-tomentose throughout, becoming less so with age, with a
tomentose petiole up to 2 cm. long. The 4-6 pairs of lateral nerves are
rather obscure although slightly elevated above, but, like the costa, are
prominently elevated beneath, and diverge from the costa at an angle of
between 20 and 30 (—45)°. The short densely ferruginous-tomentose
axillary inflorescence, up to 6.5 cm. long, consists of several depauperate
panicles which are shortly branched and almost corymbose, borne on long
peduncles up to 4 cm. in length. The densely tomentose flowers are about
6 mm. long, subsessile or borne on pedicels slightly more than 1 mm. long.
The outer perianth-lobes are ovate and acuminate, with 5 prominent veins.
+ 3.5 mm. long, and the inner are rather elliptic, acute, and + 5 mm. long.
The two inner series of stamens are + 3 mm. long, the pubescent filaments
being slightly longer than the apiculate gi The stamens of the third
series are biglandular and measure + 4 mm., .the stipitate subreniform
glands nearly equalling the anthers. The staminodia are stipitate, = =e
mm. long in their entirety, ovate, cordate, and about one-half again as long
as the rather stout pubescent stipes . The gynaecium is glabrous and
measures + 4 mm., is ellipsoid, and almost as long as the slender style,
which is topped by a conspicuous subpeltate stigma.
As has been noted under P. Brenesii, the nearest connection seems to be
with that species.
5. Persea americana Miller, Gard. Dict. ed. 8. 1768; 2 in Jour. Wash. Acad. Sci.
10:12, fig. 1, A. 1920; Standley in Contr. U.S. Nat. Herb. 23: 290. 1922; rane
& Calderon, Lista Prelims. Pl. Salvador 85. 7935: geht in Contr. U. S. Nat.
Herb. 27: 183. 1928, in Trop. Woods 21: 17. 1950, in Field Mus. Publ. Bot. 10:
201. 1931; Skutch in Torreya 32: 85-94, figs. 1, 2. 1932; Standley & Record in
Field Mus. Publ. Bot. 12: 144. 1936; Standley in an Mus. Publ. Bot. 18: 457.
Laurus persea Linnaeus, Sp. Pl. 370. 1753.
Persea gratissima Gaertner f., Fruct. & Sem. 3: 222, pl. 221. 1805.
Persea edulis Rafinesque, Sylva Tellur. 134. 183
Persea gratissima var. @ vulgaris Meissner in DC. Prodr. 15!: 53. 1864.
Persea gratissima var. Y macrophylla nian it LG;
Piola persea rome in Bull. Torrey Bot. Club 19: 95.
Persea amplifol oo ? sensu Record in Trop. =A ry wn 21. 1927; Standley in
Trop. W roods ‘21: 217. 1930; non Standley & Calderon, 19
DISTRIBUTION: ean probably of Mexico and Central America, and possibly the
West Indies; now w planted profusely throughout these regions and all tropical and sub-
tropical countri
NATIVE NAMEs: ‘Aguacate”’ geneeel term throughout Central America, of Aztec
origin); ‘Ahuacate,”’ ‘‘Avocado,” “Aguac ate oloroso” bite Cruz, Oaxaca); “On”
(Yucatan Maya) ; “Aguacate xinene,”’ ‘\Xinene”’ (Oax “Tonalahuate” (Morelos,
Vera Cruz) . (Michoacan, Jalisco). Many noe names are reported by
Standley (1928 ) as being common in South America where the species is cultivated.
He also gives numerous Indian names (1937).
This widely cultivated tree of the tropics is outstanding for its ovate-
oblong or obovate-oblong coriaceous dark green leaves, 10-20 cm. long and
3-10 cm. broad, crowded at the branchlet-tips, slightly shining above and
1945 | ALLEN, STUDIES IN THE LAURACEAE, VI 289
dull glaucous beneath, pubescent to glabrous throughout with yellow vena-
tion prominent beneath, subtended by long (1.5—5 cm.) petioles, yellow
or yellow-green, pubescent, becoming glabrous. The axillary subterminal
inflorescences are paniculate, many-flowered, densely and shortly ae
bracteate, the pubescent bracts shortly tomentose and fugacious.
small 0.5— 1.5 cm. diam.) flower is borne on a yellowish green Seer
densely yer shortly tomentose and up to 0.5 cm. long. The lobes of the
and without, rarely glabrescent, persistent, 3-6 mm. long and 2—3 mm.
wide. The outer lobes are shorter and more narrow than the ovate-oblong
or lanceolate inner lobes. The stamens of the two outer series have ovoid
or ovoid-oblong acuminate or narrowed and obtuse anthers that are
introrse; the flat pilose filaments are 2.5-3.5 mm. long. Those of the
inner series are longer, with extrorse anthers, and the filaments bear at the
base two short-stiped ovoid glands. The staminodia are short-stiped.
The pubescent gynaecium, 4—5.5 mm. long, consists of an ovoid ovary
bearing a filiform style topped by a discoid stigma. The large usually
oblique globose or pear- -shaped edible fruits are yellowish green or ae
with purple when ripe, shining, glabrous, 7—20 cm. long and 7-10 cm. in
diameter. The rind is shining and coriaceous; the flesh is thick, a
pulpy, sweetish to taste, about 1.5—2 cm. thic
5a. Persea americana var. drymifolia apis tee & Chamisso) Blake in Jour.
Wash. Acad. Sci. 10: 15. 1920; Standley in Contr. U. . Nat. Herb. 23: 291. 1922;
Standley & Calderon, Lista Prelim. PI. Fores $8. 19
Persea isi runes & Chamisso in pais ~ 365. 1831; Meissner in
dr. 151: 53. 1864; Trabut in Rev. Hort. 1908: 296, fig. 110.
DISTRIBUTION: ae known only from the type-locality.
Mexico: Vera Cruz: Papantla, March, Schiede & Deppe 1140 (fl., isotyPE of
. drymifolia, Mo).
NATIVE NAMES: ‘“Aguacate oloroso” (Mexico).
This variety of P. americana from Mexico has smaller leaves and fruits,
the leaves giving forth the odor of anise when crushed. The fruit is sup-
ported by persistent or somewhat persistent perianth-lobes, a character
which Blake mentions as differing from that of P. americana. The true
avocado, on the other hand, according to Mez’ description, has “lobes per-
sistent or subdeciduous.”’
In the case of P. americana and allied species and varieties, further work
in the field is necessary for a comprehension of the variability that most
certainly abounds in the group. This seems to be particularly true where
cultivation has occurred.
The branchlets are blackish, angular-striate, and becoming glabrescent.
The leaves are alternate or subopposite, with long slender glabrous petioles
1-2 (-3) cm. long and 1 mm. in diameter, slightly canaliculate. The
blades are elliptic, acute or acuminate, with cuneate or sometimes somewhat
obtuse base; they measure 6-12 (—15) cm. long and 2—4.5 (—5.7) cm. broad,
are pergamentaceous, in the dried state above greenish brown and glabrous,
beneath glaucous and scattered-pilose. The costa above is impressed and
glabrous, beneath elevated and brownish with slight pilosity. The lateral
nerves vary from 6 to 9, not always in opposite pairs, and are obscure above
and slightly evaied 4a brownish-pilose beneath. They diverge from
~
290 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
the costa at an angle of about 45°. The reticulation is indistinct and loose
throughout. The subterminal inflorescence is lightly clothed with a
fulvous pubescence and is shorter than the leaves, measuring up to 9 cm.
in length, with a peduncle 2—4 cm. long. The pubescent flower is borne
on a slender pedicel up to 6 mm. long, and the subequal everywhere
pubescent elliptic lobes are membranaceous. ‘The first and second series
of stamens are pubescent and about 3.5 mm. long, the ovate obtuse
anthers with introrse cells being — longer than the slender filaments.
The two lower cells are about twice the size of the upper. This is true
also of those of the anthers of the third series, which are extrorse, whereas
the two upper cells are lateral. The filaments of the third series are once
and a half again as long as the anthers and bear at the base two lateral
stipitate glands which appear to be cordate and are equal in length to the
supporting stipes. The fourth series or staminodia are + 1.25 mm. long,
d borne on thick rice stipes half the entire length. The
gynaecium measures approximately 3.5 mm. in length and is pubescent
throughout. The ovoid to subglobose ovary is shortly stipitate and bears
a slender style which is twice its length. The stigma is obliquely peltate
and conspicuous.
6. Persea Schiedeana Nees, Syst. Laurin. 130. Bante Blake in Jour. Wash. Acad. Sci.
: 16, fig. 1, B. 1920; Standley in Contr. U.S. Nat . Herb. 23: 289. 1922; Standley
& Calder, Tinta Prelim. Pl. Salvador 85. 1925; Standley in Trop. Woods 21: 17
1930; orp! in Field Mus. Publ. Bot. 10: 201. 1931; Standley & Record in op.
cit. 12: 1936; Standley in op. cit. 18: 458. 1937.
Persea sp. a | & Chamisso in Linnaea 6: 365.
Persea gratissima var. 5 Schiedeana Meissner in DC. Prodr. 151: 53. 1864.
Persea Pittieri Mez in Bot. Jahrb. 30: Beibl. 67: 15. 1901.
DistriBUTION: Forests of Mexico and Central America.
Mexico: Cruz: Forests of Misantla, Schiede (fl., type of P. gratissima
var. 5 Schiedeana not seen); Barranca de Fortin near Cordoba, Woronow 3127 (fl
h). Mexico: Acatitlan, Hinton 3167 (fl, GH). Michoacan: Llano, Coal-
coman, Hinton 12987 (fl. GH). Oaxaca: Ubero, L. Williams 9151 (sterile, A).
Chiapas: Between Copainaba and Coapilla, Woronow & Juzepczuk 1666 (fl., Ch).
GuATEMALA: Huehu ngo: Northwest of Malacatancito, at km. 8 of the
highway from Hushuctenanso, in xe se in oak-forest, Standley 82186 (f1.,
Ch). Alta Verapaz: ar San n Chamelco, in pine-forest, Standley 92204 ({fr.,
Ch); near Coban, in wit setters Pm pee (fr, Ch). Izabal: Los Amates,
Kellerman 7145 (fl, Ch). Honpuras: Tegucigalpa: Mont. de la Flor, near the
river, C. & J von Hagen 1232 (sterile, NY). Yoro: El na Geanile, C. &
V.W. von Hagen 1005 (sterile, NY). British Honpuras: Stann Creek Middle-
sex, Stevenson V (¥8932) (fl, Y). Nicaragua: N. E. ‘jaue nh ' Baaleting
162 ie Ds Costa Rica: Guanacaste: About houses in Nicoya, Tonduz 13794
Chiriqui: Vicinity of Cerro Punta, P. H. ‘Allen 1534 (sterile, Mo); Bajo Chorro,
Boquete, Davidson 304 (fr., Ch), 427 (f1., A, Ch, Mo). Panama: Isla Taboga,
P.H. Allen 1288 (fl., Mo).
NATIVE NAMES: ‘“Aguacate” (Mexico, British Honduras, Costa Rica); “Aguacaton”
he’
(Panama); “Chalté,” ‘“Chaucte’ (Guatemala); ‘Chinini’ (Mexico) “Chuche”
(Guatemala, San Salvador); “Chuti” (Honduras); “Coyo,’ “Coyoc +” “Coyoté”
(Guatemala); “Guaco” (Honduras); “Kiyau,” “Kiyo,” “Kotyo,” ‘“Shucte” (Guate-
mala); ‘Wild Pear” (British Honduras); “Y aa (Costa Rica).
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 291
This species, once considered a variety of P. americana, seems to warrant
specific rank. The closest affinity is no doubt the common avocado, but
the differences are significant. Popenoe (U.S. Dept. Agric. Bull. 743: 37.
1919) makes the comment that, although the fruits of the two entities
resemble each other, the tree is distinct in foliage and floral characters. I
agree also with Popenoe that the avocado group needs more intensive study
in the field.
The branchlets of P. Schiedeana, as well as the lower surfaces of the
young leaf-blades, are brown-tomentose. The leaf-blades are usually half
o 3 cm. long. The blades are irregularly elliptic and often obovate-
elliptic, shining above and loosely reticulate, glaucescent and finely tomen-
tose beneath. he base is frequently almost truncate and is usually
rounded, the apex rounded, acute, or abruptly acuminate. The lateral
nerves, of which there are 8-15 pairs, ascend at a variety of angles from the
costa, 75-80° at the base and 45 or even 35° in the upper portion of the
blades. The usually heavily pubescent subterminal numerous paniculate
inflorescences bear persistent bracts and are shorter than the leaves,
pedicel almost equaling it in length. The narrowly ovate or lanceolate-
ovate perianth-lobes are subequal and very pubescent without. The outer
lobes measure 4—5.9 mm., the inner 4.5—-6.4 mm. The stamens are +
mm. long and bear ovate obtuse anthers with the two upper cells smaller,
sometimes about half the size of the two lower cells. The slender filaments
are pubescent and twice as long as the anthers. The filaments of the inner
series bear glands that are cordate-stipitate (the stipes equal to the glands),
and are about one-third the entire length of the stamens. The fourth
series consists of cordate staminodia borne on hairy stipes more than half
the length of the staminodia. The pubescent gynaecium is + 3.8 mm.
long, the slender style more than twice the length of the ovoid ovary. The
fruits, of which I have not seen mature specimens, seem to be ‘obovoid
(slender and bottle-necked), and in the early stages very pubescent, later
by the thickened and enlarged pedicels, also becoming glabrescent.
7. Persea floccosa a in Jahrb. Bot. Gart. Berlin 5: oH 1889; wore in ca Wash.
Acad. Sci. 10: 15. 1920; Standley in Contr. U. S. . Herb. 23: 922
DISTRIBUTION: ae in forests.
Mexico: oe Near Chinantla, Liebmann 85 (f1., sua of Type, Ch, NY,
US). Chiapa Ventana, near Siltepec, eee 4545 (fr., A, NY); Rodeo, in
virgin forest near Giltenes. Matuda 4559 (fl.,
NATIVE NAME: “Aguacate cimarr6én’ oa.
This species is characterized by the fulvous pubescence of the voung
branchlets, which soon become glabrescent and blackish red, with very
prominent lenticels, and the dense ferruginous-lanose (according to the
author) pubescence of the young leaves. Adult leaves are densely and
manifestly foveolate-punctate and subglabrous above, while beneath they
are glaucescent- and floccose-tomentose. They vary from 11-17 cm. long
and 4.8-7.5 cm. broad, and are usually elliptic or occasionally narrowly
292 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
elliptic, the base obtuse or cuneate, the apex somewhat acuminate, with
7 or 8 pairs of lateral nerves diverging from the costa at an angle of. 35",
the lower pairs of which are often irregularly arcuate and almost at right
angles. The erect stout pyramidal inflorescence is up to 11 cm. long, the
peduncle nearly half its length, and covered with a dense subferruginous
pubescence. The flowers are densely pubescent, up to 5 mm. long, the
pedicel 1-3 mm. The perianth-lobes are elliptic, rather acute, densely
pubescent, the inner up to 3.4 mm. long, the outer slightly less. The sta-
mens are about 2 mm. long, the slender pubescent filaments are approxi-
mately one-third longer than the ovate obtuse anthers, the two upper cells
of which are smaller than the two lower ones. ‘The stamens of the inner
6 mm. long and 5 mm, in diameter at the apex.
Persea floccosa has an affinity with P. americana and P. Schiedeana, but
the smaller depressed-globose fruits are very different. The leaves are on
the whole smaller than those of the two latter species, as are the flowers.
The inflorescences are longer, accordingly, being nearly the length of the
leaves. A study of more complete material may very well show this to be
a form of P. americana.
8. Persea flavifolia Lundell in Contr. Univ. Mich. Herb. 6: 17. 1941.
DistrisuTIoN: Known only from the type-locality and vicinity.
Mexico: Chiapas: Mt. Ovando, April 9-12, 1937, Matuda 1821 (f1., A, Ch,
TYPE — Mich), 2651 (fl., A, Ch, NY).
This tree is remarkable for the stout branchlets, which are conspicuously
angled, becoming striate-sulcate, early covered with a golden-sericeous
tomentum, which becomes fuscous and eventually glabrescent. The older
branchlets are conspicuously cicatricose. The petioles are golden-seri-
ceous, striate, and up to 4.5 cm. long. The leaf-blades are coriaceous,
elliptic-oblong or lanceolate-oblong, the base rounded or obtuse, the apex
acute or obtusely subacuminate. They measure up to 21 cm. long an
8.5 cm. broad, early sparsely pubescent, soon glabrous, the surface yellow-
green, the lower surface conspicuously golden-brown-sericeous. The costa
is slightly impressed above and elevated beneath, everywhere golden-
brown-sericeous. The lateral nerves, of which there are 7—12 pairs, are
inconspicuous above but slightly elevated beneath. The reticulation is
obscure throughout. The inflorescence is axillary and subterminal, con-
sisting of numerous short compact many-flowered densely golden-brown-
sericeous panicles not more than 4-6 (—7) cm. long. The flowers are 5-6
(-10) mm. long, including the short stout pubescent pedicel. The outer
lobes are ovate or orbicular, + 2.2 mm. long, thickly hairy within and
densely sericeous without, those of the inner cycle being lanceolate-elliptic
and up to 3.5 mm. long. The stamens of the two outer series are =
2.2 mm. long. the ovate roundish anthers being shorter than the pubescent
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 293
filaments. Those of the inner series are + 3 mm. long, the filaments bear-
ing conspicuous reniform stipitate glands. The upper cells of the anthers
staminodia are + 1.25 mm. long, cordate, ovate, the pubescent stipes being
more than one- sae the entire length. The glabrous to pubescent gynae-
Cin is. = °372 long, the subellipsoid glabrous or pubescent ovary
shorter than hes ead style topped by a large conspicuous capitate
stigma. The hues of which I have seen no specimen, is globose. black,
1 cm. in diamet r, the perianth-lobes being persistent.
Lundell es this striking species to P. Benthamiana from Brazil. It
appears also to have a close relationship to P. podadenia, also from Mexico,
in spite of the occasional presence of pubescence on the ovary. e per-
sistent sericeous pubescence of the leaf-blades and their shape separate it
from the latter at once. Mexia 1632, a young fruiting specimen from
Jalisco, may possibly belong here. The fruit seems very small and the
infructescence still heavily pubescent. The leaves are on the whole small
for this species, but the infructescence is shorter than that of P. podadenia,
the only other possibility, and the lower part of the branchlet is heavily
cicatricose — a characteristic of P. flavifolia.
9. Persea es Smithii “ect in Donnell ee egies Pl. Guatem. 2:67. 1891,
nom. nud.; in Arb. Kgl . Gart. Breslau 1: . 1892
ie tes Se a — in sealant acti at 1220-2000 meters
altitude
Mexico: Vera Cruz: Near Mirador, Liebmann 11 (syNtypE, not seen).
Chiapas: Coapilla, Juzepczuk 1699 (fr., Ch); Pinada, Siltepec, Matuda 1942 (Ur.,
near Rancho Fenix, Hacienda Monserrate, Purpus 10526, 13080, 14302 (f1., Ch).
GuaTeMALA: Alta Verapaz: Chicoyonito, alt. 1310 m., April, 1889, J. D. Smith
1718 (fl., IsosyNTYPE, Mo) ; between Coban and San Cristébal, Cook & Doyle 89 (fl.
Ch); Coban, pila 646 (fr., Ch); near Coban, Pears 69166, 69315 (fl., Ch);
large swamp east of Tactic, Standley 92394, 92520 (fl, Ch). Baja Verapaz:
Region of Patal, oe 69565 (Af1., ;
NATIVE NAMES: ‘‘Aguacate,” ‘“Sacsi’’ (Guatemala).
The nearest relative of this species seems to be P. Chamissonis, from
Mexico. A short discussion of the points of difference is given under that
species. Mez places P. Donnell-Smithii near P. floccosa, which to me
seems, on account of its fruit characters, to be allied with the P. americana
complex.
This species is striking because of the large (up to 18 cm. long and 12
[-15]| cm. broad) aerrapag: elliptic or obovate leaves. the lower surface
of which is covered with a dense ferruginous tomentum, as are the stout
petioles which measure an to 4 cm. in length and are frequently as thick
mm. in diameter. The blades are often unequal at the base, cuneate
or rounded, the apex usually rounded. The inflorescence, up to 10 cm.
long though often only 4 cm. long, is narrowly subpyramidal-paniculate
and is covered with a dense ferruginous tomentum, as is the infructescence.
The flower has thick fleshy lobes that are hairy inside and densely tomen-
tose without and are definitely spreading. The outer lobes are orbicular
and up to 3 mm. long, the inner ovate-elliptic and nearly twice the length
294 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
of the outer. The two outer series of stamens are + 3.8 mm. long, the
pubescent filaments being longer than the broadly ovate anthers. The
stamens of the inner series are + 4.25 mm. long, the anthers narrowly
oblong, the two upper cells lateral, the lower laterally extrorse. The fila-
are + 1.25 mm. long, obscurely sagittate, the stipes more than one-half
the entire length, the whole densely pubescent. The seemingly glabrous
gynaecium, 4-5 mm. long, consists of a globose ovary surmounted by a
slender style twice its length and bearing a conspicuous subtriangular stigma,
The fruits are shining, globose, strongly apiculate, about 12 mm. diam.,
and borne on the persistent perianth-lobes which are somewhat enlarged,
still heavily tomentose, and spreading to a diameter of 12 mm. The short
tomentose pedicel is enlarged also to 3 mm. long and 2 mm. in diameter.
10. Persea Chamissonis Mez in Jahrb. Bot. Gart. Berlin 5: 168. 1889; Standley in
Contr, U. S. Nat. Herb. 23: 292. 1922; Lundell in Contr. Univ. Mich. Herb. 7: 12.
1942.
DistripuTion: In the forests of Mexico.
Mexico: Vera Cruz: Chiconquiaco, Schiede (fr., fragm. of syntypr, Ch).
Puebla: Near Chinautla, Liebmann 1 (photo. of syntype, Ch, NY); Chinautla, in
May 1841, alt. 2135 m., Liebmann s.n. (fl., fr., Ch, Mo).
The present species is distinguished by the ferruginous-tomentose
branchlets that eventually become glabrate and blackish brown. The leaf-
blades, with long petioles up to 3 cm. long (according to the collector),
slightly canaliculate, and cinereous-tomentose, are alternate, usually
obovate or elliptic, 9-13 cm. long, 3-7 cm. broad, chartaceous above an
densely pubescent, becoming glabrescent, scattered- and spreading-pilose
with longish hairs except for the costa, beneath somewhat glaucous and
more crisped-pilose. The blade is densely and conspicuously reticulate on
both surfaces and there are 6-8 pairs of lateral nerves diverging at an angle
of 45°, The fruits are black, globose, apiculate, about 10-11 mm, in
diameter, seated on the enlarged and rather horizontally spreading still
densely ferruginous-tomentose lobes of the perianth, which form with the
expanded tomentose tube about 1 mm. high a cupule about 12 cm. in
diameter, and are subtended by the enlarged tomentose pedicel about
3 mm. long and 1.5 mm. broad.
‘ery close to P. Chamissonis is P. Donnell-Smithii, which differs in hav-
ing its consistently larger leaf-blades (up to 17 cm. long and 12 cm. wide)
heavily coriaceous and densely tomentose. They do not show prominent
reticulations and are borne on a stout petiole up to 4 cm. long and 4—5 mm.
thick, covered with a dense ferruginous and presently cinereous tomentum.
The lateral veins of P. Chamissonis tend to ascend less arcuately than those
of P. Donnell-Smithii. 1 have seen no inflorescence of P. Chamissonis, but
presumably it is short. The inflorescence of P. Donnell-Smithii is longer
than the infructescence of the former.
11. Persea cinerascens Blake in Jour. Wash. Acad. Sci. 10: 18, fig. 2. 1920; Standley
in Contr. U. S. Nat. Herb. 23: 289. 1922.
DistripuTIon: Mexico, known only from the type.
Mexico: Vera Cruz: Zacuapan, June, 1916, Purpus 7671 (fl., TyPE not seen),
8144 (fr., not seen).
Blake has related this species to P. Liebmanni Mez, noting that it differs
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 295
in its larger flowers, longer anthers, staminodial glands glabrous on the
inner face, and larger fruits.
heavily so on the nerves, and prominently reticulate. The lateral nerves
are 14-16 pairs and diverge from the costa at an angle of 45-65°. The
petioles measure up to 3 cm. long. The axillary inflorescence is sordidly
pilose-tomentose, 5-7 cm. long, paniculate, narrowly subpyramidal. The
flowers are borne on stout pedicels up to 1 mm. long; the perianth is 7—8.5
mm. long, densely pilosulous-tomentose, with elliptic-oblong obtuse lobes,
the outer somewhat shorter than the inner. The stamens of the two outer
series are 4.5—-4.8 mm. long, those of the inner series 5.2 mm. long, with
biglandular filaments. - The pubescent filaments are almost two-thirds the
length of the stamens. The staminodia are 3 mm. long, the cordate apicu-
late apical gland, glabrous on the inner surface and pilose dorsally, present-
ing an unusual feature. The ovary is + 1.5 mm. long, ellipsoid, glabrous,
with a slender glabrous style more than twice its length. The fruits are
subglobose, glaucous-blue, about 12 mm. diam., subtended by the per-
sistent perianth-lobes which are slightly thickened to form a cupule | mm.
high and 4.5 mm. in diameter.
12. Persea Liebmanni Mez in Jahrb. Bot. Gart. Berlin 5: 1066. 1889, Standley in
Contr. U. S. Nat. Herb. 23: 292. 1922.
DisTRIBUTION: Known only from the type.
aca: Near Chinantla and Trapiche de la Concepcion, Liebmann
).
EXIC
115 (SYNTYPE not seen), 116 (f1., fr., photo. of SYNTYPE, Ch, NY
glabrous above and appressed beneath, are ovate
or elliptic or elliptic-lanceolate, cuneate and often oblique at the base, and
acuminate or subacute at the apex, measuring 14-26 3.8-10.5 cm., and
borne on a petiole up to 4.5 cm. long, plane or slightly canaliculate. The
8-12 pairs of lateral nerves diverge from the costa at an angle of 35—45°.
Above there is noticeable a prominent areolation, which beneath shows up
as a loosely prominulous reticulation. The sericeous-tomentellous inflo-
rescence is a many-flowered pyramidal panicle shorter than the leaves,
about 15 cm. long, the peduncle up to 8 cm. long. The flowers, also sub-
sericeous, are up to 5 mm. long, with the outer lobes one and one-half times
shorter than the inner. The pilose filaments almost equal the subrectangu-
Jar-oval anthers or are slightly longer. The staminodia are conspicuous,
liguliform-sagittate, and densely pilose, the stipes being about one-half the
entire length. The glabrous ovary is globose, twice as long as the. style,
which is topped with a discoid stigma. The fruit is globose, glaucescent-
blackish, up to 7 mm. in diameter, seated on the spreading persistent and
slightly enlarged perianth-lobes.
According to Blake, the only species known to which this has an affinity
is P. cinerascens, of which unfortunately no material is available.
296 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXVI
13. Persea vesticula Standley & Steyermark in Field Mus. Publ. Bot. 23: 116. 1944,
DistripuTion: Volcanic mountain-slopes of Guatemala.
Huehuetenango: Cerro Huitz, between Mimanhuitz and Yul-
huitz, Sierra ie los Cuchumatanes, Steyermark 48571 (fr., Ch). San Marcos: To
of escarpment, between La Vega ridge along Rio Vega = northeast slopes of Volcan
de Tacana, near the Mexican boundary, alt. 2500-3000 m., Feb. 20, 1940, Stevermark
36207 (fl., TypE, Ch). El Progreso: In cloud-forest, Siva de las Minas, hills north
of Finca Piamonte, Steyermark 43558 (sterile, Ch).
NATIVE NAME: “Canok’”’ (Huehuetenango).
The species so recently fisoriean by Standley and Stevermark has not
turned up in any other collectio It is distinctive at once for its sturdy
pubescent densely foliose branchlets bearing rigidly coriaceous leaf-blades
glabrous to glabrescent above and minutely pubescent beneath, borne on
stout petioles 1-2 cm. long, the pio slightly recurved, particularly at
the base, oblong- elliptic, 10-17 & 3.5-6.5 cm. The lateral nerves are 9
or 10 (— 12) pairs and faintly visible above and definitely elevated beneath,
diverging at an angle of approximately 45°. he subterminal profusion
of robust inflorescences, almost forming a single corymbose structure at the
tips of the branchlets, is many-flowered and ferruginous-tomentose. The
flowers are 12-14 mm. in diameter, the outer perianth-lobes subtriangular,
thickly pubescent. + 3.4 mm. long, the inner + 7.25 mm. long, elliptic.
The stamens are + 4.7 1 long, the ovate obtusely apiculate anthers
almost as long as the sede The anthers of the third series are more
one-fourth the length of the entire stamens. The staminodia are irregu-
larly petaloid, sagittate, stipitate, dorsally pubescent, measuring + 2.2 mm.
in length, the fides ge about equal to one-half the length of the staminodia.
ynaecium is + 5 mm. long. The slender Ay is twice as long as the
glabrous ellipsoid to subglobose ovary and is topped by a capitate conspicu-
ous stigma. The subglobose fruit measures 3.5 cm. in diameter and is
borne in a thick subverrucose cupule 12 cm. in diameter, 5 mm. long, and
+ 2mm. deep. The ens pedicel is about 5 mm. long and 8 mm.
in diameter at the a
Standley and Siva ik relates this species to P. longipes, but in my
opinion it is more closely allied to the P. americana complex, the inflores-
cence and large fruit indicating this. The thickened and ligneous fruit-
cupule, however, is certainly not typical of the Persea species of this region.
14. Ha pallida Mez & Pittier ex re in Bull. Herb. Boiss. II. 3: 231. 1903; Standley
ield Mus. Publ. Bot. 18: 1937
ae TION: Costa Rica, —. as — the type.
Costa Rica: Puntarenas: Valle de Coto, alt. 1400 m., Pittier (Herb. Inst.
Costa Rica 11111) (fl., Type not seen) (flowers in February).
Inly a flowering specimen of this species was described. The stout
branchlets, early pruinose with short yellowish pubescence, later become
glabrous. The chartaceous sparse csomnaoeih leaf-blades, 15 & 6 cm.,
are densely prominulous-reticulate, pale almost glaucescent beneath,
and are subtended by slightly canaliculate eaeririe up to 2cm. long. The
corymbose inflorescence is entirely covered with a remarkable white indu-
ment. The flowers are about 4 mm. long and borne on very short and
thick pedicels 2-3 mm. long. The lobes are elliptic, thick and fleshy,
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 297
broadly acute; the filaments of the stamens are shorter than the broad well
rounded and emarginate or obtuse anthers. The anthers of the stamens
of the third series are laterally dehiscent and are large, fleshy, and sessile.
The slender style is three times as aa as the subglobose glabrous ovary
and bears a large often triangular sti
The nearest affinity of this species is ie following new species, also from
Costa Rica and found in adjacent Panama as well. Mez remarks that
superficially one is reminded of the habit of Phoebe glaucescens from India,
but that a careful study reveals relationship to the Andean species Persea
ferruginea and P. vestita, neither of which latter are available at present.
15. Persea rigens, sp. nov.
O m. alta ramulis robustis, brunneis vel maculato-brunnescen-
tibus, sulcatis, glabris. Folia alternata vel subverticillata, petiolis brunneis
vel fuscis, crassis haud canaliculatis, glabris, ad 1.5 cm. longis et 3 mm.
latis, laminis utrinque glabris, juventute membranaceis, mox rigide cori-
aceis, in sicco olivaceo-viridescentibus, ellipticis, 20-25 (33) cm. longis et
ad 12 cm. latis, basi cuneatis, apice obtusis vel haud obtuse subacuminatis,
penninerviis, nervis 7- vel 8 8(—9)-paribus supra leviter subtus aaa
elevatis, angulo 45—55° divergentibus, costa supra subplana subtus co
spicue elevata, utrinque satis minute reticulatis. Inflorescentia paniculls
numerosis composita, paniculis ad 5 ex axilla quave folii decidui summi,
dense griseo- vel fulvo-albescentibus, ad 10 cm. longis et 2 cm. latis, pedun-
culis ad 6 cm. longis. Flores ad 4 mm. longi, campanulati, dense tomen-
tosi, pedicellis ad 4 mm. longis pubescentibus, lobis subaequalibus crassis
tomentosis ad 3 mm. longis; staminibus ser. T& I plus minusve variabi-
libus saepe subpetaloideis, + 1.25-2.2 mm. longis filamentis pubescen-
it Z
tibus, ser. III -: 2.2 mm. longis biglandulosis, glandulis stipitatis;
staminodiis stipitatis subcordatis, -+ mm. longis; gynaecio ovario
ovoideo glabro + 2.8 mm. longo, quam stylo 2% longiore. Infructescentia
robusta eee vel slaved ad 15 cm. longa, rubescens. Fructus igno-
tus. Pedicellus incrassatus, rugosus, ad 7 mm. longus et 5 mm. latus, apice
loborum reliquis, ad 7 cm. diam. insignitus.
DistRIBUTION: Costa Rica and adjacent Panama.
Costa Rica: Without locality, in 1943, Little 6075 (fl, TYPE, Ch). PANAMA:
cas del Toro: Daytonia Farm, region of Almirante, Feb. 2, 1928, G. P. Cooper
458 (¥12076) (fr., Ch, Y) (tree 18 m. high, 35 cm. diam., said to reach 30 m. and 95
cm. diam., with stout horizontal branches at top; et for boards and rough lumber).
NATIVE NAME: ‘Timber Sweetwood” (Panama).
The dense many-panicled inflorescences of this species present a striking
appearance, clothed with a heavy grayish or fulvous-white tomentum. As
many as four panicles up to 10 cm. in length arise from a single axil of
leaves that are early deciduous. Unfortunately, Cooper’s number has no
fruit attached, but the enlarged peduncles and pedicels of the infructescence
bearing the remains of the perianth-lobes leave no doubt that the specimen
is conspecific with the Costa Rican tree.
The species is near P. pallida, but differs in not having corymbose inflo-
rescences and in its large leaf-blades, 33 12 cm., only slightly paler be-
neath and not glaucescent, with petioles not more than 1.5 cm. long. ‘The
perianth-lobes are not elliptic but broadly ovate, and the pedicel is equal
298 JOURNAL OF THE ARNOLD ARBORETUM (VOL. XXVI
to if not longer than the flowers. The ovary is ovoid instead of subglobose
and is two-thirds the length of the entire gynaecium, instead of one-fourth
as in P. pallida. The type of stamen is not typical for Persea, but the
species seems to belong in the genus
16. Persea Skutchii, sp. nov.
Arbor ad 21 m. alta, ramulis rubescentibus striatis, juventute Sar et
adpresse ferrugineo-tomentosis, mox glabris. Folia alternata vel subver-
ticillata petiolis gracilibus leviter capalicilatie brunneis a 3,5 cm, longs,
laminis supra glabris subtus sparse pubescentibus, pergamentaceis, in sicco
supra pallide viridescentibus subtus pallide brunnescentibus, ellipticis vel
late ellipticis, 10-17 c m. longis et 3 (—7.5) cm. latis, basi rotundatis vel
cuneatis, interdum obliquis, apice obtusis vel acutis vel acuminatis, saepe
utrinque plus minusve planis, angulo 67—56° divergentibus, utrinque rete
venularum minuto et obscure prominulo. Inflorescentia axillaris paulo laxe
paniculata, minute sericeo-fulvo-pubescens, ad 12 cm. longa. Flores ad 6
mm. longi, campanulati, dilute RG pedicellis ad 5 mm. longis
gracilibus pubescentibus, lobis exterioribus late ovatis membranaceis pu-
bescentibus + 1.5 mm., interioribus lanceolatis pubescentibus + 5 mm.
longis; staminibus ser. I & II + 3.4 mm. longis antheris oblongis ‘ay
filamentis pubescentibus plusquam triplo longioribus, ser. III ad + 3.5 mn
longis biglandulosis glandulis stipitatis; paises crassis utrinque ani
yescentibus subcordatis, stipitatis, + 2.7 mm. longis; gynaecio glabro +
3.4 mm. longo, ovario ovoideo basi leviter pee longitudine quam stylo
breviore, stigmate conspicue triangulari-peltato. Fructus nigrescens lu-
cidus, globosus, inconspicue apiculatus, ad 7 mm. diametro, glaber, pedicello
aliquid incrassato apice ad 2 mm. diametro sparse pubescente apice plus
minusve 6 mm. diametro, loborum reliquis ornato
Disrrinution: Costa Rica and Panama, 300 to 1675 meters rai
Costa Rica: Puntarenas: Buenos Aires, nae 6680 (f1., Ch). Alajuela:
Edge of wooded ravine in : Pacific tropical zone, Atenas, A. Smith yee (f1., A). San
osé: In clearings, vicinity of El General, Skutch Se (fl., A, NY); basin of El
General in clearing, alt. 675-900 m., March 1940, Skutch 4812 (f1., rvpe, A) (tree 15 m
flowers yellowish) ; Cerro de Pratli, Escazu, Solis 183 (fl., Ch); Santa Maria de Dota,
Stork 2416 (fr.,Ch). Panama: Coclé: Vicinity of El Valle, south rim (dry), P. H.
Allen 1781 (fr., Mo, NY Vs hills south of El Valle de Anton, P. H. Allen 2498 (fr., A)
NaTIVE NAME: “Aguacatillo’”’ (Costa Rica).
This species is akin to P. caerulea, from South America, but may be sep-
arated from it by the inflorescences being shorter than the leaves and by the
less erect lateral nerves. The leaves are usually acute or subacuminate
rather than rounded-obtuse, as those of P. caerulea. The blackish (in the
dried state) fruits are depressed-globose, lack entirely the bloom of those
of the latter species, and do not measure more than 8 mm. in diameter at
the most.
17. Persea Hintonii, sp. nov.
Arbor magna |?]|, ad 5 m. alta, ramulis flavo- vel fulvo-tomentosis mox
sparse griseo-tomentosis demum glabrescentibus fusco-brunneis, subsul-
catis. Folia alternata, petiolis flavo-tomentosis ad 3.5 cm. longis canalicu-
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 299
latis, laminis supra glabris costa pubescente excepta subtus utrinque sparse
cano- vel flavo-tomentosis, coriaceis, supra in sicco pallide alent
subtus glaucis, oblongo- -ellipticis, 15-20 cm. longis et 5—6.5 (—8) cm. latis,
basi rotundatis vel cuneatis et obliquis, apice rotundatis, obtusis vi acutis
et minute apiculatis, penninerviis, costa plus minusve conspicua leviter im-
ub
venularum supra obscuro subtus satis conspicuo. Inflorescentia axillaris,
compacte paniculata vel subcapitata, 3-5.5 (-10) cm. longa, dense fulvo-
vel flavo-tomentosa, pedunculo ad 5.5 cm. longo. Flores ad 7.5 mm. longi,
confertissimi, perianthio ee lobis exterioribus crassis ovatis extus
pubescenti ibus + 2.7 mm. longis, interioribus lanceolatis ie utrinque
intus apice pseu: + 4.7 mm. longis; staminibus ser. I & II + 2.15
mm. longis, antheris ovatis saepe ae breviterque acuminatis variabilibus
saepe quam filamentis pubescentibus plusquam duplo longioribus, ser. II#
ay ‘ : :
rioribus parvissimis, ser. I & II introrsis, ser. III laterali-extrorsis; stami-
nodiis anguste ovatis graciliter stipitatis + 1.7 mm. longis pubescentibus;
career glabro + 3.8 mm. longo, ovario subgloboso leviter stipitato, ao
tvlo 13 breviore, stigmate conspicuo peltato. Fructus immaturus(?)
aa violascens ellipticus, apiculatus, 5-8 mm. diam., glaber, cupula lobo-
rum tomentosorum incrassatorum subtentus, pedicello nee breviter
flavo-pubescente ad 2 mm. longo et lato.
DistRIBUTION: Known only from Mexico.
Mexico: Jalisco: Stream-side, San Sebastian, trail from Hacienda del Ototal
to Hacienda La Indrilla, Mexia 1699 (fl, GH). Puebla: Near Honey Station
(Trinidad), Pringle 8938 (fl, GH). Mexico: Temascaltepec: Temascaltepec, on
hill, alt. 1700 m., March 27, 1933, Hinton 3533 (fl., Type, GH) (large tree), Hinton 4024
(fr., GH); Llano, Tejupilco, Hinton 3980 (fr. GH). Guerrero: San Antonio-
Buenos Aires, Montes de Oca, hill, Hinton 14027 (fl., GH); by stream in oak-forest,
Chilacayote-Soledad, Mina, Hinton 14195 (fl. Ne
NATIVE NAME: “Laurel cimarrén” (Temascaltepec).
The new species is conspicuous for the fulvous or golden-brown pubes-
cence found on the young branchlets, the lower surface of the usually very
pale green leaves, and the inflorescence itself.
This species seems to stand alone. There are features that apparently
relate it to P. longipes and P. podadenia, i.e., the inflorescence-characters,
but a glance at the infructescence destroys this connection. The shape of
the leaves, the difference in stigma and style, and the enlarged perianth-
lobes persistently surrounding the fruit without being reflexed also separate
this species from Persea podadenia.
18. Persea veraguasensis Seemann, Bot. Voy. Herald 193. 1854; Meissner in DC.
P ee ne 51. 1864, as P. veraguensis; Standley in Contr. U. S. Nat. Herb. 23:
291.
Persea one sensu Standley & Calderon, Lista Prelim. Pl. Salvador 85. 1925;
Standley in Contr. = S. Nat. Herb. 27: 183. 1928, in Field Mus. Publ. Bot. 18:
Persea laevigata var. B caerulea Meissner in DC. Prodr. oe 49. 1864, excl. spec.
S. Amer.; sensu Hemsley, Biol. Centr. Am. Bot. 3: 72.
300 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
DistripuTION: In woods of Costa Rica and south into Chiriqui, Panama, at alti-
tudes of 600-2300 meters.
Costa Rica: Without locality, Oersted 7 (cited by Meissner under P. laevigata 8
caerulea, not seen); Patarra, Orozco 341 (fl, Ch). Alajuela: Open woodland of
subtropical zone in semi-shade, Palmira, Alfaro Ruiz, 4. Smith H.964 (fl., A, Ch, NY).
Panama: Chiriqui: Bajo Mono, Boquete, Davidson 516 (fr., A, Ch, Mo) ; savan-
nas of Boquete, Davidson 753 (fl., A, Ch, Mo); Volcan de Chiriqui, Seemann 863 (fr.,
TYPE of P. veraguasensis not seen) ; Rio Chiriqui Viejo Valley near El Volcan, P. White
213 (fr., Mo); in the open Ilanos, about a mile from Bambite, valley of the upper Rio
Chiriqui Viejo, P. White 334 (fr., Mo).
NATIVE NAMES: “Aguacatillo,” “Aguacate de Mico” (Standley ).
This tree varies from 3 to 25 m. high, with the base 50 cm. diam., accord-
ing to the collectors. The branchlets are heavily striate, angled, early
densely subferruginous-sericeous, presently glabrescent and atro-rubescent
or mottled green-brown. The chartaceous leaf-blades are oblong-elliptic
or elliptic, often with a rhomboid base, acute, acuminate, or obtuse, dark
green with lustre above, glabrescent and pale beneath or glaucescent with
sericeous pubescence which may not persist. The costa above is slightly
elevated and canaliculate, and the 8-10 pairs of slender lateral nerves are
obscure above and conspicuous beneath. The petiole is slender to robust
and rusty-pubescent to glabrescent, only faintly canaliculate, and measures
to 2.5~3 cm. in length. The inflorescence consists of axillary few-flowered
ferruginous-sericeous-tomentose panicles shorter than the leaves, borne on
fairly stout sericeous peduncles up to 6 cm. long. The densely pubescent
flowers are up to 4 mm. long, sessile or borne on pubescent pedicels less
than 1 mm. long; the fleshy lobes, buff to tawny-olive according to the
collectors, are unequal, the elliptic outer lobes measuring + 1.25 mm. and
the lanceolate-elliptic inner ones + 2.15 mm. The stamens measure up
Ce ea 5 mm., the hairy filaments slightly longer than the ovate or oblong
anthers of the first two series. The anthers of the third series are oblong
and the filaments are biglandular. The fourth series or staminodia are
thin, flat, ligulate, and hairy, + 1.7 mm. long. The gynaecium is + 3.4
mm. long, the glabrous ovoid ovary only slightly stipitate, being nearly as
long as the sparsely pubescent style, which is to ped by a smallish spread-
ing discoid stigma. The fruits are globose, apiculate, with a bluish bloom,
and measure 10-11 mm. in diameter. They are subtended by a cupule
formed by the persistent spreading and slightly enlarged pubescent perianth-
lobes (pale green), which are supported by the slightly enlarged pubescent
dark red pedicel now measuring up to 3 mm.
This species has been masquerading under the name of P. caerulea, which
species was illustrated by Ruiz & Pavon as Laurus caerulea (Laurographia
t. 2. 1802) from Peru, which is said to occur in Colombia, Venezuela, and
Bolivia as well. This latter species, however, has inflorescences many-
flowered, branched and spreading, and longer than the leaves. The leaf-
blades are 8-23 cm. in length, with the petioles and nerves red. The fruits
are less than 1 cm. in length and are seated on the persistent perianth-lobes,
which are scarcely enlarged, but finally spreading or reflexed, supported by
the hardly enlarged pedicel.
The nearest affinity in our locality is P. podadenia, found only in Mexico.
19. Persea podadenia Blake in Contr. Gray Herb. ns. 52: 62. 1917; Standley in Contr.
. S. Nat. Herb. 23: 291. 1922.
Persea podadenia var. glabriramea Johnston in Contr. Gray Herb. n.s. 70: 69. 1924.
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 301
DistrRIBUTION: In the woods of northern Mexico, predominantly up to
altitude, and also in the aeehe set of Oaxaca and Vera Cruz; possibly in Ais ve
uras.
Mexico: Sonora: Oak canyon near water, in the upper sige cae region, Tepopa,
Rio Mayo, Gentry 2235 (fl., A); oak-forest in moist canyon, Curohui, Rio Mayo,
Gentry 3661 (fl., A, Ch); vicinity of Alamos, Rose, Standley & Russell 13084 (fl., GH).
Chihuahua: Side of a canyon in the oak-country of the upper Sonoran region,
ay ae Rio Mayo, Gentry 2620 (fr.. A). Tamaulipas: La Vegonia, vicinity
of San José, Bartlett 10034 (fr... GH). Nuevo Leon: Scattered along arroyo in
pine-oak wood, Potrero Redondo west to Puerto a Laguna Sanchez and beyond, Muni-
cipio de Villa Santiago, C. H. Mueller 2119 (ir., A); along arroyo bottoms on trails
between Potrero Redondo and Laguna Sanchez, Municipio de Villa Santiago, C. #1.
Muller 2736 (sterile, GH), C. H. Muller 2751 (fr.. GH). Durango: San Ramon,
April 21-May 18, 1906, E. Palmer 119 (fl., isorypE of P. podadenia, Ch, Mo). Vera
uz: Orizaba, Botteri 81 (type of P. podadenia var. glabriramea not seen).
Oaxaca: Zempoaltepec, Liebmann 768 (Lauraceae 79) (fl., Ch.). ? BRITISH Hon.
purAS: Stann Creek: All Pines, Schipp 566 (fr., Ch, GH, NY).
N E NAMES: “Laurel” (Jalisco); “Laurel,” ‘Laurel de la sierra” (Sonora) ;
“Salsafras” (Nuevo Leon).
Persea podadenia, a small tree or shrub, may be recognized at once by
its lanceolate acutish to acuminate greenish leaves 6-15 cm. long and 2-4
cm. broad, that are glabrous and very smooth in texture, except for the
slightly impressed costa on the upper surface, and beneath elagem with
the heavy costa elevated and the delicate lateral nerves slightly so. The
nerves number up to 14 pairs and usually diverge from the costa a an angle
of about 45—55°. The slender pubescent to glabrous petioles are striate
and canaliculate, measuring up to 2 cm. long and 1—1.5 mm. wide. The
short inflorescences barely reach to one- “third: the length of the leaves.
They are paniculate or subcapitate and completely covered with a dense
golden- brown-sericeous pubescence. The flowers are about 5 mm. long
including the short pedicels. The broadly ovate acute outer perianth -lobes
are hairy throughout, + 2.15 mm. long, the inner elliptic-ovate, + m.
long. The stamens of the two outer series are + 1.7 mm. long, the ovate
roundish anthers about equalling the thick pubescent style and with the
cells all introrse. Those of the inner series have anthers that are oblong,
the upper cells lateral, the lower laterally extrorse, and the filaments bear
two subsessile reniform glands near the base. The staminodia are + .8
mm. long, subovate, very pubescent. The stipes equal one-half the entire
length. The glabrous gynaecium is + 2.6 mm. long, the ovary ellipsoid,
almost equalling the rather thick style, aya is topped by a conspicuous
subcapitate stigma. I have not seen the fruit of the type, but additional
material shows an ellipsoid to subglobose eaten purple-glaucous fruit
subtended by perianth-lobes which have become somewhat enlarged and
lignified. The lobes persist and are dichily reflexed, or their tips slough
off leaving only the tube showing the line of decurrence of the lobes.
The variety described by Johnston seems from the description to be
merely a variation of the species not worthy of varietal rank. I have not
seen the type, which is from Orizaba, nor have I seen any material from
that region which would seem to answer the description. Possibly the
British Honduras specimens may match the type.
20. Persea Standleyi, sp. nov.
Arbor 7.5—12 m. alta, ramulis atro-rubescentibus striatis glabrescentibus
302 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
dense foliosis. Folia alternata vel dd lager petiolis leviter pubescen-
tibus striatis rubescentibus satis gracilibu s, ad 3.5 cm. longis et 2 mm. latis,
laminis utrinque glabris coriaceis in sicco Orient. brunneis, lanceolatis
vel oblanceolatis, ad 20 cm. longis et 4.5 cm. latis, basi obtusis saepe obli-
quis, apice obtusis vel attenuato-acutis vel acutis vel subacuminatis, penni-
nerviis, costa supra leviter impressa subtus valde elevata, nervis 10—12-
paribus supra obscuris subtus leviter elevatis angulo 35—45° divergentibus,
rete venularum supra obscuro subtus plus minusve prominulo subareolato,.
Inflorescentia axillaris subcapitata, ioliis brevior, ad 5 cm. longa, brunneo-
fulvo-sericea, pauciflora, pedunculo ad 3 cm. longo pubescente. Flores
immaturi, brevipedicellati, perianthio sca puticonseniee: lobis dis-
tincte 5-nervatis ovatis extus dense pubescentibus exterioribus + 2.15 mm,
longis interioribus + 3 mm. longis; staminibus ser. I & IT + 2.15 mm.
ngis, antheris ovato-oblongis apiculatis, quam filamentis pubescentibus
duplo longioribus, ser. III + 2.6 mm. longis, antheris anguste ovatis obtusis
filamentis pubescentibus re espe conspicue biglandulosis; staminodiis
ovatis pubescentibus + 0.8 mn ; gyna ecio glabro ad 3 mm. longo, ovario
subgloboso longitudine '3 gyn aecii aequante brevistipitato, stigmate con-
spicuissime triangulari angulis decurrentibus. Fructus immaturus(?) viri-
dis, globosus, apiculatus, 9 mm. diam., perianthii lobis pubescentibus non-
nihil rigidis incrassatis (?) plus minusve persistentibus subtentus, pedicello
incrassato pubescente.
DistripuTion: In forests on volcanic slopes of Guatemala at an altitude of
1500-2100 m., and possibly in the region of volcanoes of Vera Cruz, Mexico
Mexico: Vera Cruz: Nogales, Matuda 1118 (fl, A). GuatremMaLa: Chiqui-
mula: Volcan ie lasiass 3-4 miles northeast of Quezaltepeque, Stevermark
31491 (fr oe Solola: Trail between slopes of Volcan Santa Clara and town of
re Paine. alt. 1900-2100 m., June 6, 1942, Stevermark 47130 (f1., TYPE, Ch) (tree 12
; leaves boar membranous, deep green above, paler green beneath)
Persea Standleyi approaches P. podadenia, but may be separated from it
by the generally larger non-glaucescent longer-petioled leaf-blades, which
are subareolate-reticulate beneath, and by the lesser number of lateral
nerves diverging at a smaller angle from the costa. It is possible that more
complete material may prove P. Standley: to be an extreme variation of
P. podadenia.
The species is named for Dr. Paul C. Standley, who, through his collec-
tions and floristic work, has contributed greatly to our knowledge of Mexi-
can and Central American trees and shrubs.
21. Persea sessilis Standley & Steyermark in Field Mus. Publ. Bot. 23: 115. 1944.
DistrRIBUTION: Guatemala, in the mountains, at 2100-2400 m. altitude.
GUATEMALA: Zacapa: Upper slopes, Sierra ot se Minas along oy asi to
summit of mountain, alt. 2100-2400 m., Jan. 12, 13, 1942, Steyvermark 42487 ({fr., TYPE,
Ch) (shrub 1.5 m., leaves firmly thartaceous. i eer dull green pad with
yellow midrib, blue-silvery beneath; peduncle dull rose; fruit green, shining )
his shrub, with its stout terete densely foliose branchlets, may be easily
distinguished by its lanceolate-oblong coriaceous leaves, acute to acuminate
e apex and attenuate but distinctly cordate at the base, borne on thick
stout petioles so short as to make the leaves appear sessile. In the dried
state, the leaf-blades are greenish brown above, castaneous-brownish_ be-
neath, the stout costa bright-castaneous, slightly impressed above and ex-
ceedingly prominent beneath. The lateral nerves, which number up to 20
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 303
face is less so on the lower. The infructescence shows the remnants 0
sericeous pubescence, is shorter than the leaves (about 10-12 cm. long),
and seems to be cymose-paniculate with ascending branchlets. The fruits
are immature, according to Standley, green, shining, globose, about 1
in diameter, and borne on the expanded and patent perianth- -lobes that still
persist and are pubescent. The expanded pedicel is dull rose, according
to the collector.
There seems to be a relation between this species and Persea Standleyi,
also from Guatemala. The most striking difference is the occurrence of
sessile leaves in the former.
DouBTFUL SPECIES OF PERSEA
Persea psychotrioides Sprengel, Syst. Veg. 2: 269. 1825.
The type of this Mexican species has not been seen, but Mez says that it
belongs with Phoebe psychotrioides.
SPECIES EXCLUDED FROM PERSEA
Persea amplexicaulis se aie & Chamisso = PHOEBE AMPIEXICAULIS (Schlech-
tendal & Chamiss
Persea eee Mer i in Standley & Calderén = PuHoreBe AMPLIFOLIA Mez & J. D.
Persea ae ste the Standley = BEILSCHMIEDIA S
Persea effusa Bentham & Hooker = OcCOTEA EFFUSA Decent) Hemsley.
Persea effusa eee PHOEBE EFFUSA Meissne
Persea Gentlei Lundell = NECTANDRA S
Persea Hartwegii (Meissner) Hemsley = PHOEBE PACHYPODA (Ness) Mez.
Persea Matudai Lundell = NectANpRA SINUATA Mez.
Persea mexicana Hemsley = PHOEBE aa Meissner.
Persea ? hide bae — ns & Galeotti = LirsEA GLAUCESCENS var. SUBSOLITARIA
(Meissner) Hem
Persea salicifolia ao Pr reuieias PHOEBE SALICIFO".IA Nees.
2. Phoebe Nees
Phoebe aug: Syst. Laurin. 98. 1836; ka in DC. Prodr. 151: 29. 1864; Mez in
a . Bot. Gart. Berlin 5: 180.
TRIBUTION: In this een in oom and subtropical regions; found in
beccieo, Central America, the West Indies, and South America. Many species occur in
eastern Asia, the number of species caer in India, and also in Malaysia and the
Philippines, with a few species in New Guin
The trees or shrubs of Phoebe in Mies and Central America have alter-
nate or subverticillate leaves that are penninerved or triplinerved. They
vary in shape, size (not more than 30 cm., usually below 12 cm. long), and
texture of the leaf-blades, as well as in pubescence. The inflorescences are
paniculate with no involucre. The flowers are perfect, the tube short or
lacking entirely. The usually equal perianth-lobes are thin or occasionally
fleshy, persistent or deciduous. The stamens of the two outer series bear
304 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
anthers that are usually regularly obtusely ovate, the large introrse cells
occupying the anther almost completely, the two lower cells touching the
upper at their bases. The stamens of the inner or third series bear anthers
that are extrorse or the two lower cells are extrorse and the two upper cells
are laterally extrorse. The staminodia are conspicuous, stipitate-cordate,
the stipes often pubescent. The glabrous (in this area) gynaecium con-
sists of a subglobose or ellipsoid ovary, with a conspicuous style that is
equal to or shorter than the ovary. The stigma is variable, capitate, discoid
or obtuse. The fruit is ellipsoid or subglobose, and borne in a shallow
cupule that frequently bears the remnants of the perianth-lobes on the
margin. The cupule is seated on an enlarged pedicel which is usually ex-
panded considerably at the apex.
KEY TO THE SPECIES OF PHOEBE
A. — blades triplinerved or subtriplinerved.
B. Leaf-blades subtriplinerved or triplinerved, not more than 11 cm. long.
= Leaf- —— elliptic, usually definitely caudate-acuminate.
ung branchlets, leaves, and inflorescence densely ochraceous-tomen-
rapt largest ng blades very broadly elliptic......... 1. P. Tonduzii.
D. Young branchlets, leaves, and inflorescence not ochraceous-tomentel-
lous; largest leaf- blades elliptic.
. Inflorescence longer than leaves.
F. Subtriplinervation obscure ; . axils glandular- pubescent ; leaf-
mee subcoriaceous, not more than 13 cm. long; branchlets gla-
Sat maturity........0....000..... ccc eee ee ee. 2. P. effusa.
F. ‘Triplinervation conspicuous nly lowermost axils glandular-
pubescent; leaf-blades usually pyres coriaceous, 15-17 cm
long; branchlets fulvo-tomentellous, becoming glabrescent... ...
Gia se 4 Bea O48 ew He eG oad a4 bow ue ask 3. P. costaricana.
E. ER shorter than —
F. Pet o 9 mm. long; leaf-blades subcoriaceous to chartaceous;
Aare oe fully oe on enlarged perianth-tube...........
iM Ed RAR GSD Ae ble G aOte Cbs bad Slate ho cee 4. P. Barbeyana.
F. — to 2.5 cm. long; leaf-blades coriaceous; base of fruits
sed in shallow cupule..................... . Brenesii.
C. Leaf- i lanceolate, oblong - lanceolate or ovate- lanceolate, acutish or
acuminate, not caudate-acuminate
D. Apex of the sleaf-blades long- ricate: acuminate; largest blades up to
BE TI. Eo cana ea cioe hii bewecese ohare latina 6. P. longipes.
D. Apex o the leaf-blades not falcate-acuminate; largest blades not more
than 13 cm. long.
E. Lane never more than 6 cm. long; largest leaf-blades not longer
than 9 cm. long and 2-3 cm. broad.
F. Leaf-blades glabrous, concolorous; reticulation elevated above
pase GSP Relat 8 POA Gre bed ahd ee ae ote oe i subtriplinervia,
F. Leaf-blades rufescent beneath and glaucous; ticlation minute,
the veins deeply impressed above............... salicifolia.
FE. Panicles 6.5-12 cm. long, long-pedunculate; largest kat ‘blades n not
less than 11 cm. long, usually not less than 15 cm. lon
F. ser} .. Fi SIGDIOUS. 4 occ eccitic ee is-ne04 . P. areolata.
F, f-blades not areolate, softly pubescent beneath. = P. Arsenei.
B. Leat- . rae definitely sh peated largest leaf-blades not less than 11
m. long, usually not less than 15 cm. long.
C. ide se shorter than ne leaves.
1945]
ALLEN, STUDIES IN THE LAURACEAE, VI
D. Inflorescence composed of numerous subterminal and axillary spike-
like racemose panicles up to 15 cm. long with numerous pean gray-
MUDESCENt MO WELSt tet se cya, orele ait vensianeieleiste wise ele oe . P. mexicana.
D. Inflorescence composed of axillary rather few- Howered cee loose
Soa whee less than 10 cm. ong, more or less glabrous.
eae ae chartaceous, the base cuneate; largest
n 15 cm. lo 1
E. Leaf-blades elliptic, acuminate or caudate-acuminate, loosely and
inconspicuously reticulate mene scalariform), chartaceous or sub-
coriaceous, rounded or obtuse toward the base, - extreme base
narrowly cuneate; largest leaf-blades 18-20 cm. Hg ee
C. Inflorescence longer than the leav
B. L
D.
Subtriplinervation obscure; a as glandular-pubescent; leaf-blades
subcoriaceous, not more than 13 cm. long; branchlets glabrous at ma-
WELL yates 9 Daas 4nd BOT eter e eee P. effusa.
Lb; Serene conspicuous; only lowermost axils glandular- pubescent ;
leaf-blades usually definitely coriaceous, 15-17 cm. long; branchlets
FE
ulvo-tomentellous, becoming glabrescent............ 3. P. costaricana.
A. Leaf-blades penninerved, never triplinerved or subtriplinerved.
argest
leaf-blades 15-18-25 cm. long, never linear-lanceolate; leaves fre-
quently subverticillate
oF ni of leaf-blades onmente, the apex caudate-acuminate ; blades never more
14. P.
n 6 cm. broad, elliptic or lanceolate............. chinantecorum.
C; aa of sae er obtuse, rounded or cordate or auriculate, the apex not
caudate-acu
D. Broadest ee ot the leaf- — below the middle, the blades more or
less attenuate toward the a
Leaf-blades glabrous, are nd amplexicaul...15. P. eens
mollis
E. Leaf-blades pubescent beneath, shortly petiolate.......
E.
co)
middle.
Leaf-blades oblong, oblong-ovate, elliptic, obovate-elliptic, or
obovate-oblong, never auriculate at the base, but obtuse, rounded or
cordate or recur
F. Apex of the ioe blades obtuse or rounded.
. Inflorescence not more than 11 cm. long, pubescent or gla-
brescent; petioles usually 5 mm. long......... . P. Smithii.
G. Inflorescence not less than 15 cm. one, glabrous or pubescent ;
H. Leaf-blades not more than 6 cm. broad and oblong-ovate
or obovate-elliptic; inflorescence branching from the base,
many-flowered and glabrescent.......... 18. P. obtusata.
H. Leaf-blades not less than 7—8 cm. broad, obovate: inflo-
peat not sceneicyie from the base, coiiparatively few-
wered and pubescent. .....66<22..4. 1 . Valeriana.
F. Ape ae the leaf-blades Ror hie or rounded. 20. P. helicterifolic.
Leaf-blades elliptic, not more than 17 cm. long, auriculate at the
base, rigidly coriaceous, above shining green, glabrous, loosely and
conspicuously reticulate, beneath densely minutely appressed brightly
ferruginous-tomentellous.............-.- eee eeeeees 21. P. Salvini.
. Leaf-blades elliptic, recurved at the base, up to 32 cm. long, the
nervation tomentellous and areolate above, ferruginous-tomentose
tlc ears ccee eee ee ee ieteie eee erare eS arem = 22. P. amplifolia.
B. Largest leaf-blades usually not more than 13-14 cm. (rarely 15) cm. long;
lea
ves never subverticillate.
305
306 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
C. Leaf-blades obovate or elliptic
D. Lower surface of the leaf- blades closely appressed-ferruginous-tomen-
tellous, the base auriculate.........00 0.0.00. .0 00 cee eee. 21. P. Salvini.
. Lower surface of the leaf-blades not closely appressed-ferruginous-
tomentellous, the base not auriculate.
E. Leaf-blades 13.5 x 5.5 cm.; apex abruptly and ras acuminate,
er ee ee a ee ee P. padiformis.
digas npud-q a ee wardnkg.caleare's oy tana vow eleals . P. psvchotrioides.
. Leaf-blades lanceolate, linear- or broadly lanceolate or elliptic, never
ovate.
D. cea eer attenuately linear-lanceolate, the largest not more than 2.3
cm. broad at mos
E. Leaf- “blades sialonuks lateral nerves 5—7 pairs; branchlets glabrous
or at most only slightly pubescent................ 25. P. chiapensis.
E. pe blades minutely pubescent beneath; lateral nerves ai 16 pairs;
anchlets minutely pubescent.................. 26. P. tampicensis.
D. Leaf. blades not attenuately ae lanceolate, the largest an less than
.3 cm. broa
E. Lateral nerves 7-11 pairs.
F. Leaves glabrous... 0.00... 0sce0ee0eveeeeens 27. P. pallescens.
. Leaves sericeous beneath. +. si swvees goes 28. P. acuminatissima.
E. Lateral nerves 3-5 pairs (7 at most).
Pubescence of ura nchlets and ae leaf-surface soft.
G. Largest leaf-blades not more than 2.3 cm. broad............
‘o)
cons aars Gata Kae aele are eae bas @aaasa ee 0. P. Boa rial enn
F. Pubescence of branchlets and lower leaf-surface rough; reticula-
tion conspicuous throughout.................... 31 irvine
“— Tonduzii Mez in ire oad 30: Beibl. 67: 15. 1901; Standley in Field
. Publ. Bot. 18: 460. 19
Missense Known only _ the forests of central Costa Rica, at an altitude
of 1800
O Rica: San José: Tree of the forests near El Copey, alt. 1800 m., Feb.
1898, Tondus 11735 (fl., soTyPE, C
NATIVE NAME: ‘‘Aguacatillo aine? (Costa Rica).
This species cannot be confused with any other known from this region,
because of the minute mustard-yellow (ochraceous) tomentum clothing the
young branchlets and the entire inflorescence. The leaves are similar to
pubescence which soon disappears, broadly elliptic, subcaudate-acuminate
at the apex, cuneate at the base. The blades are strongly triplinerviate,
the venation impressed above and prominently elevated beneath. The
the axils densely pubescent glands. e inflorescence, with the flowers still
in bud, consists of few-flowered oe ena or subterminal panicles up to 13
cm. long clustered at the tips of long peduncles up to 8 cm. long. The
flowers have thick hairy ovate lobes, the stamens of the two outer series
with the elliptic mucronate anthers equalling the slender filaments. The
stamens of the inner series bear two reniform stipitate glands at the base of
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 307
the filaments. The staminodia are broadly ovate, cordate, and sharply
caudate-acuminate, hie in size to the stout pubescent stipes. The gla-
brous ovary is subglobos
2. Phoebe effusa Meissner in DC. Prodr. 15!: 33. 1864; Standley in Contr. U.S. Nat.
erb. 23: 295. 1922; Standley & Calderon, Lista Prelim. Pl. Salvador 85. 1925.
Persea effusa Hemsley, Biol. Centr. Am. Bot. 3: 71. 1882, non Benth
Phoebe effusa Y parvifolia Meissner, |.c.
DistriBuTION: In forests of Mexico, in Vera Cruz and Michoacan, presumably very
common, a 2000 m. altitude.
EX ra Cruz: About Vera Cruz and Jalapa, Linden 13 (fl., SYNTYPE of
je Hae not Oe 14 (fl., syntypE of P. effusa not seen, photo., GH), 78 (fl., SYNTYPE
of P. effusa not seen, fragm., NY); Orizaba, in 1854, Botteri 1039 (isosyntype of P
effusa Y parvifolia, NY) ; oe namibia 776 (Lauraceae 44) (fl., Ch); Totutla,
Liebmann (Lauraceae 48) (fl., - Rancho & Barranca Remuda Seve. Purpus 11163,
14184 (fl., Ch) ; open forests cc ae banks, Zacuapan, Purpus 14320 (fl., Ch), 16364
Ghs= Gh). Michoacen: Mt. San Miguel, Tancitaro, Leavenworth & ougsrag!
NaTIVE NAMES: “Pimento,” “Canelito” (Standley, 1925).
This species is one which is difficult to separate from P. costaricana. In
both of these species there is a triplinerviate condition of the leaves, which
is. however, much more pronounced in P. costaricana than in P. effusa.
The latter has faintly striate terete branchlets which are fulvo-tomentellous,
becoming glabrescent with age. The branchlets of P. costaricana are
striate, sulcate, or even angled, and become quite glabrous at maturity.
The leaves of P. effusa are variable in shape and size, the blades chartaceous
to subcoriaceous, 6-13 cm. (according to the original description) long and
—5.5 cm. broad, the base cuneate, the apex obtuse or acute to caudate-
acuminate or abruptly acuminate, all variations of the apex frequently
occurring on the same sheet. The lateral nerves vary from 3-6 pairs, all
of equal size and all bearing pubescent glands in their axils, or the lower-
most may be subtriplinerviate. The blades are glabrous throughout except
for the midrib, supported by slender, canaliculate, pubescent petioles
usually up to 1 cm. in length. The leaves of P. costaricana are also vari-
able, having thickly coriaceous blades which measure 8-17 cm. in length
and 3-6 cm. in width, with about the same variation in the apex that occurs
in the leaf-blades of P. effusa. The base of the leaf-blades of P. costaricana
is more attenuately cuneate. The lateral nerves usually number 4 pairs,
the lowermost pair often being extremely conspicuous and bearing pubes-
cent glands in their axils, whereas the upper pairs are obscure and glandless.
The petioles are usually stout, glabrous or glabrescent, and are up to 15
mm. in length. In both species under discussion the slender loosely flow-
ered paniculate glabrous inflorescences are usually longer than the leaves
(up to 18 cm. long), and are borne on slender peduncles which make up at
least half their entire length. There is a tendency for the pedicels of the
flowers of P. effusa to be longer than the perianth; whereas those of P.
costaricana are about equal to the perianth, rarely surpassing it. In spite
of the closeness of the two entities. they seem to be separate species. No
fruiting specimens are yet known of P. effusa.
308 JOURNAL OF THE ARNOLD ARBORETUM [VOL, XXVI
3. Phoebe costaricana Mez & Pittier ex Mez in Bull. Herb. Boiss. II. 3: 230. 1903;
Standley in Field Mus. Publ. Bot. 18: 459. 1937.
DistRIBUTION: Known in Costa Rica, from the type-locality at 640-1200 m. alti-
tude and in Alaiiicla, and in Chiriqui Province, Panama, at 1000-1700 m. altitude.
Costa Rica: Puntarenas: Canas Gordas, alt. 1200 m., Feb. 1897, Pittier 11107
(fl., isorypE, GH, NY). Alajuela: Hills above Santiago, near San Ramon, Brenes
14404 (fl, GH). Panama: Chiriqui: Bajo Mono, Boquete, Davidson 583 (f1.,
Ch, Mo); forests about Boquete, Pittier 2998, 3146 (f1., Ch) ; Finca Lérida to Bonasie.
Woodson, Allen & Seibert 1099 (fr., A, Ch, NY, Mo).
NaTIVE NAMEs: “Sigua,” “Sigua blanca” (Panama).
Under P. effusa there is a discussion of its relationship to P. costaricana.
The fruiting specimens from Chiriqui show loose spreading infructescences
that are still slender, bearing large ellipsoid definitely apiculate fruits about
2 cm. long and 1 cm. broad. The cyathiform or subhemispherical some-
times urceolate cupule bears the remnants of the enlarged persistent
perianth-lobes. The enlarged pin is frequently constricted ek below the
lob The cupule is up to 9 mm. long, rarely 1 cm. in diameter at the
widest portion (below the apex 4 the tube), and about 5 mm. deep, and
the pedicel is 6-10 mm. long and expanded at the apex to 3 mm. in diameter.
4. Phoebe oe Mez in eres Bot. Gart. Berlin 5: 209. 1889; Standley in Contr.
U5.
. Herb. 23: 295.
micttniven Known gus eae the type-locality.
Vera Cruz: ear Orizaba, May 12, 1866, Bourgeau 2436 (fl., fr., TYPE
Mexico:
not seen, fragm. & photo., me ee NY); Orizaba, August 1866, Bourgeau, s.n. (fr.
Ch).
