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HARVARD UNIVERSITY
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MAR 4 1974
HARVARD
UNIVERSITY
Brigham Young University
Science Bulletin
NEW SPECIES OF
AMERICAN BARK BEETLES
(SCOLYTIDAE: COLEOPTERA)
by
Stephen L. Wood
BIOLOGICAL SERIES — VOLUME XIX, NUMBER 1
JANUARY 1974 /ISSN 0068-1024
BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN
BIOLOGICAL SERIES
Editor: Stanley L. Welsh, Department of Botany,
Brigham Young University, Provo, Utah
Members of the Editorial Board:
Vernon J. Tipton, Zoology
Ferron L. Andersen, Zoology
Joseph R. Murdock, Botany
Wilmer W. Tanner, Zoology
Ex officio Members:
A. Lester Allen, Dean, College of Biological and Agricultural Sciences
Ernest L. Olson, Chairman, University Publications
The Brigham Young University Science Bulletin, Biological Series, publishes acceptable
papers, particularly large manuscripts, on all phases of biology.
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.^"^^ ^
v."
Brigham Young University
Science Bulletin
NEW SPECIES OF
AMERICAN BARK BEETLES
(SCOLYTIDAE: COLEOPTERA)
by
Stephen L. Wood
BIOLOGICAL SERIES — VOLUME XIX, NUMBER 1
JANUARY 1974 /ISSN 0068-1024
TABLE OF CONTENTS
ABSTRACT 1
INTRODUCTION 2
SYSTEMATIC SECTION , 2
Cnesimis reticulum, n. sp 2
Cnesinus licairri. ii. sp. 2
Cnesinus terctis, n. sp 3
Cnesinus alienti.^, n. sp 3
Cnesinus depcrditus, n. sp 4
Cnesinus fulf^ens, n. sp 4
Cnesiiiiw fulgidus, n. sp 5
Cn^iim/s lucaris, n. sp 5
Cnesinus triangularis, n. sp 5
Cnesinus hrigliti, n. sp 6
Cnesinus coracinus, n. sp 6
Bothrosternus lucidus, n. sp. . 6
Hylastes niger, n. sp 7
Xi/lechinus mexicanus, n. sp 7
Phloeotrihus nanus, n. sp 8
Chramesus coruiger, n. sp 8
Chramesus disparilis, n. sp 9
Chramesus variabilis, n. sp 9
Chramesus microporosus, n. sp 10
Chramesus aipiiliis. n. sp. 10
Chramesus uisteriae, n. sp 1 1
Chramesus ma'ginatus, n. sp 11
Carphohorus piceae, n. sp 11
Carphohius cupressi, n. sp 12
Cladoctonus atrocis, n. sp 12
Seoh/todes eanalis, n. sp 13
ScoUjtodes cosiabilis, n. sp 13
Pseudothi/sciniies contrarius, n. sp 14
Pseudolhi/saniH's recaius, n. sp 14
Thijsanues granulifer, ii. sp 15
Micracisella mimetica, ii. sp. 15
Micracisella oeellata, n. sp 16
Htjloeurus riialis, n. sp 16
Jli/loeurus hinodatus, n. sp 17
Cn/phalomorpJius panalis, ii. sp 17
Cryphalomorphus sclifer, n. sp 18
Cn/phalomorphus hirtus, n. sp 18
C rt/ plwlomarphus rustieus. n. sp 18
Criiphuli)morphus trueis, n. sp 19
Hypothenemus apiealis, n. sp 19
Hypotlienemus indigens, n. sp .' 20
Hypothenemus trivialis, n. sp 20
Iltipothencnius dolosus, n. sp 21
Ht/putlicnemus solocis, n. sp 21
Hi/pothenernus veseulus, n. sp 21
Hypothenemus suspectus, n. sp 22
Periocrijphahis sohrinus, n. sp 22
Dendrocranidus limatus. n. sp 22
Dendrocranuhis red it us, n. sp , 23
Dendrocraiudus eonditus, n. sp 23
Dendrocranulus consimihs, n. sp 23
Dendrocranuhis vinralis, n. sp 24
Dendrocranulus vicinalis, n. sp 24
Dendrocranulus securus, n. sp 25
Dendrocranulus fulgidus, n. sp 25
Dendrocranulus vicinus, n. sp 25
Dendrocranulus rudis, n. sp , 26
Dendrocranulus conjinis, n. sp 26
Ips borealis lanieri, n. subsp 27
Gnathophthorus art us, n. sp 27
Dryocoetoides Hopkins 28
Dryocoetoides monachus ( Blandf ord ) , n. comb 28
Dryocoetoides verrucosus, n. sp 28
Dryocoetoides pileatus, n. sp 29
Dryocoetoides velutinus, n. sp 29
Dryocoetoides rusticus, n. sp 29
Dryocoetoides severus, n. sp 30
Dryocoetoides insculptis, n. sp 30
Dryocoetoides indolatus, n. sp 31
Sampsonius expulsus, n. sp 31
Sampsonius detract us, n. sp 31
Sampsonius usurpatus, n. sp 32
Xyleborus pristis, n. sp 32
Xyleborus micarius, n. sp 33
Xyleborus bicornutus, n. sp 33
Xyleborus carinitulus, n. sp 34
Xyleborus pandulus, n. sp 34
Xyleborus varulus, n. sp 35
Xylehonts sharpi lenis, n. subsp 35
Xyleborus palatus, ii. sp 35
Xyleborus exutus 36
Xyleborus rusticus, n. sp 36
Xyleborus ocellatus, n. sp 37
Xyleborus opimus, n. sp 37
Xylebo.us lacunatus, n. sp 37
Xyleborus merit us, n. sp 38
Xyleborus aclinis, n. sp 38
Xyleborus dissimulatus, n. sp 38
Xyleborus concentus, n. sp 39
Xyleborus tribulatus, n. sp 39
Xyleborus vismiae, n. sp 39
Xyleborus demissus, n. sp 40
Xyleborus meritus, n. sp 40
Xyleborus prolatus, n. sp 41
Xyleborus dissidens, n. sp 41
Xyleborinus dirus, n. sp 41
Xyleborinus tribulosus, n. sp 42
Xyleborinus protinus, n. sp 42
Xyleborinus celatus, n. sp 43
Araptus insinuatus, n. sp 43
Araptus interjectus, n. sp 44
Araptus accinctus, n. sp 44
Araptus delicatus, n. sp 44
Araptui genialis, n. sp 45
Araptus dentifrom, n. sp 45
Araptus facetus, n. sp 46
Araptus cuspidis, n. sp 46
Araptus placatus, n. sp 46
Araptus decorus, n. sp 47
Araptus blanditus, n. sp 47
Araptus mediatis 48
Araptun comlilus, n. sp 48
Araptus fnigalis, n. sp 49
Araptti.1 laudutus, n. sp 49
Araptus vesculus, n. sp 50
Araptus exigialis, n. sp 50
Araptus refertus, n. sp 51
Araptus trcpidus, n. sp 51
Araptus frontalis, n. sp 52
Araptus nigrellus, n. sp 52
Araptus vinnulus, n. sp 53
Araptus furvus, n. sp 53
Araptus funesccns, n. sp 53
Araptus Icpidus, n. sp 54
Araptus nwudicus, n. sp 54
Araptus uauulus. n. sp 54
Pseudopiti/ophthorus festivus, n. sp 55
Giiallidtrichus obscurus, n. sp 55
C'.nalliDtricltus omissus, ii. sp 56
Gnathotrupes dilutus, n. sp 56
Gnathotrupes crcccntus, n. sp 56
iricolus simplicis, n. sp ST
Tricolus inornatus, n. sp 57
Tricolus inaffectus, n. sp 57
Tricolus cecrojiii, n. sp 58
Tricolus intrustis, n. sp 58
Tricolus ardis, n. sp 58
Tricolus parsus 59
Tricolus rufithorax, n. sp 59
Tricolus hadius, n. sp 60
Tricolus partilis, n. sp 60
Tricolus fcnoris, n. sp 60
Tricolus frontalis, n. sp 61
Tricolus capitalis, n. sp 61
Tricolus nacvus, n. sp 61
Tricolus scitulus. n. sp 62
Tricolus peltatus. n. sp 62
Tricolus aciculatus. n. sp 62
Tricolus hicolor, ii. sp 63
Tricolus amplus, n. sp 63
Atnphicrantts inirandus, u. sp 63
Autphicranus lornatilis, ii. sp 64
Amphicranus macellus, n. sp 64
Amphicranus spinescens, n. sp 65
Amphicranus spinosus, n. sp 65
Amphicranus mucronatus, n. sp 66
Amphicranus acus, n. sp 66
Paracorthi/lus mutilus, n. sp 66
Paracorthi/lus concisus, n. sp 67
NEW SPECIES OF AMERICAN BARK BEETLES
(SCOLYTIDAE: COLEOPTERA)^
by
Stephen L. Wood=
ABSTRACT
The following 156 species of American Scolv-
tidae are described as new to science: Cnesinus
reticiihts (Venezuela), C. beaveri (Brazil), C.
teretis (Venezuela), C. aUentis (\'enezuela),
C. (leperditus (Colombia), C. fulgcns (Vene-
zuela), C. fiilgidus (Colombia), C. lucaris
(Venezuela), C. triangularis (Colombia), C.
brighti (Mexico), C. coracimis (Mexico), Botli-
Tostcrnus lucidtts (Brazil), Ilylastcs niger (Mex-
ico), Xylcchimis mexicanus (Mexico), Pidocotri-
hiis nanus (Brazil), Chramesus corniger (Mex-
ico), Ch. dispardis (Mexico), CIi. variabilis
(Mexico), Cli. inicroi)()rosus (Mexico), f7(. aq
udus (Mexico), Ch. uistcriae (Mississippi), Cdi.
marginatus (Mexico), Carphoborus piceae (Ore-
gon), Carphobius cupressi (Guatemala), Cda-
doctonus atrocis (Brazil), Scohjtodes canalis
(Mexico), S. costabilis (Mexico), P.scudothijsa-
iwes contrarius (Mexico), P. recavus (Mexico),
Thijsanoes gramdifer (Mexico), Micracisella mi-
metica (Mexico), M. ocellata (Mexico), Ihjlo-
curus rivalis (Mexico), H. binodatus (Missis-
sippi), Cnjphalonwr pints parvatus (Honduras,
Costa Rica), Cr. setifer (Guatemala), Cr. hirtus
(Mexico), Cr. rusticus (Mexico), Cr. trucis
(Mexico), Ht/pothcncmus apicaUs (Mexico),
Htj. indigens (Mexico), //i/. trivialis (Costa Rica,
Panama, Venezuela), Ily. dolosus (Costa Rica),
IIij. solocis (Mexico), Ily. vesculus (Mexico),
Hy. .ntspcctus (Costa Rica, Panama, Venezuela),
Periocrypludus sobrinus (Brazil), Dcndrocranu-
lus limatus (Venezuela), D. reditus (Venezue-
la), D. conditus (Venezuela), D cotmmilis
(Mexico), D. vinealis (Honduras), D. vicinalis
(Costa Rica), D. securus (Costa Rica), D. ful-
gidus (Panama), D. vicinus (Honduras), D.
rudis (Mexico), D. confinis (Panama), Gna-
thopthorus artus (Brazil), Dryocoetoides verru-
cosus (Venezuela), Dr. pileatus (Venezuela),
Dr. velutinus (Venezuela), Dr. ru.sticus (Vene-
zuela). Dr. sevenis ( X'enezuela), Dr. insculptus
(Colombia), Dr. indolatus (Venezuela), Samp-
â– sonius expulsus (Colombia), Sam. detractus
*Mosl of the field work that led to the discovery of these insects
-Department of Zoology. Brigham Young University. Provo, t'tah
(Panama), Sam. usurpatus (Costa Rica), Xy-
leborus pristis (Costa Rica), X. micarius (Costa
Rica), X. bicornutus (Venezuela), X. crinitulus
(Venezuela), X. panduhts (Costa Rica, Pana-
ma), X. varidus (Venezuela), X. palatus (Mex-
ico), X. extitus (Costa Rica), X. rusticus (Mex-
ico), X. ocellatus (Colombia), X. opimus (Flor-
ida), X. lacunatus (Costa Rica), X. meridcnsis
(Venezuela), X. aclinis (Panama), X. dissimu-
latus (Costa Rica), X. concentus (Costa Rica,
Venezuela), X. tribulatus (Costa Rica), X. vis-
miae (Costa Rica), X. demissus (Costa Rica),
X. meritus (Costa Rica), X. prolatus (Costa
Rica), X. dissidens (Mexico), Xyleborinus dirus
(Costa Rica), Xy. tribulosus (Panama), Xy. pro-
tinus (Costa Rica), Xy. celatus (Colombia),
Araptus insinuatus (Guatemala), A. inter jectus
(Guatemala), A. accinctus (Mexico), A. dcli-
catus (Mexico), A. geiiialis (Guatemala), A.
dentifrons (Mexico), A. facetus (Costa Rica),
A. cuspidus (Mexico), A. placatus (Mexico),
A. decorus (Costa Rica), A. bkinditus (Mexico),
A. medialis (Costa Rica), A. conditus (Costa
Rica), A. frugalis (Costa Rica), A. laudatus
(Costa Rica), A. vesculus (Costa Rica), A. ex-
igialis (Panama), A. refertus (Guatemala), A.
trepidus (Guatemala), A. frontalis (Guatemala),
A. nigrcllus (Costa Rica), A. vitimdus (Costa
Rica), A. furvus (Panama), A. furvescens (Gua-
temala), A. lepidus (Costa Rica), A. mendicus
(Costa Rica), A. nanulus (Mexico), A. festivus
(Mexico), Pseudopiti/ophthorus fc.itivus (Mex-
ico), Gnathotrichus obscurus (Mexico), G. omis-
sus (Costa Rica), Gnathotrupes dilutus (Costa
Rica). Gp. concentus (Costa Rica), Tricolus
simplicis (Guatemala), T. inornatus (Costa
Rica), T. inaf jectus (Costa Rica), T. cecropii
(Costa Rica), T. intrusus (Venezuela), T. ardis
(Costa Rica, Panama), T. parsus (Costa Rica),
T. rufithorax (Costa Rica), T. badius (Costa
Rica, Panama ) , T. partilis ( Costa Rica ) , T.
fenoris (Costa Rica), T. frontalis (Mexico), T.
capitalis (Panama), T. naevus (Costa Rica), T.
was sponsored by the National Si lenie Foundation
S4lin2. Scolytidae contribution no 4.S.
Bricham Young University Science Bulletin
scituhis (Costa Rica, Panama), T. peltatits (Pan-
ama, Costa Rica), 7'. dcicuhitu.'i (Mexico), T. hi-
color (Costa Rica), T. amplus (Mexico), Amphi-
cranus iniranclus (Costa Rica), Am. tornatUis
(Costa Rica), Am. mcicellu.s (Costa Rica), Am.
spinescens (Costa Rica), Am. .spinosus (Costa
Rica), Am. mucromittis (Panama), Am. acus
(Venezuela), Paracortlujhi.i mtitihi.'i (Panama),
and Par. concisus ( Costa Rica ) . New subspecies
include Ips borealis lanieri ( Colorado, South Da-
k(jta), and X. shcirpi lenis (Mexico).
INTRODUCTION
A large number of species new to science
were discovered during the preparation of a
monograph of the Scolytidae of North and Cen-
tral America. Since it will be several years be-
fore the monograph is concluded, the new names
are being published in order to stabilizi- no-
menclature and to facilitate identification. On
the following pages 157 species and 2 subspecies
are described as new to science. The new spe-
cies represent the following genera: Cnesinus
(11), Bothrosternus (1), Hijlastes (1), Xtjlcchi-
mts (1), Phloeotribtis (I), Chrame.su.'i (7), Car-
p])ohorus (1), CarpJwhius (1), Clacloctomis (1),
Scohjtodes (2), Pseudot}njsanoes (2), Thijsanoe.s
(1), Micracisella (2), Hi/locuriis (2), CTijpJi-
aJoimirpims (5), Ihipotlwmnmis (7), Pcrio-
cniphalus (1), Dindrocranulu.s (11), Gnatlwpli-
thorus (1), Dryocoetokles (7), Samp.sonius (3),
Xyleborus (22), Xtjleborimis (4), ATaptus (28),
P.seu(h)f>itij(>])hthoius (1), Gmitliotrichus (2),
Gnutliotrupes (2), Tricolus (19), Amphicriimis
(7), and Paracorthijlus (2). One new subspe-
cies in Ips and one in Xijlcborus are also named.
The new species are from the following areas:
United States (4), Mexico (42), Guatemala
(10), Honduras (2), Costa Rica (45), Panama
(10), Colombia (7), Venezuela (20), and Brazil
(6). In addition, one species each also occurred
in the following combinations of countries: Hon-
duras/Costa Rica, Costa Rica/Venezuela. Two
species are from Costa Rica Panama/Venezuela;
four are from Costa Rica/Panama.
Illustrations, ke\s, and supplemental taxo-
nomic notes useful in identif\ing these species
will be presented in the monograph.
SYSTEMATIC SECTION
Cnesimis reticidus, n. sp.
This species is distinguished from the \-ery
closeK' related rctifcr Wood b\ the slightlv largiT
size, b\ the much more coarsel\- punctured pro-
notal disc, and bv the slightK' shorter declivital
setae.
Female.— Length 2.0 mm (paratype 1.9 mm),
2.5 times as long as wide; color very dark brown.
As in retifer except punctures on posterior
third of pronotum at k'ast twice as wide, inter-
spaces between punctures less than half as wide
as a puncture, punctures reduced in size an-
teriorly but in all areas much larger than in
retifer. Interstrial setae on declivity slightly
finer and shorter than in retifer.
Type Locality.— Thirty km E Palmar. Boli-
var, Venezuela.
Type Material.— The female holotxpe and
one female parat\pe were collected at the t\pe
locality on 12-VI-70, 200 m, No. 529, from Vi.wua
caijennensis, by S. L. Wood.
The liolotypt' and paratype are in m\' col-
lection.
Cncsimts beaveri, n. sp.
This spi'cies is distinguished from the allied
blackmani Schedl b\' the smaller size, by the
more widely separated eves and more broadly
convex frons, by the much more elongate, stri-
gose pronotal punctures, and by the finer, more
widely spaced elytral vestiture.
Female.— Length 1.4 mm (paratypes 1.35-
1.45 mm), 2.7 times as long as wide; color dark
brown.
Frons evenh' convex above shallow, trans-
verse impression at level of antennal bases; sur-
face strongly reticulate, with veiy fine, moder-
ately sparse, somewhat obscure punctures; -vesti-
ture of sparse, short hair; eyes separated b\ 2.0
times width of an eye.
Pronotum 1.1 times as long as wide; outline
as in blackmani; surface smooth, shining, punc-
tures fine and longitudinally striate, striations
Biological Series, Vol. 19, No. 1 New Species of American Bark Beetles
about 2-S or more times a.s long as wide, often
longitudinalK- confluent. Vistiture confined to
anterior third, of course, short, rather sparse
setae.
Elytra l.S times as long as wide, 1.9 times as
long as pronotum; sides almost straight and
parallel on slightK less than basal three-fourths,
rather narrowly rounded behind; anterior mar-
gins narrowly elevated, crest shallowly marked
into separate crenulations, no submarginal cren-
ulations; striae 1 moderatelv, others weakly im-
pressed, punctures rather coarse, distincth- im-
pressed; interstriae distinctly wider than striae,
shining, almost smooth, but with short, obscure,
subtransverse lines indicated, punctures fine,
shallow, uniseriate, close. Declivity steep, con-
vex; strial punctures slightK' smaller and deeper
than on disc, narrowly impressed, impression
narrower than punctures; interstriae 1 distinctly,
others weakh' convex. Vestiture largely confined
to declivity; consisting of slender interstrial bris-
tles, each with its apical third apparently flat-
tened; bristles two-thirds as long as distance
between rows, spaced within a row b\ length of
a bristle.
Type LoC'Vlity.— About 260 km N Xavantina,
Mato Grosso, Brazil ( 12 M9' .S .5r 46" W).
Type Material.— The female holotvpe and
three female parat\pes wen- taken at the type
locality on l-XII-68, No. F40, by R. A. Beaver;
one female paratype bears the same data except
13-X-68, No. B105; and one female paratype the
same data except 26-XI-6S, No. D71G.'
The holotype and one paratype are in the
British Museum (Natural Histor\), two para-
types are in the Museu de Zoologia, Universi-
dade de Sao Paulo, and two paratypes are in mv
collection.
Cnesinus teretis, n. sp.
This species is distinguished from the allied
l)Iackmani Schedl by the smaller size, b\- the
larger eyes, by the different frontal sculpture in
both sexes, and by the less strongly impressed
striae with smaller strial punctures.
Female.— Length 1.5 mm (paratypes 1.3-1.5
mm), 2.7 times as long as wide; color dark
brown, pronotum almost black.
Frons very narrow, weakly convex from ver-
tex to epistoma, median two-thirds on lower half
a slightly elevated plateau, this area to vertex
smooth, polished and entireK devoid of punc-
tures or setae; lateral areas below obscureK'
reticulate, with moderately abundant, coarse,
short setae, a row of setae continued along me-
dian margin of eye almost to narrowest point
between eyes; eyes separated by less than half
greatest width of eye, eyes ver)' large, coarsely
faceted.
Pronotum 1.15 times as long as wide; as in
hiackmani except sides more strongK' constricted
on basal half, punctures slightly smaller and
more elongate. X'cstiture confined to anterior
third, of fine, short, recumbent hair.
Elytra 1.6 times as long as wide, 1.6 times
as long as pronotum; outline as in hkickniani;
striae 1 moderately, others feebly impressed,
punctures small, shallow; interstriae twice as
wide as striae, almost flat, smooth, shining,
punctures minute, unisi-riate. those bearing setae
usualh' miinitel\- granulate. Declivity steep, con-
vex; striae 1 impressed; interstriae 1 narrowl)-
convex; interstrial punctures replaced by small,
rounded, setiferous granules. Vestiture confined
to declivity, discal interstriae and posterior half
of 3, 5, and 7; consisting of coarse bristles up to
slightly less than twice as long as distance be-
tween rows, only slightl\- longer than distance
between bristles within a row.
Male.— Similar to female except frons weak-
1\', transverseK- impressed on lower lialf more
strongly convex on upper half, surface reticu-
late and finely, sparsely punctured, vestiture
more generally distributed; disc with rows of
very small, fine interstrial setae.
Type Locality.— Seven km NW Socopo, Ba-
rinas, Venezuela.
Type Materl\l.— The female holotype, male
allot\pe, and 35 parat\pes were taken at the
type locality on 13-11-70, 200 m. No. 322, from
Nectandra twigs, b\ S. L. Wood.
The liolot\pe, allotype, and paratypes are in
mv collection.
Cnesinus alienus, n. sp.
TlTis species is distinguislied from the distant-
ly allied nitidus Eggers by the very differently
sculptured frons, by the larger, shallow pronotal
punctures, by the very shallow, smaller strial
punctures, and by the much more deeply sul-
cate el\ tral declivity.
Male.— Length 2.3 mm (allot\'pe 2.5 mm),
2.7 times as long as wide; color dark brown, al-
most black.
Frons basically convex with central third
rather deepK' concave, lower margin of concav-
ity at level just abo\e antennal insertion armed
by a pair of small, pointed, rather wideh sepa-
rated denticles; upper margin of concavity with
a median prominence; surface reticulate, almost
Brigham Young University Science Bulletin
rugose, a few small granules in lateral and lower
areas, punctures minute, obscure; vestiture of
fine, sparse hair.
Pronotum 1.2 times as long as wide; outline
as in iiitidus- surface mostU' dull, obscurely re-
ticulate to niiuutcK , longitudinalK etched, punc-
tures rather small, two to three times as long as
wide, larger than in nitidtis. Glabrous except
for a very few setae on anterior fourth.
Elvtra 1.7 times as long as wide, 1.6 times as
long as pronotum; outline as in nitidiis; striae
feeblv impressed, punctures sm;dl, shallow; in-
terstriae about three times as wide as striae, al-
most smooth, subshining, punctures obsolete.
Declivity steep, r;ither broad]\ sulcate; strial
punctures minute, distinct; sutural interstriae
moderately elevated, 2 strongly, broadly im-
pressed, 3 abruptly, moderately elevated on me-
dian side, devoid of granules, fine, uniseriate
punctures distinct except on 2. Vestiture con-
fined to decli\'it\ consisting of inti'rstrial rows
of rather short bristles.
Female.— Similar to male except frontal de-
pression less well developed, callus at upper
margin of concavity not evident; minute, con-
fused interstrial punctures indicated on disc.
Tyi'k Loc.^Lirv.— Forty km SE Socopo, Ba-
rinas, Wnezuela.
Tyi'e Matehial.— The male holot\ pe and fe-
male allotype were taken ;it the t\ pe localitN on
25-1-70, 150 m. No. 27.3, from a li;ina known
localK' as bejuco bianco, by S. L. Wood.
The holot\ pe and allotype are in my col-
lection.
Cnesinus deperdittis. n. sp.
This species is distinguislud from the allied
reticiihittis Chapuis In the smaller size, by the
different frontal sculpture, b\ the very much
more finely, obscurely pimctured discal inter-
striae, and b\- the- shorter, nuich less strongly im-
pressed elytral declivity.
Female.— Length 2.7 mm (paratype 2.5 mm),
2.6 times as long as wide; color ver\- dark red-
dish brown.
Frons as in rcticulatus except upper half
much more strongly comcx; transverse elevation
just above epistoma unarmed, triangular patch of
erect setae rather large, extending to deepest
point in impression; lateral margins of frons
abrupt but not acutely elevated.
Pronotum 1.1 times as long as wide; widest
just behind middle, sides rather strongly arcu-
ate, rather narrowly rounded in front; surface
subshining, sculpture similar to but slightly
finer than in reticulatus, punctures rather shal-
low, moderately coarse, manv or most at least
parth , longitudinally confluent. Vestiture of
moderately abundant, coarse, short setae.
Elytra 1.9 times as long as wide, 2.1 times
as long as pronotum; sides almost straight and
parallel on slighth- less than basal three-fourths,
rather broadl\ rounded behind; striae deeply,
;ibruptly impressed, punctures rather obscurely
indicated; interstriae about twice as wide as
striae, I'vidently finely rugose, punctures fine, ob-
scure, confused, rather abundant. Declivity
steep, con\('x; striae narrower and less strongly
impressed than on ilisc except 1 on right side
wider, interstriae 1 on right side flattened; inter-
strial punctures largely replaced by fine
granules.
Type Locality.— Piedras Blancas, 10 km E
Medellin, Antioquia, Colombia.
Type Matehial.- Tlie female holotype and
one female parat\pi- were taken at the type lo-
calit\ on 15-VII-70, 2500 m. No. PS5, Qricrcus
humlwldti. S. L. Wood.
The holot\pe and paratype are in m\ col-
lection.
Cnesinus fulgen.s, n. sp.
This species is distinguished from the very
closely ri'lated ni<^er Wood by the larger size,
1)\ the slighth' wider male epistomal elevation
with nuich shorter setae, by the more finely
punctured pronotum, b\' the slightly smaller,
less deepK impressed strial punctures, and by
the more deeply impressed declivital striae.
Mali:.— Length 2.7 mm (paratypes 2.6-2.9
mm), 2.7 times as long as wide; color d;irk red-
dish brown.
Frons as in niiier except epistomal elevation
wider, nonpiibescent area distinctly wider, setae
on its upper portion about half as lout;.
Pronotum as in Ji/i,'er except punctures iniich
smaller, more numerous, less strigose.
EKtra as in (i/f,'cr except strial punctures
smaller, not as deep; interstriae more than twice
as wide as striae, punctures confused; declivital
interstriiie 2 and 3 more strongly convi'x.
Female —Similar to male except epistomal
('le\ation reduci'd to a transverse callus with
one row of finer, lontjer setae on its upper mar-
KiJi-
Type Locality.— I^a Carbonera I'lxperimen-
tal Forest, 50 km (airline) NW Merida, Merida,
Venezuela.
Type Matehial.— The male holot\pe, female
allotype, and 25 paratypes were collected at the
Biological Series, Vol. 19, No. 1 New Specie.s of Amehic.\n B.\hk Beetles
type locality on 14-XI-69, 2500 m. No. 136, from
Rubus sp., by S. L. Wood. Thirty-two paratypes
are from La Miiciiy Expcrimontal Forest, 20 km
NE Merida, Mcrida, Venezuela, 22-.\II-69,
2500 m. No. 205, from Ruhus ,sp., by S. L. \Vood.
The holotype, allotype, and paratype.s are in
mv collectioiL
Cnesinus fulgidus, n. sp.
This species is distinguished from tlu' closely
allied juJ<iens Wood by the smaller size, by the
sculpture of the male epistomal ele\ation, by
the weakly impressed elytral striae, and hv the
feebly convex declivital striae.
Male.— Length 1.2 mm (paratypes 2.1-2.4
mm), 2.6 times as long as wide; color dark red-
dish brown, proiiotum often black.
Frons as in ful^ens except median longitu-
dinal axis of epistomal elevation half as great as
transverse axis, its upper margin bearing a nar-
row band of very short setae similar to those in
fulgens, glabrous area triangular, its surface
strongly reticulate.
Pronotum as in fuIgcns exet'pt punctures
more ncarlv o\al, dei'per.
Elvtra as in fuIgcns i-xcept striai' 1 moderate-
ly, others ven,' weaklv impressed, punctures
small, moderately deep; interstriae flat, smooth,
shining, twice as wide as striae, punctures mi-
nute, confused; declivity as in fuJgcns except
striae 2 and '3 not impressed, interstriae feebly
or not at all convex, 3 with a row of small,
rounded granules; decli\ital \estiture distinctly
longer.
Female.— Similar to male except epistomal
elevation reduced to a trans\'erse callus with one
row of longer setae on its upper margin.
Type Locality.— Piedras Blancas, 10 km E
Medellin, Antiocjuia, Colombia.
Type Material.— The male holotvpe, female
allotNpe, and 57 paratypes were taken at the
t)pe locality on 15-\'n-'70, 2500 m. No. 654 and
685, from Querciis htim]>ohUii twigs, bv S. L.
Wood.
The holotvpe, allotype, and paratypes are in
m\' collection.
Cnesinus hicaris, n. sp.
This species is distinguished from the closely
allied perplexus \\'ood by the nnich less exten-
sive male epistomal elevation and smaller brush
of epistomal setae, by the less coarsely sculptured
pronotum, and by the much more slender elytral
vestiture.
Male.— Length 2.3 mm (paratypes 2.2-2.5
mm), 2.8 times as long as wide; color dark
brown, elytra sometimes reddish brown, prono-
tum often abiiost black.
Frons essentially as in perplexus except epis-
tomal elevation much smaller, occupying slightly
more than median third, its longitudinal axis
about etjual in length to width of pedicel, its up-
per slope ornamented by about three rows of
compressed bristles, these bristles covering a
smaller area and finer than in perplexus.
Pronotum as in perplexus except grooves
slightly narrower and longer, ridges not as
strongly convex, finer. Vestiture finer than in
/)c'r/)/t'.v(/.v.
EKtra as in perplexus except both ground
cover and erect bristles much more slender,
slightly longer on declivity.
Female.— Similar to male except epistomal
elevation smaller, its bristles finer, reduced to
one row.
Type Locality.— Merida, Merida, Venezuela.
Type Matehial.— The male holotype and five
parat\pes witc taken at the type locality on 29-
\II-69, 1700 m. No. 210, from a small liana, by
S. L. Wood. The female allotype and 14 para-
types bear the same data except 22-XI-69, and
either No. 7 from a twig, or No. 6 from Vismia;
one parat\pe bears the sanu' data I'xcept 8-.\I-69,
No. 119 from Ruhus.
The holotvpe, allotype, and paratypes are
in my collection.
Cnesinus triangularis, n. sp.
This species is distinguished from the closely
related gihhulus Wood by the smaller average
size, by the smaller female epistomal elexation
with its setae less numerous and longer, by the
less deeply impressed striae, by the much smal-
ler interstrial punctures, and by the shorter,
stouter decli\ital pubescence.
FEMALE.-Length 2.2 mm (paratypes 2.1-2.4
mm), 2.7 times as long as wide; color black.
Frons as in gihhulus except epistomal eleva-
tion smaller, triangular, epistomal margin of tri-
angle almost straight, median angle of triangle
about 60 degrees, bristles more slender, much
longer; frons sparsely pubescent, particularly in
lateral areas.
Pronotum as in gihhulus.
Ehtra 1.9 times as long as wide; as in gih-
hulus except striae very weakly impressed, punc-
tures small, deep; interstriae three times as wide
as striae, shining, with a few very shallow, ob-
Bricham Young University Scien< e Bulletin
scuR", transverse lines, punctures very fine, inocl-
eratelv confused on 2 and 3, almost uniseriate
on others. Vestiture confined to declivity, con-
sisting of stout bristles, each bristle very slightly
longer than distance between rows.
Male.— Similar to female except frontal ele-
vation replaced by a small, low, transxersc, cpis-
tonial callus, specialized bristles absent.
Type Locality.— Piedras Blancas, 10 km E
Medellin, Antioquia, Colombia.
Type Materl\l.— The female holotNpe, male
allotype, and 12 paratypes were taken at the type
locality on 15-VII-70,' 2500 m. No. 656, from a
twig of a shrub known locallv as Uvo de Monte,
by S. L. Wood. Eighty-eiglit paratopes bear the
same data except \o. 65S, taken from a twig of
a small tree known locally as Graptero, bv S. L.
Wood.
The holotype, allotype, and paratypes are in
my collection.
Cnesinus l)ii'Jiti. n. sp.
This species is distinguished from cl('i:,(intis
Wood bv the smaller size, bv the smaller female
epistomal tubercles, by the coarser frontal x'esti-
ture, by the coarser pronotal striations, b\ the
fine hair covering the pronotum, b\- the less
strongly impressed striae, and b\' tiie discal pu-
bescence.
Feniale.— Length 2.5 mm (paratvpes 2.2-2.6
mm), 2.7 times as long as widi"; color rather
dark reddish brown.
Frons as in ele^antis but with epistomal den-
ticles much smaller and basally contiguous, \'es-
titure stouter and slightly more abundant.
Pronotum 1.1 times as long as wide; stria-
tions coarser and wider than either e/eganfi.s or
coracinus, punctures not evidi'ut. X'estiture of
rather abundant, fine, short hair on disc, longer
and coarser anteriorh'.
Elytra 1.9 times as long as wide; as in cleii'in-
tis except vestiture extends to base of disc, con-
sisting of rather abundant, short, coarse, con-
fused setae of about uniform length, not longer
on declivitv, each seta about equal in length to
width of an interstriat'.
Male.— Similar to female exci'pt epistomal
tubercles absent.
Type Locality.— Nine miles (14 km) SE
Teopisca on highway 24, Chiapas, Mexico.
Type Material.- The female holotype, male
allotype and 65 paratypes were taken at the t\ pe
locality on 14-\'-69, by D. E. Bright. Three para-
types have identical data except thev are .30-V-
69; three paratypes are from Lagos des Colores,
17-V-69, D. E. Bright.
The holotype, allotype, and most paratypes
are in thi' Canadian National Collection, some
paratypes are in my collection.
Cnesinus coracinus. n. sp.
This species is distinguished from elc^antis
Wood by the smaller size, bv the stouter body,
by the finer, closer female epistomal tubercles,
by the finer frontal \'estiture, b\- the sliglitly
coarser pronotal striation, bv the coarser strial
punctures, and by the shorter, stouter deelivital
setae.
Female.— Length 2.2 mm (paratvpes 2.1-2.3
mm), 2.5 times as long as wide; color black.
Frons as in elefiantis except epistomal tuber-
cles smaller, much closer, concavity not quite
as deep, extending slightly nearer upper level of
frons, surface obscureh' punctured, vestiture
finer, less abundant.
Pronotum 1.04 times as long as wide; much
as in elef^antis but with striations more distinctly
punctured and wider.
Elytra 1.6 tinii'S as long as wide; strial punc-
tures slighth' larger and deeper than in clcL!,(intis.
interstriae feel)ly convex; declixity less strongly
impressed, interstriae each with a row of gran-
ules. Vestiture confined to declivity, consisting
of sparse, short, ground \'estiture of rather fine
hair, and rows of interstrial bristles; each bristle
about as long as distance between rows, more
closely spaced within a row.
Type Locality.— Five miles ( 8 km ) S Simo-
liovc'l, Chiapas, Mexico.
Type Material.- The female holotype and
four female paratvpes were taken at the type
locality on 4-\'II-69, by D. E. Bright.
The holotype and two parat\pes are in the
Canadian National Collection; two paratvpes
are in my collection.
Bothrost emits lucidus, n. sp.
The eh tral decli\itv of this species has
moderatily long, uniseriate interstrial setae as
described for truncatus Eichhoff; it differs, how-
ever, bv the absence of a median frontal tubercle,
bv the pronotum being wider than long, bv the
smoother, more regularly puncturid pronotum,
and probabh' bv other characters.
Male.— Length 2.3 mm (paratvpes 2.3-2.5
mm), 2.2 times as long as wide; color reddish
brown.
Biological Series, Vol. 19, No. 1 New Species of Americ.\n Bark Beetles
Frons as in male brevis Eggers except upper
area slightly more strongK' convex, lower area
less strongly, less extensively impressed, with no
indication of a transverse calhis, punctures
smaller, less conspicuous; vestiture similar hut
coarser.
Pronotum 0.92 times as long as wide; outline
as in brevis except more distinctly constricted on
basal third; surface almost smooth, shining,
feebly reticulate at base and in lateral areas;
punctures small, close, moderately deep, round
to oval; lateral margin with a fine, acutely ele-
vated line. Glabrous. Lower three-fourths of
anterior propleural area excavated and filled by
a dense brush of white hair.
Elytra 1.4 times as long as wide, 1.6 times as
long as pronotum; sides almost straight and par-
allel on basal three-fourths, al)ruptl\- rounded,
somewhat narrowly rounded at apex; basal mar-
gins almost smooth, more distinctly elevated than
in brevis; striae narrowlv, distinctlv impressed,
punctures clearh', shallowh' indicated; inter-
striae twice as wide as striae, almost flat, smooth
except a few wrinkles on basal half, punctures
fine, distinct, confused. Declivity steep, broadlv
convex; striae reticulate, distinctlv wider than
on disc, punctures larger, deeper; interstriae as
wide as striae, reticulate, punctures replaced b\'
small, shining, uniseriate granules. X'estiture
abraded on disc; on declivitv consisting of uni-
seriate inti'istrial rows of moderatelv long, rather
stout bristles, each bristle as long as distance
bet\\een rows, very slightly closer within a row.
Female.— Similar to male except frontal im-
pression not quite as deep or as extensive, a
feeble, transverse, elevated line indicated on one
specimen.
Type Locality.— About 260 km N Xavantina,
Mato Grosso, Brazil ( 12 49' S SIMfi' W).
Type Material.— The male holot\pe, female
allotype, and six parat\pes were taken at the
t\pe locality, in 196S, by R. A. Beaver; the holo-
type and five paratvpes were taken 18-XI-68,
No. D06, the allotvpe on 24-L\-68, No. 17.3, and
one paratype on 36-XI-68, No. D98.
The holotype, allot) pe, and one paratype are
in the British Museum (Natural Historv), two
paratvpes are in the Museu dv Zoologia, Unixer-
sidade de Siio Paulo, and tliree parat\ p;'s are in
my collection.
Htjlastes niger, n. sp.
This species is distinguislied from mcxicanus
Wood by the narrow Iv impressed decli\'ital striae
with the punctures much smaller, and by the
much more closely spaced granules on the decli-
vital interstriae.
Male.— Length 4.3 mm, 2.7 times as long as
wide; color black.
Frons and pronotum as in mcxicanus except
frons less distincth' rt'ticulate, pronotum with
punctures distinctly smaller.
Elytra as in mexicanus except strial punc-
tures much smaller, interstriai' twice as wide as
striae on disc, almost three times as wide on
declivity; decli\itv not as steep; interstriai gran-
ules on declivity much more closely spaced,
spaced by distances equal to less than half width
of an interstriae.
Female.— Similar to male except frons more
finch' punctured; anterior discal area of prono-
tum with punctures retluci'd to almost obsolete.
Type Locality.— Thirty-one km (19 mi) E
Tulancingo, Hidalgo, Mexico.
Type Materl\l.— The male holotype and fe-
male allotype (damaged) were taken at the type
locality on 10-\'II-67^ 2100 m. No. 185, from 'the
same tunnel in a Finns log 60 cm in diameter, by
S. L. Wood.
The holot\pe and allotype are in my collec-
tion.
Xijlechinus nwxicantis. n. sp.
This species is distinguished from marmora-
tus Blandford by the more slender, more irregu-
larly sculptured pronotum which lacks scalelike
setae, by the slightly coarser strial punctures,
and by the much more slender interstriai
bristles.
Male.— Length 1.9 mm (paratypi's 1.6-1.9
mm), 2.7 times as long as wide; color brown,
vestiture pale.
Frons as in marnioratus but broader, vesti-
ture finer, without scales.
Pronotum 1.0 times as long as wide; about
as in marmoratus but anterior constriction not
as strong; surface shining, irregular throughout,
punctures fine, shallow, most subvulcanate. Ves-
titure of short, coarse hair of moderate abun-
dance; scales absent.
Elytra 1.7 times as long as wide; as in mar-
moratus except bases of interstriae 2-.5 each
bearing 1-4 submarginal crenulations, strial
punctures very slightlv larger. Ground vestiture
shorter than in marmoratus, apparently less
abundant, much more slender on sutural inter-
striae than elsewhere; erect bristles slender,
blunt or pointed, their length and spacing as in
marmoratus.
Brigiiam Y'oung University Scienc:e Bulletin
Female.— Similar to male cxctpt siihmai-
ginal crenulations at hasvs of cK tia mosth- al)-
sent.
Type Locality.— Yt-rha Biicna, 20 mi (32
km) N Bochil, Chiapas, Mexico.
Type Mateiual.— The male holot\'pe, female
allotype and nine paratypes were taken at the
type locality on 21-V-69, <S000 feet elevation, bv
D. E. Bright.
The holotvpe, allot\pe, and five paratypes
are in the Canadian National Collection, fonr
paratypes are in mv collection.
Phloeotiihus nanus, n. sp.
This species superficialh' resembles hiistrix
Wood, although the relationship is remote. It
differs from lu/strix by the smaller size, by the
subobsolete stria! pnnctnres, by the more
abnndant interstrial scales, and by the absence
of spines and ele\ati'd ari'as on the elytral de-
clivity.
Male.— Length 1.4 mm (parat\pes 1.4 1.6
mm), LS times as long as wide; color yellowish
brown.
Frons as in Iv/slrix cxei'pt smface senlpturc
finer, less regnlar. Antennae as in hi/strix.
Pronotum 0.81 times as long as wide; outline
about as in ht/strix: surface shining, very dense-
ly, rather coarsely, deeply punctured, each
puncture bearing a short, stout, subseal("Iike
seta; anterolateral margin armed b\ about tlnce
small crenulations.
Elytra LI times as long as wide, L5 times
as long as pronotum; sides almost straight and
parallel on basal halt, broadly roinuled ]);'l'ind;
striae strongly impressed, their margins slighth
beaded indicating positions of punctures, but
punctures obsolete; interstriae twice as wide as
striae, surface shining, finel\- punctati'-granu-
late and with a median row of slightly larger
granules. Decli\ity rather steep, broadlv con\'e.\;
interstriae slightly narrower and more convex
than on disc; devoid of tubercles or other ele-
vations. \'estitiue confined to interstriae, con-
sisting of a di'use ground cover of \ery short,
stout, subscalelike, pointed setae, each slighth
longer than wide, and interstrial rows of slightly
longer, similar setae; longer setae less than
twice as long as ground ciner, about four to
six times as long as w idi-.
Fexiale.— Similar to male I'xcept broadly
convex, more coarsely granulate-punctate, de-
void of denticles; anterolateral areas of prono-
tum with about 20 small crenulations on each
side.
Type Locality.— About 260 km N \avan-
tina, Mato Crosso, Brazil ( 12 49' S 5L 48' \V).
Type NLvteiual.- The male holot\pe, female
allotype, and eight paratypes were taken at the
type locality, in 196(S, by R. A. Beaver. The
liolotype was taken LXII-6S, No. C20, the allo-
type and one parat\pe 1-.\II-6.S, No. F27, two
paratypes 1-.\L68, No. F19, four parat\ pi's 1-.\II-
68, No. F26, and one paratype 28-L\-68, No.
A07.
The holot\pe, allof\pe. and two parat\'pes
are in the British Museum (Natural History),
two paratopes are in .Museu de Zoologia, Uni-
versidade de Sao Paulo, and four paratypes are
in m\ collection.
Chraniestts corni^er, n. sp.
This species is unique in the genus. The
male frons is \ery weakly impri'ssed. the lateral
margins are not elevated at all, the lower frons
in the male bears a pair of largi' denticles near
hut not on the epistomal margin. The elytral
ground \estiture is alisent.
Male.— Length 1.4 mm (parat\'pes 1.3-1.4
mm), 1.7 times as long as wide; color \erv dark
brown, almost black, \estiture pale.
Frons yer\' shallowly conca\'e from epistoma
almost to upper level of eyes, lateral margins
rounded, not elevated; a pair of rather large
tul)ereli's just above epistomal margin; their
bases separated by about one-third width of
frons; surface subshining, finely rugulose, a few
small, indistinct granules on upper half; vesti-
ture of sparse, fine, inconspicuous hair. An-
teniKil club mocU'rateh' large.
I'roiintnm 0,80 times as long as wide; widest
near base, sides arcuatelv eon\crging to rather
weak constriction just before rather narrowly
rounded anterior margin; entire surface strongly
reticulate; punctures small, widely spaced, those
on anterior half granulate to very finely asper-
ate. X'estiture of short, stout bristles of moderate
abundance.
l']l\ tra 1,1 times as long as wide; sides al-
most straight and parallel on slightly more than
basal half, broadly rounded behind; basal mar-
gins of elytra each armed b\ about 15 crenula-
tions, one submarginal eienulation on inter-
striae 2; striae feebly impressed, punctures
moderati-ly coarse, close; interstriae s'lightly
wider than striae, uniseriatelv, finely granulate
except slighth confused on 2, punctures not evi-
dent. Declivity rather steep, conxex; sculpture
about as on disc. Vestiture of interstrial rows
of stout bristles, each 8-10 times as long as
Biological Series, Vol. 19, No. 1 New Species of .Amehic.^n Bahk Beetles
9
wide, slightly shorter than distance between
rows, slightly confused on discal interstriae 2.
Female.— Similar to male e.xcept frons weak-
ly convex, unamied; pronotal granules smaller
in median area, two or three of those on antero-
lateral angles crenulate.
Type LocALiTi'.— Lago Catemaco, Veracruz,
Mexico.
Type Material.— The male holotype, female
allotype, and si.\ paratypes were taken at the
type' locality on 16-20-\T-69, by D. K. Bright.
The holotype, allotype, and tlni'e paratypes
are in the Canadian National Collection; three
paratypes are in my collection.
Chramesus disparilis, n. sp.
This species superficially resembles acacico-
lens Wood, but tlie male frons is entirely dif-
ferent; the clytral ground yestiture and erect
bristles are broad. It is not closely related to
any species from North or Central America.
Male.— Length 1.6 mm (paratypes 1.4-1.7
mm), 1.9 times as long as wide; color dark
brown, yestiture pale.
Frons \erv deeply, broadly conca\'e from
epistoma to slightly above eyes; lateral mar-
gins subacutely ele\ated, amied just abo\e le\el
of antennal insertion by a pair of low, blunt,
sub(juadrate denticles haxing bases displaced
mesad from crest of lateral margins; surface
fineh' rugose-reticulate, shining; \'estiture of
sparse, minute hair in concavity, of a few stout
setae of moderate length on margins. Antennal
club small for this gtnus, apex narrowh
roimded.
Pronotum 0.76 times as long as wide; widest
at base, sides and anterior margin almost form-
ing a semicircular arc, anterior constriction al-
most obsolete; surface finely reticulate, shining,
small granules of moderate abundance extend-
ing from anterior margin to base. N'estiture
rather abimdant, short, scalelike, each scale
about two to three times ;\s long as wide.
Elytra 1.3 times as long as wide; sides al-
most straight and paralk-1 on slightly more than
basal half, rather broadly rounded behind; basal
margins each anned by 13 crenulations, about
six submarginal crenulations scattered from in-
terstriae 2-4; striae distinctly, weakly impressed,
punctures rather coarse, deep; interstriae slight-
ly wider than striae, each with a uniseriate
row of fine granules and minute, obscure punc-
tures. Declivity rather steep, convex; as on disc
except interstriae 2 on lower half devoid of
granules and feebly impressed. Yestiture con-
sisting of ground cover of short, recumbent,
interstrial scales, each scale about twice as long
as wide; and interstrial rows of erect, scalelike
bristles, each about six times as long as wide,
each slightly more than half as long as distance
between rows or between bristles within a row.
Female.— Similar to male except frons feebly
convex, lateral margins rounded and unanned;
scales in ehtral ground cover only slightly
longer than wide.
Type Locality.— Lagos dc Colores, Chiapas,
Mexico.
Type Material.— The male holotype and 22
paratypes were taken at the type locality on 14-
VI-69, from Acacia, by D. E. Bright. The female
allotype and 25 paratypes are from seven miles
(11 km) SE Teopisca, on highway 24, Chiapas,
Mexico, 31-\'-69, from Acacia, by' D. E. Bright.
The holot\ pe, allotype, and most of the para-
types are in the Canadian National Collection;
the other paratypes are in nu' collection.
Chramesus variabilis, u. sp.
This species is distinguished from vincalis
Wood b\ the more extensi\e, more deeph' im-
pressed male frons, with lateral armature
higher, by the smaller pronotal punctures, by
the larger scales in the clytral ground yestiture,
and by the much stouter erect interstrial
bristles.
Male.— Length 1.9 mm (paratypes 1.8-2.3
mm), 1.6 times as long as wide; color dark
brown, vestiture fonning a slightly \ariegated
pattern in most specimens.
Frons broadly, deeply concave from epistoma
to slightly above eyes, lateral margins acutely
rather strongh' elexated, armed just abo\i' level
of antennal insertion by a large triangular denti-
tion; surface reticulate, epistoma and large pre-
mandibnlar lobe smooth, shining; vestiture of
sparse, minute hair. Antennal club large.
Pronotum 0.74 times as long as wide; out-
line as in disparilis; surface finely reticulate,
punctures small, shallow, close, spaced by dis-
tances e(|ual to diameter of a puncture, dc\oid
of granules. Wstiture of short hair and equal
numbers of scales, each scale four to six times
as long as wide; central and anterior setae
darker.
Elytra 1 0.5 times as long as wide; sides al-
most straight and parallel on basal half, broadly
rounded behind; twelve pairs of crenulations
on basal margins, six submarginal crenulations
scattered on bases of interstriae 2-4; striae dis-
tinctly impressed, punctures small, rather
10
Brigham Young University Science Bi'lletin
shallow; interstriae tlirce times as wide as striae,
smooth, bristle-bi'aring pimetiires small, almost
uniseriate, puneturi's hearing groiiiul scales
minute. Deeli\'ity rather steep, convex; sculpture
as on disc. N'estiture of ground cover oF small
scales, each scale one to two times as long as
wide; and rows of erect bristles, each bristle
about twice as long as ground cover, half as
long as distance between rows, as long as dis-
tance between bristles within a row, each bristle
about si.x times as long as wide; in an obscuri'
variegated pattern.
Female.— Similar to male except froiis weak-
ly convex, lateral margins rounded, unarmed,
surface rugose-reticulate; anterolateral areas of
pronotum sparsely asperate.
Type Locality.— Lago Catemaeo, X'eracruz,
Mexico.
Tyi'E Mateuial.— The male holotype, female
allotype, and 24 paratypes were taken at the
type' locality on 16-20-VI-69, by D. E. Bright.
The holotype, allotype, and 15 paratypes are
in the Canadian National Collection; nine para-
types are in mv colleetioii.
C'lirauicsiis uiicrojjorosiis, n. sp.
This species is distinguislied from pcriosiis
Wood by the deeper male frontal concavity
which extends slightly above the upper level of
the eyes, bv the finer pronotal punctures and
granules, by fewer submarginal erenulations on
the elytral bases, by tlie minute strial punctures,
and by the more slender, nonsubplumose scales
of the elytral ground vestiture.
Male.— Length 2.2 mm (paratypes L8-2.4
mm), 1.6 times as long as wide; color \erv dark
reddish brown.
Frons as in pcriosus except eonca\ity ex-
tending slightly above eyes, much deeper on
upper half. Pronotum as in periosiis except
punctures less than one-third as large, granules
much smalk'r and less numerous.
Elytra as in periosus except submarginal
erenulations near base of elytra about half as
numerous, strial puncture very minute to en-
tirely obsolete, striae smooth, shining, inter-
strial punctures also minute; scales in ground
cover about four times as long as wide, not
subplumose; bristles about three times as Ion"
as ground cover, scalelike, each about six to
eight times as long as wide.
Female.— Similar to male except frons con-
vex, foveate at center; lateral areas of pronotum
asperate; strial puncture very small, but dis-
tinctly larger; interstriae each with a row of
moderately large, pointed tubercles.
Type Locality.— El Sumidera, 15 miles (24
km) N Tuxtia Cutierrez, Chiapas, Mexico.
Type Matekial.— The male holotype, female
allotype, and 15 parat\ pes were taken at the
t\pe' locality on 7-\ 1-69, by D. E. Bright.
The holotype, allotype, and nine paratypes
are in the Canadian National Collection; six
paratN pes are in mv collection.
Cliramcsus (Kjuihis, n. sp.
.'\mong Central American forms, this species
is most nearly allii'd to deniissits Wood, but it
is distinguished by the more deeply concave
male frons with the lateral margins more
strongly elevati'd, by the more distincth punc-
tured pronotal disc, by the more rounded inter-
strial tubercles, and bv the different elvtral
vestiture.
Male.— Length l.S mm (paratypes 1.5-1.8
mm), 1.5 times as long as wide; color I)lack,
xt'stiture pale.
Frons broadlv, ratlier deeplv coneaxc from
epistoma to slightly beknv upper level of eyes,
lateral margins acutely, rather strongly elevated
with highest point just below level of antennal
insertion, de\did of dinticles; surface strongly
reticulate, smooth on epistoma, minute, obscure
punctures on upper half of concavity; vestiture
inconspicuous. Antennal club large.
Pronotum as in deini.'i.stis except granules
smaller, posterior fouith in median ari'a with
sparse, shallow, moderateh large punctures.
X'estiture slightly more slender than in demissiis.
Elytra 1.02 times as long as wide; sides al-
most straight and paralK-l on slightly more than
basal half, broadly rounded behind; basal mar-
gins each armed by 11 erenulations, one or more
submarginal erenulations on interstriae 2-5;
striae strongh impressed, punctures moderately
coarse, deep; interstriae about one and one-half
times as wide as striae, moderately convex,
smooth, with rows of lather large, narrowly
rounded tubercles, tubercles somewhat confused
on 2 and 3. Declivity rather steip, convex;
sculpture about as on disc. Vestiture of ground
ciner of rows of scales on both margins of each
interstriae, each scale up to twice as long as
widi'; and erect bristles in interstrial r(A\s ex-
cept mod(>ratelv confused on 2 and 3 on disc,
each bristle two-thirds as long as distance be-
tween rows, spaced within a row by length of
a bristle, each about eight times as long as
wide, ef|ual in width to scales in ground cover.
Biological Series, Vol. 19. No. 1 New Species of Americ-^n B.\hic Beetle.s
11
Female.— Similar to male except frons weak-
ly convex, lateral margin.s rounded; pronotal
granules absent, entire surface with shallow,
.sparse punctures of moderate size.
Type Locality.— Eight miles ( 1.3 km ) N
Ocasingo, Chiapas, Mexico.
Type Materl\l.— The male holotype, female
allotype, and 19 paratypes were taken at the
type' locality on 2-\a-69, by D. E. Bright.
The holotype, allotype, and ten parat\pes
are in the Canadian National Collection; nine
paratypes are in my collection.
Chramcsus uisteriae, n. sp.
Tliis species is distinguished from acacico-
lens Wood by the more coarsely, more deeplv
punctured pronotum, by the more deeply im-
pressed striae and smaller strial punctures, and
by the slightly more slender, erect interstrial
bristles.
Male.— Lengtli 1.8 mm (paratypi's 1.6-l.S
mm), 1.6 times as long as wide; color very dark
brown, vestiture pale.
Frons as in acacicolens except surface with
sparse, minutely granulate, small, obscure punc-
tures. Pronotum as in acacicolens except gran-
ules eyidenth smaller, less conspicuous, punc-
tures much larger, deeper, closer, punctures dis-
cernible from base to anterior fourth; \estiture
siiglitly more abundant.
Elytra 1.2 times as long as wide; as in
acacicolens except striae weakly impressed,
punctures larger, more distinctly impressed; in-
terstriae slightly less than twice as wide as
striae; erect interstrial bristles \'er\- slighth
shorter and more slender than in acacicolens,
each bristle six to eight times as long as wide,
half as long as distance between rows.
Female.— Similar to male except frons weak-
ly convex, lateral margins not elevated or
armed, foveate at center; pronotal asperities in
lateral areas larger; interstrial tubercles distinct-
ly larger.
Type Locality —Bay St. Louis, Mississippi.
Type Material.— The male holotype, female
allot^qie, and three parat\pes were tak(-n at the
type locality on 12-V-45, in dead Wisteria
stems, No. 45-9773, presumably by W. 11.
Anderson.
The holotype, allotype, and one paratype
are in the U.S. National Museum; two para-
types are in mv collection.
Chramesus mar^inatus, n. sp.
This species is distinguished from setosus
Wood b\' the larger size, by the much more
finely sculptured pronotum, and by the more
nearly hairlike elytral setae.
Male.— Length 2.5 mm (paratype 2.5 mm),
1.65 times as long as wide; color dark brown,
vestiture pale.
Frons similar to setosus but not as deeply
concave on upper half, punctures on upper half
slightly larger. Pronotum outline as in setosus;
surface reticulate, a few asperities in lateral
areas, a few minute granules on median part
of anterior third, punctures small, moderately
close, very shallow on anterior half, somewhat
deeper in posterior area. Vestiture of short,
slender bristles of moderate abimdance.
lillvtra outline and basal ann;iture as in
setosus; striae distinctly impressed, punctures
\'ery small, moderately deep; interstriae at least
four times as wide as striae, weakh- convex,
smooth, with a central row of fine granules
and a row of minute punctures on each margin.
Decliyit\' rather steep, broadly convex; sculp-
ture as on disc. \'estiture of sparse, short, erect,
slender, liristlelike ground cover in iipproxi-
niate rows on margins of interstriae, and rows
of interstrial bristles arising from granules, each
bristle twice as long as ground cover, two-thirds
as long as distance between rows, as long as
distance between setae within ;i row.
Type Locality.— Mexico.
Tyi'e Material.— The male h()lot\pe, female
allotxpe, and one male par;itvpe were inter-
cepted at Brownsville, Texas, on 26-1-49, No.
67.3.33, in dead wood 49-2948, by D. J. Smith,
in material coming from Mexico.
The holotvpe and allotype are in the U.S.
National Museum; the paratype is in my
collection.
Carphohorus piceae, n. sp.
This species is distinguished from the closely
allied perplexus Wood by the absence of gran-
ules on discal interstriae 3, by the less strongly
elexated, more finch' dentate alternate decli\'ital
interstriae, and by the smaller, flattened area on
the female frons.
Male.— Length 1.7 mm, 2.2 times as long as
wide; color almost black.
Frons as in perplexus. Antennal club 1.3
times as long as wide. Pronotum and elytral
disc as in perplexus except discal interstriae 3
devoid of granules; elvtral declivity as in per-
12
Bnic.HAM Young University Science Bulletin
plexus except alternate intersthae less strongh
elevated and more finely dentate, interstriae 2
wider.
Female.— Similar to male except frons flat-
tened on little more than median half from
epistoma to very sliglitly above eyes, epistomal
margin distinctly elevated.
Type Locality.— Dixie Pass, Malheur Na-
tional Forest, Oregon.
Type Matehlal.— The male h()lot\pe, dam-
aged female allotype, and one damaged female
paratype were taken at the type localit\ on
23-\T-61, from an unthriftv branch of a recently
fallen Picea enp^clnuinni. hv S. L. NN'ood. The
elytral decli\it\' is missing from both females.
The holot\pe, allot\pe, and paratype are in
my collection.
C,(ir])}Hil>'nts niprcssi. n. sp.
This is tlie second species assigned to this
genus. It is distinguished from arizonicus Black-
man bv the larger size, by the more finely
punctured frons, I)\ the convex elvtral declixity,
by the absence of denticles on the decli\ity,
by the brightlv polished appearance, bv the
finer \i'stitiire, and 1)\' man\' otlii'r cliaracters.
Male.— Length 2.8 mm ( parat\pes 2.2-.3.2
mm), 2.3 times as long as wide; color ver\
dark brown, almost black, e]\lra dark reddisli
brown.
Frons strongh', tiansxciseh- impressed just
below middle, impression extending to upper
level of eyes, epistonia strongly raised and bear-
ing a broad premandibular lobe, a small median
tubercle at base of lobe; surface of impressed
area ver\ smooth, l)rightly shining, \ crN fincK
punctured on lower third, punctures slightK
larger above; \ertex reticulate, dull. Eye shal-
lowly emarginate, finch' granulate. Antennal
scape elongate; funicle 6-segmented. longer
than scape; club as long as scape, L5 times as
long as wide, witli tluee straight, transverse
sutures.