This little known species bears branchlets that are vellowish-pubescent
at the apex, becoming glabrous, brown and early angled, presently becoming
terete. he leaves are subcoriaceous, elliptic, the base cuneate, the apex
beneath opaque and glabrous except for the pubescent glands in the axils
of the lowermost lateral nerves and sometimes the upper pairs. The leaves
are penninerved, with a seal Ani aa condition obscure on the upper
surface but very conspicuous benea The lateral nerves number 3—5
pairs and are more or less obscure ee but delicately prominent beneath,
the lowermost pair being the most conspicuous. The costa is impressed
above and more prominent beneath. The blade, 8-11 cm. long and 3.6—4.5
cm. broad, is subtended by a canaliculate minutely pubescent petiole up to
9mm. long. Mez described the inflorescence as glabrous and subracemose-
paniculate, shorter than the leaves. The flowers are glabrous with a short
perianth-tube. The ovate acute outer lobes are slightly shorter than the
inner. The glabrous filaments aig one-third shorter than the anthers,
which are broadly siast rounded at the apex, manifestly constricted at the
margin over the lower cells. The staminodia are very large, broadly
cordate, acuminate, ee with pilose stipes. The ovary is glabrous,
subglobose, attenuate into a style which is almost longer, thick-cylindrical
at the apex, with a subdiscoid obtuse stigma. The fruit, Pens ellipsoid,
measures 11 mm, long and 8 mm. in diameter, and is s seated on the lobes o
late perianth, distended at the base, and the subhemispherical thickened
pedic
Phoebe Barbeyana seems to be related to P. neurophylla, from Costa
Rica, but has much smaller though more coriaceous leaves. The fruits of
the two species are not unlike.
1945 | ALLEN, STUDIES IN THE LAURACEAE, VI 309
5. Phoebe Brenesii Standley in Field Mus. Publ. Bot. 18: 459. 1937.
DistrRIBUTION: Central Costa Rica at 600-1000 m. altitude, and in Chiriqui Prov-
ince, Panama, at 1140 m. altitude
Costa Rica: Alajuela: Near San Ramon, Brenes 377 (538) (fl., Ch) ; Piedades
near San Ramon, Brenes 4896 (fr., Ch); “San Pedro,” near San Francisco de San
Ramon, Brenes 6675 (fl, Ch); La Palma de San Ramon, Brenes 6810 (fl., Ch);
between San Miguel and La Palma de San Ramon, Feb. 14, 1933, Brenes 17048
(fl., TypE, Ch); Atenas, in sielowers tropical zone, in open pasture, clay-loam, A. Smith
P.2445 (fl, A). Panama: Chiriqui: Bo oquete, Davidson 641 (fr., A, Ch, Mo).
This tree has slender angled branchlets that early lose their fulvo-tomen-
tellous pubescence and become striate and glabrescent. The aera
elliptic or oblong-elliptic (rarely hens elliptic) abruptly acuminate o
caudate-acuminate leaves are triplinerved and except for the i Saeae
venation become glabrous very early. ‘they are borne on slender glabrous
petioles up to 2.5 cm. long, and they are up to 10 cm. long and 4 4 (-5) cm.
broad. They bear axillary pubescent glands in the lowermost pair of
lateral nerves. The inflorescence is loosely paniculate, glabrous or gla-
brescent, up to 12 cm. long, and usually shorter than the leaves. The
The ovate perianth-lobes are thinnish and + 2.5 mm. long. The two outer
series of stamens measure + 1.7 mm. long, the oblong somewhat emarginate
anthers being twice the length of the slender filaments. The inner stamens,
m. long, with their more narrow anthers equalling the filaments,
bear two reniform stipitate glands near the base of the filaments that almost
equal the anthers in length. The cordate staminodia are + 1.4 mm. long,
the pubescent Annee being almost one-half the entire length. The glabrous
gynaecium, measuring + 2.4 mm., consists of a subglobose ovary topped by
a stout style slightly longer than the ovary and bearing a conspicuous capi-
tate stigma. The fruit is oblong- ellipsoid, 12 * 5 cm., seated in a shallow
cupule 5 mm. long and 5 mm. broad at the apex and me mm. deep, the rem-
nants of the partially deciduous perianth-lobes forming the shallow cupule.
The pedicel is thickened and up to 5 mm. long, expanding slightly toward
the apex.
This species is similar to P. mexicana in foliage, but is at once separated
by the loose spreading inflorescence, P. mexicana having a more spike-like
strict panicle. It also recalls P. costaricana, but the leaves are much
smaller, and the inflorescence less heavily flowered as well as shorter.
6. Phoebe longipes Johnston in Contr. Gray Herb. n.s. 70: 69. 1924.
DistrRIBUTION: Mexico, known only from the type-locality.
Puebla: Along the river near Honey Station (Trinidad), May 6, 1904,
Pringle 8829 (fl., rypE, GH).
This species belongs with the triplinerved group with inflorescences
shorter than the leaves, but it could never be confused with any other
known species in this complex. The entire plant seems to be glabrous,
except for the suggestion of pubescence in the axils of the lowermost pairs
of lateral nerves. The branchlets are conspicuously alate and expanded at
the nodes, with a glaucous bloom which early disappears. The leaf-blades
are pale green above, paler beneath, the reddish brown venation frequently
standing out in contrast, like the petioles, branchlets, peduncles, and pedi-
cels. The blades are lanceolate or oblong or narrowly ovate-lanceolate, the
310 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
tips falcate-acuminate or eee acuminate, the — cuneate or rounded
(rarely), measuring 12—22 cm. lon ng an nd 4-5.5 ¢ broad. The lateral
veins, except for the ieeaeacal pairs which are triplinerved, are rather
inconspicuous, numbering up to 6 pairs. The reticulation is obscure above
but conspicuous and almost areolate beneath. The petioles are sturdy
enough, early glaucous, and measure up to 2.5 cm. The few-flowered very
slender inflorescences are axillary, up to 7 cm. long, borne on filiform
peduncles up to 3 cm. long. The flowers, also with a pruinose bloom, are
quently as long as 1.5 cm. The perianth-lobes are thin, membranaceous,
— acute, + 2.8 mm. long. The stamens of the two outer series are
+ 2.8 mm. long, and consist of slender filaments pubescent at the base and
somewhat longer than the ovate obtuse anthers. The stamens of the inner
series (+ 2.6 mm. long) bear two large — basal glands which
equal the oblong obtuse anthers. The filaments are about one and one-
third the length of the entire stamens. The ses et are triquetrous and
conspicous, about 1.25 mm. long overall, borne on stout stipes that are
pubescent and equalling one-half the entire length. The glabrous gynae-
cium measures + 3 mm. The ovoid-ellipsoid ovary about equals the
slender style, which is topped by a conspicuously three-parted stigma.
— subtriplinervia (Meissner) Standley in Contr. U. S. Nat. Herb. 23: 294.
San subtriplinervia Meissner in DC. Prodr. . 125. 1864.
Ocotea subtriplinervia Hemsley, Biol. Centr. Am. Bot. 3: 74. 1882.
Phoebe Galeottiana Mez in Jahrb. Bot. Gart. Berlin : 200. 1889.
DistriBUTION: Known only from the syntypes, from Mexico.
ear Jalapa, Galeotti 7061 (fl., fragm. of IsosyNTYPE of
Oreaduphe alii Sig Mirador, Linden 17 (fl., syntyPE of O. subtripli-
nervia n e
pei to the original description, this is a very glabrous species.
The single leaf available is coriaceous, lanceolate, acuminate, obscurely sub-
triplinerviate, and with the exception of the more prominent costa, slightly
elevated above and very conspicuously so beneath, the venation is every-
where of about the same size and prominence. The blade is glabrous
throughout except for the pubescent axillary glands of the lowermost lateral
nerves. The margin is undulate an e blade is up to 6.5 cm. long and
1.5 cm. broad, the broadest portion below the ape It is subtended by
a slender canaliculate glabrous petiole up to 1 cm. long. The entire blade
throughout is prominently reticulate. iene describes the racemes as
axillary and subterminal, about 2.5 cm. long, 5-flowered, with nodding
pedicels (peduncles?) up to 6.5 mm. long. The white or yellow flowers are
1 line long, and the lobes of the perianth are deciduous. From the cited
fragment of the Galeotti number, I find the flower to be about 2 mm. long,
supported by a slender almost glabrous pedicel nearly twice its length.
The broadly ellipsoid perianth-lobes are + 1.7 mm. long and thickish, with
a suggestion of — near the apex. ‘The two outer series of stamens
are + .8 mm. long, the almost square anthers nearly once and a half times
the length of the fairly slender filaments. The stamens of the inner series
are 1 mm. long and bear at the base of the filaments two very conspicuous
and spreading sessile glands. The gynaecium is glabrous, + 1.25 mm.
long, consisting of a scarcely stipitate subglobose ovary equalling the taper-
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 311
ing style, ae is capped with an inconspicuous stigma decurrent down one
side of the
With ae es material at hand it is impossible to say with certainty
what the affinity of the entity may be. Possibly it is somewhere near P.
salicifolia or its allies.
8. Phoebe ae ae in Linnaea 21: 488. 1848; Standley in Contr. U. S. Nat.
Herb. 23: 295.
Persea salicifolia es. Biol. Centr. Am. Bot. 3: 72. 1882.
DistrisuTION: Central Mexico; very little material available.
Mexico: State unknown, near Regla, Ehrenberg 875 (fl., fragm. of TYPE, Ch) +
Hidalgo: Jacala, in deep ravine, Chase 7311 (tr., Ch, GH
This species, so little known, has branchlets that hecuue glabrescent and
are first somewhat angled, later terete and finely striate. The leaves are
lanceolate to oblong-lanceolate, acuminate or subacute, subtriplinerved,
glabrous above except for the costa, subcoriaceous, glaucous, ve green in
dried state, paler beneath, the blades up to 9 cm. long and 2 (—3.2) cm.
broad. The reticulation is minute, the veins deeply pn above and
Ls hea elevated beneath. The costa is impressed above and con-
peduncle, few-flowered. The flowers are glabrous, stare ah up to 3.3
mm. long, with a slender pedicel + 4.25 mm. long. The ovate perianth-
lobes measure + 2.15 mm. long and are thinnish and sree inside near
base. The two outer series of stamens measure + 1.2 , the oblong
mucronate anthers equalling the slender ee The emer of the
inner series are slightly longer, + 1.5 mm., and bear two reniform short-
stipitate glands at the base of the eae a4 the length of the anthers.
The staminodia are + 1 mm. long, triangular, with pubescent stipes more
than half the entire length of the staminodia. The gynaecium is glabrous,
+ 1.7 mm. long, the globose ovary not quite equalling the slender style in
length. The stigma is conspicuous, almost peltate. The fruit is sub-
globose-ellipsoid and dark purple i in the dried state, 13 10 mm., and
seemingly slightly stipitate. It is seated on a cupule which is infundibuli-
form and measures 6 mm. long and 6 mm. broad at the apex. The pedicel
is thickened and about 4-5 mm. long. The perianth-lobes, persisting at
the apex of the shallow cupule for about 2 mm. in depth, presently break off
irregularly, leaving a flat disk to which the stipe of the fruit is attached.
The nearest relatives of the species are P. areolata and P. Arsenei. Their
differences will be discussed under the latter species.
9. Phoebe areolata Lundell in Contr. Univ. Mich. Herb. 7: 13. 1942.
DistrIBUTION: In the mountains of Mexico and in British Honduras.
Mexico: Chiapas: Saxchanal, Sierra Madre, alt. 2700 m., is 1, 1941, Matuda
4296 (fl, A, TypE— Mich, NY). British Honpuras: Stan Creek: Mountain
ridges, 19 mile, Stann Creek Valley, Schipp 971 (fr., Ch, GH, NY); Pine Peak, Cocks-
comb branch of South Stann Creek, Stevenson XII, XVI (fl., Ch)
Outstanding are the glabrous conspicuously areolate ovate-lanceolate
leaves of this species. The blades are rounded at the base and acuminate
at the apex, up to 11 cm. long and 4 cm. broad, supported by glabrous
312 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
canaliculate petioles up to nearly 1 cm. long. The costa is slightly im-
pressed, but brownish and conspicuous against the green blade above, and
the lateral nerves are obscure. Beneath both are elevated. The lateral
nerves are 4—6 pairs, all bearing pubescent glands in their axils, the lower-
most pair simulating a triplinerviate condition. The axillary rather few-
flowered paniculate inflorescences, up to 12 cm. long, are very sparsely
pubescent, and are borne on rather slender reddish brown peduncles up to
: he campanulate flowers, about 3.5 mm. long, are subtended
by long (7 mm.) saeipe and the ovate lobes are fairly fleshy. The two
outer series of stamens are + 1.9 mm. long, the ovate apiculate anthers
being only slightly mee than the Glaments The anthers of the inner
series of stamens, which measure + 2.15 mm., are oblong, and the filaments
bear two small subbasal stipitate glands. The ovate acuminate staminodia
are + 1 mm. long, with pubescent thick stipes somewhat less than half the
entire length. The glabrous ellipsoid ovary nearly equals the style, which
is crowned by a conspicuous peltate stigm
The closest affinities of the species are with P. salicifolia and P. Arsenei.
A discussion will be found under the latter species.
10. Phoebe Arsenei, sp. nov.
Arbor, ramulis fulvo-tomentosis deinde glabrescentibus valde striatis.
cuneatis vel Laviker obtusis, apice Bera vel obtuse acuminatis raro
emarginatis, penninerviis vel subtriplinerviis, costa supra impressa pubes-
cente laa elevata, nervis lateralibus 5-7 paribus: leviter elevatis, rete
venularum supra minuto et valde impresso, subtus conspicue prominulo.
fitpecnta axillaris et subterminalis paniculata ad 10 cm. longa, glabra,
pauciflora, longipedunculata, pedunculo gracili ad 7 cm. longo leviter
pruinoso, Flores ad 3 mm. longi, brevipedicellati, pedicellis ad 2.5 mm.
longis gracilibus, perianthio campanulato, lobis ovatis acutis -iphagh yous
mf is ad basin incrassatis; staminibus ser. I & I 1.1 mm.
longis, antheris ovato-ellipticis obtusis filamentis crassis ener ue
duplo longioribus, ser. III + 1.7 mm. longis biglandulosis, glandulis reni-
formibus stipitatis antheris oblongis emarginatis aequalibus; staminodiis
ovatis membranaceis paps avai basi pubescentibus, + 1 mm, longis;
gynaecio glabro + mm. longo, ovario nal stylo subaequali,
stigmate subcapitato . Fructus ignotus
DistrRisuTION: Known only from the type-locality.
ExIco: Michoacan: Vicinity of Morelia, Rincon, alt. 1950 m., April 11, 1909,
Arséne 2448 (fl., TypE — GH, NY).
This species seems to be very near P. areolata and P. salicifolia. It is
separated from the former by its softly pubescent lower leaf-surface and
branchlets and its conspicuously reticulate lower leaf-surface. Phoebe
areolata is practically glabrous throughout, the axils of the very prominent
triple-nerves at the base show conspicuous pubescent glands, and the blades
are conspicuously areolate. The shorter inflorescence and leaves of P.
salicifolia separate the two species.
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 313
11. Phoebe mexicana Meissner in DC. Prodr. 151: 31. 1864; Standley in Contr. U. S..
Nat. Herb. 23: 294. 1922, in Trop. Woods 21: 17. 1930, in Field Mus. Publ. Bot.
10: 201. 1931; Standley & Record in op. a 144. 1936; Standley in op. cit.
18: 459. 1937; Yuncker in op. cit. 9: 290.
Persea cinnamomifolia Hemsley, Biol. Centr. pen ‘Bot. 3: 71. 1882, non H.B
Phoebe mexicana var. Bourgeauana Mez in Jahrb. Bot. Gart. Berlin 5: 214. ne
TIoN: Generally throughout southwestern Mexico and Central America, at
varying cuiiae: from 15 m. along the coast to 2800 m. in the mountains of Guatemala.
Vera Cruz: In the valley of the city of Cordoba, April 20, 1866,Bour-
geau 2302 (ir: ae of P. mexicana var. Bourgeauana, Ch) ; Fortuho, Coatzacoalcos
River, L. Willig: 8920 (fr., A, Se NY ), 9142 Sipe ree near Jalapa, Galeotti 7026
(fl., Ch), 77 (fl., GH) ; Zacuapan, rocky plains, Purpus 5991 (fl., GH), 14328 (fl., Ch).
Oaxaca: Vicinity of Cafetal Concordia, Morton & Patt 2444 (young fruit,
Ch). Chiapas: Escuintla, Matuda 1082, 2632 (fl, A, NY). Guatemara: Hue-
huetenango: Cerro Canana, between Nucapuxlac and Canana, Sierra de los Cuchu-
matanes, Stevermark 49045 (fr., Ch). dena Vicinity of Retalhuleu, Standley
88624 (fl, Ch). Honpuras: ‘Cond agua: River bank on plains near Siguatepe-
que, Yuncker, Dawson & Youse 5811 (fr, GH). Tegucigalpa: Mont. de la Flor,
pine and oak r region near river, C. & V. W. von Hagen 1227 (sterile, NY).
Subirana, in open semi-tropical valley, C. & V. W. von Hagen 1051 (sterile, NY);
vicinity of Coyoles, in the Aguan River valley, Yuncker, Koepper & Wagner 8630 (fr.,
Ch, GH, NY). pais ad In deep forest near Lancetilla, Yuncker 4945 (fr., Ch).
BritisH HONDURAS: El a 0: Pepsi vaey on hillside, Gentle 2499 (fl., A, NY).
Big Eddy Ridge, ‘a Broken ae “Gentle 3549 (fr, A, NY). ‘eledo: lope Creek,
along river-bank, Schipp 281 (fr., Ch, GH, NY). Costa Rica: San José: Near the
River Virilla, on old trail from Heredia to San José, Brenes pe (fr., Ch); between
the graveyard Calvo and “ Rio Maria Aguilar, Rojas 404 Ch); vicinity of El
General, Skutch 4042 (fr , NY); vicinity of San José, Baers 47344 (fl., Ch).
Alajuela: Atenas, in nee tropical zone in clay-loam pasture, A. Smith P.2440
ee A); Alajuela, in the mountains, Torres 115 (fl, Ch). Panama: Canal Zone:
Near Culebra, Pittier 3438 (fl, GH) ; hospital grounds at Ancon, Pittier 3957 (fr., GH).
NaTIVE NAMES: ‘“Aguacatillo” (Honduras, Costa Rica); “Aguacate negro” (Hon-
duras) ; “Boy Job,” “Wild Pear” (British Honduras).
This species is distinct because of the numerous strict raceme-like panicles
blades are pean with varying degrees of pubescence, usually entirely
glabrous above and often so beneath except for the pubescent glands in the
axils of the lower pair of lateral nerves. The blades are elliptic, lanceolate
or oblong-lanceolate, cuneate at the base or somewhat obtuse or rounded,
usually ae to caudate-acuminate at the apex, up to 15 (—26) cm.
long and 5 (—10) cm. broad. The margin is very often slightly undulate,
and the costa is impressed above and very prominently elevated beneath.
The lateral nerves are except for the lowermost pair rather obscure, less so
beneath than above, and they number up to 7 pairs. The lowermost are
conspicuous and diverge from the costa at an angle of about 25°, ascend-
ing nearly to the middle of the blade before reaching the margin. ‘The
uppermost pairs diverge at an angle of about 55°. The petioles are stout,
canaliculate, glabrescent to glabrous, and measuring up to 2 cm. in length.
314 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
The numerous inflorescences are many-flowered, pubescent to glabrescent,
reddish, with peduncles not more than 5 cm. ‘and usually much less in
length. The small white flowers are sessile to shortly pedicellate and
fulvous-pubescent. The flowers are 3-4 mm. long, campanulate, with
seca pe -lobes thin, ovate, and +2.6 mm. long. The stamens are + 2.15
m. long, the slender filaments about equalling the elliptic anthers. The
i eleairs of the inner series of stamens bear two subreniform stipitate
glands at their bases, equalling about one-third the entire length of the
stamens. The glabrous gynaecium, + 2.6 mm. long, consists of a sub-
globose to obovate substipitate ovary topped by a slender style one and a
third times longer than the ovary. The stigma is conspicuous, dilated, and
discoid. The glabrous fruits are slightly obovoid, apiculate, about 1 cm.
long and 7 mm. broad, seated on a glabrous cupule that is campanulate,
about 5 mm. long, 7 mm. broad, and + 2 mm. deep, which bears the rather
brittle remains of the perianth-lobes. The pedicel is enlarged somewhat to
about 5 mm. lon
The variety, according to Mez, differs from the species in bearing leaf-
blades which are always obtusish at the base, perianth-lobes which remain
erect in fruit, pedicels less thickened, and fruits that are definitely sub-
globose, 8 mm. long and 6 mm. broad. This variation does not seem to
warrant a varietal status. To my mind it is merely another manifestation
of the species proper.
12. Phoebe Ehrenbergii Mez in vines = Gart. Berlin 5: 201. 1889; Standley in
Contr. U. S. Nat. Herb. 23: 294.
Phoebe Hartmanii Johnston in one a Herb. n.s. 70: 69. 1924.
STRIBUTION: Known only from northwestern and central Mexico, 450-1960 m.
re in arroyos.
Mex Sonora: Arroyos of the tropical Sonoran regions, Vinata, Rio Mayo,
Gentry 1878 (fr., ra Ch); deep canyon-bottom in short-tree forest, Arroyo Gochico,
Rio Mayo, Gentry 3629 (fl., A, Ch), 3630 (fr. & young fr., A, Ch). Chihuahua:
Batopilas, April, 1892, Hartman 1029 (fl., young fr., isotype of Phoebe Hartmanii, Mo) ;
Oca River, Rio Mayo headwaters, LeSueur 1246 (abnormal fl. G
Sinaloa: Ortega 4617 (abnormal fi., Ch). es Arroyo de los Hornos, Hari.
enda del Ototal, w. of San Sebastian, Sierra Madre Occidental, common along stream-
side, Mexia 1803 (fr., A, Ch, GH). Mexico: Ph eres gs Ehrenberg 712 (fl.
fragm., peo of P. Ehrenbergii, Ch, photo., Ch, ; Rincon, Hinton 378 (abnormal
fl., GH, NY), 3032 (abnormal fl., GH) ; aaa Hinton 3895 (abnormal fl., fr.,
H); Hide Hinton 5771 (Al., $6 , GH) ; Carboneras, Hinton 6826 (abnormal fl., GH).
NaTIVE NAMES: “Aguacatillo” (Mexico); “Bebelama” (Sonora).
Some of the cited specimens have been placed in Persea, and many sheets
have also been identified as Sassafridium macrophyllum, the type of which
is none other than the widespread Vectandra globosa. An examination of
the type-material of the two species P. Ehrenbergii and P. Hartmanii seems
to place the entity in the triplinerved group of Phoebe.
The entire tree seems to be glabrous, the young branchlets striate, some-
what angled, losing their early tomentellous condition. The elli iptic or
uring up to 15 cm. long and 4-5 cm. broad. They are usually cuneate at the
ase, occasionally obtuse, and the apex is acute or acuminate. The midrib
and lowermost lateral nerves are more conspicuous than the delicate upper
lateral nerves, which number as many as 5 pairs. The minute reticulation
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 315
is obscure on the upper surface and less so on the lower. The inflorescence
is slenderly paniculate, up to 12 cm. long, few-flowered, and more or less
pruinose. The somewhat pruinose flowers are nearly 5 mm. long, sub-
tended by slender pedicels of equal length or slightly longer. The subequal
perianth-lobes are elliptic, thin, slightly pubescent at the inner surface near
the base, and + 2.8mm. long. The stamens are + 1.7 mm. long, the ovate
or oblong anthers equalling the stoutish filaments. Those of the inner
series bear two reniform subsessile glands at the base about one-third the
length of the entire stamen. The staminodia are nearly sessile, ovate,
cordate, and + 0.8 mm. long. The glabrous gynaecium is 2.5—-3 mm. long,
the globose ovary being about 24 the entire length. The slender style is
topped by a conspicuous capitate stigma. The fruit is ellipsoid, minutely
apiculate, measuring 12 X 8 mm., subtended by a shallow spreading red-
dish pruinose cupule 4-6 mm. long and 8 mm. broad at the flat disk-like
apex. The pedicel is about 3 mm. long and slightly enlarged. The
perianth-lobes approaching anthesis develop an abscission layer about one-
half their length, and the upper half is early deciduous. The lower portion
of the lobe persists well into the fruiting stage, and finally when the fruit is
mature falls off. The specimens from Jalisco are somewhat like P. longipes
in leaf-shape, and possibly may eventually prove to be the fruiting stage of
that species. More material is needed to be able to determine this, however.
13. Phoebe neurophylla Mez & Pittier ex Mez in Bull. Herb. Boiss. I. 3: 231. 1903;
Standley in Field Mus. Publ. Bot. 18: 460. 1937
DISTRIBUTION: Costa Rica, in forests and on river-banks at varying altitudes up to
1900 m. Possibly in Mexico at very low altitudes.
MeExIc Oaxaca:(?) In llanos, Tuxtepec, Chiltepec, and vicinity, Martinez-
Calderon 119 (fl., A). Costa RIca: untarenas: On the banks of the River Hur,
valley of the Diquis, alt. 800 m., March 15, 1898, Pittzer 12054 (fr., isoTYPE, US) (small
tree). Alajuela: Continental divide 2 miles n.w. of Zarcero, at edge of forest in
rich humus, A. Smith 142 (fl, Ch). San José: Vicinity of El General, Skutch 3844
(fl., A, NY), 4329 (fr., A, Ch, NY).
This species is very near P. costaricana in vegetative characters. The
chartaceous to coriaceous elliptic leaf-blades, rounded or obtuse toward the
base, the extreme base narrowly cuneate, the apex acuminate or acuminate-
caudate, up to 21 cm. long an (—8) cm. broad, are glabrous except for
the distinctly pubescent glands in the axils of the lowermost pair of lateral
nerves. The upper pairs of lateral nerves are obscure, numbering 3 or 4
and obscurely camptodromous, giving the impression of a typically tri-
plinerviate leaf. Transverse venation is usually fairly inconspicuous. The
axillary and subterminal inflorescence, unlike that of P. costaricana, is
short, not longer than 6 cm., and is glabrous throughout and comparatively
few-flowered, with peduncles not more than 2 cm. long. The whitish
campanulate flowers are about 4 mm. long, the thin ovate acute lobes almost
equalling the tube, and are subtended by slender filiform pedicels up to 5
mm. long. The two outer series of stamens are approximately 1.9 mm.
long, the rounded ovate anthers almost as long as the slender filaments.
The inner series measure + 2.15 mm. long, the large conspicuous glands
almost equalling one-half the length of the entire stamens. The large con-
spicuous staminodia are subcordate, + 1.25 mm. in length, the thick stipes
equalling one-half the entire length. The glabrous gynaecium is about 3
mm. long, the style one and one-third times the length of the subglobose
316 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXVI
The stigma is spreading, discoid, and conspicuous. ‘The fruit is a
whi ellipsoid nana berry, 18 x 12 mm., subtended by a flaring shallow
red cupule, more or less minutely rugose, ‘with an entire margin. The
cupule is less tha § 5 mm. long, 1.3 cm. in diameter, and not more than 2—3
mm. deep. The — pedicel is up to 12 mm. long and is expanded to
5 mm. at the apex
14. Phoebe chinantecorum Schultes in Bot. Mus. Leafl. Harvard Univ. 9: 170. 1941.
isan Known only from the type-locality.
Ex1co: Oaxaca: Dark forest, San Juan Lalana, Choapam, alt. 550 m., May 8,
rr Schultes & Reko 827 (fl., isotype, GH) (small slender weak tree).
NaTIVE NAME: ‘‘Mo-gwu” (Oaxaca).
This little known species has slender branchlets, as well as petioles,
clothed with a dense ferruginous tomentum, the former bearing at the apex
a subverticillate whorl of large elliptic or lanceolate-elliptic chartaceous
cuminate at the apex. e blades measure up to 20 cm. in length and 6
cm. in width, the upper surface shining and glabrous gigi) for the pubes-
cent costa, the lower densely pubescent on the venation e few-flowered
subpyramidal paniculate inflorescences are up to 6.5 cm. long (or more?),
the slender pubescent peduncle up to 4.5 (—6) cm. The campanulate flowers
are about 3.5 mm. long, equalling the slender glabrescent pedicels. The
oblong lobes are fleshy, + 2.5 mm. long. The somewhat petaloid stamens
of the two outer series measure + 1.25 mm.; the anthers are subsessile and
more or less oblong, the connective tissue occupying the upper third of the
anthers. ‘Those of the inner series have anthers obovate, longer than the
stout pubescent filaments bearing two sessile glands equalling them in
length. The staminodia are cordate, + 0.7 mm. in — the stipes
almost the entire length. he glabrous gynaecium is + 1.5 mm., the ovoid
or subglobose ovary shortly oo and bearing a short thick style with
the subcapitate stigma at its
Very probably the species is but a variation of the widespread P. helic-
terifolia. Further collections may show a line of development toward this
type.
15. Phoebe amplexicaulis (Schlechtendal & Chamisso) Mez in — Bot. Gart. Berlin
5: 216. 1889; Standley in Contr. U. S. Nat. Herb. 23: 293.
Persea one plewbccalls ponggraaee & Chavniase in Linnaea 5: a 1830; Nees, Syst.
Laurin. 137, 672. 1836 (excl. var.), in Linnaea 21: 490. 1838; Meissner in DC.
Prodr. 151: 54. oni ey var.) ; Hemsley, Biol. Centr. Am. Bot. 3: 71. 1882
DistRIiBUTION: Known only from the mountains of Mexico.
Mexic exico: Cerro Colorado, in mountains, Schiede & Deppe 87 (fr., TYPE
of Persea ‘aeeplesicnslis not seen, photo., GH).
The single collection of this species may very possibly be a fruiting speci-
men of P. mollis which has become glabrous with age. The variation of
leaf-base shape in this complex is so great that one might reasonably expect
to find within a species sessile amplexicaul leaves grading into shortly
petiolate cordate leaves. Until the type of P. amplexicaulis is available for
direct comparison, I shall maintain it as a distinct species
Schlechtendal and Chamisso describe the leaf-blades as glabrous, cori-
aceous, as opposed to those of the latter species P. mollis, subsessile, oblong,
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 317
cordate-amplexicaul at the base, long-acuminate into an obtuse acumen
with a slight margin. The leaves measure nearly 12 cm. long and about 2
cm. broad. The axillary few-flowered cymes are long-pedunculate, with
the peduncles filiform, the entire cymes measuring nearly 6.5 cm. long.
The fruit is less than 1 cm., and the flowers unknown.
16. Phoebe see Mez in Jahrb. Bot. Gart. Berlin 5: 192. 1889; Standley in Contr.
U.S. 1 Herb. 23: 293. 1922
Daten Known aa from Mexico and western Guatemala at 1300-2100 m.
altitude.
Without locality, Haenke s.n. (fl., TyPE not seen), possibly an isotype
(labeled 1562 in Ch). GUATEMALA: an Marcos: Above Finca El Porvenir, on
“Todos Santos Chiquitos,” lower sear facing slopes of Volcan Tajumulco, Steyermark
37201 (fl, Ch).” Suchitepéquez: Upper forested slopes of barranco by Loma
Grande, above Finca El Naranjo, on Volcan Santa Clara, Steyermark 46854 (fl., Ch).
Phoebe mollis is near P. amplexicaulis in leaf-shape, and eventually, with
more material for study, may prove to be a flowering specimen of that
species. The branchlets are heavily fulvous-tomentose. he coriaceous
or subcoriaceous leaf-blades are oblong- lanceolate, rounded or cordate at
the base, and long-acuminate at the apex, measuring up to 18 cm. long,
although usually 10-15 cm., and 5—6 cm. broad, subtended by Stout tomen-
tose petioles up to 1 cm. long. The impressed costa is visible above and
frequently tomentose; the lateral nerves are obscure and number upwards
of 10 pairs, diverging at an angle of 60—80°, and are inconspicuous like the
ive aes Beneath, the costa and lateral nerves are conspicuously
ated and tomentose, and the reticulation for the most part prominent.
The hiciae. few-flowered paniculate inflorescence is early heavily fulvous-
tomentose, later becoming glabrescent. The slender peduncle is up to 8
. Th
1.7 mm. Their perianth-lobes are broadly rotund-ovate, subacute, rather
thick and + 1.7 mm. long. The rotund-ovate anthers ‘of the two outer
series of stamens are about twice the length of.the filaments, the entire
length being + 1 mm. long. Those of the inner series are more or less
obtusely ovate, with two sessile basal glands the length of the filaments.
The original description mentions sagittate stipitate staminodia. The
flowers I have dissected are not fully developed and show at most thin strap-
shaped structures. The glabrous ovary is topped by a short style and more
or less inconspicuous stigma.
17. Phoebe Smithii, sp. nov.
Arbor ad 15 m. alta, ramulis sulcatis dense fulvo-tomentosis vel glabres-
centibus. Folia alternata vel subverticillata, juventute membranacea plus
minusve molliter pubescentia, mox coriacea et sparse pubescentia dense
praesertim nervationibus, petiolis robustis dense fulvo-pubescentibus ad
5 mm. longis, laminis ellipticis vel obovatis vel oblongo-ellipticis, 11—15
(—20) cm. longis et 5-6 (—8) cm. latis, basi cordatis, obtusis, vel rotundatis,
apice rotundatis, obtusis, a obtuse subacuminatis, penninerviis, costa
supra leviter subtus conspicue elevata, nervis lateralibus 8—10-paribus supra
leviter subtus prominule elevatis, angulo 35—45° divergentibus, rete ven-
ularum supra obscuro subtus laxo et conspicuo. Inflorescentia axillaris et
ultimis et floribus glabris glaucis. Flores magni 5—6 mm. longi et 15 mm.
318 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
diam., pedicellis gracilibus ad 7 mm. longis; perianthii lobis ellipticis car-
nosis intus papillosis ad 4.5 mm. longis; staminibus ser. m.
longis, filamentis brevibus, antheris subreniformibus et subpetaloideis ;
staminibus ser, III antheris subrectangularibus filamentis conspicue big-
landulosis aequantibus; staminodiis conspicuis ovatis brevistipitatis, + 1
mm. longis; gynaecio glabro + 2.5 mm. longo subgloboso ovario brevisti-
pitato stylo subaequali; stigmate conspicue discoideo. Fructus ellipsoideus
minute apiculatus 3 cm. longus et 1.8 cm. latus, cupula hemisphaerica 1 cm.
longa et 1.5 cm. diam. et 6 mm. alta subtentus, pedicello robusto 8 mm.
longo et utrinque 4 mm. diam.
DistripuTION: Known only from Alajuela and San José in Costa Rica, at
1800-2200 m. altitude.
Costa Rica: Alajuela: Hills above Baranca de Zarcero, at edge of forest, March
22, 1938, A. Smith P.C. 367 (fl., type, Ch) (tree 15 m.; leaves coriaceous, shining,
flowers cream-yellow; fruit green; cupule crimson), H. 523 (f., Ch); Zarcero, Alfaro
Ruiz, A. Smith 10072 (fr., A). San José: El Tablazo de San José, Valerio 34 (ir.,
Ch)
This species, with its fulvous-tomentose sulcate branchlets, has charac-
teristics very like those of P. Valeriana. It also recalls the variable P. helic-
terifolia, but is easily separated by the much shorter inflorescences with a
glaucous bloom.
18. Phoebe obtusata Lundell in Contr. Univ. Mich. Herb. 6: 21. 1941.
DistrRIBUTION: Known only from type-locality.
Mexico: Chiapas: Buena Vista, near Escuintla, Jan. 1938, Matuda 1887 (f1.,
TYPE, Mich).
This species is related to the P. helicterifolia complex but may be sep-
arated by the longer very full-flowered inflorescences, which are subterminal
and branch profusely. The usually smaller leaves with obtuse or rounded
apices and the shorter less harsh pubescence also serve to distinguish Lun-
dell’s species.
e stout branchlets are densely fulvous-tomentose. The leaves are
alternate or subverticillate at the apex of the branchlets, the blades oblong-
ovate to obovate-elliptic, up to 16 cm. long and about 6 cm. broad, rounded
or obtuse at the base, occasionally subcordate, the apex obtuse or rounded.
The petioles are stout, densely tomentose, up to 1.2 (—1.5) cm. long. The
costa and nerves are impressed above and would be obscure were it not for
the pubescence persisting on them after the upper surface of the leaf be-
comes glabrous or glabrescent. Beneath, they are conspicuously elevated
and densely tomentose. The lateral nerves number to 9 or 10, the extreme
upper pairs obscure, and diverge from the costa at an angle of about 45°.
The axillary or subterminal inflorescence consists of a main terminal stalk
or panicle (to 22 cm. long) with 1-3 shorter panicles (8-15 cm. long)
branching from the base of the peduncle, which is short, stout and pubes-
cent. The pubescence disappears toward the apex of the inflorescence.
The entire inflorescence has a purplish cast extending to the perianth-tube.
The flower is similar to that of P. helicterifolia. In fact, it is probable that
more abundant collections will prove the two entities to be conspecific, with
only geographical or ecological variation.
19. Phoebe Valeriana Standley in Field Mus. Publ. Bot. 18: 460. 1937.
DistRIBUTION: Costa Rica, in forests at 1050-1800 m. altitude.
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 319
Costa Rica: Alajuela: Region of Zarcero, upper tropical zone within Atlantic
cloud-forest, A. Smith H. 268 (fl., Ch, NY); La Pefia, Alfaro Ruiz, Caribbean cloud-
forest, A. Smith P. 2120 (fl., A); woods of Los Angeles de San Ramon, near La Paz,
Brenes 4185 (197) (fl., Ch); Los Angeles and La Paz de San Ramon, Brenes 6099 (fl.,
fr. Ch). San José: Forests of El Copey, Feb. 1898, Tonduz 11746 (fl., Type, Ch,
fr.. US); Santa Maria de Dota, Stork 2405 (ir., Ch):
NATIVE NAME: “Quizarra” (Costa Rica).
The species has stout angled sulcate branchlets covered with a dense
vellowish or brownish tomentum, eventually becoming more or less glabres-
cent. The petioles are canaliculate. thick, and equally tomentose, up to
2.5 cm. long. The subcoriaceous blades are obovate or occasionally oblong,
long, the stout peduncles up to 8 cm. long, the subtending leaves frequently
deciduous. The flowers are large. up to 5 mm. long and 8.5 mm. in diam-
eter, the lobes spreading. The lobes are thin, elliptic, acute or obtuse, up
to 4.25 mm. long. he stamens of the two outer series are + 1.7 mm.
long, the anthers almost sessile, ovate. apiculate. Those of the outer series
are + 2.15 mm. long, and the subrectangular anthers are only slightly
longer than the stout filaments that bear two conspicuous subglobose sub-
sessile glands. The staminodia are + 0.8 mm. long, conspicuous, ovate or
3-4 mm. long, and about 6 mm. in diameter at the apex.
Standley relates the species to P. betazensis, a synonym of P. helicteri-
folia. It is also closely allied to the Mexican species P. obtusata, which
has smaller leaf-blades and a more copiously flowered inflorescence
branched from the base and glabrescent.
20. Phoebe helicterifolia (Meissner) Mez in Jahrb. Bot. Gart. Berlin 5: 193. 1889;
Standley in Contr. U. S. Nat. Herb. 23: 294. 1922, in Trop. Woods 21: 17. 1930;
Standley & Record in Field Mus. Publ. Bot. 12: 144. 1936; Yuncker in Field Mus.
Publ. Bot. 9: 290. 1940.
Oreodaphne mexicana Meissner in DC. Prodr. 151: 118. 1864.
Oreodaphne mexicana var. @ subsessilis Meissner, ].c.
Oreodaphne mexicana var. 8 longipes Meissner, l.c.
?Oreodaphne mexicana var. ¥ diminuta Meissner, l.c.
Oreodaphne helicterifolia Meissner in op. cit. 123.
Ocotea helicterifolia Hemsley, Biol. Centr. Am. Bot. 3: 73. 1882.
Ocotea mexicana Hemsley, I.c.
Ocotea mexicana var. a subsessilis Hemsley, l.c.
Ocotea mexicana var. 8 longipes Hemsley, l.c.
Ocotea mexicana var. y diminuta Hemsley, |.c.
320 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXVI
Phoebe betazensis Mez in op. cit. 192.
Phoebe nectandroides Mez in op. cit. 194.
DistRIBUTION: Southern Mexico and Guatemala, i in melanie yon at low altitudes
of 250-450 m., and higher up in pastures or in pine-forests up t
Mexico: Ve a Cruz: Orizaba, Botteri 1018 (f1., type > ee mexicana
var. B longipes “al seen, fragm., NY). Guerrero: Montes de Oca, Barranca, San
Antonio-Buenos Aires, Hinton 14014 (fl, GH). Oaxaca: In Sierra San Pedro-
Nolasco, Talea, Galeotti 7004 (f1., isosyntype of Oreodaphne mexicana var. « subsessilis,
(#1, syntype of Oreodaphne mexicana var. @ subsessilis not seen); in forest between
Ozumazin and Rio Chiquito, Schultes & Reko 726 ({r.,GH). Chiapas: San Bartolo,
alt. 150 m., Feb. 1840, Linden 1641 (fl, isotype of Oreodaphne helicterifolia, photo.,
GH, NY); along stony brooks or mountain- -sireams, Fenix, Purpus 10086 (fl, NY),
10356 (fi., GH). GUATEMALA: Alta Verapaz rar H. Johnson 578 ru Ch),
Rubeltein, Steyermark 44207 (f1., Ch); lowland forest in valley, “pantano,” 214 miles
west of Cubilgiiitz, Steyermark 44280 (f1., vie Cerro de Agua Tortuga (Sahacoc),
vicinity of Cubilgtiitz, Steyermark 44648 (f1., fr., Ch).
The present species is variable in scale every locality where it is
known to occur. It varies in leaf-shape and size as well as in density of
tomentum, but there can be no doubt that all of the specimens cited fall
into the same species. The branchlets are densely or with age more
sparsely clothed with a soft yellowish- or brownish-fulvous tomentum.
The oblong, elliptic or obovate, obovate- elliptic or obovate-oblong usually
membranaceous leaves are alternate or in whorls about the apex of the
branchlets. The leaf-blades have one consistent characteristic: they are
usually broadest through or above the middle. The blades are se ea
The base is cordate to rounded, occasion-
ally merely obtuse, and the apex is acuminate or abruptly acuminate, the
acumen very short or up to 1.5 cm. long, obtusely or very sharply caudate.
The blades are 9-15 (—25) cm. long and 2—9 (-12) cm. broad, and are
subtended by a short stout densely tomentose peduncle at most up to 1 cm.
long, frequently so short on the upper leaves as to make them appear ses-
sile. The costa and lateral nerves, of which there are 8 or 9 (rarely —12)
pairs, diverging at an angle of about 45° (the short lowermost almost at
right angles), are impressed above but conspicuous because of the pubes-
cence. They are elevated and densely pubescent beneath. The reticula-
tion is obscure above and slightly more prominent beneath. The paniculate
inflorescence is axillary and loosely few-flowered, usually up to 25 cm. lon
with a purplish color conspicuous on the stem and flowers, the latter fre-
quently exhibiting a purplish bloom on the entire aun The inflorescence
is scattered-pubescent toward the apex and borne on a long comparatively
slender pubescent peduncle up to 13 cm. long. The flower is 5-6 mm. long,
borne on a slender pedicel of about the same length or shorter. The strap-
like or narrowly elliptic rather thick perianth-lobes are about 3 (-4) mm
long, spreading and reflexed at anthesis. The two outer series of stamens
stances, + 1.5-2.5 mm. long, with very short filaments. Those of the
inner series are more or less oblong, almost equalling the a bearing
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 321
two sessile or slightly stipitate conspicuous sometimes spongy glands nearly
the size of the anthers. The staminodia vary, stipe-like to ovate, up to
+ 0.8 mm. long, the slender stipe one-half their entire length. The gla-
brous ovary is more or less globose, usually slightly longer than the rather
thick style with its usually conspicuous subcapitate stigma, the entire
gynaecium measuring up to 3.2 mm. he infructescence has become thick-
ened, occasionally longer, and often less pubescent. The fruits are black,
glabrous, ellipsoid, apiculate, 2.4 1.7 cm., and are seated on a shallow
cyathiform red cupule 3-4 mm. long, 7 mm. broad, and 2-3 mm. deep.
The thickened pedicel is up to 9 mm. long expanded to about 4 mm. at its
apex.
The nearest relationships are apparent in the species P: amplexicaulis,
P. mollis, and P. obtusata, all from Mexico, for discussion of which see these
species.
21. Phoebe Salvini (Mez) Lundell in Contr. Univ. Mich. a 6: 23. 1941.
Ocotea Salvini Mez in Jahrb. Bot. Gart. Berlin 5: 264.
DistRIBUTION: Guatemala and southern Mexico, up to 2800 m. altitude.
Mexico: Chiapas: Rodeo, near Siltepec, in pre forest, Matuda 4578 (fr., A,
NY). Guatemata: Without locality, Aguilar 492 (fr., Ch). Quiché: In remnant
of cloud-forest, Nebaj, Skutch 1679 (fl., A, Ch). Pircarevn eee Volcan de
Fuego, near Calderas, alt. 2590 m., 1873-4, Salvin (fl., TYPE of Ocotea Salvini not seen,
fragm., Ch); region of Los Positos, above Las Calderas, Standley 80191 (fl., fr., Ch).
Solola: Volcan San Pedro, north-facing slopes toward Lago de Atitlan, above village
of San Pedro, in damp cloud-forest dripping with mosses and hepatics, Steyermark
47215 (sterile, Ch).
From the leaf-fragment at Chicago, it is apparent that Standley’s number
80191 is without question the species concerned. e minute ferruginous
leaf-surface, petioles, and inflorescence is in striking contrast to the bright
shining green glabrous or slightly scattered pubescent upper leaf-surface
which is very conspicuously reticulate. The coriaceous leaf-blades are
elliptic, auriculate at the base and acute to acuminate at the apex, 9-17 cm.
long and 4—9.5 broad, subtended by stout canaliculate pubescent petioles
usually less than 8 but sometimes up to 10 cm. long, with its stout branches
and peduncle (to 4.5 cm. long) conspicuously striate. The tomentellous
flowers are 3-5 mm. long, subtended by pedicels of equal length. The
elliptic lobes are fleshy and more or papillose, up to 3 mm. long. The
two outer series of stamens are + 1.5 mm. long, with short stoutish fila-
ments slightly pubescent at the base ne one-half the length of the broadly
oblong anthers. e stamens of the inner series are + 1.7 mm. long, the
somewhat squarish anthers equalling the filaments which bear at their bases
two compressed sessile glands about one-third the length of the entire
stamens. The glabrous gynaecium, + 2.5 mm. in length, consists of a
subglobose or broadly ellipsoid ovary more than twice the length of the
stout style, which is topped by a triangular rather inconspicuous stigma.
The fruit is ellipsoid, 3 x 2.3 cm., seated on flattened disk-like cupule 3
322 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
mm. long, 14 mm. broad, and 1-2 mm. deep, which is very woody with a
epee almost double margin. The subtending pedicel is also enlarged
o 1.5 cm. in length and 6 mm. in width at the apex.
22. Phoebe amplifolia Mez & J. D. Smith in Bot. Gaz. 19: 261, t. 24.
Persea amplifolia Mez in Standley & Calderon, Lista Prelim. PI. CEE 85. 1925.
DistripuTION: Guatemala and from Costa Rica.
aura Re El] Jute, alt. 3050 m., April, 1892, Heyde & Lux 3033 (fl,
isoTYPE, GH, NY El Progreso: Between Finca Piamonte and top of Montana
Piamonte, prea ae Pacayal, Stevermark ree (fl., Ch). Costa Rica: San Jose:
Near Rancho Redondo, near San José, Popenoe 984 (f1., Ch).
This species has very stout a that are deeply sulcate and clothed
with ; aes close ferruginous tomentu The canaliculate petioles are also
sulcate, and are minutely tomentose sat up to 3.2 cm. long. The coriaceous
blades are elliptic, the base cuneate and recurved, the apex shortly an
ruptly acuminate, 28-30 cm. long and 13-16 cm. broad. The upper
ees is glabrous except for the persistent pubescence scattered along the
venation and densely so on the costa and lateral nerves.The costa is slightly
elevated above near the base and conspicuously so throughout on the lower
surface. The lateral nerves, of which there are 8-12 conspicuous pairs
diverging at an angle of 45-60", are impressed above and prominently ele-
vated beneath. The reticulation is obscure and impressed above and con-
spicuously elevated beneath. The inflorescence is axillary, densely tomen-
tose, rather few-flowered, up to 13 cm. long, the stout peduncle up to 9 cm.
The large flowers are nearly 5 cm. long and densely tomentose, with a short
pedicel not more than 3 mm. long. The thick fleshy lobes are spt
pubescent and eset and are + 3.8mm. long. The stamens of the tw
outer series are + 1.7 mm. long, the almost quadrate rounded anthers onus
three times the length of ye filaments. Those of the inner series are + 1.
mm. long and the filaments bear two conspicuous sessile glands at the base.
The staminodia are + 1.25 mm. long, ovate, the pubescent stipes almost
one-half the entire length and often bearing near the apex two small
roundish sessile glands. The glabrous gynaecium is + 2.5 mm. long, the
subglobose ovary slightly longer than the slender style, which bears a con-
spicuous triangular stigma. The ellipsoid fruit is 3 cm. long and 2 cm
broad and, according to the authors, is subtended by a thick sibpaterifrin
cupule that is obscurely double-margined and gradually narrows into t
strongly enlarged pedicel.
The nearest relative of the species is P. Salvini, from which it is easily
distinguished by the usually smaller leaves of the latter, which are covered
on the lower surface with a finely appressed dense bright ferruginous tomen-
tum, and the base of which is conspicuously auriculate rather than merely
recurved.
23. Phoebe padiformis Standley & Steyermark in Field Mus. Publ. Bot. 23: 117. 1944.
DisTRIBUTION: Known only from the eastern part of Guatemala, at an altitude of
570 m
GUATEMALA: Quezaltenango: sie eine Skutch 1358 (fl., A, Ch, NY, US),
alt. 570 m., Oct. 4, 1934, Skutch 1367 (f1., A, Type — Ch, NY, US) (tree 15 m. high and
about 32 cm. diam.; crown full and rounded; we ‘panatls, brown; flowers greenish
white).
This grayish species has striate fulvous-tomentose branchlets which are
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 323
quickly glabrous. The chartaceous leaf-blades are elliptic or oblong-elliptic
or occasionally emanne e and obovate-elliptic, more or less obtusely
acuminate, and supported by pubescent or glabrescent sulcate petioles up
to 12 mm. long. The blades are up to 13 cm. long and 5.5 cm. broad,
greenish on the upper surface and brownish on the lower. A slight pubes-
cence is apparent on both surfaces when very young, but it quickly disap-
pears except for occasional remnants on the midrib above and beneath.
The base is cuneate and the apex acute or acuminate to subcaudate-acumi-
nate. The costa is impressed above and elevated beneath as are the 6 (—8)
pairs of lateral nerves, which bear pubescent glands in their axils on the
lower surface, and which diverge from the costa at an angle of about 45°.
The loosely paniculate axillary and subterminal inflorescences are pubes-
cent, becoming glabrous, shorter than the leaves (usually not longer than
8 cm.), and the branches are very narrow, giving a spicate appearance. In
this respect the species resembles P. mexicana, but it differs in the smaller
leaves which are penninerved rather than triplinerved, and in the more
glabrous inflorescence. The flowers are about 3.4 mm. long and are borne
on pedicels only slightly longer. The ovate acute thin perianth- ee
measure + 2.5 mm. long. The two outer series of stamens are +
long, the filaments shorter than the oblong rounded anthers, The s ns
of the inner series are + 2.4 mm. long, the anthers more narrowly ane
and shorter than the thick filaments, which have two conspicuous stipitate
reniform glands at their bases. The conspicuously ovate cordate stami-
nodia are + 1.25 mm. long, the thick pubescent stipes almost one-half the
entire length. The glabrous gynaecium is + 2.4 mm. long, with the very
slender style, topped by an inconspicuous discoid stigma, equalling the sub-
globose ellipsoid ovary.
24. ee ame ia (H.B.K.) Mez in Jahrb. Bot. ne baa 5: 191, t. 3, fig.
889; Standley in Contr. U. S. Nat. Herb. 23: 293.
os pegrarnenys H.B.K. Nov. Gen. & Sp. 2: 129 rez. ae
DistriBpuTION: Eastern Mexico, in Vera Cruz.
Mexico: Vera Cruz: Near Jalapa, in moist vaileys, alt. 1244 m., ci &
Bonpland 4434 (fl., type of Ocotea psychotrioides not seen); near Vera Cruz, Sumi-
chrast 940 (fr., NY); Orizaba, Botteri 1040 (fl., NY); une pemetinel A ae
aceae 69) (fl., GH); Mirador, Liebmann [14941] (1, Ch); n damp forests, Zacua-
pan, Purpus 2959 (fl., Ch, GH, NY); forests near Temes ae 10460 (fr., NY);
rocky slopes, Barranca de Tenawnas Purpus 16400 (fl., Ch).
This species may eventually be found to be a variation of P. Pittieri or
of P. Bourgeauviana. Seemingly it differs from these in that its leaves are
more elliptic than lanceolate and occasionally are obovate-elliptic. The
flowers seem to be on the whole much smaller, although they show a tend-
ency toward reflexed lobes typical of the two latter species. A study of the
type may settle the question of relationship.
This very tall tree has terete glabrous branches and slender branchlets
that are ae and pale subferruginous-pubescent. The slender petioles,
also pubescent, are short, not more ian 5 mm. long at most. The leaf-
blades are lanceolate, the base cuneate and the apex shortly and abruptly
subacuminate or subacute, 5 (—7) cm. long and 2 (—2.5) cm. broad, char-
taceous or subcoriaceous, the upper surface smooth and shining, the lower
pubescent to glabrous, the venation only slightly elevated above and promi-
nently so beneath, everywhere more or less pubescent. The costa and lateral
324 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXVI
are pubescent at the base. Those of the inner series are + 1.7 mm, long,
the rectangular anthers equalling the filaments that bear two subsessile
globose glands at the base. The staminodia are usually up to + 0.8 mm.,
occasionally longer, are ovate and borne on stipes equalling one-half their
entire length. The glabrous gynaecium is + 1.7 mm, long, the ovoid ovary
about equalling the style, which bears a stigma which is subcapitate or flat
but dilated, in any case conspicuous. The fruit of Purpus 10460, which
seems to agree with the flowering material in foliage-characters, is ellipsoid-
subglobose, minutely apiculate, up to 12 mm. long and nearly 10 mm.
broad, subtended by a flat disk-like cupule (bearing remnants of the lobes
of the perianth) about 1.5 mm. long and 8-9 mm. in diameter, the support-
ing pedicel slightly enlarged to sometimes nearly 15 mm. in length and
expanded at the apex to 4 mm. in diameter.
25. Phoebe chiapensis Lundell in Contr. Univ. Mich. Herb. 6: 21. 1941.
DistriBUTION: Known only from the type-locality.
Mexico: Chiapas: Mt. Ovando, Dec. 1937, Matuda 2064 (fl., isotypr, A, NY)
(tree 7-8 m. high, 20 cm. diam.).
This species is striking with its slender reddish branchlets, petioles, and
inflorescences, that are sparsely glabrescent. idvein of the almost
linear-lanceolate acuminate glabrous leaves is also reddish in contrast to the
green color of the blade. The costa is prominent beneath, whereas the
delicate lateral nerves, of which there are 5—7 pairs, are fairly obscure
throughout. The reticulation is apparent on both surfaces. The leaves
up to 5 cm. long and glabrous. It bears flowers which have ovate acute
perianth-lobes and are borne on pedicels that are sometimes 8 mm. long.
The stamens of the two outer series are + 1.25 mm. long, the pubescent
very short pubescent stipes. he glabrous gynaecium consists of a sub-
globose ovary, slightly longer than the tapering style, and a conspicuous
capitate stigma. The fruit is unknown.
Lundell has related this species to P. pallescens and P. salicifolia. The
former has leaves that are lanceolate or elliptic, with the lateral nerves num-
bering upwards of ten. The inflorescence of P. pallescens is twice as long
as that of P. chiapensis. The latter species may be easily separated from
P. salicifolia by its penninerved leaves which show no trace of a tendency
to a triplinerved condition. The convex reticulation of the leaf-blades of
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 329
P. salicifolia is not to be found in the leaves of P. chiapensis. Floral char-
acters also separate the latter from the two above-mentioned species. Just
where one would place the species is a question that may not be solved until
more and better material is available. The type-specimen is injured by
galls, which in many cases are apt to distort the structure typical of a
given species.
26. Phoebe tampicensis (Meissner) Mez in eg i Gart. Berlin 5: 2300. 1889;
Standley in Contr. U. S. Nat. Herb. 23: 294. 1
LOSS tampicensis Meissner in DC. ee 78 136. 1864.
Ocotea tampicensis Hemsley, Biol. Centr. Am. Bot. 3: 74. 1882.
Phoebe angustata Blake in Contr. Gray Herb. n.s. 52: 63. 1917. (sphalm., in Ind.
Kew. Ocotea)
DistripuTIon: Mexico, chiefly in Tamaulipas and San Luis Potosi, from 300-500
m. altitude.
ulipas: Near Tampico, Berlandier 132 ae fragm. of TYPE of
Oreodaphne tampcensi NY); vicinity of Victoria, alt. 320 m., Feb. 1—April 9, 1907,
E. Palmer 80 (fl., type of P. angustata, GH, isotype, NY), Sierra near Victoria,
Jaumave, von ae 719 (fl., Ch, NY), 750 (fl., Ch) ; Cuesta del Abra, LeSueur 147
(fr., Ch). San Luis Potosi: Valles, San ee Potosi, Kenoyer A. 606 (fr., Ch);
limestone hills, Las Palmas, Pringle 3794 (ir.. GH). Oaxaca: Choapam, coffee
plantation to noneli: Yaveo, Mexia 9140 (fl.. GH, NY).
FP. tampicensis, like the preceding species, has attenuately linear-
lanceolate leaf-blades cuneate at the base and acuminate at the apex, which
to 2.3 cm. broad. They are early pubescent but soon glabrous throughout
except for occasional pubescent glands in the axils of the lateral nerves.
They are subtended by a minutely pubescent soon glabrous slender petiole
which may be light yellowish and may measure up to 1.3 cm. long. The
midrib is visible above but conspicuously elevated beneath, whereas the
lateral nerves and reticulation are everywhere obscure. The lateral
nerves number 12-16 pairs, diverging at an angle of about 40° The
inflorescence consists of axillary loose many- ae slightly pubescent
panicles not more than 3 (—7) cm. long, with a peduncle not more than 2
cm. long. The flowers are broadly campanulate, about 3 mm. long, equal-
ling te slender slightly pubescent pedicels. The perianth- lobes are elliptic,
rather thick, and + 2.5 mm. long. The two outer series of stamens are
+ 1.25 mm. long, the subquadrangular anthers twice the length of the
rather thick filaments. The stamens of the third or inner series are + 1.5
mm. long, bearing conspicuous stipitate glands which, as well as the fila-
ments, equal the eatin pe emarginate anthers. The glabrous gynae-
cium measures + 1.9 mm. long, the ellipsoid ovary nearly twice the length
of the rather thick style, which is topped by a slightly enlarged onice
stigma. The fruit is broadly ellipsoid, apiculate, olive- greenish, glabrous,
up to 1.8 cm. long and 1.3 cm. broad, very smooth on drying, seated on a
flat disk-like cupule about 1 mm. long and 5 mm. in diam., which is sub-
tended by the i pedicel to about 6 mm. in length and 3 mm. in diam-
eter at thea apex
The species is most closely related to P. acuminatissima Lundell, from
which it differs in its linear-lanceolate leaves with more numerous and more
obscure lateral nerves and larger fruit. The floral structure of the two
species is very dissimilar.
326 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
27. Phoebe pallescens Mez in Jahrb. Bot. Gart. Berlin 5: 218. 1889; Standley in
Contr. U. S. Nat. Herb. 23: 293. 1922.
Distrisution: Known only from the type-locality, in Mexico.
Mexico: Vera Cruz: Near Orizaba, in 1855, F. Mueller 90 (fl., fr., IsoTYPE, NY).
This species stands out because of the glabrous character of its slender
reddish striate branchlets and its glabrous lanceolate or elliptic leaves,
which are glaucous beneath. The leaf-blades are chartaceous, greenish
brown above. the base obtuse to cuneate, often oblique, the apex subacute to
acuminate, with a slender canalicuate petiole up to 17 mm. long, and meas-
ure up to 10 cm. long and 3.3 cm. broad. The midrib is reddish yellow,
impressed above and elevated beneath. The lateral nerves are indistinct
above and only delicately elevated beneath, numbering up to I] pairs. The
reticulation is everywhere apparent. The glabrous loose subpyramidal
paniculate few-flowered slender inflorescences are up to 10 cm. long, equal-
ling or slightly longer than the leaves, the peduncle being as long as 8 cm.
The subcampanulate flowers are up to 2.15 mm. long, and outstanding be-
cause of the long slender pedicels reaching a length of 8 mm. The ovate
lobes are very thin, measuring + 1.9 mm. long. The stamens of the two
outer series are + 1 mm. long and the ellipsoid apiculate anthers are twice
the length of the filaments. The stamens of the inner series are + 2.15
mm. long, with the oblong-ovate anthers equalling the stout filaments, which
bear two conspicuous reniform shortly stipitate glands at the base, not quite
equalling the filaments in length. The glabrous gynaecium measures +
1.5 mm., the subglobose ovary longer than the slender style, which bears a
discoid subtriangular stigma at its apex, according to Meissner. He notes
that the fruit is abnormally large, seated on the greatly enlarged cylindrical
pedicel with the lobes deciduous. I have not seen a fruiting specimen.
The affinity seems to be with P. acuminatissima and with P. mollicella,
from the first of which species it is separable by the glabrous leaves and
from the second by the larger number of lateral nerves.
28. Phoebe acuminatissima Lundell in Contr. Univ. Mich. Herb. 6: 19. 1941.
Phoebe saxchanalensis Lundell in op. cit. 7: 14. 1942.
Distripution: In the mountains of Chiapas, Mexico, at 2700 m., and Guatemala
at 250-2000 m. altitude.
1co: Chiapas: Mt. Ovando, Dec. 1937, Matuda 2107 (fl., isoTyPE of P.
. cm. diam.) ; Sierra Madre, Saxchanal, in virgin
forest, Matuda 4311 ({fr., isotype of P. saxchanalensis, A, NY) (small tree 5 m. high
GUATEMALA: Quezaltenango: Lower south-facing slopes of Volcan Santa Maria,
between Finca Pirineos and Los Positos, between Santa Maria de Jess and Calahuaché,
Stevermark 33750, 33752 (fr., Ch); slopes and ridges between Quebrada Chicharro and
Montafia Chicharro, on southeast-facing slopes of Volcan Santa Maria, Steyermark
34356 (fr., Ch). Sacatepéquez: Wooded slope above Duenas, Standley 63141 (f.,
Suchitepéquez: At edge of ravine, Finca Moca, Skutch 2099 (fl., A).
_
burro, along trail to San Francisco Tecuamburro on summit of volcano, north of Chi-
quimulilla Steyermark 33156 (fl.,
The small tree of 5 m. described from Chiapas by Lundell in Guatemala
reaches a height of 20 m. The very slender branchlets are early covered
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 327
but persists on the lower. The membranaceous lanceolate long-acuminate
leaf-blades (up to 12.5 3 cm.) are supported by slender pubescent
petioles 10-17 mm. long. The base is cuneate and often unequal. The
costa is impressed above and conspicuously elevated beneath, whereas the
slender lateral nerves are scarcely visible above and hardly more so beneath.
There are upwards of 19 obscurely visible pairs of lateral nerves diverging
from the costa at an angle of 45—55-70°. Frequently the pubescent glands
present at their junction are made conspicuous by the infiltration of ex-
traneous matter. The few-flowered axillary minutely pubescent panicles,
shorter than the leaves (4—5 [—7] cm.), arise on almost filiform peduncles
up to 3.5 cm. long. The thick fleshy heavily papillose-tomentose perianth-
lobes are + 2.5 mm. long, oblong and reflexed at anthesis. he stamens
of the two outer series are ++ 1.25 mm. long, the ovate obtuse anthers some-
what longer than the slender filaments. Those of the inner series are about
1.5 mm. long, the subquadrate anthers equalling the filaments bearing at
the base two subsessile subglobose glands. The ovate staminodia are
+ 0.4-0.8 mm. long, borne on slightly longer pubescent stipes. The sub-
globose glabrous ovary is slightly longer than the style, which is topped by
a conspicuous peltate stigma. The glabrous fruits are ellipsoid-obovoid,
up to 12 mm. long and 7 mm. broad, subtended by a cupule which is about
2 mm. long, 5 mm. in diameter, and about 1 mm. in depth. Lundell gives
larger measurements, so presumably the sheet at Michigan has fruit in a
more advanced stage. The pedicel is enlarged and about 5 mm. long, the
apex expanded to about 3 mm. in diameter.
The nearest relative is P. mollicella, from Costa Rica, which may be at
once distinguished by its smaller leaves with fewer pairs of lateral nerves
and shorter petioles. The pubescence of the latter species is consistently
more yellowish than gray, a feature which alone would not carry much
weight. Also the flowers are greater in number, and their structure differs
materially.
29. Phoebe mollicella Blake in Contr. Gray Herb. n.s. 52: 64. 1917; Standley in Field
Mus. Publ. Bot. 18: 459. 1937.
DistRIBUTION: Known only from the forests of Costa Rica, up to an altitude of
2135 Mm:
osta Rica: Alajuela: Zarcero, Alfaro Ruiz, in clay-loam, in half-shade on
hill-slope of open forest of upper tropical zone, A. Smith P.2309 (fl., A), edge of wood-
land, A. Smith P. 2392 (fl., A); Guadaloupe de Alfaro Ruiz, A. Smith 4178 (fre Ch):
Zarcero, A. Smith 10068 (fl., fr. A). San José: Vicinity of Santa Maria de Dota,
Standley 42421, 42426 (fl., Ch); forests of El Copey, alt. 1800 m., Feb. 1898, Tonduz
7353 (Herb. Nat. Costa Rica 11676) (fl., Ch, rypE— GH, NY). Cartago: South-
eastern slope of Irazu, Stork 3177 (fr., Ch).
NaTIVE NAMES: “Quizarra,” “Quizarra amarillo,” “Madera buena” (Costa Rica).
The species is distinguished at once by the soft pale ferruginous tomentum
which clothes the young branchlets and leaves and the lower surface of the
mature leaf-blades. The branchlets are conspicuously striate and slender.
The leaf-blades are chartaceous, smooth and shining above, elliptic, occas-
ionally lanceolate-elliptic, cuneate at the base and acuminate to obtuse at
the apex, measuring 5—8.5 cm. in length and 1.3—2.3 cm. in width. The
lateral nerves, of which there are usually 6 pairs, diverge at an angle of
about 35—45°. The venation above is pubescent and conspicuous though
scarcely elevated. Beneath it is elevated and heavily tomentose. The
328 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
short few-flowered tomentose inflorescences, not more than 5 cm. long, are
borne on peduncles up to 2.5 cm. The lobes of the perianth are spreading
and reflexed, about 2.5 (—3) mm. long, rather heavy in texture, elliptic and
obtuse. The two outer series of stamens are about 1.3—2 mm. long, the
ovate obtuse anthers slightly longer than the filaments. The stamens of
the inner series measure about 1.5 mm. long, the oblong anthers equalling
the slender filaments, which bear at the base two sessile glands about one-
third the length of the entire stamen. The glabrous gynaecium is + 2.8
mm. long, the ellipsoid ovary with a style one-third its length topped by a
more or less triangular conspicuous stigma. The fruit is oblong-ellipsoid,
glabrous, 2 cm. long and 1.3 cm. broad, black, with primrose yellow base or
“eve” (according to the collector Stork), the campanulate cupule green or
reddish to scarlet, about 2 mm. long, 6 mm. broad at the flaring undulate
apex, and 1 mm. deep, — the pedicel about 5 mm. long enlarged at the
tip to 3 mm. in diamet
The nearest species is SP. acuminatissima, from Mexico and Guatemala.
The relationship has been discussed under that species.
30. sent Bourgeauviana Mez in ies Bot. Gart. Berlin 5: 194. 1889; Standley in
Con S. Nat. Herb. 23: 294.
sees aes Mez in Jahrb. a: an Berlin 5: 196. 1889.
DistrIBUTION: Southern Mexico peel Guatemala, at an altitude of 1300-3000
m., to Honduras at a lower altitude of 1050
exico: Vera Cruz: Vallée de Mh: April 9, 1866, Bourgeau 2234 (fl., fr.,
isoTyPE of P. Bourgeauviana, Ch, GH). GuaTeEMALA: Quiché: San Miguel Uspantan,
He rs & Lux 3089 (fl, GH). Huehuetenango: Jacaltenango, mountain-slopes,
. Seler 2598 (fl, GH). Alta Verapaz: Mountains east of Tactic, on roa
tbe Standley 71282 (f1., Ch) ; large swamp east of Tactic, in forest, Standley 92314
(fl., Ch), Stevermark 43966 (f1., Ch) ; in mountain-forests near! Laraxquia, von Tuerck-
hein 371 (f1., photo. of rin of P. purpurea, Ch, GH); Coban, von Tuerckheim 1651
Baja Verapaz: Mountain-side nave of divide north of Santa Rosa,
Siindley 69883 (f1., Ch). San aera Rio Vega, near — Rafael and Guatemala-
Mexico boundary, Volcan Tacana, Steyermark 36267 (f1., Ch). El] Progreso: Be-
tween Calera and middle slopes of quebradas of Volcan Sins, Stevermark 43020 ({fr.,
Ch). Zacapa: Upper reaches of Rio Sitio Nuevo, Steyermark 43237 (fr., Ch).
Chiquimula: Middle slopes of Montana Norte to El Jutal, on Cerro Brujo, south-
east of Concepcion de las Minas, Steyermark 31072 (fr., Ch); Volcan Quezaltepeque,
34 nes northeast of Quezaltepeque, Stevermark 31442 (fr., ONDURAS:
Comayagua: Vicinity of Siguatepeque, Standle y 56055 (fl., Ch); on river-banks,
ie Yuncker, Dawson & Youse 5777 (fr.,
This species is a variable one, the branchlets clothed with a ferruginous
subcaudate-acuminate, glabrous to glabrescent and somewhat shining
above, except for the pubescent costa, and softly tomentose beneath. The
blades measure 7-10 (—12) cm. long and 2-3.5 (—5) cm. broad, and are
subtended by pubescent petioles up to 11 mm. long. The costa and lateral
nerves, which number no more than 7 pairs diverging at an angle of 45-35”
from the costa, usually are visible above but very prominently elevated be-
neath. The loose reticulation is obscure above and apparent beneath.
Pubescent glands are borne beneath in the axils of the lateral nerves. The
pubescent to glabrescent few-flowered inflorescences are much shorter than
the leaves, measuring not more than 7 cm. in length, the slender peduncle
1945 | ALLEN, STUDIES IN THE LAURACEAE, VI 329
not more than 5 cm. long. The flowers are purplish on drying, about 4-5
mm. long, borne on pedicels of almost the same length. The spreading or
at times reflexed lobes of the perianth are elliptic, obtuse, pubescent, up to
3 mm. long. The two outer series of stamens vary from + 1.25—2.7 m
long, with anthers that are petaloid, subrhomboid to ovate, obtuse, sub-
sessile or with short filaments. The stamens of the inner series are + 1.25
mm. long, with oblong anthers equalling the filaments and only slightly
exceeding the conspicuous sessile glands at the base. The staminodia vary
from triquetrous to cordate, + 0.8—1.25 mm. long, the stipes equailing
one-half the entire length. The glabrous gynaecium varies in size up to
+ 2.15 mm. long, the ovoid or elliptic ovary being more than twice the
length of the stout short style, which is topped by an often somewhat de-
current triangular conspicuous stigma. The fruits are ellipsoid-subobovate,
apiculate, up to 2 cm. long and 1 cm. broad, green turning purple-black,
seated on a cupule which is rose-red at maturity. The glabrous cupule is
x
The nearest relative of the species is P. Pittieri from Costa Rica and
northwestern Panama. The latter differs from P. Bourgeauviana in the
rough quality of the pubescence on the leaves, which are never more than
9 cm. long, usually around 7 cm., and 2.5—3 cm. broad. The reticulation
of the leaves is far more prominent and the lateral nerves never more than
5 pairs. The flowers are similar to those of P. Bourgeauviana. ‘The fruits
are smaller and frequently borne on a longer pedicel.
31. Phoebe Pittieri Mez in Bot. Jahrb. 30: Beibl. 67: 16. 1901; Standley in Field Mus.
Publ. Bot. 18: 460. 1937; Lundell in Contr. Univ. Mich. Herb. 6: 22. 1941.
DistrRiBuTION: Woods of Costa Rica and Panama, at an altitude of 1050-2200 m.