Pronotuin 0.90 times as long as wide; wid("st
near base, sides rather weaklv arcuate and con-
verging slightly on basal half, rather strongly
constricted laterally just before broadlv rounded
anterior margin; surface smooth, brightly shin-
ing, punctures coarse, deep, close. Vestiture of
fine, moderately long, rather abundant hair.
l']lytra 1.6 times as long as wide, 2.1 times
as long as pronotiun; scutellum not exposed;
sides almost straight and parallel on basal two-
thirds, ratlier narrowly loimded behind; cacli
b.asal margin strongly arcuate and armed b\
about 16 crenulations, submarginal crenulations
poorly developed; striae not impressed, punc-
tures coarse, deep; intiTstriai' smooth, shining,
as wide as striae, punctures fine, deep, con-
fused, rather numerous. Declivity rather steep,
convex; sculpture about as on declivity except
strial punctures slightlv smaller; interstrial
punctures de\'oid of granules. Vestiture of fine,
rather short, abundant strial and interstrial hair,
and interstrial rows of similar but slightly
longer hair. Third tarsal segments broad,
bilobed.
Female.- Similar to male except fions con-
vex, with a slight central impression.
Type Locality.— Ele\'en km N San Marcos,
San Marcos, Guatemala.
Type Matehial.— Tiie male liolotspe, female
allotype, and 17 par;it\pes were taken on 23-
11-72, from Ctiprcssus Uicitonka branches, by
F. \V. Clark.
The holot\pe, aIlot\pe, and parat\pes arc
in my collection.
Cf'uloclonus afrocis. n. sp.
This species is distinguished from th:' ratlier
closeK allied scnttis (Wood) b\ the smaller
size, b\ till' stouter bod\' form, bv the coarser,
more irrcgul u' pronotal and strial punctures,
bv the irr(".iular interstriae, and by the presence
of denticles on (lecli\ital interstriae 6 and S.
Male.— Length 1.6 mm ( parat\pes 1..5-1.S
mm), 2.1 times as long as wide; color reddish
brown.
Frons as in senilis except slightly narrower,
piuictures larger, less numerous; impression on
lower area slightlv stronger; eye larger, more
eoarseh' faceted.
Pronotum 0.84 times as long as wide; out-
line as in senftis; surface smooth, shining, punc-
ttni's aN'eraging larger than in senilis, of irregu-
lar size and shape. X'estiture consisting of a
verv few, vvrx widely spaced bristles.
Flvtra L4 times as long as wide, 1.7 times
as long as pronotniri; essentialh' as in senilis
except strial puncturt'S averaging larger, sub-
quadrate; interstriae narrower than striae, punc-
tures fine, uniseriate, surface strongly undulat-
ing, particularly near declivit\. Declivity very
steep, coinev; punctures smaller than on disc,
confused, positions of interstriae determined
from positions of tubercles, each interstriae
with up to 10 tubercles, except 2 entirelv un-
armed, largest tubercles near base. 9 acutely,
not stronglv elevated, its crest cur\'ed toward
Biological Series, Vol. 19. No. 1 New Species ok Amebican Bahk Beetles
13
and joininp; lateral margin at position in lini'
with intcrstriac 3 (as in sentus). X'ostiturc con-
fined to dt'clivital area, consisting of rather
widely spaced interstrial bristles, each slightly
longer and stouter than in sentus.
Female.— Similar to male except frons
Ijroadly convex, with large central area (two-
thirds of area between eyes) smooth, shining,
impunctate, remaining areas bearing several
long hairlike setae; pronotum less deeply punc-
tured, interstriae less irregular, decli\ital tuber-
cles slightly smaller.
Type Locality.— About 260 km N Xaxantia,
Mato Cro.s.so, Brazil, at 12'49" Soutli 51 46'
West.
Type Mateiual.— The male holot\pi', ft'male
allotvpe, and two paratypes were taken at the
type' localit)- on 19-X-68, No. B47/S, by H. A.
Beaver. One paratype bears identical data ex-
cept 18-X-6S, No. B38c; four paratvpes bear the
same data except 25-X-6S, No. B94/3; and one
parat\pe bears the sanu- data except 24-XI-6S.
D51/1.
The holot\pe, allotvpe, and two paratvpes
are in the British Museum (Natural Ilistorv),
two parat\pes are in the Museu de Zoologia,
Universidade de Sao Paulo, and four paratvpes
are in m\ collection.
Scoliitodes ccinalis. n. sp.
This species is not eloselv related to anv
described species, although it is placed in the
genus near chisiae Wood and parvuhis Wood.
Distinguishing characters include the slighth
protuberant, lower female frons wliich has a
narrow, shallow, median sulcus, with the frf)ntal
vestiturc confined to the upper margins; the
discal interstrial punctures are obsolete; and the
pronotal and strial punctures are moderatelv
coarse.
Female.— Length L7 mm (paratvpes L4-1.7
mm), 2.5 times as long as wide; color black.
Frons rather bn^adly convex, slightlv pro-
tuberant on lower half, median fourth iust below-
upper level of eyes shallowly concave, impres-
sion continued on median sixth as a shallow
sulcus to epistoma; surface of upper half almost
smooth, rather fineh', deeph' punctru'ed, becom-
ing finelv granulate on lower half I'xcept reti-
culate in impressed area; \estiture apparenth-
restricted to margins of upper half, tips of
some long, \cllow, hairlike setae arising on ver-
tex reaching to epistoma.
Pronotum 1.1 times as long as wide: sides
almost straight on more than basal two-thirds.
converging slightly to anterolateral angles,
broadly rounded in front; anterior third weakly
decli\ous; surface reticulate, subshining, pimc-
tures on posterior half moderately coarse, deep,
not close, decreasing in size on anterior half,
most of them replaced by minute granules on
anterior sixth or obsolete. Glabrous.
Elvtra 1.6 times as long as wide; sides
straight on basal two-thirds, very slightly wider
at base of tlecli\'itv, rather narrowly rounded
behind; striae not impressed, punctures mod-
erately deep; interstriae as wide as striae,
smooth, shining, punctures obsolete, some with
one to three minute granules. Declivitv steep,
con\'e\; strial punctures smaller than on disc;
a few minute interstrial punctures usualh' pres-
ent. V'estiture of fine sparse, erect, interstrial
liair of moderate length on odd-numbered in-
terstriae, much shorter to obsolete on even-
numbered interstriae.
Mali:.— Similar to female except frons even-
Iv, more strongK con\ex, surface reticulate,
with scattered punctures, subglabrous.
Type Localiti.— Mt. Tzontehult/,, Chiapas,
Mexico.
Type Material.— The female holotvpe, male
allotype, and 20 paratvpes were taken at the
t\pe' locality on 26 and 29-V-69 and 12-VI-69,
9500 ft (3100 m) elevation, by D. E. Bright.
The holotN'pe, allotype, and 14 paratypes
are in the Canadian National Collection; six
p;iratvpes are in n\\ collection.
Scoli/tpcles costal)iUs. n. sp.
This species is distinguished from melano-
cephahis Blandford by the different female
frontal sculpture, b\ the \erv fine punctures
on the p.onotal disc, and In- the \t'r\- small
strial punctures.
Female.— Length 1.8 mm (paratypes 1.8-2.0
mm), 2.6 times as long as wide; color brown.
Frons rather broadh' flattened from epi-
stoma to \t'rtex; a pair of low. subparallel. longi-
tudinal carinae from Ie\'el of antennal insertion
to epistomal margin; surface smooth and shin-
ing aboN'e le\el of antennal insertion, a row of
punctures around margin; feebly bisulcate be-
tween carinae, finely reticulate-punctate in
sulci, smooth and shining between; vestiture of
long, yellow hair on margins above, tips of
longest setae reach level of antennal insertion,
sulci on lower third with fine, short hair.
Pronotum 1.1 times as long as wide; sides
straight on basal two-thirds, converging very
slightlv to anterolateral angles, broadlv rounded
14
in front; surface reticulate, anterior fourth
rather strongly declivous and finely asperate,
fine, almost obsolete punctures behind each as-
perity; punctures on posterior areas \er\ small,
shallow, moderately close. Glabrous.
Elytra 1.4 times as long as wide, 1.2 times
as long as pronotum; sides almost straight and
parallel on basal two-thirds, rather broadly
rounded behind; basal margins not carinate,
abrupt; striae not impressed, punctures very
fine, distinctly impressed, spaced within a ro\\
by about twice diameter of a puncture; inter-
striae smooth, shining, punctures \er\- fine,
three or more times as wide as striae, uniseriate
except moderately confused in some specimens.
Declivity steep, convex: sculpture essentially as
on disc.
Type Locality.— Lago Catemaco, X'eracruz,
Mexico.
Type Material.— The female holotvpe and
four female paratxpes were taken at the t\pe
localitN on 16-20-\'l-69 (holot)pe) and l-.3-\'-69
( paratypcs ) by D. E. Bright.
The holotype and two paratypes are in the
Canadian National Collection: the other two
paratxpes are in mv collection.
Psctidotluisnnocs conlrarius, n. sp.
This species is distinguished from havloni
Bruck by the more slender, recurved spines on
the male deelivital interstriae 2, .3, 5, and 7.
by the coarser strial punctures, by the broader
interstrial scales, and by the more broadly
roimded anterior margin of the pronotiwn.
Male.— Length 1.0 nun (paratypes 1.0-1.2
mm), 2.1 times as long as wide: color black.
Frons convex, a weak transverse impression
on lower half, subfoveate at centtT; surface
rugose-reticulate, punctures not evident: vesti-
ture confined to epistomal brush.
Pronotum 0.89 times as long as wide: widest
near base, sides moderately arcuate, conxcrning
toward rather broadly rounded anterior margin:
anterior margin armed by ten small denticles.
Vestiture of sparse, short, stout bristles, a few-
small scales in lateral area.
Elytra 1.3 times as long as wide; sides al-
most straight and parallel on basal two-thirds,
rather broadly roimded behind; striae not im-
pressed, punctures coarse, deep; interstriae as
wide as striae, smooth, shining, with uniseriati'
rows of small granules. Declivity convex, grad-
ual, beginning at middle of elytra; strial punc-
tures gradually reduced in size until minute
near apex; granules on interstriae 1 smaller tlum
Bricham Young University Science Bulletin
on disc, obsolete on 2 except one or two near
base enlarged into slender, sharply pointed, re-
curved spines, 3 with seven to nine similar
spines, 5 with three, and 7 with about five
spines; spines on 3 slightly recurved, longest
spines equal in length to width of an interstriae,
slightly shorter than interstrial scales. Vestiture
of rows of minute, fine, interstrial hair, and
rows of erect, interstrial scales; each scale about
two-thirds as long as distance between rows,
about three to four times as long as wide, those
arising from posterior surface of spines often
slightly larger.
Type Locality.— Lagos des Colores, Chia-
pas, Mexico.
Type NLvteiual.- The male holotype and
three m:»le paratxpes were taken at the t\pe
locality on 14-\'i-69, from Acacia, b\ D.' E.
Bright.
The holotype and two paratypes are in the
C-'an;idian National Collection: one paratype is
in m\ collection.
Pseudathi/.mtuH's rrcavtis, n. sp.
This species is closely allied to dimorphtis
(Schedl), but it is distinguished bv the larger,
deeper, strial punctures, by the more deeply ex-
ca\ated female frons, with the impri'ssion ex-
tending only slightly above the e)es, by the
presence of a dense fringe of short setae on
the upper margin of the female frontal concav-
ity, and by the stouter elytral scales.
Female.— Length 1.3 mm (paratypes: males
1.2 mm; females 1.2-1.4 mm), 2.2 times as long
as wide; color very dark brown, almost black.
Frons deeply, rather broadly concave from
epistoma to slightly above eves; premandibular
epistomal lobe rather well de\eloped; surface
of eoncayitv reticulate on upper two-thirds,
smooth below; upper margin of concavity bear-
ing a dense fringe scalelike setae on median
two-thirds; epistoma with a few long, sub-
plumose setae in lateral areas. Antennal scape
strongly triangular, two and one-half times as
wide as long, bearing a dense brush of long
hair somewhat more elaborate than in dimor-
phu.s; club moderateh' large, elongate-oval,
minutely pubescent, entirely devoid of sutures.
Pronotum 0.91 times as long as wide; as in
dimorphtis except posterior areas more finely
reticulate, and granules between summit and
basal margin slightly larger.
Elytra 1.4 times as long as wide; outline as
in diinoipluis: striae not impressed, punctures
rather fine, distinctly impressed: interstriae
Biological Series, Vol. 19, No. 1 New Species of Amehican Bahk Beetles
15
rather smooth, shining, almost twice as wide as
striae, punctures very fine, uniseriate. Declivity
steep, convex; interstrial punctures replaced bv
rounded granules of similar size and appear-
ance to those of dimorphus, but more widely
spaced. \'estiture of rows of minute, fine, re-
cumbent, strial hair, and rows of erect, inter-
strial scales; each white scale about five times
as long as wide, almost as long as distance be-
tween rows, slightly longer than distance be-
tween scales within a row.
Male.— Similar to female except i)od\ 1.9
times as long as wide; frons rather weakly con-
vex, rugose-reticulate, vestiture sparse, less con-
spicuous; anterior margin of pronotum anued
by eight small teeth; interstrial pimeturi'S on
disc fineh' granulate; interstrial scales less than
one and one-half times as long as wide.
Type Locality.— Five miles (8 km) S Simo
jo\el, Chiapas, Mexico.
Type Matehial.— The female holot)pe, male
allotype, and 62 paratypes were taken at the
type locality on 4-\TI-69, by D. E. Bright.
The holotype, allotype, and most of the
paratvpes are in the Canadian National Collec-
tion; the remaining paratvpes are in my
collection.
Thi/sanoes grannhfer. n. sp.
This species is distinguished from hersche-
iniae Blackman b\' the larger size, by the
smaller diseal intiTstrial tubercles, b\' the mucli
larger declivital interstrial tubercles, and by the
much larger declivital scales.
Male.— Length 2.2 mm, 2.9 times as lonsj
as wide; color dark reddish brown.
Frons largely concealed bv pronotum, e\i-
dently as in I>erschcmiiic. Pronotiuu as in
herschemiae.
Elytra LS times as long as wide; outline as
in herschemiae; striae not impressed, punctures
moderately coarse, deep; interstriae smooth,
shining, slighth' narrower than striae, punctures
fine, slightly granulate, uniseriate, granules
coarse near declivity. Declivit\' steep, con\ex;
strial punctures slightl\' smaller than on disc,
deep; interstrial tubercles as wide as diameter
of a strial puncture, as high as wide, on inter-
striae 1-7, very slightly smaller on lower half of
j declivit\'. Vestiture of rows of minute, fine strial
hair, and rows of erect interstrial scales; scales
on disc about three times as long as wide,
shorter than distance between rows, on decli\itv
four to eight times as long as wide, one to one
and one-half times as long as distance between
rows.
Type Locality.— San Cristobal de las Casas,
Chiapas, Mexico.
Type Materl\l.— The unique male holotype
was taken at the type locality on 7-\'-69, by
J. E. H. Martin.
The holotype is in the Canadian National
Collection.
Micracisella mimetica, n. sp.
This species is distinguished from kimlh
Blackman by the larger size, b\' the frontal
granules, bv the shorter, broader elytral scales,
and bv the smaller denticles on the ele\ated
dt'cli\ital interstriae 3.
Male.— Length 2. .3 mm (paratypes 2.3-2.4
mm), 2.9 times as long as wide; color dark
reddish brown.
Frons moderately convex except rather
abruptly, strongK', trans\'ersely impressed on
lower fourth; surface rugose-reticulate in mar-
ginal areas, more finely, rather obscureh' ru-
gose-reticulate in central area; upper two-thirds
with about 30 small, high, isolated granules,
two near center much larger; \'estiture of uni-
fonnlv distributed, short, coarse setae of mod-
erate abundance. Eye oval, shallowly emargin-
ate; twice as long as wide. Antennal club 1.0
times as long as wide; suture 1 reaching middle
of club.
Pronotum 11 times as long as wide; outline
and ;isperities as in kmilU; anterior margin
amied bv six teeth, lateral pair much smaller;
posterior areas rugose-reticulate, dull, with
small, low, shining granules of moderate abun-
dance. \'estiture on disc of recumbent scales,
each scale ;ibout four times as long as wide;
a few bristles in asperate area.
Elytra 1.9 times as long as wide; outline as
in kntilli, disc as hniilli except surface irregular,
with numerous transverse lines or wrinkles. De-
clivity as in kintUi except interstriae 1 and 9
slightly more strongly convex but with denticles
on summit much smaller. Vestiture recumbent,
of interstrial scales except declivital interstriae
2, 4, and 8 glabrous; scales evidently slightly
confused on odd-numbered interstriae, uniser-
iate on even-numbered interstriae; each scale
two to three times as long as wide.
Female —Similar to male in all respects;
distinguished externally only by terminal terga
of abdomen.
Type Locality.— Three miles (5 km) N
Suchixtepec on Highway 175, Oaxaca, Mexico.
16
Bhigham Young University Science Bulletin
Type Material.— The male holotypc, female
allotype, and three paratypes were taken at the
type locality on 4-VI-71, 9500 ft elevation, in
mistletoe on oak, by D. E. Bright.
The holotype, allotype, and one paratype
are in the Canadian National Collection; two
paratypes are in my collection.
Micracisella oceUata, n. sp.
This species is remotely allied to nitidula
Wood, but it is distinguished by the very deeply
impressed declivital striae and by the moder-
ately large pimcturcs on the discal striae, each
of which has a small, elevated, central point
giving the appearance of an eye.
Male.— Length 2.2 mm (paratypes 1.9-2. .3
mm), 3.5 times as long as wide; color dark
reddish brown, pronotum iisualh' darker.
Frons broadlv convex, a slight, transverse
impression immediately above epistoma; surface
finely rugose-reticulate, a few fine punctures in
lateral areas, a few small granules in median
area of upper half; vestiture of very short,
stout, subplumose setae laterally and on upper
half, a few longer setae along epistoma. Eve
feebly emarginate; twice as long as wide. An-
tennal club as in allied species.
Pronotum LIS times as long as wide; es-
sentially as in nitidula except scales on disc dis-
tinctly wider.
Elytra 2.4 times as long as wide; outline
about as in nitidula; striae not impressed, punc-
tures large, distincth- impressed, each with a
small, central, cle\atcd granule giving appear-
ance of an eye; interstriae as wide as striae,
shining, almost smooth, punctures fine, uni-
seriate, rather close. Declivit\' rather steep, con-
vex, with apex slightly produced; striae deeply
impressed, pimctures small, obscure; interstriae
1, 2, and .3 equally convex, almost half as high
as wide, each bearing a row of low, rounded
nodules on its lateral half, granules more ob-
scure and smaller on 3; surface on lower half
somewhat rugose-reticulate. Vestiture of rows
of fine, short, strial hair, and interstrial rows
of recumbent scales; each scale on disc eight
or more times as long as wide, about four times
as long as wide on declivity.
Female.— Similar in all respects to male;
presumably those specimens with a few more
setae on scape are females.
Type LocALiTi'.— Three miles (5 km) N
Suchixtepec on Highwa\- 175, Oaxaca, Mexico.
Type Material.— The male holotype, female
allotype, and 34 paratypes were taken on A-W-
71, 9500 ft elevation, from Arbutus, by D. E.
Bright. Two paratypes bear identical data to
the type but were taken from mistletoe on oak.
Nine paratypes are from 20.5 km N Oaxaca, 31-
V-71, 9000 ft, Arbutus, D. E. Bright. Eighteen
paratypes are from .37 miles (59 km) S Valle
Nacional, 24-V-71, 8500 ft, from Arbutus, by
D. E. Bright; one paratype is from 15 mi S
(24 km) Valle Nacional, 20-V-7L 4000 ft, from
Arbutus, by D. E. Bright.
The holotype, allot\pi', and most of the
paratypes are in the Canadian National Collec-
tion; the remaining paratypes are in m\' col-
lection.
llt/lucurus ricalis, n. sp.
This species is distinguished from sclncarzi
Hlackman b\' the serrate male decli\ital inter-
striae 9, with the nodules on all interstriae
higher and slightly closer, b\- the numerous,
strong, irregularly transverse, interstrial lines,
and b\ the more strongly rugose-reticulate pro-
notal disc. It is probabK' more closely related
to effeminatus Wood.
Male.— Length 2.6 mm (paratypes 2.1-2.7
mm), 2.8 times as long as wide; color very
dark reddish brown, some specimens almost
l)lack.
Frons as in cffcuiinalus. including transverse
elevation. .Antenna! clul) distinctly larger than
in effeminatus. Pronotinn as in effeminatus.
Elytra 1.8 times as long as wide; outline
similar to effeminatus, striae not impressed,
punctures moderately large, deep; interstriae
slightly wider than striae, shining, with numer-
ous, irregular, coarse, transverse lines giving in-
terstriae a subcrenulate appearance, some of
low, trans\erse ridges continuing across striae;
moderatelx high, roundi'd nodules near decliv-
it\ on all interstriae. Declivity steep, convex,
contours about as in schuarzi; striae 1 and 2
continuing to base of mucro; base of each inter-
striae with three to five rounded nodules,
largest as high as wide, 1, 3, and 7 with small,
pointed tubercles to middle of declivity, tuber-
cles on 2 and 4 extending to upper third, those
on 5, 6, and 8 ending on upper fourth, 9 mod-
erately ele\ated about as in sclmarzi but with
small nodules to apex of elevation. N'estiture
more abundant than in schicarzi, of rows of
minute strial hair, and rows of delicate, coarse,
pointed bristles of same texture as schuarzi;
Ijristles longest near base of declivity, each of
longest bristles longer than distance between
rows, more closelv spaced within a row.
Female.— Similar to male except frons de-
void of transverse ele\ation; transverse lines
Biological Series, Vol. 19. No. 1 New Species of American Bark Beetles
17
and subcrenulate ridges on elytral disc poorly
developed; interstrial nodules much less than
half as high, extending almost to middle of
disc; declivital tubercles smaller; vestiture finer.
Type Locality.— Three miles (5 km) N
Suchixtepec on Highway 175, Oaxaca, Mexico.
Type Material.— The male holotype, female
allotype, and 52 paratypes were taken at the
type localit)' on 4-\T-71, 9500 ft ele\ation, from
Pinus, presuinabK- by D. E. Bright.
The holot\pe, allotype, and most of the
paratypes are in the Canadian National Collec-
tion; the remaining paratypes are in my col-
lection.
Hylocurus hinodatus, n. sp.
This species has the impressed frontal area
with a pair of raised spongy patches somewhat
reminiscent of the rttdis group of species, but
with the elytral declivity similar to female
harncdi Blackman.
FEMALE.-Length 2.1 mm (paratypes 1.8-2.0
mm), 2.7 times as long as wide; color verv dark
brown, almost black.
Frons with a large, transversely reniform
concavity from halfway between epistoma and
upper level of eyes to vertex, widest point at
upper level of eyes, occup\ing three-fourths of
area between eyes; concavitv rather abruptlv
impressed, moderately deep; central area of
each half of concavity occupied bv a protu-
berant, oval, spongy area, occupying about half
of concave area; spongy areas rather narrowly
separated from one another; general sculpturt'
as in rudis: subglabrous.
Pronotum as in rudis except granules on disc
smaller. Elytral disc as in rudis. Declivity as in
nidis except tubercles distincth- larger; inter-
striae 1 with a moderately large protuberance
at middle of declivity, slightly displaced from
suture, almost as high as wide, similar to but
smaller than female harnedi: interstriae 9 not
more strongly elevated than in rridis. \'estiture
as in rudis.
Type Locality.- Nicholson, Mississippi.
Type Material.— The female holotvpe and
one female paratope were taken at the type
locality on 15-1-45, No. 45-2357, under hickory
bark, apparently by W. H. Anderson. One fe-
male piiratype is from Algiers, Louisiana, 18-
IV-45, No. 45-8751, on pecan, by Ran. Slide
mounts of genitalic parts designated bv Ander-
son Scolytidae Slide No. .367 (holotvpe) and
No. 453 (Algiers parat\pe) are in the U.S.
National Museum.
The holotype and one paratypc are in the
U.S. National Museum; the other paratype is
in my collection.
Crijphalomorphus parvatis, n. sp.
This unique species is characterized by the
small size, by the unarmed anterior margin of
the pronotum, and by the uniseriate rows of
strial hair and interstrial scales. It is much more
likely to be confused with species of Uijpo-
thenemus than with other Crijphalomorphus
species.
pEMALE.-Length 0.8 mm (paratypes 0.8
mm), 2.6 times as long as wide; color ver\ dark
brown with pale scales.
Frons moderatel\- convex, almost smooth,
with rather abundant, very small punctures;
vestiture inconspicuous. Eye elongate, \'ery
shallowly, broadly emarginate. Antennal scape
short; club subcircular, septum poorly devel-
oped, almost transverse.
Pronotum 1.0 times as long as wide; widest
slightly behind middle, sides moderately arcuate
but con\'erging onh- slightly to anterolateral
angles then rather abrupth' converging to form
subangulate, broadly rounded anterior margin;
anterior margin unarmed; summit distinct, in
front of middle; anterior slope rather finely as-
perate; posterior areas obscureh- subreticulate,
with rather fine, isolated granules of moderate
abundance in lateral and dorsal areas. X'estiture
of equall\- abundant short hair and scales in
posterior areas, hair only on anterior half.
Elytra 1.6 times as long as wide, 1.6 times
as long as pronotum; sides almost straight and
parallel on basal two thirds, rather narrowly
rounded behind; striae not impressed, punctures
uniseriate, small, rather shallow; interstriae as
wide as striae, rather smooth, punctures slightly
smaller than those of striae, not clearly defined.
Declivity narrowly convex, rather steep; punc-
tures slightly smaller than on disc. \'estiture of
uniseriate rows of fine, short, recumbent strial
hair, and uniseriate rows of erect interstrial
scales; each scale about four times as long as
wide, widest at its apex, spaced within a row
and between rows by distances slightly shorter
than length of a scale.
Type Locality.— La Lima, Cortez, Honduras.
Type Material.— The female holotype and
one damaged female paratype were taken at
the t\pe locality on 5-V-64, at 100 m elevation.
No. 581, from Oestrum scandens. by S. L.
Wood. One female paratype is from Cuapiles,
Limon, Costa Rica, 22-Vni-66, 100 m. No. 103,
leguminose vine, S. L. Wood.
18
BllIGHAM VOUNG UNIVERSITY SciENCE BULLETIN
The holotype and paratypes are in my col-
lection.
Cnjphalomorphtis setifer, n. sp.
This species is distinguished from hirtiis
Wood by the smaller size, by the presence of
four to eight teeth on the anterior margin of
the pronotuni, by the less strongly impressed
striae, and by the more slender interstrial
scales.
Female.— Length 1.7 mm (paratypes 1.4-1.7
mm), 2.5 times as long as wide; color vers
dark brown, with pale vestiture.
Frons broadly convex above, somewhat flat-
tened below, a weak transverse impression just
above epistoma; surface strongly reticulate,
punctures fine, rather obscure, moderately
close; vestiture inconspicuous. Eye entire, not
at all sinuate. Antennal scape elongate; club
rather large, ovate, suture 1 septate on lateral
half, its procurved groove extending to mi'dian
margin.
Pronotum 1.0 times as long as wide; widest
just behind middle, sides weaklv arcuate on
posterior two-thirds, broadh' rounded in front;
anterior margin armed by four to eight small
teeth; summit very slightly in front of middle;
posterior areas coarsely, very closely punctured.
Vestiture of short, stout, abundant hair.
Elytra 1.6 times as long as wide, 1.7 times
as long as pronotum; outline as in parvatus;
striae very feeblv impressed, punctures rather
coarse, moderately deep; interstriae very slight-
ly wider than striae, smooth, punctures fine,
deep, close, strongly confused. Declivity rather
narrowly convex, steep; strial punctures slightly
smaller, deeper; interstriae each with a imi-
seriate row of very fine granules. X'estiture of
abundant, short ground cover, setae hairlike at
base becoming scalelike on declivity, and rows
of erect interstrial bristles; each erect bristle
as long as distance between rows and between
scales within a row, those on declivity, stouter,
almost scalelike, about eight times as long as
wide.
Type Localitv.— Volcan Pacaya, Esquintla,
Guatemala.
Type Material.— The female holotype, male
allotype, and 24 parat\pes were taken at the
type locality on l-VI-64, 1300 m ele\ation. No.
665, from a cut liana, by S. L. Wood. Thirty-
seven additional paratypes are from Guatemala
City, ;30-V-64, 1300 m,' No. 641, cut liana, S. L.
Wood.
The holotype, allotype, and paratypes are
in my collection.
Cnjphalomorphtis hirtus, n. sp.
This species is distinguished from setifer
Wood by the larger size, by the presence of
only two teeth on the anterior margin of the
pronotum, by the more strongly impressed in-
terstriae, and by the stouter interstrial scales.
Female.— Length 1.8 mm (paratypes 1.7-1.9
mm), 2.3 times as long as wide; color dark
brown, almost black.
Frons as in setifer except transverse impres-
sion on lower half much stronger, and punc-
tures much coarser but shallow. Antennal club
as in setifer but slightly wider. Pronotum as in
setifer but anterior margin armed by only two
teeth and punctures in posterior area not as
deep; a few scales sometimes present on margin
in front of scutelhmi.
Elytra 1.4 times as long as wide; sides al-
most straight and parallel on basal three-fourths,
broadly rounded behind; striae moderately im-
pressed on posterior half of disc, punctures
rather coarse, deep; interstriae slightly uider
than striae, moderately convex towaid declivity,
punctures on disc small, close, confused, armed
near declivity b\' median rows of small gran-
ules. Declivity very broadly convex, steep;
sculpture as on posterior part of disc except
interstriae slightly narrower, granules much
larger. Vestiture of abundant ground cover of
short scales and median interstrial rows of erect
scales; erect scales little longer than ground
cover on basal half of disc, more than twice
as long on decli\itv, each scale almost as long
as distance between rows and between scales
within a rcw, each about lour to fi\'e times as
long as wide.
Mali: —Similar to female I'xcept transverse
frontal impression slightly deeper.
Type Localitv.— Sixteen km (10 mi) south
of Oaxaca, Oaxaca, Mexico.
Type Matehial.— The female holotype, male
allotype, and five parat\pes were taken at the
type locality on 6-\'II-.53, by S. L. Wood. One
paratype is from 6 km S Atlixco, Puebla, Mex-
ico, i4-\'I-67, 2300 m, S. L. Wood. All speci-
mens were taken from cut branches of a small
tree having a looseh' layered structure of woody
tissues, rather than uniformly hard wood.
The holot\pe, allotype, and paratypes are
in my collection.
Crt/phalomorpfius rusticus, n. sp.
Distinguished from kriahi Hopkins by the
two teeth iuming the anterior margin of the pro-
Biological Series, Vol. 19. No. 1 New Species of American B.vhk Beetles
19
notiini, h\' the much wick'r, stalflike ground \i's-
titure on the anterior half of the elytral disc,
and by the absence of interstiial granules on the
disc.
Female.— Length 1.5 mm (paratypes 1.2-1.5
mm), 2.6 times as long as wide; color dark
brown, almost black.
Frons as in kna]>i except transverse impres-
sion much less strongly de\eloped, punctures
coarser, deeper. Eye, antenna, and pronotum as
in knabi except anterior margin of pronotum
armed by two coarse teeth and puncturt's on
pronotal disc distinctb' larger.
Elytra as in kmibi except discal interstriae
larger, deeper; declivital striae greatly reduced;
interstrial ground vestiture at base of elytra of
slender scales, each scale as wide as those in
erect rows, on decli\'it\ each scale as wide as
long; erect scales on declivity slightly longer
than in knabi.
Male.— Similar to female in all respects.
Type Locality. -Thirty-three km (21 mi)
north of Juchitlan, Jalisco, Mexico.
Type Material.— The female holot)pe, male
allotype, and 19 paratypes were taken at the
type locality on 3-\TI-65, 1300 m, No. 184, by
S. L. \\'ood, from the same iiost as hirtus Wood
(described above). Six paratypes were labeled
Lagos, Guanajuato, Mexico, 'll-\T-65, 2000 m.
No. 44, S. L. Wood, from the same host. A
series not included in the t\pe series is from
11 km SE Tuxpan, Mielioacan, Mexico, 16-\'n-
53, 2000 m, S. L. Wood, also from the same
host.
The holotype, allotype, and paratypes are
in mv collection.
CniphaJotnoiphus Iritcis. n. sp.
This species is distinguished from nisticus
Wood b\' the coarser, strial punctures on the
elvtral declivitv, Iiy the more slender discal
interstrial ground xcstiture, and by the different
armature on the anterior margin of the prono-
tum.
Female.— Length 1.6 mm (paratypes 1.3-1.5
mm), 2 6 times as long as wide; color very
dark brown, almost black, \ cstiture pale.
Frons, eye, and antenna as in rusticus ex-
cept frontal punctures larger, deeper. Pronotum
as in rusticits except posterior areas reticulate,
and anterior margin armed by about four to
eight irregularl) placed teeth.
Elytra as in mstkus except strial punctures
slightly larger, deeper, interstrial punctures
minute but evident; discal ground vestiture
near base of elytra proportionately much nar-
rower than adjacent erect scales; strial punc-
tures on declivity average much larger.
Male.— Similar to female.
Type Locality.— Sixteen km (10 mi) S Oax-
aca, Oaxaca, Mexico.
Type Material.— The female holotype, male
allotype, and two paratypes were taken at the
t\pe localitN on 6-\II-53, 1700 m elevation,
S. L. Wood. Seventeen parat\pes are from 11
km (7 mi) S Atlixco, 13-VII-53, S. L. \\'ood;
eiglit paratypes are from 6 km (4 mi) S Atlixco,
14-\I-67, 2.300 m. No. 31, S. L. Wood; and eight
paratypes are from 19 km (12 mi) SE Mata-
moros, Puebla, 3-\TI-53, S. L. ^^'ood. All are
from the same host as hirtus Wood (above).
The holotvpe, allot) pe, and paratypes are in
m\' collection.
Hijpothenemus (//)/<■«/«, n. sp.
This species is distinguished from rotumli-
collis Eichhoff and erectus LeConte by the
much more slender, erect, interstrial bristles and
by the stronglv elev;ited interstriae 9 at the
elytral apex.
Female.— Length l.S mm ( paratypes 1.6-2.0
mm), 2.3 times as long as wide; color dark
brown, vestiture pale.
Frons essentiallv as in iDlundicoUis. Prono-
tum as in wtundicoUis, with 10 to 14 asperities
on anterior slope, posterior area more strongly
reticulate, anterior margin armed b\' four teeth,
lateral pair usually smaller.
Elytra as in wtiiiulicollis except declivity
not as steep, its outline with a distinct lateral
constriction near middle; declivital striae more
strongb- impri'ssed, punctures shallow, larger,
interstriae 1-3 distinctly convex, 9 distinctly
convex to junction with 3 then strongly elevat-
ed from there to apex. Vestiture of short, con-
fused ground setae, almost hairlike on disc, be-
coming scalelike on declivity, and rows of erect
interstrial bristles; each bristle longer on de-
clivitv, on declivity each as long as distance
i^etween rows or between bristles within a row,
each about eight times as long as wide.
Type LocALiTY.-Three km (2 mi) E Ar-
meria, Colima, Mexico.
Type MATERiAL.-Thi' female holotype and
seven f(>inale paratvpes were taken at the type
locality on 2S-\T-65, 70 m elevation. No. 130,
and on 21-\T-67, 100 m elevation. No. 72, from
an unidentified liana, bv S. L. Wood.
20
Bmc.HAM Young University Science Bulletin
The holotypo and parat\pi'.s arc in iii\ col-
lection.
Htjpothencnius indi<^ens. n. sp.
This species is distinguished from rotundi-
coUis Eichhoff by tlie ininiitel\' granulose cK tral
surface and by the niucli more slender, more
wideK' spaced, erect, intcrstrial bristles.
Female.— Length 1.7 mm (paratvpes 1.4-1.7
mm), 2.4 times as long as wide; color dark
brown.
Head and pronotum as in rotundicollis ex-
cept pronotal asperities sliglitly smaller, nar-
rower, posterior areas more strongly reticulate.
Elytra 1.6 times as long as wide, as in rotun-
dicollis, except surface minutely granulate,
bristle-bearing intcrstrial punctures on disc
granulate, decli\ital ground xcstiture more near-
ly scalelike, and erect intcrstrial bristles mucli
luore slender and more widely spaced within a
row; each bristle about eight times as long as
wide; spaced within ;» row b\- distances sliglith
greater than length of a liristle.
Male.— Similar to female except 1.2 mm
long; eye reduced in size; antennal club more
slender; most characters less sharpK formed;
vestiturc longer.
Type Locality.— Nine km (6 mi) S ITuajua-
pan, Oaxaca, Mexico.
Type NL\teiual.— The female holotxpe, male
allotype, and one male parat\pe were t;iken at
the type locality on 16-\ 1-67, No. 45, from
Serjanid, by S. L. Wood. Two paratypes arc
from 17 km (11 mi) N Iluajuapan, i.5-\T-67,
No. 4-3, Psitltutinfhtis: 4 paratypes from 1.3 km
(8 mi) SE Cameron, 21-VL67, No. 76, shrub;
3 paratypes from 5 km (3 mi) S Matamoros,
Puebla, 14-VI-67, 2000 m. No. 33, Toxicoden-
dron: 6 paratypes from \'olcan Ceboruco. Naya-
rit, 5-\TI-6,5, 1000 m. No. 189, Serjania; and oni'
parat\pe from \'olcan Colima, Jalisco, 23-\'L
65, 500 m. No. 103, shrub; all by S. L. W'nod.
The holotypc, allot\pe, and paratypes are
in mv collection.
Ilt/pofhcncmtts trivicdis. u. sp.
This species is distinuuished from holivitiniis
Eggers by the smooth, shining clytral disc and
smoother pronotal disc, by the more slender,
slightly longer intcrstrial bristles, and by the
much more strongly confused pmictures and
bristles on interstriae 2 and 3 on the liasal half
of the disc.
Fentale. —Length l.S mm (paratypes 1.5-l.S
mm) 2.2 times as long as wide; color black.
Frons conyex; a small median impression at
upper leyel of eyes; surface rather finely rugose-
reticulate, except smooth and shining in median
area from epistoma to impression.
Pronotum 0.90 times as long as wide; widest
on basal third, sides rather strongly, arcuately
conyerging to rather narrowly roundi'd anterior
margin; anterior margin armed by four teeth,
lateral pair smaller; anterior slope armed by
more than 35 moderately large asperities; pos-
terior areas subreticulate, shining, granulate be-
iiind summit, granules sparse and intermixed
with fine, shallow pimctures laterally. \'estiture
of hair, intermixed in posterior areas with slen-
der scales.
Elytra 1.4 times as long as wide; outline as
in related species; striae not impressed, punc-
tures moderately large, shallow, their inner sur-
faces reticulate-granulate; interstriae smooth,
shining, twice as wide as striae, punctures fine,
uniseriate on 1 and 4-10, confused on 2 and 3
except near declivity. Decliyity commencing
near middle of ehtra, rather gradual, convex;
striae weakly impressed, punctures smaller,
deeper; interstriae weakly convex, punctures
uniseriate, finely granulate. Vestiture of rows
of minute, fine, strial hair and rows of erect
intcrstrial bristles; each bristle slightly longer
than distance between rows or between bristles
within a row; each bristle on disc slightly flat-
tened, usvialK not flattened on declivity.
Tyte LocALrry.— Santa Ana, San Jose, Costa
Rica.
Type Mati-:hial.— The female holotype and
four female paratypes were taken at the type
locality on l-\'III-63, 1.300 m elevation. No. 92,
from Cnparia <inatemulcnsis. by S. L. Wood.
Other female paratypes were taken in Costa
Hica as follows: 17 same locality as type, 30-
\TII-63, 1.300 m. No. 152, unidentified tree
branches; one at Rio Damitas in the Dota Mts.,
San Jose. 22-\'ni-6.3, 2.50 m. No. 129, Vismia
iS^iiianrnsis; sc\en at BcyerlcN. Limon, 26-VIII-
63, 7 m. No. 154, \ine; 14 at Pandora, Limon,
23-VIII-63, 50 m, Nos. 141 and 149, tree branch-
es: two at Finca Gromaco on Rio Coto Brus,
Pimtarenas, 14-\'II-63, 500 m. No. 76, seedling;
one at Rincon de Osa, Puntarenas, ll-\TII-66,
50 m. No. 25; one at Caiias, Guanacaste, 13-
VII-66, .30 m. No. 92, Ochroma. Three paratypes
are from Barro Colorado Island, Canal Zone,
Panama, 27-.\II-63, 70 m. No. .342, tree branch;
three are from Ft. Clayton, Canal Zone, Pan-
ama, 22-.\II-63, .30 m, Nos. 319 and 330. Ser-
jania: nine are from 10 km SE Miri, Barinas,
N'cnezucla, 811-70, No. 305, Serjania: three are
from 8 km S Colonia (near Buena\'entura),
Biological Series, Vol. 19, No. 1 New Specie.s of American Bahk Beetle.s
21
Valle de Cauca, Colombia, 9-VII-70, Nas. 617
and 620, Fictis; and three arc from 37 km N
Matias Romero, Veracruz, Mexico, 29-\'I-67,
100 m. No. 125, tree branch. All were taken by
me.
The holotype and paratypes are in m\ col-
lection.
Hijpothenemus dolosus, n. sp.
This species is distinguished fioin opucus
(Eichhoff) by the absence of pits bcliind the as-
perities on the pronotal summit, b\ the more
coarsely, deepK- punctured pronotal disc, by tlie
smooth, shining elytral surface, and by the ab-
sence of scalelike setae on the pronotum.
Female.— Length 1.6 mm (paratypes 1.6-1.7
mm), 2.3 times as long as wide; color dark
brown to black.
Head and pronotum as in opants except
asperities at pronotal summit not associated
with deep pits, some punctures lietween sum-
mit and base fomi deep pits but without as-
sociated asperities; posterolateral areas subshin-
ing, deeply, closely, coarseh' punctured. Vesti-
ture all hairlike.
Elytra as in opticus except surface of in-
terstriae smooth, shining; punctures on discal
interstriae 2 usually uniseriate; decli\ital sur-
face minutely granulate.
Type Locality.— Pandora. Limon, Costa
Rica.
Type Material.— The female holotvpe and
seven female paratypes were taken at the type
locality 23-\TII-63' .50 m elcxation. No. 140,
from Mimosa, by S. L. Wood; two other para-
types bear identical data except they are No.
142, from an unidentified branch; and one para-
type No. 154 is from an unidentified liana.
Two paratypes are from La Lima, Cortez, Hon-
duras, 5-\'-64, 200 m. No. 575, Sucitcnia planta-
tion, S. L, Wood.
The holotype and parat\pes are in my col-
lection.
Hijpothenemus solocis, n. sp.
This species is distinguished from squamosus
Hopkins by the more strongly impressed discal
striae, by the slightly larger interstrial tubercles
on both disc and dccli\ity, by the more nar-
rowly conyex declivity with the costal margin
near the apex much more strongly elevated,
and by the reduction of tubercles toward the
apices of interstriae 2, 4, 5, 6, and S.
Female.— Length 1.3 mm (paratypes 1.3-
1.5 mm), 2.3 times as long as wide; color dark
reddish brown.
Head and pronotum as in squamosus except
surface of posterior areas more coarsely granu-
late.
Elytra 1.5 times as long as wide; sides al-
most straight and parallel on basal two-thirds,
distincth' constricted near middle of declivity,
rather narrowly rounded behind; striae distinct-
ly impressed, punctures moderately large, shal-
low, indistinct; interstriae as wide as striae, dis-
tinctly impressed, punctures moderately large,
shallow, indistinct; interstriae as wide as striae,
distinctly convex, entire surface granulate,
punctures uniseriate, granulate. Declivity rather
narrowly convex, steep; interstriae more nar-
rowly convex, particularly 1 and 3, granules
larger except obsolete to\\ard apices of 2, 4, 5,
6, and 8, posterior part of 9 with costal margin
rather strongly elevated. Vestiture of rows of
minute, fine, strial hair, and rows of erect, in-
terstrial scales; each scale on declivity slightly
shorter than distance between rows, slighth'
longer than distance between scales within a
row, each about two to three times as long as
wide.
Type Locality.— Three km ( 2 mi ) west of
Aniieria, Colima, Mexico.
Type Material.— The female holotype was
taken at the type locality on 28-\T-65, 70 m
elevation, No. 135, from a Phorademhon twig,
by S. L. Wood. Paratypes were taken in Mex-
ico as follows; two from Concha, Sinaloa, 20-
VII-53, 15 m, S. L. Wood; three from 27 km
(17 mi) E San Bias, Nayarit, 25-VII-53. 10
m, S. L. Wood; one from 8 km S Rio Santiago
Ferry, 27-XI-4S; two from 48 km (30 miles) N
Tepic, ll-VH-65, 500 m. No. 226 in In^a (one
specimen). No. 228 in Serjania (one specimen),
S. L. Wood; and one from Los Corchos, Nay-
arit, lO-MI-65, 7 m. No. 206, tree branch, S. L.
Wood.
The holotype and paratypes are in my col-
lection.
Hijpothenemus vesculus, n. sp.
This species is distinguished from naneUus
Wood by the smooth, shining posterolateral
areas of the pronotum, with the punctures en-
tirely devoid of granulation, by the unimpressed
striae, with small, shallow, strial punctures, by
flat, smooth interstriae, and by the much more
slender interstrial scales.
Feniale. —Length 1.0 mm, 2.2 times as long
as wide; color dark brown.
Frons as in nanellus but surface below
upper level of eyes smooth, shining, punctures
more distinct. Pronotum as in naneUus except
22
Bbigham Young Univehsitv Science Bulletin
surface smooth, shining, no trace of reticulation,
punctures hrrger, deeper, devoid of granules;
scales in posterior area more slender.
Elytra about 1.5 times as long as wide; out-
line as in naneUtis; striae not at all impressed,
punctures small, shallow, distinct; interstriae
flat, smooth, shining, tw(j to three times as
wide as striae, punctures very fine, uniseriate.
Declivity rather steep, convex; essentially as on
disc. \'estiture of rows of fine, short, strial hair
on disc with similar supplemental interstrial
hair on lower declivity, and rows of erect inter-
strial scales; each scale as long as distance be-
tween rows, more closelv spaced within a row,
each about four to six times as long as wide.
Type Locality.— Ocosingo X'alley, Chiapas,
Mexico.
Type MAXEniAL.— The unicjue female holo-
type was taken at the type locality on 7-MI-50,
by L. J. Stannard.
The holotvpe is in mv collection.
Ht/pothenemus suspecttis, n. sp.
This species is distinguished from the closeh
allied ci/Undricus Hopkins by the less extensive
frontal impression, and by the much more slen-
der interstrial scales. It could easily be confused
with areccae (Hornung) but it is smaller, and
the frontal impression is much less extensive
and more shallow.
Female.— Length L2 mm (paratypes L1-L3
mm), 2.4 times as long as wide; color almost
black.
Frons as in ci/lindricus except lower frons
shallo\\'ly, transversely impressed, not concave.
Pronotnm and elytra as in ci/Iinchicus except in-
terstrial supplemental hairlike setae in ground
vestiture more abundant on and near decli\it\ ,
and erect interstrial scales much more slender,
each scale about eight times as long as wide.
Type Locality.— Pandora, Liinon Prov..
Costa Rica.
Type Material.— The female holot\pe was
taken at the type locality on 23-\Tn-63, 50 m
elevation. No. 148, from a cucurbit \'ine, by
S. L. Wood. Paratvpes were taken as follows:
Seven bear data identical to the tvpe except
for collection No. 144, No. 147, or No. 149,
unknown tree branch; one from Finca Taboga
near Caiias, Cuanacaste, Costa Rica, 8-II-67.
Cccropia petiole; two from Finca La Lola.
Limon, Costa Rica, 10-1-63, Theobroma cacao.
J. L. Saunders; one from Rincon de Osa, Pun-
tarenas, Costa Rica, ll-\TI-66. 50 in. No. 58
Cecwpia petiole, S. L. Wood; one from 8 miles
(13 km) S El Hato del Volcan, 7-1-64, 1000 m,
No. 371, tree seedling, S. L. Wood; two from
20 km SW El Vigia, Meiida, \'enezuela, 10-XII-
69, 50 m, No. 188 \ine, S. L. Wood; one from
17 km SE Miri, Barinas, Venezuela, 17-XII-69,
150 m, No. 196, Albizzia carihaea, S. L. Wood;
one Ironi 3 km NE Creole, Barinas. Venezuela,
18-.\II-69, 150 m, No. 203, In<^a. S. L. Wood.
The holotvpe and paratypes are in m\' col-
lection.
Periocnjphalus sohrimis. n. sp.
This species is distinguished from puUus
Wood b)' the larger size, by the more strongly
reticulate pronotal disc, by the more strongh',
more narrowlv produced costal margin of the
el\ tra at and near the suture, and b\ th(> more
widclv spaced, uniseriate, decli\'ital interstriae.
Female.— Length 1.25 mm (paratvpe 1.2
mm), 2.3 times as long as wide; color black.
Frons and pronotmu as in piilhis except pro-
notal disc much more strongly reticulate.
Elytra as in pulhis except apical margin
slightlv more stronglv, more narrowlv produced;
minute strial hair, in rows, indicate positions of
obsolete strial punctures (similar setae present
on pulltts but supplemented by a few similar
interstrial setae); erect interstrial scales in uni-
seriate rows, each slightlv sliorter than in ptilltis
and much more widely spaced, spacing within
a row slightly greater than length of scale on
both disc and declivity (scales on declivital in-
terstriae 2 in ptillus confused, almost forming a
double row ).
Type Locality.— About 260 km N Xavantina,
Mato Crosso, Brazil.
Type Mateiual.— The female holotvpe was
taken at the t\pe locality on 13-.\-68, No. B07,
by R. A. Bea\'er. One female paratype bears the
same data except 7-XI-68, No. F03.
The holotvpe is in the British Museum ( Nat-
ural History); the paratvpe is in mv collection.
Dciulrocranithi.s limatus, n. sp.
This species is \-erv closely related to limns
Wood, but it is distinguished by the smaller size,
bv the more strongh granulate pronotal disc,
and bv the more denseU' pubescent female frons.
Male— Length 1.6 mm (paratypes 1.5-1.8
mm), 2.6 times as long as wide; color dark
reddish brown.
Frons as in liimts except more coarsely, close-
ly granulate-punctate, not at all reticulate; vesti-
Biological Series, Vol. 19, No. 1 New Species of American Bahk Beetles
23
ture less abundant, shorter. Pronotuin as in
limus except declivital striae witli punctures
slightly larger, interstriae 1 weakly elevated, 2
weakly depressed; vestiture slightly stouter.
Female.— Similar to male except frons slight-
ly flattened, rather finely, closely punctured, in-
cluding central area, ornamented by abundant,
long, yellow hair uniformly distributed from
epistoma to well above upper level of eyes.
Type Locality.— Bumbuin Forest Station,
Barinas, Venezuela.
Type Material.— The male holotype, female
allotype, and 75 paratypes were taken at the
type locality on 29-1-70, 150 m elevation, No.
276, cucurbit vine, by S. L. Wood.
The holotvpe, allotype, and paratypes are in
my collection.
Dendrocranulus reditus, n. sp.
This species is distinguished from vincuUs
Wood by the slightly larger size, by the much
stronger frontal impression of the male, with
the tubercle on the summit much more strong-
ly developed, by the finer declivital punctures,
and by the more slender interstrial setae.
Male.— Length 1.5 mm (paratypes 1.3-1.7
mm), 2.8 times as long as wide; color very dark
brown.
Frons as in vinealis except more extensively,
slightly more deeply impressed, median summit
on vertex much more strongly developed, fomi-
ing a rather narrowly rounded elevation. Prono-
tum as in vinealis except granules in median
area of disc much smaller, almost obsolete, ob-
scure reticulation present in some specimens.
Elytra as in vinealis except declivital punctures
smaller, interstriae 2 not as strongly impressed;
interstrial bristles more slender, very slightly
longer.
Female.— Similar to male except frons con-
vex, feebly impressed on lower half, summit on
vertex feebh' elevated, surface more coarsel)',
more unifonnly punctured, vestiture sparse; pro-
notal reticulation more commonly present.
Type Locality.— Nine km S Barrancas, Ba-
rinas, Venezuela.
Type Material.— The male holotype, female
allot\pe, and 42 paratypes were taken at the
type locality on l-X-69, 150 m elevation, No.
34 (some paratypes No. 35), cucurbit vine, S. L.
Wood. Other paratvpes were taken in Venezuela
as follows: 12 from 8 km SW Bumbum, Barinas,
11-11-70, 150 m. No. 311; 20 from 20 km SW El
Vigia, Merida, lO-XII-69, 50 m. No. 185; and 24
from 5 km W El Pino ( near shore of Lake Mari-
caibo), Merida, 20-XI-69, 10 m. No. 142; all from
the same host and collector.
The holotype, allotype, and paratypes are in
my collection.
Dendrocranulus conditus, n. sp.
This species is distinguished from the verv
closelv allied consimilis \\'ood by the more finely
punctured, subreticulate frons, by the much less
strongly granulate, reticulate pronotal disc, by
the larger strial punctures, and by the slightly
depressed declivital interstriae 2 which is en-
tirely without granules.
Male.— Length 1.4 mm (allotype 1.3 mm),
2.6 times as long as wide; color reddish brown.
Frons as in consimilis except surface obscure-
ly reticulate, punctures slightly smaller and less
granulate. Pronotum as in consimilis except sur-
face of disc reticulate, distinctly, rather coarsely
punctured, each puncture with a small, trans-
\'ersely elongate granule lateral to each punc-
ture. Elytra as in consimilis except strial punc-
turc^s larger; interstriae as wide as striae, im-
punctate; declivity with interstriae 2 slighth' de-
pressed, interstriae with uniseriate, fine punc-
tures, devoid of granules; acute ventrolateral
margin of declivity less well developed.
Female.— Similar to male except elytral de-
cli\'ity very slightly more convex.
Type Locality.— Bumbum Forest Station,
Barinas, \'enezuela.
Type Material.— The male holotype, female
allotype, and one callow, male parat)pc with
collapsed frons were taken at the type locality
on 29-1-70, 150 m. No. 276, from a cucurbit vine,
by S. L. Wood.
The holotype, allotype, and paratype are in
my collection.
Dendrocranulus consimilis, n. sp.
This species is allied to lirribus Wood, but it
is distinguished by the smaller size, by the more
slender form and by the shorter setae on the
declivity.
Male.— Length 1.4 mm (paratypes 1.2-1.5
mm), 2.5 times as long as wide; color yellowish
l)rown.
Frons convex, with a slight transverse im-
pression on lower half; surface shining, closely,
deeply, rather coarsely punctured at sides and
above, becoming almost impunctate on median
24
BniGHAM Young Univebsity Science Bulletin
fourth of lower half; vestiturc of fine, sparse,
long hair uniformly distributed.
Pronotum 1.2 times as long as wide, sides on
more than posterior two-thirds subparallel,
feebly arcuate, anterior margin broadly
rounded; anterior third moderately declivous,
fine asperate, asperities decreasing in size but
attaining base in lateral areas; disc shining,
rather coarsely punctured, most punctures part-
ly granulate; surface on posterior half very
slightly subreticulate. Vestiture of sparse hair.
Elytra 1.6 times as long as wide, 1.4 times
as long as pronotum; sides almost straight and
parallel on more than basal two-thirds, abruptly,
very broadly rounded behind; striae not im-
pressed, punctures rather small; interstriae al-
most smootli, twice as wide as striae, punctures
uniseriate, almost as large as those of striae.
Declivity very steep, transversely flattened be-
tween interstriae .3, longitudinally slightly con-
vex; posterolateral margin from suture to a])out
interstriae 3 subacutely elevated; striae 1 and 2
distinctly impressed; interstriae 1 weakly ele-
vated, 2 very slightly depressed, punctures of
2 and 3 finely granulate, obscurely granulate
in lateral areas. V^cstiture of rows of minute,
obscure, recumbent, strial hair and rows of
erect interstrial bristles; each bristle about one
and one-third times as long as distance between
rows or within a row, except shorter, slightly
more than half as long on declivital interstriae
1 and 2.
Female.— Similar to male except frons more
broadly, evenly convex, with frontal pubescence
about twice as abundant.
Type Locality.— Los Corchos, Navarit, Mex-
ico.
Type Material.- The male holotype, female
allot)pe, and 40 paratypes were taken at the
type locality on lO-VII-65, 7 m elevation. No.
207, from an unidentified, climbing cucurbi-
taceous vine, hv S. L. Wood. Other specimens,
not included in the type series, are from
Honduras.
The holotype, allotype, and paratypes are
in my collection.
Dendrocranulus vinealis, n. sp.
This species is distinguished from the closely
allied pumilus Wood by the shining pronotal
disc, by the subcrenulate granules accompan\-
ing each puncture on the pronotal disc, and
by the much more strongly impressed male
frons.
Male.— Length 1.3 mm (paratype 1.2 mm),
2.7 times as long as wide; color very dark
brown.
Frons convex above upper level of eyes,
strongly, transversely impressed below that
level; surface smooth and shining, with mod-
erately fine, siibgranulate punctures in convex
area, almost impunctate in impressed area, ves-
titure of sparse hair.
Pronotum outline as in pumilis, anterior area
more coarsely asperate; surface shining, punc-
tures moderately coarse, deep, rather close, each
with a low, lateral, subcrenulate, transverse
granule. Vestiture confined to margins, sparse,
hairlike.
Elytra! outline about as in ptimilus; striae
not impressed, punctures small, moderately
deep; interstriae smooth, twice as wide as
striae, punctures slightK' smaller than those of
striae, rathi-r widely spaced. Decli\'ity steep,
rather broadly flattened; strial punctures
deeper, slightly larger than on disc; interstriae
1 weakly ele\ated, 2 weakly depressed, punc-
tures not at all granulate; \entrolateral margin
rounded. Vestiture of interstrial rows of flat-
tened bristles, each bristle slightly longer than
distance between rows or within a row.
Type Locality.— La Lima, Cortez, Honduras.
Type Material.— The male holotype and
one male parat\pe were taken at the type local-
ity on 5-\'-64,' 200 m elevation. No. 579, in
Caijaponia niicrodonta, by S. L. Wood.
The holotype and paratxpe are in mv col-
lection.
Dendrocranulus vuinalis, n. sp.
This species is distinguished from vinealis
Wood by the much smaller strial and interstrial
punctures, by the more gradual elytral declivity,
and by the interstrial granules on the declivity.
Male.— Length 1.5 mm (paratypes 1.6-1.7
mm), 2.9 times as long as wide; color yellowish
brown.
Frons as in vinealis except impression not as
abrupt, impunctate area much smaller, small
granules more conspicuous in lateral areas. Pro-
notum as in vinealis except discal punctures
slightly smaller, granules not as slender. Elytra
as in vinealis except strial and interstrial punc-
tures smaller, very shallow; punctures " on de-
clivital interstriae 1-3 replaced by small gran-
ules.
Female.— Similar to male except frontal im-
pression not as strong, sculpture finer, vestiture
finer, more abundant but not conspicuous.
Biological Series, Vol. 19, No. 1 New Species of American B.\hk Beetles
25
Type Localiti.— Rio Damitas in the Dota
Mountains, San Jose, Costa Rica.
Type Material.— The male holotype, female
allotype, and one male paratype were taken at
the type locality on 18-11-64, 250 m elevation.
No. 441 (paratype 440), in an unidentified
climbing vine, by S. L. Wood.
The holotype, allotype, and paratype are in
my collection.
Dendrocranulus securus, n. sp.
This species is distinguished from rnaurus
( Blandford ) by the partly reticulate postero-
lateral areas of the pronotum, by the reticulate,
more finelv punctured frons, and by the shorter,
stout setae on declivital intcrstriac 1 and 2.
Male.— Length 1.8 mm (paratypes 1.7-2.1
mm), 2.5 times as long as wide; color reddish
brown.
Frons convex, a shallow, transverse impres-
sion from epistoma to upper level of eyes; sur-
face reticulate, punctures moderately fine, deep,
close, not at all granulate, less abundant near
median line on lower half; vestiturc sparse,
hairlike.
Pronotum 1.1 times as long as wide; widest
well behind middle, sides moderatelv arcuate
on posterior half, feebly constricted one-third
length from anterior margin, anterior margin
rather narrowly rounded; asperities small,
abundant; surface of disc and lateral areas sub-
reticulate, punctures rather small, deep, a low,
transverse, rounded granule lateral to each
puncture. Vestiture fine, sparse, hairlike.
Elytra 1.5 times as long as wide, 1.4 times
as long as pronotum; sides almost straight and
parallel on less than basal two-thirds, posterior
margin straight on median half; striae not im-
pressed, punctures rather small, deep, close; in-
terstriae almost smooth, twice as wide as striae,
punctures fine, uniseriate, rather widelv regu-
larly spaced. Declivity steep; striae 1 weakly
impressed; interstriae 1 weaklv elevated; inter-
stri;\l punctures minutely granulate; central half
flattened, broadly convex in all marginal areas
as in niaurus. Vestiture of rows of minute, re-
cumbent strial hair, and rows of erect, inter-
strial bristles; each bristle almost as long as
distance between rows and between bristles
within a row, slightly shorter and coarser than
in rnaurus.
Female.— Similar to male except frontal im-
pression less distinct, vestiture very slightly
more abundant.
Type Locality.— Rio Damitas in the Dota
Mountains, San Jose, Costa Rica.