Costa Rica: Alajuela: Woods, La Palma de San Ramon, Brenes 5334 (483)
(fr., Ch), 5405 (549) (sterile, Ch), 5416 (fl., Ch) ; moist woods on the hills of Santiago
near San Ramon, Brenes 14464 (sterile, GH); Palmira, Alfaro Ruiz, in sub-tropical
zone, in wet loam at edge of swamp, in semi-shade, 4. Smith H.522, H. 679 (fl., Ch).
osé: Forests of El Copey, Tonduz 11893 (fl., photo. of TYPE, GH). PAnaMa:
Chiriqui: Cloud- and rain-forest of Cerro Horqueta, C. & V. W. von Hagen 2031,
2070 (fi., fr.. A, Mo), 2022 (fl, A, Mo).
The branchlets of this species are fulvous- or yellowish-tomentose, the
very young ones somewhat sericeous, becoming fuscous with age, and harsh
to the touch. ‘The leaf-blades rarely exceed 7 cm., occasionally 9, in length
and 2.5—3 cm. in breadth; they are lanceolate or elliptic with cuneate base,
acuminate to obtusely acute apex, and a canaliculate pubescent petiole up
to 10 mm. long. The upper surface is smooth and almost shining, the lower
roughly pubescent. The lateral nerves, diverging from the costa at a
35-45° angle, usually number 5 pairs and are inconspicuous above but ele-
vated beneath, frequently with inconspicuous axillary glands. The loose
reticulations are slightly more prominent beneath than above. The few-
flowered pubescent to glabrescent axillary inflorescence is not more than 5.5
cm, long, the peduncle not more than 2. cm, The white (purplish in dried
state) flowers are somewhat like those of P. Bourgeauviana, being spread-
ing-campanulate, up to 5 mm. long, with slender pedicels to 8 mm. The
elliptic lobes are thick, papillose, up to 3 mm. long and often reflexed in
330 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
anthesis. The two outer series of stamens measure + 1.7 mm. long, the
roundish anthers equalling the filaments. The stamens of the inner series
have oblong anthers equalling the filaments and the large sessile spongy
glands at the base of the latter. The staminodia are + 1 mm. long, ovate,
subcordate, the stipes slightly more than one-half the entire length. The
glabrous gynaecium is + 2.8 mm. long and has an obovoid or subglobose
ovary slightly narrowed at the base and one-third longer than the style,
which bears an inconspicuous discoid stigma. The fruits are greenish to
purple, ellipsoid, glabrous, 12 x 9 cm., and seated . a flaring undulate
fluted cvathiform reddish cupule about 5 mm. long, 9 mm. diam., and 3 mm.
deep. The ar pedicel is upwards of 1 cm. ne er ei to 4
mm. at the a
The ane claties of the species is P. Bourgeauviana, under which
species is a discussion concerning the two entities.
DouBTFUL SPECIES AND VARIETIES OF PHOEBE
Phoebe effusa 8 areolata Meissner in DC. Prodr. 151: 33. 1864.
Mez cites Linden 19, the type, from Jalapa, Mexico, under Nectandra
sanguinea. The type is not available at this time.
Phoebe granatensis 6? Oerstedii Meissner in DC. Prodr. 151: 32. 1864.
Mez cites the type of this, from Volcan Barba, Costa Rica, Oersted 4,
under Phoebe mexicana Meissner. 1 have not seen the type.
Phoebe mayana Lundell in Amer. Midl. Nat. 29: 473. 1943.
I have not been able thus far to match this fruiting specimen with any
material at hand. It is somewhat doubtful whether or not this is the cor-
rect genus. The generic lines of fruiting characters are not vet as clear-cut
as it is to be hoped that future study may render them. For this reason,
it is difficult to determine the genus of a specimen on fruiting characters
alone. At present, since I cannot relate the species to any known entity,
I am not including it in the general treatment or the key. Possibly it may
belong to Beilschmiedia.
SpecIES EXCLUDED FROM PHOEBE
Phoebe Benthamiana Mez = PERSEA PACHYPODA Nees.
ebe Hartwegit Meissner = PERSEA PACHYPODA Nees
Phoebe insularis Meissner = OcOTEA INSULARIS on Mez.
Phoebe pachypoda Mez = Persea PAcHYPODA Nee
3. Ocotea Aublet
— oe Pl. Guian. 2: 780, 370. 1775; Hemsley, Biol. Centr. Am. Bot. 3: 72
; Mez in Jahrb. Bot. ee Berlin 5: 219. 1889.
Oreadaphne Nees, Pl. Laurin. [ Progr. | 1833, in Linnaea 8: 39, 1833, Svst. Laurin.
; Meissner in DC. Prodr. 151: 111. 1864.
Plein Beurling in Vet. Akad. Hand oo 1854: 145. 1856.
Sassafridium Meissner in DC. Prodr. 151: 171.
DistriBuTION: Throughout the American nn cs from Mexico — Central
America, with the greatest number of species occurring in South Americ
Trees or shrubs usually with alternate leaves varying is membra-
naceous to rigidly coriaceous, the blades from 9 cm. to 55 cm. long, and
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 331
equally as variable in shape and pubescence. The reticulation of the leaf-
blades of this genus is an important diagnostic character. Occasionally the
decurrence and recurving of the leaf-bases into the broad petioles is mani-
fest. The inflorescences are usually paniculate, and bear usually in this
area flowers that are perfect. There are three species, as far as can be
ascertained from the material at hand, that have dioecious flowers. In the
perfect flowers the perianth-tube may be conspicuous or almost entirely
absent. The lobes may be fleshy or thin in texture, and are, for the most
part, equal or at most subequal, usually deciduous, although they may per-
sist in some species as remnants crowning the expanded perianth-tube in
fruit, Generally speaking, the stamens of the two outer series bear varl-
ously shaped anthers that are sessile or borne on glabrous or pubescent fila-
ments of varying length. The anthers may be petaloid and papillose,
exhibiting a substantial connective, or they may have no perceptible con-
nective tissue present. Usually the four introrse cells are arranged in two
planes, one above the other. Those of the inner series have longer fila-
ments accordingly, which always bear two sessile or stipitate glands which
vary in size and shape, often so large that they appear confluent. The four
cells of the anthers are sometimes all extrorse; sometimes the two upper are
lateral or lateral-extrorse, the two lower being wholly extrorse or rarely
lateral-extrorse. If staminodia are present they are usually stipitate,
linear-lanceolate, or very occasionally ovate, pubescent or glabrous but
invariably thin in texture. The gynaecium is usually glabrous, the style
rarely being slightly pubescent, and of varying length. The stigma may be
small, inconspicuous, and discoid, or large, conspicuous, peltate or triangular,
and slightly decurrent at the angles. In species with dioecious flowers, the
9 flower shows staminodia replacing the first three series of stamens found
in the perfect flowers. The staminodia may have well developed anthers,
but usually they are smaller, not much broader than the filaments and with
the cells evident though often malformed. The gynaecium develops as in
the perfect flowers. The 4 flowers have well developed stamens, but
either lack the gynaecium entirely or have an aborted small linear struc-
ture, the different parts being indeterminate; or the ovary is as slender as
the style, but topped by an extremely large stigma. The fruit is ellipsoid
or globose, borne in a usually simple-margined cupule, although in a very
few species the margin is inconspicuously double. The cupule may be
flat and disk-like, the margin undulating, or hemispherical and plane, or
variously 6-lobed, the lobes being the remnants of the persistent perianth-
obes.
KEY TO THE SPECIES OF OCOTEA
A. Largest leaf-blades not less than 20 cm. long
Leaf-blades heavily coriaceous and densely and conspicuously reticulate and
shining above.
C. Leaf-blades pubescent beneath.
D. Leaf-blades decurrent at the base, strongly recurved and _ closely
appressed-pubescent ; petioles not canaliculate........ 1. O. stenoneura.
D. Leaf-blades not decurrent at the base nor recurved, but pubescent ;
petioles shallowly canaliculate...............00- eee eed. O. operi
332 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
C. Leaf-blades glabrous beneath and heavily coriaceous........3. O. Seibertii.
Leaf-blades, if densely reticulate, not conspicuously so and not shining above;
texture various.
.
sad
Leaf-blades beneath conspicuously ferruginous- or subferruginous-tomen-
tose; venation conspicuously pubescent; inflorescence densely ferruginous-
or subferruginous-tomentose.
D. Leaf-blades coriaceous; reticulation usually obscured beneath by a fine
; branchlets stout; lateral nerves 8 or 9 pairs. .4. O. palmana.
D. Leaf-blades membranaceous, with long subferruginous pubescence on
venation beneath; reticulation conspicuous; branchlets slender; lateral
SPANO DOS iiiindc ns sissies wxes da xevee$-40sa8e 6a OO. eel,
: bax "hades glabrous or glabrescent ranese not conspicuously ferrugi-
nous- or subferruginous- or — ish-tomento
D. Leaf-blades definitely obova
E. sie
io)
of leaf-blades ae. sometimes very conspicuously so and
CUORVER es sees ei wane BGs 2b 6.6 Se eee bade aeSSd oaas 6. O. Tra.
E. ae a ie leaf-blades never conspicuously oe or recurved.
F. Leaf-blades not more than 9 cm. broa
G. Axillary glands absent or so ane and inconspicuous as to
be unobserved.
H. Branchlets glabrous; cupule of ps — brittle about
the margin; fruit globose, 1 cm. diam..... 7. O. Wedeliana.
H. Aura Shape slemgas pulbesient: ane wood Sige ellip-
| IAM hae obo csedesndaw ee eas subalata.
G. Kciiasy ats conspicuous
H. Cupule very saeoarae lobed: 7 mm. diam.; fruit oblong-
ellipsoid, to 2 cm. long; branchlets frequently dark brown-
ish-pubescent; style twice the length of ovary; leaves not
shining beneath..............0..00.-05. 9. O. eucuneata.
H. Cupule thick, woody, entire, 1.5 cm. diam.; fruit ellipsoid,
at pe 2.5 cm. long; branchlets glabrous, style shorter
vary ; leaves shining beneath. . . verapazensis.
F. Leaf-blades a less than 10 cm. broad nanan 15— 20 cm.).
G. Branchlets sturdy, heavily angled; blades shortly obtusely
acuminate
ct
sip-aigierein eggs Siu gas aa oe wp om 1. O. nicaraguensis.
G. Branchlets terete, only young slightly angled; apex of th
blades rounded or very obtuse or rarely a ca or shortly
AG cicrecty b24 8 hecuwes . O. Standle vi.
D. a blades elliptic or oblong or oblong-elliptic or cancel
eaf-blades not less than 12 cm. broad............ . O. Standlevi.
Saas geeg paw ee0m 13. O. atirrensis.
G. Leaf-blades coriaceous, brownish or castaneous. .14. O. Paulii.
G. Leaf-blades chartaceous, greenish.
H. Leaf-blades smooth on upper surface, eae See eT ee eee
Sale Ateusleraia owe ate dig e's bile dare wk eae 15... 0, ndrodaphne.
H. Leaf-blades with conspicuous reticulation Ppieee ee not
pale green above.................... 10. O. verapazensis.
A. aja leaf-biades not more than 17 cm. long (rarely 20, in O. eucuneata
Leaf-blades variable, even not obovate but elliptic or rece ae or
variations of these shapes, acute or rounded, always chartaceo ma al-
ways finely undulate, aenenee crisped on drying; petioles ce blackish
MTN BOG sia 050s Pew 4 NSs eS he SSee xb agewsd 16. O. veraguensis.
B. Leaf-blades obovate; margin not consistently finely undulate.
C. Leaf-blades with bases decurrent or recurved.
1945]
ALLEN, STUDIES IN THE LAURACEAE, VI 333
D. Leaf-blades membranaceous and with 3-6 pairs of ne ee ellipsoid
densely grayish-pubescent axillary glands............ 17. O. chiapensis.
Leaf-blades not membranaceous; glands, if present, few ie and ver
inconspicu
E. Reaeantion everywhere prominent; leaves shining above, appressed-
sericeous, becoming glaucescent and glabrescent beneath............
pea hanetetue notte sen CL) ac shGs® ‘ahi wih meee Stee Keane 18. O. Austinit.
Ee oyun not prominent; leaves not shining above nor ferrugi-
us-pubescent beneath.
F. Leaves sessile, the nae broadly decurrent and recurved; cupule
entire, pubescent; axillary glands not nana fap age appar-
ent petiole 2-3 cm. Gia, lt ca Re Reta od eee . Endresiana.
F. Leaves shortly petiolate, the blades narrowly ee and re-
curved; pubescent axillary glands not narrowly fina Pay
6-lobed; apparent petiole sie more than 1.5 cm. (rarely 2.5 cm.).
eee i ee ee ay 20:50: Foeuiaie
F. Leaves shortly petiolate, the blades narrowly decurrent and re-
curved; cupule entire; pubescent axillary glands when present
eh narrowly ellipsoid; apparent ge to: 2.5.cm.
YQ esa ares wks # Bia BX ike nade a eee os yen pe . O. Skutchii.
. Leaf-blades with bases not decurrent or recurved.
D. Largest leaf-blades not less than 15 cm. long............ ata
E. Leaf-blades coriaceous; reticulation minute but conspicuous; nerva-
tion very prominently elevated beneath and ae yellow........
Sec ease ere napa here een ows iia ope ayo al a creirs eas ate ma Op dallas
E. Leaf-blades not coriaceous; reticulation not ee nervatio
elevated but not strikingly iim eo ben
F. Lateral nerves up to 7 pai ee eee buff-gray ; cupule
thick, woody, the margin swdniate 13 mm. diam.; flowers dioe-
GIOUSIN( Paar aie ete etonctte starserenaeis sa atet nace 3. .O. subsericea
F. Lateral nerves 7-12 pairs; branchlets fuscous; cupule shallow,
cyathiform, lobed, of thinnish texture; flowers monoecious......
Edi sie hie Hh Te Oe OM, 8's < Sete waa 9. O. eucuneata.
B. Leaf-blades not obovate
oF
C
Leaf-blades not more pee 2.5 cm. broad.
Dp: Sener and lateral nerves prominent throughout; apex of blades
isi EA EE ke cb 5.8 24. O. Klotzschiana.
D. cite and lateral nerves very obscure; apex caudate: eee
ee ee SOO Ce Oe oe EEO ITE 25.0. vac,
Leaf-blades not less than 3.5 cm. broad (usually broader).
D. Leaf-blades definitely oblong and rigidly coriaceous; inflorescence up to
16 cm. or more long.
E. Leaf-blades not more than 11 cm. a densely ferruginous-sericeous
beneath; inflorescence to 25 cm. long............... 18. ustinii.
E. Leaf-blades not less mn 12em: ee glabrous opeuuhauts inflo-
rescence up to.16°¢m. long cms, oc sees vse siete sg ovate ace eos 4. O. Paulii
D. Leaf-blades not oblong, or, if oblong, oo, not rigidly coriaceous;
inflorescence usually not more than long
E. Leaf-blades oblong or oblong-elliptic, hen eous or chartaceous
or subcoriaceous; apex always caudate-acuminate or abruptly ob-
tusely acuminate.
F. Flowers perfect; inflorescence few- to many-flowered..........
ee ee eco ren ee ee 26:0: randy anete
F. Flowers dioecious; inflorescences usually many and many-flowere
G. Fruit 1.5 cm. long; cyathiform cupule covering 4 its length. .
5h ahs Seg cas or asa ara cas alo rela aha ee Tt teas rams, eel . O. cernua,
334. JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
G. ng 2. : cm. long; disk-like cupule with undulating margin
c ANS TEU smo eosin Go one courte oes s 28. O. tenera.
Leaf- aie elliptic or lanceolate-elliptic or oblong-elliptic, never
membranaceous; apex abruptly acuminate to obtuse but not usually
caudate.
Branchlets at the apex ferruginous-tomentellous, becoming
brownish and glabrous; buds ferruginous- or pom wish-tomen-
CNDUS ssid ano cease ate caskets eee O. paradoxa.
Branchlets at the apex not ferruginous- Sas apie buds not
ferruginous- or yellowish-tomentellous.
Leaf-blades subcoriaceous, shining, paler ae everywhere
prominently reticulate...................5. . O. Matudai.
G. Leaf-blades not subcoriaceous or shining or ene beneath,
the reticulation various.
. Lateral nerves 4-6 pairs; inflorescence ia cm. long......
ae ie err eee ee Pee O. Meziana.
H. Lateral nerves 8-10 pairs; inflorescence pele 12-15 cm.
oI
ie
eo)
ong.
I. Flowers dioecious; branchlets blackish...............
SS Sieh wanes wa SS dies Sank atee les atetaee data.
I. Flowers perfect; branchlets greenish. 33. 0. phere
1. Ocotea stenoneura Mez & Pittier in Bull. Herb. Boiss. I]. 3: 233. 1903.
DistripuTION: Known only from the type-locality and adjoining areas in Costa
Rica, 650-975 m. altitude, in forests.
Rica: San José: at near Las Vueltas, alt. 650-700 m., Feb. 1899,
Tonduz 13377 (fr., photo. of LEcroTyPE of O. stenoneura, Ch, GH); forests of Las
Vueltas, Tucurrique, Tonduz oF (fr., GH); forest in vicinity of El General, Skutch
3014 (f1., A, NY)
NaTIVE NAME: “Ira Zopilote” (Costa Rica).
This species has stout conspicuously angled densely ferruginous-tomen-
tose branchlets bearing leaves densely spiraled near the apex. The unusual
petioles are actually very short and stout, less than 5 mm. long, but the
conspicuously decurrent and recurved leaf-base forms an apparent petiole
that is up to 2.5 (-3) cm. long, 6-7 mm. broad and closely and minutely
appressed-ferruginous-tomentose throughout. The leaf-blades are cori-
aceous, obovate to elliptic, with the base cuneate (due to the recurved lower
portion) and the apex shortly, abruptly, but very sharply acuminate, meas-
uring 18—24 cm. in length and up to 9cm. in breadth. The upper surface is
shining and conspicuous for its pattern of regular venation and the minute
conspicuous reticulation. e lower surface is glaucous and densely
clothed with a soft fine ferruginous pubescence. The costa above is im-
pressed and glabrous except for the lowermost portion; beneath it is thickly
and prominently elevated and pubescent. The lateral nerves, of which
there are 10-14 pairs, ascend without curving to the margin, and are evenly
spaced and almost parallel with each other. They diverge from the costa
at an angle of about 35°. The inflorescence is robust, densely ferruginous-
tomentose, up to 25 cm. long, heavily flowered, the flowers densely tomen-
tose throughout, practically sessile, campanulate, 5—6 mm. long, the fleshy
and hairy perianth-lobes being up to 2.15 mm. long. The tube is very hairy
within. ‘The stamens of the two outer series are up to + 1.25 mm. long,
with the roundish anthers nearly equalling the stocky pubescent filaments.
Those of the inner series are squarish, the filaments bearing laterally two
round conspicuous basal glands that are almost equal to the anthers in size.
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 335
The glabrous gynaecium is about + 2.8 mm. long, the ellipsoid ovary nearly
twice the length of the style, bearing an inconspicuous stigma. The stout
infructescence may measure up to 25 cm. in length, still densely ferru-
ginous-tomentose, according to the authors, but only grayish-pubescent and
not as thickly so as is the case in the flowering stage. The fruit is sub-
globose, apiculate, roughish and minutely tuberculate near the apex, up to
2 cm, in diameter, and supported by a flat disk-like cupule about 12 mm.
in diameter, the margin undulate and (according to Mez and Pittier) fre-
quently bearing the remnants of the perianth-lobes. The gynophore is per-
ceptible in the bottom of the empty cupule, although it is less than 1 mm.
high. The stout woody pedicel is 5 mm. or more long and 3.5 mm. in
diameter.
The relationship of this species will be discussed under O. Seibertii and
also under O. Cooperi. In connection with the latter species, I discuss my
selection of Tonduz 13377 as the lectotype of O. stenoneura.
2. Ocotea Cooperi, sp. nov.
Ocotea stenoneura Mez & Pittier in Bull. Herb. Boiss. II. 3: 233. 1903, quoad Cooper
10217, excl. Tonduz 13377; sensu Standley in Field Mus. Publ. Bot. 18: 456. 1937.
Arbor(?), ramulis valde angulatis dense ferrugineo-tomentosis deinde
griseo-tomentosis sulcatis. Folia alternata, petiolis robustis pubescentibus
leviter canaliculatis, 1.5 mm. longis et 3 mm. latis, laminis supra glabris
basi costae excepta, lucidis, subtus glaucescentibus molliter tomentosis,
membranaceis, in sicco supra brunneo-olivaceis, oblongis, (15—) 35 cm
longis et (5—) 9.5 cm. latis, basi rotundatis vel obtusis, apice acuminatis,
penninerviis, costa supra impressa basi pubescente subtus robusta valde
elevata utrinque pubescente, nervis 10—15-paribus supra subobscuris subtus
satis elevatis pubescentibus angulo 45—60° divergentibus, rete venularum
utrinque prominulo, Inflorescentia axillaris et subterminalis, paniculata,
foliis apicalibus deciduis, ad 20 cm. longis, ferrugineo-tomentosa, mu i-
flora, pedunculo ad 7 cm. longo ferrugineo-tomentoso. Flores ad 5 mm.
longi, pedicellis 2-3 mm. longis perianthii tubo urceolato intus glutinoso,
lobis late ovatis vel subtriangularibus subexpansis crassis vel sublignosis
+ 1.7 mm. longis; staminibus ser. I & II late spathulatis ad + .6 mm.
longis antheris oblongo-truncatis filamentis aequalibus, ser. III + .8 mm.
longis conspicue biglandulosis, glandulis sessilibus et antheris filamentis
aequalibus; staminodiis lanceolatis gracilibus pubescentibus + . m.
longis; gynaecio glabro + 2.5 mm. longo, ovario conspicue stipitato, late
ovoideo gynaecii dimidium longitudine parum excedente, et stylo robusto
duplo longiore, stigmate triangulari subdiscoideo conspicuo styli lateribus
decurrente. Fructus ellipsoideus, minute apiculatis, 3.5 cm. longus et 1.8
cm. latus, cupula lignosa subhemisphaerica rugosula ad 1.5 cm. longa, 2 cm.
diam., et 11 mm. alta, margine suberosa tenui subtentus, pedicello crasso
ad 1 cm. longo apice 5 mm. expanso.
DistriBpuTION: Costa Rica and adjacent Panama at 675-900 m. altitude.
Costa Rica: Limon?: Santa Clara, J. J. Cooper 10217 (fr., isosyntype of
Ocotea stenoneura, US). San José: El General, Skutch 4757 (fl., A, Ch, NY)
naMaA: Bocas del Toro: Changuinola Valley, G. P. Cooper & G. M. Slater 96
(¥10277) (fl., Ch, type — GH, Y); Cricamola, region of Almirante, G. P. Cooper 498
«J
NATIVE NAMES: “Quizarra amarillo” (Costa Rica) ; “Sweetwood,” “Vaya” (Panama).
336 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
The urceolate tube and spatulate anthers of the rather lignified and often
exceedingly large inflorescence distinguish this species from others known
from this area. It is of course most nearly related to O. stenoneura Mez &
Pittier. Since their description agrees with Tonduz 13377 rather than with
the other syntype, Cooper 10217, the former was retained as the lectotype
of O. stenoneura, and the latter placed with Costa Rican and Panamanian
specimens with which it seems to belong. The long apparent petioles with
the blades decurrent for almost their entire length, and conspicuously re-
curved, separate O. stenoneura from O. Cooperi, as well as the smaller
globose fruit subtended by the shallow disk-like undulate-margined cupule.
3. Ocotea Seibertii, sp. nov.
Arbor 22.5-30 m. alta, ramulis griseis mox griseo-rubescenti-maculatis
angulatis mox striatis vel sulcatis juventute pubescentibus mox glabrecen-
tibus vel glabris. Folia alternata, glabra, petiolis + gracilibus vel robustis,
glabris, canaliculatis, ad 12 (— 17) mm, longis et 2 (—2.5) mm. latis, lami-
nis utrinque glabris interdum basi subtus costa nervisque exceptis, coriaceis,
in sicco iar aacemingne: supra valde subtus haud nitidis, ellipticis, 12- 15
(—21) m. longis et 4.5— 6 (~9) cm. latis, basi cuneatis interdum leviter
obliquis apice leviter obtuso-acuminatis, margine recurvatis, penninerviis,
costa satis crassa supra leviter subtus valde elevata rubescente vel flaves-
cente, nervis 6 (—9)-paribus supra haud subtus conspicuiore elevatis an-
gulo 35-45° divergentibus, rete venularum juventute utrinque satis
conspicuo mox supra laxe subtus valde minute conspicuo. Inflorescentia
axillaris, paniculata, ad 13 cm. longa, juventute pubescens mox glabrescens,
rubescens, longipedunculata, pedunculo ad 6 cm. longo. Flores ad 3 mm
Pade pedicellis ad 3 mm. longis, gracilibus, perianthio vadose infundibuli-
for obis + oblongis, subacutis, crassis, extus pubescentibus intus dense
papiloo. -pubescentibus, + 2.8 mm. longis, staminibus ser. I & IT +
1.25 mm. longis, riers oblongo- -subrectangularibus filamento duplo lon-
pee ies ser. II 1.7 mm. longis, conspicue biglandulosis, glandulis
staminibus peries tertiam aequantibus; epg aioe subsessilibus -+
mm. longis ovatis; gynaecio glabro + 2.15 m m. longo, ovario subgloboso
vel obovoideo, stylo aequali, stigmate discoideo conspicuo. Fructus viridis,
fide coll. globosus vel leviter obovoideus, a egos (hip stoner 2.3 X 2cm.,,
cupula subcampanulata rugosula glabra ‘rubra, 5 mm. longa, 3 mm. diam., et
3 mm. alta, margine undulata, gynophorio bene ae ad 2 mm. longo
et 6 mm. diam. subtentus, pedicello incrassato glabro, 4-8 mm. longo.
DistRIBUTION: seihidciotia forests of Costa Rica and Panama.
Costa Rica: Alajuela: Lecheria on Poas, alt. 2285 m., July 30, 1932, Stork 3377
(fl., rype, Ch) (tree ae 5-30 m.; felled for lumber) ; forests of El Copey, Tonduz 11939
(fr.. GH). Panama: Chiriqui: Valley of the Upper Rio Chiriqui Viejo of Monte
Lirio, R. J. Seibert 308 (fr., A, Ch, Mo, NY); Chiriqui Viejo Valley, G. White 96 (ir.,
Ch, GH 0).
ious NAMES: “Quisarra,” “Quizarra” (Costa Rica) ; “Sigua amarilla” (Panama).
The nearest affinities of O. Seibertii seem to be O. stenoneura and O.
Cooperi, both of which are easily separable because of the tomentum on the
lower leaf-surface. The fruit of O. Cooperi is ellipsoid and borne in deep
subhemispherical cupules. That of O. stenoneura is globose and borne on
more disk-like cupules, than that of O. Seibertii.
g
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 337
4. Ocotea oe Mez & J. D. Smith in Bot. Gaz. 33: 258. 1902; Standley in Field
Mus . Bot. 18: 455. 1937
DIstRIBUTION: Costa Rica and Peas at about 1500 m. altitude.
Costa Rica: Alajuela: Los Angeles and La Paz de a a Brenes 6092
(fl., Ch). San José: In the forests of La Palma, alt. 1500 m., Nov. 1898, Tonduz
7374 cand Herb. Nat. Costa Rica) (fl., isotype, GH, NY) ; Platanares nel W.
W.& H. E. Rowlee 210 (sterile, NY); La Palma, W. W. & H. E. Rowlee 233 (fl., Ch).
Lee Bocas del Toro: Fish Creek Mts., vicinity [i Chiriqut Lagoon, von
Wedel 2264 (fr., A).
NATIVE NAME: “Ira mangle” (Costa Rica).
The stout‘branchlets of this species are angled and cae shortly, and
minutely ferruginous-tomentose. The leaves are borne on thick petioles
that are also pubescent, striate, and up to 3 cm. long ae 4 mm. broad.
The blades are rigidly coriaceous, obovate, cuneate at the base, rounded,
and very shortly, abruptly, and obtusely acuminate to acutely acuminate
at the apex, measuring up to 25 cm. long and 14 cm. broad. The blades
are glabrous above, except for the venation, sg beneath are covered with
a dense fine close ferruginous tomentum. e broad costa is scarcely
elevated above, though prominently so seer 4 The lateral nerves, of
which there are 8-10 pairs, are slender-and delicately and slightly elevated
above and conspicuously so beneath and diverge at an angle of about 55
The inflorescence is stout, densely ferruginous-tomentose, axillary, panicu-
late, rather few-flowered, about 8 cm. long, the peduncle not more than
2 cm. long. The flowers are large, 5-6 mm. long and about 12 mm. in
diameter, and densely tomentose, the ovate sabes almost leathery, + 3.8
mm. lon ‘he stamens of the outer series have short filaments and the
anthers are ovate or rounded; those of the inner series are longer, with the
filaments nearly equalling the oblong rounded anthers and completely
covered by the large sessile roundish basal glands equalling the anthers in
size. The glabrous gynaecium consists of an ellipsoid ovary topped by a
rather stout style of nearly equal length bearing a capitate stigma. The
fruiting specimen from Panama has branchlets that are less ferruginous-
than tawny-brown-tomentose, and thicker shorter petioles subtending
blades that are more rounded at the base and less pubescent. ‘The fruit
is green, ellipsoid, oblong, up to 3.5 cm. long and 1.6 cm. broad, subtended
by a pink cupule that is subcampanulate = very thin- margined, up to
6 mm. long, about 15 mm. in diameter, and about 5 mm. deep, the sup-
porting pedicel nearly 2.5 cm. nea and expanded to nearly 1 cm. in diam-
eter at the tip.
The species is related to O. mollifolia, under which it is further discussed.
S Ney pace ‘ae & yee in ee Herb. Boiss. II. 3: 233. 1903; Standley in
ield Mus. Publ. . 18: 455
nmin Known only oe the type-locality.
jn: Hacienda Victoria, forests of the plains of Zent, alt. 200 m.,
Feb., 1900, Pittier 16031 (fl., fr., TYPE not seen), 16030 (fl., GH).
This species has slender striate terete branchlets densely clothed with a
fine close short ferruginous tomentum, which persists on the short rather
thick petioles measuring not more than 7 mm. in length. The chartaceous
blades are subelliptic or usually obovate-elliptic, with the base cuneate an
the apex abruptly short- or long-caudate-acuminate, measuring up to 24
cm. long and to 9 cm. broad. They are glabrous above except for the
338 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
midrib and covered beneath with a soft pubescence which is conspicuously
ferruginous on the venation. The costa and lateral nerves, of which there
are about 10 pairs, diverging at an angle of 35—45°, are very slightly ele-
vated above and prominently so beneath. The reticulation, visible above,
is very conspicuous beneath. The very slender inflorescence is axillary
or subterminal, paniculate, few-flowered, shortly densely ferruginous-
pubescent throughout, up to 9 cm. long, the weak peduncle not more than
5 cm. long at most. The flowers are about 4 mm. long, subtended by a
slender pedicel of equal length, both densely ferruginous-pubescent. The
equal perianth-lobes (subequal according to Mez) are broadly ovate, acute
(almost round according to Mez), thick, fleshy and slightly papillose with-
in, nearly 3 mm. long. The stamens of the two outer series are + 1.25 mm.
long, the ovate anthers papillose at their round or obtuse tips, and only
slightly longer than the broad pubescent esa which gradually expands
toward its Sapo with the anther. ‘Those of the inner series are + 1.7
mm. long, the anthers oblong and equal to the pubescent filaments, which
are covered one-half their length by large subsessile glands that are almost
confluent. The gynaecium is + 2.15 mm. long, the ellipsoid glabrous
ovary slightly longer than the tapering pubescent style, which bears a
conspicuous distinctly capitate stigma. e fruit, according to Mez, is
large, ellipsoid, up to 3.5 cm. long, and about 1.5 cm. in diameter
Although the type is not available for study, the number collected by
Pittier presumably at the same time in essentials agrees precisely with the
tvpe. Mez mentions the shorter flower and the unequal os of the
perianth, both characters which may vary in the same inflorescence. e
mentions O. Salvini and O. palmana as being near O. mollifolia, hut differ-
ing because of the strongly pilose style. The former has been transferred
by Lundell to the genus Phoebe. ‘The latter is abies a far more robust
species in foliage and inflorescence-characters
6. ~~ Ira Mez & Pittier ex ro in rae Herb. Boiss. II. 3: 232. 1903; Standley in
eld Mus. Publ. Bot. 18: 193
pismo Costa Rica wes vee Panama, in the lowlands, or at not more
than 20 e:
oo. ane Limon?: Llanuras de Santa Clara, alt. 200 m., June, 1899, Pittier
7608 (13999 Herb. Nat. Costa Rica) (fl., Lecrorypr, US). Panama: Bocas del
Toro: Region of Almir ae Cricamola Valley, G. P. Cooper 532 (fr., Ch, NY);
Water Valley, von Wedel 7 se , A); vicinity of Chiriqui Lagoon, von Wedel 1382
(fl, A). Chiriqui: in ands, Cooper a Slater 218 (¥ 10571) (sterile, Y);
Progreso, Cooper & Slater Pit ye 10660) (f1., Y).
NATIVE NAMES: “Ira” (Costa Rica); “ epee (Panama).
This species has branchlets that early lose their closely appressed brown-
ish almost sericeous pubescence, becoming glabrous, angled, and darkish.
The leaves are borne on petioles that are strongly or in some cases slightly
winged, the blade being decurrent and recurved for nearly 4 cm.,, forming
an apparent petiole; or the distinctly winged petiole may measure up to
1 cm. in length. The blades are eae to rigid, glabrous to gla-
brescent, with frequently inconspicuous axillary glands, obovate, attenu-
ately cuneate at the base and recurved, the apex abruptly and obtusely
cuneate. They are u re 25 cm. long and 9.5 cm. broad. The costa is
broad and conspicuous above, although slightly impressed, and prominently
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 339
elevated beneath. The slender lateral nerves, of which there are 9-12
pairs, are very slightly elevated above and more so beneath, diverging at
an angle of about 35—45°, curving toward the marginal region, the reticula-
tion being inconspicuous ‘above and conspicuous and frequently pubescent
beneath. The inflorescence is axillary and subterminal, paniculate, many-
flowered, brownish-pubescent, becoming glabrous, up to 15 cm. long, the
stout peduncle up to 7 cm. long. The flowers are small, + 2.15 mm. long,
pubescent without, the slende : ‘pubescent pedicel about ae same length;
the rather thick papillose ovate acute lobes are + 1.4 mm. long. The
stamens of the two outer series are + 1.25 mm. long, the ovate obtuse
meee sander! longer than the slender filaments. Those of the inner
eries are + 1.7 mm. long, the subrectangular truncate anthers not quite
Ponalling 4 the pubescent filaments, with two subglobose subsessile glands
about half the length of the filaments. The glabrous gynaecium measures
up to + 2.4 mm., the subellipsoid ovary slightly longer than the slender
style ea a subtriangular subdiscoid stigma. Mature fruit of the species
is unkno
Mez & Pittier cite two syntvpes, the number cited above, Pittier 7608
(73999), and the number 10415. The latter certainly corresponds not to
the description of O. /ra, but to that of O. cuneata (O. Tonduzii). It does
not match the only other number cited. Mez himself says that no. 10415
differs from O. cuneata (O. Tonduzii) and is more nearly related to O.
pedalifolia, O. pentagona, and O. insularis, but differs from them in shape
of leaves, the inflorescence, and smaller flowers. The species seems to me
to be closely allied with the group of specimens of which O. Tonduzii is a
member, because of the decurrent leaf-bases and the shape of the blades,
although it does differ presumably in floral characters and the shape of the
cupule. The leaves of the O. Tonduzii group (according to the original
description) are on the whole much smaller than those of O. /ra.
7. Ocotea a, sp. nov.
Arbor 3-12 m. alta, ramulis minute et inconspicue pubescentibus mox
glabris, Pte minusve angulatis vel alatis, brunneis, rubescentibus vel griseis.
caudato-acuminatis, ad 26 cm. longis et 9.5 cm. latis, penninerviis, costa
supra impressa subtus elevata, nervis lateralibus 8 vel 9 paribus, supra
leviter subtus prominenter elevatis, angulo 35—45° divergentibus, rete
venularum supra obscuro subtus satis prominulo. Inflorescentia axillaris
vel subterminalis paniculata ad 12 cm. longa, glabrescens, pauciflora, pedun-
culo gracili ad 7.5 cm. longo. Flores fulvo-pubescentes, ad 3.5 mm. longi,
pedicellis gracilibus ad 4.25 mm. longis, perianthii tubo conspicuo, lobis
late ovatis acutis crassis + 2.15 mm. longis; staminibus ser. I & II + 1.25
mm. longis antheris ovatis obtusis 4-plo filamentis robustis pubescentibus
long ee ser. III + 1.5 mm. longis antheris basi pubescentibus anguste
ovatis filamentis Giglanduloce duplo longioribus, glandulis reniformibus
conspicuis sessilibus filamentis aequalibus; staminodiis nullis; gynaecio
.7 mm. longo, ovario subgloboso stylo gracili aequali, stigmate
conspicuo discoideo. Fructus globosus, glaber, viridis, ad 1 cm. diam.,
340 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXVI
cupula vadosa undulata rubra ad 2 mm. longa et alta et 8 mm. diam. sub-
tentus, pedicello rubro glabrescente ad 8 mm. longo et 2 mm. diam. apice
expanso.
DistriputioN: Known only from Bocas del Toro, northern Panama, at 375 m.
altitude
— Bocas del Toro: Without a Wedel 388 (fr., A), G. P. Cooper
399 (fl., fr., Ch, type — GH) (tree 12 m., diam. cm., with long — very slender
bole; nae reddish, fruit green with white aad pulp inside), ie Bi Ch, NY);
Buena Vista Camp, on Chiriqui Trail, Cooper 603 (¥ 12236) (fr., Ch,
NaTIVE NAME: “Sigua” (Panama).
This species is not, in foliage and floral characters, so very different from
O. nicaraguensis, the principal difference being in the texture and size of the
leaf-blades.
8. Ocotea subalata Lundell in Lloydia 4: 48. 1941.
DistrrpuTIoN: Known only from the type-locality.
Chia Volcan rage on north side, alt. 2100 m., April 2, 1939,
Matuda 2057 (fl., ri isoTyPE, A, Ch,
This species, known only con te type-material, has branchlets gira oe
brownish-hirsute, becoming glabrescent, angled (subalate), and stri
The leaves have petioles that are hirsute, canaliculate, and up to 2 cm. ‘sn
The blades are glabrous above and glabrescent beneath, chartaceous, nar-
rowly obovate or broadly oblanceolate, the base cuneate or almost obtuse,
the apex usually abruptly and obtusely acuminate, 8-21 cm. long and 4—8.3
cm. broad. The broad costa is only slightly depressed above and promi-
nently elevated beneath; the lateral nerves, of which there are 6—9 pairs
are obscure above and also prominently elevated beneath, the reticulation
being minute and conspicuous throughout. The inflorescence is axillary,
few-flowered, paniculate, pubescent to glabrescent, up to 12 (—?) cm. long,
the blackish ‘glabrescent peduncle measuring up to 7 cm. in length. The
flowers are brownish-hirsute, about 2.5 mm. long, and borne on pedicels to
4 mm. long. The specimens at hand bear flowers that are too little de-
veloped for satisfactory dissection, but the author describes a short
perianth-tube, ovate obtuse lobes, filaments about half as long as the ovate
obtuse anthers and sparsely appressed-puberulent, the filaments of the
cence expands in the fruiting stage to a rather robust structure up to 25 cm.
long with a stout peduncle. The shining black ellipsoid fruits measure up
to 2.5 cm. long and 1.7 mm. in diameter, and are oad by an exceed-
ingly shallow undulating woody cupule 3-4 mm. long and deep and about
1 cm. in diameter. The sturdy pedicel is glabrous, aa tedty rugose,
and up to 2.5 cm. long, expanded equally throughout to a diameter of
The nearest relative of O. subalata seems to be O. verapazensis, but the
latter is easily distinguished by the differences in fruit-characters, particu-
larly the deeper and even more woody cupule, and the presence of conspicu-
ous glands. There is a superficial resemblance to O. Wedeliana, but again
the small globose fruit of the latter, supported by a small unlignified cupule,
unquestionably separate the two entities.
1945 | ALLEN, STUDIES IN THE LAURACEAE, VI 341
9. Ocotea eucuneata Lundell in Contr. Univ. Mich. Herb. 6: 16. 1941.
DistriBUuTION: From British Honduras eine high ridges south to Guatemala, along
rivers and in mountains up to 1200-2000 m. alti
GUATEMALA: Huehuetenango: Cerro ate between Finca San Rafael and
Ixcan, ee de los Cuchumatanes, Steyermark 49183 (f1., Ch). Izabal: Along Rio
Tameja, Steyermark 41769 (fr., Ch), 41797 (fr., Ch). British Honpuras: Stann
Creek: Middlesex, in high ridge, Oct. 27, 1939, Gentle 3068 (fl., isorypE, A, NY)
(a large tree about 75 cm. in diameter; flowers creamis
NaTIVE NAME: ‘Timbersweet”’ (British Honduras).
A species with slender branchlets minutely, shortly, and inconspicuously
pubescent, becoming glabrous and angled. The petioles also are slender
and minutely pubescent, earns up to 1.5 cm. long at most. The
blades are very thinly m mbranaceous, obovate, the base attenuately
cuneate, the apex abruptly eon. up to 16 (occasionally 23) cm. long
and 5 (occasionally 8) cm. broad, glabrescent on both surfaces. The costa
and lateral nerves, of which there are 8 or 9 pairs diverging at an angle of
about 35°, with inconspicuous pubescent axillary glands, are very slightly
elevated above and very prominently so beneath. The inflorescence is
axillary-subterminal, paniculate, grayish-pubescent to glabrous, up to 9 cm.
long, the comparatively stout peduncle no more than 2.5 cm. long. The
pubescent flowers are small, not more than 2.5 mm. long, the slender pedicel
3 mm. or less long. The perianth-tube is conspicuous, the ovate acutish
lobes thinnish, up to 1.8 mm. long. The stamens of the two outer series
are almost 1 mm. long, the ovate roundish anthers slightly longer than the
pubescent filaments. Those of the inner series are somewhat longer, the
anthers more oblong, the filaments bearing two subreniform conspicuous
stipitate glands at the base. The glabrous gynaecium is + 2.15 mm. long,
the subglobose ovary surmounted by a slender style twice its length and
bearing a conspicuous capitate stigma at its apex. The infructescence
often retains the pubescence of the early stage. The fruit is oblong, slender,
dull green, according to the collector, up to 2 cm. long an and 6-10 mm. broad,
subtended by a shallow lobed brittle cupule that is dull red, measuring up
to 2 mm. long and deep and 8-9 mm. in diameter. The pedicel is enlarged
to nearly 5 mm. long and is expanded to about 3 mm. in diameter at the
apex. The Sranciilets of the fruiting oa are much more densely
pubescent than those of the flowering materi
The species is most nearly related to O. Beis It may be distin-
guished from this species by its leaves reaching a length of not less than
15 cm., as opposed to 12 cm. in O. subsericea, and by its much smaller
thinner cupule.
10. Ocotea verapazensis Standley & Steyermark in Field Mus. Publ. Bot. 23: 114. 1944.
DistRiBUTION: Known only from Guatemala, usually in wet forests at an altitude
of 50 m. along river- oe in Izabal, and at 300-2000 m. in the mountains of Alta
Verapaz and San Marc
GUATEMALA: rae se apaz: Dense wet forest in mountains east of Tactic, on
road to Tamahu, Standley 71163 (fl., Ch), April 9, 1939, 71421 (fl., fr., Type, Ch) (tree
m.; flowers green; cupule red). Iza ae lo ie Rio Tameja, Sieveemark 41789
(fr, Ch). San Marcos: Between Finca E] Porvenir and Loma Corona on forested
slopes 9 miles northwest of El Porvenir, uhectieas: slopes of Volcan Tajumulco,
Steyermark 37752 (fl., Ch).
The branchlets of this species are distinctly angled and in the early stages
342 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
are glabrescent and dark brown, shortly becoming glabrous, mottled dark
brown, and pale buff-brown. The petioles are, on the mature a
yellowish brown, canaliculate, and sturdy, measuring up to 1.5 cm. lon
The chartaceous blades are opaque above, shining beneath, oblong- “elliptic
or obovate-lanceolate, the base cuneate and often very narrowly and ob-
scurely decurrent and recurved for half a millimeter’s length, the apex
abruptly and usually shortly obtusely acuminate, 14-24 (—27) cm. long
and 4.5—7 (—8) cm. broad. The costa is often yellowish or reddish, slightly
elevated but conspicuous above and more prominently so beneath,
lateral nerves, numbering up to 9 at most, are inconspicuous above and
prominently elevated beneath, diverging arcuately at an angle of about
35—45°, sometimes with rather inconspicuous axillary glands. The minute
reHeitish is obscure above but very conspicuous beneath. The rather
weak axillary or subterminal inflorescence is very slender, rather few- to
many-flowered, up to 15 cm. long, glabrous for the most part, with a
peduncle that is not more than 5 cm. long, usually less. The glabrous
flowers are up to 2.5 mm. long, the slender pedicel less than 3.5 mm. page:
the perianth very narrowly campanulate, the lobes elliptic, acutish,
mm. long, rather thick. The stamens of the two outer series are + 0. ry mm.
long, the anthers ovate, nearly twice the length of the filaments, with the
short connectives obtusely but shortly acuminate. Those of the inner
series are + ong, the anthers ovate with short obtuse connectives,
the filaments bearing two conspicuous basal —- glands. The slender
linear attenuate pubescent staminodia are + 0.55 mm. long. The glabrous
gynaecium is about 1.4 mm. long, the ellipsoid ovary ie twice the length
of the slender style bearing an shine dour stigma. The fruit is shining,
ellipsoid, up to 2.5 cm. long 1.8 mm. broad, subtended by a red woody
fluted cupule that is 6 mm. pa up to 1. ie cm. in diameter, and about 4 mm.
deep. The pedicel is ae grooved, about 1.5 cm. long, and ex-
panded to 8 mm. in diameter at the apex
The species in foral prairie recalls O. laetivirens, but has a much more
robust habit and coarser bark and infructescences.
11. Ocotea nicaraguensis Mez in Jahrb. Bot. Gart. eoeeal - 238. 1889,
Ocotea pentagona Mez in Bot. phere 30: Beibl. 67: Ol.
Ocotea pedalifolia Mez in op.
DistripuTion: Costa Rica and ies Nicaragua, at 200-915 m. altitude.
Juan del Norte: Near San Juan, along the southern bounda-
ries, Friedrichsthal 627 (fr., photo. of Type of Ocotea nicaraguensis, Ch, NY). Costa
Rica: Alajuela: Banks of the Rio Naranjo, near San Mateo, alt. 200-250 m.,
March, 1893, Tonduz 7613 (fr., isosyntype of Ocotea pentagona, US); margins of “La
oe. Machu uca, in “La Calera” de San Ramon, Brenes 22644 (7) (fl., Ch).
mon: Forests of Shiroes, Talamanca, alt. 100 m., Feb., 1895, Pittier & Tonduz
9172, 9179 (f1., fr., isosyntypes of Ocotea pedalifolia, US). San José: Vicinity of El
“sie Turrialba, Tonduz 7106, 8362 (fl., syntypes of O. pentagona aot sane) 3 bank of
Rio Tuis, Pittter 11347 (fl., GH).
The present species has usually very characteristically angled rather
stout fee branchlets that are early minutely ferruginous-tomentellous,
petioles which are tomentose on the ventral surface and measure up to 1.5
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 343
m. long. The blades are heavily coriaceous, more or less opaque above,
slightly darker beneath, obovate-elliptic or oblanceolate, the base obtuse,
subrounded, or cuneate, ‘often attenuately so, the apex abruptly and shortly
acutely or obtusely acuminate. They measure up to 55 cm. long, although
they are usually about 30 cm., and Riad from 6-15 cm. in breadt The
costa is exceedingly broad (u up to 5 mm.), usually only slightly clevated
above and more prominently so fe The lateral nerves, of w
there are 10-12 (—15) pairs, with the lowermost and uppermost Spake;
are slightly elevated above and more prominently so beneath, and diverge
from the costa at an angle of 35-55°. The reticulation is very prominent
beneath, slightly less so above. The axillary and subterminal paniculate
inflorescence, 15-30 cm. in length, is glabrous or glabrescent, borne on a
stout brownish or reddish brown angled striate peduncle up to 10 cm. long.
The flowers are up to 2.5 mm. long, glabrescent without; the glabrous to
glabrescent slender pedicel is up to 5 mm. long, the tube well defined, the
ovate or broadly ovate acute thinnish lobes + 1.7 mm. long, and frequently
broader than long at the base. The stamens of the two outer series have
the anthers ovate, frequently with a definite connective, measuring + 1.
mm. overall, about twice the length of the pubescent filaments. Those of the
inner series are + 1.7 mm. long, the more narrowly ovate anthers three
times the length of the filaments, which bear at the base two conspicuous
ovate cordate short stipitate glands which are about half the length of the
anthers. The glabrous gynaecium is + 2.4 mm. long, the ovoid ovary
about equal to the stout style topped with the discoid stigma. The ellip-
soid smooth shining fruit, about 17 mm. long sey 1 cm. broad, is borne on
a shallow irregularly lo bed cupule not more than 2 mm. long and deep and
8 mm. in diameter. The expanded deeply aciculate pedicel is up to 5 mm.
long and 3—4 mm. broad at the ti
Tonduz 7613 is annotated, presumably by Mez, as “Ocotea paradoxa
Mez, n. sp.,” but it is the syntype of O. pentagona and is a very good match
for Pittier & Tonduz 9172, the fruiting syntype of O. pedalifolia. Although
only a photograph of the type of O. nicaraguensis is at hand, I am reason-
ably certain of the numbers being conspecific.
12. Ocotea Standleyi, nom. nov
sie i a as Standley & Steyermark in Field Mus. Publ. Bot. 23: 116. 1944,
eae aan Known at present only from Guatemala, at an altitude of 1200-2700
m., usually in wet forests.
GuateMaLaA: Alta Verapaz: Southeast of Tactic, in wet forest, alt. about 1500
m., March 30, 1939, Standley 70009 (fl., fr., TyPE, Ch); wet forest near Tactic, above
the bridge across Rio Frio, Standley 90429 (sterile, Ch) ; wet mixed forest, mountains
along road between Tactic and the divide on road to Tamahu, Standley 91326 (fl., Ch).
n Marcos: Wet forest, Barranco ee road between San Marcos and San
Rafael Pie de la Cuesta, in upper part of t ae between Finca La Lucha and
Buena Vista, Standley 86538 (sterile, Ch). ee ezaltenango: Dense mixed forest,
along old road between Finca Pirineos and Patzulin, ad 86927 (fr., Ch)
This large tough-leaved tree or large shrub has stout branchlets that are
slightly angled, striate, glabrous, and castaneous or reddish brown. The
heavy glabrous subcanaliculate petioles are the same ees sa continues
up into the stout broad costa, and they measure up to 2.5 cm. long and
about 3 mm. in thickness. The blades are heavily coriaceous, glabrous
344 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
throughout, elliptic or obovate-elliptic, the base obtuse or rounded and
occasionally with a slight tendency toward being cordate, the apex rounded
or obtuse or possibly shortly abruptly subacuminate, up to 30 cm. long an
12 cm. broad. The costa is conspicuous above, although impressed slightly,
the lateral nerves, of which there are upwards of 12 pairs, diverging at an
angle of about 45°, are slightly elevated above, and both costa and nerves
are prominently elevated beneath. The reticulation is apparent although
not prominent above, but conspicuously so beneath. The inflorescence is
surprisingly slender for such leathery foliage, ee of axillary panicles
paratively few-flowered, the peduncle up to 5.5 cm. long. The flowers and
pedicels are each about 3 mm. long and pubescent. ‘The tube is well de-
= and the lobes are broadly ovate (broader than long), thick, and
e stamens of the two outer series are + 1 mm. long,
the aoe anthers oa slightly longer than the slender filaments. Those
enormous lobed glands arising from the base. The staminodia are slender
stipitate bodies, pubescent toward the base, the apical portion subrhomboid,
measuring in their entirety about 0.6 mm. long. The glabrous gynaecium
measures + 1.7 mm. long, the subglobose ovary equalled by the tapering
style, which bears a subtriangular stigma that is decurrent. The fruit is
green (subcastaneous in the dried pc ener Parry apiculate,
measuring up to 2 cm. long and 1.5 broad, ted by a shallow
cupule that is woody, Satna, ae at ue cite and
measuring 5—7 mm. long, 12 mm. in diameter, and 3 mm. deep; the pedicel
is thickened and also verruculose, measuring up to 1 cm. long and about
4 mm. in diameter at the apex.
The size and texture of the leaves suggest O. nicaraguensis, but the
smaller fruit of the latter has a lobed less woody cupule, and the floral struc-
ture differs.
13. Ocotea atirrensis Mez & J. D. Smith ex pee in igs Jahrb. 30: Beibl. 67: 18. 1901;
Standley in Field Mus. Publ. Bot. 18: 454.
DistriBuTION: Costa Rica oe — faa enue at an altitude of 700-850
m. bling it is found as a as 1
Costa Rica: Alajuela: ees uis de Zarcero, in shady forest of Pacific cloud-
Paes zone, A. Smith HS77 (fl., Ch); Villa Quesada, San Carlos, shade of Caribbean
ae forest, A. Smith H.1737 (fl., A, Ch); La Palma de San Ramon, Brenes 20596 (fl.,
ae near Rio Naranjo, Pittier (Herb. Inst. Costa Rica 7592 b) (fl., syNTYPE not seen).
Cartago: cinity of Pejivalle, in forest, Skutch 4591 (fl., A); Atirro, alt. 700 m.,
Ma on , J. D. Smith 4930 (fr., sosyNTYPE, US). PANAMA: Bocas del Toro:
Vicinity ne pict Lagoon, von Wedel 1399 (fl., A).
NATIVE NAME: “Quizarra” (Costa Rica).
This species is characterized by rubescent angled branchlets that are
early covered with a dense minute ferruginous pubescence which quickly
disappears, pahaess the branchlets glabrescent or glabrous. The es are
borne on stout canaliculate petioles, glabrescent except for the residue of
oblong or oblanceolate-elliptic, the base roundish, obtuse or no encd
cuneate, the apex caudate-acuminate, up to 38 cm. long and 10-13 ¢
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 345
broad. The broad costa is impressed above and the lateral nerves, of which
there are 12—16 pairs, diverging at an angle of 45°, are rather obscure, but
all are elevated prominently beneath. The a sete is visible, although
only slightly raised above but prominently so beneath. The axillary pani-
culate inflorescence is 15-20 cm. long, pubescent, feescane inetd the
peduncle up to 8 cm. long. The glabrescent flowers are up to 3 mm. long,
with a short glabrescent pedicel not more than 2 mm. long. ‘The lobes are
thinnish, ovate-elliptic or even broadly ovate, obtuse, papillose within,
ee 2 mm. long. e stamens of the two outer series measure about
0.8 mm., the ovate subrotund or obtuse ee wees twice the length of
the eee Those of the inner series are ovate, obtuse, + 1.25 mm
long, the anthers supported by filaments slightly iene and bearing two
subreniform sessile glands at the base. The glabrous gynaecium is + 1.7
mm. long, the ovoid to subglobose ovary equalling the style, that bears an
obtuse usually inconspicuous stigma at its apex. The black fleshy ellipsoid
fruit measures up to 3.3 cm. long and at least 1.5 cm.
seems to be a shallow disk-like undulate-margined structure measuring up
to 8 mm. in length and diameter, and less than 2 mm. deep. The support-
ing pedicel is very short and slightly enlarged to 2 mm. at the apex.
better prepared fruiting specimen of the species may change this concept
of the cupule-structure and dimensions.
Ocotea atirrensis stands out because of the very long oblanceolate leaves,
which are membranaceous. The species is perhaps more nearly related to
O. Pauli than to any other of this region, but it is easily separated by the
texture of the leaves and the different type of reticulation.
14. Ocotea Paulii, sp. nov.
Arbor ad 15 m. alta, ramulis angulatis brunneis mox griseis glabris sul-
catis. Folia alternata, glabra, petiolis robustis glabris atro-rubescentibus
esetaie a eae multiflora, pedunculo rubescente, ad 6 cm - Jongo.
Flores ad 2 mm. longi, pedicellis + 2 mm. longis, gracilibus, perianthio
campanilato pallide iivecente lobis late ovatis obtusis membranaceis
a m. longis; staminibus ser. I & II + 1 mm. longis, antheris ovatis
Rents. duplo longioribus, ser. III + 1.25 m longis, antheris ovato-
quadratis, filamentis conspicue biglandulosis, ia alle stipitatis antheris
subaequalibus; staminodiis nullis; gynaecio glabro, + 1.7 mm. ongo,
ovario late ovoideo vel ae stylo aequali, stigmate conspicuo.
Fructus atratus (in sicco), oblongus, 18-20 * 10-11 mm., cupula_ sub-
anes al giana rugosula, glabra, ++ 2 mm. longa et 5—6 mm. diam., 1
mm. alta, margine undulata, subtentus, pedicello satis incrassato, glabro,
3-4 mm. longo.
DistriBuTION: Panama, at an altitude up to 1100 m., and Costa Rica, up to
450 m.
346 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
Costa Rica: Alajuela: San Luis de Zarcero, edge of Pacific watershed cloud-
forest, A. Smith H.633 (fl., Ch) ; woods, La Palma de San Ramon, Brenes 5590 (181)
, Ch). Panama: Chiriqui: Between Cerro Vaca and Hato del Loro, Pittier
5395 (fl., Ch). Coclé: Vicinity of El Valle, alt. 600-1000 m., Dec. 8, 1938, Paul H.
Allen 1211 (fl., Ch, type —GH, Mo) (tree 15 m.; flowers pale yellow) ; south rim
(dry), Allen 1775 (fr., Ch, GH, Mo, NY); region south of El Valle de Anton, Allen
2848 (fl, A)
The nearest affinity of O. Paulii is O. atirrensis, but the new species is
quickly distinguished by the coriaceous leaf-blades which are bright brown
on drying. The apex of the blades of O. atirrensis is definitely caudate.
15. Ocotea Dendrodaphne Mez in Jahrb. Bot. Gart. Berlin 5: 238. 1889; Standley in
Contr. U. S. Nat. Herb. 27: 183. 1928.
Dendrodaphne macrophylla Beurling in Vet. Akad. Handl. Stockholm 1854: 145.
1856.
Ocotea Quisara Mez & J. D. Smith in Bot. Gaz. 33: 259. 1902; Standley in Field
Mus. Publ. Bot. 18: 456. 1937.
Ocotea ovandensis Lundell in Contr. Univ. Mich. Herb. 6: 16. 1941.
DisTRIBUTION: Mexico through Guatemala, Honduras, and Costa Rica to Panama,
from 200 to about 1250 m. altitude.
hiapas: Mt. Ovando, Dec. 17, 1936, Matuda 444 (fl., isotype of O.
ovandensis, A, NY), 1839 (fr. A, NY). GUATEMALA: Quezaltenango: Colomba,
in coffee plantation, Skutch 1980 (f., A, Ch, NY), by roadside, Skutch 2007 (fl., A, Ch).
Honpuras: Province unknown, in forest near Bolet’s plantation, P. Wilson 351
(sterile, GH). Atlantida: Tela, near Lancetilla, Bangham 214 (fr., Ch). Costa
rca: Alajuela: Villa Quesada, San Carlos, in Caribbean rain-forest, A. Smith F.
1765 (sterile, fl, A, Ch), 1771 (fl., A, Ch); Suere, A. Smith F. 1849 (sterile, fl., A, Ch).
Limoén?: La Emilia, Llanuras de Santa Clara, alt. 250 m., April, 1896, J. D. Smith
6751 (fl., isosyntype of O. Quisara, GH) ; La Concepcion, Llanuras de Santa Clara, alt.
250 m., Feb. 1896, J. D. Smith 6756 (ir., isosyntype of O. Quisara, GH). Cartago:
Volcan de Turrialba, Tonduz 9020 (fr., Ch); Atirro, alt. 600 m., April, 1896, J. D.
Smith 6753 (syntype of O. Quisara not seen). Panama: Colon: Near Puerto Bello,
Billberg s.n. (fl., type of Dendrodaphne macrophylla not seen). Coclé: Hills north
of El Valle de Anton, vicinity of La Mesa, P. H. Allen 2299 (Ir., Ch).
Native NAMES: “Quisara” (Costa Rica) ; “Sigua” (Panama, fide Standley ).
This species is striking because of the usually silvery-gray angled branch-
lets, early closely and minutely pubescent but soon glabrous. The blackish
stout canaliculate glabrous petioles up to 2 cm. long offer a contrast to the
branchlets. The blades are chartaceous, glabrous throughout, or with a
few dark hairs persisting at the base on the lower surface, usually pale
green to light brown in the dried state, elliptic or oblong-elliptic, the base
obtuse or abruptly cuneate, rarely rounded and abruptly cuneate, the apex
variable, obtuse, or acutish, or even acuminate to caudate-acuminate. The
blades measure up to 30 cm. long and up to 10 cm. in breadth. The robust
costa is slightly impressed above and prominently elevated beneath. The
lateral nerves number 9 or 10 pairs and are delicately and imperceptibly
elevated above and more prominently beneath, diverging at an angle of
about 45°. The upper surface of the blades is very smooth, showing no
flowered, the peduncles not more than 4.5 cm. long and usually less. The
flowers vary in size from 4-6 mm. long, the elliptic or oblong or ovate-
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 347
elliptic fleshy papillose lobes rounded or obtuse and up to 5 mm. long. The
stamens of the two outer series are + 2.8 mm. in length, the ovate acutish
or obtuse petaloid anthers borne on exceedingly short stout filaments, the
upper third of ba anthers being connective tissue. Those of the inner
series are longer, + 3.2 mm., and the filament is also longer, being about
one-third the ie. length of the stamen, and bearing two small subsessile
glands at the base of the anther. Wherever staminodia have been ob-
served, they are cordate, subsessile, and + 0.6 mm. long. The glabrous
gynaecium is + 2.4 mm. long; the ovary is probably ovoid, there being
seemingly no distinct line of demarcation between it and the stout style,
which is topped by a usually conspicuous stigma varying from capitate to
decurrent. The fruit is ellipsoid, black, + 2.3 cm. long and 1.2 cm. broad,
ee ina glabrous woody cyathiform cupule that is 1 cm. long, 6 mm. deep,
14 mm. in diameter, with a shallowly and obscurely lobed and slightly
afiiitate double margin, subtended by a thickish striate pedicel up to 6 mm.
long and pubescent to glabrous.
There are differences between the material collected in Mexico and
Guatemala and that of Costa Rica. For example, the flowers of specimens
collected in the former area are smaller than those of the southern area.
The fruits, also, are smaller and the cupules subtending them are shorter.
Nevertheless, it seems advisable to combine the two, since the differences
are merely of degree
The nearest relative of the species appears to be the smaller-leaved O.
veraguensis, which has essentially the same floral structure, the style being
more pronounced, the same incipiently double-margined cupule, and
blackish petioles contrasting with the grayish angled branchlets and bearing
leaf-blades that show no reticulation on their upper surface.
16. Ocotea veraguensis eee Mez in Jahrb. Bot. Gart. Berlin 5: 240.
Standley in Contr. U. S. Nat. Herb. 23: 295. 1922; Standley & Spreiiee Lista
Prelim. Pl. Salvador he 1925; eyes in sg er bet Bot. 12: 45
Sassafridium veraguense Meissner in DC. Ley ou:
Ocotea Bakeri Blake in Contr. Gray Herb. n.s. 52: 65. ae
Ocotea escuintlensis Lundell in Contr. Univ. Mich. aon 6: 15. 1941.
T a General in forests throughout the ee along the Pacific coast
(Pacific tierra caliente) from Mexico to Panama, usually a very low altitude; occa-
sionally found inland at higher altitudes (up to 1200 m. ). . wood is used in some
localities for construction. Some parts of the tree are used by the natives of Nicaragua
as a tonic
Mexico: Sinaloa: El Castillo, Mazatlan, Ortega 6913 (ir., Ch). Nayarit
(Tepic): San Blas, Lamb 611 (fr..GH, NY). Guerrero or Michoacan: Botella,
Langlassé 656 (fl. GH). Guerrero: Mountains around Acapulco, Gamon 6 (sterile,
Y). Chiapas: Tonala, in undergrowth along the banks of the river, C. & E. Seler
1886 (fr., GH); near Tapana and La Junta in the high forest of the river-bed, C. & E.
Seler 1889 (fl., GH, NY); Escuintla, May 3, 1936, Matuda 654 (fl., type of O. escuint-
lensis, Mich) ; San Pedro, Mell 558 (fr., NY). GUATEMALA: ‘Cuutena la: Rio
Ocosito, J. D. Smith 2608 (fl., fr., GH, NY). Escuintla: Below Las Lajas, common
in damp forest, Standley 64785 (fl., A). Ex Satvapor: Dept. unknown, Cerro del
Guayabal, S. Calderon 1971 (fl, GH). Ahuachapan: Sierra de Apaneca, in forest
in the region of Finca Colima, Standle Z 20178 (sterile, GH). La Libertad: Coma-
sagua, S. Calderén 1390 (fr., GH). n Jacinto: Cerro de San Jacinto, S. Cal-
deron 2258 (fl., fr., GH, NY) ; in ee near summit of Cerro de San Jacinto, near San
348 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
Salvador, Standley 20638 (fl, GH, NY). San Vicente: In quebrada, vicinity of
San Vicente, Standley 21668 (fl. GH, NY). Usulutan: Berlin, S. Calderén 2146
(fl, GH, NY). Nurcaracua: Without locality, Wright s.n. (ir., NY); Prov. unknown,
Volcan Marabacks, Baker 2493 (fl., type of O. Bakeri, GH). Costa Rica: Guana-
caste: Roadsides south of Liberia, Dodge & Thomas 6312 (fl, NY); near Nicoya,
Near Nee op Bridges (SYNTYPE of Sassafridium veraguense not seen), Warscenice
(fl., photo. of syNTYPE of ee veraguense, Ch, GH). ocle: enonome
ee vicinity, R. S. Williams 236 (fr.. NY). Panama: Rio Las Lajas, P. H. Allen
11 (fi., fr., GH, Mo, NY).
ATIVE NAMES: ‘‘Aguacatillo’” (Costa | “Canelillo” (Costa Rica) ; “Canelito”
(El ae dor) ; “Canelo” (Costa Rica, El Salvador) ; “Laurel” (Mexico) ; “Palo colo-
rado” (Nicaragua); “Pimientillo” (El Salvador); ‘“Pimiento” (El Salvador) ; “Qui-
sarra” (Costa Rica) ; “Sigua canelo” (Panama).
This coastal species has branchlets that are finely gray- or tawny-seri-
ceous, very quickly becoming pale gray, glabrous, and striate. The black-
ish stout canaliculate glabrescent petioles, up to 1 (sometimes to 1.5) cm
long, often make a striking contrast with the smooth grayish branchlets.
The blades of the leaves are chartaceous to subcoriaceous, early soft
grayish-pubescent below, becoming entirely glabrous, the margin in the
dried state usually conspicuously minutely undulate or crisped. The color
is uniformly a pale grayish green and the upper surface is very smooth.
The blades are variable in shape, being elliptic, lanceolate-elliptic, oblong,
oblong-elliptic, and finally rarely obovate, the base obtuse or cuneate, the
apex rounded-obtuse or obtusely acute. The costa is very prominent above
and beneath, although only slightly elevated above and more conspicuously
so beneath. The lateral nerves, of which there are 6 or more obscure pairs,
are not visible above and only faintly so beneath. The reticulation is not
visible above and is very inconspicuous beneath. The inflorescence is
axillary, paniculate, grayish-pubescent to glabrescent, many-flowered, the
ultimate branchlets frequently giving the appearance of ieee a head-
like formation. The entire length of the panicle varies from 3 to 13 cm
the peduncle sometimes attaining a length of 9 cm. ‘The flowers are pu-
bescent, up to 5 mm. long and more than twice that in diameter at anthesis,
are supported by slender pubescent pedicels to 6 mm. long, and show a well
developed tube. The lobes are oblong, obtuse or acutish, thick, fleshy,
papillose, up to 3 mm. long. The stamens of the two outer series are + 2.4
mm. long, the petaloid ovate anthers nearly sessile, the protruding papillose
connective occupying at least the upper third of the anther. Those of the
inner series are often slightly longer than the outer ones, the anthers being
subtended by short filaments which bear at the base two conspicuous short-
stipitate or sessile glands that are nearly the length of the anthers. The
staminodia are variable, being absent or, if present, linear or ae i
a definite stipitate ovate head, always pubescent, from + 0.6 to + 0.9 mn
long. The glabrous gynaecium also varies, sometimes Sree sey a anaes
of + 2.15 mm., the ellipsoid to subglobose ovary usually twice the length
of the tapering style with its usually conspicuous triangular stigma. The
fruit of the species is very poorly represented in collections. It seems to
be (immature?) greenish or light brown, not very firm in texture, ellipsoid,
apiculate, up to 1.7 cm. long and about | cm. broad, subtended by a shallow
thinnish cupule about 5 cm. long, 12 mm. in diameter, and about 3-4 mm.
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 349
deep. The margin is distinctly double, as in the genus Licaria — in fact
supe t
mm. The aie is up to 1 cm. long, enlarging toward the apex to about
3 mm. in diam
The grayish aha the blackish petioles, and the smooth gray-green
leaf-blades recall to mind the larger-leaved species O. Dendrodaphne. The
floral structure is of the same type, and the fruits are subtended by double-
margined cupules. The consistently extremely large leaves of O. Dendro-
daphne, however, preclude the possibility of its being a variety of the more
widely spread O. veraguense.
Meissner cites as syntype Oersted 10, which is a Litsea, but Mez cites
Oersted 12 and 13, from Costa Rica. I have seen neither of these “praia
Most of the specimens from El Salvador show larger stamens, + 1.25 m
in length, and frequently these are equipped with glands similar to nae
of the stamens of the third or inner series. Occasionally a specimen (i.e.,
A. Smith P.2464) will have flower-parts that are extra large throughout.
17. Ocotea chiapensis (Lundell) Standley & Steyermark in Field Mus. Publ. Bot. 23:
Nectandra chiapensis Lundell in Contr. Univ. Mich. Herb. 6: 12. 1941.
DISTRIBUTION: ee only from type-locality and vicinity in Mexico and from
adjacent ee
Mexico: picnic: At Cerro Laguna, Mapastepec, January, 1938, Matuda 2042
(fl., isotype, A, Ch, NY); Rodeo, near Siltepec, Matuda 4579 (fi., A). GUATEMALA:
San Marcos: Along river, between La Vega ridge along Rio Vega and northeastern
slopes of Volcan Tacana, to 3 miles from Guatemala-Mexico boundary, in vicinity of
San Rafael, Steyermark 36195 (fl., Ch).
This species has stout cicatricose aera that are angled at first and
brownish-sericeous, becoming glabrescent and eventually grayish brown.
The leaves are supported by eh apparent petioles up to 2 and some-
lower leaf-surface and the exceedingly conspicuous ellipsoid pubescent
glands that occur in nearly all of the axils of the lateral nerves. The blades
are usually obovate-elliptic, occasionally oblong-elliptic, up to 15 cm. long
and 7.5 cm. broad, the base (not including the decurrent petiole) rounded
or cuneate, the apex acutish or subacuminate. The broad costa is only
slightly elevated above and conspicuously so beneath. The tenuous lateral
nerves, of which there are 9-12 pairs, are faintly outlined above but slightly
elevated beneath. The reticulation is obscure above and slightly promin-
ulous beneath. The inflorescence is axillary, often few-flowered, brown-
ish-sericeous, up to 15 cm. long, the stout pedicel attaining a length of 5 cm.
The flowers are + 3.5 mm. long and about 6 mm. in diameter, the pedicel
+ 1.7 mm. long, the subequal thick ovate acutish papillose lobes reflexed
and + 2.15 mm. long. The stamens of the two outer series are about
1.25 mm. long, the elliptic rounded anthers lags the length of the fila-
ents. Those of the third series are + 1.5 mm. long, the oblong rounded
anthers slightly pubescent at the base and only slightly longer than the
350 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
filaments that bear conspicuous basal glands. The staminodia are pubes-
iad == 0.6 mm. long, either st -like or occasionally ovate. The glabrous
gy ium measures + 2.15 mm., the ellipsoid ire ad longer than
the ite style topped by a sanecus peltate sti
The species probably belongs near O. Austinii, on may be separated by
the smaller less robust and less pubescent inflorescence and the more promi-
nent reticulation of the leaf-blades, as well as by the presence of very large
conspicuous axillary glands.
18. Ocotea Austinii, sp. nov
Arbor 10-18 m. alta, ee striatis, juventute minute brunneo-sericeo-
paige reat mox gris €0- vel atrato- pubescentibus Goinde sea apapiigentirs
ticis, 7.5-9 ~11) cm. lance et 3-4.5 cm. latis, basi cuneatis in ire
decurrentibus ibique valde recurvatis, apice obtusis vel leviter et obtuse
subacuminatis, penninerviis, costa supra satis subtus conspicue elevata,
utrinque pubescente, nervis 7-10 paribus supra aeque et regulariter leviter
elevatis concoloribus, subtus pubescentibus obscuris angulo 45° srt ae
tibus, interdum glandulis obscuris in nervorum lateralium axibus, r
larum supra aeque et regulariter elevato subcancellato. =a ametored
axillaris anguste paniculata, subcapitata, pauciflora, robusta, dense sericeo-
ferrugineo-pubescentia, ad 8 cm. longa et a cm. lata, longipedunculata,
pedunculis 3—5 cm. longis. Flores + 2.5 mm. longi, sessiles vel pedicellati,
pedicellis + 2.5 mm. longis, perianthio campanulato (urceolato-campanu-
lato, fide coll.), sulfureo-flavescente, fide coll. (vel viridescenti- flavescente),
lobis late ovatis acutis crassis, dense pubescentibus, 3 mm. longis; stami-
nibus ser. I & I + 1.9 mm. longis, antheris filamentis aequalibus, ser. IIT
+ 2.15 mm. longis, antheris filamentis aequalibus, conspicue biglandulosis,
Sananli sessilibus, antheras aequantibus; staminodiis nullis; gynaecio
glabro, + 2.5 mm. longo, ovario subgloboso vel obovoideo, longitudine
ultra 14 gynaecii aequante, stigmate triangulari, conspicu o. Fructus late
obovoideus atro-olivaceus, minute canescenti- vel griseo-canescenti-puncta-
tus, fide coll. 28 20 mm., cup ula vadosa rubra subcampanulata vel ma-
cente rugosula, 6-7 cm. longa et ad 1.5 cm. lata, + 3 mm. diam. subtentus,
pedicello glabro rugosulo aciculato + 1 cm. longo.
DistrisuTIoN: In cloud-forests or cleared pasture-land of Costa Rica and in rain-
and cloud-forests of the adjacent areas of Panama, at an altitude of 1765 to sa m. in
Costa Rica and at 1980 m. in Panama. The wood is used for lumber in Panam
Costa Rica: Alajuela: Alfaro Ruiz, Palmira, region of Zarcero, in - clay -
(forest tree 15 m., base 1 m; bark light brown, roughened, cambium-layer rufous-
brown, sapwood cream-colored; leaves coriaceous, polished on upper surface, lighter
green below; flowers sulphur-yellow, with an expanse of 4 mm., urceolate-bell-shaped,
stigma and anthers ecten, maturing to dark brown), A. hear A.504 (fl., Ch), 4.1181
(fl, Ch). San José: Near Camp Empalme in oak cloud-forest, along the w. slope
of the Continental Divide, oak 29 km. S. of Cartago, rites 6007 (F.S.95042) (fr., “
Panama: Chiriqui: In rain-forest, Bajo Chorro, Boquete, Davidson 268 (fr., A,
Ch, Mo); cloud-forest, Cerro Horqueta, C. & V. W. von Hagen 2128 (tr., A, Mo).
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 351
NATIVE NAME: “Sigua Canela” (Panama).
The exceedingly prominent and regular more or less cancellate reticula-
tion of the leaf-blades of this species separate it from the related species O.
Endresiana, O. Tonduzii, and O. Skutchii. The blades also differ in the
sericeous pubescence and succeeding glaucescent aspect of the lower surface.
19. Ocotea Endresiana Mez in Jahrb. Bot. Gart. Berlin 5: 257. 1889; Standley in Field
Mus. Publ. Bot. 18: 455. 1937.
DistTRIBUTION: Known only from Costa Rica.
Costa Rica: Without locality, Endres 223 (fl., TYPE not seen). Alajuela: La
Palma de San Ramon, Brenes 6317 (fi., Ch) ; Los Angeles de San Ramon, Brenes 16227
(fl., Ch); Piedades Sur (La Palma) de San Ramon, Brenes 17172 (fr., Gh):
NaTIVE NAME: “Maranon” (Costa Rica).
A species that is very difficult to separate, without dissection of the
flower, from Aiouea. The stout cicatricose branchlets are dark brown,
cm. long, the peduncle 2-3 (—6) cm. long. The flowers are + 5 mm.
long, glabrous, supported on slender pedicels 2 (—4) mm. long. The ovate
acute lobes are fleshy and up to + 2.5 mm. long. The stamens of the two
outer series are up to -- 1.25 mm, long, the ovate roundish anthers only
slightly longer than the somewhat stout filaments. Those of the inner
series are -- 1.7 mm. long, the more narrowly ovate anthers surpassed
slightly by the filaments which bear two large subglobose sessile glands at
the base. The glabrous gynaecium is + 2.4 mm. long, the ellipsoid ovary
only slightly exceeding the stout style in length, the whole topped by a
discoid inconspicuous stigma. (Mez describes the ovary of the type-speci-
erect style with discoid-obtuse stigma.) The fruit (that of Brenes 17172,
the vegetative characters of which match Mez’ description of O. Endre-
siana) is ellipsoid, apiculate (immature?), up to 1 cm. long and 7 mm.
broad, and is borne in a shallow woody cupule with an entire margin, 3—5
mm. long and about 5—6 mm, in diameter and not more than 1 mm. deep;
the thickened pedicel is not more than 2 mm. or so long and 3-4 mm. in
diameter at the apex.
The species is separated from O. Tonduzii by the very broad decurrent
and recurved sessile blades, forming a much longer apparent petiole, and
in fruit by the entire cupule.
20. Ocotea Tonduzii Standley in Field Mus. Publ. Bot. 18: 456. 1937.
Ocotea cuneata Mez in Jahrb. Bot. 30: Beibl. 67: 17. 1901, non Gomez, 1894.
DistripuTiIon: Costa Rica, in forests from 640 to 1200 m. altitude.
NATIVE NAME: “Ira” (Costa Rica).
Costa Rica: Province unknown, near Rancho Flores, Tonduz 2142 (fl., syNTYPE of
352 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
O. cuneata not seen). Heredia: On banks of the Rio Segundo, alt. 2000 m., Tonduz
(Herb. Inst. Costa Rica) 1739 (fl., syNTYPE of Ocotea cuneata not seen). Sen José:
Vicinity of El General, Skutch 4303 (fr., A, Ch, NY). Cartago: Forests of Que-
brada Honda, near Juan Vinas (Atlantic), Tomius 10415 (fr., isosyntype of O. Jra, US).
The fruiting material above matches very well with the description given
by Standley. It is with little hesitation that the numbers are placed here.
The branchlets are glabrous, grayish, almost checkered, very thick, and
angled toward the tips. The leaves are subsessile, the base of the blade
decurrent and strongly recurved, making an apparent petiole up to 2.5 cm.
long. The coriaceous glabrous blades are opaque, obovate, attenuately
cuneate at the base and rounded or very inconspicuously and shortly acumi-
nate at the apex. They measure up to 15 cm. long and 7 cm. broad (those
of Tonduz 10415 are much smaller). The costa is impressed above but
prominent beneath. The lateral nerves, of which there are upwards of 8
pairs, are delicately elevated above and more prominently so beneath. The
reticulation is obscure throughout. The inflorescence, according to Mez, is
stoutly se ie paniculate, the pedicels 0.5—1.5 mm. long, the bracteoles
deciduo The flowers are 2.5-3.5 mm. long, the perianth-tube barely
distinct ya the pedicel and gradually merging into it. The lobes are
broadly ovate, rotund or acutish. The filaments are glabrous and those of
the inner series have two large glands at the base, irregularly and incised-
fruits are 1.5 cm. long and 8 mm. broad, borne in woody cyathiform red
cupules still crowned with the remnants of the perianth-lobes, 4-6 mm.
long, about 8 mm. in diameter, and about 4 mm. deep. The pedicel i is very
short, not more than 2 mm. long and expanded to the same width at the
ex.
Ocotea Tonduzii is very close to O. Endresiana, but may be separated
from the latter by the very short petiole, narrowly decurrent blades, the
6-lobed cupule, and the pubescent axillary glands. The fruiting material
of this species is often confused with that of Aniba, and it is verv difficult to
enumerate the points of difference.
The confusion of numbers cited by Mez under O. /ra is taken up under
that species.
21. Ocotea Skutchii, sp. nov.
Arbor ad 27 m. alta, ramulis brunneis angulatis mox griseis sulcatis.
Folia alternata, juventute fulvo-sericeo- pubescentia mox glabrescentia vel
glabra, petiolis conspicue alatis, alis recurvatis, inconspicue pilosis vel gla-
bris, ad 2.5 cm. longis et basi 2 mm. apice ad 8 mm. latis, laminis supra
glabris, subtus sparse et inconspicue pubescentibus, satis coriaceis, in sicco
supra viridescentibus subtus pallidioribus vel subglaucescentibus, ellipticis
vel leviter obovato-ellipticis, 10.5 (—13) cm. longis et 4—4.5 (—5.5) cm. latis,
basi cuneatis, in petiolum decurrentibus ibique valde recurvatis, apice
abrupte obtuse acuminatis, penninerviis, costa supra haud impressa subtus
valde elevata, costa nervisque utrinque conspicue flavescentibus vel alb
centibus, interdum costa subtus rubescente, nervis 8 vel 9 paribus supra
leviter impressis subtus elevatis, angulo 45° divergentibus, interdum
a
culata, ad 16 cm. longa, pubescens, pedunculo ad 6 cm. longo. Flores
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 353
2.5-3 mm. longi, pedicellis ad 5 mm. longis, gracilibus, Paar gee campanu-
lato, cing fide coll., lobis late ovatis, crassis 1.7 mm. longis;
staminibus ser. 1 & 1+ 1 mm. longis antheris ovatis eruneael aimee
duplo longioribus, ser. III + 1.5 mm. longis conspicue biglandulosis,
glandulis subglobosis stipitatis; staminodiis nullis; gynaecio glabro + 1.8
mm. longo ovario ellipsoideo stylo gracili duplo longiore, stigmate parvis-
simo inconspicuo. Fructus non visus viridis, fide coll., cupula rubra vadosa
leviter campanulata minute verruculosa crassa glabra ad 1 cm. longa, 9 mm.
diam., 3 mm. alta subtentus, pedicello glabro, striato, ad 7 mm. longo.
Dicemeueey: Known only from Costa Rica, at 700-1680 m. altitude.
Costa Rica: Heredia(?): Vicinity of Vara Blanca, north ae of ie Cor-
dillera, between Pods and Barba volcanoes, Skutch 3755 (fr., A, NY). an José:
Saige of El General, alt. 700 m., Dec. 1936, Skutch 3062 (f1., Sear NY) (tree
27 m.; flowers yellowish).
at NAME: “Ira rosa” (Costa Rica).
This species may be separated from O. Tonduzii and O. Endresiana by a
combination of characters, such as the short petiolate leaves with the blades
only narrowly decurrent on the petioles, the apparent petiole up to 2.5 mm.
long, axillary glands either absent or if present conspicuous for being nar-
rowly ellipsoid, and the entire cupule.
22. Ocotea rubrinervis Mez in Jahrb. Bot. Gart. Berlin 5: 351. 1889.
DistRIBUTION: Panama, and south to Peru and Bolivia; probably So ag in the
intervening areas. Not mentioned by Kostermans in Pulle’s Flora of Sur
PANAMA: Without locality, Duchassaing (SYNTYPE not seen). San fae Island,
Perlas bree Gulf of Panama (about 55 miles SSE. of Balboa), es beageis forest
along South Road (S° area), ie 111, 133, 250, 415, 497 (fl., A), , 563, 604,
655 (fr., A). ipa San Martin: Near Tarapoto, pee 4580 aaa of syN-
The species is described as having young branchlets pilose at the apex
the adult ones glabrous, cinereous, terete. The branchlets of the Johnston
length cinereous. The short canaliculate petioles are also pubescent, be-
coming glabrescent, and measure to 1 cm. long as opposed to Mez
tion. The blades of Mez’ description are foveolate and glabrous above,
pilose beneath. Those of Johnston’s collections are scattered- pubescent
near the base, rather reticulate above, glabrescent beneath except for the
pubescent axillary glands. The blades are elliptic or broadly elliptic, the
base subrounded, obtuse or cuneate, the apex shortly aarsie acuminate or
obtusely acute, measuring to 11 cm. long and up to 7 cm. in breadth, larger
on the whole than the specimens of Mez’ description, The ci and lateral
nerves, of which there are 4—6 pairs, are impres sed above and prominently
elevated and yellowish beneath. The inflorescence is axillary or sub-
terminal, slender, the branches of the ecuale narrowly racemose, glabres-
cent for the most part, the flowers grayish-pubescent rather than ferru-
ginous-pilose. The staminate flowers are yellow, about 3 mm. or less long,
and sessile or borne on a short pedicel less than 1 mm. long. The ovate
acutish lobes are rather thick and + 2.15 mm. long. The stamens of the
two outer series are + 1.5 mm. long, the ovate obtuse to roundish anthers
slightly longer than the filaments. Those of the inner series are slightly
larger and have anthers more narrowly ovate, supported by filaments com-
354 JOURNAL OF THE ARNOLD ARBORETUM [VOL, XXvI
pletely covered on the dorsal side by the conspicuous fused sessile glands
borne at the base. The slender transparent coating: less than one-half
a millimeter in length, are oblanceolate. ez says that all staminodia
are aborted in his material. The aborted eee gynaecium is + 1.
mm. long, subcylindrical, the basal part bulging only slightly, and the apex
siaetieg a conspicuous (triangular according to Mez) stigma depressed
bov
Nous there are discrepancies and Johnston’s material does not pre-
cisely match the description of Mez’ species, the two are so close that it
seems unwise, at least until the types are available, to describe Johnston’s
collections as new.
23. Ocotea subsericea Standley in Field Mus. Publ. Bot. 18: 456. 1937.
DistripuTION: Costa Rica and Panama, at an altitude of about 1250 m.
C Alajuela: La Palma de San Ramon, 1250 m. alt., March 30, 1929,
Brenes 6789 (665) (f1., re Bret Panama: Coclé: North rim sf El Valle, P. H.
Allen 1907 (fr., Ch, GH , NY
This species was rete — a specimen which seemingly is not only
from a branch that is very young, but growing in an unexposed situation.
The branchlets are minutely but densely appressed-yellowish-buff-sericeous-
tomentose, angled, becoming grayish, glabrous, terete, and striate-sulcate.
those of the adult leaves more thickly membranaceous, elliptic, the base
cuneate, the apex abruptly and obtusely acuminate, up to 12 cm. long and
4.5 cm. broad. The costa is minutely elevated above and conspicuously so
beneath. The lateral nerves, of which there are 6-8 pairs, diverge at an
angle of about 45°, and bear more or less inconspicuous axillary glands.
The inflorescence is axillary, paniculate, rather few-flowered, densely but
closely pubescent, up to 6.5 cm. long, the glabrescent flowers up to 3 mm.
long, ii agra acter A the pedicel 2-4 mm. long. The ovate acutish
lobes are fleshy papillose vino + 2.15 mm. long. The stamens of
the two outer series are + 1.25 m n. long, and the anthers are ovate, trun-
cate, almost equalled by the stout pene Those of the inner series are
oblong, the filaments covered with conspicuous sessile glands. The slender
(undeveloped, or maybe this is the staminate flower of a dioecious species )
glabrous gynaecium is + 2.15 mm. long, the narrow ovary blending imper-
ceptibly into the stout style that bears a conspicuous peltate stigma. Th
fruit is pointedly ovoid, black, up to 2.5 cm. long and 1.5 cm. broad near
the base, supported by a flat almost disk-like woody cupule separated from
the fleshy fruit, at least on drying, about 1.5 cm. in diameter and slightly
undulate. The enlarged pedicel (as well as the cupule) is pale brownish-
buff, deeply furrowed and verruculose, measuring up to 1 cm. in length an
5 mm. in diameter throughout.
The relationship of the species to O. eucuneata is discussed under that
species.
24. pairs Klotzschiana (Nees) ngrees Biol. Centr. Am. Bot. 3: 73. 1882; Mez in
Bot. Gart. Berlin 5: 273.
eas Klotzschiana Nees in ican 21: 523. 1848.
DistRIBUTION: Known only from Mexico.
1945 | ALLEN, STUDIES IN THE LAURACEAE, VI 355
Mexico: State unknown, “Bei El Banco,” Ehrenberg 943 (fl., fragm. of TyPE, Ch) ;
Hartweg 383 (fl., cited by Mez, photo., Ch).
This entity (a tree according to Ehrenberg; a shrub 2—3 m. high accord-
ing to Hartweg) has terete branchlets that are sparsely pilose near the apex,
auc becoming glabrous and black-brown. ‘The petioles are up to 15
long and canaliculate. From the fragment the blades appear to be
able elliptic (or in the photo. lanceolate or elliptic- lanceolate), the base
cuneate, the apex acuminate (long-acuminate in the photo.). They are
chartaceous, shining above, and everywhere very prominently and coarsely
reticulate, measuring up to 9 cm. long and 2.8 cm. broad. ‘The costa is
conspicuous though not elevated above, and very prominent beneath. The
lateral nerves are obscured by the prominent reticulation but appear to
number 4-6 pairs, diverging at an angle between 30° and 50° and bearing
beneath conspicuous axillary pubescent glands. The axillary paniculate
many-flowered inflorescence, up to 5 cm. long, is described as pubescent,
and the peduncle is 1 cm. or ‘less in length. The flowers are about 2.5 mm.
long and pubescent, the slender pubescent pedicel not more than 2—3 mm.
long. The thinnish ovate acutish lobes are + 2.15 mm. long. The stamens
of the two outer series are + 1 mm. long, the anthers are almost rectangu-
lar-ovate and twice the length of the filaments. Those of the inner series
are + 1.7 mm. long, the almost equal filaments bearing conspicuous ovate
glands that are stipitate and almost the len oe of the filaments. The
staminodia are linear, acuminate, pubescent, + 0.6 mm. long. The gla-
brous gynaecium is + 2.25 mm. long, the subglobose- ovoid ovary topped
by a ae style of equal length which flares into a discoid very conspicu-
ous stigm The fruit is unknown at present.
This littie collected and little known species is one of the smallest-leaved
species found in this area, sharing this character with O. effusa, from south-
ern Mexico, adjacent Guatemala, and British Honduras. In addition it is
outstanding because of its shining prominently reticulate leaf-blades, which
are chartaceous.
. Ocotea effusa leauge Hemsley, Biol. Centr. Am. Bot. 3: 73. 1882; Standley
. Herb. 23: 296. 1922; Lundell in Contr. Univ. Mich. Herb.
6:15. 1941.
Oreodaphne effusa Meissner in DC. Prodr. 151: 120. 1864.
DISTRIBUTION: Southern Mexico, Guatemala, and British Honduras, in forests.
x Oaxaca: Near San Pedro Nolasco, Talea, etc., 1843-44, Juergensen
906 (fl., fragm. of Type, Ch, photo., Ch, NY). (Probably Oaxaca): Monte Mistan,
184-, Galeotti sn. (fl, Ch, GH, US). Chiapas: Damp forest, mountains east of
Fenix, Purpus 10309 (fl., NY), 10690 (fl, GH). Guatemata: Huehuetenango:
Between Ixcan and Rio Ixcan, Sierra de los Cuchumatanes, Steyermark 49373 (fr., Ch).
British Honpuras: Stann Creek: In high ridge on hillside, Middlesex, Gentle
2926 (fr., A, NY); Mountain Cow Ridge, Gentle 3266 (fi., A, NY
This slender dainty-leaved species has slender terete dpscutay striate
branchlets that are early sericeous but quickly lose their pubescence, be-
coming glabrous almost at once. The petioles are delicate, canaliculate,
and attenuately cuneate, the apex attenuately and obtusely caudate-acumi-
The costa is slightly but conspicuously elevated above and beneath.
356 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XXVI
The numerous pairs of lateral nerves are so obscure that it is difficult to
count them. The reticulation is not visible above and although present
beneath is very obscure. The tenuous weak inflorescence is axillary o
subterminal, paniculate, glabrous, few-flowered, up to about 12 cm. long,
and the peduncle is up to 5.5 cm. he tenuous flowers are about 3 mm,
long subtended by filamentous pedicels that are sometimes as long as 5 mm.
The perianth- -tube is well developed, and the ovate acutish lobes are thin in
texture, + 1.7 mm. long. The stamens of the two outer series are + 1.25
mm. long, the Ravn ovate roundish anthers nearly twice the length of
= slender sometimes pubescent filaments. Those of the inner series are
1.7 mm. long, the anthers more slender, and with the subequal filaments
wee. two distinct subglobose ecu glands at the base. The staminodia
are linear, pubescent, and + 0.7 mm. long. The glabrous gynaecium is
+ 1.7 mm. long, the ellipsoid- gst oe ovary about equalling the at-
tenuate te bearing a subcapitate obtuse stigma. ‘The fruit is ellipsoid,
apiculate, up to 14 mm. long and 9 mm. broad, subtended by a shallow
—s cupule up to 4 mm. long, 6 mm, in diameter, and not more than 1.5
m. deep, the margin only slightly undulate. The pedicel is almost 1 cm.
ae expanded at the apex to 4 mm. in diameter.
e slender aspect of the eilewaues and the dainty thread-like pedicels
of the small flowers of this species separate it immediately from the only
other near relative, O. Klotzschiana. Also the leaf-blades, with prominent
costa above and the reticulation very obscure, are in contrast to the con-
spicuous reticulation of O. Klotzschiana throughout.
26. Ocotea Bernoulliana Mez in Jahrb. Bot. Gart. Berlin 5: 275. 1889.
DistripuTION: Guatemala, Honduras, and British Honduras, up to 2000 m. altitude.
GuaTEMALA: Huehuetenango: Cerro Chiblac, between Finca Rafael and
Ixcan, Sierra de los Cuchumatanes, Stevermark 49/81 (fr. Ch). Alta Verapaz:
Damp forest, region of Chelac, northeast of Carcha, Standley 70367 (fl., Ch), 70523
(fl., Ch). Quezaltenango?: Near Mujulia (Majulia), Bernoulli & Cario 2590 (fi.
fr., TYPE not seen). Honpuras: Yoro: oe Progreso, Farm 42, Hottle 83 (fl,
Ch); C. & V. W. von Hagen 1118 (fr., Ch, NY). Atlantida: Lancetilla Valley,
near Tela, Standley 56721 (fl., Ch). Britrisu esewiuia: eee Walk: Roaring
Creek, Lundell 325, 370 (fr., Ch). Stann Creek: In high ridge on hillside, Big
Eddy Ridge, Gentle 3351 (fr., A, NY); Middlesex, along river-bank, Schipp 399 (ir.,
GH, NY). Toledo: Eldorado, Schipp 398 (fr., ra Silkgrass Reserve, Hope Creek
Road, '4 m. n. Silkgrass Camp, Stevenson 167 (fr., Ch).
“Aguacatillo” (Honduras); “Canoj’” (Guatemala); ‘‘Laurel,”
A Piaharsipeet” (Br.tish Honduras
The present species has dark reddish brown glabrous branchlets that are
rather slender and eae The slight petioles are dark, canaliculate, and
glabrous, measurin o 1.5 cm. long. The blades are membranaceous,
rarely subcoriaceous, pane usually obiong-elliptic, the narrowly
cuneate or seein roundish and abruptly cuneate at the very base,
the apex usually caudate-acuminate, 13 (—18) cm. long and °s (-7) cm.
2
broad. The costa is slightly elevated above and conspicuous, more promi-
(—8 rarely) pairs, are slightly elevated above, more conspicuously so be-
neath, the upper pairs diverging at an angle of 25-35°, the lower rae 45°.
The minute and regular reticulation, although very obvious above,
more conspicuous beneath. The inflorescence is glabrous, axillary, ae
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 357
ing of usually not more than one (occasionally three) few-flowered panicles,
not more than 10 cm., borne on long slender peduncles not more than 6.5
cm. long. The small flowers are perfect, glabrous, and up to 3 mm. long
neon to Mez (the ones at hand, being in bud, do not measure more
than 2.15 mm.). The perianth- tube is conspicuous, the thickish lobes
nae or ae + 2.15 mm. long. The stamens of the two outer series
+ 1mm. ee with long ovate obtuse nearly sessile anthers. Those o
the inner series are + 1.25 mm. long, with rectangular or square anthers
borne on short filaments about one-half their length and having at the base
two short-stipitate cordate glands equalling the filaments in length. The
staminodia, when present, are attenuately linear and pubescent, + 0.6 mm.
long. The labrous gynaecium is + 1.7 mm. long, the globose ovary
equalling the rather sturdy style topped by a triangular- peltate stigma.
The fruit is practically identical with that of the preceding species, which
see for a discussion of relationships.
27. Ocotea tenera Mez & J. D. Smith ex Mez in neta Herb. Boiss. II. 3: 234. 1903;
Standley in Field Mus. Publ. Bot. 18: 456. 1937.
DistRIBUTION: Known only from Costa ana for the most part from the Atlantic
tierra caliente, at 200 m. altitude along the coast to 1250 m. inland.
Costa Rica: Alajuela: Region of Zarcero, in rm on humid hill-slope of
ears cloud-forest, A. Smith 159 (fl., Ch), 4.228 (9 fl., A, Ch); San Pedro de San
Ramon, Brenes 3617 (153), 4200 (212) (fr., Ch), 4439 (224) (@ fl, Ch). Limon:
Forest of Shiroes, Pate cice, Pittier & Tonduz 9184 (fr., isosyNtyPE, US); along
Banana River (Bananita), near Port Limon, Pittier 3633 (9 fl., Ch); Llanuras de
Santa Clara in the plantations at La Colombiana, Pittier 7607 (Herb. Nat. Costa Rica
13396) (Q fl, Isosyntype, Ch, US). Cartago: Turrialba, alt. 570 m., June, 1895,
Tonduz 8330 (SYNTYPE not seen).
NATIVE NAMES: ‘“Aguacatillo,” “Quisarra lantisco” (lentisco?) (Costa Rica).
This species, also belonging in the O. cernua complex, has dark gla-
brous branchlets that on the whole seem to be somewhat stouter than those
of O. cernua. The petioles are dark, slender, Capalic uate and glabrous,
up to 1 cm. in length. The membranaceous blades are elliptic-oblong or
Senay. elliptic, the base cuneate or broadly so E espneard obtuse or
undish), the apex caudate-acuminate, up to 16 cm. long and usually not
more than 5 cm. broad. The broad costa and dc: lateral nerves, of
which there are usually 6 (—8) pairs diverging arcuately at an angle of
about 45°, are obscure above and elevated beneath. The delicate a
tion is also rather obscure above and prominent beneath. The inflores-
cence is axillary, slender, glabrous, paniculate, usually less than 8 cm. long,
occasionally to 12 cm., the slender peduncle usually not more than 2 cm.
long, occasionally to 4.cm. The flowers, seemingly @ , are glabrous, up to
acutish, and + 1.7 m m. long, thinnish or occasionally thick. The stami-
nodia of the two outer series are + 1 (—2.15) mm. long and usually peal
poorly developed anthers, as do those of the third series, which have small
ovate sessile basal glands. The filaments are usually very broad and as
ong as if not longer than the anthers. The staminodia of the fourth: series,
when present, are scale-like and lanceolate. The glabrous gynaecium is
+ 1.7 mm. long, the ellipsoid ovary narrowing gradually into the tapering
ie that is only about half its length and topped by a conspicuous sub-
capitate stigma. The fruit is 2.5 cm. long and about 1.3 cm. broad. shining-
358 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
black and glabrous, subtended by a flat flaring disk-like undulate-margined
cupule not more than 3 mm. long and 1 cm. in diame yet fia pedicel seem-
ingly up to 1.5 cm. long and expanded to 5 mm. at its
There is a discrepancy between the description of Mez a the specimens
cited here. Also, some of the Costa Rican sheets seem to show flowers that
appear almost perfect. It is possible that examination of the type in con-
nection with the types of the other two species of this group, namely O.
cernua and O. Bernoulliana, may clarify the delimitation of the flowering
phase of these three entities. Certainly the fruit of O. fenera is distinct
from that of the other two species.
28. — —— (Nees) ae = bt Bot. Gart. — 5: 377. 1889; Standley in
. Nat. Herb. . 1922, in op. cit. 27: 183. 1928, in Trop. Woods
- c ron in Field ree ee Bot. 10: 201. 1931; Standley & Record in Field
Mus. Publ. Bot. 12: 143. 1936; Standley in op. cit. 18: 454. 1937; Yuncker in op.
cit. 9: 290. 1940.
Oreodaphne cernua Nees, Syst. Laurin. 424. 1836.
Oreodaphne Sieberi biaeinee in DC. Prodr. 151: 137. ees at least in part.
Ocotea Sieberi Hemsley, Biol. Centr. Am. Bot. 3: 73.
DistrIBUTION: Southern alll Central America, a a Indies, and, according
to Meissner, in South Ameri
exico: Vera Cruz: Sa Coatzacoalcos River, L. Williams 8441 (2 fl,
A, Ch), 8953 (& fl, A, Ch, NY). Tabasco: Teapa, alt. 90 m., May, 1840, Linden
601 (¢ fl, fragm. of syntype of Oreodaphne Sieberi, NY), 1607 (syntype of Oreo-
daphne Sieberi not seen). GuaTEMALA: Without locality (eastern portions of Verapaz
and Chiquimula), Watson 450 (é& fl. GH). Honpuras: Atlantida: In open
forest, foothills near the Cangrejal River, Yuncker, Koepper & Wagner 8788 (fr., Ch,
NY). British Honpuras: Stann Creek: On hillside, Stann Creek Valley, 15
‘esti Gentle 2120 (fr., A, NY); in high ridge, Middlesex, Gentle 2767 (¢ fl., A, NY);
Big Eddy Ridge, Gentle 3310 (¢ fl, A, NY); Stann Creek Railway, 15 mile, Schipp 161
(é fl, Ch, GH, NY). Costa Rua Puntarenas: Playa Blanca, Golfo Dulce,
Valerio 397 (& fl., Ch). Alajuela: Guadaloupe de Zarcero, A. Smith A.556 (6
. San José: Forests of Las Vueltas, Tucurrique, Tonduz 13366 (2 fl., GH,
NY); vicinity of El General, Skutch 4270 (é fl., A, NY), 4738 (¢ fl, A, Ch).
Cartago: Tuis, near Turrialba, Pittier 11251 (2 fl, GH). PANAMA: ree del
Toro: Changuinola Valley, Cooper & Slater 105 (Y 10286) (4 fl., Ch, GH, eee
Island _ Dunlap 568 (4 fl., Ch, GH); Water Valley, von Wedel 803 ({ir.,
Mo). Chiriqui: Without locality, Cooper & Slater 262 (Y 10615) (fr., Y). ooo
Barro Colorado Island, Shannon Trail, — 535 (fr., Ch), Drayton Point,
Shatiuck 1140 (fr., Ch) ; shore of cove s. from Lock site, Woodworth & Vestal 471 (6
Darién: Along the Sambu River, brian tide-limit, Pittier 5692 (3
Native NAMEs: “Aguacatillo” (Mexico, Honduras, British Honduras); “Laurel”
(Tabasco); ‘“Laureo de bajo” (Campeche); “Sigua,” “Sigua amarillo” (Panama) ;
“Timbersweet” (British Honduras).
This species has branchlets which are early pubescent but quickly gla-
brous, angled or striate, terete. The petioles of the leaves are slender,
canaliculate, glabrous, up to 12 mm. long. The blades are chartaceous or
subcoriaceous, glabrous da, usually oblong-elliptic, the base, gen-
erally spe eaking, roundish or obtuse with the extreme portion abruptly
eh the apex abruptly or gradually acuminate, usually caudate-acumi-
nate. They measure up to 16 cm. long and 6.5 cm. broad. ‘The costa is
conspicuous above, although not as prominently elevated as beneath. The
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 359
speared etched lateral nerves, of which there are 4—6 pairs, are obscure
ove but more prominent beneath, the lowermost pairs usually being the
net The lowermost pairs diverge at an angle of about 25—35° from
the costa, and the upper at an angle of about 45°. The reticulation is
obscure above and less so beneath. The inflorescence consists of numerous
The glabrous ¢ flowers are yellowish, very small, not more than 2 mm.
long at most, and ee on slender sometimes thread-like pedicels up to 4
mm. long. The thin perianth-lobes are ovate, obtusish or acute, + 1.7 mm.
long. The stamens of the two outer series are + 0.8 mm. long, the ovate-
triangular obtuse anthers almost sessile. Those of the inner series are
more narrowly ovate and borne on short filaments which have two basal
suborbicular compressed glands that are sessile. The glabrous aborted
gynaecium is linear, not measuring more than 0.8 mm, in length. The ¢
flowers superficially seem very much like the ¢ flowers. The stamens,
however, are much smaller and seemingly sterile (not more than 0.8 mm
long), ie of the inner series biglandular. The glabrous gynaecium is
m. long, the ellipsoid-subglobose ovary attenuate into an extremely
brief style which bears a conspicuously three-parted stigma. The fruit is
woody cupule that is about 6 mm. long, 11 mm. in diameter, and 3-4 mm
dee The supporting pedicel is up to 7 mm. long and expanded at the
tip to 2 mm. in diameter
Neither Meissner’s syntypes (except Linden 1601) of Oreodaphne Sieberi
nor Nees’ syntypes are available, and so for the present Mez’ interpretation
will be accepted, and O. cernua considered as a widespread species extend-
ing to the West Indies and South America.
The two nearest relatives seem to be O. Bernoulliana, with perfect
flowers, from Guatemala, which is so similar that one suspects that it may
be a different manifestation of O. cernua with dioecious flowers; and O.
tenera, which also is dioecious, but whose fruits are larger and subtended
by a flat shallow cupule with an undulate margin.
29. ai paradoxa Mez in is Jahrb. 30: Beibl. 67: 16. 1901; Standley in Field
. Publ. Bot. 18: 455. 19
Paes ats Known only oe the type-collection, from Costa Rica.
Costa Rica: Alajuela: In woods near River Naranjo, alt. 200-300 m., Tonduz
(Herb. Inst. Costa Rica) 7648 (fl., TyPE not seen).
Mez described the species as a tree or shrub with the branchlets slightly
ferruginous-tomentellous at the apex, becoming quickly glabrate, brown,
terete, with ferruginous- or yellowish-tomentellous buds, the cortex slightly
aromatic. The petioles are up to 10 mm. long, incised-canaliculate, gla-
brous, supporting sparse, glabrous, chartaceous leaves entirely glabrous in
the adult stage and dull opaque olivaceous-green, elliptic, the base short-
or long-acute, the apex shortly or sometimes very shortly acuminate, meas-
uring up to 13 cm. long and 6.5 cm. broad, with an almost plane margin.
The venation is almost smooth or minutely prominulous-reticulate above
and manifestly prominulous-reticulate beneath. The inflorescence is loose,
ne flowered, subcinereous- or subferruginous-tomentellous. ‘The flowers
e 1.2 mm. in diameter, Rone! on pedicels 5-10 mm. long, the bracts de-
360 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
ciduous. The lobes spread like those of Nectandra, and are ligulate, nar-
rowing gradually toward the apex, finally becoming roundish. The stamens
have foliaceous anthers which are acute, the connective long-produced be-
tween the cells. Those of the inner series have shortly stipitate pulvinate
glands that are for a conspicuously long distance connate with the filaments.
The staminodia are aborted; the ovary is ovoid, drawn out into a short
conical style; the stigma is large, subdiscoid. The fruit is unknown.
Mez notes that this species is to be put in the subgenus near O. Dendro-
daphne and O. veraguensis, conspicuously differing in entire habit, few-
flowered and racemose inflorescence, and floral glands. It keys out to be
near the last three discussed species above, but nonetheless, from the de-
scription it would seem to be a variant of O. veraguensis,
30. Ocotea Matudai Lundell in Bull. Torr. Bot. Club 69: 388. 1942.
DIstTRIBUTION: Known only from type-locality.
Mexico: Chiapas: Mt. Ovando, western slope near Escuintla, alt. 1800 m., July
1-16, 1940, Matuda 4221 (fl., A, type — Mich, NY, US) (tree, 7 m.; flowers white).
Another species known only from the type-collection, with very slender
branchlets early angled, glabrescent, and reddish, presently terete, striate,
glabrous, and brownish or maculate-reddish gray. The petioles are dark
reddish, slender, canaliculate, glabrous, and up to 2.cm. long. The blades
glabrous and shining above, paler beneath, subcoriaceous, oblong-elliptic or
oblong-lanceolate, the base obtuse to cuneate, the apex obtuse or obtusely
acute, to 15 cm. long and up to 6 cm. broad. The costa is impressed above
and elevated beneath. The lateral nerves, of which there are 6-9 pairs,
are slightly elevated above and very prominently though slenderly so be-
neath, diverging at an angle of about 45—55° and bearing axillary pubescent
glands beneath that are fairly conspicuous. The reticulation is conspicu-
ous above and beneath. The inflorescence is axillary and subterminal,
paniculate, glabrous to glabrescent, many-flowered, u
usually branched to the base, at most the peduncle not more than 1 cm.
long. The pubescent flowers measure up to 3 mm. long, spreading to 7 mm.
in diameter, borne on slender pedicels up to 4 mm. The elliptic or oblong
lobes are up to 3 mm. long, pubescent within and without. The stamens
£
=]
o
=
ge
stigma.
The species-relationships are discussed under O. laetivirens.
31. Ocotea Meziana, sp. nov.
Arbor 7-30 m, alta, ramulis viridescentibus glabris striatis angulatis.
Folia alternata, juventute canescenti-sericeo-pubescentia subito glabra,
petiolis gracilibus glabris leviter canaliculatis ad 1.3 cm. ongis et 1 mm.
Jatis, laminis utrinque glabris chartaceis, in sicco viridescentibus ellipticis,
ad 15 cm. longis et 5 cm. latis, basi attenuato-cuneatis, apice abrupte acumi-
natis vel plus minusve acutis vel obtuse acutis. penninerviis, costa utrinque
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 361
inconspicue et leviter elevata, nervis 4—6-paribus utrinque inconspicue et
leviter elevatis angulo 35-45° divergentibus, rete venularum supra sa is
subtus minute perprominulo. Inflorescentia axillaris, paniculata, ad 9 cm.
antheris subreniformibus stipitatis subaequalibus; staminodiis nullis vel
linearibus pubescentibus ad + 1 mm. longis; gynaecio glabro ad + 2.4
mm. longo, ovario gynaecii dimidium longitudine parum excedente, stig-
DistripuTion: In the upper tropical or Caribbean rain- or cloud-forest of Costa
Rica, at an altitude varying from 825 to 2100 m
Costa Rica: Alajuela: La Pefia de Zarcero, A. Smith H.309 (fl., Ch); Zarcero,
edge of woodland, in semi-shade, in stiff clay-loam, upper tropical zone, continental
divide, alt. 1615 m., A. Smith H.359 (fl., TYPE, Ch) (tree 10.5 m., base 34 cm., bark
brown, slightly roughened; leaves thin but firm, dark green above, glabrous, shining;
flowers cup-shaped, oil-yellow), A. Smith H.407 (fl., Ch); La Brisa de Zarcero, A.
Smith H.443 (fl., Ch); Villa Quesada, San Carlos, A. Smith F.1774 (fl., Ch), P.2596
(fl., A); Tapeseo, Alfaro Ruiz, A. Smith P.2615 (fl., A). Heredia: Vicinity of Vara
Blanca, north slope of Central Cordillera, between Poas and Barba volcanoes, Skutch
3745 (fi., A).
This species is, as mentioned above, related to O. laetivirens.
32. Ocotea pyramidata Blake ex T. S. Brandegee in Univ. Calif. Publ. Bot. 7: 326. 1920.
DistripuTioN: Known only from the type-locality in Mexico.
Mexico: Vera Cruz: Zacuapan, Nov., 1919, Purpus 8456 (fl., ISOTYPE, GH,
NY) (large tree).
The present species has branchlets dark reddish black, angled, deeply
sulcate, early covered with a thin ferruginous tomentum which quickly
disappears, leaving them entirely glabrous. The leaves are supported by
dark slender glabrescent petioles up to 2.5 cm. long. The blades, unfor-
~—
delicately traced above and elevated beneath, diverging at an angle of about
2 ae e very fine pattern of the minute reticulation is present but not
conspicuous throughout the blade. The numerous axillary and subterminal
racemose-paniculate inflorescences are many-flowered, subferruginous-
tomentose, shortly becoming glabrescent, up to 8 (—12) cm. long, the short
peduncle not more than 1 cm. long. The glabrescent to glabrous flowers
are up to 4 cm. long, subcampanulate, and subtended by a slender glabres-
cent pedicel not 2 mm. long. The perianth-lobes are up to 3 mm. long,
very thin, elliptic, obtuse or rounded. The stamens of the two outer series
are 5 mm. long, the ovate subrotund anthers twice the length of the
362 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
stout filaments. Those of the inner series are + 2.15 mm. long, the sub-
rectangular anthers equalled by the filaments, bearing two subreniform
spreading subsessile glands that are almost their entire length. The aborted
ovary is very narrowly lanceolate, up to + 0.5 mm. long and topped by a
subcapitate stigma.
lake did not relate this species to any known from Mexico. It is very
possible that some of the collections later described from this region may be
conspecific, but it is difficult to match diseased foliage such as is found in
the type of this species.
33. Ocotea laetivirens Standley & Steyermark in Field Mus. Publ. Bot. 23: 114. 1944.
Distripution: Guatemala, and the forests of Costa Rica, at altitudes up to 2000 m.
ATEMALA: Huehuetenango: Cerro Chiblac, between Finca San Rafael and
Ixcan, Sierra de los Cuchumatanes, alt. 1200-2000 m., July, 1942, Steyermark 49189
(fl, type, Ch) (vine; leaves membranous, deep green above, paler beneath), 49378
(fl., fr., Ch). Costa Rica: Alajuela: Region of Zarcero, in forest, A. Smith H.517
(fl., Ch) (clay-loam, in forest below oak-level in shade) ; Guadaloupe de Alfaro Ruiz,
A. Smith 4182 (abnormal fl., Ch).
This species has the young branchlets early pubescent but very quickly
glabrous, conspicuously greenish, striate and angled. The leaves are sup-
ported by stout or slender (in the type) petioles 1-1.7 mm. long, often yel-
lowish, sometimes dark, canaliculate. The blades are chartaceous, con-
spicuously greenish, elliptic or oblong-elliptic, the base cuneate, the apex
acuminate or acutish or obtuse, usually 16 (—18) cm. long and 7 cm. broad.
The costa is rather conspicuous and yellowish above, although slightly ele-
vated, but beneath is prominently elevated. The lateral nerves, of which
there are 8 or 9 pairs, are obscure above but conspicuously prominently ele-
vated and yellowish beneath, diverging at an angle of about 45-55°. The
loose reticulation is rather obscure throughout. The inflorescence is axil-
lary, paniculate, many-flowered, early finely and inconspicuously grayish-
pubescent, becoming glabrous, about 15 cm. long and borne on a peduncle
up to9 cm. long. The flowers are pubescent to glabrescent, 2—3 mm. long,
the pedicels not more than 4 mm. long. The perianth-lobes are elliptic to
ovate, obtusish, thick, + 2.5 mm. long. The stamens of the two outer
series are + 1.7 mm. long, the broadly ovate anthers obtuse or rounded and
not quite twice the length of the stout filaments. Those of the inner series
have anthers that are more narrowly ovate, the connectives rounded an
slightly projecting, the almost equal filaments bearing conspicuous globular-
squarish glands about one-third the length of the entire stamen. The
staminodia are ++ 0.8 mm. long, very slender, attenuately linear-lanceolate,
and pubescent. The glabrous gynaecium is + 2.5 mm. long, the ellipsoid
ovary nearly twice the length of the slender style that bears a small capitate
stigma. The fruit is unknown, but the cupule attached to Steyermark
49378 is woody (rose-red according to the collector), rather thinnish, cam-
panulate, and up to 5 mm. long and 1.2 cm. in diameter, the pedicel grooved
and expanded to 1.5 cm. long and 6 mm. in diameter at the apex. Bot
the cupule and the pedicel are brownish-verruculose.
The species, like the preceding, seems to have no near relatives, although
it has characteristics in common with O. Matudai and O. Meziana. It is
easily separated from both because of foliage-characters and the habit of
the branchlets.
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 363
DouBTFUL SPECIES AND VARIETIES OF OCOTEA
Ocotea insularis (Meissner) Mez in Jahrb. Bot. Gart. Berlin 5: 271. 1889; Standley in
Field Mus. Publ. Bot. 18: 455. 1937.
Phoebe insularis Meissner in DC. Prodr. 151: 33. 1864.
DistrisuTION: Cocos Island, according to Pittier, characteristic of the forests of
the northern and western part of the island, at an altitude of 10-200 m.
Costa Rr cos Island: Without data, Menzier (fi., fr., TYPE not seen),
Pittier 16257 (fr., GH); Wafer Bay, Howell 10185 (fl., Ch).
This species, which seemingly inhabits only Cocos Island, has stout red-
dish brownish glabrous branchlets that are pronouncedly alate and striate.
The petioles are about 5 mm. long, stout, canaliculate, and glabrous. The
blades are subcoriaceous or chartaceous, glabrous except for the pubescent
axillary glands, obovate, the base cuneate and decurrent, forming apparent
petioles sometimes up to 1 cm. longer than the actual petioles, the apex very
slightly and abruptly obtusely subacuminate. They measure up to 19 cm
long and 7 cm. broad, the stout costa deeply impressed above, especially
near the base, and conspicuously elevated beneath. ‘The lateral nerves, o
which there are about 9 pairs, are impressed and very slightly elevated
above and more conspicuously elevated beneath, diverging at an angle of
35-40°. The reticulation is fairly conspicuous on the upper surface and
more prominently so on the lower. The inflorescence is axillary and sub-
terminal, paniculate, many-flowered, and, according to Mez, scarcely fer-
ruginous-tomentellous, up to 10 cm. long, the glabrous alate striate peduncle
up to 2.5 cm. long. The flowers are 3 mm. long, minutely and sparsely
pubescent, the short pedicel + 2.15 mm. long. The tube is conspicuous
and the thinnish ovate acute lobes are + 2.15 mm. long. The stamens of
the two outer series are + 1.7 mm. long, the ovate obtuse or roundish
anthers shorter than the broad pubescent filaments. Those of the inner
series are -+ 1.9 mm. long, the anthers subrectangular, the filaments bear-
ing sessile subglobose glands at the base. The glabrous gynaecium is
+ 2.4 mm. long, the subglobose (subobovoid) ovary only slightly longer
than the cylindrical style with a rather conspicuous discoid (obtuse) stigma.
The fruit is ellipsoid, glabrous, 13 cm. long and 6 cm. broad, and irregularly
subtended by a 6-lobed rugose cupule about 7 mm. long, 9 mm. in diameter,
and about 5 mm. deep, the subalate pedicel being 5 mm. long and expanded
to 3.5 mm. diameter at the apex.
The affinity of this species seems to be with the group containing O. /ra,
O. Tonduzii, O. Endresiana, etc., but the narrowly decurrent recurved
blades, the variation in size and texture of the blades, among other char-
acteristics, segregate the species. It is not included in the key, for the flora
of Cocos Island is quite distinct from that of the mainland.
Ocotea psychotrioides Billberg ex Meissner in DC. Prodr. 151: 150. 1864.
This species is treated as a synonym of Nectandra globosa by Mez.
Along with the type, Billberg 321, from Costa Rica, he also cites Oersted
16, 17, and 18, also from Costa Rica, which numbers may very well be
syntypes of Nectandra amazonum var. 8 Oerstedii. None of these have
been seen for study.
Ocotea puberula Nees var. truncata (Meissner) Mez in Jahrb. Bot. Gart. Berlin 5: 344.
Strychnodaphne puberula var. Y ? truncata Meissner in DC. Prodr. 151: 143. 1864.
364 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
Ocotea puberula Nees, Syst. Laurin. 472. 1836, p.p.; Standley in Contr. U. S. Nat.
Herb. 23: 295. 1922.
DistrIBUTION: Known only from the type-locality.
eExico: Vera Cruz: Without locality, a 1142 (syntype of Ocotea pu-
berula, and cited by Mez under the variety, not s
Ocotea sanguinea J. S. Presl, [Wseobecny] Rostl. 2: 60. [1846].
This species, according to Mez, is synonymous with Nectandra sanguinea.
Index Kewensis follows Mez and lists it also from Mexico and the West
Indies. I have supplied the bracketed portion of the citation. The
volume in question does not contain Ocotea sanguinea, although two other
Ocoteae are mentioned. Possibly the following reference is meant, even
though the entity to which Mez has referred Ocotea sanguinea does not
occur in Bohemia, the locality taken up in Berchtold and Presl: Berchtold,
F. G, von, & Presl, J. S., Prirez. Rostlin, 3 vols. 195. pl. 1823 [-35].
SPECIES AND VARIETIES EXCLUDED FROM QOCOTEA
Ocotea Brenesii Standley = NECTANDRA sp.
Ocotea campechiana Standley = LicariA CAMPECHIANA (Standley) Kostermans.
Ocotea Cufodontisii O. C. Schmidt = NECTANDRA sp.
Ocotea globosa Schlechtendal & ee NECTANDRA GLABRESCENS Bentham
Ocotea helicterifolia (Meissner) Hemsley = PHOEBE HELICTERIFOLIA (Meissner) Mez.
Ocotea latifolia H.B.K. = Nectanpra LATIFOLIA (H.B.K.) Mez.
Ocotea mexicana (Meissner) Hemsley = PHOEBE HELICTERIFOLIA (Meissner) Mez.
Ocotea mexicana var. a subsessilis (Meissner) Hemsley = PHOEBE HELICTERIFOLIA
Ocotea mexicana var. 8 ? longipes (Meissner) Hemsley = PHOEBE HELICTERIFOLIA
Ocotea mexicana var. Y diminuta (Meissner) Hemsley = PHOEBE HELICTERIFOLIA
(Meissner) Mez.
Ocotea perseifolia Mez & J. D. Smith = NectTan sp.
Ocotea psychotrioides H.B.K. = PHorse psycuotrioieEs (H.B.K.) Mez:.
Ocotea rubriflora Mez = Sieeraswns sp.
Ocotea salicifolia H.B.K. = NECTANDRA GLABRESCENS Bentham.
Ocotea Salvini Mez = Daskey SALVINI (Mez) Lundell.
Ocotea striata Buck = Myropia cf. FUNEBRIS L. gp Giirke), fide Mez
Ocotea subtriplinervia (Meissner) Hemsley = PHOEBE SUBTRIPLINERVIA (Meissner)
Standley.
Ocotea tampicensis (Meissner) Hemsley = PHOEBE TAMPICENSIS (Meissner) Mez.
Ocotea Whitet Woodson = NECTANDRA Sp.
(To be concluded)
JOURNAL
OF THE
ARNOLD ARBORETUM
VoLt. XXVI OCTOBER, 1945 NuMBER 4
STUDIES IN THE LAURACEAE, VI
PRELIMINARY SURVEY OF THE MEXICAN AND
CENTRAL AMERICAN SPECIES
CAROLINE K. ALLEN
Concluded from page 364
4. Nectandra Rolander
Bact Rolander ex Rottboell in Act. Univ. Hafn. 1: 267. 1778; Grisebach, FI. Brit.
. 281. 1860; Meissner in DC. Prodr. 151: 146. 1864; es Biol. Centr.
ie Bot. 3: 74. 1882: Mez in Jahrb. Bot. Gart. Berlin 5: 393.
DistriBuTion: Tropics of America, the bulk of the species occurring in South
America, particularly in the Andes, with about 38 in Central America and Mexico, and
a few in the West Indies, the fringe of the latter just touching the mainland of Florida.
The genus consists of trees or shrubs with various types of glabrous or
pubescent foliage. The alternate leaves have blades that are membrana-
ceous to rigidly coriaceous, lanceolate, elliptic, or obovate, often with
variously expanded and recurved leaf-bases. The blades are usually
penninerved, rarely subtriplinerved, the reticulation being obscure to
extremely conspicuous. Pubescent axillary glands are frequently con-
spicuous on the lower surface. The inflorescences are usually paniculate,
axillary or subterminal, with peduncles of varying length, the bracts
deciduous, with the pubescence variable. The flowers are always perfect
in the species found in this area. The tube may be conspicuous or almost
entirely lacking. The equal lobes are lanceolate to elliptic, ovate or
occasionally obovate, usually fleshy and papillose, occasionally membrana-
ceous, almost always spreading or reflexed at anthesis, and usually
deciduous. The stamens of the outer series are either fleshy, petaloid,
papillose, and ovate, or quadrate or orbicular, with conspicuous connective-
tissue, or they are reniform or subreniform, frequently emarginate, with
no apparent connective tissue, the cells occupying the entire anther. The
four cells are always introrse and usually are arranged in an arc-like forma-
tion, very rarely this arc is obscure. The anthers are sessile or borne on
filaments of varying length and thickness, often pubescent, particularly at
the base. The stamens of the third or inner series are usually quadrate;
366 JOURNAL OF THE ARNOLD ARBORETUM [ VOL, XXVI
in the flowers bearing the petaloid outer series, the inner also may be fleshy
and papillose, with truncate connectives; in other cases, they are not fleshy
or papillose and the connectives are inconspicuous. The four cells are
arranged in two horizontal planes; those of the upper plane are lateral or
laterally extrorse; those of the lower plane are usually extrorse.
staminodia when present are for the most part stipe-like; occasionally they
bear well developed heads of varying shape. The gynaecium is, except in
a very few instances, com letely glabrous, the ovary globose or depressed-
The
oO
sionally may be even the length of the ovary, and bears a discoid or
triangular or occasionally a peltate conspicuous stigma. The fruit is
ellipsoid, globose, or oblong, borne in a more or less shallow usually woody
cupule formed by the enlarged perianth-tube usually with simple margin,
occasionally bearing the remnants of the perianth-lobes. This is uniformly
subtended by an enlarged pedicel.
KEY TO THE SPECIES OF NECTANDRA
A. Leaf-blades not recurved at all at the base, or decurrent, not auriculate or cordate
or even rounded at the base generally
B. Largest leaf-blades never more than 9 cm. long.
C. Largest leaf-blades up to 7 cm. long and 2 cm. broad, lanceolate-elliptic, var-
nished-shining above, the reticulation somewhat obscure..1. N. Davidsoniana.
OF ri leaf-blades up to 9 cm. long and 3.5 cm. broad, elliptic, not varnished-
above, the reticulation gd oe SN ee er eee 2. N. Smithii.
B. Largest a blades never less than
C. Anthers of the two outer series i erwin fleshy, petaloid, papillose, never
emarginate, the upper third consisting of connective tissue, the remaining
space occupied by the cells.
D. Anthers of two outer series ovate or quadrate, the cells not forming a per-
fect arc; leaves membranaceous.
E. Anthers of the two outer series definitely ovate; largest leaf- — not
more than 14 (usually 12) cm. long; lateral nerves 4 or 5 pairs........
ET Pe ee a ee ee ee ee 3. N. Br renesit.
ern less than 15 (up to 26) cm. long; lateral saci 7-12 pairs.
F. Leaf-blades shining above, densely and prominently reticulate
throughout, onan the base cuneate.......+.s.s0> 4. N. —
F. Leaf-blades what shining seen with loose very obscure re-
ticulation lett elliptic, oblong-elliptic or ovate- elliptic, the
e roundish or obtuse penn Pe PaO ere 5. . rubriflora.
, the cells
o
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E. Young branchlets, lower surface of igh des, and_ inflorescence
densely and conspicuously ferruginous-tomentose; leaf- — elliptic.
SOSH Moie Mais e bateege eae anes Sie bla aie eg 4 Weise ang Galese beak . Schippii.
E. Young branchlets, lower surface of leaf-blades, and Rei ce not
densely and conspicuously ferruginous-tomentose.
F. Young branchlets, young leaves, and inflorescence densely fulvous-
or brownish-tomentose; blades obovate; anthers of two : series
truncate; cupule hawisphereal ipaawneesotewewoneat 7. N. Austiniz.
F. Young brwatbletie, young vied ei or inflorescence mt densely
fulvous- or brownish-tomentose; blades not obovate; anthers of
two outer series not truncate; pi peice
1945 ]
ALLEN, STUDIES IN THE LAURACEAE, VI 367
G. Leaf-blades pale green, usually with the venation and reticula-
tion showing whitish, often with large ellipsoid axillary pubes-
ae glands on the lower surface, but conspicuous on pc sur-
8. N. menstis.
HACOSH Mp tstsre A occ ef ovee arate wins onshoyece Sek eet enens
G. eye blades not pale green with conspicuous whitish venation
and reticulation; axillary glands if present sae inconspicuous.
H. Leaf-blades with lateral nerves 8-12 pairs..... 9. N. globosa.
H. Leaf-blades with lateral nerves 4-6 (- a pairs
1
J. ramonensis.
ee ae oe ee ee ee
C. Anthers of two outer series of stamens not fleshy, petaloid, or papillose, but
quadrate or reniform, subreniform, or suborbicular, and frequently emargi-
ae the cells occupying the entire anther
. Anthers quadrate, sometimes apiculate...............-: 11. N. Heydeana.
6 Anthers reniform or subreniform or suborbicu
E. Greatest width of leaf-blades at and below ae middle, ee blade taper-
ing only toward the apex............. cess eee cece . Gentle.
E. Greatest width of leaf-blades exactly at the middle, the as tapering
toward the base and the apex equally or the a blades obovate.
F. Largest leaf-blades not less than 7 cm. broa
Pubescence (when present on the bedi “branchlets: lower sur-
face of the leaf, buds, petioles, and inflorescence) predominantly
canescent, not sericeous; petioles not longer ra 1.5—2 cm.
Peder tele UP. inde S18 .6 ke eee Oe ay era
G. Pubescence (when present on young cece leaf-buds,
etioles, and inflorescence) fulvous-sericeous ; gt to 3 cm.
ong; flowers white and fragrant............-- _N. Lundellii.
F. Largest leaf-blades not more than 6.5 cm. ee ee Gras
rarely 8 cm.
G. ees nerves 3—5 (-6) pairs; leaf- a often hae aae he
H. Leaf-blades concen not more tha 15 . lon
ete less, the minate..... 15: i savannarum,
H. Leaf-blades not rors irene 12-14 cm. long, the apex
LONE 2CaUGates eee eeeieree eet cieas 16. N. longicaudata.
G. Lateral nerves not less than 6 pairs, usually 7-9.
lel prereset usually glabrous; leaf-blades usually lanceolate-
rps —14 (-20) cm. long and 5 (-6.5) cm. bro
Leaf- fas not caudate at the apex.
J. Leaf-blades always lanceolate-elliptic, not more than
2-3.5 cm. broad, always long-acuminate at the apex;
reticulation prominent on the lower surface of the
blades.
K. nee rae glaucous above, reddish green beneath ;
frui x 6 mm.; cupule ee BO reese sic cs
vosa.
K. Leaf-blades grayish green tvoughou, or brownish,
paler beneath; fruit 20 cupule not
WELLUCULOSG. sete etecy terse steps! ate . salicina.
Leaf-blades usually elliptic or lanceolate- elliptic, not
less than 4 cm e apex variable; reticulation
prominent throughout.
K. Leaf-blades coriaceous and pale, the ise
obscuring lateral nerves on the upper s
19
ba
K. Leaf-blades euereacan brownish o
reticulation not obscuring as nerves on the
UIPPeL, SUTIACE te ere etes esis sos 20. N. salicifolia.
368 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
. Leaf-blades caudate at = apex...... 21. N. fuscobarbata.
H. Inflorescence variously pubes
branchlets ar or ferruginous-sericeous or
subferruginous-tomentose
J. Reticulation scarcely apparent on upper surface of
leaf-blades.
K. Leaf-blades chartaceous; axillary glands absent;
young branchlets and young leaf-blades golden-
~-tomentose.s .6s464045 042% 22. N. nitida.
K. Leaf-blades coriaceous; axillary glands conspicu-
ously pubescent; young oe >. young
leaves subferruginous-tomentose.. .23. NV. perdubia.
J. Reticulation exceedingly eer ‘these hout.
oblong-elliptic, caudate- acuminate ;
“dealt Blobose.......... 0. ccec ences 24. N. latifolia.
K. rea elliptic, acuminate; fruit peti ellip-
psi Sembee eS Alek welaniata 25. N. Cufodontisii.
I. Young genuihye not golden- or ferruginous-sericeous or
subferruginous-tomentose, sometimes glabrous.
niga and branchlets grayish- or fulvous-
SCTICCOUS). 6.66.00 00d dda dace eae eax 26. N. tabascensis.
T: = MOREE shortly pilose; branchlets oa te Saas
OGG SAGAN CHR GENS ee Ae OeRaeeeee 27. N. Loeseneri.
A. Leaf-blades recurved at the base and decurrent, or auriculate Be recurved, or
cordate and recurve
B. Base of-leaf-blades decurrent and recurved, not auriculate.
C. Largest leaf-blades about 20-22 cm. long, membranaceous or coriaceous.
D. Lateral nerves 8-12 pairs; petiole not thickened............ 8. N.
D. Lateral nerves 4-6 pairs; petiole less than 1 cm. long, only slightly thick-
ene
E. Liat blades not more than 4.5 cm. pes blades ered ovate-
lanceolate, attenuately acuminate at the apex......... entlei.
E. Leaf-blades not less than 5 cm. broad; blades oblong- pk ins
sely acuminate at the apex...................0.. 28. N. Skutchii.
C. Largest leaf-blades not more than 17 (—18) cm., usually less than 15 cm. , Jong,
coriaceous, subcoriaceous, or chartaceous.
D. — chlets, lower leaf-surface, and inflorescence densely woolly ferru-
NOUS-LOMENEOSE:.. 6 ce eee ee ccaccacecsteseueances es N. Schippii.
D. “enter t Ht lower leaf-surface, and inflorescence not densi woolly fer-
ruginous-tomentose.
E. Petioles oe thickened; leaf-blades only slightly decurrent at the base
and very slightly recurved for less than 5 mm,
F. Anthers more or less ovate, obtuse, with fleshy — connec-
tives; fruit ellipsoid.........0...0..0.......20.. 9. N. ramonensis.
. Anthers subreniform, subemarginate ; fruit globose. 29. N. lignan
E. Petioles variously thickened; leaf-blades conspicuously decurrent and
pace ved at the base up to 4—5 cm.
. Largest leaf-blades not more than 5.5, usually 4, cm. broa
aie of leaf- pee dahil and recurved, making an aeueent
iole 4-5 cm. long................000.-005. . N. producta.
G. ee of leaf- “blades narrowed and a sei an apparent
iole not more than 3 cm. long at most......... es N. Whitei.
‘nibiedes usually shining ginies and aoe ner anthers
quarter of the anthers; stigma borne on a well defined style;
cupule and pedicel 2.5 cm. long............. 32. N. hypoglauca.
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 369
G. Leaf-blades not shining, inconspicuously reticulate above;
anthers amp the cells occupying the entire anther;
SEIS INA, SESSLL Co estre-cterersie nk grnce tree cl alaysyevenere) ohemerete Me 33. N. Paulit.
B. Base of leaf-blades cordate or ee conspicuously or sometimes oe re-
curved as well, never decur
C. Leaf-blades oblong- ake or elliptic, rounded, or subcordate; young
branchlets and inflorescence, if pubescent, ferruginous- or subferruginous-
tomentose.
D. Leaf-blades chartaceous, subferruginous-tomentose beneath, the surface
not Sasa reticulate; inflorescence softly and loosely ig ened
COMET LOSE saree nities ooo oo oare Hie SUS 2 6 ans bo Gea aR omer enamels 34. N. belizensis.
D. Leaf. ane rigidly coriaceous, glabrous throughout, scree reticulate
above and shining; inflorescence glabrous..............-+-+- 35. N. rudis.
Leaf-blades obovate to obovate-oblong, cordate or at least rounded, not re-
curved usually; pubescence of branchlets and inflorescence never ferruginous-
tomentose, but tawny, if present at all.
D. Leaf-blades membranaceous, glabrous or glabrescent pra AtAMOSE. cays.
O
Atte ee eA bat AL ee ee IP en eee rE Eos
D. ree blades coriaceous or subcoriaceous, tawny- nee aes fruit
sely pubescent on lower half........... 6. cess eee eee es 7. N. sinuata.
B. Base of ieaf- blades definitely auriculate and recurved, the ae auricles often
overlapping conspicuously beneath............ 002 eee eee eee eed _N. reticulata.
1. Nectandra Davidsoniana, sp. nov.
Arbor, ramulis foliosis fulvo-sericeo-pubescentibus celerrime glabres-
centibus ee glabris, ramulorum cortice revera colore rubescenti-brunneo
epidermate tenuissimo secedibili griseo velato. Folia alternata, petiolis
Se "ate aas ad 8 mm. longis, laminis utrinque glabris mem-
branaceis, supra lucidis, in sicco viridescentibus vel brunneis subtus pal-
lidioribus, lanceolatis vel elliptico-lanceolatis, 5-6 (—7) cm. longis et 13-17
(- 22) mm. latis, basi cuneatis, apice obtusis vel obtuse acuminatis, pennin-
erviis, costa supra inconspicua subtus leviter elevata, nervis plerumque
4-par ribu us supra obscuris subtus leviter elevatis angulo 35—45° divergentibus,
rete venularum supra rT iaecapeaeny ee haud conspicuo. Inflo-
minibus ser. I & Il +1 mm. 1. longis antheris oblongo- clits. filamento
gracili triplo longioribus, ser. III + 1.25 mm. longis, antheris oblongis big-
landulosis, glandulis brevistipitatis antheris et ear ees aequalibus; stami-
nodiis “triangularibus pubescentibus +0.6 mm. longis; gynaecio glabro
m. longo, ovario ovoideo stipite triplo longiore, stylo duplo longiore,
Goa iecaaics conspicuo. Fructus ignotus, niger, fide coll. , receptaculo
rubro, fide coll., glabro, hypocrateriformi, disco plano 1 cm. diam. sub-
tentus, pedicello subverruculoso, ad 1 mm. longo et utrinque 4 mm, lato,
margine integro.
DisTRIBUTION: Known only from the type-locality.
: Chiriqui: Chiquero, Boquete, alt. 1830 m., April 11,
Davidson 564 (fl., fr.. TypPE—A, Ch) (tree; flowers cream; fruits ee with red
receptacle).
The new species is somewhat like N. salicifolia in aspect, but the leaves
are smaller than any known representative of that species. The floral
370 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
structure too is not unlike, the greatest divergence being the subglobose
anthers of the two outer series, as opposed to the subreniform anthers
characteristic of N. salicifolia and its allies. The fruit unfortunately is
lacking on the specimens of NV. Davidsoniana found in the herbaria of the
Arnold Arboretum and the Chicago Museum, but the cupule is present.
According to the collector, the fruit is black, and it is subtended by a red
woody cupule that is hypocrateriform.
2. Nectandra Smithii, sp. nov.
Arbor 10-17 m. alta, ramulis foliosis minute subferrugineo-pubescentibus
celerrime griseis striatis deinde griseis verruculosis. Folia alternata, petiolis
gracilibus glabrescentibus canaliculatis (5—) 7-10 (—12) mm. longis,
laminis utrinque glabris, basi costae excepta, membranaceis, in sicco utrin-
utrinque conspicua subtus elevata, nervis plerumque 5-paribus supra paulo
subtus valde prominulis angulo 35—45° divergentibus, glandulis conspicuis
fulvo-pubescentibus in nervorum lateralium axillibus, rete venularum utrin-
que perconspicuo. Inflorescentia axillaris, brevis, paniculata, 2-3 (—5.5)
cm, longa, glabrescens, pauciflora, brevipedunculata, pedunculo gracili 1—2
cm. longo. Flores ad 3.5 mm. longi, pedicellis 1.5—2 mm. longis gracilibus,
perianthio subcampanulato mage vel gilvo, lobis oblongis obtusis crassis
papilloso-tomentosis, 2.5-3 mm. longis; staminibus ser. I & II +-0.8 mm.
longis antheris subreniformibus filamento duplo longioribus, ser. ITI] + 1.25
mm. longis antheris filamentis conspicue biglandulosis glandulis aequalibus;
staminodiis oblanceolatis acutis +-0.6 mm. longis; gynaecio glabro + 1.7
mm. lon ngo, ovario late ovoideo, stylo brevi robusto, stigmate triangulari.
Fructus niger, subglobosus, minute gesagt ad 1 cm. diam., cupula
vadosa tumescente ad 4 mm. longa, 6 m , et 2 mm. alta subtentus,
pedicello 4-5 mm. longo apice ad 3 mm. as senting
DisrRiBUTION: Costa Rica, in the Caribbean cloud-forest at 1600-1700 m. altitude,
and in Panama up to 800 m
Costa Rica: Alajuela: La Palma de San Ramon, Brenes 6825 (fl., Ch); Zapote
de San Carlos, region of Zarcero, growing at edge of woodland in semi-shade in Carib-
bean cloud-forest, alt. 1600 m., March 26, 1938, . bap H541 (fl, type—A, Ch)
round buds pink or red; petals white) ; La Pefia de Zarcero, A. Smith H.590 (fl., A, Ch)
PanaAMA: Coclé: Vicinity of El Valle, P. H. Allen 774 (fr.,Ch, GH, Mo). Panama:
Residual forest in boa grassland, trail from Campana to Chica, Cerro Campana,
P.H. Allen 2652 (fr., A).
This species, also, seems to have its affinity with the variable NV. salicifolia.
The foliose branchlets, the consistently few pairs of lateral nerves, and the
prominent reticulation of the small leaves set it apart from the well known
species. The bulging cupule which fits snugly about the very base of the
fruit is another differentiating character.
3. Nectandra Brenesii (Standley), comb.
Ocotea Brenesii Standley in Field Mus. Publ. Bot. 18: 454. 1937.
DistrRIBUTION: Known only from Costa Rica.
1945 | ALLEN, STUDIES IN THE LAURACEAE, VI 371
Costa Rica: Alajuela: La Palma de San Ramon, Brenes 5535 (127) (fi., Ch),
woods and pastures; Cataracts of San Ramon, March-April, 1931, Brenes 13653 (f1.,
Type, Ch); Caribbean cloud-forest, Zapote de San eee region of Zarcero, A. Smith
H.469 (fl., Ch, NY); continental divide, Zarcero, A. Smith H.516 (fl., Ch), A.571 (fl,
Ch), 4102 (fl., Ch), 4171 (fl., Ch); Suere, San Carlos, Caribbean rain-forest, on edge
of ravine, A. Smith H.1687 (f1., fae Tapeseo, Alfaro Ruiz, A. Smith P.2620 (fl., A);
Palmira, in fog-zone, A. Smith 4202 (fl. Ch). Heredia(?): Vicinity of Vara
Blanca, north. slope of Central a ueeee between Poas and Barba volcanoes, Skutch
3730. ¢i1., it., A; NY).
The branchlets of this species are slender, somewhat angled, striate, and
early clothed with a close sericeous pubescence varying from buff or tawny
to a deeper almost subferruginous color. The leaves have slender grayish-
pubescent petioles that are canaliculate and measure up to 12 mm. long.
The blades are membranaceous, elliptic, shining above, dull beneath, the
base obtuse or almost rounded, the apex ee to abruptly i paaed
acuminate or subcaudate. They measure up to 8-12 (—14) cm. long an
3.5—6 (—8) cm. broad, and at maturity show a ae close persistent pubes-
cence, sparse on the blade as a whole but dense near the costa and nerves.
The costa is impressed above and very prominently elevated beneath. The
lateral nerves, of which there are 4 (or 5) pairs, diverging from the costa
at an angle of 35—45°, are slightly elevated above and more prominently so
beneath, bearing only a slight suggestion of axillary glands. The inflo-
rescence is a short few-flowered cymose panicle, sparsely and inconspicu-
ously pubescent, becoming glabrescent, up to 6(—8) cm. long, with a slender
peduncle up to 4cm. long. The flowers are large, measuring almost 5 mm.
long and 12 mm. in diameter, the tube short, the lobes thick, fleshy, elliptic,
up to 4.25 mm. or more long. The two outer series of stamens are + 1.9
mm. long, with anthers ovate, obtuse, borne on stout filaments one-quarter
the length of the stamens. The connective is well developed, about one-
third the entire length of the anther. The stamens of the third series are
more or less oblong, bearing at the base of the filaments two sprawling some-
what depressed sessile glands that are about the height of the filaments,
only wider. The staminodia are ovate, +0.8 mm. long, borne on pubescent
stipes slightly more than half the entire length. The glabrous gynaecium
is 2.7 mm. long, the depressed-globose ovary twice the length of the rather
stout style which bears a triangular slightly decurrent flat stigma. The
only fruiting material cited does not exactly match the remainder of the
specimens in foliage-characters. The leaves are shorter, more narrowly
elliptic, and are definitely caudate. The fruit is lacking on the specimen,
but the subtending cupule (seemingly hegenwie is smooth, purplish, very
shallow, about 7 mm. long and 7—8 mm. in diameter at the apex; the long
pedicel is 1.5 cm., smooth ad oa e join the cupule with no visible
line of deareaian: The young fruits are very similar to those of NV.
savannarum, but the flowers on the same branch show the characters of NV.