Type Material.— The male holotype, female
allotype, and 19 paratypes were taken at the
type locality on 18-11-64, 250 m elevation. No.
446, from an unidentified climbing vine, by
S. L. Wood. Four paratypes are from Playon,
Puntarenas, Costa Rica, 22-11-64, 50 m, Nos.
454 and 455, and three paratvpes are from Tur-
rialba, Cartago, 9-III-64, 700 m, Nos. 460, 458,
same host and collector.
The holotype, allotype, and paratypes are
in niv collection.
Dendrocrcmtilus fiilgidus, n. sp.
This species is distinguished from the closely
allied rnaurus (Blandford) by the discal gran-
ules on the pronotum, by the narrower elytral
declivity, by the more strongly impressed male
frons, and by the more coarsely punctured fe-
male frons, with less abundant vestiture.
Male.— Length 2.0 mm (paratypes 1.8-2.3
mm), 2.8 times as long as wide; color dark
reddish brown, elytra lighter.
Frons as in rnaurus except transverse im-
pression distinctly deeper, median summit on
vertex slightly higher. Pronotum as in rnaurus
except granules extend to impunctate median
line; posterolateral areas reticulate. Elytra as in
rnaurus except more slender, declivity narrower,
strial punctures much smaller, not as deep, in-
terstriae 2 more distinctK' impressed, surface
brightU' shining; granules as in maurus; de-
clivital setae on interstriae 1 and 2 very short,
less than half as long as on 3 or on disc.
Female.— Similar to male except frons plano-
convex, more coarsely punctured on lateral
thirds than in female rnaurus, vestiture fine,
long, slightly less abundant than in female
maurus.
Type Locality.— Volcan de Chiriqui, near
Cerro Punta, Chiriqui, Panama.
Type Material.— The male holotype, female
allotype, and 97 paratypes were taken at the
type locality on 11-1-64, 1800 m elevation, Nos.
378 (holotype and allotype), 408, and 418, from
a cucurbit vine, by S. L. Wood.
The holotype, allotype, and paratypes are in
my collection.
Dendrocranulus vicinus, n. sp.
This species is distinguished from the very
closely related schedli Wood by the less strong-
ly impressed male frons, by the more coarsely
26
Brigham Young University Science Bulletin
punctured, nongranulate female fions, witli a
tuft of abundant, fine, long, yellow hair, and
by the larger granules in the median area of
the pronotal disc.
Male (allotype).— Length 1.5 mm (para-
types 1.4-1.7 mm). 3.0 times as long a.s wide;
color yellowish to reddish brown.
Frons as in female schecUi. Pronotum as in
schedli except granules in median area of disc
slightly larger and evidently never associated
with punctures. Elytra as in schedli except
striae 1 and 2 and interstriae 2 usualh less
strongly impressed.
Female (holotype).— Similar to male except
frons more broadly convex, more closely punc-
tured, punctures not at all granulate, orna-
mented by a tuft of long, fine, abundant, )ellow
hair.
Type Locality.— La Lima, Cortez, Hondu-
ras.
Type Material.— The female holotype, male
allotype, and 20 paratypes were taken at the
type locality on 5-\'-64, 200 m elevation. No.
579, from Cayaponia microdonta, by S. L.
Wood. Four parat\pes are from La Ceiba, At-
lantida, Honduras,' 20, 29-V-49, at light, E. C.
Becker.
The holot\'pe, allotype, and parat\pis are
in my collection.
Dendrocranitlits rudis, n. sp.
This species is allied to cucurhitae (Le-
Conte), but it is distinguished by the slightly
larger rugae on the pronotal disc, with the
punctures obscure to obsolete, and by the less
deeply excavated male frons with the median
elevation on the vertex less well de\eloped.
Male.— Length 1.8 mm (paratypes 1.6-1.9
mm), 2.8 times as long as wide; color ver\'
dark brown.
Frons somewhat flattened, transversely im-
pressed from epistoma to upper level of eyes,
weakly ascending above; median elevation on
summit rather well developed, highest at its
dorsal extremity, descending below; surface
smooth with an occasional fine puncture or
minute granule; vestiturc sparse, hairlike.
Pronotum 1.2 times as long as wide; widest
just behind middle, sides rather strongly arcu-
ate on posterior two-thirds, rather narrowU-
rounded in front; anterior third rather coarsely
asperate, low, transverse crenulations continu-
ing to base, some crenulations in median area
associated with obscure punctures. X'estiture
fine, long, moderately abundant.
Elytra 1.7 times as long as wide, 1.5 times
;is long as pronotum; sides almost straight and
parallel on basal three-fourths, very broadly
rounded behind; striae not impressed, punctures
rather small, moderately impressed, spaced by
about two diameters of a puncture; interstriae
smooth, twice as wide as striae, equal ia size
and spacing to those of striae. Declivity steep,
broadly flattened; striae 1 and 2 feebly im-
pressed; interstriae 1 weakly elevated, 2 weakly
impressed, lateral areas about as high as suture;
intcrstrial punctures not granulate. X'estiture of
rows of fine, short, recumbent, strial h;iir, and
rows of erect intcrstrial bristles; each bristle
rather slender, about one and one-fourth times
as long as distance between rows, slightly
siiorter on declixital interstriae 1 and 2.
Female.— Similar to male except frons rather
coarsely punctate-granulate, granules mostly ar-
ranged in rugae, \'ertex devoid of median ele-
vation; declivity more nearly convex, impressions
and elevations obscure.
Type Locality.— Nineteen km E Carapan,
Michoacan, Mexico.
Type Material.— The male holotype, female
allotype, and 50 paratypes were taken at the
type locality on 18-\I-65, 2300 m elevation, No.
75, in cucurbit vine, by S. L. Wood. One para-
type is from 5 km W El Salto, Durango, Mex-
ico, 7-\'L65, No. 32, cucurbit vine, S. L. Wood.
The holotype, allotype, and paratypes are
in my collection.
Dendrocramdus confinis, n. sp.
This species is distinguished from the very
closel)' related cucurhitae (LeConte) by the
subreticulate pronotal surface between crenu-
lations on the disc and by the slightly shallower,
more extensive frontal excavation of the male.
Male.— Length 2.0 mm (paratypes 1.5-2.0
mm), 2.7 times as long as wide; color ver\' dark
reddish brown, pronotum black in some speci-
mens.
Frons as in cucurhitae but very slightly less
strongly excavated, impression extending higher
in lateral areas, median projection on vertex
ecjual in size. Pronotum as in cucurhitae except
discal punctures slightly larger, surface subreti-
culate, evidently less irregular. Elytra as in
cucurJntae except discal punctures \erA' slightlv
larger.
Female.— Similar to male except frons
broadly convex, a slight transverse impression
Biological Series, Vol. 19, No. 1 New Species of Amehic.'vn B.-vrk Beetles
27
just above epistoina; surface subreticulate,
punctures rather small, shallow, vestiture
sparse; decli\itv more evenh' convex, interstriae
2 not as strongly impressed.
Tyi'e Locality.— Volcan dc Chiriqiii, near
Cerro Punta, Chiri(jiii, Panama.
Type Material.— The male holotype, female
allotype, and 38 paratypes were taken at the
type' locality on 11-1-64, 1800 m. No. 378, from
a cucurbit vine, by S. L. Wood.
The hoIot\pe, allotype, and paratypes are
in my collection.
Ips liorealis Janieri, subsp. n.
This subspecies is distinguished from hore-
alis borealis Swaine by the almost evenly con-
vex female frons, which is sparseh' granulate
and fineh' punctured below the upper le\el of
the eyes, and by the transverse row of epistomal
granules. It is much more similar to b. thomasi
Hopping, but it is distinguished bv the shorter,
finer, less abundant \estiture of the female
frons, which is also less stronglv inflated, and
the granules average much smaller in size.
Female.— Length 3.5 mm (paratypes 3.0-3.8
mm), 2.6 times as long as wide; color v(tv dark
brown to almost black.
As in />. borealis except for sculpture and
ornamentation of female frons as cited in the
above diagnosis; entire frons with small, dis-
tinct punctures, on lower half intermixed with
several small granules.
Male.— Similar to female except frontal
punctures and granules much larger, more
numerous.
Type Locality.- One mile south of Browns-
ville, Lawrence County, South Dakota.
Type Material.— The female holotype, male
allotype, and 61 parat\pes were taken at the
type localitN' on lS-\T-68, No. 47, Picea <ihnica,
by S. L. Wood. Additional parat\pes were taken
as follows: 21 from 2 miles SW Lead, Lawrence
Co., South Dakota, 18-VI-68, No. 48, Picea
Ulauca. S. L. Wood; 7 from SE;4 86, T4N, R3E.
Black Hills, South Dakota, 17-MII-67, Picea
glauca, J. M. Schmidt; 5 from 6 miles SW Buf-
falo, Johnson Co., W\omin5, 20-M-68. No. 58,
Picea enfiehnanni, S. L. Wood; 6 from about 20
miles NW Pagosa Spring, Hinsdale Co., Colo-
rado, 28-\T-68, Picea engelmanni, W. G. Har-
wood.
The holotype, allotype, and paratypes are
in my collection.
Gnathophthorus artus, n. sp.
This species is distinguished from the five
previously named species by the smaller size,
by the simple declivity, and by the very broadly
impressed female frons with much more abun-
dant and longer frontal \estiture.
Female.— Length 1.05 mm (paratypes 0.9-
1.05 mm), 4.3 times as long as wide; color of
pronotum and declivits rather dark vellowish
brown, basal area of el\ tra pale \ellowish
brown.
Frons rather shallowly, subcireularlv con-
cave from eve to eye, from epistoma to vertex;
margins of upper half bearing a dense row of
verv long hair, lower area appearing spongy
and bearing a few moderately long setae; sur-
face largelv concealed by \Tstiture. Antennae
small; club distinctlv longer than scape, an-
terior face glabrous, clearly marked bv two al-
most straiglit, parth' septate sutures.
Pronotum 1.84 times as long as wide; sub-
quadrate, sides straight and parallel more than
three-fourths of their lengtli, anterior margin
feebly arcuate, weakh serrate; anterior tenth
weakly declivous, anterior third finely asperate;
posterior areas smooth, shining, minutelv, shal-
lowlv, ratlicr closelv punctured. ClalHous.
Elvtra 2.4 times as long as wide, 1.4 times
as long as pronotum; sides straight and parallel
on basal three-fourths, moderately rounded be-
hind; striae not impressed, punctures minute;
interstriae four to five times as wide as striae,
punctures similar in size and spacing to those
of striae. Declivity moderately steep, narrow;
sutural interstriae distinctly elevated, convex;
punctures of striae 2 and 3 fine, distinct, rather
widelv spaced on 3; interstriae 2 broad, flat,
smooth, shining, with only four or five minute
punctures, 3 not elevated or serrate. Vestiture
confined to declivit)', sparse; consisting of
rather fine interstrial bristles on interstriae 3
and on lateral areas, smaller, similar setae on
interstriae 1.
Male —Similar to female except frons mostly
convex, a short, transverse carina at upper level
of eyes and immediately below this a small,
shallow concave area on median fourth, sur-
face shining, rather closely, coarsely punctured,
vestiture of sparse, short hair; ten serrations on
anterior margin of pronotum slightly larger;
elytral declivity, shallowly, broadly bisulcate,
interstriae 3 veiy feebly elevated.
Type Locality.- About 260 km N Xa\an-
tina, Mato Grosso, Brasil (12°49' S 4r46' W).
Type Material.— The female holotype, male
allotype, and 12 parat\pes were taken at the
28
type locality on X-1968, No. F41, by R. A.
Beaver. One paratype bears the same data ex-
cept 23-IX-68, No. 166.
The holotype, allotype, and five paratvpes
are in the British Museum (Natural History),
two paratypes are in the Museu de Zoologia,
Universidade de Sao Paulo, and si\ paratypes
are in my collection.
Dn/ocoetoides Hopkins
Dn/ocoetoide.s Hopkins, 1915, U.S. Dept. Agric. Rept.
99:52 (Type-species: Dnjocoetoides guatematensis
Hopkins = Xi//efcori/.s capuciims Eichhoff)
This neotropical genus \sas placed in syn-
onymy by Schedl (1952, Ent. Bliitt. 47-48:161)
and, presumably because of the lack of material
or interest, it has not been seriously reconsid-
ered since then. The type-species clearly is a
representative of the Xyleborini. Contrary to
the action of Schedl of placing virtually all rep-
resentatives of this tribe in one enormous genus.
Xijleborus, I prefer to elevate certain clearly
marked groups to generic rank in order to make
them more meaningful and to make Xijlehoms
less unwieldy.
Drijocoetoidcs has the prothoraeic tibia in-
flated and tuberculate on its posterior face; the
antennal club is rather strongly flattened, with
the basal area slightly thickened and moder-
ately corneous, two clearly marked, almost
straight sutures are on the basal half of the
anterior face and both continue to and arc
clearly marked on the distal half of the pos-
terior face; in most species the eyes are rather
large and coarsely faceted; the scutellum is
flat and moderately large: the pronotum is
rather stout, about as long as wide, and armed
bv several serrations on the anterior margin;
and till' elvtrul disc and declix'ity usually ha\e
contrasting sculpture.
In addition to Xijleborus capuciiuis Eichhoff,
the following species are transferred into Dry-
ocoetoides: Xiikhorus monachtis Blandford,
Bostrichus fhivtis FalMieius. Bosfrkhus cristuttis
Fabricius (=Xtilchonis soUtarius Hagedorn, X.
urichi Eggers, X. crenattis Eggers), and Xyle-
1)orus pseudosnlitaiius Eggers. Seven pre\iously
undescribed species are added below.
Driiocoetoides monachus (Blandford), n. comb.
Xxjleborus monachus BUindford, 1898, Biol. Centr. Amer.,
Coleopt. 4(6):204 ( Syntypes, males; Cerro Zunil,
Guatemala; British Mus. Nat. Hist.).
This species w as n;\med from two male syn-
types and has not been reported in the litera-
ture since then.
Brigham Young University Science Bulletin
iMJur females were collected at P;ui]in, Es-
quintla, Guatemala, 19-V-64, 300 m, No. 584,
from Inga branches; 21 females from Rancho
Grande, Pittier National Park, Aragua, Vene-
zuela, 9-IV-70, 1100 m, No. 408, tree seedling;
one female from 24 km E Barbosa, Antioquia,
Colombia, LS-\TI-70, 1200 m, No. 696, from a
Rubiaceae sapling; all were taken by me. The
Guatemalan specimens and several males of the
very closely related capucinus (Eichhoff) were
compared to both of Blandford's male syn-
types. In general fonn and sculpture the types
(2.7-2.9 mm) were exactly ;\s the male capu-
cinus (2.3-2.5 mm) except for the body size and
det;iils of the elytral structure. The el)tral
structure and larger size conformed to that of
the Guatemalan females. Since there are no
other known species in Guatemala with which
moiochus could be confused, I am confident
the association is correct even though males
were not taken in my series.
Females of this species differ from those
of capucinus by the larger size (2.8-3.2 mm,
compared to 2.2-2.6 mm for capucinus), by the
longer elvtral disc ( 50 percent of elytral length,
compared to 40 percent), by the more shining
elytral declivitv with some reticulation in the
posterolateral areas, and by the more nearly
uniseriate interstrial granules on the elytral de-
cli\ity.
Dn/ocoetoides verrucosus, n. sp.
This species is distinguished from flaws
(Fabricius) by the more slender form, by the
smaller strial punctures, by the rugose-reticu-
late, dull, dark, elytral declivity, and by the
verv different declivital sculpture.
Fi-MAi.E.-Length 2.5 mm (paratype 2.4
mm), 3.2 times ;is long ;is wide; color reddish
brown, elytra much dl^rker.
Frons and pronotum as in flavus except
pronotum 1.2 times as long as wide.
Elvtra 1.9 times as long as wide, 1.6 times
as l(Hig as pronotum; sides almost straight and
parallel on basal two-thirds, tapered then nar-
rowK' rounded behind; striae not impressed,
punctures \'ery small, very shallow, spaced
within a row by two diameters of a puncture;
interstriae smooth, shining, about six times as
wide as striae, punctures almost as large as
striae, uniseriate except slightly confustxl on 2.
Declivitv occupying posterior tliird of elytral
length, convex; surface minutely rugose-reticu-
late; striae feebly impressed, punctures slightly
larger than on disc, feebly impressed; interstriae
uniseriately, rather coarsely tuberculate, except
Biological Series, Vol. 19, No. 1 New Specie.s of American Bauk Beetles
29
at base and lower half of sutiual intcr.striae;
sutural interstriae from just above middle to
just before apex very strongly, irregularly ele-
vated, elevation as high as wide; interstriae 2
and 3 slightly sinuate to accommodate sutural
elevation. \'estiture confined to declivitv, con-
sisting of minute strial hair and confused, semi-
recumbent interstrial setae, each slightly shorter
than width of an interstriae.
Type Locality.— Thirty km E Palmar, Boli-
var, Venezuela.
Type Material.— The female holotype and
one female parat>pe were taken at the type
locality on 12-\T-70, 200 m, No. 579, from a
tree known locally as Rosada (Moraceae), by S.
L. Wood. One female parat\ pe is from 3 km NE
Creele, Barinas, Venezuela, 18-XII-69, 150 m.
No. 203, Inga, S. L. Wood.
The holotype and parat\pes are in m\- col-
lection.
Dnjocoetoides pileatu.s, n. sp.
This species is distinguished from the closeh'
allied capucinus (Eichhoff) by the much larger
size, by the much shorter clytral disc, by the
more gradual elytral decli\'it\, and by the ab-
sence of strial punctures on the declivity.
Female.— Length 3.2 mm (paratypes 3.2-
3.3 mm), 2.1 times as long as wide; color almost
black.
Frons and pronotum as in capucinus except
surface of disc less strongly reticulate, punc-
tures deeper.
Elvtra LI times as long as wide, L3 times
as long as pronotum; disc occupying 30 percent
of elytral length, transition from disc to decliv-
ity rather abrupt; striae not e\ident, surface
smooth, shining, punctures varialile, small to
very small, close, confused. Decli\'ity gradual,
convex; surface minuteh' rugulose, dull; striae
and strial punctures entirelv obsolete, numer-
ous small, roimded, confused, setiferous gran-
ules; subapieal posterolateral margin acutely,
subcrenulately elevated from suture to position
of interstriae 8. Vestiture confined to decli\it\',
consisting of moderately abundant, short hair.
Type Locality.— Fortv km SE Socopo, Bar-
inas, Venezuela.
Type Material.— The female holotype and
one female parat^â– pe were taken at the t\'pe
locality on 25-L70, L50 m elevation, No. 263,
in Inga limb, bv S. L. Wood; four female para-
types are from 10 km SE Miri, Barinas, Wne-
zuela, 8-II-70, 150 m elevation, No. 295, in Inga
limbs, bv S. L. Wood; one female paratvpe is
from 17 km SE Miri, Barinas, Venezuela, 17-
.\II-69, 150 m elevation. No. 197, from a palm
bole 15 cm in diameter, by S. L. Wood.
The holotype and paratypes are in my col-
lection.
Drijocoetoides vehttinus, n. sp.
This species is placed in a species group
with pseudosolitarius. although the relationship
is not close. It is distinguished by the larger
size, by the stouter body form, and b)' the
steeper elvtral declivity that has different sculp-
ture.
Fe.male.— Length 2.8 mm (paratypes 2.7-
2.9 mm), 2.4 times as long as wide; color red-
dish brown.
Frons and pronotum as in j)sctidosolitorius;
pronotum 1.1 times as long as wide.
Elytra 1.4 times as long as wide, 1.3 times
as long as pronotum; disc occupying basal two-
thirds of elytra; striae not impressed except 1
slightlv near decli\'itv. punctures very small,
shallow, spaced witliin row bv three diameters
of a puncture; interstriae smooth, shining, about
six times as wide as striae, punctures minute,
half as large as those of striae, confused; de-
clivital sculpture commencing on posterior third
of disc. Declivity steep, broadly convex; sur-
face minuteh- rugulose; strial punctures small
but larger and deeper than on disc; interstriae
with abundant, rather coarse, pointed, confused
tubercles; lower half of sutural interstriae mod-
eratelv protuberant; subapieal posterolateral
margin subacutelv subcrenulate from suture to
interstriae 8. \'estiture of fine, abundant, mod-
eratelv long hair on disc and declivity.
Male— Length 2.8 mm; essentially as in
female, but \\ ith all characters less perfectly
formed except pronotum 1.3 times as long as
wide, excavated as in male perebeae ( Ferrari )
except median tubercle on anterior margin
\erv broad.
Type Locality.— Thirtv km E Palmar, Boli-
var, \'enezucla.
Type M.^terial.- The female holotype, male
allot\pe, and 21 female paratypes were taken
at the type locality on 12-\T'-70, 200 m. No.
538, from the bole 30 cm in diameter of a tree
localK- named Pandanga, by S. L. Wood.
The holotvpe, allotvpe, and paratypes are
in mv collection.
Dniocoetoides rusticus, n. sp.
This species is distinguished from velutinus
U'ood bv the larger size, bv the confused punc-
30
BmcuAM VouNc University Science Bulletin
turcs on the discal interstriae, by the more
gradual elytral declivity, with much finer inter-
strial tubercles, and bv the much coarser ves-
titure.
Female— Length 3.5 mm (estimated, elvtra
spread), about 2.4 times as long as wide; color
reddish brown.
Frons as in vehitinus but much narrower
due to enlarged eyes. Pronotum as in veltitiwis
except punctures on disc closer.
Elytra about 1.4 times as long as wide;
elytral disc as in vehitinus except all punctures
deeper, interstrial punctures more numerous,
confused, minute, irregular surface lines pres-
ent. Declivity much as in vehitintis, surface
rugulose, more ex'enlv, more gradually coincx;
strial punctures larger than in vehttinus, inter-
strial tubercles more numerous, much smaller,
weakly protubrant toward apex of suture; sub-
apical ventrolateral margin acutely ele\ated
from suture toward but becoming obsolete on
interstriae S, not connected to but continuing
toward base of declivity as a row of subcrenu-
late tubercles. Vestiture confined to decli\it\
and sides; consisting of \'erv stout, short
bristles on central and lower parts of declivity,
some setae on lateral and upper parts of de-
clivity at least twice as long and tapered.
Type Locality.— Thirty km E Palmar, Boli-
var, Venezuela.
Type Material.— The unique female holo-
type was taken at the type localit\ on 12-\I-70,
100 m ele\ation, \o. 5S2, from an unidentified
tree limb, b\- S. L. Wood.
The holotvpe is in my collection.
Dn/ococtoich's scvcnis. n. sp.
Among known forms this species is allied
to rusfictis Wood, ])ut it is distinguished b\' the
large size, by the subijuadrate pronotum, and
by the very different sculpture of the elytral
declivity.
Female.— Length 5.0 mm, 2.6 times as long
as wide; color reddish brown; head and pio-
thorax detached and mounted separately on the
same paper point as boch .
Frons as in rustictis but narrower, 1.8 times
as wide as width of eve. Pronotum 1.0 times as
long as wide; subquadrate; sculpture essentially
as in rustictis.
Ehtra 1.7 times as long as wide, 1.6 times
as long as pronotum; sides straight and parallel
on basal three-fourths, obtusely subangulate be-
hind; disc occupying basal three-fourths; b;isal
half of disc as in rusticus: posterior half of disc
with surface smooth, shining, punctures re-
placed by rounded granules. Declivity steep,
broadly eonx'ex; a large, low, conspicuous,
rounded prominence at basal margin on inter-
striae 2; decli\ital surface smooth, almost shin-
ing; sutural interstriae slightly elevated on
lower half, impressed toward base; area of striae
1 impressed to middle, are;i lateral to striae 1 on
upper half gradualh- elexated toward prom-
inence at base on interstriae 2; interstriae 1
with small confused tubercles to apex, others
with a few tubercles on basal half only; strial
punctures much larger and slighth' dei'per tlian
on disc; subapical \entroIateral margin acutely.
su])crenulately elevated from apex to apex of
interstriae S. \'<\stiture confined to posterior
half of ehtra; consisting of fine long hair ex-
et"pt mueli slioiter on lower luilf of di'cli\itv.
Type Locality.— Thirt\ km
\ar, \'enezuela.
Boli
Type Material.— Thi' unicjue female holo-
typ(> was taken at the type loealit\- on 12-\'I-70,
200 m. No. 54S, from Esclincilcra suliislandu-
losa. bv S. L. \\'ood.
rile ]iol()t\pe is in m\' collection.
Dnjocoetoides insculptis, u. sp.
This species is distinguislied from tlie rather
remotely related iclulintis Wood b\' the larger
size, b\' tlie strongK impressed deeli\ital striae,
by the much less abundant elvtral \'estiture,
and b\' other charactcTS.
P'emale— Length 4.2 nun ( parat\pes 4.0-
4.2 mm). 2.6 times as long as wide; color red-
dish brown.
Frons esseutial!\' as vehitinus except not as
wide due to enlarged e)es.
Pronotum 1.2 times as long as wide; about
;is in vehitinus except sides almost straight and
pai;illel on basal two-thirds, broadly rounded in
front; antt'rior margin anned bv 14 low serra-
tions; disc shining, lateral areas reticulate, punc-
tures very small, deep, moderately abundant.
Elvtra 1.6 times as long as wide, 1.5 times
as long as pronotum; disc occupying basal
three-fourtlis; sides straight and parallel on basal
three-fourths, rather narrowly rounded behind;
striae feeblv impressed on basal half, punctures
small, rather deep; interstriae on basal half of
disc smooth, sliining, al)out three times 'as wide
as striae, punctures \-ery small, confused; de-
clivital sculpture commencing just behind
middle of disc. Decli\it\- confined to posterior
fourth of elytra, steep, conxex; strongly reticu-
late, dull; striae ver\' deeply impressed, punc-
Biological Series, Vol. 19. No. 1 New Species ok A.\ieiuc.\n Bahk Beetles
31
tures slightly larger tliaii on disc, shalluw; inter-
.striae rather strongly convex, nniscriately rather
coarsely serrate, tubercles much smaller on
lower half of declivity; sutural interstriae mod-
erately protuberant on lower half of declivity.
Vestiture confined to reticulate area; consisting
of fine, short, strial hair and an occasional simi-
lar interstrial hair, and rows of coarse, erect
bristles arising from posterior margins of ser-
rations, each bristle about as long as distance
between rows.
Type Locality.— Eight km S Colonia ( near
Buenaventura), Valle de Cauca, Colombia.
Type Material.— The female holotype and
eight female paratypes were taken at the type
locality on 9-VII-70, 30 m elevation. No. 605,
in limbs of Htimirisfnivi excehum, b)' S. L.
Wood.
The holotype and paratypes are in my col-
lection.
Dnjocoetoidcs iiulolatus, n. sp.
This species is distinguislied from the allied
insctilpttis Wood by the slightly smaller size,
by the more nearly suhquadrate pronotum, by
the unimpressed d('cli\ital striae, b\- the finer
interstrial tubercles, and b\' tlie vestiture.
Female.— Length 3.7 mm, 2.6 times as long
as wide; col(;r reddisli l)rown.
Frons as in insculptus but narrower. Pro-
notum subquadrate, serrations on anterior mar-
gin poorly developed; sculpture essentially as in
irisciilptiis.
Elytra L5 times as long as wide, L4 times
as long as pronotum; disc occup\ing basal half
of elytral length; striae not impressed, punc-
tures very small, spaced within a row by one
or two diameters of a puncture; interstriae
smooth, shining, with shallow, irregular lines,
punctures about equal in size to those of striae,
confused. Declivity very gradual on its basal
half, steep and convex on its posterior half;
entire surface closely, finely rugose, dull; striae
not impressed, punctures twice as large as on
disc, very shallow; interstrial punctures replaced
by fine, confused tubercles, slightly larger on
basal half of convex area; sutural interstriae
weakly protuberant toward apex; subapical pos-
terolateral margin acutely costate near suture,
becoming subcrenulate toward interstriae 8.
Vestiture confined to sides and decli\it\', con-
sisting of fine, long, moderately al^undant, con-
fused, interstrial hair; minute strial hair also
present on declivity.
Type Locality.— Thirty km E Palmar, Boli-
var, Venezuela.
Type Material.— The unique female holo-
type was taken at the type locality on 12-\'I-70,
200 111, No. 578, from Parinari cxcelsa, by S. L.
Wood.
The holotype is in iii\- collection.
Sdinpsonitis expuhtis. n. sp.
This species apparently is more closely allied
to detnictus Wood than to other described spe-
cies, but it is readily distinguished by the longi-
tudinally concave elytral declivity and by the
absence of declivital spines.
Female.— Length 5.1 mm, 3.4 times as long
as wide; color reddish brown.
Frons, pronotum and elytral disc as in de-
tractus. Elytral dechvity slightly longer than
disc, its surface reticulate; base of declivity
rather abruptly impressed, basal and lateral
margins continuously, finely serrate from inter-
striae 2 to near sutural apex; face of declivity
longitudinally concave, broadly flattened to ser-
rate margins, suture rather strongK' ele\ated
particularly on lower half; interstriae 3 weakly
ele\ated on lower half. W'stiture of fine, mod-
erateh' long, rather al)und;mt liair; onl\' slightly
longer on declivity.
Type Locality.— Eight km S Colonia (near
Buenaventura), \'alle de Cauca, Colombia.
Type Material.— The female holotype was
taken at the type locality on 9-\'II-70, at 30 m
ele\ation. No. 619, from a Poutcria branch, bv
S. L. Wood.
The holotype is in mv collection.
^anipsonius dctravtus. n. sp.
The large size and the presence of one pair
of long slender spines near the apex of declivi-
tal interstriae 3 distinguish this species from
other representatives of the genus.
Female.— Length 5.6 mm (paratype 5.8
mm), 3.6 times as long as wide; color reddish
brown.
Frons narrow, broadly convex; siuface reti-
cuhite, \\ ith small, low, rather abundant, shining
granules; vestiture of sparse, fine, long hair.
E\e large, emarginate, very coarsely faceted, 1.9
times as long as wide. Antennal club subcir-
eular, rather strongly flattened, sutures 1 and 2
rather strongh' procur\ed, 2 reaching slightb
beyond middle; one strongly procurved suture
near apex on posterior surface.
Pronotum 1.4 times as long as wide; sides
feebly arcuate and subparallel on posterior two-
thirds, narrowly produced on anterior third to
32
BnicnAM Young Univkbsiiv .Sf:iENf:E Bulletin
two very coarse, close serrations arming an-
terior margin, two pairs of small serrations lat-
eral to major ones; summit indefinite, about
one-third length from anterior margin; anterior
third moderately asperate; posterior areas
smooth, shining, minutely, rather sparsely punc-
tured; lateral margins abrupt. Vestiture of
sparse, fine hair, slightly coarser and longer on
asperate area.