Brenesii.
The same reason may be given for transferring this species from Ocotea
to Nectandra that is offered in the case of the next, Phoebe ambigens. Tam
at a loss to suggest the true affinity of this species.
4. Nectandra ambigens (Blake), comb. n
Phoebe ambigens Blake in Contr. U. S. an Herb. 24: 3, pl. ] 1922; Record in Trop.
Woods 10: 21. 1927; Standley in Trop. Woods 21: 719. 1930.
DistriBuTion: Eastern Guatemala and Honduras.
372 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
GUATEMALA: Without locality, Kuylen G.54 (1) (¥ 8885) (sterile, Y) ; Hopi Farm,
Kuylen G.149 (Y 10508) (fl, NY, Y). Izabal: Las Playitas, Whitford & Stadt-
miller 32 (sterile, GH). San Marcos: Finca El Porvenir, on Potrero Matasan along
Rio Cabts, Volcan Tajumulco, Steyermark 37643 (fr., Ch). Honpuras: Copan:
Rodezno, Whitford & Stadtmiller 7 (fl, isorypr, GH, Y). Colon(?): El Limon,
Whitford & Stadtmiller 26 (sterile, Y).
Native NAMES: “Aguacatillo” (Guatemala); “Ajio,” “Guambo” (Honduras).
This species has branchlets that are finely and sparsely grayish-pubescent,
becoming glabrous, and angled, becoming grayish-striate. The leaves have
stout petioles 2 (—2.8) cm. long and sparsely pubescent to glabrescent.
The blades are subcoriaceous, shining above and beneath, usually elliptic,
cuneate at the base, obtuse or shortly obtusely acuminate, 9.5—26 cm. long
and 3.5-10 cm. broad. The thick costa and lateral nerves, of which there
are 7 or 8 (—9) pairs diverging at an angle of about 45°, are slightly ele-
vated above and more so beneath. Blake mentions axillary tufts being
frequently present, but on the material at hand they are almost imper-
ceptible. The dense and prominent reticulation on both surfaces of the
leaf is one of the most striking characteristics of the species. The glabres-
cent inflorescence is axillary, few-flowered, up to 15 cm. long, the stout
peduncle up to 7.5 cm. long. The most arresting feature of the species is
the presence of large flowers up to about 8 mm. long and 15 mm. in diam-
eter at anthesis, densely grayish-pubescent without, the spreading lobes
bright- castaneous at anthesis, and subtended by a slender pubescent pedicel
up to 9 mm. in length. The tube is short, the lobes broadly elliptic, up to
7 mm. long and about 5 mm. broad, eshy and papillose within. ‘The
stamens of the two outer series are ro} 5 mm. long, the almost quadrate
anthers are rounded at the apex and nearly sessile. Those of the inner
series measure nearly 3 mm. long, the anthers square, truncate at the apex,
the filaments nearly half the entire length and bearing near the base two
conspicuous subglobose sessile glands. The staminodia are ovate, sub-
sessile (in the type), and +1.25 mm, long. The glabrous gynaecium is
mm. or more long, the depressed- globose ovary equalling in length the
stout style topped by a broad conspicuous subtriangular decurrent stigma.
The Steyermark specimen from Guatemala shows a large fruit which is
green (probably not fully ripe), ellipsoid, apiculate, the entire apex con-
spicuously smooth, 2.5 (—3) cm. long and 1.7 cm. broad, subtended by a
sharply lobed woody verrucose cupule 1 cm. long, over 2 cm. in diameter,
and 8-9 mm. deep. The enlarged pedicel is 1.5 cm. long, expanded to
nearly 1 cm. diameter at the apex.
Kuylen G. 149, from Guatemala, shows staminodia that are more stipitate
than those of the type, but in other respects it matches the type-material.
In spite of the presence of staminodia, the other characters of the flower
indicate that Nectandra is the genus to which the entity belongs. The
large flower recalls that of V. globosa, but the anthers have a less developed
connective, and the style is longer. Also the foliage-characters are entirely
different.
5. Nectandra rubriflora (Mez), comb.
Ocotea rubriflora Mez in Jahrb. Bot. Ge Berlin 5: 279. 1889; Standley in Contr.
U.S. Nat. Herb. 23: 296. 1922
Ocotea perseifolia Mez & J. D. Smith in Bot. Gaz. 20: 10. 1895; Standley in 1. c.
Ocotea persicifolia Mez & J. D. Smith, in Index Kewensis, Suppl. I. 1906, sphalm.
1945 | ALLEN, STUDIES IN THE LAURACEAE, VI 373
Siete eal Eastern Mexico, Guatemala, and British Honduras, at low altitudes,
from 20 to
Mexico: ae Chiltepec and vicinity, Tuxtepec, in llanos, Martinez-Calderén
49, 469 (fl, A). Tabasco: Teapa, Linden in 1840 (fl, photo. of type of Ocotea
rubriflora, Ch); San Sebastian, Rovirosa 475 (fl, Ch). Guatemata: Izabal:
Izabal, alt. 36 m., April, 1889, J. D. Smith 1807 (f1., isotype of Ocotea perseifolia, US).
BriTIsH Honovnas: Stann Creek: Middlesex, along river-bank, Schipp 381 (fl.,
Ch, GH, NY
This species has branchlets that are angled and clothed with a close short
pubescence which verges on a ferruginous shade, or perhaps more tan. The
leaves are borne on petioles up to 3 cm. long, canaliculate and glabrescent.
The blades are chartaceous to subcoriaceous, broadly elliptic or oblong-
elliptic, roundish to obtuse or even cuneate at the base, acuminate to shortly
acuminate or subcaudate at the apex, up to 26 cm. long and up to 12 cm.
broad, shining above, glabrous or very minutely and inconspicuously pubes-
cent beneath, greenish brown throughout in the dried state. The costa is
conspicuous though impressed above, frequently slightly pubescent to-
ward the base on both surfaces, and very prominently elevated beneath.
The 10-12 pairs of lateral nerves are often reddish, only slightly elevated
above but more so beneath, and diverge from the costa at an angle of about
45° or occasionally 60°. The reticulation of the blade is loose, prominent
above and beneath. The inflorescence consists of axillary or subterminal
panicles 9 (—15) cm. long, clothed with the same minute pubescence that
covers the branchlets, and borne on a stout peduncle 4 (—9) cm. in length.
The flowers, usually with a reddish tinge, are large, up to 4. ; mm. in length
and 8-9 mm. in diameter, the ovate obtuse perianth-lobes fleshy and papil-
lose, +3-3.8 mm. long. The stamens are +1.25 mm. long, and the anthers
are thick, papillose, square or parabolical, with the apex often emarginate,
borne on stout very short filaments pubescent at the base. The filaments
of the inner series of stamens bear at the base sessile glands, which are
seemingly compressed-subreniform or subglobose. No staminodia are pres-
ent. The glabrous gynaecium is +1.7 (—2.15) mm. long, the ellipsoid or
subglobose ovary topped by a very fies stout style with a subcapitate
stigma at its apex. No fruit is kno
The species resembles ania N. Lundellit. The flowers of N.
rubriflora, as the name indicates, are usually reddish, whereas those of
N. Lundellii are white and, the collectors note, very fragrant. The squarish
stamens — more or less petaloid and densely papillose — present a further
distinguishing character.
6. Nectandra Schippii, sp. nov.
r 10.5 m. alta, ramis griseis glabris, ramulis dense ferrugineo-tomen-
fgets ” Folia alternata subverticillata, juventute utrinque ferrugineo-tomen-
tosa, petiolis robustis ferrugineo- tomentosis, ad 5—10 (—15) mm. longis et 2.5
mm. latis, laminis supra glabris costa et nervis exceptis, subtus ferrugineo-
tomentosis, coriaceis, in sicco viridescenti-brunneis, ellipticis, (6—) 12-14
cm. longis et (2.5—) 5 (—6) cm. latis, basi acutis, rotun datis, vel subcordatis,
apice subobtusis vel rotundatis et emarginatis, saepe longe vel abrupte
subtus conspicuissime elevata et ferrugineo-tomentosa, nervis (6—) 8- vel
9-paribus supra impressis subtus elevatis et pubescentibus, angulo 45°
374 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXVI
divergentibus, rete venularum supra satis impresso subtus elevato et pubes-
cente. Inflorescentia axillaris, juvenili bracteis adhuc onusta, ad 6.5 cm
longa, ar gl te cymoso-paniculata, pauciflora, longipeduncu-
lata, pedunculo ad 4.5 cm. longo. Flores leviter fragrantes, immaturi,
pedicellis brevibus ACRE perianthio albo, lobis crassis, papilloso-
tomentosis; staminibus ser. I & II subreniformibus apice rotundatis fila-
mentis crassis et latis ae ser. III subreniformibus biglandulosis,
glandulis magnis sessilibus antheris aequalibus, staminodiis brevistipitatis
bene conspicuis ; gynaecio glabro, ovario stylo duplo longiore, stigmate
parvo inconspicuo. Fructus ignotus
DistrIBUTION: Known only from the type-locality.
British Honpuras: Stann Creek: Rare in dense shade on bank along Big
Creek, alt. 30 m., Dec. 3, 1931, Schipp 856 (fl., Ch, rype— GH, NY) (tree 10.5 m.,
diam. 23 cm., wood cream-colored, close-grained; flowers white, slightly fragrant).
The flower-structure of this species is very similar to that of V. globosa.
The branchlets, however, with their dense conspicuous woolly covering of
ferruginous-tomentose pubescence, differ widely. The bright tomentum
after a period wears off to a thin rather straggling layer of grayish incon-
spicuous pubescence that dwindles to a mere glabrescent state. The
fragrant flowers resemble those of V. globosa.
7. Nectandra Austinii, sp. nov.
Arbor 7.5—13 m. alta, ramulis valde sulcatis fulvo- vel pallide brunneo-
tomentosis. Folia alternata, juventute utrinque dense fulvo-tomentosa vel
brunneo-tomentosa, petiolis robustis tomentosis canaliculatis, 1-2 cm.
longis et ad 4 mm. latis, laminis supra glabris basi costae excepta, subtus
es inde consperso- -pubescentibus nervatione dense fulvo-pubescentibus,
oriaceis, in sicco supra pallide olivaceis, subtus atro-brunnescentibus,
obovatis, A3 (—22) cm. ea et 7 (-12 5) cm. latis, basi cuneatis saepe
leviter elevatis angulo (35—) 45 (—55)° divergentibus, rete venularum supra
obscuro subtus conspicuo atrato-pubescente delineato. Inflorescentia axil-
laris anguste paniculata, 4-8 cm. longa, dense fulvo-tomentosa, pauciflora,
longipedunculata, pedunculo ad 4 cm. longo satis gracili. Flores immaturi,
ad 4 mm. longi, glabri, pedicellis brevibus ad 1 mm. longis pubescentibus,
gracilibus, perianthio subcampanulato(?), lobis ellipticis nonnihil crassis
glabris +2.15 mm. longis, staminibus ser. I & II +1 mm. longis, antheris
subsessilibus, connectivo conspicuo truncato, ser. IIT +1.25 mm. longis,
DistrIBUTION: Known only from the cloud-forests of Alajuela in Costa Rica, at
an altitude of 1600-2345 m
Costa Rica: Alajuela: Alfaro Ruiz, La Pefia, A. Smith P.2114 (fl., A) ; Tapeseo,
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 375
in mold and loam in half-shade in Caribbean cloud-forest, gn 1650 m., Jan. 6, 1940
Austin Smith P.2226 (fl., Type, A) (tree 11 m., trunk base 38 cm.; bark nectral: bow
with raised dots; crown broad; leaves cheriareous: thickened, Gacad opaque green,
paler below with fine reticulation; drupe green, fig-form) ; region of Zarcero, A. Smith
A.240 (fr., Ch).
The new species superficially has characters in common with J. sinuata,
but the flowers are smaller and more delicate, with their anthers truncate
and the ovary glabrous; the leaf-blades are less densely tomentose and
their bases are not cordate or conspicuously rounded.
8. Nectandra panamensis Mez in Jahrb. Bot. Gart. Berlin 5: 443. 1889.
DistRIBUTION: Central part of Panama, at 30-100 m. altitude.
PanaMA: Canal Zone: Near Gorgona and Maume, Wagner s.n. (fl., photo. of
type, GH); Darién, MacBride 2703 (fr. Ch, Mo). Panama: Around Alhajuela,
Chagres Valley, forests, on dry limestone, Pittier 2398 (fl., Ch, GH, NY); vicinity of
Pacora, P. H. Allen 1126 (fl., Ch, GH, Mo), 2033 (f1., A
The branchlets of this species are glabrous with only a faint suspicion of
pubescence at the apex, greenish, becoming brownish, angled, becoming
terete and striate. The leaves have glabrous canaliculate petioles up to 1
cm. long. The blades are membranaceous or chartaceous, pale greenish in
the dried state, glabrous, narrowly elliptic or elliptic- lanceolate, the base
cuneate, the apex acutish or subacuminate or often obtusely long- acuminate,
15-20 cm. long and not more than 5.5 cm. broad. The costa is plane above,
though conspicuous because of its yellowish color, and is elevated beneath.
The lateral nerves, of which there are 5—7 (—9) pairs, are yellowish and
slightly elevated above but more conspicuously so beneath, diverging at an
angle of 25-35 (—45)° and usually bearing in their axils very conspicuous
ret ellipsoid eee glands which are conspicuous on the upper surface
the blade as well. The glabrescent paniculate axillary or subterminal
ees is - to 12 (-19) cm. ng: much-branched near the base, with
a very short peduncle. The white flowers are pubescent, and measure
4 (-6, according to the author) mm. long. The perianth- lobes are fleshy,
hairy, and up to 3 mm. lon The two outer series of stamens are 0.8
mm. long, the anthers are broadly ovate, obtuse or somewhat depressed-
globose, the connective tissue is apparent, and the filaments are very short
and thick. The stamens of the inner series are 1.25 mm. long and consist
of subrectangular anthers that are broader than long and borne on rather
stout filaments with two ae aa sessile basal glands that are almost
larger Asie the anthers themselves. The staminodia are very thin, ovate,
+ m. long, the stipe mele eaunilitie half the entire length.
brous pias is +1.7 mm. long, the ovary ellipsoid with a very on
stout style topped bya oaeneae triangular stigma. The fruit (probably
immature) is borne in a cyathiform cupule up to 6 mm. long, 9 mm, in
diameter, and 4 mm. deep. The supporting es is a to 5 mm. long
and scarcely broader at the apex than at the
The species is reminiscent of V. Gentlezt, ie to be found in this area.
The leaf-shape, however. and the type of venation distinguish it from the
above-mentioned species. There are points of difference between the
species-description and the specimens cited, but I can find no major
features which bar the numbers from inclusion in Mez’ species.
376 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
9. Nectandra globosa (Aublet) Mez in eae “i Gart. Berlin 5: 415. 1889, excl. syn.
Laurus globosa Aublet, Pl. Guian. 1: 364
Sassafridium macrophyllum Rose in sey fg S. Nat. Herb. 1: 355. 1895.
DistriBuTION: West Indies, Mexico, and Central America.
Mexico: Sinalo San Ignacio, Arroyo del Agua Colgada, Ortega 715 (fl., Ch).
Nayarit (Tepic) : Santiago, Lamb 6 614 (fl, GH). Colima: Manzanillo, E. Palmer
the Rio Tecpan, Langlassé 740 (fl, GH). Oaxaca: Tuxtepec, Chiltepec, and vicinity,
in llanos, Martinez-Calderén 292 (fl., Ren Pinotepa to Jamiltepec, Nelson 2347 (fl.
GH) ; Jami iltepec, Conzatti 4404 (fl, NY). Guatemara: Without locality, Brigham
sn. (fl, GH). Petén: Above El pate along Rio Cancuen, between La Concordia
and El Pos a 45892 (fl, Ch). Alta Verapaz: Cubilgiiitz, von Tuerck-
heim 7964 (fl, GH, NY); wet wooded ravine, along Rio Carcha, between Coban and
San Pedro se Siendliaw 89878 (fl., Ch), at edge of river, 90010 (fl., Ch). Izabal:
Near mouth of Rio Polochic, Maxon & Hay 3788 (fl., NY), von Tucrckhein ieee (hh,
N Guatemala: Aguilar 499 (fr., Ch). Chimaltenawec: Iztapa, J. R. John-
ston 1169 (fl, Ch). Retalhuleu: In wet thicket, Standley 66695 (f1., Ch). Suchi-
tepéquez: Vicinity of Mazatenango, stream-bank, Maxon & Hay 3477 (fl., Ch);
near Santo Domingo, south of Mazatenango, stream-bank, Standley 88900 (fr., Ch).
Escuintla: Concepcion, J. D. Smith 2095 ({fr., GH) ; Escuintla, Hayes s.n. (fl., GH) ;
south of Rio Burridén, northeast of Escuintla, road-side, Standley reba (fr., Ch).
Santa Rosa: Rio de La Plata, Heyde & Lux 4261 (fl., GH, NY) ; about Guazacapan,
in damp forest, Standley 78677 (fl., Ch); wet forested uebeade. Rio de la Cruz, east
of Taxisco, Standley 79005 (fl., Ch) ; road-side, region of Capulin, south of Los Cerritos,
on road to El Ahumado, Standley 79623 (fi., el Honpuras: Tegucigalpa:
Mont. de la Flor, river-bank in pine-region, C. & V. von Hagen 1135 (sterile, NY).
Yoro: Near Progreso, Farm 42, Hottle 85 (fl. a 106 (fl., Ch). Britisn Hon-
purAs: Toledo: ‘Forest Home,” Punta Gorda, in open pasture, Schipp 1037 (f1.,
GH, NY); forest, Moho River, Peck 553 (fl., GH); Temash River, Stevenson & Smart
126, 130 (¥ 19785) (fl., Ch, Y). Ex Satvapor: Ahuachapan: Vicinity of Ahu-
achapan, Standley 19883, 20280 (fl., GH). Sonsonate: Vicinity of Izalco, Standley
22225 (fl, GH, NY). La Libertad: Santa Tecla, S. Calderén 1419 (fl, GH, NY).
Usulutan: Triunfo, Shannon 5003 (fl, GH). Nicaracua: Chinandega: Chin-
andega, Baker 2015 (fl., Ch, GH, NY). Costa Rica: Guanacaste:Near Nicoya,
Tonduz 13806 (fl., GH), Valevio 495 (fr., Ch). Puntarenas: Santo Domingo,
Tonduz 7153 (Herb. Nat. Costa Rica 10047) (fl., GH); between Puerto Jiminez and
S mingo, Brenes 12278 (757) (fr. Ch). Alajuela: Cataracts of San Ramén,
Brenes eke (fl., Ch) ; Alajuela, J. D. Smith 6755 (fl., GH) ; in forest of Pacific tropical
zone, Atenas, A. Smith 2 465 (f1.,, A). San José: Vicinity of El General, Skutch 3836
(fl, A, NY). Panama: Without locality, Hayes 1021 (fl., NY), Mell sn. (fr., NY).
Bocas del Toro: Changuinola Valley, Dunlap 215 (fl. Cooper & Slater 86
(Y¥ 10267) (fl, Ch). Chiriqui: San Felix, — etd ‘AL, Ch). Coclé: Above
Penonome, R. S. Williams 257 (fl., NY), 379 (fl, fr.; NY), 530 (fl, NY). Canal
one: Gatun Lake, near laboratory, Wetmore & He 17 (A, Ch, GH), 43 (f1., A,
of Miraflores Lake, G. White 191 (fl., Ch, GH), P. Mua’ 276 (AL, Ch, GH); pen
P. White 95 (fr., Ch, GH, Mo); Batra. ‘Colorado Island, M. Brown 40, 68, 72, 18
Ch), Shattuck 314, 458, 807 (fl., Ch), C. L. Wilson 2 (fl, Ch); along Rio tere in
forests and thickets, Pittier 3873 (fr.. NY). Panama: Vicinity of Arraijan, P. H
Allen 1622 (fl., Ch, GH, Mo, NY); Chepo, ae 27 (fl, Ch). Darién: Chepigana,
Tucuti, M. FE. & R.A. Tery 1391 (fl., A, Ch,
NAMES: “Aguacate del monte” ne Salvador) ; fis eemmrenal (Mexico,
haa British Honduras, Costa Rica); “Canelon” (El Salvador); ‘“Quizarra,”
“Quizarra quina” (Costa Rica); “Sangre blanco” acaiaale “Sweetwood”
(Panama) ; ‘“‘Timbersweet,” “Wild Pear” (British Honduras)
mm,
ra
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 377
This species, as it occurs in Mexico and Central America, has branchlets
that are closely and minutely subferruginous-pubescent, in the early stages
angled and flattened at the nodes, later becoming reddish brown, striate,
and glabrous. The leaves are borne on stout petioles up to 1.5 cm. long,
minutely pubescent to glabrous and canaliculate. The blades are cori-
aceous, shining above and dull beneath, with obscure reticulation, early
minutely pubescent, becoming glabrescent to glabrous, elliptic to oblong-
elliptic, the base rounded or sharply cuneate, the latter appearance due to
the lower centimeter of the leaf-base being recurved, the apex long- -acumi-
nate. The costa is slightly impressed above and rather conspicuous and
elevated beneath. The lateral nerves are inconspicuous above and slightly
elevated beneath, numbering 8-12 pairs and diverging at an angle of
30-45°, frequently bearing pubescent glands in their axils. The stout
paniculate inflorescence measures up to 20 cm. long, is minutely subfer-
ruginous- -pubescent to glabrescent, many-flowered, usually widely branch-
ing, the peduncle varying from very short to 8 cm. in length. The flowers
are very large and conspicuous, up to 12 mm. in diameter, with reflexed
lobes that are very fleshy, papillose within and oo without , 4-5 mm.
long, elliptic or obovate-elliptic, acutish or obtuse at the apex. ’ The two
outer series of stamens are +1.25 (—1.5) mm. long, with the sessile or sub-
sessile anthers rounded or broadly rounded-ovate, the fleshy papillose con-
nective often one-third the entire length. The stamens of the inner series
are +1.7 mm. long, the broad anthers narrowing slightly into thick fila-
ments almost one-half their length and bearing at the base two conspicuous
spreading sessile glands almost the length of the anthers. The almost
triquetrous staminodia are +1 mm. long, the thick stipe nearly one-half the
entire length. The glabrous gynaecium measures +1.7 mm. long, the a
globose ovary two-thirds the entire length. The short style is topped by
pg discoid or obtuse stigma. The globose tees sed hon
1 n diameter, is subtended by a very shallow cupule up to 2 mm. lon
8 mm. in diameter, and 2 mm. deep, with the margin alle aera and
thin. The pedicel is up to 5 mm. in length and expanded to 3 mm. in diam-
eter at the apex
It is not certain at the present moment that NV. globosa is the correct
binomial for the species involved. Kostermans (Meded. Bot. Mus.
Utrecht 25: 19, 54. 1936) believes that Aublet’s name globosa should
replace NV. antillana of Meissner, the West Indian entity not occurring in
Surinam, and that Mez’ V. globosa should be reduced under NV. Pisi. Until
types are available for study, the species from Mexico and Central America
may continue to go under the name of NV. globosa.
There can be no doubt that Sassafridium macrophyllum Rose belongs
with this species. The earlier described Sassafridium veraguense is an
cotea.
10. Nectandra ramonensis Standley in Field Mus. Publ. Bot. 18: 453. 1937.
DistriBuTION: Costa Rica and adjacent Panama.
Costa Rica: Alajuela: Near San Ramon, Brenes 353 (478) (fl., Ch); San
Pedro de San Ramon, on the side toward San Francisco, Brenes 6612 (478) (fl., Ch);
on the trail to San Francisco, Brenes 6660 (fl., Ch); San Francisco and San Pedro de
S amon, Feb. 7, 1933, Brenes 17018 (fl., TypE, Ch); Rio Jesis de San Ramon,
Brenes 14988 (fl., Ch); La Calera de San Ramon, on the trail to La Calera, Brenes
378 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
21990 (fl., Ch) ; between Santiago and San José de San Ramon, Brenes 6834 (fr., Ch) ;
La Palma de San Ramon, Brenes 6819 (fl, Ch). Panama: Prov. unknown, Sabana
de la Tortuga, Pittier 3307 (fr.. Ch). Chiriqui: Boquete, Davidson 566 (fl., Ch,
Mo). Coclé: Vicinity of El Valle, P. H. Allen 1635 (fl., Ch, GH, Mo, NY); north
rim of El Valle, Alston (Allen?) 1858 (ir., Ch, GH, Mo, NY).
This species is very close to NV. globosa, occurring insofar as is known at
present in Costa Rica and Panama. A similar pubescence clothes the
young branchlets and leaves. The branchlets however at maturity are
usually grayish, the outer cortex sometimes flaking off to reveal a dark red-
dish brown color. The leaves are subtended by short slender pubescent
petioles less than 1 cm. long. ‘The blades are sericeous beneath in the early
stages, later becoming minutely pubescent. The adult leaf-blades are
shining above, dull beneath, elliptic, occasionally oblong-elliptic, the base
obtuse and/or the lowermost portion attenuately cuneate, frequently almost
decurrent and recurved. The apex is obtuse to acutish or acuminate, the
costa and lateral nerves slightly impressed above and somewhat obscure,
slightly elevated beneath. The lateral nerves number not more than 4 or §
pairs, diverging from the costa at an angle of 35—45° and bearing pubescent
glands in their axils. The inflorescence consists of fewer- flowered axillary
or subterminal panicles minutel sericeous-t and
up to 8 cm. long at most, with long stout ‘peduncles frequently up to 6 cm.
long. The flowers are similar to those of NV. globosa in structure. The
fruits are ellipsoid rather than globose, 1.5 cm. long and 9 mm. in diameter,
subtended by deeper cupules 4—5 mm. long, 8-10 mm. in diameter, and
—3 mm. deep. The pedicel is about 5 mm. long and 3—4 mm. in diameter
at the apex.
11. Nectandra Heydeana Mez & J. D. Smith in pe ‘ae 19: 262, t. 25. 1894; Standley
Calderon, Lista Prelim. Pl. Salvador 84.
DistripuTIon: Guatemala and Honduras, at 900-1360 m. altitude.
GuateMaLA: Alta Verapaz: Along Rio Carcha, near San Pedro Carcha, Stand-
ley 92158 (fl, Ch). Santa Rosa: Santa Rosa, alt. 900 m., Nov. 1892, Heyde & Lux
4260 (fl., IsoSYNTYPE, GH, NY), Jan. 1893, Heyde & Lux 4578 (fl., IsosyytyPE, GH,
NY). Honpuras: Tegucigalpa: Along river, Mont. de la Flor, Guarabuqui,
C. & V. W. von Hagen 1275 (fr., Ch, NY).
NATIVE NAME: ‘“Aguacatilla’” (Honduras).
The present species, so little collected to date, has branchlets which are
early close fulvous-pubescent, later glabrescent, angled, becoming striate.
The leaves are borne on slender petioles up to 2 cm. in length which are
subcanaliculate and fulvous- or grayish-pubescent. ‘The blades are gla-
brous except for the pubescent axillary glands beneath, elliptic, sometimes
oblong-elliptic, the base cuneate or obtuse or even almost rounded, the apex
abruptly acuminate or obtuse, up to 19 cm. long and to 8 cm. broad. ‘The
costa is impressed and conspicuous above and the lateral nerves, of which
there are usually 7 or 8 pairs, are very slightly elevated and conspicuous
above, whereas both are conspicuously elevated beneath. ‘The lateral
nerves diverge from the costa at an angle of 35—45°, and the reticulation is
rous, paniculate, up to 7 cm. long, and is borne on a slender peduncle up
to 5 cm. long. The flowers are large, almost 4 mm. long and up to 7 mm
in diameter, the slender filamentous pedicels glabrous and up to 5 mm. long.
The broadly elliptic lobes are rather fleshy, + 3.8 mm. long, and papillose
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 379
on the inner surface. The stamens of the two outer series are +1.25 mm.
long, with anthers that are almost square or broadly ovate, truncate, ae
sessile, sometimes mucronate. Those of the inner series are eet, 7
long, the squarish anthers somewhat emarginate, the two upper cells defi.
nitely lateral. The filaments are one-half as long as the anthers and bear
laterally at the base two broad spreading glands about half their length and
twice as broad as long. The staminodia are ovate, +0.6 mm. long and
densely shaggy-pubescent. The glabrous gynaecium is +2.5 mm. long;
the subglobose or broadly ovoid ovary is not quite twice the length of the
style with its almost peltate conspicuous stigma.
The fruit of the Honduran specimen is ellipsoid, greenish black in the
dried state, up to 2.5 cm. long and 1.8 cm. broad, subtended by a shallow
aiid flaring cupule not more than 6 mm. long, 1 cm. in diameter, and less
than 3 mm. deep.. The expanded oe also woody, reaches a length of
9 mm. ee a breadth of 5 mm. at its a
The species may be said to show a ight relationship to V. Woodsoniana,
but the grayish pubescence of the many-flowered inflorescence of the latter
separates it immediately. The fruits of the two species are entirely different,
that of V. Woodsoniana being much smaller and with a more shallow cupule.
12. Nectandra Gentlei Lundell in Contr. Univ. Mich. Herb. 6: 13. 1941.
DistrRIBUTION: Southern Mexico, Guatemala, Honduras, British Honduras, and
Panama
Mexico: Oaxaca: Ubero, L. Williams 9377 (fr., A, Ch, NY). GuATEMALA:
Petén: La Libertad, Lundell 2578 (fr.. Ch, NY). Huehuetenango: Wooded
slopes bordering Rio Lacandon, between Ixcan and Ixcan, Sierra de los Cuchuma-
tanes, Steyermark 49354 (fl. Ch). Alta Verapa Cubilgiiitz, von Tuerckheim
8578, 8579 (fl. GH, NY); woods between Finca Cubilgiiitz and Hacienda Yaxcabanal,
Steyermark 44827 (fr., Ch); wet forest, above Tamahu, Standley 70932 (sterile, A).
British Honpuras: Belize: Forest near Manatee Lagoon, Peck 196 (fl., GH).
ert Creek: Miullins River, in high ridge on river-bank, Gentle 3456 (fl., ISOTYPE,
NY); rae Schipp 337 (f1., Ch, GH, NY); Stann Creek Railway, 10 mile,
ci, 164 (fr. GH, NY). Toledo: In forest, Peck 574 (fl, GH). Honpuras:
Bs ear Puerto ae P. Wilson 558 (fr., Ch, GH, NY). Panama: Province unknown,
a and ae R.S. Williams 797 (fr., NY). Chiriqui: El Pedregal de David,
Ee 5117 (fl, Ch). Coclé: Above Penonome, R. S. Williams 617 (fr., NY).
Canal Zone: icity of Miraflores Lake, P. White 243 (fl., Ch, GH, Mo); Ancon
Hill, Killip 3032 (fl., Ch), Standley 26376 (fr., GH); eo Quarantine Station, Pittier
2076 (fr., Ch) ; Hospital Grounds, Pittier 2733 (fr., GH) ; een Corozal and Ancon
Pittier 2639 (fr.. NY); Balboa, hillside, west side of ao poe & Stork 987 (fi.
NY). San José Island: Perlas archipelago, Gulf of Panama (about 55 miles SSE
of Balboa), Johnston 82 (fl., A), 221 (fl., A), 270 (young fr., A), 521, 699, 716 (fr., A).
This species has angled branchlets that early are covered with a die
short subtomentellous pubescence of a pale ferruginous or brownish color,
presently becoming fuscous, and acer the branchlets being glabrescent
to glabrous, dark reddish brown, and striate. The leaves are supported by
petioles up to 1.5 cm. long, canaliculate, and brown-tomentellous. The
chartaceous lanceolate leaf-blades appear cuneate at the base but actually
are rounded or even subauriculate, the extreme bases being usually tightly
recurved. The apex of the blade is attenuate into a slender acumen which
may or may not be caudate. The blades measure up to 20 cm. long and to
4.5 (—5) cm. broad, the broadest part of the blade being at or below the
middle. The blades are early sericeous throughout, but soon become gla-
380 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
brescent to glabrous above, remaining shortly but persistently pubescent
e costa and lateral nerves alike are slightly impressed and
rather inconspicuous above, being prominently elevated beneath. The
lateral nerves number 4-6, occasionally 7 or 8, and diverge ea ie from
the costa at an angle of about 45°. Ata distance of 1-l.5c rom their
origin they ascend abruptly and follow the outline of the leaf rene parallel
with the midrib. The inflorescence is axillary, paniculate, papa Speed
or brown-tomentellous, becoming guameate beri up to 8 (—12) cm
in length, the peduncle reaching a length of 4 (—6) cm. P The flowers are
yellow or white, sometimes fragrant, up to 3 mm. long and 5.5 mm. in diam-
eter, the perianth- -tube being well defined, constricted at the apex, and about
1 mm. long. The lobes are usually elliptic, +1.7 (—2.15) mm. long, rather
thick and papillose at the tip. The stamens of the two outer series are
+0.6-.8 m m. long, the anthers agen globose, almost sessile. Those
of the inner series are +0.8 (—l m. long, with almost. square anthers
slightly emarginate, with large ti cra and contiguous glands borne at
the base of the short filaments. The slender, stipitate, often pubescent
staminodia are variable, lanceolate, oblanceolate, or even ovate. The gla-
brous gynaecium is + 1.7 mm. or r le ess s long, the style slightly longer ‘i
the ovoid ovary, and topped by a conspicuous triangular slightly decurrent
stigma. The fruit is subglobose, about 8 mm. in diameter, glabrous and
black at maturity. It is subtended by a thin shallow cupule not more than
3 mm. long, 6 mm. in diameter, and usually less than 2 mm. deep, glabrous
and with an entire margin. The pedicel is = than 5 mm. long and ex-
panded at the apex to about 2 mm. in diamet
Nectandra Pichurim (H.B.K.) Mez, in which Mez included NV. cuspidata
Nees, was interpreted by him to iaclucle specimens collected from Mexico
south to Brazil and Venezuela. Kostermans (Meded. Bot. Mus. Utrecht
25: 21. 1936) believes NV. cuspidata to be distinct from NV. Pichurim because
of the difference in the type of pubescence, the number of primary nerves,
the cupule-shape, etc. It is my belief that the Mexican and Central
American entity is to be separated from 1. cuspidata also on floral charac-
ters as well as foliage. The apex of NV. cuspidata is attenuate but always
obtuse. Further collections may show an intergradation of all of these
characters. Nectandra membranacea, from the West Indies, has also been
confused with the species under discussion. A glance at the original
description of the latter immediately precludes the possibility of their being
identical, for Grisebach gives the leaf-blades as ovate-oblong or elliptic
with an abruptly acuminate apex. The texture of the leaf and the recurved
base recall NV. globosa, but the small flowers separate it at once from the
atter.
13. Nectandra Woodsoniana, sp. no
rbor 7-15 m. alta, ae ineniicg adpresse fulvo-tomentellosis, mox
glabrescentibus brunnescentibus demum angulatis striatis glabris griseis.
Folia alternata, petiolis plus minusve atris ad 1.5 (—2) mm. longis canalicu-
latis pubescentibus vel glabrescentibus, laminis glabrescentibus vel glabris
glandulis axillaribus exceptis, in sicco griseo-viridibus, ellipticis vel oblongo-
ellipticis, ad 24 cm. longis et 8 cm. latis, basi cuneatis, apice obtusis acutis
vel acuminatis, penninerviis, costa supra conspicua leviter impressa subtus
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 381
elevata, nervis lateralibus 7 vel 8 (—10)-paribus supra leviter subtus valde
elevatis angulo 45 vel 35° divergentibus, rete venularum supra leviter ee
conspicue elevato. Inflorescentia axillaris vel subterminalis, ad 20
multiflora, pedunculo ad 10 cm. longo. Flores ad 3 mm. longi, pedicellis
2-3 mm. longis pubescentibus, perianthio albo, lobis ellipticis vel ovatis vel
anguste obovatis, +2.5 mm. longis, carnosis, ha papillosis, extus pubes-
centibus; staminibus ser. I & II +0.6-.8 mm. longis, antheris subreni-
formibus filamento robusto duplo long Sie ser. III
longis, antheris quadratis filamentis pialandulose aequalibus, glandulis
antheris longitudine aequalibus; staminodiis +0.6 mm. longis ae obtusis
stipite robusto dimidio breviore: gynaecio glabro +1.7 mm. longo, ovario
ovoideo longitudine %4,8 gynaecii aequante, stylo brevi stigmate discoideo
conspicuo, Fructus ad 15 mm. longus et 10 mm. latus, in sicco atro-
rubescens, ellipsoideus, cupula vadosa discoidea mae glabra vel glabres-
cente rugosula, ad 2 mm. longa, 5-6 mm. diam., et 1 mm. alta subtentus,
pedicello incrassato striato glabrescente ad 2 mm. longo.
Distr1BuTION: El Salvador, Costa Rica, and Panama, from an altitude of 1800 m.
in Costa Rica to 670 m. farther south, and finally in central Panama as low as 20-90 m
L SALVADOR: Ayutuxtepeque, S. Calderén 1117 (fl, GH, NY). Costa Rica:
Province unknown, Las Nubes, Valerio 1451 (fl, Ch). Guanacaste: Obrededores
de Tilaran, Brenes 15617 (fl., Ch). Alajuela: Hills about San Pedro de San Ramon,
Brenes 5826 (fr., Ch), 16824 (fr., Ch), 19203 (fl., Ch) ; San Francisco de Guadaloupe,
Pittier 11490 (fr., Ch). Limon: Monte Verde, Stork 1688 (fr., Ch). San José:
Rio Virilla, Brenes 14296 (fl., Ch) ; Escasi, Valerio 1327 (fl., Ch) ; vicinity of El General
in a bushy clearing, Skutch 2634 (fl, GH, NY), 4374 (fl, A, NY); Potrero of Don
José, Barrantes near S. Isidro del General: alt. 720 m., June 30, 1932, Stork frig
(A “Types «Ch: ao. Canal Zone: Mamei Hill, pre 3803 (fl. Hs
sing of Salamanca Hydrographic Station, Rio Pequeni, Woodson, Allen & on
0 (fl., A, Mo, ys
NATIVE NAMES: ‘Tepeaguacate” (El Salvador) ; “Laurel,” “Quisarra” (Costa Rica).
Nectandra Woodsoniana is striking because of its gray-green foliage and
long grayish-pubescent inflorescence. The species is very near N.
martinicensis, from the West Indies, but the latter has leaves that are
18 x 5.3 cm., the margin of the blades recurved. The filaments are
pilose, the anthers depressed-orbicular and slightly papillose, the apex
rounded, and there are large staminodia present.
The species is named for Dr. Robert E. Woodson, Jr., whose contribu-
tions toward making known the flora of Panama are of the greatest value.
14. Nectandra Lundellii, nom. nov.
Persea Gentlet Lundell in Contr. Univ. Mich. Herb. 6:18. 1941, not Nectandra
)
Phoebe Gentlei Standley & Steyermark in Field Mus. Publ. Bot. 23: 117. 1944.
DistrisuTIoN: Known only from British Honduras.
ITIsH HonpurAs: Stann Creek: Middlesex, Hope 3 (Y 4798) (fl., Ch, _Y),
A). Toledo: In open places on river-bank, Rio Grande, Schipp 1164 (fr., Ch, GH,
NATIVE NAMES: ‘“Ca’ca’woung,” “Timbersweet,” “Wild Pear” (British Honduras).
This tree has young branchlets that are densely tawny-sericeous, becom-
382 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
ing more darkly pubescent and finally glabrescent with age. The leaves
are borne on stout petioles up to 2 cm. long and up to 4 mm. in width. The
blades are coriaceous or subcoriaceous, shining above and glabrous, beneath
dull and minutely but definitely pubescent at maturity, elliptic or oblong-
elliptic, cuneate at the base or obtusely cuneate, the apex usually abruptly
acuminate, up to 32 cm. long and to 14.5 cm. broad. The costa is deeply
impressed above and thickly and prominently elevated beneath. The 8-13
pairs of lateral nerves are very lightly elevated above and more so beneath
and slightly pubescent, diverging from the costa at an angle of about
45-55°. The reticulation is very prominent above and more delicately so
beneath. The inflorescence is axillary or subterminal, paniculate, tawny-
sericeous, up to 18 cm. long, the stout glabrescent peduncle up to 8 cm.
long. The fragrant white flowers are shortly sericeous-tomentose, becom-
ing glabrescent, up to 4 mm. long. The obovate-orbicular lobes are +3.5
mm. in length, reflexed at anthesis, thick and papillose. The stamens of
the two outer series are -+-0.8 mm. long, the squarish anthers subtended by
very short stout filaments. Those of the third series are 1 mm. long, with
anthers almost rectangular, about twice the length of the filaments, which
bear subsessile conspicuous glands at the base equalling the anthers in size.
The staminodia are fairly conspicuous, +-0.8 mm. in length, the subtrique-
trous tips equalling the rather stout stipes. The glabrous gynaecium meas-
ures +1.25 mm. in length, the globose ovary topped by a nearly sessile
rather inconspicuous discoid stigma. The fruit is blackish, ellipsoid, gla-
brous, up to 2.5 cm. long and 1.3 cm. broad, borne on a flaring irregularly
lobed woody glabrous cupule 8-10 mm, long and to 17 mm, in diameter at
the apex, and about 5 mm. deep. The supporting pedicel is not more than
5 mm. long and expanded to 4—5 mm. at the apex.
This species is very similar to V. rubriflora,; for a further discussion of
the similarity to that species see the preceding pages. The cupules of the
Schipp number are entire and not lobed, and the leaves are less coriaceous,
but other than this the specimens match V. Lundellit.
15. Nectandra savannarum (Standley & Steyermark), comb. nov.
Phoebe savannarum Standley & Steyermark in Field Mus. Publ. Bot. 23: 118. 1944.
DistripuTIoN: In Guatemala, at an altitude below 400 m. except for the specimens
collected in the cloud-forest in Chiquimula, where the altitude is given as 500-1500 m.,
and mountain-slopes and coastal plains of Honduras and British Honduras.
GuatemMaLa: Alta Verapaz: Along stream bordering forest south of savanna
between base of Cerro Chinaja at Sachaj and Sacacao, alt. 150-180 m., April 6, 1942,
Steyermark 45712 (fl., Type, Ch) (tree 9 m.; leaves firmly membranous, deep green
and shining above, dull and paler green beneath; flowers white) ; near the top of wooded
ridge, vicinity of Cubilgiiitz, 14-2 miles south of Cubilguitz, Steyermark 44428 (fl,
Ch); south of Cubilgititz, Steyermark 44499 (fl, Ch). Izabal: Along stream be-
tween Milla 49.5 and ridge 6 miles from Izabal, Montana del Mico, Steyermark 38625
(fl., Ch) ; wet thicket near Puerto Barrios, Standley 73043 (sterile, Ch). Chiquimula:
Cloud-forest on top, Cerro Tixixi (Tishishi), 3-5 miles north of Jocotan, Steyermark
31567 (fr.. Ch). Honpuras: Atlantida: On bank of the Danto River, lower
slopes of Mt. Cangrejal, back of La Ceiba, Yuncker, Koepper & Wagner 8762 (fl., Ch,
NY). British Honpuras: Belize: Mullins River Road, Schipp 79 (fl. in, Ch;
GH, NY); Gracie Rock, Sibun River, Gentle 1572 (fl, NY). Stann Creek: Big
Eddy River, in high ridge, Gentle 3488 (fl., A).
NATIVE NAMES: “Laurel” or “Lavrel” (Guatemala) ; “White laurel” (British Hon-
duras).
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 383
This species, originally described under Phoebe, has slender branchlets
early covered with a close brownish pubescence that presently disappears,
leaving the branchlets dark brown or eventually grayish. The leaves are
borne on slender petioles that are pubescent and slightly canaliculate,
reaching a length of 6, rarely 8, mm. The blades are glabrous, chartaceous
to subcoriaceous, elliptic to oblong-elliptic, occasionally ovate-elliptic, ob-
tusely caudate-acuminate, the acumen as long as 1 cm., the base cuneate or
obtusely cuneate or almost r ounded, up to 8.5 (—11. 5) cm. long and 3.5
pairs of lateral nerves inconspicuous above, except the basal pair, all ele-
vated beneath. The lower pair or pairs of lateral nerves usually diverge
from the costa at an angle of about 35°, whereas the upper less conspicuous
pairs diverge at an angle of 45° or more. The appearance simulates a sub-
triplinerved condition. The reticulation is loose and obscure above but
more closely knit and prominent beneath. The glabrescent few-flowered
inflorescence consists of an axillary panicle up to 4 cm. long, subtended by
a slender glabrescent peduncle up to 2 cm. in length. The white flowers are
pubescent and up to 3 mm. long, the diameter 8.5 mm., the filamentous
pedicel 2-3 mm. long. The elliptic thick lobes are about 3 mm. long and
papillose within. The two outer series of stamens are +0.8 mm. long, con-
sisting of subreniform often emarginate anthers borne on thick very short
filaments. Those of the third series are +1 mm. long, with squarish
anthers slightly longer than the filaments, which bear two roundish short-
stipitate glands at their bases. The staminodia are usually stipe-like, +0.6
mm. long. The glabrous gynaecium measures +1 mm. long, the depressed-
globose short-stipitate ovary nearly twice the length of the style, which
bears at its apex a peltate conspicuous stigma. The immature (?) fruit is
subglobose, apiculate, borne in a cyathiform coral-orange, according to the
collector, cupule, up to 4 mm. long, to 8 mm. in diameter, and 3—4 mm. deep,
undulate or often bearing the persistent remains of the enlarged lobes of
the perianth. The pedicel measures up to 8 mm. long and has expanded
to 3 mm. in diameter at the apex.
Although the flowers of the t type possess staminodia that are conspicuous,
a character of the genus Phoebe, the shape of the anthers and the gynaecium
as a whole place the species under Nectandra.
16. Nectandra peeve (Lundell), comb.
Phoebe longicaudata Lundell in Bull. Torrey Boe Club 64: 548. 1937.
DISTRIBUTION: Se Honduras, Guatemala, and eastern Mexico, usually in ad-
vanced forests
Mexico: Chiapas: Javalinero, Palenque, Matuda 3643 (fl, A, Ch, NY).
GuaTEMALA: Izabal: Bay of Santo Tomas, between Escobas and Santo Tomas,
Steyermark 39231 (fr.. Ch). British Honpuras: El Cayo: On hillside, Vaca,
Gentle 2474 (fr., A, NY); in advanced forest, on limestone hill, Valentin, Lundell 6401
(fr.. GH, NY); in riparian and marginal forest near San Augustin, Mountain Pine
Ridge, August, 1936, Lundell 6757 (fr., SYNTYPE not seen), 6833 (fl., SYNTYPE not seen).
Stann Creek: All Pines, in broken forest-clumps, Schipp 571 ‘Gr, Ch; GH; NY).
Toledo: Near — Creek in hammock in pine-ridge, north of Monkey River,
Gentle 4094 (f1.,
NATIVE NAME: “Aguacatillo” (British Honduras).
seen no type-material of this species, but Lundell 6401, cited by
Gutidell doubtless represents the species described. The branchlets are
384 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXvI
early pubescent (rufous-pilose) , becoming glabrescent to glabrous with ase,
and slightly sulcate. The leaf-blades, with flat petioles up to 1 cm
are coriaceous, elliptic, up to 14 cm. long and 4.5 (—6.2) cm. broad, ma
often oblique cuneate bases and acuminate to caudate-acuminate apices.
The costa and 4 or 5 pairs of lateral nerves are impressed above and elevated
beneath, the lateral nerves bearing pubescent glands in their axils and
diverging from the costa at an angle of 35—45°. The blades are everywhere
reticulate. The few-flowered inflorescences are not longer than 4.5 cm.
and are pubescent, becoming glabrescent. The flowers are about 4 mm.
sessile and subreniform. Those of the inner series are rectangular, slightly
longer than the filaments, which are sparsely pubescent at the base and bear
conspicuous sessile glands the length of the filaments. The staminodia are
cordate, up to 0.7 mm. long, the slender pubescent stipes more than half
the entire length. The glabrous gynaecium is +1.5 mm. long, the sub-
globose ovary slightly stipitate, with a short style eae by a discoid
stigma. The ellipsoid or subglobose apiculate black fruit, 12 * 9-10 mm.,
is seated on a shallow cyathiform cupule not more than 2 mm. long and
6 mm. broad at the apex, and about 1 mm. deep. The subtending pedicel
is enlarged to about 7 mm. in length and 2.5 mm. in width at the apex.
The nearest relatives of the species may be found, generally speaking,
in the NV. salicifolia complex. Specifically NV. longicaudata resembles N.
savannarum, {rom the same general area, from which it is readily dis-
tinguished by the persistent ferruginous pubescence of the young branchlets
and the rather decided tendency toward a subtriplinerviate condition. The
leaves of NV. savannarum are on the whole smaller, not more than 11.5 cm.
long at most. Schipp 571 has leaves that are more heavily coriaceous,
varnished-shining above, and more heavily and more conspicuously reticu-
late. The fruit is oblong-elliptic, consistently 16 < 8 mm., the infructes-
cence as a whole being more robust than that of the other specimens.
Gentle 1572, cited by Lundell under Phoebe longicaudata, seems to belong
with NV. savannarum.
17. Nectandra nervosa Mez & Pittier ex Mez in Bull. Herb. Boiss. I]. 3: 235. 1903;
Standley in Field Mus. Publ. Bot. 18: 453. 1
Disrripution: Costa Rica, known only from the type-locality.
Costa Rica: Puntarenas: Terraba, at the os of the river on the banks,
Tonduz (Herb. Inst. Costa Rica) 6758 (fr., TYPE not s
Slender glabrous branchlets and_ shortly ee buds characterize this
species, known only from the type. The glabrous leaves are borne on
slender canaliculate petioles up to 8 mm. long. The blades are broadly
lanceolate or elliptic-lanceolate, acute at the base, very acuminate at the
apex, chartaceous, 13 cm. long and 3.5 cm. broad, above in the dried state
glaucous, below reddish green, everywhere densely and very prominently
reticulate. The inflorescence is unknown, but the infructescence is longer
than the leaves (three or more times) and is glabrous, with the fruit-bearing
erie sg up to 10 mm. in length. The ellipsoid fruit is 10 mm. long
. in diameter, subtended by a semi- foe ag acutely simple-
ie cupule which in the dried state is verruculose
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 385
The affinity of the species is noted under JN. salicina.
18. Nectandra salicina, sp. nov
Arbor 5—8 m. alta, ramis — foliosis, aa sulcatis, ae ramulis
brunneis mox peas striatis, angulatis. Folia alternata vel s aepe subop-
oie See alatis gracilibus cee canaliculatis, aes mm. pee et
nervis 6 vel 7-paribus utrinque obscuris angulo 25—35° divergentibus, rete
venularum nunc utrinque conspicuo nunc supra inconspicuo. Inflorescentia
axillaris, paniculata, 8-9 (—12) cm. longa, glabra, tea gracilis, longi-
pedunculata, pedunculo 6-8 cm. longo, glabro. Flores ad 3 mm. longi,
pedicellis ad 5 mm. longis, tenuibus, perianthio alee Gib crenaulate
cano, lobis oblongis recurvatis crassis intus et see ee dense papilloso-
tomentosis, ad 3 mm. longis; staminibus ser. I & II +0.9 mm. longis et
latis, antheris reniformibus plerumque filamento duplo longioribus, ser. III
gs mm. longis, conspicue biglandulosis, glandulis et filamentis antheris
subaequalibus; staminodiis conspicuis triangularibus stipitatis +
longis; gynaecio glabro +1.5 mm. longo, ovario subgloboso stylo duplo
longiore, stigmate plerumque triangulari conspicuo. Fructus viridis, fide
coll., subglobosus, apiculatus, 20 18 mm., cupula rubra minute verru-
culosa, fide coll., glabra, ad 6 mm. longa, 10-12 mm. lata, et 2-3 mm. alta,
margine undulata, subtentus, pedicello incrassato glabro, in sicco aciculato,
d 1 cm. longo
DistrinuTion: In Alajuela Province of Costa Rica at 850-1000 m. altitude, in
Guanacaste Province at an altitude of 500-600 m., and in the cloud-forest of Cerro
Horqueta, Boquete District, Chiriqui, ea at 1980 m. altitude.
Costa Rica: Guanacaste: Vicinity of Tilaran in moist forest, Standley &
Valerio 44506 (fl, Ch). Alajuela: Near San Ramon, Brenes 371 (515) (f1., Ch);
hedges, along road between San Miguel and La Palma de San Ramon, alt. 900-950 m.,
Feb. 7, 1925, Brenes 4206 (218) (fl., TYPE, ea pee 5-7 m. with conical-globose crown;
flowers small, white, in terminal bee of 2 3-flowered cymes) ; woods, Piedades of
San Ramon, alt. 1000 m., June 21, 1925, poke pee (fr., Ch) (small tree 5-6 m.,
fruits green, pendent; cupule ey red, same length as the peduncle, round or heany
so, small); hedges, woods, San Miguel de San Ramon, Brenes 5393 (537)> (i..-Chi):
Ch); between “Pata de Gallo” and Santiago de San Ramon, Brenes 6650 (fl., Ch) ;
Camino at Calera and Calera de San Ramon, Brenes 18942 (fl., Ch); [hills of San
Pedro de San Ramon, Brenes 20332 (fl., Ch)]; Barranca of San Ramon, A. Smith
P.2351 (fl, A). Panama: Chiriqui: Cerro Horqueta, Boquete, C. & V.
von Hagen 2118 (fr., fragm., A, Mo).
NATIVE NAMES: ‘“Aguacatillo” (Costa Rica); “Sigua blanca” (Panama).
his species seems to be r N. nervosa Mez & Pittier. The latter,
however, has ily iene nates or elliptic-lanceolate leaves up to 13 cm.
long, glaucous peas and reddish green beneath. The a is
squarrosely tripinnately paniculate. The fruit is ellipsoid, mm.,
with a semi-globose acutely simple-margined cupule, ies aaa in the
dried state. These characters separate the two entities at once.
386 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
19. — coriacea (Swartz) Grisebach, Fl. Brit. W. Ind. 281. ee ie in Jahrb.
. Gart. Berlin 5: 459. 1889; Allen in Addisonia ix he pl. 709
mci coriacea Swartz, Prodr. 65. 1788; Fl. Ind. Occ. . 1800.
— Willdenoviana Nees, Syst. 321. 1836; Lyrae in DC. Prodr. 151: 165.
cord in Field Mus. Publ. Bot. 12: 143.
Sie aac: Florida, West Indies, Yucatan ak British Honduras, and
Guatemala.
[Jamarca: Without locality or collector, (rypE of Laurus oo not seen).]
Mexico: Yucatan: without locality, Gawmer 23980 (fl., Ch, GH, NY), 24248 (11,
Ch, GH), 24274 (fl., GH, NY), 24290 (fl., Ch), 24337 (f1., Ch) ; rile bee 85
(fl., Ch); Chichen Itza, off Kaua road in advanced deciduous forest, C. L. & A. A.
Lundell 7435 (fl., A); Lake Chichankanab, Gaumer & Sons 23655 (fl., and 23668
(fl., Ch, GH). Campeche: Hacienda San Pablo, near Champoton, Morelos, Collins
47a (fl., NY); Tuxpena, Lundell s.n. (fl., Ch), 1071 ({r., Ch), 1367 (fl, NY). Guate-
MALA: Petén: Lake Petén, Lundell 3195 (fl., Ch), Ixlu ruins, Lake Petén, June 15,
1933, Lundell 4359 (fl. type of — Lundellii, Ch) ; Uaractin, Bartlett 12335 (fl,
A, Ch, NY), 12547 (fl. A, Ch, NY). Alta Verapaz: Woods southeast of Finca
Fslemmncle. near Alta Verapaz- Sah icintaie line, Steyermark 45216 (sterile, Ch).
Izabal: Rio Dulce, between Livingston and 6 miles up river on north side (right
hand side going up river), Steyermark 39402 (fl., Ch). British Honpuras: Orange
Walk: C i Camp, Lundell 423 (fr., Ch, NY); Belize:
Maskall,Gentle 1206 (fl., A, Ch, GH, NY), 1216 (fl, A, Ch, GH, NY). Toledo:
Jacinto Hills, in shaded fe Schipp 1206 (fl., Ch, GH, NY
NATIVE NAMES: “Laurel” ene aninis “Sweetwood” (Jamaica, British Honduras).
This species, = it occurs in our has branchlets that vary from
brownish and striate ee a striking silver -gray. The alternate leaves are
borne on herr that are usually 1 cm. or less in length (7-14 mm.), cana-
liculate, and glabrous. The 5 Ae are coriaceous, glabrous, lanceolate-
elliptic or lanceolate-oblong, the base cuneate, the apex obtusely acute or
obtusely acuminate, the acumen varying in length from very short and
abrupt to nearly 1 cm. The blades measure up to 12 cm. long and 6 cm.
broad (usually 7-9 & 3.5-4cm.). The venation of this species is the most
erin Lunde Standley in Carnegie Inst. hire Publ. 461: 56. 1935; Stand-
ey 936
@
°
p
wn
ae
2
ed
ig:}
a
—_ 2
ms
iY)
°
‘<
nerves diverge from the costa at an angle of 45° and are slightly arcuate.
The lateral nerves, however, are usually obscured by the very conspicuous
over-all rather loose elevated reticulation of the blades. This reticulation
usually draws the attention before the lateral nerves are note The inflo-
rescence varies from few to many axillary or subterminal panicles not more
than 6 (—8) cm. ne, either sparsely or densely flowered. The white
flowers are 5—8 mm. in diameter, the elliptic lobes heavily papillose, + 3.6
mm. in length. The two outer series of stamens consist of nearly sessile
anthers which are reniform and slightly emarginate, measuring +-0.6 mm,
in length. The stamens of the inner series are longer, +-0.8-1 mm. long,
the squarish anthers borne on filaments almost their equal in length, which
bear two large subreniform sessile glands nearly the width of the anthers.
The staminodia are triquetrous, 0.8 mm. long, borne on slender pubescent
filaments that are nearly two-thirds the length of the entire structure. The
gynaecium is glabrous, +1.7 mm. long. The subglobose or subovoid ovary
is about twice the length of the stout style, which is topped by a conspicuous
capitate stigma. The fruit is black, ovoid or subglobose, apiculate,
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 387
1 (—1.5) cm. long and 6 mm. wide, at maturity supposedly nearly ay same
width. The fruit is subtended by a shallow cyathiform cupule
long and 6-7 mm. in diameter at the undulating apex, and 2.5 mm. a deep.
The eee pedicel is up to 8 mm. long, expanding to 2 mm. in diameter
at the a
The et affinity of the species is the entity which heretofore has
apart. Standley notes that Ocotea Lundelli is most nearly related to
Ocotea Catesbyana Sargent from southern Florida. The latter was based
on Laurus Catesbyana Michaux, which was once included erroneously under
N.coriacea. Although the superficial resemblance is striking, the structure
of the flowers, particularly the reniform anthers, immediately places the
species in Nectandra
20. Nectandra salicifolia (H.B.K.) Nees, Syst. Laurin. 302. 1836, in Linnaea 21: 506.
1848.
mee salicifolia H.B. a ae Gen & Sp. 2: 132 [166]. 1817; Hooker & Arnott, Bot.
Voy. Beechey 309
Ocotea globosa sensu a renia & Chamisso in Linnaea 6: 366. 1831.
Nectandra sanguinea sensu Nees, Syst. Laurin. 318. 1836, quoad spec. Mex., non
Rolander ex Rottboell; Meissner in DC. Prodr. 151: 164. 1864, quoad spec. Mex
Hemsley, Biol. Centr. Am. Bot. 3:75. 1882; Mez in Jahrb. Bot. Gart. Berlin
¢ C. Am
Weesnnira Viabrestens Bentham, Bot. Vo se: Sedphiie 161. 1846; Walpers, Ann. 1: 575.
1849; Meissner in DC. Prodr. 15!: ve 1864; Hemsley, Biol. Centr. Am. Bot.
3: 74. 1882; Mez in Jahrb. Bot. Gart. Beilin 3: 425. 1889, excl. spec. S. Am. & W.
Ind.; Standley in Contr. U. S. Nat. Herb. 23: 297. 1922; Standley & Calderon,
Lista Prelim. Pl. Salvador 84. 1925; Record in Trop. Woods 10: 21. 1927; Standley
in Field Mus. Publ. Bot. 10: 200. 1931; Standley & Record in Field Mus. Publ.
Bot. 12: 142. 1936; Yuncker & Record in Field Mus. Publ. Bot. 17: 363. 1938.
Nectandra sanguinea var. angustifolia Schlechtendal in Linnaea 19: 257. 1847.
Nectandra sanguinea var. 8 lanceolata Meissner in DC. Prodr. 151: 164. 1864; Hems-
ley, Biol. Centr. Am. Bot. 3: 75. 1882
Disraipe iow: Mexico and Central America, at varying altitudes and in diverse
habit
oe Without locality, Galeotti B.8 (fl., Ch), [718] (fl., Ch) ; Haenke [1540]
(fr., NY); Leibold 108 (fl. type of N. sanguinea var. angustifolia not seen). San
Luis Potosi: El Banito, 7 mi. south of Valles, Leavenworth 189 (fl., Ch, GH) ;
semi-desert road-side aie of Valles, Leavenworth 223 (fl., Ch, GH, NY); Tamasopo
Canon, Pringle 3541, 3725 (f1., GH, NY); Tamazunchale, ands 602 (young fr., Ch).
Vera Cruz: Pacieada de la Laguna, Schiede 57 (fr., Mo); Papantla, Schietle &
Deppe 1144 (fl., cited by Schlechtendal and Chamisso as O. globosa, Mo); road from
Papantla to Zamora, Goldman 88 (fl. GH); Colipa, Liebmann s.n. (fl., GH, NY);
Jalapa, [Schiede?] (fr.. GH); Mirador, Liebmann s.n. (fr., GH) ; Sea Purpus
2023 (fr., Ch, GH), 14151, 14327 (fl., Ch); Barranca de Peace: Purpus e893.
(fl., GH) ; Remulatero, Purpus 8741 (fl. GH, NY); Remudadero, Purpus eh a GH,
NY), 10965 (fl., Ch); banks of Rio de los Pescados, near Puente Nacional, Purpus
11166 (fl., Ch) ; Orizaba, Botteri 34, 302 (fl, GH), 986 (fl., NY), 1081, 1193 (fl., GH),
jel sae Herb. sn. (fl., Ch), Purpus 1324 (fl. GH, NY); Cordoba, Bourgeau 1961
( NY), Orcutt 3344 (fl., Ch); Cuitlahuac, Matuda 1417 (fl., A). Mexico:
loa Salitre-Canitas, wet barranca, Hinton 3816 (fl. A, NY). Guerrero:
La Aguila, Langlassé 250 (f1., GH); Acapulco, Humboldt & Bonctond s.n. (fr. ?, TYPE of
Ocotea salicifolia not seen), E. Palmer 164 (fr., GH), 582 (fl, GH, NY). Oaxaca:
388 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
Tuxtepec, Chiltepec and vicinity, in llanos, Martinez-Calderén 376 (fl., A); Piedras
Negras, Pochutla, Conzatti, Reko & Makrinius 3203 (fl. GH); Yaveo, trail west to
Rio Yaveo; Choapam, understorey in open forest, Mexia 9175 (fl., Ch, GH, NY);
Vunmitedl inl a Rio Verde, Conzatti 4385 (fr.. NY). Tabasco: eevedea Balan-
can, Matuda 3027 (fl., A, Ch) ; Tenosique, Matuda 3402 (fl., A, Ch, NY). Chiapas:
Santa Rosa, near Escuintla, in advanced forest, Matuda 4264 (fl, A, NY); Malpaso,
near Siltepec, in advanced forest, Matuda 4522 (fl., A). GUATEMALA: Pe
Paso, Lundell 1528 (fl, NY); Uaxactun, near acivada: Bartlett 12355 (fl., Ch); Santa
Cruz, 12371 (fl., Ch), 12724 (f1., Ch); 7 Libertad and vicinity, Aguilar 91 (fl., A),
280, 387 (f1., NY), Lundell 2113, 2176 (fl., Ch) ; along Rio Santa Izabal, between mouth
of Rio Sebol and El Porvenir, Steyermark 45847 (f1., Ch) ; along Rio Cancuen, between
El Cambio and mouth of Rio Machaquila, Steyermark 45921 (fl., Ch). Alta
Verapaz: Between Sachaj and Sacacac, Steyermark 45130 (fl., Ch); along Rio
Semococh, between Semococh and Chajamayic, Steyermark 45731 (f1., Ch); along Rio
Sebol, between Sebol and Carrizal, north of Sebol, Steyermark 45757 (11., Ch); Cubil-
guitz, von Tuerckheim 8577 (fl, GH, NY). Baja Verapaz: Sierra de las Minas,
San Augustine, Kellerman 7628 ({fr., Ch). Izabal: Rio Dulce, between Livingston
and 6 miles up river, on north side (right hand side going up river), Stevermark 39397
(fl, Ch); bank of Rio Dulce, C. L. Wilson 406 (f., Ch). Quezaltenango
Colomba, Skutch s.n. (fl, Ch), 1286 (f1., A, NY), 7363 (f1., A, Ch). Honpuras:
Without a forest along Highland Creek, Puerto Sierra, P. Wilson 145 (f1., NY).
Comayagu Thicket above the plains of Siguatepeque, Vuncker, Dawson & Youse
6144 (il., Ch, GH, NY). Yoro: In open forest on margin of small stream on
mountain-slopes near the village of Los Flores, in the Aguan River valley, near Coyoles,
Yuncker, Koepper & Wagner 8161 (f1., Ch, GH, NY), 8762 (fl, Ch). Atlantida:
On the mountain-slopes and coastal plains, in forest, foothills near the Cangrejal River,
in the vicinity of La Ceiba, Yuncker, Koepper & Wagner 8680 (fr., Ch, GH, NY).
Britisu Honpuras: Without locality, Castillo 16 (11., Ch, Y), Gentle 174 (f1., Ch),
Northern River, Gentle 1035 (f1., NY). Corozal: High ridge, San Joaquin, Gentle 8
(fr., Ch), 4748 (fr., Ch, GH, NY). Orange Walk: Roaring Creek, Lundell 448
(f1., Ch); Honey Camp, Meyer 54 (fr., Ch). El] Cayo: Bank of Belize River, Gentle
2232 (fl, A, NY). Belize: Little aan ar Belize River, Lundell 4361 (fr., Ch),
md
4362 (f1., Ch, NY), 4363 (fr., Ch); kall Pine es Gentle 1055 (f1., GH, NY),
1275 (fr., Ch, NY); Sibun River, Sern 1410 (fr., , NY), 1511 (fl., NY), Gracie
Rock, 1524 (fl. NY). Stann Creek: io a Freshwater Creek, Kelly 2
(f1., Ch, GH, NY); Moho River, in wet , Peck 737 (fl. GH). Costa Rica:
San José: Vicinity of El General, Shuck aie (fl, A, NY), 4328 (fl, A, Ch, NY),
4905 (fl., A, Ch, NY).
Native NAMEs: ‘‘Aguacate’” (Vera Cruz); ‘“Aguacatillo” (Michoacan); ‘Laurel’
(Vera Cruz, Yucatan, British Honduras); ‘“Piesito de Paloma” (Mexico); ‘“Timber-
sweet” (British Honduras)
This is one of the most variable of the Nectandrae, which has been known
heretofore as N. sanguinea Rottb., and is of all species in our area the most
widely collected. Originally the species Vectandra sanguinea was described
from Surinam. Nees extended the range to Martinique, other islands of
the West Indies, and Mexico, describing a variety from St. Vincent. He
included Laurus globosa Aublet from Guiana in the synonymy. Meissner
followed Nees, adding another variety from Mexico which included in part
Schiede 57 and in toto Ocotea salicifolia H.B.K. Mez segregated Laurus
globosa Aublet, under the binomial Vectandra globosa. According to
Kostermans (Meded. Bot. Mus. Utrecht 25: 20. 1936), Mez did not see the
type of V. sanguinea, based on Rolander’s specimen. Kostermans intimates
that the latter name belongs to NV. Pisi from Surinam, and that V. sanguinea
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 389
as interpreted by Mez (presumably the Mexican, Central American, and
West Indian material) does not occur in Saran A complete under-
standing of the synonymy involved must perforce await examination of the
types of the species in question, when they are once more available for
study. Insofar as may be ascertained from the specimens at hand, the
correct epithet for the Mexican and Central American material appears to
be N. salicifolia (H.B.K.) Nees. There can be no question of Mez’
segregation of Laurus globosa Aublet as Nectandra globosa, for the shape
of the anthers of the latter places it in a different category entirely from
N. salicifolia.
Nectandra salicifolia, then, may be described as a tree or shrub with
branchlets that are angled, becoming terete, striate, brownish and becoming
grayish, or sometimes reddish and becoming brown. They are early whitish-
pubescent, later becoming glabrous and often shining. The leaves vary con-
siderably, with the petioles up to 1 cm. long, canaliculate, glabrous or gla-
brescent. The blades are chartaceous to coriaceous, lanceolate, lanceolate-
elliptic, or elliptic, cuneate at the base and either acute or obtusely acute or
obtusely acuminate at the apex. They measure (4—) 5.5-16 (—20) cm.
long and (2—) 3-5 (—9) cm. broad and are glabrous except for pubescent
glands in the axils of the lateral nerves on the lower leaf-surface, which are
sometimes very conspicuous or often entirely lacking. The costa is im-
pressed above, very prominent beneath; the 5—9 pairs of lateral nerves are
slightly elevated beneath and diverge at an angle of 35—45° from the costa.
The reticulation of the blade is usually very conspicuous throughout. The
subterminal or axillary inflorescences are shorter or occasionally longer than
the leaves and are up to 10 cm. long. They may be comparatively few-
flowered single panicles or, as is usually the case, many-flowered broad and
branching subcorymbose panicles. In the early stages they are whitish-
pubescent but shortly become glabrescent or glabrous. The flower is
usually between 3 and 4 mm. long (occasionally 5 mm.), with scarcely any
noticeable tube, and is supported by a slender pubescent pedicel 3-4 mm.
long. At anthesis the perianth-lobes are spreading (the flower being some-
times to 1 cm. in diameter) and reflexed. The lobes are lanceolate-elliptic,
occasionally elliptic, usually obtuse, up to 3 (sometimes 5) mm. long, fleshy,
pubescent on the outer surface and papillose within. The stamens of the
two outer series are +0.6 (rarely +0.8) mm. long and nearly half again as
broad. The anthers are reniform, frequently emarginate, sessile or with
very short filaments. The stamens of the third series are longer, with the
anthers almost square and emarginate at the tip. The two upper cells are
lateral, whereas the two lower are extrorse. The filaments bear at their
base two large conspicuous glands almost as long as the anthers. The
slender small triangular or triquetrous staminodia are usually +0.6 mm.
, borne on stipes nearly 24 their entire length and almost the same
width as the staminodia. The glabrous gynaecium is +1.25 (sometimes
+1.9) mm. long. The subglobose ovary is usually three times the length
of the short style, which is topped by a flat inconspicuous stigma. The
fruit is subglobose to (immature?) ellipsoid, minutely apiculate, to 11 mm.
long and 8 mm. in diameter, glabrous, black, subtended by an almost disk-
like shallow woody cupule about 1 mm. long and 5 mm. in diameter, gla-
_brous and only slightly undulate. The cupule is supported by the enlarged
390 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
pedicel about 5 mm. long, expanded to 3 mm. in breadth at the apex.
Most specimens collected are in the flowering stage and show variations
which are definitely within the limits of the species. Occasionally speci-
mens are found in which no staminodia are seen, but this is not usual. The
length of the short filament supporting the anther varies considerably.
The greatest amount of variation occurs in the specimens from Acapulco,
which have larger flowers than the other Mexican specimens.
21. Nectandra fuscobarbata (Mez),
Nectandra glabrescens var. ee Meri in Jahrb. Bot. Gart. Berlin 5: 425. 1889.
Arbor 6-12 m. alta, ramulis brunneis minute sparse pubescentibus mox
griseis glabris striatis. Folia alternata, juventute sparse pubescentia mox
glabrescentia deinde glabra, petiolis tenuibus, canaliculatis, supra sparse
pubescentibus, ad 1 cm. longis et | mm. latis, laminis utrinque glabris basi
costae subtus excepta, membranaceis, in sicco brunneis vel viridescenti-
brunneis, lanceolato-ellipticis, ad 15 cm. longis et ad 5 cm. latis, basi per-
attenuato-cuneatis, apice longe caudato-acuminatis, penninerviis, costa
supra leviter subtus conspicue elevata, nervis 4-6 (—8) -paribus utrinque
plus minusve obscuris angulo 55° divergentibus, glandulis inconspicuis in
nervorum lateralium axillis, rete venularum utrinque leviter prominulo.
Inflorescentia axillaris laxe paniculata, ad 15 cm. longa, sparse pubescens,
pedunculo brunneo sparse pubescente, ad 5 cm. longo. Flores ad 3 mm
longi, pedicellis pergracilibus pubescentibus, ad 3 mm. longis, perianthio
campanulato albo vel pallide viridescente, lobis oblongis reflexis papilloso-
pubescentibus, +2.5 mm. longis; staminibus ser. I & II +0.6 mm. longis,
antheris subreniformibus longitudine 24 filamentis gracilibus basi pubes-
centibus aequalibus, ser. III +1 mm. longis conspicue biglandulosis,
glandulis stipitatis antheris oblongis et filamentis aequalibus; staminodiis
ovatis stipitatis, basi pubescentibus, +0.6 mm. longis; gynaecio glabro,
+1.25 mm. longo, ovario ovoideo-globoso longitudine plusquam stylo
minusquam gynaecii aequante, stigmate capitato conspicuo, Fructus
abnormalis(?) corsaggeriea Sackett conspicue griseo-sericeus, minute
papillosus, +7 mm. diam., cupula vadosa glabra minute verruculosa sub-
campanulata 3 mm. longa et 5 mm. diam. subtentus, pedicello ad 5 mm.
longo apice ad 3 mm. diam. expanso.
DISTRIBUTION: Known only from Panama.
Panama: Without locality, Hayes 487 (vype of N. glabrescens var. fuscobarbata
not seen). Bocas del Toro: Fish Creek Hills, vicinity of Chiriqui Lagoon,
von Wedel 2431 (f1., A); Isla Colon, von Wedel 2866 (fl., A), 2969 (fl., A) ; Flat Rock,
region of Almirante, G. P. Cooper 551 (fr., Ch, GH, NY)
NATIVE NAME: “Rock Sweetwood” (Panama).
This entity, seemingly a segregate of NV. salicifolia, because of the single
character of fuscous barbate hairs in the axils of the leaves, has flower-
structure similar to that of the latter species. Examination of a large
amount of material from Mexico and Central America of the species proper
shows many numbers with pubescent glands in the axils of their lower
leaf-surfaces. The bases of the filaments as well as the stipes of the
staminodia are pubescent. The ovary is glabrous, but the fruit is
pubescent. I am inclined from the general aspect of the fruiting material
rom Bocas del Toro to believe that these are not normal fruits.
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 391
22) pe cue nitida — in seg Bot. Gart. Berlin 5: 461. 1889; Standley in U. S.
erb. 23: 29
Puan oe and Panama.
Mexico: Without locality, western part, Haenke sm. (SYNTYPE not seen).
PANAMA: ana one: Near Barbacans station, Hayes 133 (SYNTYPE not seen).
This species seems to stand out for its golden-‘‘glittering’’-tomentose
young branchlets, ae becoming cinereous, glabrous, and terete, and
its golden-lanuginose buds. The petioles are not more than 6 mm. in
length, are ee subtending chartaceous leaf-blades that in young
stages are sericeous-lanuginose, golden, and glittering. See V. latifolia for
discussion.
23. Nectandra perdubia Lundell in Lloydia 4: 47. 1941.
DistripuTion: Mexico and Central America, for the most part at fairly low alti-
tudes. Frequently to be found along stream-banks.
Michoacan: Coalcoman, Hinton 13667 (fr.,. GH), 13668 (fl., GH),
13856 (fl., GH), 13918 (fl, GH). Oaxaca: Vicinity of San Juan Guichicovi, Nelson
2725 (fi., fr., Ch) ; Ubero, ZL. Williams 9492 (fl., A, Ch, NY). Tabasco: Boca Cerro,
Tenosique, vals 1-5, 1939, Matuda 3576 (fl., isorypr, A, Ch, NY). GwuaTEeMALa:
Uaxacttin, Bartlett 12708 (fl., Ch); Santa Teresa, Subin River, Lundell 2735
(fr., Ch); La Libertad, Lundell 3348 (fr.. Ch), 3716 (fl, Ch). Alta Verapaz:
Vicinity of Secanquim, Pittier 178 (fl., Ch); vicinity of caves, southwest of Lanquin,
Steyermark 44073 (fr., Ch); along route no. 5 between Chirriacté and Semococh,
Copan: Along trail from El Paraiso to La Florida, Pittier 8468 (fl., NY). Britisu
Honpuras: Without locality, hill-bank, Record B.H.30 (¥ 8798) (fr. ¥Y). El Cayo:
El Cayo and vicinity, Chanek 58, 61 (fl, Ch). Belize: Gracie Rock, Sibun River,
ae 1692 (fl. A, NY). Stann Creek: In valley, Big Eddy Ridge, Gentle 3543
(He, NY). Costa Rica: Puntarenas: Puerto Jiminez de Osa (Golfo Dulce),
Brens 12163 (642) (fl., Ch); woods on sea-shore, Golfito de Osa, Brenes 12314 (793)
(fl., Ch). San José: Near San José, edge of Rio Virilla at Uruca, Tonduz 7271
(Herb. Nat. oe Rica 10104) (fl, GH, NY). Cartago: Rio Turrialba, J. D
Smith 4932 (f1., GH).
NATIVE NAMES: ‘‘Aguacatillo”’ (Mexico) ; ‘‘Bastard Timbersweet,” “Laurel” (British
Honduras).
A species near NV. nitida, according to the author, but distinguished from
it by the narrower leaves which are less reticulate and more smooth and
shining on the upper surface. The younger branchlets and inflorescences
retain their pubescence longer than do those of NV. nitida. ‘The latter, as
well as NV. perdubia, has the same basic floral structure as is found in NV.
salictfolia.
he species has striate branchlets which are clothed with a minute close
ee ee tomentum which rubs off early, leaving them dark
ray or occasionally darker brown. ‘The leaves are borne on stout tomen-
tose scarcely ae petioles up to 1 cm. long. The lanceolate to
elliptic, acuminate to subcaudate, leaf-blades are cuneate at the base and
measure up to 17 cm. (oe and 6 cm. broad. The upper surface is early
pubescent, but shining at maturity, the 7—9 lateral nerves diverging at an
angle of about 35°, and the costa impressed and not prominent. The lower
surface is everywhere minutely pubescent at maturity, with scattered floccose
pubescence in addition, particularly about the region of the veins, which
392 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXvI
bear more or less pubescent inconspicuous axillary glands beneath. The
lateral nerves and costa are prominently elevated beneath and pubescent.
The lower surface is definitely but loosely reticulate. The inflorescences
are axillary and subterminal many-flowered panicles, densely covered with
a minute ferruginous tomentum which becomes less with age. They are
subtended by sericeous lanes bracts which presently are deciduous. The
flowers are 5.5—-7.5 mm. in diameter, the tube short, the elliptic perianth-
lobes rather thin or siinniace thickish, fleshy and papillose, up to 3.5 mm.
long, tomentose without. The stamens of the two outer series are +0.8
mm. , the subreniform anthers once and a half or twice the length of
the stout ‘filaments, ~— are sometimes pubescent. Those of the inner
series are +-1.25 mm. long and have anthers which are almost square,
emarginate, the two upper cells lateral, the lower extrorse. The filaments
nearly equal the anthers in length and bear at the base two large subreni-
form sessile glands nearly the length of the anthers. The staminodia are
+0.6 mm. long, variable, often triquetrous, the usually somewhat slender
stipe nearly half the entire length. The glabrous gynaecium measures
+ 1.25 mm., the subglobose ovary constricted at the base and about three
times the length ot the slender style which bears at its apex a usually con-
spicuous subcapitate stigma. The fruit (immature?) appears to be small
(6.5-10 mm. long and 5.5-8 mm. broad), subglobose, apiculate, subtended
by a shallow % athifor m cupule 1.5-3 mm. long, up to 7 mm, in diameter,
and less than 2 mm. deep. The peel enlarged pedicel is up to 5 mm.
long and anne to 3mm. at the ap
24. Nectandra pigeons (HL. B.K.) Mez in Jahrb. Bot. ae Berlin 5: 454. 1889.
9]. 1817.
Ocotea latifolia . Nov. Gen. & Sp. 2: 133 [16
DistripuTIoN: Central America from Nicaragua pens Panama, south through
Colombia to Brazil, ke to Mez.
PAN > Canal Zone: a! locality, Christopherson 132 (f1., NY); hills
near ei Station, Panama R.R., Feb. 7, 1860, Hayes s.n. (fl, GH, US), 59 (fr., GH) ;
Gatun Lake at turning point from canal. Bangham 425 ti Ch); Chagres, Pesdioe J4
(f1., Ch, GH, Mo, US); Barro Colorado Island, Aviles 113 fe. Ch), tgs _ Ch),
L.H.& E. Z. Bailey 92 (fl., Ch), 307 (fr., Ch) ; Woodworth & Vestal 692 (fl., A, Ch).
(Corompia: Cundinamarca: High plains of Bogota, kaise & Rowslead
({r., TYPE not seen). |
This species is very distinct because of its foliage characters. The leaves
have petioles 6-12 mm. long, pubescent to glabrous, and canaliculate. The
blades, becoming glabrous except for inconspicuous axillary glands, are
chartaceous to subcoriaceous, very shining above, dull beneath, elliptic.
the base cuneate to roundish, the apex caudate-acuminate, and they measure
about 12 (—-15) cm. long and 3.5 (—6) cm. broad. The costa and lateral
nerves (4-6 pairs) are somewhat obscure above, elevated beneath, and
diverge from the costa at an angle of 35-50 (~55)°. The minute reticula-
tion likewise is exceedingly prominent above and beneath. The sub-
corymbose-paniculate axillary and subterminal re inflorescences,
minutely pubescent, becoming glabrous, measure up t (—13) cm. in
length, the peduncle up to § cm. long. The flowers measure about 6 mm.
in diameter, with thick papillose perianth-lobes, haa are elliptic- ovate
and + 2.15—-2.5 (-3.4) mm. long. The stamens of the two outer series are
+0.6-0.8 (—1) mm. long, the anthers subreniform, slightly emarginate,
twice the length of the rather stout filaments. Those ‘of the inner series are
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 393
+.8 (-1) mm, long, the squarish anthers about bear eres yee filaments,
which bear conspicuous sessile glands nearly the size of the anthers.
staminodia are ovate, ++0.6 mm. long, the stipes pee pense to two-
thirds their entire length. The glabrous gynaecium is + 1.25 mm. long, the
subglobose ovary nearly three times the length of the thick short style that
is topped by a subcapitate stigma. The subglobose black fruit is ea
1 cm. in diameter and is seated on a shallow cupule not more than 2 n
long, 6 mm. in diameter, and 0.5 mm. deep, the margin slightly and fnely
undulate. re pedicel is enlarged to 3-4 mm. long and 2.5 mm, in diam-
eter at thea
Mez ser Fendler 54 under N. latifolia, vege al from
Colombia, and it seems to agree fairly well with the descripti c
species N. nitida, described from Mexico and the Ca Zone as well, seems
to be very close to the so-called V. latifolia, differing in leaf-blades that are
prominently reticulate above and in the young stage scarcely sericeous
beneath, the absence of axillary glands, the branchlets that are sparkling
golden-tomentose and the buds that are golden-lanuginose. The flowers
also are golden-brown-tomentellous, but their structure is not strikingly
different from those of N. latifolia. Examination of the types of both
species in question is necessary before an accurate disposal may be made of
the material at hand.
25. Nectandra Cufodontisii (O. C. Schmidt), comb. nov
Ocotea Cufodontisii O. C. Schmidt in Arch. Bot. Forli 11: 50. 1935.
DiIstRIBUTION: Known only from Costa Rica.
Costa Rica: Alajuela: Alfaro Ruiz, La Brisa de Zarcero, in loam and mold in
semi-shade of forest in subtropical zone, A. Smith H.969 (f1., Ch, NY), H.972 (f1., A,
; Palmira, A. Smith P.2036 (fl., fr.. A). Cartago: On the southwest slope of
Velen Irazu, at the mouth near San ees alt. 2000 m., May 30, 1930, Cufodontis 315
(fl., fr., isoTyPE, Ch, photo., Ch, NY) (tree 10 m., with broad crown; flowers greenish).
The young branchlets of this species are angled or almost sulcate, at first
subferruginous-sericeous, becoming glabrescent, presently glabrous and
grayish. The leaves are borne on ene petioles pubescent and up to
1.5 cm. in length. The blades are elliptic or, according to the author,
subovate- aa rae acuminate, the base cuneate, See ae to subcori-
aceous, up to 13 cm. long and 5 cm. broad. The e upper surface, according
to the author, is nea opaque, whereas the lower is almost shining, the
entire surface glabrous except for the pubescent axillary glands. The costa
and lateral nerves, of which there are 7—9 pairs diverging from the costa at
an angle of 35—45°, are very slightly elevated on the upper and more so on
the lower surface. The entire surface is conspicuously reticulate. The rather
few-flowered axillary or Oe panicles are minutely subferruginous-
pubescent, up to 12 cm. long, and borne on slender pubescent peduncles up
to 7 cm. long. The greenish ieee are spreading- campanulate, the heavy
fleshy cries oblong-ovate perianth-lobes measuring up to 3 mm. in
length. The stamens of the two outer series are +1.5 mm. long, the
rounded ovate-globose anthers supported by short stout filaments pubescent
at the base and one-quarter = entire length of the stamens. Those of the
third series measure -+-1.7 mm. and have anthers that are almost square or
subtrapezoid, the filaments dae equalling the anthers and bearing two
large conspicuous sessile glands at the base that equal them in length.
394 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
The author does not mention the staminodia, which are conspicuous,
cordate, +0.6 mm. in the type, with the stipes one-half the entire length.
Occasionally the staminodia are also biglandular, and occasionally only
scale-like. The glabrous gynaecium measures + 2.15 mm., the subglobose
or broadly ovoid ovary more than equalling the stout style, which bears an
often conspicuous capitate stigma. The fruit is ovoid-elliptic, up to 2.5 cm.
long and about 1.5 cm. broad, borne in a shallow almost disk-like cupule,
which q glabrescent, brown, striate (in the dried state), about 2 mm. long,
7 mm. in diameter, and 0.75 mm. deep, the margin undulate. The sup-
porting mera is glabrescent, also brown and striate, not more than 4 mm.
long and expanded to 2 mm. at the apex.
A fruiting specimen, Standley 47364 from near San José, has young
fruits that resemble those of the type, but the leaves do not show the minute
reticulation of those of the latter. Schmidt relates the species to Ocotea
insularis. ‘To my mind, it is more nearly related to Nectandra latifolia,
differing in the elliptic acuminate rather than oblong-elliptic caudate leaf-
blades, the shorter, ovate-oblong rather than strap-shaped perianth-lobes,
and the ovoid-ellipsoid instead of globose fruits.
26. Nectandra tabascensis Lundell in Lloydia 4: 48. 1941.
DistrRIBUTION: Known only from Mexico.
Mexico: Jalisco: Santa Cruz de Vallanta, wooded ravine on mountain-side,
Mexia 1264 (fr.. GH, NY). Guerrero: Montes de Oca, by the river, Hinton 10589
(f1., GH), 77473 (fr.. GH). Tabasco: La Palma on the San og de Martir River,
near the Petén border, Balancan, Matuda 3299 (fl., isorype, A, GH an
NATIVE NAMES: “Aguacatillo,” ‘“Aguacatillo blanco” ae
This species is similar in floral structure to V. salicifolia and also to V.
Woodsoniana. The young branchlets, early fulvous-sericeous-tomentellous,
later become glabrescent, dark brown, and striate. The petioles are slen-
der, up to 1 cm. long and sericeous, becoming glabrescent. The leaf-blades
are chartaceous, lanceolate-elliptic or lanceolate-oblong, cuneate at the base,
the apex acute or subacuminate, sparsely grayish-pubescent throughout,
more conspicuously so along the midrib, up to 20 cm. long and 5 cm. broad.
The costa and lateral nerves, of which there are 11—13 pairs, are slightly
elevated on the upper and more prominently so on the lower surface. The
reticulation is loose and conspicuous throughout. The inflorescence is
axillary, whitish-pubescent, becoming glabrescent, subcorymbose-paniculate
or paniculate, up to 11 cm. long, and subtended by a slender sparsely pubes-
cent peduncle up to 6 cm. long. The flowers are whitish-tomentellous,
about 3 mm. long and about 6 mm. in diameter at anthesis, subtended by a
slender pubescent pedicel up to 3 mm. long. The perianth-tube is con-
spicuous and the lobes are slightly unequal, rather fleshy, the outer broadly
elliptic or ovate and the inner oblong, +2.5 mm. long. The stamens of the
outer series are roundish-subreniform, sometimes emarginate, +0.75 mm.
long, the anthers slightly longer than the stout filaments. Those of the
inner series are +-0.8 mm. long, the squarish anthers about equalling the
filaments in length. The filaments bear at the base two subglobose sessile
glands nearly the length of the anthers. The broadly ovate staminodia
measure +0.4 mm. and are borne on short stipes that are about one-third
the entire length of the staminodia. The glabrous gynaecium is +1 mm.
long, the subglobose ovary twice the length of the short stout style that is
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 395
topped by a rather inconspicuous stigma. The material from Guerrero and
Jalisco does not exactly match the type-collection, but the variation appears
to be intra-specific. From Hinton 11473 and Gentle 1264, the fruits seem
to be black with a bloom, glabrous, broadly ellipsoid, apiculate, and borne
in a shallow sparsely pubescent cyathiform cupule 2 mm. long at most,
7 mm. in diameter, and about 1 mm. deep, with a minutely undulate margin.
The subtending pedicel is pubescent, short, and thick, 2-3 mm. long and
about 1.5 mm. in diameter at the apex.
The narrower leaves with more numerous lateral nerves and more
persistent pubescence separate this species from the Costa Rican and
anamanian N. Woodsoniana. The floral structure is very like that of
N. salicifolia, but the foliage characters are quite different.
27. Nectandra Loeseneri Mez in Bull. Herb. Boiss. II. 5: 243. 1905.
STRIBUTION: Reported only from type-locality and vicinity in Vera Cruz and
adjacent Tamautas along the northeastern coast of Mexico
MExIc Tamaulipas: Coastal dunes north of Pacuiee: LeSueur 146 (f1., ee
Pringle a (fl, GH); vicinity of Gomez Farias, E. Palmer 272 (fl, GH). Ver
Cruz: Island of Juana Ramirez, about 56 km. south of Tampi ico, E. Palmer 458 (A,
GH); vicinity of Pueblo Viejo, 2 km. south of Tampico, £. Palmer 360 (irs, Ch, (GH,
NY); in the primeval forest, between Cazones and Tuxpam, Jan. 4, 1903, C.&G E
Seler 3696 (fl., photo. of type, GH); Cazones, Mell s.n. (fl., NY); Conteaegalcos,
isthmus of Tehuantepec, C. L. Smith 985 (fl., GH, NY), 1116 (fl., NY).
NATIVE NAME: “Laurel” (Vera Cruz).
The branchlets of this species are slender, glabrous, terete, and, according
to Mez, shining. The leaves are borne on slender petioles which are gla-
brous, canaliculate, and about 1 cm. long. The blades are chartaceous,
shining above, elliptic, cuneate at the base and obtuse or shortly
obtusely acuminate a t the apex. They measure up to 11 (rarely wi Pye
long and 3-4 (rarely a cm. broad, the loose reticulation being prominent
on both surfaces. The costa and lateral nerves, of which there are 5—7
pairs, are very slightly elevated above and prominently so beneath. The
lateral nerves diverge from the costa at an angle of 25-35 (—45)° and bear
in their axils conspicuous pubescent glands. The inflorescence consists of
more or less subcorymbose panicles 8-10 cm. long, with few or many
eae peed pubescent and borne on long peduncles up to 6 cm. The
flow e about 3 mm. long with a diameter up to 8 mm. ‘The perianth-
sibe - ce short, the lobes oblong-elliptic, 3.5 mm. long, rather thick,
and papillose on the inner surface. The stamens of the two outer series
are +0.8 mm. long, with subreniform anthers nearly twice as long as the
stout pubescent filaments. Those of the third series are + 1 mm. long and
have anthers which are squarish, emarginate, the two upper cells lateral,
the two lower lateral-extrorse, the anthers equalling the filaments in length.
The filaments bear two large sessile glands nearly the size of the anthers.
The staminodia are small, +0.8 mm. long, more or less triquetrous, sub-
tended by stipes pubescent at the base, which are nearly two-thirds the
entire length. The glabrous gynaecium is +1.4 mm. long, the ovary ellip-
soid-ovoid, four times the length of the short stout style with its flat triangu-
lar stigma. The fruiting specimen Palmer 360 seems to match very well
the other material. The fruit is globose, apiculate, black in the dried state,
about 12 mm. in diameter, subtended by a shallow cyathiform cupule 3 mm
long, 5 mm. broad, and about 1 mm. deep, with a slightly undulating
396 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
margin. The pedicel is somewhat slender, up to 5 mm. long and expanded
to about 1.5 mm. at the apex.
Although there is only a photograph of the type for comparison, there
seems no doubt that the cited numbers may be safely referred to the above
species. The nearest relative is .V. salicifolia. The shape of the leaves
and the rather more conspicuous reticulation recall the characters of
NV. coriacea, but in the latter the leaf-blades are more heavily coriaceous
and the venation obscured by the reticulation. The floral characters are
definitely those of .V. salicifolia.
28. Nectandra Skutchii, sp. nov.
Arbor ad 23 m. alta, ramulis sparse et minutissime ferrugineo-pubescen-
tibus atro- rubescentibus glabris sulcatis angulatis. Folia alternata,
petiolis — adpresse subferrugineo- pubescentibus canaliculatis, ad 1
cm. longis et 2 mm. latis, laminis supra glabris subtus basi costae excepta
glabris, Seer aon in sicco ae pallide viridescentibus subtus minus
pallidis, oblongo- -ellipticis, ad 20 cm. longis et 6.5 (—7) cm. latis, basi
attenuato-cuneatis, in petiolum aoe ibique recurvatis, apice sub-
caudato-acuminatis, penninerviis, costa utrinque castanea supra impressa
subtus valde elevata, nervis 4—6-paribus castaneis, subtus obscuris impressis
subtus elevatis arcuatis angulo 35—45° divergentibus, rete venularum utrin-
que obscuro et leviter subcancellato. Inflorescentia axillaris anguste pani-
culata, ad 8.5 cm. longa, minute adpresse pubescens, pauciflora, shige nee
ad 2.5 cm. longo. Flores ad 2.5 mm. longi, pedicellis ad 2 m ongis,
pubescentibus, perianthio infundibuliformi, albo, lobis ao ey obtusis
reflexis crassis papilloso-tomentosis, +1.7 mm. longis; staminibus ser. I &
Il +0.7 mm. longis, antheris subreniformibus emarginatis filamentis flabel-
liformibus aequalibus, ser. III +0.8 mm. longis, obovatis emarginatis big-
landulosis, glandulis et antheris filamentis aequalibus; staminodiis +0.4
n longis, stipitatis, subtriquetris; gynaecio glabro + +1.7 mm. longo,
ovario globoso 1% stylo longiore, stigmate conspicuo discoideo. Fructus
glabra 5-6 mm. longa, 10-12 mm. lata, et 3 mm. alta subtentus, pedicello
incrassato verrucoso glabro 7-10 mm. longo.
DIstrRIBUTION: Known only from Costa Rica, at altitudes of 670-850 m.
Costa Rica: San José: Vicinity of El General, alt. 850 m., Agent 1936, Skutch
2668 (fl, Type — GH, NY) (tree 23 m.; flowers white), 4182 (fr., A, NY).
The affinity of this species is with the V. globosa bene in spite of the
difference in floral structure, namely the subreniform anthers and the longer
style
29. Nectandra Standleyi, sp. nov.
Arbor 6-13 m., alta, ramulis novellis minute et dense fulvo- vel subferru-
gineo- -pubescentibus, angulatis, mox glabrescentibus, striatis griseis vel
atro-brunneis. Folia alternata, petiolis satis robustis fulvo-tomentosis
canaliculatis ad 1.5 cm. longis, laminis supra minute et inconspicue pubes-
centibus mox glabris, subtus utrinque minute adpresse pubescentibus, cori-
aceis, in sicco brunneis, ellipticis, ad 16 (—17) cm. longis et 4.5 (—6) em
latis, basi obtusis vel subrotundatis imis recurvatis et ut videtur cuneatis,
apice attenuato-acuminatis vel caudato-acuminatis, penninerviis, costa
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 397
supra impressa subtus elevata, nervis 4 (raro —8)-paribus, supra impressis
subtus elevatis, angulo 25585° divergentibus, rete venularum utrinque
obscuro. Inflorescentia axillaris vel subterminalis fulvo- vel griseo- sid
cens, paniculata, multiflora, ad 15 (—20) cm. longa, pedunculo ad 6 m
longo. Flores ad 3 m m. longi, pedicellis 2 mm. longis pubescentibus, peri.
anthio albo vel fulvo- ean fide coll., campanulato, lobis reflexis
carnosis ellipticis rotundatis +1. 7 mm. longis extus pubescentibus intus
papillosis; staminibus ser. I & II +-0.6 mm. longis, antheris subreniformibus
subemarginatis subsessilibus, ser. III +1 mm. longis, antheris truncatis,
subemarginatis, filamentis aequalibus lateraliter glandulis basi Soin
subglobosis filamentis subaequalibus; staminodiis subovoideis +0.6 n
longis, stipitibus dimidio aequalibus; gynaecio glabro, ovario stylo eal
oe aes subtriangulari discoideo conspicuo. Fructus subglo-
bosus 1 (—1.3) cm. diam. (viridis, fide coll.), cupula campanulata glabra
verrucosa margine irregulariter vadose lobata, ad 6 mm. longa, 12 mm.
diam., et 3 mm. alta subtentus, pedicello robusto verrucoso ad 8 mm. longo
et 43 1 mm. diam. apice expanso
DistrRIBUuTION: Costa Rica and adjacent Panama.
Costa Rica: Alajuela: San Luis de Zarcero, A. Smith 164 (fl., Ch); region of
Zarcero, A. Smith A.243 (fl, A, Ch, NY); (La Cidra) de San Ramon, Brenes 3850 (3)
(fr., Ch) ; without a locality, presumably San Ramon, Brenes 4061 (f1., TyPE, Ch) ;
Piedades near San Ramon, Brenes 4344 (fl., Ch); hills of San Pedro de San Ramon,
shi 4780 (565), 4794 (579), 5019 (175) (fl, Ch), 5443 (36) (fr., Ch); Cerro de
de San Ramon (El Socoro), Brenes 5714, 5733, 5831 (423), 16833 (fl., Ch).
San ary La Caja, Valerio 1326 (fl, Ch). Cartago: El Mufeco, on Rio
Navarro, Standley & Torres 50908 (fr., Ch). Limén(?): ta Concepcion, Llanuras
de Santa Clara, J. D. Sm . 6757 (fr. GH). Panama: Bocas del Toro: Crica-
mola, near Almirante, G. P. Cooper 488 (fl., Ch, NY); vicinity of Chiriqui Lagoon,
Big Bight, von Wedel Le Isla aa von Wedel 2967 (fl., A)
Native NAMES: “Aguacatillo,” “Quizzera,” “Guizarra quina” (Costa Rica); “Sigua”
(Panama)
This species is very similar to V. globosa and N. ramonensis in foliage-
characters, differing from the former in having fewer pairs of lateral nerves,
from the latter in having larger leaf-blades, and from both by its smaller
flowers with the anthers devoid of the large papillose connective character-
istic of the two above-mentioned species. The long stout style, which
equals or subequals the length of the ovary and is topped by a conspicuous
somewhat triangular discoid stigma, presents further differentiating charac-
ters. The oe cupule is less shallow in NV. Standleyi and has a tendency
to be subglobos
The ee from Chiriqui Lagoon are doubtfully referred to this
species, because of the variation in the leaf-shape and the presence of
pubescence on the small staminodia.
30. Nectandra producta, sp. nov.
Arbor ad 30 m. alta, ramulis argenteo-fulvo-sericeo-pubescentibus striatis
valde angulatis. Folia alternata longipetiolata, ee conspicue alatis
Sas vadose canaliculatis pubescentibus ad 5 cm. longis, ut videtur, et
3.5 cm. latis, laminis juventute conspicue puncticulatis, lucidis, supra
ane subtus minute moliiter et inconspicue pubescentibus, mem mbranaceis
vel chartaceis, in sicco viridibus, elliptico-lanceolatis vel leviter oblanceo-
398 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
lato- -ellipticis, ad (8—) 12 cm. longis et 2.5—4 cm. latis, basi attenuato-
cuneatis, in petiolum longum decurrentibus ibique valde recurvatis, apice
acutis vel attenuato-subacuminatis, penninerviis, costa supra flavescente
conspicua subplana et basi pubescente subtus elevata pubescente, nervis ad
10—12- paribus supra tenuibus conspicuis flavescentibus subtus obscuris
angulo 45—55° divergentibus. Inflorescentia axillaris paniculata, ad 15 cm.
longa, oe pubescens, multiflora, pedunculo ad 4 cm. longo. Flores
2.5-3 . longi, pedicellis ad 3 mm. ibneis gracilibus glabrescentibus,
ae. campanulato viridescente, lobis late ellipticis obtusis crassis
dense papilloso-tomentosis, +-2.15 mm. longis; staminibus ser. I & IT +0.8
mm. longis, antheris longis aequidem ac latis filamento duplo longioribus,
ser. IIT +1.4 mm. longis biglandulosis, glandulis et antheris oblongis oF
mentis aequalibus; staminodiis anguste oblanceolato pubescente ad +
mm, longo; gynaecio glabro, +2.4 mm. longo, ovario ellipsoideo mee
gracili aequali, stigmate discoideo satis conspicuo. Fructus ignotus
DistriBuTION: Known Raed from the type-locality, in Costa Rica.
: In forest in nae vicinity of E] General, alt. 700 m., Jan.
1939, Skutch 3006 (fl., type — A, NY) (tree 30 m., with prop-roots; flowers greenish).
Nectandra producta, so-named one of the long apparent petioles,
belongs in the same general vicinity with V. Whitet and N. hypoglauca.
The branchlets are rather like those of the latter in that they are thick,
angled, and fulvous-sericeous, becoming grayish and striate with age. The
leaf-blades are similar in many respects to those of the former species, but
are borne on extremely long apparent petioles of 5 cm. in length, formed by
the narrowly and conspicuously decurrent leaf-base, being sericeous
throughout. Even though the fruit is unknown, the species stands out for
its unusual leaves, with the blades narrowly decurrent, forming an apparent
petiole of so great a length.
31. Nectandra Whitei (Woodson), comb. nov.
Ocotea Whitei Woodson in Ann. Mo. Bot. Gard. 24: 188. 1937.
DistripuTion: Western Panama and Costa Rica, up to 2000 m. altitude.
Costa Rica: Without locality, Little 6059 (fr. Ch). Alajuela: Pastures of
La Paz de San Ramon, Brenes 4262 (47) (fr., Ch). Limén: Inland from Sequirres,
Stork 2809 (Y 38450) (fr. Y). Panama: Chiriqui: Valley of the upper Rio
Chiriqui Viejo, vicinity of Monte seve alt. 1300-1900 m., June 27-July 13, 1935,
Seibert 307 (fl., A, rypE— Mo, NY) (tree 30 m.; flowers eit greenish yellow; fruit
sae pes red; aromatic) ; trail from ‘Paso hors to Monte Lirio, P. H. Allen 1486
(fr., Ch, GH, Mo, NY
neh NAMES: “Ira,” “Ira rosa” (Costa Rica); “Bambito” (Panama).
The e present species has slender branchlets that are minutely and closely
fulvo-sericeous, becoming glabrous, grayish, and striate. The actual
petioles are slender, glabrous or glabrescent, and about 1 cm. long.
coriaceous blades are oblanceolate, narrowly attenuate at the base and
recurved, giving the appearance of a long winged petiole of 3 cm., up to
12 cm. long and 3.5 cm. broad, with the broadest portion above the middle
of the blade; the apex is obtuse or obtusely abruptly acuminate. The
young leaves are early fulvo-sericeous, becoming glabrous above and less
conspicuously pubescent beneath. The costa is slightly elevated above and
more prominently so beneath, but conspicuous everywhere. The lateral
nerves, of which there are up to 12 pairs, are not very conspicuous and
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 399
diverge at an angle of 35-45°. The axillary or subterminal paniculate
inflorescence is up to 13 cm. long, subtended by a stout reddish black
peduncle up to 6 cm. long. The pubescent light greenish yellow flowers are
about 3 mm. long, supported by a slender pubescent pedicel not more than
2mm. long. The ovate-elliptic lobes are rather thick and +2.15 mm. long.
The two outer series of stamens are --1 mm. long, the subglobose anthers
twice the length of the stout filaments. The stamens of the inner series
are +1.25 mm. long, the squarish anthers almost equalled by the filaments
and the two basal sessile subglobose glands. The glabrous gynaecium
measures +2.15 mm., the ovoid ovary slightly exceeding the rather stout
style with its subtriangular flat decurrent stigma. The fruits are oblong
or in the younger stages presumably ellipsoid, the apex in the dried state
remaining conspicuously shining and unwrinkled, drying in a more or less
regular star-shaped pattern, black at maturity, up to 4 cm. long and 1.5 cm.
in diameter, the surface frequently tuberculate. The cyathiform red verru-
cose subtending cupule is up to 6 mm. long, 13 mm. in diameter, and 2-3
mm. deep, the margin gently undulating. The pedicel is verrucose, striate,
up to 15 mm. long at times, and 6 mm. in diameter at the apex.
The long fruits distinguish this species from any other from the area
under study. It is near N. Paulii and N. producta, distinguished from the
latter by shorter apparent petioles, and from the former by its chartaceous
leaf-blades that are much smaller and with more prominent reticulation,
and by its shorter inflorescences.
32. Nectandra hypoglauca Standley, sp. nov.
Arbor 15—21 m. alta, ramulis fulvo-sericeis, mox griseis, striatis, juventute
angulatis. Folia alternata, petiolis alatis robustis pubescentibus, 2 (—2.5)
cm. longis et ad 4 mm, latis, laminis supra nitidis glabris basi costae excepta,
subtus pubescentibus, glaucis, coriaceis, in sicco olivaceo-brunneis, obovato-
ellipticis, ad 18 cm. longis et 8 cm. latis, basi attenuato-cuneatis, in petiolum
decurrentibus ibique plus minusve valde recurvatis, apice rotundatis leviter
obtuse et abrupte acuminatis, penninerviis, costa supra leviter impressa
subtus satis elevata robusta, nervis 6-9 (—10)-paribus supra haud subtus
nonnihil elevatis plus minusve castaneis angulo +40° divergentibus, rete
venularum supra utrinque conspicuo. Inflorescentia axillaris, juvenili
bracteis adhuc onusta, paniculata, ad 18 cm. longa, dense fulvo-tomentosa,
longipedunculata, pedunculis robustis striatis angulatis ad 8 cm. longis.
Flores dense fulvo-tomentosi, +3.5 mm. longi, pedicellis ad 4 mm. longis
tomentosis, perianthio campanulato, canescente fide coll., fulvo-tomentoso,
lobis ovatis obtusis crassis, +2.5 mm. longis; staminibus ser. I & If +1
mm. longis, antheris ovatis, connectivo antherae 13 longitudine aequante
filamento duplo longioribus, ser. III +1.25 mm. longis biglandulosis,
antheris oblongis truncatis glandulis filamentisque duplo longioribus; stami-
nodiis lato-lanceolatis 0.75 mm. longis; gynaecio glabro + 1.7 mm. longo,
ovario ovoideo-globoso stylo duplo longiore, stigmate peltato conspicuo.
Fructus in sicco flavescenti-brunneo-maculatus, obovoideo-ellipticus, con-
spicue et obtuse apiculatus, 2.2 * 2cm., cupula campanulata rugosa verru-
cosa glabra margine leviter undulata, ad 1 cm. longa, 1.7 cm. diam., et
5 mm. alta subtentus, pedicello incrassato glabro, ad 15 mm. longa apice ad
5 mm. lata.
DistriBUTION: Known only from the type-locality, in Panama.
400 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
PANA Chiriqut: eee mua Boquete, in rain-forest, alt. 1340 m., April 9,
1938, Pawson pe (a, Ch, — Mo: f r., A, Ch) (tree 15-21 m.; flowers white).
Nectandra hypoglauca is ‘similar to NV. Whitei and N. producta, which
see for discussion.
33. Nectandra Paulii, sp. nov.
Arbor ad 30 m. alta, ramulis robustis sulcatis minute fulvo-tomentosis
mox griseo-pubescentibus vel glabrescentibus. Folia alternata juventute
sparse adpresse minute fulvo-pubescentia mox glabrescentia demum glabra,
petiolis robustis pubescentibus vadose canaliculatis ad 2 cm. longis et 4 m
latis, laminis supra utrinque glabris basi costae srg subtus minute
pubescentibus, juventute membranaceis mox coriaceis, in sicco supra viri-
descenti-brunneis, subtus seg gamete subglaucis, ellipticis vel obovato-
ellipticis, ad 15 cm. longis et 7 cm. latis ;, basi cuneatis, in petiolum decur-
rentibus ibique satis recurvatis, aia acutis, acuminatis, vel rotundatis,
raro emarginatis, margine leviter te penninerviis, costa robus sta
supra conspicua subtus valde elevata, nervis (6—) 7—8 (—9)-paribus supra
conspicuis sed leviter impressis, subtus elevatis angulo 35—45° divergenti-
us, rete venularum utrinque obscuro. Inflorescentia axillaris, late panicu-
lata, ad 30 cm. longa, SNR multiflora, longipedunculata, pedunculo
a pubescente, ad 10 m. longo. Flores ad 3 mm. longi, pedicellis
1—3 mm. longis gracilibus, Seem ts hypocrateriformi albo, fide coll., lobis
shiodats reflexis, crassis, papilloso- tomentosis, +2.5-3 mm longis: sta-
minibus ser, I & II + +0.6 mm. longis, antheris subreniformibus vel depresso-
globosis filamento duplo longioribus, ser. II] +0. longis conspicue
biglandulosis, glandulis et antheris quadratis ner me aequalibus; stami-
nodiis oblanceolatis pubescentibus +0.6 mm. longis; gynaecio glabro,
+1.25 mm. longo, ovario subgloboso, stylo subnullo, stigmate ergy
rotundato conspicuo. Fructus ellipsoideus, apiculatus, 28 X mm.,
cupula rubra campanulata crassa verrucosa glabra ad 8 mm. longa, ef mm.
diam., et 5 mm. alta subtentus, pedicello incrassato pubescente striato, ad
I cm. longo et apice 8 mm. lato.
DistripuT1IonN: In Costa Rica, up to 915 m. altitude, and in Chiriqui Province of
Panama, at 1500-2000 m. altitude.
Costa Rica: San José: Forests in the vicinity of El haope alt. 915 m., Feb.
1936, Shek 2005 (fl., type — A, GH, NY) (tree 30 m.; flowers white). PANAMA:
Chiriqui: Bajo Mono, mouth of Agee Chiquero, along Rio Caldera, Woodson,
Allen & Seibert 1022 (fr., A, Ch, , NY); vicinity of Cerro Punta, Paul H. Allen
1572 (fr., Ch, GH, Mo)
The lack of well developed connective tissue of the outer series of anthers,
and the sessile stigma and globose ovary distinguish the flowers of NV. Paulii
from those of V. hkypoglauca. The fruiting pedicel is shorter than that of
N. hypoglauca, and the fruit is ellipsoid rather than obovoid-ellipsoid.
34. Somes belizensis (Lundell), comb.
Phoebe belizensis Lundell in Contr. Univ. "Mich. Herb. 6: 20. 1941.
DISTRIBUTION: Known only from the type-locality and vicinity, in British Hon-
ras.
Bri rng Honpuras: Stann Creek: Creek-side, Mountain Cow Ridge, Gentle
3281 (fr. A, NY); Mountain Cow Ridge, in high ridge, March 31, 1940, Gentle 3304
(f1., isotrypr, A, NY) (tree 25 cm. diam.; flowers white) ; Big Eddy Ridge, Gentle 3346
(fr., A, NY).
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 401
NATIVE NAMES: ‘“Timbersweet,” “White laurel” (British Honduras).
This tree, described under the genus Phoebe, certainly resembles the
latter genus in habit-characters, particularly the species P. helicterifolia.
The softly ferruginous- tomentose young branchlets and petioles are char-
acteristic, as well as the oblong chartaceous to subcoriaceous pale leaf-
blades with the rounded bases and abruptly and sharply acuminate apices.
The blades are 11-22 cm. long and up to 8.5 cm. broad. The costa is
impressed above and retains its early pubescence, whereas the lateral nerves,
of which there are 6—8 pairs, are merely impressed, diverging at an angle of
45-55°. The reticulation, which is also impressed above and not too con-
spicuous, stands out prominently beneath. The few-flowered axillary inflo-
rescences are up to 10 cm. long and borne on slender sae ee
peduncles up to 5 cm. long. The white flowers are about 8-9 mm. in diam-
eter, the thick ovate lobes up to 3 mm. long and pubescent corners The
stamens of the two outer series are +0.8 mm. long, the broadly obovate
anthers are nearly sessile, and the connective is slightly prominent. Those
of the inner series are +1.25 mm. long, subrectangular, the large conspic-
uous glands equalling the filaments and anthers in length. The staminodia
are thin, triangular, less than 0.4 mm. long. The glabrous gynaecium
measures ++ 1.25 mm. long; the subglobose ovary is topped by a very short
style bearing an peu wate discoid stigma. The fruit is glabrous,
ellipsoid, apiculate, up to 1 (—1.3) cm. long and 6 (—?) mm. broad, the thin
shallow pubescent to ne cupule not more than 4 mm. long and about
5 mm. in diameter, the slender pubescent to glabrescent pedicel from 5 mm
to 1 (—1.72) cm. long.
The lack of conspicuous staminodia, the short-styled globose ovary, the
form and development of connective-tissue in the anthers, and the shape
of the anthers themselves all seem to point to the fact that this species
should be included under Nectandra. The fruits of the numbers cited
above appear to be immature.
35. Nectandra rudis,
Arbor (?) aes ee atro-brunneis verruculosis minute et obscure
ee Folia alternata, petiolis robustis glabrescentibus haud can-
aliculatis, ad 3 cm. longis et 4 mm. latis, laminis utrinque glabris percori-
aceis in sicco brunneis, supra nitidis, ellipticis, ad 20 cm. longis et 8 cm.
latis, basi cordatis recurvatis, apice ignotis, penninerviis, costa supra rubes-
cente et leviter impressa subtus conspicue elevata, nervis ad 10-paribus
supra rubescentibus leviter elevatis subtus elevatioribus angulo 35—45°
divergentibus, rete venularum supra conspicuo subtus obscuro. — Inflores-
centia axillaris et subterminalis late paniculata, ad 13 cm. longa, minute
adpresse pubescens, multiflora, pedunculo robusto ad 5 cm. longo. Flores
ad 6 mm. longi, dense ferrugineo- pubescentes, pedicellis pubescentibus satis
robustis ad 5 mm. longis, perianthio late campanulato, lobis late ellipticis
obtusis (mox reflexis) crassis papilloso- tomentosis ad 4 mm. longis; stamin-
ibus ser. I & IJ +1.25 mm. longis antheris subglobosis filamento basi
pubescente duplo longioribus, ser. IIT +1.25 mm. longis biglandulosis,
antheris oblongis, glandulis aequalibus, longitudine a pusaoe aequanti-
bus; staminodiis nullis; gynaecio glabro +1.7 mm. longo, ovario globoso
brevistipitato ee approx. duplo longiore, bana parvo discoideo.
Fructus ignotu
ee de iano only from type-locality.
402 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
Mexico: Chiapas: Mt. Ovando, Dec. 29, 1936, Matuda 470 (fl., TYPE, A).
This robust species, known only from the flowering specimen, seems to be
distinct enough to warrant description without fruiting material. It is
perhaps most nearly related to NV. sinuata, in spite of the glabrous leaves
and ovary and the roundish stamens with very little connective tissue pro-
truding at the apex of the anthers.
36. Nectandra platyphylla (Lundell), comb. n
Phoebe platyphylla Lundell in Contr. Univ. Mich. Herb. 6: 23. 1941.
DistrIiBUTION: Known only from the type-locality, in Mexico.
xico: Chiapas: Finca Suiza near Montecristo, Jan. 1938, Matuda 1930 (fl.,
isoTyPE, A, NY).
The branchlets of this tree are angled, at first dark and minutely puber-
ulous, later becoming glabrous, terete, striate, and often grayish. The
pedicels are rather slender, canaliculate, puberulous, and up to 1.5 cm. long.
The membranaceous leaf-blades are yellowish green, at first minutely seri-
ceous, becoming quickly glabrescent on both surfaces except for the frequent
presence of pubescent axillary glands beneath. They are oblong-elliptic
or obovate-elliptic, the base rounded and often abruptly cuneate with a
tendency toward recurving, the apex obtuse or obtusely acuminate or occa-
sionally acutish, and measure 9-25 cm. long and 4.5—11.5 cm. broad. The
costa is slender, impressed above and somewhat elevated beneath. The
lateral nerves, of which there are 6-12 slender pairs, are only slightly ele-
vated above and little more beneath, visible but not conspicuous, and
diverge at an angle of about 45°. (The isotype has leaves on the whole
much smaller than the length and breadth given by Lundell.) The inflo-
rescence is axillary, glabrous or at least glabrescent, up to 10 (—15) cm.
long, few-flowered, the slender dark brownish red peduncle up to 6 cm.
long. The white flowers are aree, about 1 cm. in diameter, supported by
filamentous pedicels up to 1 cm. long. The lobes are 4 (—5) mm. long,
broadly elliptic, obtuse to uae. fleshy, papillose, the tube short. The
stamens of the two outer series are + 2.15 mm. long, with sessile anthers
elliptic, obtuse, and petaloid. Those of the inner series have anthers that
are subrectangular, rounded, and borne on filaments about one-third their
length and completely covered by two conspicuously large glands that are
as large as the anthers. The peers are subtriquetrous, stipitate, =: 1
mm. long. The glabrous gynaecium is +2.15 mm. long, compressed -
globose, with a very short thick ae topped by a triangular conspicuous
stigma.
Lundell places this near Phoebe ambigens, from which it may be
distinguished by leaf-blades that are more frequently obovate-elliptic than
elliptic, membranaceous rather than subcoriaceous, and by smaller flowers,
the styles of which are extremely short.
37. — — Mez in Jahrb. Bot. Gart. Berlin 5: 402. 1889; Standley in
. . Herb. 23: 297. 1922; Standley & apie: Lista Prelim. PI.
Salvador 84. on Standley in Field hi Publ. Bot. 18: 453.
Persea Matudai Lundell in Lloydia 4: 49. 1941.
DistriBUTION: Southern Mexico through Central America.
Mexico: Oaxaca: Vicinity of Cafetal Concordia, Morton & Makrinius 2486
(fr., Ch); Cafetal San Antonio, Pochuntla, Reko 6039 (fl, GH). Chiapas: Las
1945] ALLEN, STUDIES IN THE LAURACEAE, VI 403
Cadenas, Escuintla, January 5, 1938, eee 1880 (fl., isotype of Persea Matudai
A; Ch; ie GUATEMALA: uezaltenango: Coffee plantations, Colomba, Shuich
1978 (fl. NY); Finca Pirineos, ane south-facing slopes of Volcan Santa Maria,
between see Maria de Jesus and Calahuaché, Steyermark 33234 (fl. Ch). Zacapa:
Trail between Santa Rosalia de Marmol and Vegas, Steyermark 42961 (fl., Ch).
Chiquimula: Around crater-lake, Volcan de Ipala, Pittier a (fl, GH).
Guatemala: Near Finca La Aurora, Aguilar 279 (fr., Ch). Sacatepéquez:
Near Barranco Hondo, a het of Alotenango, dry thicket, Standley fas (fl., A,
Ch). Retalhuleu: Near Retalhuleu, Bernoulli & Cario 2581 (fl., TYPE of Nectandra
sinuata not seen). Suchitepéquez: Finca Moca, Skutch 1489 (fl., Ch); Bequaert
55 (fr., Ch, GH). Santa Rosa: Chiapas, Heyde & Lux 4374 (fl, GH, NY).
Honpuras: Dept. unknown, Cerro del Guayabal, S. Calderén 2012 (fl, GH).
Ahuachapan: Vicinity a eae along stream, Standley 19959 (fl., GH, NY);
Sierra de Apaneca, in the on of Finca Colima, Standley 20095 (fl., GH, NY).
L SALVADOR: San Salv ae Volcan de San Salvador, S. Calderdén 473 (fl., GH),
Standley 22968 (fl., GH, NY); cerro de San Jacinto, S. Calderén 2248, Standley 20629
(fl., GH ae La Paz: Zacatecoluca, S. Calderén 308 (fr.. GH, NY). Costa
Rica: ae uela: San Pedro de San Ramon, Brenes 6817, 15090 (fl., Ch); San
Francisco rf Guadaloupe, Pittier 12348 (fr., Ch, GH); Naranjo, Cerro del Espiritu
Santo, in thicket in reddish clay-loam of Pacific tropic zone, A. Smith P.2409 (fl., A);
Alajuela, J. D. Smith 6754 (fl, GH, NY). San José: About the Hacienda Belmira,
near Santa Maria de Dota, Tonduz 11651 (fl, Ch, GH). Cartago: Near Cartago,
Skutch 4687 (fl., A, NY)
NaTIvVE NAMES: “Aguacamico,” ‘“Aguacate amarillo,” ‘“Aguacate de mico” (Fl
Salvador); “Aguacatillo” (Mexico); ‘“Chipinahuaca,” “Palo de Chipinahuaca” (El
Salvador) ; “Palo de Tejon” (Mexico) ; “Quisarra,” “Quisarra hedionda” (Costa Rica) ;
“Tepeaguacate rojo” (Guatemala) ; “Trompillo,”’ “Trompito” (El Salvador).
This is a very well known species occurring throughout most of Central
America and adjacent Mexico. The stout angled branchlets are clothed
with a fulvous or grayish tomentum. The densely tomentose pedicels are
somewhat canaliculate, robust, and up to 3 cm. long. The oblong-elliptic
or obovate blades are cordate at the base, the apex subobtuse, rounded or
abruptly subacuminate or abruptly and sharply acuminate, 25 (—30) cm.
long and 12 (—19) cm. broad, chartaceous, densely grayish- or fulvous-
tomentose beneath and more sparsely pubescent above, the costa and nerves
heavily tomentose. The costa and slender lateral nerves, of which there
are 9-11 (—15) pairs diverging at an angle of 45—70 (—80)° (the lowermost
almost at right angles), are conspicuous above because of their pubescence
and are prominently elevated as well as pubescent beneath. The inflo-
rescence is a stout subcorymbose or pyramidal many- or few-flowered axil-
lary panicle, usually densely pubescent, up to 25 cm. long, the robust
peduncle up to 16 cm. long. e large flowers are up to 1 cm. long and
within, broadly ovate or elliptic, subacute to obtuse or round, reflexed at
anthesis and in the dried state castaneous or occasionally dark brown. The
two outer series of stamens are 3 mm. long, the elliptic or ovate anthers
sessile or subsessile, usually variously petaloid, heavily papillose, the fleshy
truncate, the cells sublaterally extrorse and the filaments with conspicuous
sessile basal glands about one-third the length of the stamens. The usually
ensely pubescent (sometimes glabrous) gynaecium is +3 mm. long, the
404 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXVI
subglobose ovary equalling in length the stout style with its ee
subtriangular eae stigma. The ellipsoid fruit measures up to
long and 17 mm. in width, the supporting campanulate glabrescent to ie
brous cupule ee up to 13 mm. long, 2.5 cm. in diameter, and about
10 mm, deep, the pedicel enlarged to 13 mm. long and about 11 mm. diam-
eter at the apex. In the dried state the cupule flares away from the fruit
instead of surrounding it closely. The upper half of the fruit is glabrous
except for the entire tip, which is covered with persistent pubescence.
Approximately the lower third to half, which is almost entirely hidden by
the cupule, is densely fulvous-sericeous-tomentose.
The above description is applicable to the majority of specimens of the
species. All of the material from San Salvador shows flowers with glabrous
ovaries. Some of the Guatemalan specimens are less tomentose as to
foliage and inflorescence in general. Nonetheless, it is apparent that they
all belong under the same widely variable species
38. Nectandra reticulata (Ruiz & Pavon) Mez in Jahrb . Gart. Berlin 5:
1889; Standley in Contr. U.S. Nat. Herb. 23: 297. tg in on Mus. Publ. ane
Laurus reticulata Ruiz & Pavon, Fl. Peruv. Chil. 4: t, 348, & Laurogr. t. 23. 1802.
DisrripuTion: Tropical America from Mexico into Central and South America.
Mexico: Puebla: Zonquimistlan, C. & E. Seler 3643 (fl, GH). Oaxaca:
Chiltepec and vicinity, Tuxtepec, in anos, Martinesz- Calder 488 (fl, A). Chiapas:
Escuintla, Matuda 383 (fl, A, "). GUATEMAL Alta Verapaz: South
Cubilguitz, in thickets, Steyermark 44561 (fl., Ch); Hacienda Vancabnat, cielo
45093, 45094 (fl., Ch); Cubilgilitz, von Tuerckheim 7965 (f1., GH, NY). Quezalte-
ngo: Pinata: in Saliee plantations, Skutch 1988 (f1., A, acl a annie
Retalhuleu, Aellerman 6587 (fl., Ch); vicinity of Las Delicias, south of Retalhuleu,
Standley 88121 (f1., Ch); vicinity of Retalhuleu, Standley 88822 (fl., Ch). Honpuras:
Mosquitia(?): Black River Valley, Record & Kuylen H. 69 (Y¥ 10015) (fl, Y).
Nicaracua: Bluefields: Region Braggman’s Bluff, Englesing 65,123 (fl., Ch).
Costa Rica: Alajuela: San Pedro de San Ramon, Brenes 6586 (f1., Ch), Tonduz
22a (fl., Ch); Phionwanls Valley, G. P. Cooper 32 (Y ieee (fr., Ch); region of
Almirante, Cricamola along river, oe & Slater 512 (fr.. Ch). Canal Zone:
Forest along the Rio Indio de Gatun, Pittier 2775 (fl., GH, NY); in swampy woods,
Lion Hill Station, Hayes 467 (fl, US); Mindi, Cowell 182 (fl, NY). [Peru: With-
out locality, Pavon 504 (fl., isotype of Laurus reticulata, Ch).|
Native NAMES: “Aguacatilla” (Honduras); ‘“Chuala” (Guatemala) ; “Sweetwood”
(Panama).
his widespread species is conspicuous for its densely ferruginous-tomen-
tose branchlets and inflorescences. The stout petioles, up to 1.5 cm. long,
as well as the lower surface of the leaf-blades, are densely tomentose. The
blades are coriaceous, glabrous above except for the venation, lanceolate-
elliptic or oblong-elliptic, the base auriculate and strongly recurved, the
apex attenuate-acuminate, sometimes up to 30 cm. long, and as broad as
9 cm. The costa and lateral nerves are impressed and pubescent above,
prominently elevated and densely pubescent beneath. ‘The lateral nerves
number up to 12 pairs and diverge at an angle of 35—45°. The reticula-
tion is pronouncedly impressed above and conspicuously elevated beneath.
The inflorescence is stout, axillary, many-flowered, ferruginous-tomentose,
paniculate, usually with long peduncles up to 10 cm. long. The densely
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 405
tomentose flowers are large, up to 7 cm. or more long and 10-15 mm. in
diameter, the pedicel 4—5 mm. long, often less. The inner surface of the
tube is frequently densely hairy. The fleshy tomentose lobes are ovate,
obtuse or rounded, and papillose within, measuring nearly 6 mm. long, the
inner being slightly shorter than the outer. The stamens of the two outer
series are variable in size, up to +2.4 mm. long, and the almost sessile
anthers vary in size from depressed- globose to ovate and almost petaloid,
the connective tissue occupying usually about half the length of the anther.
The stamens of the inner series are also variable in length, up to +2.5 mm.,
the anthers often squarish and emarginate or occasionally ovate; the fila-
ments are sometimes almost equal in length to the anthers and always bear
conspicuous spreading sessile glands at the base, nearly equal to the anthers
in length. The linear-lanceolate staminodia are usually hairy, and are
+0.8 mm. long. The gynaecium is 3 mm. or under in length, and is for
the most part densely pubescent, although in some cases it may be glabres-
cent or even glabrous. The ovate or ellipsoid ovary is slightly longer than
the stout style, which is topped by a conspicuous subcapitate stigma. he
fruit, of which I have no material at hand, is presumably ellipsoid, up to
13 mm. long and 8 mm. broad, subtended by a simple- ea ee that
is slender and subpateriform, and one-third the length of the
Although the specimens from Mexico and Central penta oS con-
siderably from each other and from the type from Peru, there seems no
reason for setting them up as a new species. Kostermans (Meded. Bot.
Mus. Utrecht 25:19, 1936) mentions the close relationship between
N. Laurel, N. rigida, and N. reticulata, giving the densely sericeous-hirsute
inner surface of the tube of the latter as a diagnostic character. Many of
the specimens cited above are intermediate in this as well as other characters.
All gradations of leaf-base, shade of pubescence, and internal floral structure
are apparent in the material at hand. Eventually further study of South
American collections may reveal different trends and make another disposi-
tion of the North and Central American specimens advisable.
DouBTFUL SPECIES AND VARIETIES OF NECTANDRA
Nectandra mollis Y venosa Meissner in DC. Eee 151: 149. 1864.
This variety, described from Mexico, Central America, and South
America, Mez reduces to Nectandra reticulata (Ruiz & Pavon) Mez.
None of the syntypes are at hand for comparison; they include Oersted’s
collection from Costa Rica and Schiede & Deppe 243 from Mexico.
Nectandra mollis 6 villosa Meissner in DC. Prodr. 151; 148. 1864.
This also is reduced by Mez to Nectandra reticulata (Ruiz & Pavon)
Mez. The syntypes, from Mexico (Schiede & Deppe 241, 1145) and
South America, are unavailable at present.
Neciandra polita 8? Oerstedii Meissner in DC. Prodr. 151: 164. 1864.
e type, Oersted 12, from Nicaragua, has not been seen; Mez includes
the variety under Nectandra latifolia.
Nectandra turbacensis Y mexicana Meissner in DC. Prodr. 151: 152. 1864.
Mez reduces this to Nectandra rectinervia Meissner. The syntypes,
Linden 16 and Galeotti 7101, from Mexico, are not available.
406 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
Nectandra amazonum var. 5 Oerstedii Meissner in DC. Prodr. 151: 150. 1864.
The syntypes presumably are the numbers cited by Mez, under Nectandra
globosa, Oersted 16, 17, 18, from Costa Rica. So far, they have not been
available for study.
SpEecrES EXCLUDED FROM NECTANDRA
Nectandra chiapensis Lundell = Ocorea cu1apensis (Lundell) Standley & Steyermark.
Nectandra striata Nees = Myropta cf. ruNEBRIS L. (ex Giirke), fide Mez.
5. Litsea Lamarck
Litsea papel Dict. 3: 574. 1791; Hemsley, Biol. Centr. Am. Bot. 3: 76. 1882; Mez
hrb. Bot. Gart. Berlin 5: 474. 1889; Bartlett in Proc. Amer. Acad. 44: 597.
Tetranthera oraiae Hort. Schoenbr. 1:59, ¢. 113. 1797; Meissner in DC. Prodr.
7.
Malapoenna em Rev. Gen. 2: 571. 1891.
DistrIBUTION: In mountain-forests of Mexico and Central America.
Small trees or shrubs not more than 6 m. high, with leaves lanceolate to
elliptic with all intermediate forms, not more than 13 and usually less than
10 cm. long, or leaves ovate to orbicular and not more than 7 cm. long.
The diagnostic character, however, lies in the dioecious flowers borne in
small axillary subumbellate clusters on peduncles of varying lengths, usually
not more than 2 cm., surrounded by an involucre that is quickly deciduous
at anthesis. The floral parts are extremely variable in this unstable group,
but in the main there are six perianth-lobes, occasionally aborted to fewer,
equal or subequal in length. The ¢ flower usually bears three or four
series of fertile stamens, the two inner of which are biglandular. The
anthers are four-celled and are all introrse, though very occasionally in the
inner cycles the lower locules appear to be lateral. A small aborted ovary
may occur or be absent entirely. The ? flower has three or four series of
three each of staminodia, the first two of which are usually without glands.
The third and fourth, if present, bear at the base of the filament two glands
which are often stipitate. The ovary is well developed in the @ flower.
The fruit is a more or less globular berry, seated on a small scarcely changed
perianth-tube, or on one that is thickened and enlarged to form a cupule
or disc, which is seated on the enlarged pedicel.
The genus Litsea in America presents a pattern different from that of
other genera of this hemisphere. There are two morphologically distinct
divisions: first the group consisting of three species with leaves ovate or
ovate-lanceolate to orbicular-ovate, showing bases rounded or subcordate,
found only in Coahuila, Nuevo Leon, and San Luis Potosi; second, embrac-
ing the remainder of the genus, the group with leaves elliptic, lanceolate,
linear, or oblong and all variations thereof, their bases not subcordate nor
rounded, occurring from Chihuahua and Sinaloa south and east throughout
Mexico and Central America.
KEY TO THE SPECIES OF LITSEA
A. Leaf-bases rounded or subcordate.
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 407
B. Young branchlets saad ferruginous-pubescent ; lower surface of leaves tomen-
tose; petioles pubescent; inflorescence densely ferruginous-pubescent; infruct:
cence not more than 1.5 cm. long; fruit =5 mm. long............. 1. L. Mue ee
B. Young branchlets sebreis or at most glabrescent; lower surface of leaves and
petioles glabrous; hates Lannen or pitreseat infructescence not less
than 1.5 cm. long; fruit 1 cm. ong.
C. Leaf-blades ovate- eee the ae up to 7 cm. long, the ie acute..
Stee) et aicaer ead oureHor say ey ares ete nar tes Se ib tr Veal wisn Sopweme er eu enema euete erate me Pringlei.
C. Leaf-blades ovate or aa -ovate, the largest up to 4 cm. ane the apex
acute, obtuse, or rounded............... 0. cece cece eeceaee 3. L. parvifolia.
A. Leaf-bases not rounded or sbcordat Gs ls. ost x na ean Renee ee ees 4. L. glaucescens.
B. rae pees, petioles, and venation not conspicuously flaves
-blades usually Cre’ elliptic or ovate-elliptic, ike us to dense
eee Peychs ants eee aro eee Se wha css eis eles 4a. L. glaucescens var. haere
C. Leaf-blades linear or at most linear-lanceolate to oblong, always glabrous..
SO OI nr ee eae 4b. L. glaucescens var. Schafneri.
B. Branchlets, petioles, and venation conspicuously flavescent.............00..008
Waite steiarei arouse ee ePanuantora aye heels 6268s eo gieea'e 4c. L. glaucescens var. flavescens.
al ras ee Rehder in Jour. Arnold Arb. 16: 449.
Litsea Tharpiana Standley in Field Mus. Publ. Bot. 7 ie 1937.
ai minihes Mountains of Nuevo Leon, Mexico, at an altitude of 1500-2600 m.
Leon: Sierra Madre Oriental, common in dense woods east
side of dive between San Francisco Canyon and Pablillo, 15 miles s.w. of Galeana,
alt. 2600 m., May 14, 1934, C. H. & M. T. Mueller 379 (4 fl., rype of L. Muelleri — A,
Mich, NY, Te. US); Hacienda Pablillo, Galeana, August 18, 1936, Taylor 188 (fr.,
type of L. Tharpiana, 8 fl., Tex); cafion above Alamar, C. H. & H. T. Mueller 689
(fr., A, Mich, NY, Tex, US).
This species is easily separated from the other species of Litsea with
rounded or subcordate leaf-bases by the occurrence of a dense persistent
pubescence on the young branchlets and petioles and a tomentum on the
lower surface of the leaf-blades. The nearest relatives are to be found in
the L. parvifolia complex, from Nuevo Leon and Coahuila, but separated
by the ferruginous (sometimes becoming fuscous) pubescence on the young
branchlets, and by the venation of the lower leaf-surface. On the whole,
the lateral nerves are more ascending in L. Muelleri, the petioles longer and
densely hairy, and the leaf-bases less pronouncedly cordate. The two
entities described undoubtedly represent the same species.
Zz ay ee Bartlett in ts Amer. Acad. 44: 598. 1909; Standley in Contr.
a Herb. 23: 287.
re cae asts Bartlett in op. ann 601; Standley in op. cit. 288.
DistrIBuTION: Ridges in the Sierra Madre near Monterrey, Nuevo Leon, and. in
southwestern San Luis Potosi
Mexico: Without locality, April, 1926, Runyon & Tharp 893, 1008 (sterile, fr.,
Te: Leon: Municipio de Villa Norse an Cafion Marisio Arriba, Rancho
Las Adjuntas, C. H. Mueller 2042 (fr., GH); Cafion Diente, near Monterrey, C. H.
Mueller 2662 (fr., GH); ae Madre Oriental, waterway below Alamar, about 15
Ze
=
<
Pringle 2078 (fr., GH), 2837 fs type of L. novoleontis, GH) ; limestone ledges, Sierra
Madre above Monterrey, alt. 800 m., March 8, 1906, Pringle 10238 (4, @ fl., TYPE of
L. Pringlei— GH, NY, US) (shrub 1.3-2 m.), Tharp 1835 (Q fl., Tex, US). San
Luis Potosi: Mivares , Sept. 5-10, 1902, E. Palmer 62 (6 fi, fr.. GH, NY, US);
mountains, San José Pass. August 5, 1890, Phage 3146 (fr., NY, US).
NATIVE NAMES: “La Brel” (Nuevo Leon); “Laurel” (San Luis Potosi).
408 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXVI
This shrub, 1.3-2 m. high, bears furcate branchlets which are densely or
sparsely leafy and are reddish black or olivaceous in color. The coriaceous
reticulate ovate-lanceolate leaf-blades are extremely variable in size, be-
coming as long as 7 cm., their tips often mucronulate, and their bases
rounded to subcordate. The ¢ inflorescences are variable, which fact
accounts for the description of the two species L. Pringlei and L. novoleontis
from the same locality. In the former the 1—4 erectly pedunculate subum-
bellate ¢ inflorescences occur principally in the axils of terminal leaves,
giving an appearance of dense flower-clusters. The ¢ inflorescences of the
specimens described as L. novoleontis are single, axillary, and borne on
slender nodding peduncles. The fruit is black, globose, about 11 mm. in
diam., subtended by an expanded pedicel about 5 mm. long, and +4 mm.
in diam. The peduncle is only slightly enlarged.
3. Litsea parvifolia (Hemsley) Mez in Jahrb. Bot. Gart. Berlin 5: 481. 1889; are
Amer. Acad. 44: 601. 1909; Standley in Contr. U. S. Nat. Herb. 23
1922
Umbellularia parvifolia casera” Biol. Centr. Am. Bot. 3: 77. 1882.
1891.
Malapoenna parvifolia Kuntze, Rev. Gen. 2: 571
Litsea Byesnge-s Bartlett in Pro. reas erg 44: 598. 1909; Standley in Contr.
U. erb. 23: 287.
jee Mountains near on Coahuila, and Monterrey, Nuevo Leon.
Mexico: Coahuila: Abundant in mountains near Saltillo, July 30, 1848, Gregg
314 (9 fi, fr., isotype of Umbellularia parvifolia, GH) (shrub 1.5-3 m. tall); April
15-30, 1898, E. Palmer 68 (2 fl., GH), alt. 2100 m., April 12, 1906, Pringle eee
(3S fl., type = L Sipser Reig NY, US) tahirub 1.3-1.5 at Nuevo Leo
Monterrey, C. H. & M. T. Mueller 301 (Q fl, fr., Tex).
This is ren earliest species of Litsea to be described from northern
Mexico. The shrub, 1.3-3 m. in height, has ovate-lanceolate to orbicular
leaf-blades, minutely mucronulate at the apex and rounded, subcordate or
cordate at the base, more or less coriaceous, and heavily reticulate, not more
than 4 (usually 3) cm. long. Characteristic is a bloom discernible on leaf-
blade, stem, buds, and fruiting calyx, due to the presence of a thin film of
minute, rapidly disappearing pubescence. ‘The situation existing in Litsea
parvifolia and L. pedicellata parallels that in L. Pringle and L. novoleontis.
The ¢ inflorescence of L. parvifolia is a single axillary subterminal umbel
borne on slender more or less recurving peduncles. The flowers, 3 or 4 per
umbel, are subtended by densely tomentose pedicels 2 mm. long. The ¢
inflorescences of L. pedicellata are more densely aggregated near the tips of
the flowering branchlets, occurring in a few subumbellate axillary clusters
and a heavily flowered terminal panicle up to 3 cm. long. ere is a
slender grace perceived in the branchlet of L. pedicellata that is wholly
lacking in the more coarse and rather stiff aspect of L parvifolia. The leaf-
blades of the type of the former are less heavily coriaceous and of a bluish
green, with reticulations more prominent. In spite of these differences,
there seems to be no sound reason for keeping up both species.
4. Litsea glaucescens H.B.K. Nov. Gen. & Sp. 2: 133 [168]. 1817.
Small aromatic tree, the branchlets smooth, dark or reddish brown or
olive, terete. Petioles slender, canaliculate, 5-20 mm. long, --
1945 | ALLEN, STUDIES IN THE LAURACEAE, VI 409
diameter; leaf-blades variable, usually thinly coriaceous, sometimes mem-
branaceous, olive-green to dark brown above when dried, glaucous to
glaucescent or at least paler beneath, usually elliptic- lanceolate or lanceo-
late, ae oblong or ovate- lanceolate, (4—) 7-9 (—12.5) cm. long and
(1—) 2-3 (—3.5) cm. broad, attenuate to cuneate or obtuse at base, acute
to acuminate or sometimes obtusish at apex, usually with a mucro of vary-
ing length, glabrous throughout or sometimes in young stages slightly
pubescent above and more densely pubescent beneath, the venation yel-
lowish or red-brown and except for the costa obscure above, the lateral veins
usually ee slightly prominent beneath, occasionally with the mar-
ginal vein prominent, the entire surface frequently conspicuously areolate.
3 ae axillary, solitary or racemose umbels less than 2.5 cm.
long, the peduncles 1 cm. or less in length, usually glabrous, the pedicels
up to .5 cm. long and pubescent. Flowers 3-5 per umbel, often slightly
pubescent, up to 4 mm. long; perianth-lobes 6, membranaceous, oblong or
elliptic, usually 3 mm. long, the tube very short, the stamens 9. @ inflo-
rescences axillary, solitary umbels up to 2 mm. long, the peduncles ne to
1.5 cm. long; flowers 3-5 per umbel, 2—3 mm. long, often slightly pubescent,
the perianth-lobes 6, membranaceous, usually elliptic, the tube very short,
the staminodia usually 9. Fruit a globular berry up to 12 mm. in diameter,
subtended by the enlarged usually glabrous tube and pedicel up to 7 mm.
long, flaring at the tip to 2 mm. in breadth usually, sometimes the tube
expanding to form a shallow cupule or disc up to 4 mm. in diameter.
DistripuTion: Northwestern Mexico, south and east to Guatemala, Honduras,
and Costa Rica. The species occurs in every state of Mexico except Sonora and
brond sense occurs generally at an altitude of between 2000 and 3000 m. in forested
mountainous regions, although to the north, where it abounds in the plains and along
the coast, it descends to a much lower altitude. It is locally used for seasoning or
flavoring for meats, soups, and other foods, and as a substitute for tea; it is used also
as a medicine for colic pains, etc.
This widespread species, as Tetranthera glaucescens, was split into four
varieties by Meissner (1864), the separation being made principally on the
type of inflorescence. Hemsley (1882) listed also a second species Litsea
Neesiana with two varieties. Mez (1889) submerged L. glaucescens var.
subsolitaria (Meissner) Hemsley into the species proper, and elevated var.
major Meissner to specific rank as L. guatemalensis. Bartlett (1909) set
apart the Costa Rican collections as L. flavescens, and those from San Luis
Potosi and Tamaulipas (L. glaucescens var. subsolitaria) as L. Schaffneri,
in which he included Parry & Palmer 798, cited by Hemsley under L.
glaucescens var. subcorymbosa.