Elytra 2.1 times as long as wide, 1.5 times
as long as pronotum; sides almost straight and
parallel on basal three-fourths, slightly nar-
rowed, then truncate on median two-tliirds;
discal striae not impressed, punctures ver\'
small, distinct; interstriae smooth, shining, about
four times as wide as striae, punctures almost
uniseriate, minute. Declivity occupying poste-
rior 46 percent of ehtral length, gradual basal
margin abruptly impressed and irregularly,
finely serrate from interstriae 1 to posterolateral
apical angle; surface reticulate; sutural inter-
striae not elevated or armed; interstriae .3 pos-
terior to middle broadly elevated and slightK'
higher than suture, decending to moderate im-
pression just mcsad of serrate lateral margin;
armed just before apex of interstriae .3 by a
pair of very large, subcvlindrical spines, each
four times as high as wide, equal in length to
discal distance from suture to striae 4. N^estiture
of fine, moderately long hair on disc and sides,
very much longer and slighth- more abundant
on declivit\'.
Type Locality.— Madden Forest, Canal
Zone, Panama.
Tyi'e Material.— The female holotvpe and
one female paratype were taken at the type
locality on 2-T-64, 70 m elevation. No. ,367, from
a tree limb, hv S. L. \\'()od.
The holotype and paratype are in my cf)l-
lection.
Sampsonius usuri)afus. n. sp.
This species is distinguished from tlie closeK
allied dampfi Schedl b\' the larger size, bv the
longer elytral dt'clivity, l)y the declivital suture
being lower than the lateral convexities, and by
the much smaller, contiguous spines near the
sutural apex on the el\ tral declivit\'.
Female.— Length 4.1 mm (paratypes: fe-
males 4.0-4.4 mm; males 2.S-3.3 mm), 3.8 times
as long as wide; color reddish l^rown.
Frons as in detractus Wood except slightly
narrower. Pronotum as in detractus.
Elytra 2.2 times as long as wide; outline as
in dectacttts except tapered posteriorly, subtnm-
cate on less than median half; disc and general
contour of declivity as in detractus except basal
margin gradual, di'nticles smaller, mori'
scattered; apex of interstriae 3 entirely un-
armed; sutural interstriae anned by contiguous,
subapical, pointed processes about as high as
their longitudinal base, height about equal to
width of discal interstriae, positioned as in
dampfi but much smaller. Vestiture as in de-
tractus.
Male.— Dwarfed, head concealed by pro-
longation of pronotum; pronotum longer than
cKtra, 1.7 times ;ls long as wide, its anterior
three-fifths broadly, rather deeply concave, an-
terior margin narrowly biemarginate gi\ing ap-
pearance of a median and two lateral denta-
tions; elytra similar to female but characters
poorly formed, more convex, devoid of subapi-
cal spine.
Type Locality.— Turrialba, Caitago, Costa
Rica.
Type Material —Tlu' female hoIot\pe, male
allotype, and 23 paratypes were collected at
the type locality on 9-111-64, at 700 in elevation,
No. 460B, from a cut tree seedling, by S. L.
Wood. Other paratypes were taken in Costa
Rica as follows: two from San Ignacio de
Acosta, 5-\'II-6.3, 1500 m. No. .38, tree seedling,
S. L. Wood; three from Finca Cromaco on Rio
Coto Brus, Puntarenas, 14-V1I-63, 500 m. No.
76, tree seedling, S. L. Wood; one from Rincon
de Osa, Puntarenas, ll-VHI-66, 30 m. No. 6S.
tree branch. S. L. ^\'ood. One paratype is from
El Hato del N'olean, Cliirif(iii, Panama, 11-1-64,
2800 m. No. 374, tree- branch, S. L. Wood.
The holotype, allot\pe, and paratopes are
in uw eolleetiou.
Xi/Ichonis (Thcohonis) pristis. n. sp.
The antennal club of tliis species clearly is
as in theohromae Hopkins, but tiie relationship
is nf)t close. It differs from theohromae by the
more slender form, by the less strongly serrate
antiTior margin of the pronotum, bv the more
broadly convex elytral declivity, and by the
different sculpture of the elytral declivity.
Female.— Length 1.8 mm (femak' paratypes
1.7-2.0 mm), 2.6 times as long as wide; color
very dark brown.
Frons broadly convex, a weak, narrow,
transverse impression just above epistonla; sur-
face strongK reticulate, punctures sparse, ob-
scure, coarse; \'estiture of sparse, fine, incon-
spicuous, long hair.
Pronotum 1.04 times as long as wide; essen-
tially as in theohromae except surface more
Biological Series, Vol. 19, No. 1 New Species of Amehic.\n Bahk Beetles
33
.strongh' reticulate, including in anterior area
between asperities, and anterior margin weakly
serrate. Vestiture of fine, rather short hair on
sides and asperate area, disc almost glabrous.
Elytra 1.3 times as long as wide; sides al-
most straight and parallel on basal three-
fourths, rather narrowly rounded behind, ser-
rate near weakly, narrowly emarginate suture;
striae not impressed, punctures minute, shallow,
distinct, spaced within a row by three to four
diameters of a puncture; interstriae smooth,
shining, about eight times as wide as striae,
punctures uniscriate, almost indistinguishable
from those of striae. Declivity confined to pos-
terior fourth, steep, broadlv convex; stri;ie 1
weakly, others feebly impressed, punctures
slightly larger, deeper and much closer than on
disc; interstriae with punctures closer and uni-
seriately granulate, granules often oljsolete near
apex; sutlue narrowh' emarginate, margin witli
a row of two to four coarse cusps near suture,
this row continued along margin as four to eight
somewhat smaller granules, margin rather nar-
rowly rounded to interstriae S. Vestiture con-
fined to sides and declivity; on declivity con-
sisting of rows of fine hair; strial hair short;
interstrial hair two to two and one-half times
as long as distance between rows.
Male.— Length 1.5-1.8 mm; essentially as in
female, but with characters more poorlv formed,
and pronotum 1.4 times as long as wide and
essentially as in male theohromae.
Type Locality.— Tapanti, Cartago, Costa
Hica.
Type Material.— The female holotvpe and
tliree female paratypes wxtc taken at the tvpe
locality- on 2-\TI-63, 1300 m. No. 9. from
Miconia, by S. L. Wood; three paratypes are
from the same locality, 17-1X-63, No. 184, from
Miconia caudata 10 cm in diameter. Four fe-
male paratypes were taken at \'olcan, Punta-
renas, Costa Rica, ll-XIl-63, 1000 m. No. 305
in "huarumo" and No. 308 in a tree limb, bv
S. L. ^^'ood; six additional paratvpes bear this
same data c^xcept they arc No. 308, from a tree
limb. The male allotvpe and 44 paratypes were
taken 6 km S San \'ito, Puntarenas, Costa Rica,
13-21-III-67, from a dead, standing tree.
The holotype, allotvpe, and paratypes are
in my collection.
Xylehorus (Theohorus) micarius. n. sp.
This species is closely related to pristis
Wood, but it is distinguished by the slightly
larger, more closely spaced strial punctures, by
the more gradual, more broadh' arched elvtral
declivity, by the smaller cusps near the suture
on the ventrolateral margin of the declivitv,
and bv the very stout, short, declivital setae.
Female.— Length 1.8 mm (female paratypes
1.8-2.0 mm), 2.7 times as long as wide; color ver>'
dark brown.
Frons and pronotum as in pristis except pro-
notum outline somewhat more subquadratc and
1.1 times as long as wide.
Elytra 1.5 times as long as wide; outline es-
sentially as in pristis, disc as in pristis except
strial punctures slightlv larger, spaced within a
row by two to three times diameter of a punc-
ture; interstriae about five to six times as wide
as striae, punctures uniseriate, distinctly smaller
than those of striae. Declivity occup\ing 50 per-
cent of strial length, more gradual and more
broadly convex than in pristis; sculpture as in
pristis except posterolateral margin subacute, its
summit rather weaklv crenulatc from weak
sutural emargination to interstriae 7. Vestiture
of rows of strial and interstrial setae, finer and
shorter on disc; declivital strial iiair verv fine
and short, interstrial setae coarse, blunt, each
about one to one and one-half times as long as
distance between rows.
Type Locality.— Cuapiles, Limon, Costa
Rica.
Type Material.— The female holot\pe and
seven female paratypes were taken at the tvpe
locality on 22-\TI-66, 100 m elevation. No. 120.
from a branch of Cordia sp. 5 cm in diameter,
by S. L. Wood; three female paratypes bear the
same data except No. 117 from a palm log. One
female parat\pe was taken at Tapanti, Cartago,
Costa Rica, i7-I.\-63, 1.300 m elevation. No. 182,
from a tree branch, by S. L. Wood.
The holotvpe and paratypes are in my col-
lection.
Xtilcl>orus hicornitttis. n. sp.
This species is \er\ closely related to ehenus
Wood, but it is distinguished by the larger aver-
age size, bv the more strongly impressed upper
half of the declivitv. bv the different arrange-
ment of minor elvtral denticles, and by the
larger, longer major declivital spine.
Female.— Length 3.8 mm (paratypes 3.6-4.0
mm), 2.0 times as long as wide; color dark
brown to black.
Frons and pronotum as in ehenus. Elytra
as in ehemis except upper third of decli\'ity
more strongly impressed; denticles on interstriae
3 at base of declivity larger, 2-6 in number ( 2 in
34
Brigham Young Uni\ ersity Sc:iENf:E Bulletin
ehenus), extending posterior to level of spine
of interstriae 2; major spine slightly larger, its
basal area less extensive; punctures on declivital
face more sharply defined, slightly larger, en-
tire surface smooth, shining (some impressed
points at base and extreme lateral areas in
ehenus).
Male.— Similar to male ehenus except lateral
margin of declivity aimed by four spines of
almost equal size on left side, six spines ( ab-
nonnal?) on right side.
Type Locality.— Seventeen km SK Miri, Bar-
inas, Venezuela.
Tyi'k Material.— The ftniale holot\pe and
three female paratypes were taken at the type
locality on 17-XII-69, 150 m elevation, \o. 195.
in Protium sp., by S. L. Wood; the male allo-
type, one male and 23 female paratypes are
from 30 km N Canon Zancudo, Zulia, Vene-
zuela, 4-IV-70, 10 111, No. 515, from a tree sap-
ling, by S. L. Wood. Other paratypes were
taken in Venezuela as follows: 7 from 40 km
E Canton, Barinas, S-III-70, 70 m, tree seed-
lings; 13 from 40 km SE and 7 km NW of
Socopo, Barinas, 25-1-70, 13-11-70, from hiLia.
Hirtolla friandru, Rheedia madruno, Nerlandra
sp., and Protium; 3 from 20 km SW El \'igia,
Merida, 21-XI-69, 12-VI-70, lO-XII-69, from var-
ious hosts; and 1 from Haneho CIrande, Aragua, 9-
IV-70, 1100 m, tree sapling. Tliree parat\pes
are from Campo Capote, 27 km NE Mont()\a,
Santander, Colombia, 2-VII-70, 150 m, tree
sapling. One paratype is from .30 km E Palmar,
Bolivar, 12-\T-70, 200 m, Bronnia sp. All were
taken by me.
The liolot\pe, all()t\pe, and parat\pes are
in my collection.
Xi/leI)orus crinilulus. n. sp.
Although not closely related, this species is
distinguished from the allied inicariu.1 ^^'ood
by the larger size, by the more closely, more
deeply punctured pronotal disc, by the confused
discal interstrial pimctures, and b\' the steeper
elytral declivity that has different sculpture.
Female.— Length 2.2 mm (paratypes 1.9-2.3
mm), 2.3 times as long as wide; color rather
dark reddish brown.
Frons as in micarius except with moderately
numerous, small, rounded granules. Pronotum
as in micarius except serrations on anterior
margin larger and punctures on disc more
numerous and deeper.
Elytra 1..3 times as long as \\ide, 1.2 times
as long as pronotum; striae not impressed, punc-
tures small, rather shallow, spaced within a row
by one or two times diameter of a puncture;
interstriae four to five times as wide as striae,
shining, with irregular lines, punctures fine,
moderately deep, confused ou basal two-thirds,
uniseriate toward declivity. Declivity steep,
broadly con\ex, occupying 42 percent of elytral
length; strial punctures almost twice as large
as on disc; interstriae imiseriately tuberculate,
tubercles rather widely spaced, pointed, mod-
erateb fine except rather coarse on lower half
of 1; suture slightly produced just before apex;
subapical posterolateral margin acutely, sub-
crenulatelv elevated from suture to interstriae
8. W'stiture hairlike, rather abundant, of vari-
able length, mostly rather short.
Type Locality.— Forty km SE Socopo, Bar-
inas, Venezuela.
Type Matehial.— The female holotspe and
17 female paratypes were collected at the type
locality on 2.5-1-70, 150 m elevation, No. 280,
from a dead branch of llirtella friandra 3-5 cm
in diameter, b\' S. L. Wood, One paratype was
taken at Barro Colorado Island, Canal Zone,
Panama, 27X11-63, 70 in. No. .345, from a tree
liranch, by S. L. Wood; and two paratypes are
labeled Fort Chnton, Canal Zone, Panama, 22-
XII-63, .30 m. No'. 317, tree branch, S. L. Wood.
The holot\'pe and paratypes are in my col-
lection.
Xi/lclionis pcinduhis, n. sp.
This rather common species has been incor-
rectly identified by Sclicdl and others as var-
ians (Fabricius). It differs from various by the
smaller average size, by tlie steeper elytral de-
clivity, by the flatter lower half of the declivity
with its ventrolateral margin subacutely ele-
vated and its face irregular and closely, coarsely
punctured.
Female.— Length 2.7 mm (paratypes 2.7-3.0
mm), 2.0 times as long as wide; color dark
brown, almost black.
Frons and pronotum as in pereheae ( Fer-
rari) except each minute puncture on pronotal
disc with a minute, smooth, shining spot on its
posterior margin.
Elytral 1.0 times as long as wide, 1.06 times
as long as pronotum; disc limited to basal half;
disc as in pereheae except strial punctures
slightK- smaller. Decli\ity abruptly impressed at
base, steep, rather broadly, deepK excavated on
upper half; lower half more broadly flattened
and with its abrupt lateral margin weakly ele-
vated; basal margin abrupt, aimed by about 4-8
small, pointed denticles scattered from inter-
Biological Series, Vol. 19, No. 1 New Species of American Bark Beetles
35
.striae 1-3; margin at interstriae 3 bearing large,
stout, pointed conical spine one-fourth declivi-
tal length from basal margin, a large setiferous
pore just below its ape.x; interstriae 4 with
four to six small, pointed denticles on and
just before declivital margin, 5 with similar
denticles extending along margin to or near
sutural apex; declivital face closely, deeply,
rather coarsely punctured, surface subshining,
somewhat irregular. Vestiture limited to stout,
scalelike bristles on margin and sides of de-
clivity.
Type Locality.— Fort Clayton, Canal Zone,
Panama.
Ty'I'E Material.— The female hol()t\pe and
nine female paratypes were taken at the tvpe
locality on 22-XII-63, 30 m elevation, No. 320.
from a broken tree limb, by S. L. Wood. Other
female parat\pes were taken as follows: 3 at
Madden Forest, Canal Zone, Panama, 2-1-64, 70
m, Nos. 364, 367, tree limb, S. L. Wood; 1 at
Limon Ba\-, Canal Zone. .30-XII-63, 5 m. No. 354.
tree branch, S. L. Wood; 2 at Dominical, Punta-
renas, Costa Rica, 9-XII-63, 3 m. No. 301, tree
branch, S. L. Wood; 3 at Boston, Limon, Costa
Rica, LX-64, Theohroma cacao, J. L. Saunders;
and 1 at Finca La Lola, Limon, Costa Rica,
\Tn-63, Theohroma cacao, J. L. Saunders.
The holotype, allotype, and paratopes are
in my collection.
Xijlchonis vaiuhis, n. sp.
This species is almost identical to pandulus
Wood, but it may be distinguished by the
denselv placed minute, impressed points on the
elytral declivity on the surfaces between the
usual punctures, by the larger average size, by
die slighth' smaller declivital denticles (the
major spine is usualh' more slender), and by
the shorter scales on the margins and sides of
the elytral declivity.
Female.— Length 3.2 mm, 2.1 times as long
as wide; exactly as in pandulus Wood except as
noted in the above diagnosis.
Type Locality.— Thirteen km SW El X'igia,
Merida, Venezuela.
Type Material.— The female holotype and
five female paratypes were taken at the type
locality on 22-X-69, 100 m elevation. No. 76,
from /(iga branches, by S. L. Wood. Other fe-
male paratypes were taken as follows: 4 at 20
km SW El'Vigia, lO-XII-69, 50 m, various hosts,
S. L. Wood; and 1 at 8 km S Colonia (near
Buenaventura), \'alle de Cauca, Colombia, 9-
Vn-70, 30 m. No. 636, Pwtium nervosum, S. L.
Wood; 2 at 27 km NE Montoya (Campo Ca-
pote), Santander, Colombia, 2'-Vn-70, 150 m.
No. 583, tree sapling, S. L. Wood.
It is entirely possible that this form repre-
sents only a subspecies of pandulus; insufficient
material is available to make an evaluation at
this time.
The holot\pe, allotype, and paratypes are
in m\' collection.
Xi/leborus sJiarpi Icnis, sub.sp. n.
This subspecies i.s distinguislu'd from s.
sharpi Blandford bv the al)sence of the abun-
dant, minute, impressed points on the elytral
deeli\ity, and bv the more northern distribu-
tion.
Female.— Length 3.5 mm, 2.0 times as long
;is wide; color dark brown. Essentially as in s.
sJiarpi except declivital surface almost smooth,
punctures very fine, impressed points essentially
absent. Entire decli\ity in s. sharpi densely cov-
ered bv minute impressi'd points.
Tyi>e Locality.— Twent\ -nine km or 18 miles
E Coatzoeoalcos, \'eracruz, Mexico.
Type Material —The female holotype was
taken at the type locality on 26-\T-67, 30 m,
No. 104, from a tree limb, by S. L. Wood.
The holot\pe is in my collection.
Xyleborus palatus, n. sp.
This species is remotely allied to squamula-
tus Eichhoff, but is distinguished by the slightly
stouter bodv form, by the much steeper, flat
(•1\ tral declix it\-, and b\ the arrangement ol de-
cli\ital tiibi'reles.
Female —Length 1.9 mm (paratypes 1.8-2.1
mm), 2.3 times as long as wide; color very
dark brown to black.
Frons broadlv concave, a slight, transverse
impression above epistoma; surface strongly
reticulate, punctures small, shallow, sparse.
Pronotum 1.1 times as long as wide; sides
feeblv arcuate, almost parallel on basal two-
thirds, rather broadlv rounded in front; anterior
margin anned by six to ten serrations, median
one or two pairs distinctly larger; summit at
middle; anterior area rather coarsely asperate;
posterior areas mostly reticulate with some shin-
ing areas, punctures rather small, shallow, mod-
erately close. Vestiture of moderately abundant,
fine, short hair.
Elytra 1.4 times as long as wide, 1.3 times
as long as pronotum; sides almost straight and
parallel on slightly more dian basal two-thirds.
36
Bhigham Young UNivEnsrrv Science Bulletin
rather narrowly rounded behind; disc occupy-
ing basal 60 percent of elytral length; striae not
impressed, punctures rather small, moderately
deep, spaced within a row by one to two diam-
eters of a puncture; interstriae almost smooth,
shining, about three to four times as wide as
striae, punctures very fine, slightly confused on
basal two-thirds of disc, uniseriate toward de-
clivity. Declivity rather steep, flat on median
half from rounded base to near apex; surface
shagreened, striae as on disc, apices of 2 and
3 converge toward suture; interstriae 1-3 flat; all
interstriae with rows of small, pointed granules,
those near base on all interstriae larger, those
on 1 and 2 on face of declivity and on lower
half of 5, 7, and 8 minute, others sligiitly
larger; posterolateral margin rounded, marked
by a row of small tubercles from apex to inter-
striae 9. Vestiture of minute strial iiair on and
near declivity, and interstrial bristles each as
long as distance between rows on and near de-
clivity, confused on anterior areas of disc.
Type Locality.— Twent\-fc)ur km (1.5 mi)
W Armeria, Colima, Mexico.
Type: Material.— The female holotype and
one female parat\pe were taken at the type
localitN' on .30-\'I-65, .30 m. No. 147, in flight.
by S. L. Wood; 13 female parat\pes bear iden-
tical (lata to the t\pe except Nos. 153, 154, frnin
an unidentified broken tree 1)raneh. Otlier fe-
male paratxpes were taken in Mexico as follows:
one from 53 km (33 mi) S Colima. C'olima, 27-
VI-65, 700 m. Nos, 125. 127, in flight: one from
6 km (4 mi) ,S Cihuatlan, .30-VI-a5, 70 in. No.
157, from a liana; one from \'ole.:ui C^olima, |alis-
co, 23-VI-65, 2.500 m. No. 124. from a shrub limb:
four from Lagima Santa Maria, P.-YU-^r,. 1000
m. No. 19.3, from a liana; and one from 8 km
(5 mi) E San Bias, Nayarit, 12-Vn-65, 70 m.
No. 2.32, from a leguminose tree; all were taken
by me.
The holotype and parat\pes are in m\' col-
lection.
Xr/lebortts cxiitiifi. n. sp.
This species is distinguished from the allied
tolimanus Eggers by the complete absence of
denticles on the elytral declivitv.
Female.— Length 1.9 mm, 3.0 times as long
as wide; color reddish brown, pronotum more
yellowish brown.
Frons about as in catuhis Blandford but
surface finely granulate to well above eyes.
Pronotum 1.2 times as long as wide; basically
as in catulus except anterior margin rather nar-
rowly rounded and more coarselv serrate.
Elytra 1.9 times as long as wide, 1.6 times
as long as pronotum; sides almost straight and
parallel on basal half, acutely converging to
one-fifth greatest elytral width at narrowly,
shallowly emarginate apex; striae not impressed,
punctures small, shallow, distinct, spaced within
a row by twice diameter of a puncture; inter-
striae smooth, shining, three to four times as
wide as striae, punctures fine, uniseriate, rather
widely spaced. Declivitv gradual, narrowlv con-
vex; striae feeblv impressed, punctures slightly
larger than on disc; all interstriae equally sculp-
tured, punctines distinct, regularly, rather close-
ly spaced, anterior margin of each rather
broadly, weakly granulate; interstriae 1 weakly
elevated near acuminate apex; posterolateral
margin weakh' elevated and rather narrowly
rounded near apex, becoming more broadly
numded anteriorly, entirely devoid of granules
and denticles. \'estiture entirely abraded except
for a few hairlike setae on sides.
Type Locality.— Turrialba, Cartago, Costa
Rica.
Type Matehial.— The unique female holo-
type was taken at the t\'pe locality on 9-III-64,
700 m elevation. No. 468, from a new fence
post, by S. L. Wood.
The hoIot\pe is in m\' collection.
Xylchorus rusticus. n. sp.
This species is distinguished from rubricoUis
Eichhoff by the larger size, by the proportion-
ately smaller punctures on the declivital striae,
and by the unifomily con\ex elytral striae.
Female.— Length 3.3 mm (paratype 3.5
mm), 2.3 times as long as wide; color dark
brown.
Frons broadly, irregularly convex; surface
reticulate, shining, punctures rather coarse,
close, moderately deep above, obscure below;
\estiture sparse, inconspicuous.
Pronotum 0.98 times as long as wide; sub-
circular, all margins about equally arcuate,
widi'st slightly behind middle; summit distinctly
behind middle, entire surface closely, rather
coarsely asperate to base, slightly finer on pos-
terior third; surface between asperities reticu-
late, dull. X'estiture of fine, rather long, mod-
erately abundant hair.
Elytra 1.5 times as long as wide, L6 times
as long as pronotum; sides straight on basal
two-thirds, slightly wider at base of declivity,
broadly rounded behind; disc occupying basal
two-thirds; striae 1 feebly, others not im-
pressed, punctures rather small, moderately
Biological Series, Vol. 19. No. 1 New Species of Ameh[C.\n B.mik Beetles
37
deep, spaced within row by one to two diam-
eters of a puncture; iiiterstriae about four times
as wide as striae, almost smooth, sliining, punc-
tures small, moderately confused, their anterior
margins finelv granulate. Declivity steep,
evenly, broadly convex; strial punctures slightly
wider than on disc; interstriae tliree times as
wide as striae, granules (jn all interstriae dis-
tinctly larger, pointed. Vestiture of short strial
hair and fine, long, abundant hair; interstrial
setae in almost uniseriate rows on declivital in-
terstriae 1 and 2, confused elsewhere.
Type Locality.— Ten km NE Teziutlan,
Puebla, Mexico.
Type Material.— The female holotype was
taken at the type locality on 2-VII-67,'l600 m.
No. 150, from an unidentified log by S. L.
Wood. One female parat\pe is from the same
locality and collector, taken 27-\T-.53, No. 49.
The holot)pe and paratype are in my col-
lection.
Xijlehonis occUatus, n. sp.
This species is distinguished from nistictis
Wood by the coarser strial punctures, by the
more deeply impressed declivital striae, bv the
strial punctures on tlie decli\itv, each with a
granule occupying its central half thereby giv-
ing the superficial appearance of an eye, and
by the larger interstrial granules on the de-
clivity.
Female.— Length .3..5 mm (parat\pes .3..'3-.'3.5
mm), 2.3 times as long as wide, color black.
Frons and pronotum as in nisfictts except
asperities on basal half of pronotum distincth
larger.
Elytra 1.4 times as long as wide, L5 times
as long as pronotum; outline as in rusticus;
striae 1 feebh', others not impressed, punctures
moderately coarse, rather sliallow; interstriae
three times as wide as striae, almost smooth,
punctures rather fine, finelv granulate on their
anterior margins, confused on 2 and 3. Decli\-
ity convex, steep; striae 1 distincth-, 2 feeblv
impressed, punctures sliglitlv larger than on
disc, each puncture with its central half occu-
pied by a rounded granide; interstriae feeblv
convex, each amied by a uniseriate row of
moderately coarse, pointed tubercules, 7 sub-
acutely elevated and tuberculate. \'estiture as
in ntsticits.
Type Locality.— Picdras Blancas, 10 km E
Medellin, Antioquia, Colombia.
Type Material.— The female holotvpe and
two female parat\'pes were taken at the type
locality on 15-Vn-70, 2500 m, No. 675, from an
unidentified log, by S. L. Wood. Two other
female paratypes bear identical data except
one is from No. 684 in Chtsia, and one is No.
677 taken in flight.
The holotype and p;iratypes are in my col-
lection.
Xijlehonts opinuts. n. sp.
This species is distinguished from Iccontei
(Hopkins) by the slightly smaller size, by the
more strongly rounded anterior and lateral mar-
gins of the pronotum, b\' the finer, less numer-
ous asperities on the pronotal disc, by the con-
\'ex el)'tral declivity, and by the different ar-
rangement of declivital tubercles.
Female.— Length 2.4 mm, 2.2 times as long
;is wide; color reddish brown.
Frons as in leconfei. Pronotum 1.0 times as
long as wide, as in Iccontei except antcTior ;uid
lateral margins more strongly ;ucuate, summit
slightly higher, and asperities behind summit
slightly smaller and evidently less numerous.
Elvtra 1..36 times as long as wide; outline
and disc as in Iccontei except strial punctures
not larger near disc, interstrial punctures near
declivity granulate. Decli\itv rather broadly,
evenly convex, steep; strial punctures as large
as on disc, deep, smaller than in Iccontei; inter-
striae 1 bearing a row of granules on basal half,
those near base rather coarse, 2 with several
pointed tubercles on basal half, one or two at
or sliglitlv below middle distincth larger, 3
with three to five smaller tubercles; all tubercles
smaller than major tubercles of Iccontei. X'esti-
ture as in Iccontei.
Type Locality.— Sebring, Florida.
Type Materl\l.— The female holotvpe was
taken at the t\pe locality on 20-\T-51,'at light,
by S. L. \Vood.
The holotype is in mv collection.
Xi/Iehorus lacunatus, n. sp.
This species is distinguished from commi.xtii.'i
Blandford hv details in sculpture of the elvtral
decli\itv as indicated in the description.
Female.— Length 4.9 mm, 2.5 times as long
as w ide; color dark brown.
Frons, pronotum, and elytral disc as in com-
mixtits. Elvtral declivity essentially as in com-
mixtus except rather strongly, transxerseh' im-
pressed on lower third; suture distinctly concave
on lower two-thirds; strial punctures larger; in-
terstriae about twice as wide as striae; postero-
38
Bnic.HAM Young 1'ni\ersitv Science Bulletin
lateral margin much more strongly, aciitch- ele-
vated than in comiiiixtus, rather strongly concave
from its crest to suture on lower third of decliv-
ity. \'cstiture as in commixtus.
Type Locality.— Turrialba, Cartago, Costa
Rica.
Type Material.— The female holotype was
taken at the type Iocalit\ on 18-IV-6.3, from
Theohroma cacao, by J. L. Saunders. One female
paratype is from Peralta, Cartago, Costa Rica,
10-III-64, flight, S. L. Wood.
The holotype and parat\pe are in mv collec-
tion.
Xtjlehonis iiwridemis, n. sp.
This species is distinguished from caraihicus
Eggers by the slight!)' larger size, by the larger,
less deeply impressed punctures on the striae, by
the more gradual declivity with the lower me-
dian half almost flat, and b\- the rounded postero-
lateral margin of the decli\ity.
Female.— Length 4.1 mm (paratypes .3.7-4..3
mm), 2.7 times as long as wide; color dark
brown.
Frons and pronotum as in caraihicus except
posterior areas of pronotum more distinctlv re-
ticulate.
Elytra L7 times as long as wide, L7 times
as long as pronotum; sides almost straight and
parallel on slightly less than basal two-thirds,
slightl\- tapered then rather narroulv rounded
behind; striae not impressed, punctures moder-
ately coarse, shallow; interstriae twice as wide
as striae, smooth, punctures fine, uniseriate. De-
clivity moderately steep, rather narrow, feebly
impiessed on median area partictilarl\- on central
half; suture on lower two-thirds straight; striae
not impressed, punctures on middle half larger
than on disc, interstriae one and one-half times
as wide as striae; interstriae smooth, shining,
each armed by about five small, pointed tuber-
cles; posterolateral margin rather narrowly
rounded (not carinatc), amied by a series of
rather widely spaced, small, pointed granules.
Male.— Length 3.4 mm; head and elytra es-
sentially as in female. Pronotum L2 times as
long as wide, 0.86 times as long as elytra; cjuad-
ratc, anterior third broadly, concavely excavated,
anterolateral margins abruptly angulate, acutely
elevated; anterior margin acutely elevated, slight-
ly produced into an obtuse, median point; sub-
glabrous.
Tyi'e Locality.— I^a Carbonera experimental
forest, about 50 km (airline) NE Merida, Meri-
da, \'enezuela.