It seems impossible at present to formulate any clear-cut species in this
highly variable group. Careful study of the available material shows a
single complex species which varies with each type of locality in which it
has been collected. The typical variety, var. subsolitaria (the oldest
varietal name extant applied to the type-material), was originally collected
in Guerrero and Vera Cruz. Subsequent scattered collections have been
410 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
recorded from nearly every state in Mexico, each showing slight variation
from the variety. On the periphery of the range of the typical variety are
two divergent groups of specimens which seem worthy of varietal rank.
They may be easily recognized in most cases, but frequently specimens are
found which show a gradation toward the typical variety. A treatment of
the three varieties follows.
4a. ie glaucescens var. subsolitaria (Meissner) Hemsley, Biol. Centr. Am. Bot.
1882; Mez in a Bot. Gart. Berlin 5: 477. 1889; Bartlett in Proc. Amer.
rere 509. 190
ris prises s H. as K. . Gen. & Sp. 2: 133 [168]. 1817; Mez in Jahrb. Bot.
Gart. Berlin 5: 477. 385 nee in Pree Amer. Acad. Ad: 599, 1909.
Litsea Cervantesii H.B.K., op. cit. 134 [16
Tetranthera aeeadele Sprengel Syst. Hi 2: 267. 1825.
Persea ? Orizabae Martens & Galeotti in Bull. Acad. Sci. Brux. 10%: 358. 1843;
Meissner in DC. Prodr. 151: 56. 1864; Hemsley, Biol. Centr. Am. Bot. 3: 72. 1882.
Tetranthera villosa Martens & Galeotti in Bull. Acad. Sci. Brux. 107: 359. 1843.
Tetranthera — Schauer in Linnaea 19: 712. 1847; Meissner in DC. Prodr.
151: 193.
Tetranthera en var. o subsolitaria Meissner in DC. Prodr. 151: 193. 1864, p.p.
Litsea Neesiana Hemsley, |. c.; Mez in Jahrb. Bot. Gart. Berlin 5:477. 1889; Bartlett
in Proc. Amer. Acad. 44: 599. 1909; Standley in Contr. U. S. Nat. Herb. 23: 287.
2
Litsea Neesiana var. 8 corymbifera aon lic.
Litsea Neesiana var. Y villosa Hemsley, 1.
—_ Orizabae Mez in Jahrb. Bot. Gart. Berlin 5: 479. | rides in Proc. Amer.
44: 599. 1909; Contr. U. S. Nat. Herb. 23: 922.
wae guatemalensis Mez in Jahrb. Bot. Gart. Berlin . ving en Bartlett in Proc.
ad. 44: 599. 1909.
Malapoenna glaucescens Kuntze, Rey. Gen, 2: 571. 1891.
Malapoenna Orizabae Kuntze, 1.
Litsea acuminatissima Lundell “ Coie: Univ. Mich. Herb. 4: 3. 1940.
Litsea Matudai Lundell, op. cit.
Mexico: Without locality, eee & Bonpland sn. (6 fl, type of Litsea
Cervantesii not seen), Aschenborn 349 (fl, type of Tetranthera Neesiana not seen),
(fr. A). Sinaloa: Culiacan, August 27-Sept. 15, 1891, EF. Palmer 2770 (sterile,
GH, US), ?Montez a Salazar 1688 (sterile, US); Cerro del Viejo, San Ignacio, Montez
& Salazar 82 (é fi., US). Zacatecas: Plateado, Rose 3650 (@ fl., US). Nayarit
(Tepic): In the Sierta Madre, near Santa perc Rose 3437 fsterdle, US). Jalisco:
Bolafios, Rose 3746 (fr., US). Hidalgo: 1 Chico, Lyonnet 727 (fr., A, NY, US);
Real del Monte, March 22, 1849, Gregg 639 a ., ISOSYNTYPE of Tetranthera glau-
cescens a subsolitaria, GH) ae [fl.] yellowish) ; on Sierra de Pachuca, Rose & Hay
5566 (fr, US). Vera Cruz: La Joya, Perote, Balls 5520 (9 fl., fr., A); near Jalapa,
Pringle 8156 (@ fl, GH, NY, US), Schiede & Deppe s.n. (6 d., GH, NY); on eastern
slopes of mountains near Jalapa, 1280 m. alt., Humboldt & Bonpland sn. (Q fl,
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 411
SYNTYPE of Litsea glaucescens not seen); Mirador, Liebmann (Lauraceae 60) (6 fi,
US), August 1841, Liebmann s.n. (4 fl., US), Mohr sn. (6 fl., US); San Cristobal,
Orisada [Orizaba], Mohr [434] (2 fl., US); Orizaba, Balls 4320 (é fl., A), Bilimik
359 (6 fl, GH, US), Botteri 7 (2 fl, GH, US), 708, 183 (fr., GH, US), 549 (4 fl,
GH), Bourgeau 3128 (fr., GH), Galeotti 252 (fl., type of Persea ? Orizabae not cen),
Liebmann (Lauraceae 65) (8 fl., cited by Mez a Litsea Orizabae, fragm., US),
Matuda 1211 (fr., A, type af Litsea ee various localities, Schiede
58 (fr, NY). Puebla: Chinautla, alt. 2100-2400 m., May 1841, Liebmann s.n.
(sterile, GH, US). Mexico: Temascaltepec, Pineda, Tinton ae (@ fl., NY, US);
Las Cr Ag! aes 3257 (6 fl., A, US), 7223 (@ fl., GH), Salitre- Caritas, Hinton
3940 (fr., Michoacan: Sierra Torricillas, Coule ona: Hinton 12365 (6, GH),
12800 ret By. 15746 (&, US). Guerrero: On shores near Acapulco, Humboldt
& Bonpland s.n. (Q fl., syntypE of Litsea glaucescens not seen); Piedra Ancha-Tres
Cruces, Galeana, Hinton 15418 (9 fl, GH). Oaxaca: Without locality, Cuming
(¢ fl., isotype of Tetranthera glaucescens Y racemosa, NY); Talea, alt. 900 m., a
184-, Galeotti 257 (6 fl., GH [as 2977], US) (fl. white-rose), alt. 1800 m., Oct. 18
Galeotti 258 (Q fl., US) (fl. white); Villa Alta, Schultes eh ee NY): Sierra -
San Felipe, in ene and on small wooded plateaus in the untains, alt. 2700-2900
m., on lime-ridges of Sola, alt. 2400-2600 m., and in the iene of Yavesia and
Capulapan, alt. 2100-2600 m., Galeotti 251 (fl., type of Tetranthera villosa not seen) ;
Sierra de San Felipe, Pringle 5679 (@ fl., GH), Conzatti & Gonzales 1120 (9 fl., GH,
US); mountains southeast of Miahivailan, alt. 3050 m., Nelson 2531 (9 fl., GH, US);
Cafetal San Rafael (Cerro Espino), Reko 3562 (é fl, US). Chiapas: Between
Huitztan and Oxchuc, C. & E. Seler 2149 (4 fl., GH, NY, US); Saxchanal, Sierra
US
Baja Verapaz: Montana de San Ysidro-San Jerénimo, Salas 491 (fr., US); dry,
rocky hills n. of Santa Rosa, Standley 69694 (sterile, NY). Huehuetenango:
Sierra de los Cuchumatanes, above Chiantla, Standley 65630 (9, NY). San Marcos:
Above Rio Tacana, near San Antonio, Standley 66089 Sag A). Quezaltenango:
Cerro Quemado, Kellerman 5927 (2 fl., US), 5935 (2 US); Cerro La Pedrera,
south of Quezaltenango, Standley 66450 (2 fl., A, NY): slopes of Volcan de Santa
Maria, above Palojunoj, Standley 67526 (@ fl., A). Gua ala: Volcan de Pacaya,
above Las Calderas, Standley 58448 (@ fl, A). Sacatepéquez: Near San Juan,
Pgitges 613 (¢4 fl., isotype of © ihinthers ie 5 major, US); San Rafael,
J. D. Smith 1276 (é fl., GH Aare slopes of Volcan de Agua, south of Santa
de . Standley 50447 (3 ‘AL, A, NY), Volcan Agua, Ralevecn 4953 (3, US),
Maxon & Hay 3753 (2, US); Sane Maria de la Antigua, Pittier 13 (¢, US); hills ot
Finca Cae s.e. of Antigua, Standley 63668 (sterile, US). Chimaltenango
Cerro de Tecpam, region of Santa Elena, Standley 58756 (&, NY, US); Barranco de ls
Sierra, s. e. of Patziim, Standley 61605 (3, A, NY); Chichavac, Skutch 253 (6, US),
Salas 581 (6, US). Et Satvapor: Chalatenango: La Reina, S. Calderén 2456
(3, NY, US). San Salvador: Cultivated in Santa Tecla, S. Calderén 1489 (9,
GH, NY, US). Honpuras: Comayagua: Near El Achote, hills above plains of
Siguatepeque, Yuncker, Dawson & Youse 6370 (2, GH).
NATIVE NAMES: “Laurel” (Vera Cruz, Mexico, Michoacan, Guerrero, Oaxaca,
Chiapas, Baja Verapaz, Chimaltenango, Duezalienanea) “Laurill” (Zacatecas, Nayarit
{[Tepic], Jalisco) ; “Laurillo” (Michoacan) ; “Laurel de la Sierra” (Sinaloa) ; “Ziz-uch”
(Chiapas) ; “Laurel aromatico” (Petén) ; “Laurel de especie” (El Salvador)
The specimen from Chihuahua occurs in the transition pine-oak country
and, at first glance, seems to be at variance with the typical variety as we
412 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
are most familiar with it. The leaf-blades are densely pubescent beneath,
slightly smaller, acutish rather than acuminate, and oblongish lanceolate
rather than elliptic. Otherwise, the number is similar to the other speci-
mens found in the northern Mexican states.
The collections made in Sinaloa, Zacatecas, Nayarit, Jalisco, and Mexico
are similar to those found in Hidalgo, and show a tendency toward speci-
mens of the varietal segregate Schaffneri as found in Tamaulipas.
In Vera Cruz we find the typical variety in its truest sense. The lanceo-
late-elliptic acuminate leaf-blades measure 9.5 2.5-3 cm. and are
abundant on the branchlets, bearing in their axils numerous short-peduncu-
late inflorescences, simple or branched. The original description of Persea
Orizabae, from this region, is sketchy. Mez enlarges this to indicate a tree
with large leaf-blades, comparatively speaking (7.5-3.5 cm.), ovate to
lanceolate, albescent or softly cinereous-long-tomentose beneath. Except
for the pubescence, which has been found to be extremely variable in this
group, the fragment matches perfectly any number of specimens of typical
L. glaucescens from Vera Cruz and Oaxaca.
The sterile Liebmann sheet from Puebla, not far from Orizaba, Vera
Cruz, shows leaf-blades that are certainly broader than is usual with L.
glaucescens. There can be no doubt that the sheet is lauraceous and prob-
ably a variant of this species. After one hundred years in an herbarium,
the bark still has the characteristically pungent odor typical of many
Lauraceae.
The specimens collected by Hinton in Michoacan have leaf-blades that
are slightly broader in proportion to their length, and the veins are rather
more arcuate than is the case of the majority of leaf-blades found on typical
specimens. It is unfortunate that material from one of the type-localities
of the typical variety is so scanty. The Hinton plant is not typical, for it
possesses the largest-known leaf-blades of the genus in America (13 X 3.5
cm.). There is no doubt, however, that in spite of their oversize, the sheet
belongs here. The Oaxacan material was collected very near the type-
locality of the species and is similar to specimens found there. The mate-
rial from Chiapas also ties up with that from Oaxaca and Vera Cruz. The
elliptic or lanceolate-elliptic leaf-blades of the specimens are very con-
spicuously reticulate and shining. The branchlets are densely leafy and
the inflorescences are very full-flowered.
The Guatemalan specimens differ from typical L. glaucescens in con-
sistently having on the lower surface of the leaves a pubescence of varying
density, sometimes early exhibiting a thick tomentum which later may be
reduced to a few strigose hairs persisting about the costa and veins. The
young branchlets also are pubescent in varying degrees, as are the petioles,
peduncles, and pedicels. The leaves tend for the most part to be more
elliptic than lanceolate. Again these specimens must be included in the
species in the broad sense and not maintained as a separate species.
Schauer’s description of the leaf-blades of 7. Neesiana mentions the shin-
ing upper surface scattered with stellulate-pilose dots, a characteristic not
1945 ] ALLEN, STUDIES IN THE LAURACEAE, VI 413
found in the Lauraceae. The remainder of the description is typical of the
genus, so probably the type of pubescence was erroneously reported.
4b. Litsea glaucescens var. Schaffneri (Bartlett), comb. nov
Tetranthera glaucescens var. 0. subsolitaria Meissner in DC. Prodr. 151: 193. 1864, p.p.
Litsea Schaffneri Bartlett in Proc. Amer. Acad. 44: 600. 1909; Standley in Contr.
at. Herb. 23: 288. 1922
Litsea pallens Lundell in Contr. Univ. Mich. Herb. 4: 5. 1940.
DisrripuTion: Mountains of San Luis Potosi, Guanajuato, and Tamaulipas.
Mexico: Tamaulipas: Cerro los Armadillos, vicinity of San José, Bartlett
10389 (fr., GH, US) ; in shady gorges before arriving at Palmilla from Victoria to Tula,
Nov. 1830, Berlandier 2185 (=765) (isosyntype of Tetranthera glaucescens var. o
subsolitaria, GH, NY) ; vicinity of Victoria, alt. 320 m., Feb. 1—April 9, 1907, E. Palmer
208 (fr.. GH, US), Runyon 1008 (9 fl., US); San Lateas Viereck 44, 585 (6 fi. fr.,
US) ; Sierra near Victoria, alt. 1200 m., ks 1932, von Rozynski 341 (2 fl., isotype of
L. pallens, NY, US). San Luis Vieock: Near Santa Barbara, Hartweg "382 (3S fl
US); near San Luis fee lege 12-16, 1902, E. Palmer 453 (9 fl, A, US), May 24,
1905, E. Palmer 647 (fr., GH, US), Parry & Palace 798 (Q fil., fr., GH, NY, US); in
mountains, San Miguelito, acre 1876, Schaffner 23 (2 fl., TYPE of E Schaffneri, GH),
431, 463 (6 fl, NY, US), 710 (9 fl, ¢ fi, fr., GH). Guanaiieee Santa Rosa,
Duges 231 (2 fi., US), 1000 (2 fl., GH)
NaTIVE NAMES: “Laurel,” “Sacred Laurel” (San Luis Potosi).
This aggregate is one of the extreme variants from the norm of Litsea
glaucescens. The shrub bears slender foliose ochraceous branchlets, with
narrowly lanceolate mucronulate leaves, acute at both ends, more or less
coriaceous, glabrous and glaucous beneath, the venation inconspicuous
except for the prominent midrib and reticulation. The leaf-blades vary in
size, being 2-6 cm. X 5-12 mm. The inflorescences are usually solitary
pedunculate umbels located in the axils of the leaves of terminal or lateral
branchlets. The rather large globose and black fruits are about 12 mm
‘in diameter.
The specimens from Tamaulipas differ from those from the type-locality
in their larger more acuminate leaf-blades, not glaucous but concolorous,
ith more prominent venation. Lundell notes the similarity between L.
ee and L. Schaffne
4c. Litsea glaucescens var. flavescens (Bartlett), comb. n
Litsea He asi Bartlett in Proc. Amer. Acad. 44: 599, 1800 Standley in Field Mus.
Publ. . 18:
Rea rain ee regions of Costa Rica, at an altitude of 1500-1900 m.
Costa Rica: San José: Hills above Belmira, near Santa Maria de Dota, alt.
00 m., Jan. 1898, igi 7352 (Herb. Nat. Costa Rica 11638) (4, 9 fl., TYPE of
a ae GH, NY, US), Standley 42525 (8 fi., US); Cuesta de Tarrazu, Tonduz
7796 (6 fl., US); in area of Candelaria, Ontiel 10 (é fi., US).
NaTIVE NAME: “‘Lentisco.”
This Costa Rican segregate is distinguished by the striking flavescent
venation, particularly the prominent marginal vein, and the yellowish
petiole apparent in the dried specimens, as well as the lack of glaucosity on
the lower leaf-surface. Bartlett also mentions the smaller flowers and the
tomentose pedicels as presenting specific differences. Both of these char-
acters seem of too variable a nature to be criteria of specific importance.
414 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
Pubescence may be found on many specimens of the typical variety; and
some of the Chiapas material shows a similarity of foliage-characters.
Since no specimens have been collected as yet in the intervening areas, the
entity is maintained for the present as a regional variety of L. glaucescens.
6. Beilschmiedia Nees
ee Nees in Wallich, Pl. As. Rar. 2: 61, 69. 1831, Syst. Laurin. 21, 192, 197.
eissner in DC. Prodr. 151: 62. 1864; Kostermans in Rec. Trav. Bot.
Dullantin Nees, Pl. Laurin. nea ie (n.9), 21. 1833, Syst. Vintieii: 187, 674. 1836;
Meissner in DC. Prodr. 15}:
DistriBuTION: Tropical regions ¥ pan hemispheres.
KEY TO THE SPECIES OF BEILSCHMIEDIA
A. Leaf- — on both surfaces prominently and loosely and often incompletely
reticulat
B. Leaf- ‘blades cote not less than 7 cm. in width, glaucous ne fruits to
. B. Ana
5 CM M0 TOURED o.gaik cna bo Sa Sao Ge Sa Sa eee ess eter ase ee eee a5 eeF
B. Leaf-blades
a
Qa
a
434 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXvI
E. Panicles usually less than 12 cm. long; flowers not whitish gray-
DUbEsCent.s 56 ws sess e's ove aie Mais eo ae ae asa e sees lla. P. Johnstonii.
D. Inflorescence composed of axillary and subterminal short loose few- to
many-flowered panicles less than 10 cm. long and more or less glabrous.
E. Largest leaf-blades not more than 15 cm. long; lowermost pair of
lateral nerves woe iia above the middle of the blade and less
stout than the co
F. Inflorescence many- flowered, not more than 8 (rarely 12) cm. long;
flowers not pruinose and not more than 4 mm. long...............
ee ee ee eee ee oe ee lla. P. Johnstonii.
F. anlage few-flowered, up to 12 cm. long; gee pruinose and
OUR iis ca se eewlswhee en aeaeewn P. Ehrenbergii.
E. Largest leaf - blades 18-20 cm. long; lowermost pair a lateral nerves
conspicuous 4 to % the length of the blade and as stout as the costa..
j dG ntad a cis: 4.aik Rate SS Dales aes Gua alee eee miew arate 4Hs 13. P. neurophylla.
22. Ocotea rubrinervis Mez; Allen in Jour. Arnold Arb. 26: 353. 1945.
The nine Johnston numbers cited as “in mixed forest along South Road
(S2 area),” actually were collected in various areas on San José Island.
31. Nectandra Whitei (Woodson) Allen in Jour. Arnold Arb. 26: 398. 1945.
id 6059, et as from Costa Rica, actually was collected in Panama.
riqui: Vicinity of Camp El Volcan, Little 6047 (fl., USFS), 6056
is x USFS), 6059 (young fr., Ch, USFS), 6062, 6069 (young fr., USFS).
ARNOLD ARBORETUM,
Harvarp UNIVERSITY.
1945 ] CRONQUIST, DIPHOLIS AND BUMELIA 435
STUDIES IN THE SAPOTACEAE, III
DIPHOLIS AND BUMELIA
ARTHUR CRONQUIST
A REVISION of certain groups of American Sapotaceae has been under-
taken under the auspices of the Chicle Development Co. The present
paper comprises a revision of the genera Dipholis and Bumelia. Current
interpretations of the identity of the older types have in general been
accepted, since these are in most cases not now availabie for study. In
accordance with Article 19 of the International Rules of Botanical Nomen-
clature, names which were not validly published have rigorously been
excluded from consideration. My concepts of intraspecific units have been
given in a previous paper (Bull. Torrey Club 70: 265. 1943). Although
a relatively large amount of material, measured by the standards of tropical
botany, has been available for study, I keenly feel its inadequacy in a
number of species; my confidence in the accuracy of my taxonomic interpre-
tations varies in approximately direct proportion to the number of specimens
seen.
I wish to acknowledge my thanks to Mr. B. A. Krukoff, under whose
direction this study was prepared, and to the curators of the several herbaria
who have provided essential assistance by kindly loaning specimens for
study. The abbreviations used hereinafter to indicate these herbaria are
as follows:
A — Arnold Arboretum, Harvard University.
CR — Museo Nacional de Costa Rica, San José.
Cu — Estacion Experimental Agronomica, Habana, Cuba.
F — Field Museum (Chicago Natural History Museum).
G — Gray Herbarium, Harvard University.
Mich — University of Michigan, Ann Arbor.
Mo — Missouri Botanical Garden, St. Louis.
NY — New York Botanical Garden.
PA — Philadelphia Academy of Natural Sciences.
PR — Tropical Forest Herbarium, U. S. Forest Service, Rio Piedras,
Puerto Rico.
US — United States National Herbarium, Washington, D. C.
DIPHOLIS
The genus Dipholis was founded in 1844 by Alphonse de Candolle, in the
Prodromus, vol. 8, p. 188. The single species listed, D. salicifolia, neces-
sarily becomes the type. This species has, incidentally, by far the widest
distribution of any in the genus, and is biologically as well as nomencla-
turally typical. Dipholis is antedated six years by Spondogona Raf.
(Sylv. Tellur. 35. 1838), based on Bumelia pentagona Sw., which is now
436 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
referred to Dipholis salicifolia on the authority of Radlkofer (Erg. Mon.
Serjania, pp. 55-56. 1886). Swartz described the fruit as 5-angled, and
Rafinesque, apparently without seeing the type, enlarged upon this descrip-
tion to say that it was also S-seeded. A 5-seeded or even 5- angled fruit
in Dipholis salicifolia, or any other known species of the genus, would be a
monstrosity, and any name founded on such a specimen should be rejected
under Article 65 of the Rules. I should not hesitate to recommend that
Dipholis be conserved, were it necessary, but Spondogona can be rejected
on another basis.
Dipholis is closely related to and evidently derived from Mastichoden-
dron,! from which it differs primarily in the presence of lateral lobes on
the corolla-lobes. Several other usual differences, none entirely constant,
are found in the more petaloid staminodes, more nearly basal seed-scar,
and frequently smaller leaves and fruit of Dipholis, the leaves lacking the
deeply channeled midrib and petiolar pouch so commonly found in
Mastichodeydron. One species has been described as lacking the lateral
lobes on the corolla-lobes, but in my opinion this species is better referred
to Bumelia. The relationship and distinctions between Dipholis and
Bumelia are discussed under the latter.
Dipholis is confined to the warmer parts of North America and reaches
its greatest development in the Greater Antilles, where 10 of the 14 known
species occur. Evolutionary trends in the genus are toward reduction in
size of the plant, size of the leaves, size of the fruit, and number of flowers
in a Cluster, and toward the development of a truly basal seed-scar. The
most primitive surviving species, such as D. Stevensonii and D. minutiflora,
are good-sized trees with large leaves, large (2-3 cm.) fruits, numerous
flowers in a cluster, and somewhat basilateral seed-scar. These species,
particularly D. Stevensonii, are not far removed from the generic prototype.
Dipholis nigra is closely allied to D. minutiflora, and may be nearly ancestral
to D. salicifolia. Dipholis Jubilla and, from the description, D. Bellonis
seem most likely to be related to D. nigra. Dipholis parvifolia and D.
durifolia seem to be allied to D. salicifolia. The closely related species
D. montana and D. octosepala are probably derived from the D. nigra-
salicifolia stock. Dipholis cubensis, D. repens, and D. ferruginea form a
closely knit group that is related to D. montana. Dipholis sericea is
probably allied to the D. cubensis group.
Dipholis A. DC.
Unarmed shrubs or trees; leaves alternate, exstipulate, often small; pri-
lobes or appendages near its base; staminodes present, more or less petaloid,
these and the appendages more or less erose, fimbriate, or laciniate; ovary
name Mastichodendron, as used by Lam (Rec. Trav. Bot. Neéerl. 36: 521.
oo is at this writing not yet validly published. It is intended to validate this name
in another paper, publication of which may precede this paper
1945 | CRONQUIST, DIPHOLIS AND BUMELIA 437
glabrous, very rarely shortly appressed- hairy; ovules 5, attached basi-
un
persistent style; fruit fleshy, mostly 1-seeded, not over about 3 cm. long;
seed-scar very nearly basal, rarely evidently basilateral : endosperm well-
developed
KEY TO THE SPECIES
1. Fruit about 2 cm. long; leaves 8-12 cm. long; poorly known plant of Puerto Rico...
BI Peeks Pee adeeb ane Maries tera ene mse tae ress ay Giro wee GENO bk age ORE aE Gea EE erator Dn LD eB CLL ONES
1. Fruit (where known) about 1 cm. long or less, except in some Jamaican and con-
tinental species; leaves various.
2. Inflorescences borne on peduncles up to 15 mm. long, sometimes slightly branched;
LO) a Yet sll Gin | of: peek a ween or eS ne I ese SRP e ee . D. Jubilla.
2. Inflorescences sessile, si sdccat
Flow
4. Fruit 12-30 mm. long; pedicels 4— 15 mm. long; leaves often but not always
retuse or rounded; sepals glabrous or hairy.
5. Leaves densely and loosely rufous-hirsutulous beneath, eventually more or
less glabrate; pedicels 10-15 mm. long; petioles 2.5-4 cm. long; sepals
strongly and loosely hairy, about 3.5 mm. long or more; British Honduras.
Breet eee ay eet eEe sd 0.64 305 @ 46, 9 sa ede eee ee 1. D. Stevensoniz.
5. Leaves glabrous or merely sericeous- oe beneath, the hairs appressed;
0.5
pedicels 4-10 mm. long; petiol . long; canals glabrous or wi
few appressed hairs, at bapa time ns than 2.5 mm. long.
ruit about 1.5-3 long, yellow to purplish; leaves tardily glabrate
beneath, the hairs eee continental:.. «cise ree 2. D. minutiflora.
6. Fruit 12-16 mm. long, black; ae very soon glabrate, the i. when
present often reddish; Jamaica.............. 02. c cece cece de . nigra.
4. Fruit 6-10 mm. long; pedicels 1-4 mm. long; leaves acute or scuminate sepals
distinctly hairy; general ve aees 10) 0 eS Ree wei ee eh ee 6. D. salicifolia.
3. Flowers few, commonly ab 1-7 in a cluster, borne in the axils.
4. Leaves strongly ee closely and sonemicuaieny erie rom veiny; con-
ti nental.
ban about 6-10 cm. long, borne on petioles about 1.5—2.5 cm. long; pedi-
s 5-7 mm. long; sepals eee and rather loosely ferrugineous, about
de
stim melone> British Monduraseucus. «ctor cena aso vee se D. durifolia.
. Leaves ne 2.5—4 cm. lon ae on petioles about 3-7 mm. long; flowers
"cabs the sepals plenrous or slightly strigose, Boa 2 mm. long; Costa
IRI CAS tase) oaneis ord te eee et ae ced. sk 8 8. parvifolia.
ede mostly obtuse or rounded, if evidently acuminate then not ‘gosly and
conspicuously degree veiny; Greater Antilles
5. Corolla at lea m. long, or if perhaps Someumes not so, then the sepals
8 or 9; ee comm eae but not always more than 5; Jamaica
6. Outer 2 sepals essentially glabrous; fruit about 13-18 mm. long; veiny
-
reticulum of the leaves scarcely raised; sepals 5-8......... 9. D. montana.
6. Sepals all hairy; fruit about 1 cm. long or less; veiny he cas of the
leaves ne ele beneath ; sepals BRO TRO ors as boc eens . D. octosepala.
5. Corolla not over 4.5 mm. long; sepals 5, rarely 6; not of eee,
6. Leaves en Beare sparsely hairy hea 5 roung.
7. Erect shrubs or trees; leaves 1.5-10 cm. long ; Cuba, Hispaniola, and
_ Puer to BIC On coer icnctecuaeiar een hoe eR ee sei gece 11. dD. cubensis.
Sra ne castrate eva eec Unser artesian Gi chp Maula lg uenal aetgs a amete Need ete Re pte wate ct) etal ale gas cts . repens.
6. Leaves strongly rufous-sericeous beneath, eventually paler or glabrate;
Dominican Republic
438 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
7. Leaves 4— 8 cm. long, 1.5-3 cm. wide, eventually glabrate; panne
3. D.
evident, raised............ cc cece eee e eter cece neeeeee ricea.
7. Leaves 2-4 cm. long, 1-1.5 cm. wide, acme’ Feige heer
ODECUIOs 2 i435c suds a teed oe ic ese cate passeewauaes . ferruginea.
1. Dipholis Stevensonii Stand]. Trop. Woods 11: 1927.
— Rie eavaareias Standl. Carnegie ce Wash. Misc. Publ. 461(Bot.
Maya a 4):
Tree up to 25 m. pene leaves elliptic or elliptic-obovate, rounded at the
apex, about 10-22 cm. long and 6—12 cm. wide, strongly veiny, the primary
hirsutulous-tomentose with rufous hairs at first, soon glabrate above, even-
tually more or less glabrate beneath; petioles 2.5—4 cm. long; flowers nu-
merous in clusters at defoliated nodes, the pedicels about 10-15 mm. long,
rufous-tomentose; mature flowers unknown, but maturing buds 5- or
6-merous, the sepals about 3.5 mm. long or more, the corolla-lobes 2 mm.
long, the anthers 1.5 mm. long, the staminodes ovate, scarcely erose, nearly
1.5 mm. long, the lateral appendages narrow, entire or subentire, only 0.5—1
mm. long; fruit ellipsoid, about 16-25 mm. long, resembling that of Masti-
chodendron, seed up to about 16 mm. long, the scar basilateral, ellipsoid,
5 mm. long, the seed-coat about 1 mm. thick.
TYPE COLLECTION: Stevenson s.n., British Honduras, 1927 (F—fragment, US).
Loca, NAME: Zapote faisan.
DistRIBUTION: British Honduras.
British Honpuras: Lundell 6200 (F, G, NY, US), 6252 (G, NY, US); Schipp
S674 (F, G, Mo); Stevenson s.n. (1926) (US).
The large veiny leaves, somewhat channeled midrib, relatively long
petioles, relatively large fruit, and basilateral rather than basal seed-scar of
this species are suggestive of Mastichodendron, to which it makes an
approach, but the petaloid staminodes and the definite, if smaller than
usual, lateral lobes on the corolla-lobes clearly indicate its place in Dipholis.
2. Dipholis minutiflora Pittier, Contr. U. S. Nat. Herb. 13: 464. 1912.
Sideroxylon Matudai Lundell, Phytologia 1: 221. 1937.
Bumelia tabascensis Lundell, Contr. Univ. Mich. Herb. 5: 22. 1940.
Sideroxylon Steyermarkii Standl. Field Mus. Publ. Bot. 22: 368. 1940.
Dipholis Matudai Lundell, Contr. Univ. Mich. Herb. 7: 43. 1942.
Tree up to 30 m. high; ee elliptic- oo or narrowly elliptic-
obovate, rounded to acutish at the apex, tapering to the base, about 6—20
cm. long and 2.5—-10 cm. wide, with 10-20 pairs of primary lateral veins,
glabrous above, finely white-strigose beneath, eventually glabrate; petioles
about 0.5—1.5 cm. long; flowers very numerous in clusters at defoliated
ovate, acuminate, erose-laciniate, 1.5 mm. - long on the i margin; fila-
ments firm, about 2—2.5 mm. long; anthers about 2 mm. long, sagittate to
the middle or beyond; staminodes laciniate, 2.5—3 mm. long; fruit yellow,
olive-green, or purplish, ovoid to subglobose, usually conical-acute at the
apex, about 1.5—-3 cm. long, with thin pulp; seeds about 14-18 mm. long, a
little compressed, with broadly ellipsoid basal or basilateral scar 5 mm.
long; seed-coat about 0.5 mm. thick.
1945] CRONQUIST, DIPHOLIS AND BUMELIA 439
PE COLLECTION: Tonduz 11935, forests of El Copey, Dota Mountains, Costa Rica,
aot 1800 m., February, 1898 (NY, US).
LocaL NAMES: Sapote prieto, ees
DistTRIBUTION: State of Mexico, Mexico, to the Panama Canal Zone.
Mexico: Mexico: Hinton 4923 (G), 7700 (G). Guerrero: Hinton 10416
(G). Tabasco: Matuda 3455 (Mich, NY). Chiapas: Matuda 571 (A, Mo,
US), 4175 (A, F, NY). Guatemara: San Marcos: Steyermark 37615 (F).
Quezaltenango: Steyermark 33858 (F). Suchitepéquez: Steyermark 35409
(F). Honpuras: Comayagua: Edwards P-302 (A, F). Yoro: von Hagen
1007 (F, NY). Costa Rica: Austin Smith 4184 (F); Standley & Valerio 46505 (A,
US), 46481 (US). Panama: Chiriqui: Allen 1564 (F, G); White 109b (F, G);
Woodson, Allen & Seibert 995 (A, F). Canal Zone: Allen 1314 (F, G, US)
3. Dipholis nigra (Sw.) Griseb. hr Brit. W. Ind. 400. 1861.
Bumelia nigra Sw. Prodr. -
Achras nigra Poir. in Lam. Encyc. Meth. 6: 532.
Dipholis nigra var. ea ae Oe Urb. Symb. ie rs ae 1904.
Tree up to 15 m. tall; leaves elliptic or narrowly elliptic-ovate, commonly
acute or acutish, sometim es obtuse or bluntly acuminate, glabrous, sige
with evident reticulum, commonly . 18 cm. long, 3—6.5 cm. wide, the
petioles 1.5—3 cm. long: flowers numerous in Clusters at defoliated ae
the pedicels about 4-10 mm. long, essentially glabrous; sepals essentially
eee about 1.5—-2 mm. long; corolla about 3.9-4.4 mm. long, the tube
1.1-1.5 mm.; corolla-lobes cordate-auriculate above the lateral lobes, which
are er and broader than those of D. salictfolia ; filaments up to 3 mm
Swe stout, except for the tenuous tip, as in D. salicifolia; anthers about
1. ong, more evidently sagittate than in D. salicifolia; stami-
andes ovate, petaloid, erose, about 2 mm. long or more; style 1 mm. long or
less, conical; fruit black, ellipsoid to ovoid-globose, 12-16 mm. long, up to
TYPE COLLECTION: None given; reference to Browne, Hist. Jam. 201. 1756.
LocaL NAME: Red bullet or bully.
DIsTRIBUTION: Jamaica; reported, probably incorrectly, from Cuba.
Jamaica: Alexander s.n. (1850) (G, US); Britton 1105 (NY), 3688 (NY); Harris
5388 (A, G, NY, US); Miller 1389 (US); Rehder s.n. (Feb. 13, 1903) (A); Hart 1057
(US); Taylor 220 (NY
4. Dipholis Jubilla Ekm. ex Urb. Symb. Ant. 9: 415, 1925.
Tree; leaves lanceolate or elliptic, glabrous nearly from the first, com-
only ce 15 cm. long, 2—5.5 cm. wide, acute or acuminate, the primary
lateral veins ecient beneath, the ae ones obscure, not forming an
evident reticulum; petioles about 1—2.5 cm. long; flower- clusters borne on
peduncles up to 15m m. long, the inflorescence sometimes slightly branch-
ing; pedicels about 2— i mm. long, finely ferrugineous; sepals finely and
lated ferrugineous, 2.5 mm. long in bud; mature flowers and fruit un-
known
E COLLECTION: Ekman 8324, “ad Alto de Iberia in cacumine montis cr. 1135
»’ November 12, 1916 (A, F, NY).
ooo NAMES: Jubilla, juba prieto.
DIsTRIBUTION: Oriente, Cuba.
Cu Oriente: Ekman 4249 (G, NY), 5574 (US), 9369 (NY), 15659 (G, US);
Leon 19677 (NY) ; Roig 5316 (Cu), 6209 (Cu, NY).
440 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
The leaves of D. Jubilla are somewhat similar to those of D. nigra, but
may be distinguished by the lack of an evident veiny reticulum.
5. Dipholis Bellonis Urb. Symb. Ant. 5: 137. 1904.
Leaves 8—12 cm. long, 2.5—4 cm. wide, ovate-oblong, acute at the base,
acuminate at the apex, borne on petioles 1—2 cm. med fruiting pedicels
10-15 mm. long; fruit obovoid, sometimes narrowly 18-20 mm. long,
10-15 mm. thick, violet-blackish. (Description He Si from the ma
no specimens seen.
Type COLLECTION: Bello, near Furnias, Puerto Rico.
LocaL NAME: Varital.
DistrRiBUTION: Known only from the type collection, Puerto Rico.
6. Dipholis salicifolia (L.) A. DC. in DC. Prodr. 8: 188. 1844.
Achras salicifolia L. Sp. Pl. ed. 2. 1: 470. 1762
Bumelia salicifolia Sw. Prodr. 50. 1788.
Bumelia pentagona Sw. loc. cit.
Sideroxylon pauciflorum baie Tab. Encyc. 2: 42. no. 2459. 1793.
Achras pentagona Poir. in Lam. Encyc. Meth. 6: 533. 1804.
Sideroxylon salicifolium Gaertn. {. Carp. _— 124. t. 202. 1805.
Spondogona nitida Raf. Sylv. Tellur. 35. 18
Sideroxylon pentagonum A. DC. in DC. Cah 8: 185. 1844.
Dipholis salicifolia var. jamaicensis Pierre in Urb. “eer ~ 5: 139. 1904.
Spondogona salicifolia House, Am. Midl. Nat. 7: 131.
?Dipholis leptopoda Urb. Ark. Bot. 22A(17): 70. 1929.
Large shrub, or more commonly a tree, sometimes nearly 25 m. high;
leaves narrowly elliptic to elliptic-lanceolate or elliptic-oblanceolate, acute
or acuminate at both ends, commonly 5-11 cm. long and 1.5—4 cm. wide,
villosulous when young, but very soon glabrate, the petioles short, com-
0.5-1.5 cm. long; flowers numerous in clusters at defoliated nodes
or sometimes in the axils, the pedicels finely sericeous, about 1-4 mm. long;
sepals finely hairy, about 1.4-3 mm. long; corolla about 3.3—4.5 mm, long,
the tube 1.3-1.6 mm.; corolla-lobes rounded, elliptic, narrowed to a short
claw-like base above the lateral lobes or appendages, which are acute or
acuminate, lanceolate or ovate; staminodes ovate or broadly elliptic,
ere about 1.5-2 mm. long, more or less erose-laciniate; filaments about
mm. long, very stout and firm except for the short abruptly tenuous
ree portion; anthers about 0.9-1.6 mm. long, ayes ovary 5-loculate,
glabrous or rarely slightly hairy, the style 1.5-2.5 mm. long; fruit black,
with thin pericarp, broadly ellipsoid or subglobose, about 6 6-10 mm. long,
with | or sometimes 2 or 3 seeds; seed-scar basal, circular or elliptic, 1 mm.
in greatest diameter; seed-coat sae 0.2-0.3 mm. thick, light to dark
brown.
TYPE COLLECTION: None given; references to works of Sloane and Browne on
Jamaica
Caan NAMES: White bullet or bully, mijico; for many others see Symb. Ant. 5:
138. 1904
DistripuTion: Southern Florida; Bahama Islands; Greater Antilles: Virgin Islands,
and sparingly through the Lesser Antilles to Guadeloupe and the Barbados Islands;
southern Mexico, Guatemala, and British Honduras. i ioe from Florida are so
numerous that I have cited only a small proportion of t
US.A.: Florida: Curtiss 1760 (A, NY, US), 5859 nn US); Duckett 223 (A,
F, NY, US); Moldenke 750 (NY, US), 5701 (NY); Pollard et al. 197 (NY, US);
1945 | CRONQUIST, DIPHOLIS AND BUMELIA 441
Small & ae 5509 (NY), 5652 (NY), 5760 (NY), 6553 (NY), 6591 (NY, US);
Tracy 9252 (NY, US). Banamas: Brace 3893 (NY, US), 4620 (NY), 4932 (NY),
6737 (NY); ee 10 (NY, US), 112 (NY), 3376 (G, tees 6434 (NY, US); Britton
& Brace 188 (NY); Britton & Millspaugh 2579 (NY), 0 (NY); Coker 257 (NY),
395 (NY); Cooper 3 (NY); Curtiss 148 ee G, NY, Hee Farle 50 (NY); Eggers
4106 (F, NY, US); Northrop 326 ed F, G, NY); Small & Carter 8459 (G, NY, US);
8287 (NY), 8422 (NY). CuBa: is de Pinas: Briton. Wilson & to 15270
(G, NY, US). Habana: Ledn 8524 (NY), 10674 (NY); Leén & Edmund 8764
(NY). Matanzas: Britton & Wilson 14045 (US). Santa Clara: Britton, Earle
& Wilson 5888 (NY, US) ; Britton & Wilson 5527 (NY); Ekman 18353 (A); Jack 4215
(A), 4351 (A), 4808 (A, US), 5608 (A, US), 5661 (A), 5675 (A, US), 6015 (A, US);
Leén 398 (NY); Rehder 1119 (A); Shafer 12310 (NY, US). Camaguey: Shafer
436 (NY), 677 (G, NY, US), 934 (G, NY, US), 1045 (G, NY, US). Oriente: Brit-
ton 1875 (NY); Britton & Cowell 12613 (NY, US); Britton, Cowell & Shafer 13065
1313 (NY), 1503 (NY, US); Wright 1325 (G, NY, US). Jamatca: Alexander s.n.
(1850) (NY, US); Britton 1887 (NY), 1949 (NY); March s.n. (NY); Eggers 3509
(A, US); Hines sn, (1897) (A, F, US); Harris 5511 (NY), 7070 (NY), 8642 (NY),
8949 (NY), 9616 (NY, US), 10170 (NY, US), 10806 (NY, US); Hart 611 (NY);
Miller 1363 (US), 1413 (US); Perkins 1264 (G); Rothrock 141 (NY), 231 (F).
Haiti: Holdridge 987 (US); Leonard 3204 (G, NY, US), 8870 (G, NY, US), 12376
(A, G, US), 14850 (NY, US), 15743 (A, G, NY, US). Domunican Repusiic: Abbot
1295 (G, US), 2248 (G, US), 2867 (G, US); Ekman 14380 (US); Fuertes 210 (A, G,
NY, US); Rose, Fitch & Russell ae (A, G, NY, US); Scarff 2a (F); Wright et al.
305 (F). Puerto Rico: Britton & Brown 6367 (NY); Britton & Marble 2333
(NY, US); Holdridge 321 (PR); iieuion 909 (NY), 1862 (NY); Sargent 438 (US);
Shafer 2761 (NY, US) ; Sintenis 733 (G), 733b (NY, US), 3640 (G, NY, US) ; Steven-
dingh 1168 (NY). GuapELouPE: Duss 2913 (A, F, G, NY, US); Stehlé 2870 (US).
RIE GALANTE: Stehlé 420 (NY). Barpapos: Bovell 468 (NY). Sr. ANDREW
(Barbados): Bovell & Freeman 374 (NY). Mexico: Vera Cruz: Gaumer 2150
(G); Purpus 2031 (NY, US). Puebla: Purpus 3382 (F, US). Campeche:
Lundell 1346 (F, NY, US). Yucatan: Flores s.n. (1936) (F); Gaumer s.n. (1888)
(F), 1047 (A, F, G, US), 2150 (A, F, US), 23516 (A, F, G, US), 23632 (A, F, G, US);
Lundell 7557 (A); Seler 3993 (A, F, G, US). Guatremata: El Petén: Bartlett
: a 9 (A, F, NY, US); Lundell 3058 (F, G, US), 3098 (F), 3197 (F), 3354 (A),
0 (F). Bemce Honpuras: Karling s.n. (1931) (A, F, US); Lundell 3895 (F);
pe S-650 (A, F, G, NY).
Dipholis leptopoda Urb. & Ekm. was based on some immature specimens
which have unusually few flowers for D. salicifolia, but which do not seem
to belong anywhere else. The leaves are also broader and more obtuse
than is general for the species, and it may eventually prove to be distinct.
7. Dipholis one Standl. Carnegie Inst. Wash. Misc. Publ. 461 (Bot. Maya Area
7 Oa op
Tree 6 m. ee leaves mania strongly acuminate, 6-10 cm. long,
1.5—2.5 cm. wide, quite gla at lea st at maturity, conspicuously and
finely reticulate-veiny on both "aides petioles about 1.5-2.5 cm. long;
flowers 1-6 in the axils, the stout ferrugineous pedicels 5-7 mm. long;
sepals strongly ferrugineous, about 3.2-4 mm. long; corolla about 5 mm.
ong or more, the tube about equaling the lobes; staminodes petaloid, ovate,
erose, about = 2.5 mm. long; ovary glabrous, - celled, its style about 2-2. 5
m. long: fruit unknown.
442 JOURNAL OF THE ARNOLD ARBORETUM [VoOL. XXvr
Type COLLECTION: Schipp 1202,
feet, September 8, 1933 (A, F, G,
DistrRIBUTION: Known only from the type collection, British Honduras.
This species and D. parvifolia are evidently closely related; their rela-
tionship to other species in the genus is uncertain, but D. salicifolia may be
suggested as the nearest relative.
8. Dipholis parvifolia Standl. Field Mus. Publ. Bot. 18: 909. 1938.
Tree; leaves elliptic-lanceolate, long-acuminate, 2.5—4.5 cm. long, 1—1.5
cm. wide, glabrous, firm, closely and conspicuously reticulate-veiny on both
sides, but the veins not much raised; petioles 3—7 mm. long; flowers borne
singly or in very small groups in the axils, nearly sessile, the sepals about
2 mm. long, sparsely strigose or glabrous; fruit obovoid, about 17 mm. long.
T COLLECTION: Standley & yee 45525, wet forest, Los Ayotes, near Tilaran,
Guanacaste, Costa Rica, 600-700 m., January 21, 1926 (US).
DistRIBUTION: Known only eee the type collection, Guanacaste, Costa Rica.
9. Dipholis montana (Sw.) eg e Fl. Brit. W. Ind. 401. 1861.
Bumelia montana Sw. Prodr. 788
Achras montana Poir. in Lam. geal Meth. 6: 533. 1804.
Dipholis pallens Pierre & Urb. in Urb. Symb. Ant. . 136. 1904.
Dipholis lanceolata Pierre in Urb. Symb. Ant. 5: 136. 1904.
Shrub or small tree sometimes as much as 15 m. high; leaves elliptic to
broadly or narrowly obovate, broadest at or generally above the middle,
commonly 3-11 cm. long, 2—5.5 cm. wide, obtuse to abruptly short-
acuminate, ees fais veiny, but the reticulum scarcely raised; petioles
about 3-10 mm. long; flowers in axillary clusters of about 2—7, the stout
pedicels thinly strigose, 2—6 mm. long, elongating in fruit; sepals 5—8, the
2 outer glabrous or sparingly strigose, generally shorter than the others,
about 2-4 mm. long, the inner evidently though finely sericeous-strigose,
commonly 3—4 mm. long; corolla about 6.3 mm. long, the tube and bases
of the as thick and firm, the lobes and their appendages thin;
corolla-tube about 3.2 mm. long or more; filaments about 1.5 mm. long or
a little less, stout, tapering from the base; peat about 0.8—1.1 mm. long;
staminodes orbi cular- obovate, about 1.5 m 1. long, fimbriate, thickened
toward the base; style about 1-2 mm. long, ie stigma slightly expanded;
ovary 5-loculate: fruit ellipsoid or ellipsoid-ovate, about 13-18 mm. long,
9-12 mm. thick; seed (from fruit 18 mm. long) 13 mm. long, 10 mm. thick,
with basal or elliptic scar 3.5 mm. long; seed-coat dark brown, 0.4—0.6 mm.
thick.
TyPE COLLECTION: None g
LocaL NAMES: White ne eas or hats, black bullet or bully.
DIsTRIBUTION: Jamaica.
Jamaica: Britton 270 (NY), 1107 (NY); Harris 5340 (NY), 5355 (NY), 5370
(A, G, NY), 5704 (A, F, US), 5777 (NY), 6691 (A, NY, US), 6731 (NY, US), 9742
(NY, US), 9803 (NY, US), 10118 (NY, US), 10807 (A, G, NY, US); Hart 533 (US),
642 (F, US); Miller 1267 (US), 1297 (US); Shreve sn. (May 16, 1906) (NY).
10. Dipholis octosepala Urb. Symb. Ant. 7: 324. 1912.
Tree up to 10 m. high; leaves rufous-tomentulose when young, later
glabrate, elliptic to ovate or obovate, 9-12 cm. long, 4-6 cm. wide, obtuse
to acuminate, the veins obscure above, raised and forming an evident retic-
bare hilltop, Jacinte Hills, British Honduras, 700
NY)
1945] CRONQUIST, DIPHOLIS AND BUMELIA 443
ulum beneath; petioles 1.5—2.5 cm. long; flowers about 2-4 in the axils,
the stout pedicels up to 5 mm. long, rufous-hairy; sepals about 8 or 9,
spiraled, all finely and loosely rufous-hairy, about 3-4 mm. long; mature
corolla unknown; lateral appendages of the corolla-lobes erose-laciniate;
staminodia well developed, fimbriate; fruit ellipsoid or or gael about
1 cm. long or a little less, with 1 or 2 seeds; seed-coat about 0.3 mm. thick.
Type COLLECTION: Harris 10986, “ultra Clarendon in sylvis Peckham ane July,
Jamaica (NY).
DISTRIBUTION: Jamaica, all of the anes from the same station.
Jamaica: Harris 11049 (NY), 12798 (NY, US).
In its few-flowered clusters, stout as and numerous sepals, this
species resembles D. montana, but differs in having all the sepals loosely
hairy, in having a smaller fruit, and in the differently shaped thicker leaves
with longer petioles and with raised veiny reticulum beneath.
11. Dipholis cubensis (Griseb.) Pierre in Urb. Symb. Ant. 5: 140. 1904.
Bumelia cubensis Griseb. Cat. P]. Cub. 164.
Dipholis cubensis var. oblongata ae ee = Urb. Syiih. re - ae 1904.
Dipholis domingensis Pierre & Urb. in Urb. Symb. Ant. 5: 1904.
Dipholis Sintenisiana Pierre in Urb. Sy - Ant: 32139. reg
phi angustifolia Urb. Symb. Ant. 7: 323. 1912
Dipholis Ekmaniana Urb. Symb. Ant. 9: 416. 1925,
neh branchen shrub or small tree up to 10 m. tall; leaves firm, gla-
brous, oblanceolate to obovate or elliptic, acute at the base, rounded to
acutish at the apex, about 2—10 cm. long and 6-30 mm. wide; petioles
2-10 mm. long; flowers about 1—4 in a cluster, the pedicels 2-20 mm. long,
glabrous or hairy; sepals about 1.8—2.5 mm. long, glabrous or strigose;
corolla about 2.7—3.7 mm. long, the lobes a little shorter than the tube,
Log baie bears scattered long red hairs; filaments about 0.5—1.6
- anthers about 0.8 mm. long; staminodes petaloid, erose or
ene less than 1 mm. long; style short and stout, about 0.4-0.7 mm.
long; fruit purple or reddish, or perhaps sometimes green, about 5-13 mm.
long and 3—7 mm. thick, oliviform-ellipsoid to ovoid; seed-scar basal, a
little over 1 mm. wide; seed-coat about 0.4 mm. thick.
TYPE COLLECTION: Wright 2921, near Monteverde, Oriente, Cuba (G, Mo, NY,
US — dra
Sen clonte Pinar del Rio and Oriente, Cuba, to Hispaniola and Puerto Rico.
Cu Wright 1326 (G), 1327 (G). Pinar del Rio: Ekman 17527 (NY, US);
Ledn 13200 (NY), 13837 (NY). Oriente: Ekman 2296 (NY, US), 2713 (F), 5562
(G), 6017 (NY), 9330 (NY, US), 9407 (NY, US), 9886 sen 10008 (NY), 14326 (A,
F, G, US); Ledn 10828 (NY), 11959 (NY) ; Ledn et al. 10210 (NY); Roig 47 (NY),
6550 (Cu); Shafer 3033 (NY), 3461 (NY, US), 3563 (NY, US), 3801 (NY, US);
Wright 1637 (F,G). Hartt: Buch 1857 (US), 1923 (US); Cook, Scofield & Dovyle 26
(US) ; Ekman H9420 (US), H10594 (US), H1875 (US) ; Holdridge 1384 (US) ; Leonard
7591 (US), 8069 (US), 8387 (G, NY, US), 9545 (US), 13242 (NY, US), 13430 (NY,
US), 13488 (A, US), 13489 14845 (US), 14867 (US), 15017 (US); Nash 722
(NY), 822 (NY), 840 (NY); Nash & Taylor 1415 (NY, US). Dommunican REPUBLIC:
Ekman H11079 (US), pee (US), H13846 (US), H15405 (US); Fuertes 362 (A,
G, NY, US), 577 (A), 1831 (A, NY); Valeur 459 (A, US), 804 (A, NY, US). Puerto
Rico: Britton 7836 (NY); Britton & Hess 2559 (NY, US); Gerhart 321 (NY);
Gregory 388 (PR); Sintenis 183 (G, NY, US).
There is considerable variation within this species in size and shape of
Ad JOURNAL OF THE ARNOLD ARBORETUM (VOL. XXVI
the fruit, length of the pedicels, and distribution of the leaves, and at first
I thought that several species, bolstered by geography, might be recognized
on these bases. It now seems plain, however, that these characters are
unstable, and that only one species is involved. It is possible that the
plants from Hispaniola should be distinguished intraspecifically, from those
of Cuba, on the basis of the usually smaller and less firm leaves, which
tend to be well distributed along the twigs, instead of clustered at the ends.
The Puerto Rican plants, however, have the leaves clustered at the ends
of the twigs, like those from Cuba, while retaining the texture of those
from Hispaniola.
12. Dipholis repens Urb. & Ekm. Ark. Bot. 22A(17): 70. 1929.
Creeping or prostrate shrub; leaves obovate or oblong-obovate, 1.5—2
cm. long, 6-11 mm. wide, or the upper smaller, acute at the base, rounded
at the apex, fer wee except when very young; petioles 2-3 mm. long;
flowers | or 2 in the axils, the pedicels 1-3 mm. long; sepals 5, the 2 outer
glabrous, 1.5 mm. long, the 3 inner appressed- -hairy, 2 mm. long; corolla
white, fragrant, 4 mm. long, the lobes surpassing the tube; filaments
scarcely 1 mm. long; style 1 mm. long; fruit brown- -purple, obovate-
oblong or ellipsoid, 5-6 mm. long, 3.5—4 mm. thick, l-seeded. ( Descrip-
tion abstracted from the original; no specimens seen.)
LLECTIO®: Ekman H6790, “Cordillera de Bahoruco in Sierra de los
Comisarios in pinetis ultra 1700 m. alt.,”’ August, Barahona, Sea Republic (US).
DistriBpuTION: Haiti and the Dominican Republic.
This species is obviously related to D. cubensis, but differs in its creeping
habit and smaller leaves.
13. Dipholis sericea sp. nov.
Planta lignosa; foliis firmiter chartaceis oblanceolatis vel anguste ellip-
ticis, circiter 4—9 cm. longis et 1.5—3 cm. latis, apice rotundatis vel abrupte
et breviter acuminatis, subtus primo rufo-sericeis demum glabratis, nervis
lateralibus primariis et minus secundariis subtus elevatis perspicuisque,
rete venularum laxo; petiolis circiter 3—8 mm. longis; floribus in axillis 1-4,
pedicellis crassis ferrugineis circiter 2-4 mm. longis; sepalis plerumque
6 interdum 4, subtiliter et interdum sparse ferrugineis, late et valde imbri-
catis, circiter 3. 5—3.7 mm. longis vel extimo plerumque manifeste breviore;
corolla (in alabastro tantum visa) cum appendicibus fimbriatis stami-
nodiisque cito ae ovario glabro in stylum 2 mm. longum abrupte con-
tracto; fructu igno
Type: Ekman . Cordillera Central, prov. Santo Domingo, Villa Altagracia,
top of Loma Marian Chick, mossy thicket, about 825 m., Dominican Republic, Janu-
ary 6, 1930 (US).
DistrRiBUTION: Known only from the type collection, Dominican Republic.
Although mature flowers of D. ferruginea are not yet available, maturing
buds indicate that its flowers are significantly smaller than those of
D. sericea. The leaves of the two species differ conspicuously in size,
shape, and venation.
14. Dipholis ferruginea Ekm. & Schmidt, Rep. Sp. Nov. 32: 94. 1933.
Large tree; leaves firm, narrowly elliptic, oblong, rounded at the apex,
1945 ] CRONQUIST, DIPHOLIS AND BUMELIA 445
2-4 cm. long, 1-1.5 cm. wide, glabrous, shining above, densely sericeous
ee the hairs at first strongly rufous, later pale; petioles about 2-5
m. long: flowers mostly 1 or 2 in the axils, the pedicels less than 6 mm.
ie densely rufous-hairy.
TYPE COLLECTION: Ekman H15406, “Cordillera Central, prov. de Samana, Los
Haitises, Boca del Liga ” limestone crags, light forest, characteristic, om 24, 1930,
Dominican Republic (U
DISTRIBUTION: ae. bee from the type collection, Dominican Republic, but
reported by the collector to be very common
BUMELIA
The genus Bumelia was described in 1788 by Swartz, in his Prodr. Veg.
Ind. Occ., p. 49. Of the seven species there included, four have been
removed to Dipholis and one to Mastichodendron, leaving only B. rotundi-
folia and B. retusa. Bumelia retusa, being the more common and wide-
spread of the two, may well stand as the nomenclatural type of the genus.
Bumelia is antedated 11 years by Robertia Scop., but it has been conserved
by an international botanical congress. Otto Kuntze rejected Bumelia in
favor of Lycioides L. (1738), but, since he gave neither descriptions nor
references to descriptions for his numerous transfers (other than L.
Spinosum), his names are not validly published and are here ignored.
Rafinesque proposed for B. tenax a new genus Sclerocladus, which he later
modified to Sclerozus, but this has universally and correctly been reduced
to synonymy.
In 1923 Lecomte described a new species, Bumelia Harmandii, from
Indo-China (Bull. Mus. Nat. Hist. Par. 29: 179). Its floral and vegetative
characters would seem from the description to be wholly consonant with
Bumelia, although the leaves are somewhat larger than those of any other
known species of the genus. In the absence of fruits, its true generic
position is uncertain. The genus Bumelia is otherwise strictly American.
Bumelia is related to and evidently derived from Dipholis, from which
it differs primarily in the absence of endosperm. The differences between
the two genera may be tabulated as follows:
BuMELIA
Seeds without endosperm; cotyledons
fleshy
Ovary usually hairy, occasionally gla-
rous.
Plant commonly spiny or thorny.
Young and mature fruit commonly
broadly paper to subtruncate or
even retu the apex
Lateral nee “of He earalla lobes some-
times wanting.
bm edema on the continent as well
n the West Indie
DIPHOLIS
Seeds with endosperm; cotyledons thin.
Ovary nearly always glabrous.
Plant unarmed.
Young and mature fruit amlaead
abruptly tapering to the style
Lateral lobes of the corolla-lobes al-
ways present.
Principal development in the Greater
tilles.
Ae to Record ee. Woods 59: 33. 1939) there is also some difference in the
wood-anatomy
Although Bumelia and Dipholis have almost universally been considered
446 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
nevis they were united by Baehni in his survey of the family (Candollea
: 394-508. 1938). In that and subsequent works Baehni denies the
Kane aiid of endosperm, pointing out that there are cases where the
character is known to fail. All other characters in the family also fail on
occasion, however, including the type of seed-scar, on which Baehni founds
his subfamilies. The fact is that students of the group have repeatedly
noted that the presence or absence of endosperm runs parallel to natural
groupings which are otherwise discernible, and this is also the case with
Dipholis and Bumelia. It is not ardlinarily necessary to dissect a mature
seed to distinguish between the two genera. Baehni’s objection to the
endosperm character seems also to be founded on the practical difficulties
in determining its presence in herbarium specimens, a view with which I
can sympathize but not agree.
Bumelia has a considerable number of widespread and highly variable
species, and specific lines must frequently be drawn rather broadly. Even
then constant tangible characters are all too few, although I have seen very
little evidence of hybridization. Length of the pedicels has frequently
been used as a specific character, but, except in certain cases of subsessile
versus evidently pedicellate flowers, it appears to be of very little value.
There is a great deal of variation in pubescence, and many species have
been founded on pubescence characters. It is true that the kind and color
of the pubescence are very helpful in delimiting the entities, but there is
a strong tendency for the hairs to fade and fall off progressively with
advancing maturity, so that the young leaves and twigs may be very
different in appearance from more mature ones.
The species which occur in the United States have recently been revised
by Clark (Ann. Mo. Bot. Gard. 29: 155-182. 1942), who recognized 14
species and several additional varieties. My observations as to the con-
stancy of certain characters, such as the variegation of the seeds in several
species, the glabrous ovary in B. reclinata, and the entire versus erose corolla-
lobes in “B. texana’”’ and related entities, are quite contrary to Clark’s. I
am able to recognize only 5 species in the United States; Asa Gray’s treat-
ment in the second edition of the Synoptical Flora is reasonably adequate
to distinguish these. Not a single valid species of Bumelia seems to have
been described from the United States since the appearance of Michaux’s
flora in 1803. Since Clark cites numerous specimens, and his entities
are in general readily equated with mine, I am not citing specimens for the
5 species concerned. In general, all other specimens from outside the
United States are cited, except those which belong to widespread species
and bear incomplete data
The most primitive surviving species of the genus is B. persimilis. The
general similarity of this species to Dipholis salicifolia has several times
been remarked, and it seems not to be accidental. Bumelia persimilis may
very well be only slightly removed from the prototype of the genus.
Evolutionary trends are toward reduction in size of the whole plant, in size
of the leaves and prominence of their veiny reticulum, in size of fruits, and
1945] CRONQUIST, DIPHOLIS AND BUMELIA 447
sometimes in the number of flowers in a cluster. I have found the con-
struction of a satisfactory key more than ordinarily difficult. The one
here presented follows the natural lines as nearly as possible, at the expense,
in some cases, of serviceability in identification.
Bumelia Sw.
Shrubs or trees, commonly but not always spiny; leaves alternate or
casually opposite, exstipulate, often small; primary lateral veins not very
numerous, sometimes obscure; flowers mostly 5-merous throughout, some-
sree ae hasten young and mature fruit generally broadly rounded
or subtruncate to even retuse at the apex; fruit fleshy, mostly 1-seeded, not
over soot 2.7 eM. rote seed-scar small, nearly basal; endosperm wanting.
KEY TO THE SPECIES
1. Corolla-lobes with lateral appendages (except rarely in South American B. obtusi-
olia
2. Fruit large, about 1.5-2.5 cm. long; Mexico to Venezuela.
3. Fruit smooth, not gall- like ; pubescence various.
4, Leaves ae the larger mostly 5-12 cm. long; primary lateral veins about
10-30 pai
5: Peticles aay 2-10 mm. long; style 3-7 mm. long; leaves variable in
Le ai beneath; sepals hairy or sometimes glabrous; Mexico to
MENEZ 16] avcii jac. sr ee ei eer mere ese otee sg on aeons 13.3: persons:
Si. Petioles iy 10-20 mm. long; style mostly 2-3 (rarely 4) mm. lon
leaves finely sericeous-strigose with tardily deciduous gray- white oe
b ; sepals glabrous; Mexico only................05. 2. B. laetevirens.
4, Leaves smaller, about 2—4.5 cm. long; primary lateral veins about 4—10 pairs;
@ueretaro; WexicG., Wome ieiteritns te oils soc 3. B. Altamiranot.
3. Fruit _ like, covered with short densely rufous-tomentose processes; twigs
and low leaf-surfaces finely velutinous-tomentose, at least when you
ao
o
~
pan
eae
ng;
Oaxa ( cis o's eden we acae a ln aren . B. revoluta.
1. Bumelia persimilis Hemsl. Biol. Centr. Am. Bot. 2: 298. 1882.
Heavy-trunked spiny tree or large shrub, about 5-18 m. tall; leaves firm,
ovate-oblong to elliptic or broadly elliptic-lanceolate, obtuse to more com-
monly acutish or acuminate, the larger about 5-12 cm. long and 2-5 c
wide, from conspicuously velvety or spreading-villous with rufous hairs .
finely strigose with rufous or gray hairs beneath when young, sooner or
later generally more or less glabrate; primary lateral veins about 10-30
pairs, more or less evident, but not always sharply distinct from the larger
secondary ones, the final reticulum evident or obscure, but not raised and
conspicuous; petioles mostly 3-10 mm. long; flowers several or numerous
in — clusters, from subsessile to sometimes borne on pedicels sae
c ong; sepals densely rufous-hairy to glabrous, about 1.8-3.7 m
long; “corolla about 3-6 mm. long, the tube about 1.2—1.8 mm. ee
anthers aaa or slightly exserted, 0.8-2 mm. long; staminodes about
1.7-3.8 mm. long, lanceolate to obovate, more or less laciniate; ovary short-
pilose, esecui near the base, to sometimes glabrous; style 3-7 mm. long;
fruit oblong or ellipsoid to spheroidal, about 1.5—2.5 cm. long, reputedly
sweet and edible
Type COLLECTION: Botteri 989, Orizaba, Vera Cruz, Mexico (F — fragment, G, US).
DistRIBUTION: Chihuahua, Mexico, to Federal District, Venezuela.
1A. Bumelia persimilis subsp. typica nom. nov.
Bumelia persimilis Hemsl. Biol. Centr. Am. Bot. 2: 298. 1882, sens. strict.
Bumelia leiogyna Donnell Smith, Bot. Gaz. 18: 4. 1893
Bumelia pleistochasia Donnell Smith, loc. cit.
Bumelia megaphylla Blake, Contr. Gray Herb. n. ser. 52: 76. 1917.
Bumelia guatemalensis Standl. Trop. Woods 4:9. 192
Bumelia panamensis Standl., loc. cit.
Bumelia barba-tigris Pittier, Man. Pl. Us. Venez. 125. 1926.
Bumelia Lankesteri Standl. Trop. Woods. 31:40. 193
Bumelia Austin-Smithii Standl. Field Mus. Publ. Bot. 18: 905. 1938.
Bumelia elexochitlensis Schultes & Conzatti, Leafl. Bot. Mus. Harv. Univ. 9: 190.
1941
Pubescence of the young twigs fine, appressed or a little loose, reddish or
often gray, often sparse, that of the lower surfaces of the leaves fine, strictly
appressed, and commonly soon deciduous, reddish or gray; corolla and
stvle sometimes fully as long as in subsp. subsessiliflora, sometimes only
450 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
about 3 mm. long; sepals and seed sometimes glabrous; fruit commonly
spheroidal, sometimes more elon
LocaL NAME Corpus espina, pe espino blanco, espino de crujo, ispundio,
limoncello, miorespino, yoni tree, tempisquito, zapotillo de pena, zapotillo bravo.
DistriBuTION: Vera Cruz and Oaxaca, Mexico, to Federal District, Venezuela.
Mexico: Haenke 1139 (F); Haenke 1596 (F); Haenke 1600 (F, NY); Haenke
1601 (F, NY). Vera Cruz: Purpus 13071 (A, F, Mo, NY). Guatemata: E
Petén: Egler 42-214 (F). San Marcos: Steyermark 37199 (A, F). Quezal-
tenango: Steyermark 34370 (F). Guatemala: Aguilar 193 (F). Sacatepé-
utiapa: Standley 75645 (F); Sicuidies 76223 (F); para she (F); Steyer-
mark 31791 (F). British Honpuras: Peck 756 (G); Schipp 1077 (A, F, G, Mich,
Mo, NY); Schipp 1339 (A, F, G, Mich, Mo, NY). Ex SAtvapor: Calieréa 294 (NY,
US); Calderén 313 (G, Mo, NY, US); Calderén 1554 (US). Costa Rica: Brenes
3874 (CR, F); Lankester 1194 (F); Lankester 1258 (F). Guanacaste: Standley &
Valerio 45491 (US); Standley & Valerio 46426 (US). Alajuela: Austin Smith
3 (F); Austin Smith H229 (F); Austin Smith H610 (F, US); Austin Smith 2685
(US). eet ly Oersted 315 (US); Standley 35881 (US). Panama: Panama:
Kluge 12 (F, NY, US). Corompia: Atlantico: Dugand 508-B (US); Elias 486
(NY, US) ; eee 698 (NY, US); Elias 1170 (A, NY, US). Bolivar: Killip & Smith
14487 (A, G, NY, US). Venezuera: Tamayo 1155 (US). Zulia: Tejera 57 (G,
US). epee District: Pittier 5856 (NY); Pittier 9163 (G, NY, US
1B. Bumelia persimilis subsp. subsessiliflora (Hemsl.) comb. nov.
Bumelia subsessiliflora Hemsl. Biol. Centr. Am. Bot. 2: 299. 1882.
Bumelia stenosperma Standl. Contr. U. S. Nat. Herb. 23: 1117. 1924.
Bumelia Lesueurti Standl. Field Mus. Publ. Bot. 22: 365. 1940.
Pubescence of the young twigs dense, rufous, spreading, often coarsely
velvety, that of the lower surfaces of the leaves similar but often longer,
looser, and less reddish, and commonly becoming appressed before even-
tually falling; corolla and style seldom under 4.5 mm. long; sepals and
ovary always more or less hairy; fruit commonly oblong or ellipsoid.
TYPE COLLECTION: Galeotti 7198, Guadalajara, Jalisco, Mexico, 3000 feet (F — frag-
ment).
LocaL NAMES: Bebelama, cupia, cupilla.
rics aamag Chihuahua and Durango to Michoacan and Oaxaca, Mexico.
eExico: Chihuahua: Gentry 2451 (F); Hewitt sn. (A); LeSueur 1160 (F);
Pon 1171 (G); Lumholtz 528 (G); Zing A13 (F). Sinaloa: Ortega 223 (G,
US); Ortega 4241 (US); Rose, Standley & Russell 13893 (G, Mo). Durango:
Gentry 5261 (G, Mo, NY). Colima: Palmer 1123 (A, G, Mich, Mo, NY, US).
Michoacan: Leavenworth & Hoowsriat 1393 (Mo). Oaxaca: Nelson 2548 (F,
US).
Bumelia persimilis is one of the several species in the genus with very
wide ranges. It is characterized by its large leaves, broadest at or below the
middle, with relatively numerous lateral veins, and without a conspicuous
raised reticulum, by its large fruit, and by its relatively long style.
Herbarium specimens do not indicate an unusual amount of intraspecific
variability, except in pubescence, on which two subspecies may be founded.
ome specimens from Guatemala, which have mostly been passing as
B. pleistochasia, have a slightly different aspect than most others of the
species and tend to have smaller flowers. Field study may show that
1945] CRONQUIST, DIPHOLIS AND BUMELIA 451
varietal recognition is warranted, but from the herbarium material it seems
yet quite unjustified. Bumelia Austin-Smithii Standl., from Costa Rica,
was described by the collector as having a pronounced nipple on the fruit,
but it does not seem otherwise different. Careful field investigation may
show the need for varietal recognition. Bumelia stenosperma Standl.,
described as having leaves only 1.5—3 cm. long, is merely subsp. subsessili-
flora with the leaves young and not yet fully expanded. It should be noted
that specimens of subsp. subsessiliflora with immature foliage have been
collected in August, as well as in the spring, suggesting that some factor
other than temperature, perhaps rainfall, has a controlling influence.
2. Bumelia laetevirens Hemsl. Biol. Centr. pea rig 2: 298. 1882.
Bumelia mexicana Engl. Bot. Jahrb. 12: 519.
Bumelia Palmeri Rose, Gard. & For. 7: ne pan
Achras olivacea Sesse & Moc. Fl. Mex. 91. 1894.
Spreading heavy-trunked tree much like B. persimilis, up to 15 m. tall,
the herbarium specimens, at least, nearly always unarmed; leaves elliptic
or ovate, broadly rounded to obtuse or acutish at the apex, firm, smooth
and shining above, finely strigose or sericeous-strigose with tardily decid-
uous white hairs beneath, the larger commonly 6—12 cm. long and 2.5-6
cm. wide; venation much like B. persimilis, the secondary lateral veins
sometimes a little more raised above; petioles about 10-20 mm. long;
flowers numerous in axillary Seay the pedicels commonly 2—8 mm. long;
sepals glabrous, about 2—3.5 mm. long; corolla about 3-6 mm. long, the
tube about 0.8-1.8 mm. long; anthers about 1—1.9 mm. long, slightly
faa ovary sparsely are especially below; ae mostly 2-3 (rarely
4) mm. long; fruit black, subglobose, mostly s m. long, edible.