Type Material.— The female holotype, male
allotype, and four paratypes were taken at the
type loeaiitx- on 23-IV-70, 2.500 m, Nos. 450, 45L
from an unidentified log b\' S. L. Wood. Other
paratypes were taken at the same l()c;i!itv as fol-
lows: 5 on 16-L\-69, No. 21 in Pniniis spluiero-
carpa\ 1 on 16-L\-69, No. 20 on Ficiis. 1 on 14-X-
69, No. 66; 2 on 27-.\-69, No. 92; 6 on 9-.\n-69,
No. 171; 8 on 2S-I\'-70. Nos. 450, 451, and 457,
from unidentified logs; all by S. L. Wood.
The holotype, allot\pe, and paratypes arc in
mv collection.
Xijleborus aclinis. n. sp.
This species is distinguished from incridensis
Wood by the much smaller size, by the more
strongly convex deelixitv, and by the presence
of only one tubercle on decli\'ital interstriae 2.
Female.— Li'ngth .3.1-3.3 mm, 2.9 times as
long as wide; color dark brown.
Frons and pronotum as in incridensis except
pronotum 1.14 times as long as wide.
Elytra 1.8 times as long as wide, 1.6 times as
long as pronotum; outline and disc as in meriden-
sis except interstrial punctures more widely
spaced. Decli\itv moderately steep, convex;
longitudinal axis of suture feebly convex on
lower half; striae not impressed, punctures little
if any larger than on disc; interstriae smooth,
shining, 1 and 3 each amied by about three to
six pointed granules of variable size, 2 armed
by one moderateh' coarse di'uticle one-third de-
clivital length from apex, a few small granules
in lateral areas; posterolateral margin as in meri-
densis. narrowly rounded, its crest amied by
se\('ral small, isolated granules. \'estiture eon-
fined to decli\'ity, sparse, one short bristle aris-
ing from posterior basal margin of each granule,
each about half as long as width of an inter-
striae.
Type Locality.— Cerro Punta near Volcan
de Chiriqui (Barn), Chiriqui, Panama.
Type Material.— The female holotype and
three female paratypes were taken at the type
locality on 11-1-64. 1800 m. No. 386, from a
stump 15 cm in diameter, by S. L. \\'ood.
The holotvpe and paratypes are in my collec-
tion.
Xijlehonis dissimidatus, n. sp. -
This species is distinguished from mcridensis
Wood by the much smaller size, by the smaller
strial punctures, by the much smaller granules
on declivital interstriae 2, and bv the more sub-
angulate, subserrate posterolateral margin of
Biological Series, Vol. 19, No. 1 New Species of American B.\hk Beetles
39
the declivity. It is distinguished, with difficulty,
from posticus Eichhoff by the shining, more
gradual decHvity, and by the shorter, sparse
dechvital bristles.
Female.— Length 2.2 mm (paratypes 2.2-2.3
mm), 2.6 times as long as wide; color brown.
Frons, pronotum, and elytral disc as in posti-
cus.
Elytra 1.6 times as long as wide, 1.6 times as
long as pronotum; disc occupying basal 60 per-
cent of elytral length. Declivity moderately
steep, rather broadly convex; strial punctures
much larger than on disc, very shallow; inter-
striae smooth, shining, as wide as striae, each
with a sparse row of minute granules, 1 and 3
each with about two distinctly larger granules;
posterolati'ial margin subacute, more continu-
ously subserrate than in posticus. Vestituri'
largely confined to declivity, of very short, stout
and fine, interstrial bristles, each bristle about
one-third as long as distance between rows.
Type Locality.— Tapanti. Cartago, Costa
Rica.
Type Material.- The female holotype and
three female paratypes were taken at the type
locality on 17-IX-63, No. 184, from Miconin
caudata, by S. L. Wood. One female paratNpe
bears identical data except No. 178 from a liana.
One paratype is from the same locality and col-
lector taken 2-\'II-63, No. 9, Miconia sp.
The holotype and paratypes are in my col-
lection.
XijIe])orus concentus, n. sp.
This species is distinguished from parollclo-
collis Eggers by the more narrowly con\ex
declivity on the transverse axis, by the serrate
posterolateral margin of the declivitv, and by tlu'
shorter interstrial bristles on thi' declivit).
Female.— Length 2.5 mm (parat\pes 2.5-2.7
mm), 2.7 times as long as wide; color dark
brown.
Frons and pronotum as in caraihicus Eggers
except pronotum 1.15 times as long as wide.
Elytra 1.7 times as long as wide, 1.6 times as
long as pronotum; sides almost straight and
parallel on slightly less than basal two-thirds,
tapered, then narrowh' rounded behind; .striae
not impressed, punctures rather small, distinctlv
impressed; interstriae twice as wide as striae,
punctures very fine, uniseriate. Declivity grad-
ual, transversely con\ex, suture very feebly con-
vex on lower half; surface rather dull in most
specimens; striae not impressed, punctures larger
than on disc; interstriae flat, slightlv ^^'ider than
striae, granules small, of uniform size, spaced
by distances less than width of an interstriae;
posterolateral margin abrupt, subacute, finely
serrate. Vestiture largely confined to declivity,
consisting of short, stout, interstrial bristles, each
bristle equal in length to one-third width of an
interstriae; a few similar, much longer bristles
on disc.
Type Locality.— Tapanti, Cartago, Costa
Rica.
Type Mateklal.— The female holotype and
two female paratypes were taken at the type
localitN' on 26-Xl'-63, 1.300 m. No. 265, from
Phoebe mexicana, b\' S. L. Wood. Other female
paratypes were taken as follows: 1 at Rincon de
Osa, Puntarenas, Costa Rica, ll-\'Ill-66, 30 m,
No. 90, liana; 1 at \'olcan, Puntarenas, Costa
Rica, ll-Xll-63, 1000 ni, No. 308, tree limb; 1 at
Cuapiles, Limon, Costa Rica, 22-\'II-66, 100 m.
No. 101, tree limb; and 6 at 30 km E Palmar,
Bolivar, X'enezuela, 12-VT-70, 200 m, No. 556,
Alexii imperatricis; all by S. L. \\'ood.
The holotype and paratypes are in n\\ col-
li'ction.
Xyleborus trihulattis, n. sp.
This species is distinguished from .seniipunc-
tatus Eggers except as noted in the following de-
scription.
Female.— Length 3.8 mm, 2.4 times as long
as wide; color very dark brown.
Frons and pronotum as in semipunctatus ex-
cept punctures on posterior half of pronotum
distinctlv larger, closer.
Elvtra as in semipunctatus except discal striae
2 distinctly sinuate; interstriae 2 and 3 on disc
distinctlv confused; strial punctures on disc and
dcclixity distinctly larger, interstriae on disc
three times as wide as striae ( four times as wide
in .semipunctatus). Declivitv as in .semipuncta-
tus except as noted.
Type Locality.— Rio Damitas in the Dota
Mountains, San Jose, Costa Rica.
Type Material.— The unique female holotype
was taken at the type locality on 22-\'III-63,
250 m. No. 126, from a liana 10 cm in diameter,
by S. L. Wood.
The holotype is in my collection.
Xyleborus vismiae, n. sp.
This species is distinguished from costaricen-
sis Blandford by the shorter, steeper declivity
which is strongly, transversely impressed on the
lower half, b\' the smooth, shining declivital sur-
face, and by the larger declivital granules. The
40
Brigham Young I'niveiisity Science Bili.etin
costaricensis group of species is characterized by
the shghtly protuberant, smooth, peculiarly retic-
ulate frons that is desoicl of punctures, bv the
tapered posterior half of the elytra, and b)' the
long, rather narrow declivity.
pEAfALE.— Length 3.5 mm (paratypes 3..'3-3.7
mm), 2.6 times as long as wide; color dark
brown.
Frons shallowly, transversely impressed be-
tween eyes, \\eakly inflated below; surface
smooth with very fine reticulation below, more
coarsely reticulate toward vertex, punctures
rather small, sparse, mostly confined to lateral
areas, none on lower third; vestiture very sparse,
hairlike, inconspicuous.
Pronotum 1.13 times as long as wide; sides
almost straight and parallel on basal half, rather
narrowly roimded in front; smnmit at middle;
anterior half ratlier fineh' asperate; posterior
areas very finc'ly subreticulate; punctures minute,
distinct, rather sparse.
Elytra l.S times as long as wide, 1.6 times as
long as pronotum; sides ;ilmost straight and
parallel on basal half, then rather strongly ta-
pered, rather nanowly rounded behind; striae
not impressed, punctures small, distinct, spaced
within a row by one to two diameters: interstriae
smooth, moderately shining punctures \ery mi-
nute, distinct, uniseriate. Declivity occupying
slightly more than posterior third, strongly,
transversely impressed on lower half; transverseh-
flat; longitudinalh' conca\e; striae curved toward
suture near apex, punctures almost twice as
large as on disc; interstriae shining, anned bv
rows of small granules, altematt- granules slight-
ly larger; posterolateral margin narrowly round-
ed, its crest with ;i ivw feeble granules.
Type Localitv,— Rio Dainitas in the Dota
Mountains, San Jose, Costa Rica.
Type Material.— Th(> female holotype and
three female paratypes were taken at the t\pe
locality on 22-\'Ili-63, 250 m. No. 126, from
Vismia gtiai/dnemis. by S. L. Wood. Eight fe-
male paratypes are from Rancho Grande, Pittier
National Park, Aragua, \'enezuela, 9-n'-70, 1100
m. No. 433, from a species of Guttiferae near
but probably not Vismia, by S. L. Wood, ex-
cept one of these is No. 441 from an unidenti-
fied tree limb.
The holot\pe and paratypes are in mv collec-
tion.
Xylehonis demissiis. n. sp.
This species is distinguished from dvplanatns
Eggers by the more slender body form, b\- the
shorter declivity, by the two sizes of intirstrial
tubercles on the declivity, and by the less
strongly serrate posterolateral margin of the
declivity.
Female.— Length 2.0 mm, 2.9 times as long
as wide; color dark brown.
Frons and pronotum ;is in cleplanafus except
pronotum 1.21 times as long as wide.
Elytra 1.7 times as long as wide, 1.5 times as
long as pronotum; sides straight and parallel on
basal half, then slighth' tapered, narrowK round-
ed behind, weakly emarginate at suture; disc
occupying slightly more than basal half; striae
not impressed, punctures small, shallow, distinct;
interstriae smooth, shining, three times as wide
as striae, puncturi's unisiTiate, slightly smaller
than those of striae, their anterior margins feebly
granulate at base, becoming more distinctly gran-
ulate posteriorly. Declivity rather gradual, mod-
erateK' coincx on both axes; stri;ie not im-
pressed, punctures slightly larger than on disc,
lateral margins of punctures on 2 and 3 weakly
granulate; interstriae shining, about twice as
wide a.s striae, each anned b\- a row of granules,
granules mostly small except on 2 and 3 dis-
tincth' larger granules alternate with small ones.
X'estiture of rows of fine, short, strial hair, and
interstrial rows of short, almost scalelike bristles,
eacli bristle one-third to one-half as long as dis-
tance between rows, slighth' closer within a row,
each three to fi\e times as long as wide.
Type Locality.— Rincon de Osa, Puntarenas,
Costa Rica.
Type Material.- The imique female holo-
type was taken at the type locality on 11-\'II1-
66, 30 m. No. 70, from a log 20 cm in diameter,
by S. L. Wood.
The holotvpe is in my collection.
Xylehonis mcritits, n. sp.
This spi'cies is distinguished from vismiae by
the slightly larger sizt\ by the more gradual,
more nearly flattened elytral declivit\', and b\
the much finer interstrial granules on the decliv-
ity.
Female.— Length 3.0 mm (parat\'pes 2.8-3.4
mm), 2.8 times as long as wide; color yer\' dark
brown.
Frons, pronotum (1.15 times as long as
wide), and ehtral disc as in vismiae Wood.
Elytra 1.8 times as long as wide, 1.6 times as
long as pronotum; outline as in vismiae. Decliv-
ity occupying slightly less than posterior half
of elytral length, its lower half very shallowK
concave; striae not impressed, cur\ing toward
suture near apex, punctures almost twice as
Biological Series, Vol. 19. No. 1 New Species ok Americ.vn B.-^hk Beetles
41
large as on disc; intcrstriae smooth, shining, with
widch' spaced fine grannies, those on lower half
of 2 and 3 smaller, sometimes almost obsolete.
Vestitnre sparse, of fine hair, length on declivit)'
equal to about half width of an interstriae.
Type Locality.— Tapanti, Cartago, Costa
Uica.
Type Material.— The female holotype and
one female paratype were taken at the type lo-
calit\ on 17-IX-6:3, 1:300 m. No. 178, liana, S. L.
Wood. Thirteen female paratvpes were taken at
the same locality (2) 2-\TI-63, No. 7, liana, (1)
3-VII-63, No. 17, Conostegia oerstedkina. (2)
17-IX-63, No. 182. Miconia caudato. (1) 24-X-
63, No. 184, Miconia caudata. (6) 26-XI-63, No.
265, Phochcd mexicana. Ten female paratvpes
are from 14 km SE Cartago, Cartago, Costa Ri-
ca, 1800 m, (1) 3-\TI-63, No. 17, Conostegia
oerstediana, (8) 24-IX-63, No. 204, Miconia
ghhidiflora, and (1) 24-IX-63, No. 200, Myrica
ptd)escens: all by S. L. Wood.
The holotvpe and paratvpes are in mv collec-
tion.
Xiilel>onis prolatns. n. sp.
This species is distinguished from costari-
cen.iis Blandford b\' the larger size, b\ the
longer, more strongly impressed declivity, and
by the near absence of granules on declivital in-
terstriae 1 and 2.
Female.— Length 4.2 mm (paratypes 4.0-4.4
mm), 2.8 times as long as wide; color dark
brown.
Frons and pronotuni as in costariccnsis.
Elvtra 1.7 times as long as wide, 1.7 times as
long as pronotum; outline about as in costari-
censis except more strongly tapered on posterior
half, more narrowly rounded behind; disc con-
fined to basal third, as in costaricensis. Decli\ity
verv gradual, shallowlv, broadly bisulcate; sur-
face dull; strial punctures twice as large as on
disc; interstriae almost twice as wide as striae,
smooth, 1 distinctly elevated toward suture, 2
broadlv, subconca\el\' impressed, 3 distincth-,
gradually ele\ated toward broad summit at striae
4, 3 and 4 each with a ro\\- of widely spaced,
moderately large granules, 1 and 2 sometimes
with one or two smaller granules near apex ( one
paratype with five granules on 2 on left side).
V'estiture as in costaricensis except almost en-
tirelv absent on declivital interstriae 1 and 2.
Type Locality.— Tapanti, Cartago, Costa Ri-
ca.
Type Material.— The female holotype and
eight female paratvpes were taken at the tvpe
locality on 24-X-63, 1300 m. No. 265, from a re-
cently cut limb of Phoehea mexicana 10 cm in
diameter, bv S. L. \\'ood.
The holotvpe and parat\pcs are in my col-
lection.
Xijlehorus dissidcns. n. sp.
This species is placed near sparsipilosus Eg-
gers because of the slender pronotum and steep
elytral declivity; however, it probably is more
closely related to species placed in the sub-
genus Euicallacca. In ;uldition to the slender
pronotum with its procurxcd ;mterior margin,
this species has the eKtral decli\it\' steep, con-
vex, and interstriae 1-3 eciuallv armed b\' pointed
granules.
Female.— Length 3.1 mm (paratypes 3.0-3.2
mm), 3.0 times as long as wide; color black.
Frons as in costaricensis Blandford but with
lower area less strongly inflated.
Pronotum 1.22 times as long as wide; as in
sparsipilosus except posterior areas subreticulate.
Elvtra 1.8 times as long as wide, 1.5 times as
long as pronotum; sides straight and parallel on
slightly more than basal two-thirds, rather broad-
lv rounded behind; disc occupying slightly
more than basal two-thirds; striae not impressed,
punctures rather small, distinct, spaced within a
row by diameter of a puncture; interstriae
smooth, shining, twice as wide as striae, punc-
tures uniseriate, minute, distinct, almost ob-
solete. Declivitv steep, rather broadly convex:
striae about as on disc; interstriae 1-3 each
I'cjuallv armed by six to ten pointed granules
of slightly irregular size; posterolateral margin
obtuse, ;ibrupt. Wstiture confined to decli\ity,
of rows of rather coarse, interstrial setae, each
seta about one and one-half times as long as
distance between rows, more widely spaced
within a row.
Type Locality.— Nine km NE Tezuitlan,
I'uebla, Mexico.
Type Material.— The female holotype and
thri'e female parat\pes were taken at the type
locality on 2-\TI-67, 1500 m. No. 141, from
Alnits cordwood, by S. L. Wood. Two female
paratypes liear identical data except one is No.
147, from an unidentified log, and one is No.
141, taken in flight.
The holotvpe and paratypes arc in my col-
lection.
Xtflehorinu.s dints, n. sp.
Superficiallv this species might be confused
with Xi/lehortis ferox Blandford, but the conical
42
Bhigham Young University Science Bulletin
scutellum and emarginate elytral base show the
true relationships to be very different.
Female.— Lcngtli 3.5 mm (paratypes: fe-
males 3.4-3.6 mm, males 2.5-2.7 mm), 2.7 times
as long as wide; mature color black.
Frons convex, opistoma slightlv elevated; sur-
face reticulate, punctures rather large, im-
pressed, indefinite; vestiture inconspicuous.
Pronotum 1.06 times as long as wide, widest
a third of its length from base; sides rather
weakly arcuate on basal two-thirds, con\'i'rging
very slightly, then rather strongly rounded in
front, median area rather narrowly produced
but unarmed ( in manv paratypes anterior mar-
gin rather broadly rounded); summit very
slightl)' in front of middle; posterior area reticu-
late, indistinctly so near base, punctures small,
distinct, rather close; vestiture rather long and
abundant on sides and in asperate area.
Elytra 1.6 times as long as wide, 1.5 times
as long as pronotum; sides almost straight and
parallel on basal half, then arcuatelv converging
to apex of \entrolateral spine, rather broadlv
U-shaped between spines; striae 1 weaklv, others
not impressed, punctures moderatelv large, deep,
somewhat confused on 1; interstriae mostly twice
as wide as striae, punctures similar to those of
striae but deeper, imiseriate except paith con-
fused on 1 and 2. Declivitv moderately steep,
broadly, shallowly excavated, lateral margins
armed by three pairs of major spines; spine 1 at
upper margin in line with striae 2, spine 2 on
lateral margin two-thirds declivital lengtli from
upper margin, spine 3 at posterolateral margin;
spine 1 slighth' more than half as long as 3, 2
slightlv smaller than 3, 2 ef(ual in length to
width of antennal club; one minor tooth in front
of spine 1, two others between 1 and 2; decli\'ifal
face with confused punctures similar to those on
disc. Vestiture consisting of rather long, slender
hair arising from interstrial pimctures on disc
and sides, setae minute on excavated area.
Male.— Length 2. .5-2. 7 mm; similar to fi'inale
except smaller, eye reduced, pronotum not
strongly arched, with asperities somewhat re-
duced; elytral declivity much longer, more
gradual, with spine 1 greatly reduced, minor
teeth mostly absent.
Type Locality.— Rincon de Osa, Puntarenas,
Costa Rica.
Type Material.— The female holotype, male
allotype, and 34 paratypes were collected at the
type locality on H-VIII-66, at about 30 m eleva-
tion, from cut limbs 10-20 cm in diameter in
the primary forest. Nineteen additional para-
types were taken at Rio Damitas in the Dota
Mountains, San Jose, Costa Rica, 22-\'1 11-63. 250
m, from a stump 25 cm in diameter in the pri-
mary forest; all specimens were collected by my-
self. The tunnels were of a branching type that
included several enlarged cavities.
The holotype, allotype, and paratypes are in
my collection.
XtjIeI)onnris tril)tilosiis, n. sp.
This species is distinguished from dints Wood
by the smaller size, by the more slender body
form, by the smaller, uniseriate strial and inter-
strial punctures on disc and declivity, and by
the different elytral declivitv.
Female.— Length 2.5 mm (paratype 2.4
mm), 2.S times as long as wide; color very dark
brown.
Frons and pronotum essentially as in dims
except pronotum 1.1 times as long as wide;
pronotal disc glabrous.
Elytra 1.6 times as long as wide (spines ex-
cluded), 1.4 times as long as pronotum; outline
essentially as in dims; striae not impressed,
punctures small, uniseriate, close; interstriae
smooth, shining, slightly more than twice as
wide as striae, punctures half as large as those
of striae, uniseriate. Declivitv occupying two-
thirds of elytral length; broadly flattened to
feebly impressed, its margin amied hv about
24 rather co;irse, pointed denticles on interstriae
1-6 from base to near sutural apex; interstriae 3
armed at middle and near apex by two pairs of
very large spines, each spine one and one-half
times as long as its basal width, about ec(ual in
length to discal distance from suture to striae
3; face of declivity with strial punctures in rows,
slightly larger than on disc; interstrial punctures
largely obsolete. Vestiture of rows of rather
coarse inteistrial setae, on disc each seta about
as long as distance between rows, distinctly
closer within a row, on declivity distinctly longer
and less regularly placed.
Type Locality.— Madden Forest, Canal Zone,
Panama.
Type Material.— The female holotype and
one female paratvpe were taken at the type lo-
calitv on 2-1-64, 70 m. No. .367, from a tree limb
by S. L. Wood.
The holotvpe and paratype are in mv col-
lection.
Xt/leborinus protinus, n. sp.
This odd species is not closely related to any
American species knowii to me.
Biological Series, Vol. 19, No. 1 New Species of Amehic:.\n B.\rk Beetles
43
Female.— Length 1.7 mm (paratypes 1.6-1.7
mm), 2.7 times as long as wide; color rather
light brown, prothora.x distinctly lighter.
Frons very weakly convex from upper level
of eyes to epistomal margin; surface reticulate,
rather coarsely, obscureK punctured; \estiture
inconspicuous.
Pronotum 1.25 times as long as wide; widest
one-third pronotum length from base, sides
weakly arcuate, basal and anterior angles more
strongly rounded, rather naiTowly rounded in
front; asperities fine, largely isolated; summit
indefinite, in front of middle; surface reticulate
in front of summit, niosth' smooth and brightly
shining behind, punctures small, deep, not close;
vestiture largely confined to sides and asperate
area.
Elytra 1.6 times as long as wide, 1..3 times
as long as pronotum; sides essentially straight
and parallel on basal three-fourths, then rather
abruptly rounded, shallowlv emarginate on me-
dian third behind; scutellum conical; striae not
impressed, punctures small, deep; interstriae
twice as wide as striae, punctures small, indefi-
nite; elytra arched from base to apex. Declivity
beginning on basal third of ehtra, gradual;
lateral margins on lower half gradualh' ele\'atcd
and continuing almost to apex, elevation rather
high, not at all acute, evidently entireK un-
armed; lower half transversely concave; striae 1
and 2 with punctures larger than on disc, strong-
ly impressed; interstriae 2 wider than 1; inter-
strial punctures obscure. Vestiture consisting of
stout, hairlike setae, more abundant on declivitv,
particularlv on inner slope of lateral elevation.
Type Locality.— Finca La Lola, Limon, Cos-
ta Rica.
Type Materl\l.— The female holotype and
four female paratypes were collected at the t\pe
localitv on 7-II-63, from Thcohroma cacao. h\
J. L. Saunders.
The holotype and paratypes are in in\ collec-
tion.
Xylehorimis celatu.s. n. sp.
This species differs from rcconditus Schedl
by the much finer punctures on the pronotal
disc and by the reduction of the denticles on
declivital interstriae 2.
Female.— Length 1.7 mm (parat\pes 1.6-l.S
mm), 2.5 times as long as wide; color dark
brown, almost black.
Frons and pronotum as in recouditus except
posterior areas of pronotum with punctures
much finer, less numerous.
Elytra 1.4 times as long as wide, 1.3 times
;is long as pronotum; sides almost straight and
parallel on basal three-fourths, then rather
abruptly rounded, straight on median half be-
hind; disc confined to basal third of elytral
length; striae not impressed, punctures minute;
interstriae almost smooth, subshining, at least
six times as wide as striae, punctures slightly
smaller than those of striae. Declivity gradual,
longitudinallv flat and trans\ersely very weakly
conv(>x on its basal half, slightlv steeper and
broadlv, shallow Iv sulcate on its lower half; base
commencing much more abruptly than in rccon-
ditus; lower half of declivity with contours as
in rccondiltis but strial punctures smaller and
interstriae 3 armed b\- onlv two denticles, one
just behind middle of declivity and one on its
apical fourth; interstriae on basal half armed
by row's of fine granules. \'estiture confined to
declivitv, consisting of minute strial hair and
interstrial rows of stout bristles; each bristle as
long as distance between rows.
Type Locality.— Eight km S Colonia (near
Buenaventura), \'alle de Cauca, Colombia.
Type Matehial.— The female holotype and
fi\'e female parat\pes were taken at the type lo-
cality on 9-\TI-76, 30 m. No. 646, from Zngrt sp..
In S. L. \\'ood. Nine female parat\pes bear
identical data except they are No. 628 from Idea
(dfisiiiHi or Xo. 631 from Protitim nervosum.
The holotype and paratypes are in my col-
lection.
Araptus insinuatus. n. sp.
Tliis species is distinguished from the very
closelv related fovcifrons Schedl bv the strong-
ly impri'ssed decli\ital interstriae 2, by the
longer ehtral \estiture, and by the more finely,
more closely punctured pronotal disc.
Male.— Length 1.6-2.1 mm, 2.4 times as long
as wide; color reddish brown.
Frons and pronotum as in fovcifrons except
punctures on pronotal disc distinctly smaller,
closer. Elytra as in fovcifrons except discal
punctures a\eraging slightly longer, finer than
in fovcifrons.
Female.— Similar to male except frons (con-
cealed) apparentlv as in female fovcifrons.
Type Locality.— Guatemala.
Type Material— The male holotype, female
allotype, and two male paratypes were inter-
cepted from Guatemalan seeds at San Pedro,
California, 7-III-63. One male paratype is from
44
La Ceiba, Atlantida, Honduras, 17-VI-49, at
light, by E. C. Becker.
The holotype, allotype, and paratypes are
in my collection.
Araptits interjectus, n. sp.
Thi.s species is distinguished from foveifrom
Schedl by the larger size, by the more elongate
form, by the near absence of interstrial punc-
tures, and by the different frons in both sexes.
MALE.-Length 2.4 mm (paratypes 2.4-2.7
mm), 2.9 times as long as wide; color rather
dark reddish brown.
Frons strongly, transversely impressed from
eye to eye from middle to well above upper
level of eyes, its margins obtuse, its lower lateral
angles anned b)' a pair of rather coarse tuber-
cles and with a' large, pointed, median tubercle
at same level; lower third of area below upper
level of eyes subaciculate; all surfaces shining;
vestiture sparse except along epistomal margin.
Antennal club as in foveifwns.
Pronotum 1.15 times as long as wide; widest
behind middle, sides moderately arcuate on
more than posterior half, weakly constricted on
anterior third, rather broadly rounded in front;
anterior margin armed by about eight irregular
serrations; indefinite summit one-third of prono-
tum length from ;mterior m;irgin; asperities fine,
confused; posterior areas obscurely reticulate in
some areas, numerous minute, impressed points
present, punctures rather small, deep, moder-
ately close. Glabrous except for a few setae on
asperate area and lateral margins.
Elytra 1.7 times as long as wide, 1.7 times
as long as pronotum; sides straight and parallel
on basal two-thirds, broadly rounded behind;
striae not impressed except 1 near decli\'ity,
punctures moderately coarse, spaced by diam-
eter of a puncture; interstriae twice as wide as
striae, shining, with rather numerous minute
points and moderately abundant, subtrans\erse
impressed lines, punctures obsolete except for
an occasional puncture on some specimens. De-
clivitx' steep, broadly convex; sutural interstriae
distinctlv, rather ;ibruptK- ele\ated, 2 slightly
wider than 1 or 3 and impunctate, 1 and 3 each
with a few small punctures. Vestiture largely
abraded, a few short interstrial bristles on de-
clivity.
pEXJALE.-Similar to male except frons
broadlv flattened, weakly impressed near
median line, median line with a conspicuous,
blunt carina from epistoma to well above eyes;
frontal vestiture of moderately abundant, fine.
BH1GH.\M VOUNC. UnIVERSITV SCIENCE BULLETIN
uniformly distributed long hair, distinctly longer
at margins.
Type Locality-. -N'olcan de Agua, Guate-
mala.
Type MATERtAL.-The male holotype, female
allotype, and 12 paratypes were taken at the
type locality on 19-V-64, 1000 m, Nos. 609 and
612, from pith tunnels in a cut vine, by S. L.
Wood.
The holotype, allotype, and paratypes are
in mv collection.
Araptus dcrincftis, n. sp.
This species is distinguished from foveifwns
Schedl by the more slender body form, by the
sparse discal interstrial punctures, and b\ the
very different frons in both sexes.
MALE.-Length 2.2 mm (paratypes 2.1-2.5
mm), 2.S times as long as wide; color dark
reddish brown.
Frons excavated as in foveifrom but with
lateral cusps longer, extending ventrad to upper
level of eves, a sharply ele\ated median carina
extending from deepest point of excavation end-
ing on epistomal margin in a small tooth, lateral
margins of lower half of excavation amied by
one or more pairs of small tubercles. Antennal
club as in foveifwns.
Pronotum and elytra as in interjectus Wood
except odd-numbered discal interstriae punc-
tured, even-numbered interstriae very sparsely
pimctured, and declivity as in foveifrons except
more n;ui()\\l\ eoincx, punctures smaller.
FEMALE.-Similar to male except frons
shallowlv concave from e\'e to eye from epi-
stoma to vertex, surface shining, finely punc-
tmed, subaciculate, a distinctly elevated median
carina extending from center of exca\ation to
denticulate epistomal margin; margins of frontal
excavation ornameiitt'd by a rather dense fringe
of fine, long hair.
Type Locality. -Three km (2 miles) SE
Acatlan, Puebla, Mexico.
Type MAXERiAL.-The male holotype, female
allotvpe, and five parat\pes were taken at the
type' locality on 15-\'-67, 1500 m. No. 38, from
pith tunnels in a cut vine, by S. L. Wood.
The holotype, allotype, and paratypes are
in mv collection.
Araptus delicatus, n. sp.
This species is distinguished from p,enialis
Wood bv the smaller size, by the more slender
form, and bv the \ery different sculpture of
the frons.
Biological Series, Vol. 19, No. 1 New Species of Americ.'\n B.-mik Beetles
45
Male.— Length 1.9 nun (paratype.s 1.7-2.5
mm ), 2.S times as long as wide; color dark hro\\n
except ba.sal half of elytra light brown.
Frons dcepl\' impressed on triangular area
from epistoma to vertex, upper angle of tri-
angle on vertex an inverted U-shaped area hav-
ing its margins acutely costate, floor of im-
pressed area obscurely aciculate and with a
low, long, acute carina; vestiture fine, short,
moderately abundant, with a conspicuous cpi-
stomal brush. Antennal club as in foveifrons.