TYPE COLLECTION: Galeotti i Cordillera of Oaxaca, pra at 3000 feet.
Loca NAMES: Bebalama, cupia, tempesquistle.
DistRIBUTION: Sinaloa to Tamaulipas, south to Oaxaca and Vera Cruz, Mexico.
Mexico: Liebmann 311 (US); Liebmann 312 (US); Sesse, Mocino et al. 5151 (F).
Sinaloa: Collins & Kempton 69 (US) ; Gentry 5494 (F, G) ; Ortega 795 (F) ; Ortega
4174 (US); Ortega 4503 (US); Ortega 4532 (US); Ortega 5647 (US); Ortega 5686
(US) ; Ortega 5847 (G, US); Ortega 6703 (F, Mo, US); Palmer 1513 (G, NY, US);
Rose, Standley & Russell 13897 (G, Mo). Durango: Goldman 334 (F, G, US).
ipas: LeSueur 644 (F); LeSueur 648 (F); Palmer 212 (G, Mo, NY, US).
San Luis Potosi: Palmer 48 (G, NY, US). Tepic: Maltby 99 (US). Hidalgo:
Chase 7441 (F, G, Mo, NY). Vera Crs Botteri 1014 (G, US); Palmer 364 (G,
Mo, NY, US). Mexico: ne 1229 (A, Mich, Mo). Puebla: Pringle 13872
(G, Mich, US); Purpus 5849 (G , NY, US); Rojans 4 (US); Rose et al. 10140
(NY, US). Oaxaca: Conzatti De (F); Conzatti 3459 (US) ; Conzatti 4612 (US);
Conzatti 4618 (US).
This species is obviously related to B. persimilis, and none of the charac-
ters separating it from that species, sens. amplior., shows real discontinuity.
I have seen no evidence of hybridization, however, and am convinced that
the two are properly considered distinct. It should be noted that B. per-
similis subsp. typica, which sometimes approaches B. laetevirens in pubes-
cense, does not occur through most of the range of the latter.
3. Bumelia Altamiranoi Rose & Standl. Contr. U. S. Nat. Herb. 23: 1117. 1924.
Spiny tree or shrub; young twigs loosely tomentose with gray-white or
452 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXVI
at first tawny-rufous hairs; leaves ovate to elliptic-oblong, broadly rounded
to acutish at the apex, 2.5—4.5 cm. long, 1.5—3 cm. wide, woolly-villous with
persistent white hairs beneath, more finely hairy and eventually glabrate
above; primary lateral veins about 4-10 pairs; secondary lateral veins
forming a loose scarcely raised reticulum above; petioles about 3—6 mm.
long; flowers unknown; fruit about 1.5—2 cm. long, subglobose, the pedicel
mm. long, woolly-villous, the persistent sepals about 3-4 mm. long,
similarly white-hairy.
Type CoLLECTION: Rose, Painter & Russell 9725, near Cadereyta, Queretaro, Mexico,
August 21, 1905 (NY, US).
DistriputioN: Known only from the type locality, Queretaro, Mexico. Another
collection is Altamirano 1644 (US).
This species resembles forms of the more northern B, lanuginosa in
pubescence but it has smaller, less reticulate leaves, and larger fruit. It
may also be compared to B. persimilis, which has larger leaves with more
numerous lateral veins, and in which the pubescence, if spreading, is always
rufous.
4. Bumelia eriocarpa Greenman & Conzatti, Field Mus. Publ. Bot. 2: 334. 1912.
More or less thorny; twigs coarse, finely velutinous-tomentose, at least
when young; leaves narrowly elliptic, lance-elliptic, or elliptic-oblong,
rounded at the apex, the larger commonly 5—9 cm. long and 18-30 mm.
wide, smooth and shining above, densely and sometimes very finely velu-
tinous-tomentose beneath with gray or sometimes tawny, persistent or
tardily deciduous hairs; primary lateral veins about 5-12 pairs; secondary
veins few, forming a loose reticulum above and beneath; petioles about
5-10 mm. long; flowers in clusters axillary to leaves or leaf-scars, subsessile,
the pedicels less than 2 mm. long, tawny-tomentose ; sepals about 2.5—3.5
mm. long, hairy like the pedicels; corolla about 5—6 mm. long, the tube
1.7-2 mm. long; anthers 2 mm. long; ovary densely long-hairy; style 4 mm.
long; fruit 1.5—2 cm. long, ellipsoid-globose, gall-like in appearance, covered
with very short densely rufous-tomentose processes; seed (from a fruit 2
cm. long) 1.6 cm. long, its scar nearly orbicular, basilateral, 6.5 mm. long.
E COLLECTION: Conzatti 1586, Cerro San Antonio, 1700 m., October 28, 1906,
Oaxaca, Mexico (F, NY, US).
DISTRIBUTION: Oaxaca, Mexico.
. : . : Conzatti 1772, Cerro del Tule, 1700 m. (F, G) ; Conzatti 2028,
Cerro San Antonio, 1600 m. (F); Smith 159, Mts. of Jayacatlan, 4500 ft. (G).
Although evidently related to B. persimilis, B. Altamiranoi, and B.
lanuginosa, this species is readily distinguished by its gall-like fruit. Addi-
tional differences will be noted when it is compared with any one of the
related species.
5. Bumelia lanuginosa (Michx.) Pers. Syn. 1: 237. 1805.
Shrub or tree about 1—15 m. tall, more or less thorny; leaves oblanceolate
to sometimes obovate or elliptic, broadly rounded to sometimes acute at
the apex, mostly 2—10 cm. long and 0.5—3.5 cm. wide, loosely woolly-villous
with white to tawny or rufous hairs when young, soon glabrate above, per-
sistently hairy to sometimes soon glabrate beneath, reticulate-veiny on both
sides, sometimes fascicled; flowers more or less numerous in each cluster,
1945] CRONQUIST, DIPHOLIS AND BUMELIA 453
the hairy or subglabrous pedicels 2-15 mm. long; wee strongly hairy or
nearly glabrous, 1.5—3.2 mm. long; corolla about 3—4.7 mm. long, the tube
1.3-2 m m. long: ete deltoid-ovate, about 1.9—2.7 mm. long, nearly
par ee the corolla- lobes; anthers 1—1.5 mm. long; ovary pilose; stvle
1—1.5 mm. long; fruit obovoid to broadly ellipsoid or subglobose, commonly
purplish- “black. 7-12 mm. long.
Type COLLECTION: Michaux, Georgia.
DistriBuTION: Mostly in uplands, occasionally in bottoms; Florida to sevens and
Kansas, south to Texas, southern Arizona, and the northern ‘hee of states in Mex
Key to the Subspecies
1. Larger leaves mostly 5-10 cm. long (occasionally smaller in ssp. typica) ; often over
m. tall; Florida to Missouri, Kansas, Oklahoma, eastern and central Texas, and
Coahuila anid Nuevo Leon, Mexico.
2. Pubescence of the leaves persistently tawny or rufous; almost entirely east of the
IMINSSISSID Taw iwliy Glastas ayer sere ec ele ec sieh . « o.'e ccghas-e “ayn loon tore eee et tee eee aes ssp. tvpica.
2. Pubescence of the leaves gray or nearly white, occasionally tawny at ste before
the leaves are fully expanded; almost entirely west of the Mississippi River.......
ey eR Map eee een icone rst chaa arine URL's. ais. 5! Scag spe ee ane EE SS ssp. 0 oblongifolia.
1. Larger leaves mostly 2—5 cm. long; seldom over 4—5 m. tall; southwestern Oklahoma,
through a and western Texas, to southern Arizona and the northern tier of
Sta CES: Ute Ve rake cya oe eee Soro sania:a- fe a 1085. 1798.
Sideroxylon tenax L. Mant. 48
Chrysophyllum woh tatu pial ie 323. = 54.1768.
Sideroxylon sericeum Walt. Fl. Car. 100. 17
Sideroxylon chrysophylloides Michx. FI. hig im 1: 123. 1803.
Bumelia chrysophylloides aes Fl. Am. Sept. 1: 155. 1814.
Sclerocladus tenax Raf. Syl ae en en
Sclerosus tenax Raf. Aut. Bot.
Bumelia megacocca Small, Bull. 'N. Y. in Gard. 1: 441. 1900.
Bumelia lacuum Small, Man. S. E. Fl. 1034. 1933.
Branching thorny shrub or small tree about 1-5 m. tall; young twigs
puberulent to sericeous-tomentose with mostly rufous or tawny hairs;
leaves oblanceolate to spatulate or oun nearly elliptic, rounded at the
apex, mostly 1-8 cm. long and 0.4—3.5 cm. wide, glabrous and evidently
reticulate-veiny on the upper surface, enue sericeous or sericeous-tomen-
tose with mostly tawny or rufous hairs on the lower surface, the pubescence
often obscuring the veins; flowers about 10-30 in a cluster, the clusters
sometimes closely ny as pane the pedicels rufous- eas to occasionally sub-
glabrate, commonly 6-15 mm. long; sepals finely ‘une sericeous or -stri-
gose, about 1.5-3 mm. inte corolla about 3.1-4.2 mm. long, the lobes
broadly ovate, about 1.5-1.9 mm. long; ovary shortly pilose; style about
1-1.5 mm. long; fruit aca: to ellipsoid or subglobose, about 8-14 mm.
long; seed solitary, about 6-9 mm. long, very smooth and shining, light
brown, occasionally pester variegated,
Type COLLECTION: Dr. Alexander Garden, South Carolina, probably near Charleston.
DistriBuTION: Dry, often sandy soil; coastal plain, from South Carolina to Florida.
An uncommon but widely distributed form with the pubescence of the
leaves merely silvery may be known as Bumelia tenax f, anomala (Sarg. )
comb, nov. (Bumelia lanuginosa var. anomala Sarg. Jour. Arnold. Arb.
2: 168. 1921; Bumelia anomala Clark, Ann. Mo. Bot. Gard. 29: 169. 1942).
9. Bumelia obtusifolia Roem. & Schult. Syst. Veg. 4: 802. 1819.
Large, more or less spiny shrub or small tree, up to 15 m. tall; twigs
sericeous or sometimes loosely sericeous- tomentose with pale or sometimes
1945 | CRONQUIST, DIPHOLIS AND BUMELIA 457
rufous hairs when young, soon glabrate; leaves from oblanceolate to sub-
orbicular, broadly rounded at the apex, gradually or abruptly tapering at
the base, commonly 2—5 cm. long and 1-3 cm. wide, sometimes larger, as
much as 9 cm. long and 4.5 cm. wide, sericeous or strigose beneath with
gray or faintly rufous-tinted hairs when young, later glabrate, obscurely or
sometimes more evidently veiny; flowers 1 — numerous in axillary clusters,
subsessile or on pedicels up to 8 mm. long; sepals 1.3-3 mm. long, strigose
or saat with gray hairs; corolla about 2.4—5.4 mm. long, its tube 0.5—2
1. long; anthers 0.9—2 mm. long; staminodes from narrowly oblong and
erose to ovate and ae Bene sometimes hooded, about 1.5—3.2 mm. long;
style about 1.5-4.6 mm. long: fruit ellipsoid-cylindric to subglobose, about
—15 m ies
Type coLLection: Humboldt & Bonpland, “in ripa fluminis Amazonum, ad c
ate ‘Chinchipen. alt. 200 h. (Prov. Jaen de Bracamoros)’’, at the north end of ihe
present province of Cajamarca, Peru.
DistrRIBUTION: Southern Mexico to northern Argentina.
Locat NAMES: Has toch, chi cheh chehum, sinan-che, malermo, picurero, picuyu,
caimito, piquillin, ivira-nina
. Bumelia obtusifolia subsp. typica nom.
ge obtusifolia Roem. & Schult. Syst. Ver. rm 802. 1819, sens. strict.
Leaves about twice as long as wide, subelliptic to broadly oblanceolate
or narrowly obovate; flowers in axillary clusters of about 1—5, the pedicels
2-4 mm. long; corolla 4.7—5.2 mm. long; style about 4 mm. long;
staminodes ovate or oblong, subentire, more or less hooded.
DistriBUTION: Ecuador and northern Peru.
Ecu Manabi: Haught 3377, abundant near the sea, between Salango and
a ee (NY, US).
. Bumelia obtusifolia subsp. buxifolia (Roem. & Schult.) stat. nov.
te buxifolia Roem. & Schult. Syst. Veg. 4: ine 1819.
Bumelia Dunanti A. DC. in DC. Prodr. 8: 191.
Bumelia Cruegerii Griseb. Fl. Brit. W. Ind. 401. re
Bumelia obtusifolia var. buxifolia Miq. in Mas Fl. Bras. 7: 47. 1863.
Bumelia guatemalensis Standl. Trop. Woods 4 1925.
Bumelia mayana Standl. Trop. Woods 31: 41. 1932.
Leaves mostly 1—2 times as long as wide, broadly elliptic to obovate or
suborbicular; petioles generally conspicuously exceeding the pedicels;
S :
ong style about 1.5-4 mm. long; staminodes ovate, subentire, scarcely
not at all hooded.
— COLLECTION: Humboldt & Bonpland, Cumana, Sucre, Venezuela (NY—photo).
DISTRIBUTION: Tabasco, eas to Nicaragua; northern Colombia and Venezuela
to Trinidad, chiefly near ee
Mexico: Tabasco: Matuda 3034 (A, Mich). Campeche: Lundell 1277 (A,
F, G, Mich, Mo, NY, US); Stewart 130 (G). Yucatan: Gaumer 473 (A, F, Mich,
Mo, NY, US); Gaumer 1572 (F); Gaumer 1791 (A, F, G, Mo, NY, US); Gaumer
23238 (A, F, G, Mo, NY, US) ; Gaumer 23845 (F, G, Mo, NY,US):; Landell 7501 (A);
Schott 341 (F, US); Schott 341A (F); Schott 341B (F); Seler 4937 (G, US); Steere
1674 (Mich); Stewart 258 (G). Quintana Roo: Lundell 7717 (A); Steere 2399
458 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXvI
; ch). GuaTeMALA: El] Petén: Bartlett 12290 (A, F, G, Mich, NY, US);
Bartlett 12593 (A, NY, » Bartlett 12739 (A, Mich, NY, US); Lundell 2201 (Mich).
Baja Verapaz: Kellerman 6588 (F). Zacapa: Steyermark 29363 (F). BritisH
Honpuras: Bartlett pete (Mich). Honpuras: Comayagua: Edwards 559 (A,
F, US). Tegucigalpa: Dyer 268 (US). Nuicaracua: Wright sn. (G, US).
Matagalpa: Rothschuhe 463 (F—photo & fragment, G— ph — photo)
CotrompiA: Magdalena: Herbert Smith 2740 (A, G, pee Revboi Smith 2071 (A,
G, NY, US). Venezuera: Curran & Haman 810 (G, NY); Curran & Haman 910
NY, US). Carabobo: d’Heguert 861 (NY). Federal District: Pittier 7765
(G, US) ; Pittier 9206 (G, NY, US) ; Pittier 12432 (A, NY, US) ; Pittier 13386 (A, NY,
US); Pittier 13479 (US); Rose 21830 (US); Tamayo 1139 (US). Sucre: Bond,
Gillin & Brown 44 (NY); Broadway 107 (G, NY, US); Broadway 213 (G, NY, US);
Broadway 613 (G, NY, US); Broadway 651 (G, NY, US) ; Johnston 273 (G). Trint-
pap: Britton & eee) 2240 (G, NY, US); Britton & Broadway 2633 (G, NY, US);
Britton & Hazen 817 (G, NY, US), 1725 (G, NY, US); Britton et al. 2696 (NY, US);
Miner 8085 hee Hart 2196 (NY, US).
9C. Bumelia obtusifolia subsp. excelsa (A. DC.) stat. nov.
Bumelia sartorum Mart. Herb. FI. Bras. 233. 1837-40.
Bumelia rhamnoides maa Nov. Stirp. Bras. ear 64. 1843.
Bumelia excelsa A. DC. in DC. Prodr. 8: 192.
Bumelia obtustfolia var. aah Miq. in Mart. m4 a 7: 48. 1863.
Bumelia fragrans Ridley, Jour. Linn. Soc. 27: 43, 1890.
Leaves mostly 1.5—3.5 times as long as wide, oblanceolate to elliptic or
narrowly obovate; petioles scarcely if at all exceeding the pedicels; flowers
in loose clusters of about 1-10; corolla about 3.3-4.7 mm. long, its tube
only 0.6-1.5 mm, long; staminodes narrowly oblong to lance-ovate, erose,
not hooded
Type COLLECTION: Blanchet 2162, “in maritimis ca. Bahiam,” Brazil.
DistripuTION: Piauhy, Pernambuco, and Fernando do Noronha Island, Brazil,
south to Rio de Janeiro, and inland to Paraguay, northern Argentina, and southern
Bolivia.
RAZIL: Riedel 19 (US). Piauhy: Lwuetzelburg 1643 (NY). Fernando do
Noronha Island: Ridley, Lea & Ramage 97 (G). Pernambuco: Pickel 3386
(A, G). Bahia: Blanchet 2763 (NY). Rio de Janeiro: Glaziou 11159 (NY,
US), 18439 (A, NY); Riedel 543 (US). Paracuay: Fiebrig 5392 (US); Hassler 2153
(G), 7200 (A), 7250 (G), 11507 (A), 11811 (A, G, US), 12286 (A, G, US); Kuntze s.n.
(in 1892) (NY); Malme 1032 (G, NY, US). Arcentina: Jujuy: Evyerdam &
Beetle 22547 (G), 22548 (G) ; Lillo 10792 (G), 10838 (G). Salta: Eyerdam & Beetle
22896 (G), 22929 (G); Parodi 9219 (G); Rodriguez 54 (G); Rodriguez 1129 (NY);
West 6147 (G). Tucuman: Lillo 7208 (G); Venturi 1059 (G, US); Venturi 1530
(A, US); Venturi 7588 (US). Formosa: Jorgensen 2154 (G). Chaco: Jorgen-
sen 1954 (G, US); Venturi 9768 (A, G, US). Corrientes: Parodi 11922 (G).
Bouivia (Southern): Pflanz 693 (G).
The northern and southern phases of subsp. buxifolia show different
trends of variation, but many individuals are quite indistinguishable in the
herbarium. The more northern plants have a short style (1.5-3 mm.),
often have relatively large leaves (to 9 & 4.5 cm.), and tend to have
narrowly ellipsoid fruit. The southern ones have a longer style (to 4 mm.),
seldom have the leaves over 5 cm. long, tend to have broader often subglo-
bose fruit, and sometimes have the lateral lobes of the corolla-lobes
conspicuously reduced or even obsolete. In spite of the apparent geographi-
1945] CRONQUIST, DIPHOLIS AND BUMELIA 459
cal disjunction of more than 600 miles, taxonomic segregation seems unwise
until more stable distinguishing ie are found.
Subspecies excelsa differs from ssp. buxifolia in an imposing array of
features, none of which is quite constant. In addition to the characters
given in the description, it frequently has the leaves more persistently
pubescent beneath than does ssp. buxifolia, with hairs that may be slightly
rufous-tinted, and tends to have a more nearly rotate, less campanulate
corolla.
I have not seen the type of B. obtusifolia, but from the description and
locality I think it should be associated with a single collection from Ecuador,
which has narrow leaves and few flowers, like ssp. exce/sa, but short pedicels
and ovate entire staminodia like ssp. buxifolia. The Ecuadorean specimen
also differs from both ssp. buxifolia and ssp. excelsa in having the staminodia
more or less hooded. The flowers of B. obtusifolia were originally described
as white, but, although collectors’ notes for ssp. buxifolia and ssp. excelsa
indicate white or greenish flowers, it is uncertain whether the color was
noted by Humboldt in the field, or merely taken from the dried specimen
by Kunth. The Ecuadorean specimen was noted by the collector to have
yellow flowers; it is not known whether or not this color difference is
significant.
10. Bumelia rotundifolia Sw. Prodr. Veg. Ind. Occ. 50. 1788.
Achras rotundifolia Poir. in Lam. Encyc. Meth. 6: 534. 1804.
Sumelia Purdiaei Urb. Symb. Ant. 5:,143. 1904.
Bumelia clarendonensis Urb. Rep. Sp. Nov. 13: 470. 1915.
Bumelia clarendonensis Urb. Rep. Sp. Nov. 21: 67. i not 1915.
Bumelia peckhamensis Urb. Rep. Sp. Nov. 22: 93. 1925
Small unarmed tree about 4—9 m. tall, essentially glabrous from the first,
or the young twigs occasionally with some appressed evanescent white
hairs; leaves firm, suborbicular or broadly elliptic to occasionally obovate,
about 1.5—7.5 cm. long and 1-5 cm. wide, alternate or opposite, evidently
veiny when young, less so with advancing age, often becoming very
obscurely so, borne on short petioles about 3-7 mm. long; flowers in
axillary clusters of about 3-12, the pedicels mostly 3—6 mm. long, glabrous;
sepals ‘glabrous, about 1—2.5 mm. long, firmly erect and becoming cartilagi-
nous at the base, restricting the lateral growth of the base of the fruit;
corolla about 3.2-3.9 mm. long; anthers 1.2—1.5 mm. long; staminodes about
.5-1.8 mm. long; style about 2—3 mm. long; mature fruit unknown, but
maturing fruit ellipsoid, about 7 mm. long.
TYPE COLLECTION: Swartz, Jamaica.
DisTRIBUTION: Jamaica.
Jamaica: Britton 2824 (NY), 3067 (NY); Britton & Hollick 1865 (NY), 2220
(NY); Harris 6169 (NY), 10035 (NY), 10165 (NY, US), 10386 (NY, US), 11040
(NY), 17111 (NY); Purdie sn. (G, NY).
This species is related to B. obtusifolia, from which it differs in being
essentially glabrous from the first, with the leaves becoming very firm, and
in its distribution. The only other species of Bumelia that occurs on
Jamaica is B. retusa, from which the present species likewise differs in being
glabrous, with broader leaves that rarely taper to the base. One specimen
460 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
(Walsingham s.n.— NY) seems intermediate between B. rotundifolia and
B. retusa, and may be a hybrid.
11. Bumelia retusa Sw. Prodr. Veg. Ind. Occ. 49. 1788.
Shrub or small tree about !—6 m. tall, ordinarily nearly or quite unarmed;
young twigs sericeous-strigose with rufous hairs which may later turn pale;
leaves alternate or gee narrowly to broadly obovate or occasionally
suborbicular, narrowed at the base, broadly rounded at the apex, mostly
1.5—5 (rarely 7) cm. ae and 7—40 mm. wide, rather obscurely or scarcely
axillary clusters of about 1-10, the pedicels 2—13 mm. long. sericeous-
strigose with rufous hairs that eventually fade; sepals rufous-strigose, about
1.5—3.3 mm. ees corolla about 3.3—5.5 mm. long, the tube 1-2 mm. long
anthers 0.8-1.7 mm. long; staminodes ovate or lanceolate to rotund, erose
or subentire, eae 1.3-2.8 mm. long; style about 0.7—-4.6 mm. long: fruit
black, broadly ellipsoid or subglobose, about 6—12 mm. long.
Type COLLECTION: None given; stated to come from Jamaica.
DistripuTION: Bahama Islands; Navassa Island; Jamaica; Santa Clara and Pinar
del Rio, Cuba; southern Vera Cruz to Yucatan and British Honduras. Chiefly or
entirely in coastal areas
11A. Bumelia retusa subsp. typica nom.
Bumelia retusa Sw. Prodr. Veg. Ind. Occ. 49, 1788, sens. strict.
Achras retusa Poir. in Lam. Encyc. 6: 533. 1804
Bumelia retusa var. ting ee biceagt in ‘Urb; Suns binge he 145. 1904.
Bumelia loranthifolia Britt. Bu 2 . Bot. Gard. 3: er
Bumelia oblongata —— Symb. a 6: 31. 1909.
Bumelia excisa Urb. Rep. Sp. Nov. 13: 471. 1915.
Bumelia Roigii Britt. & Small, Bull. Torrey Club 53: 461. 1926.
Bumelia navassana Urb. & Ekm. Ark. Bot. 22A(17): 71. 1929.
Flowers piguilied small, the corolla mostly 3.3-4.5 mm. long, the anthers
about 0.8—-1.3 mm., the staminodes about 1.3-2 mm. long, the style about
0.8-3.3 mm. ne Bahamas, Cuba, Navassa, and Jamaica.
BaHAMaAsS: Abaco: Brace 1565 (NY); Brace 1543 (NY). Acklin’s Island:
Brace 4366 (G, NY, US); Brace 4476 (F); Eggers 3924 (US). Andros: Brace 5034
(NY); Brace 5151 (NY); Brace 5264 (NY); Brace 5311 (NY); Brace 5323 (NY);
Northrop 544 (A, F); Small & Carter 8545 (G, NY, US); Small & Carter 8612
(G, NY, US). Anguilla Isles: Wilson 7955 (Mo, NY). Atwood Cay: Wilson
7396 (G, NY). Berry Islands: Britton & Millspaugh 2243 (NY). Caicos
Group: Millspaugh 9223 (G, NY). Cat Island: Britton & Millspaugh 5964
(F, NY); Wilson 7164 (G, NY). Crooked Island: Brace 4600 (NY, US); Brace
4698 (NY); Rothrock 246 (F). Eleuthera: Britton 6416 (NY); Britton &
Millspaugh 5412 (NY); Britton & Millspaugh 5426 (F, NY); Britton & Millspaugh
5433 (NY, F). Exuma Chain: Britton & Millspaugh 2780 (NY); Wilson 7894
NY). Great Bahama: Britton & Millspaugh 2565 (NY). Great Exuma:
Britton & Millspaugh sn. (NY). Great Ragged Island: Wilson 7818 (G, NY);
Wilson 7864 (G, NY). Inagua: Nash & Taylor 975 (NY); Nash & Taylor 1282
(NY); Nash & Taylor 1294 (NY). Little San Salvador: Britton & Millspaugh
56604 (F, NY). Long Cay: Brace 4071 (NY). Long Island: Britton & Mills-
paugh 6301 (F, NY). Mariguana: Wilson 7463 (G, NY). New Providence:
Britton 44 (NY); Britton 88 (NY); Britton & Brace 283 (G, NY, US); Britton
1945 | CRONQUIST, DIPHOLIS AND BUMELIA 461
& Brace 290 (NY, US); Britton & Brace 295 (NY, US); Britton & Brace 315 (NY);
Britton & Brace 351 (NY, US); Britton & Brace 538 (G, Mo, NY, US); Curtiss 85
G, Mo, NY, US); Wilson 8175 (NY); Wilson 8196 (NY); Wilson 8225 (NY);
Wilson 8349 (NY). North Bimini: Brace 3475 (NY) ae Island: Britton
& Milispaugh 2131 (G, NY, US). Rum Cay: Brace 6 (NY). St. George’s
Cay: Coker 315 (NY). Turk’s Islands: Millspaugh ae (G,NY). Watling’s
Island: Britton & ree 6168 (F, NY); Rothrock 295 (F, NY); Wilson 7240
(G, NY); Wilson 7261 (G, NY). Cusa: Pinar del Rio: Roig 3256 (Cu, NY, US);
Roig 3257 (NY). on ieee Acuna 10942 (Cu). Santa Clara: Ekman 18560
(A, F, G, Mo, NY, US). Navassa Istanp: Ekman 10811 (US). Jamaica: Britton
1153 (NY, US); Britton 1244 (NY); Britton 2544 (NY); Harris 9729 (NY, US);
Harris 9734 (NY, US); Harris 10380 (NY, U
Sig
11B. Bumelia retusa subsp. neglecta subsp. nov.
A subsp. typica differt floribus majoribus, corolla 4.3—5.5 mm. longa,
antheris ae mm. longis, staminodiis 2.5—2.8 mm. longis, stylo 3.3—4.6
mm. long
ay, ipp 585, open places at edge of mangroves, 5 ee elevation, All Pines,
British Tina August 20, 1930 (F); isotypes, A, G, Mich, Ye
DistriBuTION: Along the coast from southern Vera Cruz to Yucatan and British
Honduras
Mexico: Vera Cruz: Charles L. Smith 1123, Pei Saale Yucatan:
Gaumer 23210, ais (M o, US), 23338, Mina de Oro (A, F, G, ; NY, US);
Goldman 594, Progreso (US) ; Tuiel 7302, Ae (A): Schott oe Progreso
(F, Mo, US), 313a, Celestun (F). Quintana Roo: Gaumer 131, Cozumel Island
(G), s.n., in 1836, Mugeres Island (US).
A few specimens of B. retusa from the Bahama Islands have very spiny
twigs with reduced less than usually hairy leaves under 1 cm. long, thus
seemingly approaching B. glomerata. ‘They pass into the more typical
forms of the species, however, and in one case the collector noted that they
are young twigs from an otherwise apparently not unusual plant. Hy-
bridity seems out of the question, since the only other species of Bumelia
known to occur in the Bahamas is B. celestrina, which these plants do not
resemble.
Urban and Ekman differentiated B. navassana from B. retusa by its
opposite instead of alternate leaves, but it may be noted that the original
description of B. retusa called for opposite leaves. The leaves may in fact
be either alternate or opposite, even on the same plant. B. Roigii was
described as having a relatively large fruit 1.5 cm. in diameter, but the
fruit on the type may be abnormal, and those on the isotypes are smaller.
12. Bumelia socorrensis Brandegee, Zoe 5: 106. 1901.
Tree to 25 m. tall, apparently unarmed except for sometimes a few
short mie spines; twigs finely sericeous- strigose with rufous hairs when
young, soon glabrate; eee broadly oblanceolate, about 4—8 cm. long and
m. wide, tapering to the base, broadly rounded at the apex, finely
rufous- ee on both sides when young, later more or less glabrate
primary lateral ones about 8-15 pairs, not always sharply separable from
the larger secondary ones; petioles about 5-10 mm. long; flowers in
axillary clusters of about 1-5, appearing short and bulky, the pedicels
rufous-sericeous, about 1-4 mm. long; sepals more or less rufous-hairy.
462 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
about 2.3-3.5 mm. long; corolla about 4—5 mm. long, the tube thick, 1.5—2
mm. long, the lobes much thinner; staminodes about 2.5-3 mm long; fila-
ments coarse, the anthers only abo ut 1.1-1.4 mm. long; ovary slightly
5-lobed, brown- hairy, its style about 2.8-4 mm. long; fruit reputedly ellip-
soid, 12-14 mm. lon g and 8 mm. thick, dark blue.
Type COLLECTION: Anthony s.n., Socorro Island, Mexico (G).
DistriBUTION: Socorro Island.
Mex Socorro Island: Barkelew 190 (A, G, Mo, NY, US); Mason 1638
(A, G, US); Solis 82 (US).
This species, while sharply distinct, seems to be allied to B. persimilts
on the one hand and B. peninsularis on the other. It differs from the
former in its characteristically oblanceolate broadly rounded leaves with
fine appressed rufous pubescence, and in its somewhat smaller fruit. It
differs from B. peninsularis in its larger size, larger more oblanceolate leaves
with finer pubescence, and in its smaller flowers, particularly its smaller
anthers. B. socorrensis is of course eweunhiane remote from all others
of the genus.
13. Bumelia peninsularis Brandegee, Zoe 5: 107. 1901,
Shrub 3-4 m. tall, much branched from the base, provided with short
axillary spines; twigs pubescent with appressed rufous hairs when young,
later glabrate; leaves elliptic, elliptic-ovate, or elliptic-oblong, about 2.5—4.5
cm. long and 1—2.5 cm. wide, broadly rounded at the apex, firm, evidently
or obscurely reticulate, pubescent beneath, especially along the midrib,
with coarse, conspicuously malpighian, appressed rufous hairs, at least a
few of which commonly persist until maturity; petioles about 3-8 mm. long;
flowers about 2—6 in axillary clusters, the pedicels nearly or quite glabrous,
4-8 mm. long; sepals about 4.2—-5 mm. long, glabrous or nearly so, of the
long; staminodes about 3.5 mm. long; anthers about 2.5-2.7 mm. long,
slightly exserted; ovary short-hairy or reputedly glabrous, its style about
7 mm. aan ng; fruit broadly ellipsoid, about 15 mm. long.
TYPE : Bran wi sm., Sierra de la Laguna, Cape Region, Baja Cali-
rvs Mexico, March 1892 (G, US).
DistRIBUTION: Baja California, Mexico.
Mex Baja California: Brandegee s.n., in 1894 (US); Purpus 261, San
Felipe, Cape oe March, 1901 (Mo, US).
14. Bumelia cartilaginea sp. nov.
Frutex vel arbor parva usque ad 10 m. alta spinis brevibus axillaribus
praedita, ramulis juvenilibus subtiliter rufo-sericeo-strigosis cito glabratis;
hese firmiter chartaceis ellipticis vel anguste obovato-ellipticis, vulgo
5-4.5 cm. longis et 8-22 mm. latis, apice late rotundatis vel subacutis,
oe vel perspicue reticulatis, subtus praecipue secus costam pilis
adpressis rufis valde decidue strigosis (pilis paucis ad maturitatem persis-
tentibus); petiolis 2-5 mm. longis; floribus in glomerulis axillaribus 1-6,
pedicellis 2-5 mm. longis glabris vel glabratis; sepalis circiter 3—3.5 mm.
longis, glabris vel pilis paucis rufis ornatis, parte basali acute circumscripta
et cartilaginea, parte distali membranaceo- chartacea; corolla circiter 4.5—5
mm. longa, tubo 1.5—1.8 mm. longo, lobis suberectis; staminodiis 1.5—2 mm.
1945] CRONQUIST, DIPHOLIS AND BUMELIA 463
longis glabris eroso-laciniatis plus minusve cucullatis, inferne anion
superne complanatis plus minusve ieee antheris circiter
longis; ovario came stylo 3.7-5.2 . longo; fructu ignoto.
Type: Salazar 857, San Ignacio, an, Mexico, 460 ; 1919,
(US 1014209). What is evidently an a ie at US, bears the additional data
Montes and Salazar, Arroyo del Palmar La Cafia, shrub 3-4 m. high in moist shady
places
DistrRIBUTION: Sinaloa to Guerrero, Mexico.
Mexico: Sinaloa: Ortega 857, La Cafia, San Ignacio, in 1922 (F). Michoa-
errero: Langlassé 995, ee del Peregrino, palit nee 500 m.,
April 15, 1899 (G). Guerrero: Haenke 1594, Acapulco (F OE
Bumelia cartilaginea has been confused with B. pee which has
the flowers larger in all parts, with the sepals of the same texture throughout,
and the staminodia not hooded. It is in some respects transitional between
B. peninsularis and B. verruculosa, but it differs from the latter in its
glabrous and less strongly hooded staminodia, which are erose-laciniate
instead of entire-margined, in its suberect sepals and corolla-lobes, and in
its soon glabrate not at all verrucose twigs. The known ranges of these
three species are all quite distinct. An approach to the cartilaginous-based
type of sepal so conspicuous in B. cartilaginea may sometimes be seen in
B. persimilis and B. socorrensis. It should be pointed out that the bases
of the sepals do not become evidently cartilaginous until the flowers have
opened; maturing buds have the sepals apparently of the same texture
throughout.
15. Bumelia verruculosa sp. nov.
Frutex spinosus, ramulis primo rufo-tomentosis demum basibus papillatis
pilorum delapsorum verruculosis; foliis lanceolato-ellipticis vel elliptico-
costam pilis adpressis rufis strigosis mox glabratis; petiolis 2-5 mm. longis,
floribus in glomerulis axillaribus circiter 3—6, pedicellis glabris vel subglabris
3-6 mm. longis; sepalis circiter 2.7~3 mm. longis, glabris vel pilis pauc's
rufis ornatis, parte basali firma et cartilaginea, Pas ee mem
ceo- -chartacea reflexa; corolla circiter 4.8-5 mm. longa, t
longo, lobis reflexis; staminodiis cucullatis inflexis integris ‘villosis circiter
1.7—2 mm. longis; antheris 1.8—2 mm. longis; ovario glabro, stvlo 4—4
longo; fructu ignot
Type: Pringle 6984, Pedi hills, Las Palmas, near ara San Luis Potosi,
Mexico, March 7, 1899, 400 feet (US 1638968) ; isotypes at G, US:
DisTRIBUTION: Known only from the type collection, San om ies Mexico.
This species is superficially similar to B. peninsularis and B. cartilaginea,
but it differs strikingly in its hooded, villous, entire-margined, inflexe
staminodia, and in the persistent bases of the hairs of the twigs. It also
differs in its strongly reflexed sepals and corolla-lobes, and in its known
geographic distribution. Both the staminodes and the pubescence of the
twigs are quite unique in the genus.
16. Bumelia obovata (Lam.) A. DC. in DC. Prodr. 8: 191. 1844.
Shrub or small tree commonly 2-6 m. tall, unarmed or sometimes
404 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
strongly spiny; young twigs sericeous-strigose with rufous hairs, generally
soon glabrate; leaves oblanceolate to broadly ovate or suborbicular, broadly
rounded and sometimes obscurely wavy at the apex, cuneate at the base,
commonly 1—3.5 cm. long and 3—33 mm. wide, occasionally some of them
smaller, rufous-strigose beneath when young, sooner or later subglabrate,
the hairs often fading before falling; leaf-surfaces, especially the upper,
having a peculiar texture due largely to the fine slightly raised irregularly
reticulate striations trending parallel to the primary lateral veins, not other-
wise reticulate; flowers mostly about 2—10 in axillary clusters, from subsessile
to borne on pedicels as much as 10 mm. long, the pedicels rufous-strigose
to glabrous; vice glabrous or more or less strigose, about 1.2—2 mm. long;
sag about 2.7—4.3 mm. long, the so 0.7—1.3 mm. long; anthers 0.8—1.3
1. long; ovary hairy or subglabrous; style about 2—4.5 mm. long; fruit
aie to sometimes lala oe black, 5-12 mm. long.
Type COLLECTION: None giv
DistRIBUTION: Hispaniola il Puerto Rico through the Lesser Antilles to Curacao
and Aruba, also rarely in northern Venezuela.
16A. Bumelia obovata var. typica nom. nov.
Sideroxylon obovatum Lam. Tab. Encyc. 2: 42. 1793.
Bumelia cuneata Sw. FI. Ind. Occ. 1: 496. 1797.
Achras cuneifolia Poir. in Lam. Encyc. Meth. 6: 534. 1804.
Sideroxylon cuneatum A. DC. in DC. Prodr. 8: 181, 1844.
Bumelia parvifolia A. DC. in DC. Prodr. 8: 190. 1844.
Bumelia obovata A. DC. in DC. Prodr. 8: 191. 1844, sens. strict.
Bumelia myrsinifolia A. DC. in DC. Prodr. 8: 192. 1844.
Bumelia obovata var. portoricensis Pierre in Urb. Symb. Ant. 5: 143. 1904.
Bumelia obovata . thomensis ere loc. cit.
Bumelia cit ark Urb. Symb. . 7: 326. 1912.
Usually essentially unarmed, oon occasionally evidently spiny; pedicels
mostly 2-10 mm. long, sparsely hairy or glabrous; sepals and ovary sparsely
hairy or glabrous; bis ene as much as 4.5 mm. long, often much
less. Range of the sp
Haiti: Ekman a ), Nash & Taylor 1595 (NY, US). Dominican
Repusiic: Bertero sn. (Mo, NY — fragment); Ekman 14067 (US); Fairchild 2622
(A, US); Taylor 519 (NY). Puerto Rico: Britton 9106 (NY), 9648 (NY) ; Britton
& Brown 6025 (NY); Britton, Cowell & Brown 4821 (NY, US), 4879 (G, Mo, NY, US) ;
Britton, Cowell & Hess 1657 (NY, US); Britton & Shafer 1878 (NY, US); ag
306 (NY), 408 (PR); Hess 1668 (NY); Miller 1631 (US); Shafer 1998 (NY, US);
Sintenis 3400 (G, US), 3485 (Mo, NY, US), 3546 (US), 3780 (G, US), 4814b (US).
St. THomas: Britton 192 (NY); Britton & Shafer 18 (NY, US); Eggers 409 (G), s.n.
(Mo, NY, US) ; Holton s.n. (NY); Rose 3196 (NY). Sr. Jan: Britton & Shafer 526
(NY, US). Tortora: Fishlock 136 (NY), 446 (NY, US), 447 (F); Britton & Shafer
905 (NY, US). Virctn Gorpa: Beard 328 (A); Britton & Fishlock 1116 (NY, US).
AnecapA: Britton & Fishlock 960 (G, NY, US); Fishlock 10 (F, G, NY, US). Sr.
Croix: Ricksecker 383 (G, Mo, NY, US). St. Martin: Boldingh 2765 (NY).
St. BARTHOLOMEW: Forsstroem s.n. NY). BarBupa: Box 617 (US). AntTiGua:
Box 847 (US), 1401 (A, US), 1511 (A, US). GuapeLoupe: Bertero s.n. (Mo); Duss
2909 (NY). Martinique: Duss 260 (NY). Curacao: Boldingh 5312 (NY);
Britton & Sug! 3109 (NY, US); Curran & Haman 88 (G, NY), 158 (A, G, US), 205
(G, NY, US); Potter 5118 (NY); Realino s.n. (Krukoff Herb.). Arusa: Boldingh
6494 (NY); Curran & Haman 410 (G, US). Bownarre: Boldingh 7235 (NY), 7422
(NY). Vice Aragua: Ocumare de la prices Williams 12174 (US).
Sucre: Paria Peninsula, Bond, Gillin & Brown 262 (NY).
1945] CRONQUIST, DIPHOLIS AND BUMELIA 465
16B. Bumelia obovata var. Krugii (Pierre) comb. nov.
Bumelia Krugii Pierre in Urb. Symb. Ant. 5: 146. 1904.
Strongly spiny; more hairy than var. typica, the twigs, pedicels, and
sepals finely sericeous-strigose with rufous hairs; leaves averaging smaller
than in var. typica; flowers subsessile, the pedicels up to about 2 mm. long;
ovary eae short-hairy; style apparently not over 3 mm. long.
Puerto Rico.
Type COLLECTION: Not specified, from among Sintenis ies 3473, and 4813b, “in
Portorico prope Guanica in fruticetis searae ad Salinas
Puerto Rico: Britton 9115 (NY), 9327 (NY), 9996 (NY): Britton & Boynton
8295 (NY, be Britton & Cowell 1298 (NY); Britton, Cowell & Brown 4897 (US),
4905 (Mo, NY, US); Britton & Shafer 1837 (Mo, NY, US), 1908 (NY, a Gregory
650 (PR); ie 2940 (NY, US) ; Sintenis 3472 (Mo, NY), 3473 (G, US
Bumelia obovata may be distinguished from all others of ae genus by
the peculiar leaf-texture, a character which, although perhaps more readily
observed than described, is as nearly constant as any in the genus. It is
presumably related to B. obtusifolia. Except at the borders of its range,
in Hispaniola and Venezuela, no other species of Bumelia is known to occur
in its area.
The variety Krugii is well separated from the ordinary nearly or quite
unarmed forms of var. typica. However, several otherwise representative
specimens of the latter from Hispaniola, one from Puerto Rico, one from
Curacao, and one of the two known collections from Venezuela resemble
var. Krugii in being more or less spiny. Most of these specimens, it will be
noted, come from outside the known range of var. Krugzi. It is possible
that a third variety could be established for these plants, but in view of
their very spotty distribution I am reluctant to do so without further
evidence. It seems not improbable that the type of the species is of this
nature, since it was originally described by Lamarck as being spiny.
17. Bumelia glomerata Griseb. Mem. Am. Acad. II. 8: 518. 1862.
Bumelia horrida Griseb. Cat. Pl. Cub. 165. 1866.
ent microphylla Griseb. Cat. Pl. Cub. 165. 1866.
Bumelia tortuosa C. Wright ex Sauvalle, Ann. Acad. Habana 6: 288. 1870.
?Bumelia subintegra Urb. & Ekm. Ark. Bot. 21A(5): 56. 1927
Bumelia pachyclada Urb. & Ekm. Ark. Bot. 21A(5): 57. 1927
Bumelia lineolata Urb. & Ekm. hee Bot. 22A(17): 73. 1929.
More or less spiny shrub or small tree, sometimes less than 1 m. high,
sometimes reaching 12 m.; young twigs nely rufous-strigose; leaves
about 3-40 mm. long and 2-17 mm. wide, elliptic or broadly oblanceolate
(especially when larger) to broadly obovate or suborbicular (especially
when smaller), broadly rounded at the apex, thick and smooth, mostly
fascicled, strigose beneath with mostly rufous hairs when young, later
glabrate, the veins, except the midrib, generally visible only as faint fur-
rows or not at all; flowers about 1-4 in a cluster, ae the rufous-
hairy pedicels up to about 2 mm. long; sepals abou 1.1-2.2 m mm. long,
glabrous or rufous-strigose; corolla about 3.1-3.8 mm. long, the tube
1.2-1.5 mm. long; stamens included, the filaments only about 1 mm. long;
anthers about 0.7—1 mm. long; staminodes ovate to reniform, about 1-1.3
466 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
mm. long; ovary with flattish to broadly rounded glabrous top and short-
hairy to glabrous sides, the style 0.7—1.4 mm. long; fruit dark red, subglo-
bose, less than 1 cm. long.
TyPrE COLLECTION: Wright 347, eastern Cuba (G, NY).
DistriBUTION: Cuba and Haiti.
This species might conceivably be divided into 3 varieties, a typical one,
only slightly spiny, with relatively large leaves that have the midrib raised
and visible beneath nearly to the tip, one based on B. horrida, strongly
spiny, with smaller leaves in which the midrib visibly extends scarcely
beyond the middle, and an equally spiny but coarser variety based on
B. microphylla, with very small leaves that have scarcely discernible midrib
and tend to become coppery beneath. It is entirely possible, however, that
these represent mere individual responses to variations in the severity of the
habitat, and I am therefore unwilling to propose the new combinations that
would be necessary. The specimens have been annotated as ‘‘typical
phase,” ““B. horrida phase,” and “B. microphylla phrase,” and are so cited
below.
Typical phase: Cupa: Isla de Pinos: Britton, Britton & Wilson 15072
(NY, US). Pinardel Rio: Ekman 16577 (NY). Oriente: Ekman 6264 (NY),
7893 oo 9624 (A, F, US), 19184 (G, NY, US); Roig 4910 (Cu), 4992 (NY), 5185
Cu,
B. aoe phase: Cusa: Wright 1922 (NY), 2922 (G, Mo, NY, US). Isla de
Pinos: Britton, Britton & Wilson 15377 (NY, US); Ekman 12335 (US). Pinar
del Rio: Britton & Cowell 9924 (NY); Fors 4762 (Cu); Roig 3966 (Cu).
Habana: Ekman 13271 (NY); Ledén 6010 (NY), 6256 (NY), 7172 (NY), 7591
(NY); Leon & Roig 11445 (NY); Wilson 13933 (NY, US). Santa Clara: Britton,
Earle & Wilson 4595 (G, NY, US); Combs 734 (G, Mo, NY). Camaguey: Biman
15478 (US). Oriente: Britton & Cowell 12705 (NY, US); Britton, Cowell &
Shafer 13064 (NY, US); Ekman 2961 (US), 10204 (F). Harrr: Ewan H 10004
(US) ; Leonard 13267 (NY).
B. microphylla phase: Cusa: Wright 2922a (G, Mo, NY), 3623 (G). Habana:
Leon & Roig 2522 (Cu), 8150 (Cu). Matanzas: Ledn & Roig 4167 (Cu), 12950
(NY); Ekman 16499 (NY), 17181 (US), 18592 (US). Santa Clara: Ekman
18841 (G, NY). Oriente: Ekman 19103 (G, NY, US); Shafer 1234 (NY, US).
Bumelia subintegra Urb. & Ekm. was described as having the pubescence
of the twigs spreading, instead of appressed as in B. glomerata, and is only
doubtfully included here. I have not seen the type.
Bumelia glomerata seems to be related to B. obovata, the var. Krugii of
which tends toward it. The leaves of B. glomerata, while differing from
those of B. obovata in being thick, firm, smooth, and not at all veiny, as
well as smaller, frequently show a suggestion of the fine raised striations so
characteristic of the latter species. It is possible, however, that the rela-
tionship of B. glomerata is with B. retusa instead of with B. obovata, and
occasional forms of B. retusa with spiny twigs and very small leaves are
indeed difficult to distinguish from B. glomerata.
18. Bumelia occidentalis Hemsl. Biol. Centr. Am. Bot. 2: 298. 1881.
Bumelia fragrans Brandegee, Zoe 5: 106. 1901, not Ridley (1890).
Bumelia Brandegei Blake, Contr. Gray Herb. ns. 52: 76. 1917.
Bumelia cuneifolia Jones, Contr. West. Bot. 18: 63. 1933-35.
1945 ] CRONQUIST, DIPHOLIS AND BUMELIA 467
Branching spiny shrub or small tree commonly 2-5 m. tall; leaves
oblanceolate to obovate, elliptic-obovate, or rarely subrotund, tapering to
the base, broadly rounded at the apex, about 8-30 mm. long and 2—20 mm
wide, scarcely or obscurely veiny, finely gray- -puberulent especially beneath,
eventually more or less glabrate; flowers several in axillary clusters, the
pedicels about 4-20 mm. long, commonly appressed-puberulent like the
leaves; sepals about 2.3-3.5 mm. long, grayish- daira or strigose;
corolla about 4.5—5.2 mm. long, its tube 0.8—-1.5 mm. long; anthers about
1.3-2.6 mm. long; style about 4.5—-5.2 mm. long; fruit ellipsoid. -oblong,
blue-black, Ree 12-16 mm. long.
TYPE COLLECTION: Coulter 0) upper Sonora, Mexico.
Locat NAME: Bebelama.
DIstTRIBUTION: Baja California and Sonora, Mexico.
MeEx Baja California: Brandegee s.n. (April 9, 1889), San José de la
Gracia G US); Brandegee s.n. (in 1893), Pescadero (A); Brandegee s.n. (May 7,
1897), San José del Cabo (G, US) ; Gentry 4264, Los Encinos, Sierra Giganta (G, Mo);
(Cape Region) (Mo, US) ; Wiggins 5482, north of Comondu (A). Sonora: Coville
1676, Torres (US); Ferris 8729, ae Carlos Bay, north of Guaymas (NY, US) ; John-
ston 4296, San Pedro Bay (A, G, NY, US); Johnston 4367, San Carlos Bay (G, US);
MacDougal & Shreve 6, west of ae Well (US) ; Shreve 6054, near Santa Rosa (F) ;
Shreve 6195, north of Palma, between Guaymas and Hermosillo (US); White 416,
Tonibabi hot springs, near Moctezuma ‘are Mido 6167, between San Carlos and
Santa Rosa, 45 miles west of Norio (US); Wiggins 6482, > miles north of Guaymas
(Mich, US) ; Wiggins 8293, San Pedro, north of Tajitos (U
This species is related to B. celastrina, from aie it differs in its more
copious and more persistent pubescence, longer pedicels, larger flowers with
longer style and anthers, larger fruit, and more western disjunct distribu-
tion. Bumelia fragrans Brandegee ( — B. Brandegei Blake), with shorter,
less hairy pedicels, and broader sooner glabrate leaves than usual, is a
art of this species. Further collecting may conceivably warrant its
varietal recognition; the specimens now available do not
19. Bumelia celastrina H.B.K. Nov. Gen. et Sp. 7: 212. 1825.
B 831
: 49.
melia Eggersii Pierre in Urb. Symb. Ant. re ie 1904.
rain Schottii Britton, N. Am. Trees 777. 19
Bumelia affinis Blake, Contr. Gray Herb. ns. ay on 1918.
Shrub or small tree commonly 2-9 m. tall; leaves generally fascicled,
oOo
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obovate or sometimes nearly elliptic, broadly rounded at the apex, generally
acute at the base, mostly 1-4 cm. long and 3—25 mm. wide, occasionally a
little larger, firm, the veins not very prominent, the reticulum rather obscure
or indiscernible; flowers about 3-10 in a cluster, the pedicles glabrous or
with a few inconspicuous white hairs, about 2— 7 mm . long; corolla nee
3—4.5 mm. long, the short tube only 0. 8—1.2 mm. long: anthers 1-1.5 m
long; staminodes lance-elliptic to rhombic-ovate, about 2—3 mm. lon
ovary pilose toward the base, glabrous above, the style about 2.5-4 mm
468 JOURNAL OF THE ARNOLD ARBORETUM [ VOL. XXVI
long; fruit ellipsoid-cylindric, occasionally a little widened above, 7-13 mm.
long, blue-black
Tyree coLiection: Humboldt & Bonpland, ‘“crescit in declivitate occidentali
montium Mexicanorum, in convali Sopilote, inter Chilpansingo et Tasco, alt. 517 hex.”
Loca NAMES: Hapuche, caimito, coma, bebelamilla bagre.
DistrIBUTION: Southern Texas to Venezuela; Florida; Bahama Islands; Camaguey,
Mexico: Nuevo Leon: Dodge 96 (NY, US); Edwards 378 (F); Gregg 196
(G, Mo); Kenoyer 1173 (F); Lozano 13904 (G, Mich, US) ; Pringle 2787 (A, G, US) ;
Taylor 378 (Mo). Tamsulive as: Bartlett 10565 (F); Bartlett 10833 (F); Bartlett
10784 (F, US); Bartlett 11111 (F, G, US); Berlandier 3048 (Mo) ; LeSueur 340 (F, G).
Sinaloa: Ortega 6572 (US). Vera Cruz: Miller 1855 (NY); Seler 3711
(G, US); Woronow 2099 (F). Oaxaca: Reko 3677 (US); Williams 9907 (F).
Yucatan: Flores 6 (F); Gaumer 1313 (A, F, G, Mich, Mo, NY). Chiapas:
Bossé s.n. (F); Matuda 2702 (A, Mich, Mo, NY, US). Guatemara: San Marcos:
Steyermark 37776 (F). diet nege Standley 69035 (F); Standley 69042 (A, F);
Steyermark 43859 (F). Zacapa tandley 74144 (F); Standley 72004 (F). Retal-
uleu: Standley 66512 (F); aa 66643 (F); Standley 87545 (F). Chiquimula
Steyermark 31533 (F). Ev Sanvapor: Standley 20864 (G, NY, US); Standley 21893
(G, NY, US). Panama: Coclé: Pittier 4988 (G, NY, US). Coromsia: Bolivar:
Heriberto 95 (US). Magdalena: Herbert Smith 423 (A, G, NY, US). VENEZUELA:
Isla de San regina Curran & Haman 802 (G, NY, US). Falcon: Curran &
Haman 534 (A, G, NY, US). Banuama Istanps: Great Bahama: Brace 3605
(NY); Britton & Millspaugh 2737 (NY). New Providence: Britton 6592 (NY,
US); Britton & Brace 326 (NY); Eggers 4418 (A, US); Millspaugh 2492
Andros: Brace ie (NY); Brace 5291 (NY); Brace 6773 (NY); Coker 202 (NY);
Northrop 666 (F, NY). Hog Island: Wilson 8317 (Mo, NY). Rose Island:
Britton & Millspaugh 2 2157 (NY). Cat Cay: Brace 3740 (NY). South Cat Cay:
Millspaugh 2420 (NY). Cusa: Camaguey: Ekman 15476 (NY, G, US); Ekman
15523 (NY); Shafer 407 (G, NY, US); Shafer 886 (NY, US); Shafer 906 (NY, US).
The plants from Florida and the West Indies es the reticulum of the
leaf a little more obscure, on the average, than others of the species, and
are generally totally glabrous from the first, but the differences are minor
and not sufficiently constant to warrant taxonomic recognition, An imma-
ture fruiting specimen from Oriente, Cuba ( Roig 5545 — Cu) seems nearest
to B. celastrina, but has the young twigs evidently rufous-strigose.
20. Bumelia conferta “ Wright) aaa in Urb. Symb. Ant. . fg 1904.
Sideroxylon sconfert Wright in Sauvelle, Fl. Cub. 86
Unarmed tree, or ie young ae spiny; twigs seein when
5 cm and
2
-
glabrous from the first, not at all veiny, the primary lateral veins faint or
obscure at maturity; flowers about 1—4 in a cluster, the pedicels glabrous,
about 6-10 mm. long; sepals slightly strigose, becoming cartilaginous at
the base; corolla (not seen by me) about 4.5 mm. long, the lateral lobes
absent or vestigial; ovary short-hairy; style about 3—3.5 mm. long; mature
fruit unknown, but a maturing one RASEOWY ellipsoid, about 9 mm. long.
PE COLLECTION: Wright 2920, in part, “a ~ nape de los manglares Potrero
Playitas, Bahia-Hondia,” Pinar del Rio, Cuba (G, , US —drawing). An entirely
different plant, also bearing Wright’s number soe was made the type of Daphnopsis
cuneata Radlk. (Mo)
DistripuTion: Pinar del Rio and Oriente, Cuba.
1945 | CRONQUIST, DIPHOLIS AND BUMELIA 469
uBA: Wright 2928 (presumably in Pinar del Rio) (G, Mo). Pinar del Rio:
Ekman 17405 (US), 17435, Toscano (Cu, NY, US). Oriente: Sierra Maestra,
Ekman 9389 (A, F, G, NY, US), 14206 (NY); Ledn 11022 (NY).
Bumelia conferta is closely related to B. rotundifolia and, at least in the
herbarium, greatly resembles it, but it differs in the obsolescence of the
lateral lobes of the corolla-lobes, in the more evidently hairy twigs and
sepals, in having fewer flowers per cluster, and in having the leaves, on the
average, less veiny. The known distribution of B. conferta, Oriente and
Pinar del Rio, Cuba, is perplexing and unexplained, but the specimens
available from the two areas show no appreciable differences.
21. Bumelia Picardae Urb. Symb. Ant. 5: 148. 1904.
Spiny shrub about 2-3 m. tall; young twigs ere with silvery or
rufous hairs, very soon glabrate; leaves firm, oblanceolate or more com-
monly obovate to narrowly or broadly elliptic, abhi or broadly rounded
at the apex, more often narrowed at the base, about 6-30 mm. long and
4-15 mm. wide, silvery-strigose when young, very soon glabrate, the midrib
raised beneath, the primary lateral veins visible only as faint furrows,
especially beneath, or not at all, the upper surface smoother and shinier
than the lower, subsessile, the petioles less than 3 mm. long; flowers about
1—3 in the axils, subsessile, the pedicels less than 2 mm. long; sepals about
1.5-2.9 mm. long, glabrous, generally green, especially toward the base;
corolla about 3.2—-4 mm. long, the tube 1.3—1.5 mm. long, the lobes without
lateral lobes; anthers about 1—1.2 mm. long; staminodes ovate, about
7 mm. long; ovary short-hairy or subglabrous, the style about
1.2-1.9 mm. long; fruit narrowly ellipsoid, about 12-13 mm. long.
TYPE COLLECTION: Picarda 1242, “Haiti in Plaine,” June (NY).
DISTRIBUTION: Haiti and the Dominican Republic.
Haitt: Ekman H2021 (US); Leonard 10003 (G, NY, US), 14718 (A, G, US), 14899
(US), 15879 (NY, US). Dominican Repusiic: Ekman 15576 (US).
Urban established a separate section Bumeliopsis for B. Picardae, based
on the suppression of the lateral lobes of the corolla-lobes. Three other
species (and occasionally a fourth) are now known to share this character,
but it is doubtful that they are sufficiently closely related to form a natural
section. Bumelia Picardae seems related to B. obovata, B. conferta is
obviously close to B. rotundifolia, and B. revoluta and B. integra are of
doubtful affinities, perhaps allied to B. retusa. All of these species are
probably eventually derived from and not distinctly related to B. obtust-
folia, in which the lateral lobes are occasionally suppressed.
22. Bumelia integra nom. nov.
Dipholis anomala Urb. Symb. Ant. 7: 325. 1912. Not Bumelia anomala Clark.
Unarmed tree; leaves obovate or elliptic, obtuse or rounded at the apex,
tapering to the base, about 2.5—5 cm. long and 1.5—2.5 cm. wide, short-
petiolate, very finely and densely rufous-strigose on both sides at first,
very soon glabrate above, more tardily so beneath, not very veiny, only the
midrib and primary lateral veins evident beneath, only the midrib above;
flowers in dense axillary clusters of about 5—25, subsessile, the stout rufous-
strigose pedicels 2 mm. long or less; sepals nearly glabrous; corolla about
3.2 mm. long, white, the lobes double the tube, without lateral lobes;
470 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
anthers about 0.9 mm. long; ovary appressed-hairy; style about 2.5 mm.
long; fruit unknown.
Type COLLECTION: Fuertes 1039, “Barahona, ad El Hoyo,’ Dominican Republic,
700 m., Sept., 1911 (A, G, NY, US).
DistrIBUTION: Known only from the type collection, Dominican Republic.
This species is transferred from Dipholis to Bumelia because of its hairy
ovary and because suppression of the lateral lobes of the corolla-lobes, while
occurring in several species of Bumelia, is otherwise unknown in Dipholis.
More certain determination of its affinities awaits the collection of fruiting
material.
23. Bumelia revoluta Urb. Symb. Ant. 9: 417. 1925.
Unarmed shrub or small tree about 3—4 m. tall; young twigs pate
puberulent with rufous hairs; leaves firm, more or se revolute, obov
to narrowly oblong, about 7—25 mm. long a nd 2-6 wide, green and
shiny above, covered beneath with a dense tit nog sericeous- tomentose
pubescence, which becomes thinner, paler, and more strigose in age, not
at all veiny above, only the midrib evident beneath, subsessile, the petioles
about 1-3 mm. long; flowers 1 or seldom 2 in the axils, the wenden
pedicels about 2-4 mm. long; sepals rufous-hairy, about 2.5 mm
corolla (not seen by me) about 3 mm. long, the lobes without lateral he
ovary pubescent; fruit snap
TYPE COLLECTION: Ekm , ‘Sierra de Nipe prope Woodfred in fruticetis
carrascales ca. 500 ae Sacha Pac September (NY).
DistripuTIon: Oriente, Cuba.
Cupa: Oriente: Ekman 4034 (US); Shafer 3181 (NY).
DouBTFUL AND EXCLUDED SPECIES
Bumelia amazonica Krause, Notizbl. Bot. Gart. Berlin 6: 170. 1914. Dubious; possibly
a Pouteria; type-photo seen
Bumelia ambigua Ten. Sem. Hort. Neap. 1827. Nom. dub.
Bumelia argentea Roem. & Schult. Syst. 4: 499. 1819 = HEERIA ARGENTEA.
Bumelia Auzuba Roem. & Schult. l.c. = MASTICHODENDRON sp.
Bumelia borbonica Lodd. ex Loud. Hort. Brit. 69. 1830. Nom. dub.
Bumelia crenulata Spreng. Syst. 1: 665. 1825. Apparently not sapotaceous.
Bumelia cuneifolia Rudge, Pl. Guian. 1: 30. 1805 = CHRYSOPHYLLUM CUNEIFOLIUM.
Bumelia denticulata Raf. New Fl. Am. 3: 29. 1836. Not sapotaceous.
Bumelia depressa Urb. & Ekm. Ark. Bot. 22A(17): 73.1929. Not sapotaceous.
Bumelia dulcifica Schum. & Thonn. Beskr. Gui. Pl. 130. 1827 = PourTERIA DULCIFICA,
according to Baehni.
Bumelia foetidissima Willd. Sp. Pl. 1: 1086. 1797 = MAstTICHODENDRON sp.
Bumelia laurifolia Standl. Trop. Woods 18: 31. 1929 = POUTERIA AMYGDALINA.
Bumelia lucida Roem. & Schult. Syst. 4: 499. 1819. Nom. dub.
Bumelia macrantha Willd. ex Roem. & Schult. Syst. 4: 802. 1819 = ARDISIA sp.
Bumelia Manglillo Willd. Sp. Pl. 1: 1087. 1797 = MyrsinE MANGLILLO.
Bumelia Mastichodendron Roem. & Schult. Syst. 4: 493. 1819 = MasTICHODENDRON sp.
Bumelia multiflora Roem. & Schult. Syst. 4: 498. 1819. Not sapotaceous.
1945] CRONQUIST, DIPHOLIS AND BUMELIA
Bumelia nervosa Vahl. Eclog. Am. 1: 28. 1796 = POUTERIA MACROPHYLLA.
Bumelia pallida Sw. Prodr. Veg. Ind. Occ. 49. 1788 = MAsTICHODENDRON sp.
Bumelia pauciflora Roem. & Schult. Syst. 4: 493. 1819 = MAsTICHODENDRON sp.
Bumelia pubescens Ten. Sem. Hort. Neap. 1827. Nom. dub.
Bumelia punctata Roem. & Schult. Syst. 4: 498. 1819. Not sapotaceous.
Bumelia serrata Pursh, Fl. Am. Sept. 1: 155. 1814 = PRUNUS CAROLINIANA.
Bumelia serrulata Raf. New Fl. Am. 3: 29. 1836. Not sapotaceous.
Bumelia strigosa Spreng. Syst. 1: 665. 1825. Nom. dub.
Bumelia undulata Raf. New Fl. Am. 3: 28. 1836. Probably not sapotaceous.
New York Botanicat GARDEN,
New York.
471
472 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
NOTES ON SOME CULTIVATED TREES AND SHRUBS, II*
ALFRED REHDER
Juglans nigra L. f. laciniata, f. no
uglans nigra laciniata J. Cake. Price list Nut Tree Nurseries p. 4 [1937] “Cut
Leaf Black Walnut,” cum descr. angl.; p. 6 [1940] “J. Laciniata’; p. 8 [1941]
“J. Nigra Laciniata,” :
A typo speciei differt. foliis bipinnatis, foliolis primi ordinis pinnatis,
pinnulis utrinque 5-8 oblongo-lanceolatis vel lanceolatis serratis vel
interdum plus minusve raro fere ad medium pinnatifidis basi decurrentibus,
aaa terminali pinnatifida vel plus minusve serrata.
Cu ED SPECIMENS: Arnold Arboretum, no. 309-41, E. J. Palmer, Sept. 8, 1944
any sinters in 1941 from H. F. Stoke, Roanoke, Va.).
A very distinct form with bipinnate leaves finely dissected into lanceolate
to linear-lanceolate and serrate to pinnatifid leaflets 1—-3.5 cm. long an
4-il mm. wide. In the shape of its foliage and in general appearance this
form resembles somewhat Rhus typhina {. dissecta Rehder, but it is more
finely dissected and more graceful; the tree certainly merits attention as a
highly ornamental form.
With the exception of the laciniate form of the English Walnut, Juglans
regia {. laciniata (Jacques) Schneider, no other form with lacinate leaflets
was known in the whole family until the form of J. nigra described above
was raised. The cut-leaved form of the English Walnut, from which J.
regia {. heterophylla (Jacques) Schneider differs but little, is distinctly
inferior as an ornamental plant to the cut-leaved Black Walnut, having a
rather coarse, very irregularly shaped foliage, and moreover being more
tender.
According to information kindly furnished by Mr. J. W. Hershey, the
owner of the Nut Tree Nurseries at Downingtown, Pa., about thirty plants
of this form were found in 1926 at the State Nursery at Milton, Pa., among
seedlings raised from seed collected from normal Black Walnut trees in
Buffalo Valley, east of the State Nursery. Unfortunately most of the
seedlings got lost, but Mr. Hershey obtained one plant and, recognizing its
ornamental value, propagated it so that he was able in 1937 to offer it for
sale. The Arnold Arboretum received in 1941 a plant of this form from
Mr. H. F. Stoke of Roanoke, Va., with the statement that it originated in
Pennsylvania; this was probably obtained from the same source as Mr.
Hershey’s plant.
Juglans cordiformis Maxim. var. ailantifolia (Carr.), comb.
Juglans Sieboldiana Maximowicz in Bull. Acad. Sci. St. Pétersb, ‘18: 61 (in Mél. Biol.
8:63) (1873).— Miyabe & Kudo, ane For. Trees Hokk. 65, t.20 (1922). — Non
Goeppert (1854), fossil.
* For no. I see p. 67 of this volume.
1945 ] REHDER, CULTIVATED TREES AND SHRUBS, II 473
Juglans ailantifolia Carriére in Rev. Hort. as 414, fig. 85-86 (1878). — Little in
our. Washington Acad. Sci. 33: 132 (19
Juglans macrophylla Carriere in op. cit. ee nee
Juglans japonica Hort. ex Lavallée, Arb. Segrez. Teor: 1 [1880], pro syn
Juglans ailantifolia var. cordiformis (Maxim.) Rehder in Jour. Arnold Aeb 26: 68
(1945).
When Little in 1943 (1. c.) drew attention to the fact that J. Sieboldiana
Maxim. was antedated by J. Sieboldiana Goeppert of 1854 and took up the
name J. ailantifolia Carr. for the species, I accepted his disposition and, con-
sidering J. cordiformis Maxim. a variety of J. Sieboldiana, I published
the combination J. ailantifolia var. cordiformis, overlooking the fact that
J. cordiformis Maxim. of 1873 has priority over J. ailantifolia Carr. of 1878
and should have been taken up when J. Sieboldiana Maxim. was invali-
dated by the older homonym of Goeppert. As there can be little doubt
that J. cordiformis Maxim. and J. Sieboldiana Maxim. are conspecific and
represent only variations in the shape of the fruit, the name J. cordiformis
has to be taken up when J. Sieboldiana is rejected, which makes necessary
the new combination proposed above. If, however, J. cordiformis and J.
Sieboldiana Maxim. should be considered distinct species, the latter would
become J. ailantifolia Carr.
Ulmus parvifolia Jacq. f. pendens, f. nova.
A typo recedit ramis elongatis laxe pendulis.
CutLtivaATep in California: City Park, Covina, Los Angeles Co., southwest corner
of 4th Street and San Bernardino Road, tree with trunk 3’ 9” in Srenniieience. with the
head 50’ across, coll. C. R. Tower, Oct. 1944 (TYPE in herb. Arnold Arb. with photo. of
habit) ; same locality, coll. C. R. Tower, Sept. 1944 (in herb. Arnold Arb.) ; John Galvin
Park, Ontario, San Bernardino Co., coll. R. Tower, Sept. 1944, with photo. of habit
showing a wide-spreading tree with somewhat less pendulous branches than the type
tree (in herb. Arnold Arb.). There are me photographs in the collection of the Arnold
Arboretum of this form from North Clementine St., Anaheim, Orange Co., and from
the Roy F. Wilcox Nursery in Montebello, near Los Angeles.
The bark of the trunk and larger limbs is rather thin and scaly, the scales,
when shedding, exposing a smooth lenticellate pale reddish layer of bark.
The leaves are subcoriaceous, oblong-lanceolate, 3.5—5 cm. long and 9-20
mm. broad, acute or acuminate, unequal at the base, cuneate on one side
and subcordate to rounded on the other. The photographs listed above
show trees with a large spreading head to fifty feet wide or more, more or
less flattened at the top without upright or ascending leaders, but with long
and slender pendulous branches and branchlets
This pendulous form raised in California (together with the typical form
from seed received about twenty-five years ago from China) seems to be an
unusual form in China, for the several photographs in our collection of U.
parvifolia, taken in different parts of China, show trees of upright habit
except in a photograph of trees taken in ihe grounds of the Temple of
Heaven near Peking, which shows rather wide-spreading but not pendulous
branches.
The epithet ‘“‘pendens” has been chosen for this form to avoid confusion
with Ulmus parvifolia pendula Hort. ex Dippel, Handb. Laubh. 2: 27
474 JOURNAL OF THE ARNOLD ARBORETUM [VOL. XXVI
(1892), pro syn., a name which apparently had been used in nursery cata-
logues before Dippel cited it as a synonym of a form of U. pumila L.
Clematis triternata DC. f. rubro-marginata (Jouin), comb. nov
Clematis rubro-marginata Jouin in Mitt. Deutsch. Dendr. Ges. 1907(16) : 236 (1907).
Chomails Flammula var. rubro-marginata Cripps ex Jouin, |. c. (1907), pro syn
Clematis bor ii var. rubro-marginata Rehder in Jour. Arnold Arb. 1: 195 (1920) :
n. Cult. Trees Shrubs, ed. 2, 217 (1940).
According to Graebner f. in Ascherson & Graebner, Syn. Mitteleur. FI.
5,3: 65 (1935), C. triternata DC, [1817] is identical with C. violacea A.
DC. (1845), which makes necessary the transfer proposed above.
Spiraea Bumalda Burvenich f. pruhoniciana (Kriechb.), Comb. nov.
X Spiraea pruhoniciana Kriechbaum in Gartenschonh. 6: 38 (1925) “‘S. japonica ovali-
folia