Pronotum 1.2 times as long as wide; sides
on basal half almost straight and parallel,
broadly rounded in front; anterior margin
armed by about 12 low serrations; indefinite
summit one-third pronotum length from anterior
margin; ;isperities small, confused; posterior
areas shining with numerous impressed points,
punctures moderately coarse, deep, close. Ves-
titure of moderately abundant, fine, very short
hair.
Elytra 1.6 times as long as wide, 1.4 times
as long as pronotum; sides straight and par-
allel on basal two-thirds, rather broadly
rounded behind; striae not impressed except 1
weakly, punctures rather small, deep, spaced
by diameter of a puncture; interstriae two to
three times as wide as striae, shining, marked
by irregular lines, punctures almost as large as
those of striae, irregularly placed. Decli\it\
steep, broadly convex, strial and interstrial
punctures smaller than on disc; sutural inter-
striae feebly elevated, area from striae 1 to 3
flat on middle half. \'estiture of rows of short
strial and slightly longer interstrial hair, longest
interstrial setae slightly shorter than distance
between rows.
Female.— Similar to male except frons more
extensively, subcircularly impressed, U-shaped
carina as in male, vestiture on margins ;ibiin-
dant and \erv long.
Type Localitv.— Eight km S La Huerta.
Jalisco, Mexico.
Type Material.— The male holotype, female
allotype, and 62 parat)pes were taken at the
type locality on l-VII-65, 500 m. No. 161. from
axial pith tunnels in stems of a vine, by S. L.
Wood.
The holot\pe, allotype, and parat\pes are
in my collection.
Araptus genialis. n. sp.
Tin's species is distinguished from delicatus
Wood by the larger size, by the stouter body
form, and bv the sculpture of the frons.
Male.— Length 3.0 mm (paratypes 2.8-3.4
mm), 2.7 times as long as wide; color dark
reddish brown.
Frons deeply, triangularly impressed on
median three-fourths from epistoma to vertex,
upper angle more strongly impressed, its mar-
gins armed by one median and two lateral
acutely elevated cusps; floor of impression
punctate to obscurely aciculate, with a broad,
strongh' ele\ated carina from emarginate epis-
toma to deepest part of concavity, both upper
and lower ends tenninate abruptlv. .\ntennal
club ;vs in foveifrons.
Pronotum 1.16 times as long as wide; as in
delicatits except discal punctures smaller, not
as deep.
Elytra 1.7 times as long as wide, 1.4 times
as long as pronotum; as in delicatus except
discal interstriae three to four times as wide as
striae and decli\ital interstriae 3 feebly ele-
vated.
Female.— Similar to male exci'pt frontal im-
pression more extensive, its margins ornamented
b\' a dense fringe of long, yellow hair.
Type Locality— X'olean de Agua, Cuate-
m;ila.
Type Material.— The male holotype, female
allotspe, and 18 parat\pes were taken at the
t\ pe localitN' on 19-\'-64, 1000 m. No. 609 ( two
;ue 608), from axial pith tunnels in a cut vine,
b\' S. L. Wood.
The holotype, allotype, and paratypes are
in m\' collection.
Araptus dentifrons, n. sp.
This species is distinguished from confinis
( Blandford ) bv the impressed male frons, with
the carina much stronger, and with a series of
small tubercles arming the lateral and dorsal
margins of the impressed area, bv the different
female frons, and by the coarser elvtral punc-
tiues.
Male.— Length 1.8 mm (paratypes 1.6-1.9
mm), 3.0 times as long as wide; color very
dark reddish brown.
Frons subconcavely impressed to upper le\el
of eyes with median area on lower half very
strongly produced into a short, dentate, median
carina, upper and lateral margins of impressed
area anned b\' a row of about nine tubercles;
surface shining, subaciculate near epistoma;
vestiture sparse, inconspicuous except on
epistoma.
Pronotum as in confinis except posterior
areas with numerous impressed points ( obscure-
46
Uiiic.HAM Young University Science Bulletin
ly reticulate in some specimens), punctures
coarse, deep. Elytra as in confinis except discal
surface smooth, shining, with impressed points,
strial punctures larger, interstriae less than
twice as wide as striae, dccli\ital punctures
strongly reduced in size, almost obsolete.
Female.— Similar to male except frons
flattened, very shallowlv concave on median
third, with an obscure median impressed line
below, a weak carina above, marginal areas
subgranulate, not tuberculate, vestiture rather
uniformly distributed but more abundant and
longer on margins.
Type Locality.— Three km (2 miles) SE
Acatlan, Puebla, Mexico.
Type Materl\l.— The male holotype, female
allotvpe, and 21 paratvpes were taken on 15-
VI-6'7, 1500 m. No. 38,' from axial pith tunnels
of a cut vine, hv S. L. Wood; nine paratypes
are from \'olcan Cf)]ima, Jalisco, Mexico, 2'3-\T-
65, No. 104. S. L. W ood.
The holotype, allotype, and paratypes are
in mv collection.
Aroptits facetus, n. sp.
This species is distinguished from cxfitiisitus
Blackman bv the smaller size, bv the sliorter or
obsolete clvtral vestiture, and b\ the ornamen-
tation of the female frons.
Female.— Length 1..3 mm (paratvpes 1.2-1.4
mm), 2.5 times as long as wide; color reddish
brown.
Frons shallowlv concave from eye to eve
from epistoma to vertex, its surface smooth,
impunctate on lower fourth, finely, closely
punctured above, vestiture confined to area
above eyes, consisting of a dense brush of long
yellow hair.
Pronotum and elvtra as in r.\Y/t/(.vi7(;.$ except
elytra glal)rous.
Male.— Similar to female except frons as in
male exquisitus.
Type Locality.— Rio Teiiipisc(ue, Cuana-
caste, Costa Rica.
Type Material— The female holotype, male
allotvpe, and two paratvpes were taken at the
type locality on 25-III-64, 15 m. No. 501, from
Fictis twigs, by S. L. Wood.
The holotype, allotype, and parat\pes are
in my collection.
Arapttis ciispidis, n. sp.
This species and cnulitus
conspicuous median process
(Sdiedl) liavc a
at the posterior
margin of abdominal sternum 5 that distin-
guishes them from all other species in the genus.
This species is distinguislud from cnulitus by
the convex, unarmed eKtral declivity and by
the different female frons.
Female.— Length 1.3 mm (paratypes 1.2-1.4
mm), 2.6 times as long as wide; color yellowish
brown.
Frons shining. con\-ex, slightly flattened on
median half jjelow upper level of eyes, this area
punctured on its lateral and upper margins,
smooth and shining in central area, punctured
area with moderately abundant, short, incon-
spicuous hair.
Pronotum 1.0 times as long as wide; widest
on basal third, sides moderately arcuate, con-
verging toward rather narrowly rounded ante-
rior margin; anterior margin anned by about
12 low serrations; summit definite, very slightly
in frf)nt of middle; asperities moderately coarse,
confused; posterior areas strongly reticulate,
punctures small, obscure. Glabrous.
Elytra 1.5 times as long as wide, 1.6 times
as long as pronotum; sides straight and parallel
on basal two-thirds, rather narrowl)' rounded
behind; striae not impressed, punctures small,
distinct; interstriae three times as wide as striae,
almost smooth, impri'ssed points rather ob-
scure, impunctate. Declivit\- steep, convex;
sutural interstriae feebly ele\ated, 2 indistinctly
impressed; strial and interstrial punctures
minute, almost obsolete. \'estiture largely con-
fined to decli\itv, of interstrial rows of slender
bristles, each slightly longer than distance be-
tween rows, similarb' spaced within a row.
Sternum 5 with a conspicuoirs median eleva-
tion on posterior margin.
Mali;.— Similar to female except frons con-
vex, shining, surface obscurely, rather coarsely
punctured. Antennal club rather small, subcir-
cular, sutures rather strongly procurved, 1 ex-
tending one-third, 2 two-thirds club length from
base, 1 finely septate.
Type Locality.— Eight km (5 miles) E San
Bias, Na\arit, Mexico.
Type Material.- The female holotype, male
allotype, and 14 paraty^jes were taken at the
t\pe locality on 12- VI 1-65, 70 m. No. 2.33, from
;i broken tree branch, by S. L. Wood.
The holotype, allotvpe, and paratypes are
in my collection.
Araptus placatus, n. sp.
This species is distinguished from obsoletus
( Blandford ) by the larger size, by the more
Biological Series, Vol. 19, No. 1 New Species of American Bark Beetles
47
Strongly impressed, more finely punctured male
frons, by the short, sparse female frontal vesti-
ture, by the elytral vestiture, and by other
characters.
Male.— Length 1.7 mm (paratypes 1.5-1.8
mm), 2.S times as long as wide; color dark
brown.
Frons broadly impressed from epistoma to
upper le\el of eyes, upper limits attaining a dis-
tinct summit on median half; surface shining,
rather finely, deeply, closely punctured; an ob-
scure, median carina sometimes present; \esti-
ture fine, sparse, inconspicuous. Antennal club
as in ohsoletits.
Pronotum 1.13 times as long as wide; widest
just behind middle, sides weaklv arcuate, rather
broadly rounded in front; anterior margin
armed by about 10-12 low serrations; indefinite
summit one-third pronotum length from an-
terior margin; asperities small, confused; pos-
terior areas smooth to obscurely reticulate,
punctures rather fine, moderately close. Glab-
rous.
Elytra 1.7 times as long as wide, 1.6 times
as long as pronotum; sides straight and parallel
on more than basal two-thirds, rather broadi}'
rounded behind; striae not impressed except 1
near declivity, punctures small, spaced within
row by twice diameter of a puncture; interstriae
almost smooth, subshining, about four times as
wide as striae, impunctate. Declivity steep,
rather broadly conxex; striae 1 distinctlv im-
pressed, interstriae 1 distinctly ele\ated, 2
weakly impressed, interstrial punctures minute.
Vestiture confined to declivity, of rows of stout,
blunt interstrial bristles, each slightly shorter
than distance between rows, similarly spaced
within a row.
Female.— Similar to male except planocon-
vex, a median callus on lower half, area at
upper level of eyes not eknated; surface shin-
ing, rather finely, closely punctured, \estiture
very fine, moderateh- abundant, rather short.
Type Locality.— Five km (3 miles) W Jal-
tipan, X'eraciTiz, Mexico.
Type Material —The male holotype, female
allotype, and 27 paratypes were taken at the
type' locality on 25-\T-67, 50 m, Nos. 117, 100,
from the hollow central axis of a cut \'ine, by
S. L. Wood; two paratypes are from 37 km or
23 miles N Matias Romero, \'eracruz, 29-VI-67,
No. 126, S. L. Wood.
The holotype, allotype, and paratypes are
in my collection.
Araptus decorus, n. sp.
This unique species is distinguished by the
shallow declivital impression, confused elytral
punctures, slender fonu, and fine sculpture, and
by the very different female frons.
Male.— Length 1.9 mm (paratypes 1.8-1.9
mm), 2.8 times as long as wide; color yellowish
brown.
Frons convex, except weakly flattened on
central half below upper level of eyes, a small
callus near upper margin of flattened area, a
pair of calluses in ventrolateral parts of this
area; surface shining, rather finely punctured
except impunctate in median area on lower
half; vestiture fine, sparse, inconspicuous. An-
tennal club oval, sutures moderately proeurved,
suture 1 weakly septate.
Pronotum 1.25 times as long as wide; about
as in placatus \\'ood except median serrations
on anterior margin distinctlv larger, and poste-
rior area reticulate, dull, moderately fine, deep,
rather widely separated. Glabrous.
Elytra 1.6 times as long as wide, 1.3 times
as long as pronotum; outline ;is in placatus;
surface shining, with minute points, punctures
confused, ratiier small, moderately close. De-
clivity moderately steep, shallowh- impressed,
almost flat on median half; strial punctures in
rows; interstriae 1 weakh- ele\'ated, 2 distinctly
impressed; interstrial pimctures fine. N'estiture
eimfined lo decli\it\', ciHisisting of rows of
sparse, stout bristles except absent on 2, bristles
spaced in all directions by distances slightly
greater than length of a bristle.
Female.— Similar to male except frons flat-
tened almost from eye to eye from epistoma to
vertex, with an elevated, transverse, obtuse
summit just below upper level of eves, area from
summit to epistoma on median third smooth,
shining, impunctate, remaining area punctured
and ornamented by rather abundant, long,
yellow hair.
Type Locality.— Rincon de Osa, Puntarenas,
Costa Rica.
Type Material —The male holotype, female
;illotype. and seven parat\pes were taken at
the tvpe locality on ll-\Til-66, 30 m. No. 64,
from the phloem of a recently cut log 20 cm
in diameter, by S. L. Wood.
The holot)pe, allotype, and paratypes are
in mv collection.
Araptus blandittts, n. sp.
This species is distinguished from deijroUci
(Blandford) and schicarzi (Blackman) bv the
48
BlUGHAM VOUNG UNIVERSITY SCIENCE BlLIETIN
smaller size, bv the impunctate inteistriae, and
by the more deeply concave, more finely punc-
tured female frons, with longer, more abundant
frontal vestiture.
Female.— Length l.S mm, 2.6 times as long
as wide; color veiy dark reddish brown.
Frons moderately conca\e on central two-
thirds from distinctly ele\atcd epistoma to
slightly above eyes; surface smooth, shining,
very finely, closely, uniformly granulate-punc-
tate; vestiture of unifonnlv distributed, lather
abundant, fine, long hair, marginal setae dis-
tinctly longer. Antennal club oval, external
sutures obsolete, internal septum of anterior
half of suture 1 conspicuous.
Pronotum 1.1 times as long as wide; widest
on basal third, sides on basal half moderatel)
arcuate, distinctly constricted on anterior half,
rather broadly rounded in front; anterior mar-
gin aniied by aljout 12 low serrations; broad
summit near middle; asperities r;ither small,
confused; posterior areas smooth, shining, im-
pressed points very small, moderately abundant,
punctmes rather coarse, deep, moderately close.
Glabrous.
Elytra 1.55 times as long as wide, 1.65 times
as long as pronotum; sides almost straight and
parallel on l)asal two-thirds, rather narrowly
rounded behind; striae 1 moderately, others not
impressed, punctures moderately large, deep,
very close, rows slightly irregular; interstriae
smooth, shining, points obscure to absent, im-
punctate. Declivity moderately steep convex,
shallowlv suleate; sutural striae narrowly im-
pressed, interstriae 2 moderately impressed, as-
cending laterally, 3 higher than suture; strial
punctures slightly smaller than on disc; inter-
strial punctures small, obsolete on 2. Vestiture
largelx' confined to deeli\it\ . of interstri;d rows,
except on 2, of moderatel)' stout bristles, each
as long as distance between rows, similarly
spaced witliin a row.
Type Locality.— Fortin de las Flores, X'era-
cruz, Mexico.
Type Materl^l.- The female holotype was
taken at the type locality on 22-\'-65, 'at light,
bv D. Rabago.
The holotype is in mv collection.
Arapfus mcdialis. n. sp.
This species is distinguished by the im-
pressed elytral declivity, by the confused punc-
tures on the elvtral disc, and by the \ery dif-
ferent female frons.
Female.— Length 1.5 mm (paratypes 1.4-1.6
mm), 2.7 times as long as wide; color yellowish
brown.
Frons flattened almost from eye to eye from
epistoma to vertex, area below eyes dull,
densely, fineU' punctured, central area above
eyes smooth, shining, impunctate, entire lower
area and marginal areas above bearing abun-
dant, long, yellow hair, longest on upper mar-
gin. Antennal club small, oval, suture 1 weakly
procurved, septate throughout, 2 very obscure.
Pronotum 1.1 times as long as wide; widest
on posterior half, sides feebly arcuate on pos-
terior half, then distinctly constricted, rather
narrowlv rounded in front; anterior margin
arnu'd 1)V 10 coarse serrations, summit rather
Inroad, at middle; asperities rather coarse, con-
fused; posterior areas shining, almost smooth
to subreticulatc, impressed points rather ob-
scure, punctures fine, not sharjoly impressed,
rather sparse. Clabrous.
Elytra 1.5 times as long as wide, 1.4 times
as long as pronotum; sides almost straight and
parallel on slightly less than basal two-thirds,
rather broadly rounded behind; striae not im-
pressed; punctures rather small, confused on
more than basal half of disc, in rows posteriorly;
surface shining, with numerous impressed
points. Declix'itv rather steep, strongly suleate;
strial punctures in rows, small, rather obscure;
interstriae 1 weakly elevated, 2 impressed,
rather abruptly, moderately elevated laterally,
lateral margins rounded, distinct!)- higher than
suture, interstriae 1 and 3 each bearing a row
of small granules. \'estiture confined to de-
eli\itv, consisting of rows of rather long, slender
bristles except absent on interstriae 2.
Male— Similar to female except 2.5 times
as long as w ide; frons convex, shining, a small
median callus at upper level of eyes, punctures
small, shallow, rather close, \'estiture fine,
sparse, inconspicuous; decli\ital bristles stout,
blunt, shorter, each distinctly longer than dis-
tance between rows, similarh' spaced within a
row.
Type Locality.— San Isidro del Ceneral, San
Jose, Costa Rica.
Tyi'e Matkiual— The female holotype, male
allotype, and 21 paratypes were taken at the
t\pe locality on 5-XII-63, 1000 m. No. 288,
from a tree branch by S. L. Wood.
The h()lot\pe, allotype, and paratypes are
in my collection.
Araptus condittis. n. sp.
This species is distinguished from medialis
Wood bv the larger size, by the absence of the
Biological Series, Vol. 19, No. 1 New Species of American Babk Beetles
49
lateral half of the septiini of suture 1 of the
antennal club, bv the eoar.ser punctures on the
pronotum and elytra, and l)y the \ery different
frons in both sexes.
Female.— Length 2.0 mm ( paratopes 1.9-2.2
mm), 2.4 times as long as wide; color yellowish
brown.
Frons shallowly, evenly concave from eye to
eye from epistoma to vertex, denselv, finely,
uniformly punctured, co\ered bv abundant
erect, short, )elIow hair of uniform lengtii ex-
cept slightly longer at margins. Antennal club
moderatelv large, subcircular, suture 1 strongly
subangulate, its median lialf septate, suture 2
almost obsolete.
Pronotum 1.12 times as long as wide; sides
feebly arcuate and almost parallel on basal
half, slightly constricted anterioiK', rather nar-
rowly rounded in front; anterior margin acute,
subserrate; summit indefinite, at middle; as-
perities fine, confused, covering anterior two-
thirds; posterior areas shining, obscureh' reticu-
late, punctures moderatelx coarse, close, deep.
Glabrous.
Elytra 1.4 times as long as wide, 1.26 times
as long as pronotum; outline and disc as in
medialis except striae 1 impressed, punctures
rather coarse, deep. Declivity sti'ep, bisuleate;
interstriae 1 moderately ele\'ated, 2 rather
stronglv impressed, narrowly abo\'e, broadlv be-
low, lateral areas rather strongh- ele\ated,
rounded, higlier than suture, 1 and .3 each \\ ith
a sparse row of granules; punctures in rows,
rather small, shallow. \'estiture largely confined
to decli\itv, of interstrial rows (absent on 2) of
short, coarse bristles, each almost as long as
distance between rows, similarh' spaced within
a row.
Male.— Similar to female except frons broad-
ly, transversely impressed from epistoma to
upper le\-el of eyes, a rounded median summit
at its upper limits; surface smooth, finelv, \er\-
closely, uniformlv punctured; \'estiture of rather
abundant, fine, short, semirecumbent hair.
Type LocALiTi\— Puerto X'iejo, Heredia,
Costa Rica.
Type Material.— The female holot\pe, male
allotype, and 14 paratypes were taken at the
type' locality on 12-111-64, 70 m. No. 478 (except
one is No. 481), from radiate tunnels in the
cambium of a cut woody \'ine, bv S. L. Wood.
One paratype is labeled Turrialba, Costa Rica,
1970, R. I. Gara.
The holotype, allotvpe, and paratypes are
in my collection.
Araptus frugalis, n. sp.
This species is distinguished from conditiis
Wood by the minute pronotal and elvtral punc-
tures, by the much more abundant elvtral ves-
titure, bv the glabrous female frons, and b\ the
antennal club.
Female.— Length 2.1 mm (paratypes 1.9-2.2
mm), 2.7 times as long as wide; color vellowish
brown.
Frons essentially as in male condifu.s except
more brightlv polished, less closeh' punctured,
subglabrous, with a definite median tubercle
at upper level of e^â– es. Antennal club elongate-
oval, anterior half of suture 1 represented bv a
longitudinal septum extending one-third of
club length from base, 2 indicated exteniall)',
profouiidh prociu\'ed, extending three-fourths
of club length from base.
Pronotum 1.2 times as long as wide; about
as in conditus except summit distinetlv anterior
to middle, anterior margin armed h\ 18 rather
coarse serrations, posterior ar(>as with punetiu'es
much smaller. \'estiture of moderatcK- abundant
fine, short hair.
l']lytra 1.5 times as long as wide, 1.'3 times
as long as pronotmii; outline as in conilitiis;
striae not impressed, punctures verv fine, mostly
in rows; interstriae four or more times as wide
as striae, shining, with impressed points and
\erv fine, irregular lines, punctures as small or
smaller than those of striae, in rows. Deeli\itv
steep; sulcate; essentially as in conditris except
punctures on striae 1 and 2 obsolete, others
minute. Wstiture of rows of minute strial hair,
and longer, rather fine, interstrial bristles on
disc and declivity, l^ristles about as long as dis-
tance between rows, similarly spaced within a
row, absent on declivital interstriae 2.
Male.— Similar to female except frons con-
vex, rather coarsely, closely punctured, median
area weakly elevated and impunctate; ehtral
bristles slightly longer, stouter.
Type Locality.— Rincon de Osa, Puntarenas,
Costa Rica.
Type Material.— The female holotype, male
allotype, and .36 paratypes were taken at the
tvpe locality on ll-Vni-66, 30 m. No. 80, from
a cut vine 2 cm in diameter, by S. L. Wood.
The holotype, allotvpe, and paratvpes are in
my collection.
Araptus laudatus, n. sp.
This species is distinguished from frugalis
Wood by the narrow, convex frons in both
sexes, with a distinct impression just above
50
Bhigham Voung University Science Bulletin
the epistoma in the female, by the more nar-
rowly, deeply silicate elytral declivity, and by
the subtriangular antennal club.
Female.— Length 2.0 mm (paratypcs 1.7-2.1
mm), 2.8 times as long as wide; color reddish
brown.
Frons convex except kjwer third rather
strongly, transversely impressed; epistoma with
median area distinctly elevated; surface dull,
reticulate, rather coarsely, somewhat obscurelv
punctured; glabrous. Antennal club subtriangu-
lar, apex somewhat pointed; mesal half of suture
1 septate, sutures not indicated externally.
Pronotum 1.2 times as long as wide; sides
almost straight and parallel on basal half, rather
narrowly rounded in front; anterior margin
armed by 10-12 low serrations; summit rather
indefinite, one-third pronotum length from an-
terior margin; asperities rather fine, confused;
posterior areas smooth, shining, with numerous
impressed points, punctures rather small, mod-
erately close. Glabrous.
Elytra 1.8 times as long as wide, 1.6 times
as long as pronotum; sides almost straight and
parallel, then slightly tapered, bisinuateh
truncate on median half behind; striae not im-
pressed; all punctures rather small, confused;
surface smooth, shining, with minute impressed
points. Declivity narrow, steep, broadly, rather
deeph' sulcate; inti'rstriae 1 distinctly elevated,
2 impressed, lateral areas abmpth', strongh'
elevated, rounded, armed on inner margin bv
a series of two to six small denticles, 1 usuallv
with one or more small granules on basal half;
punctures on striae 1 and 2 obscure to obsolete.
Vestiture largely confined to declivity, consist-
ing of a few, short, interstrial bristles.
Male.— Similar to female except frons moix*
evenly convex, transverse impression almost ob-
solete, punctures coarser, deeper; serrations on
anterior margin of pronotum larger; decli\'ital
sulcus deeper, more abrupt, sutural interstriae
more regularly gramdate.
Type Locality.— San Isidro del General, San
Jose, Costa Rica.
Type Material.— The female holotvpe, male
allotype, and 48 paratypes were taken at the
type locality on 1.3-Xl'l-6:3, 1000 m. No. .31.3,
from a broken tree branch, by S. L. Wood.
The holotvpe, allotype, and paratvpes are
in mv collection.
Arapttts vesctilus, n. sp.
This species is distinguished from exiiiioUs
Wood bv the smoother elvtral surface and bv
the very different frons, particularly in the
female.
Female.— Length 1.2 mm (parat\pes 1.1-1.3
mm), 2.5 times as long as wide; color \erv dark
reddish brown.
Frons almost flat, median line above eves
weakly elevated, epistonial margin rather
weakly elevated medially; surface reticulate,
punctures rather coarse, close, .sparsely punc-
tured toward center; vestiture of rather sparse,
very long hair. Antennal club subcircular, suture
1 moderatelv proeurxi-d, weakly septate, 2 ob-
scure.
Pronotum 1.1 times as long as wide; as in
eruditus ( Schedl ) except posterior areas shining,
weaklv reticulate, punctures fine, shallow,
rather widely spaced. Almost glabrous.
Elytra 1.5 times as long as wide, 1.4 times
as long as pronotum; outline as in eruditus;
striae not impressed, punctures fine, deep; inter-
striae almost smooth, shining, twice as wide as
striae, impressed points almost obsolete, im-
punetate. Declivity steep, sulcate; essentially as
in eruditus. Vestiture as in eruditus except
less abundant.
Male -Similar to female except frons more
distincth', more evenlv convex, punctures
coarser, more evenly distributed, almost gla-
brous.
Type Locality.— Guapiles, Limon. Costa
Hiea.
Type Mateiual.— The female holotype, male
allotype, and two paratvpes were taken at the
type' locality on 22-Vn-66, 100 m. No. 121, from
a cut x'ine, by S. L. Wood.
The holotvpe, allotype, and paratypes are
in in\' collection.
Araptus exigialis, n. sp.
This species is distinguished from vesculus
Wood by the more abundant, more conspic-
uously impressed lines on the elvtral disc, by
the more slender form, and b\' the different
frons in both sexes.
Female.— Length 1.3 mm (paratypes 1.3-1.4
mm), 2.7 times as long as wide; color reddish
brown.
Frons shallouK, liroadK concave from epis-
toma to ab<)\(' upper level of eves; surface
shining, minutely, densely, uniformly "punc-
tured; vestiture of fine, very short, abundant
hair, very slightly longer on margins. Antennal
club as vesculus.
Pronotum 1.2 times as long as wide; as in
vesculus except posterior areas very finely.
Biological Series, Vol. 19, No. 1 Np:w Species of American Bahk Beetles
51
rather strongly reticulate, punctures very fine,
shallow, rather sparse. Glabrous.
Elytra 1.7 times as long as wide, 1.4 times as
long as pronotum; as in vescultts except discal
surface with points and fine, irregular lines.
Male.— Similar to female except frons
strongly convex, weakly impressed along upper
margin of epistoma; surface reticulate, punc-
tures rather coarse, close, shallow, vestiture
fine, short, sparse, inconspicuous.
Type Locality.— Fort Clayton, Canal Zone,
Panama.
Tyi'e Material.— The female holot\pe, male
allotype, and 10 paratvpes were taken at the
type localit)- on 22-XII-63, 30 m. No. 31S, from
a cut woody vine, by S. L. Wood.
The holotype, allotype, and paratypes are
in my collection.
Ar(ii)tus refertus. n. sp.
This species is distinguished from the pre-
ceding members of this genus by the much
coarser strial punctures, with impunctate discal
interstriae, by the rather strongly bisulcatc ely-
tral declivity, and by the frons in both sexes.
Female.— Length 2.0 mm (paratypes 1.9-2.3
mm), 2.8 times as long as wide; color light
brown (mature color probabK dark brown).
Frons broadly, weakly convex; surface shin-
ing, smooth and impunctate on triangular area
on lower, median half, finely, closely, rather
deeply punctured on lateral and upper areas;
vestiture of rather abundant, moderately long,
fine hair. Antennal club oval, suture 1 mod-
erately procurved, septate, not reaching middle
of club, suture 2 \irtuallv obsolete, near apical
margin.
Pronotum 1.13 times as long as wide; as in
vesculus Wood except posterior area smooth
shining, with numerous impressed points, punc-
tures moderately coarse, deep, close. Glabrous.
Elytra 1.7 times as long as wide, 1.5 timi's
as long as pronotum; sides straight and par-
allel on basal two-thirds, rather broadly rounded
behind; striae not impressed, punctures coarse,
deep, close; interstriae as wide as striae, smooth,
shining, with moderately numerous impressed
points, impunctate. Declivity steep, broadh',
rather strongh' blsulcate; strial punctures im-
pressed, slightly smaller than on disc; inter-
striae 1 moderately, gradually elevated, 2 rather
strongh- impressed, flat, narrow at base, rather
broadly expanded on middle half, 3 rather
abruptly, moderately elevated, 1 and 3 each
armed on basal half bv about three very small.
pointed, semirecumbent denticles, a few punc-
tures on lower half. \'estiture sparse, hairlike,
mostly on or near declivity.
Male.— Similar to female except frons
strongly, evenly convex; surface shining, rather
coarsely, deeply, closely punctured, subglabrous
except on and near epistoma; elvtral declivity
more strongly impressed, denticles slightly
larger.
Type Locality.— \'olcan Zunil, Quezalten-
ango, Guatemala.
Type Material.— The female holot\pe, male
;illotvpe, and 16 paratypes were taken at the
t>T3e locality on 27-\'-64, 1000 m. No. 625, from
a composite shrub, by S. L. \\'ood.
The holotype, allotype, and paratypes are
in in\ collection.
Araptus trepidus. n. sp.
This species is distinguished from the closely
allied poricoUis (Blandford) by the absence of
declivital denticles, and by the almost con-
tinuous trans\erse epistom;d rle\ation in the
male.
Female.— Length l.S mm (paratypes 1.6-
1.8 mm), 2.7 times as long as wide; color dark
reddish brown.
Frons broadly flattened from epistoma to
vertex, median third of lower half smooth, shin-
ing, distinctly ele\'ated, remaining area densely,
finely, deeply punctured; vestiture on punctured
area abundant, fine, rather long, unifomily
distributed.
Pronotiun 1.15 times as long as wide; as in
refertus Wood except median line of posterior
area impunctate.
Elytra 1.6 times as long as wide, 1.5 times
as long as pronotum; outline and disc as in
refertus except impressed points obscure to ob-
solete. Decli\it\- steep, bisuleati'; interstriae 1
abruptly, distinctly elevated, 2 rather strongly
impressed, flat, wider than 1 or 3, 3 abruptly,
moderately ele\'ated, 1 and 3 finely punctured,
not granulate; strial punctures small, rather ob-
scure. \'estiture of rows of very minute strial
hair, and longer, slender interstrial bristles on
disc and declivity, each slightly longer than dis-
tance between rows, similarly spaced within a
row.
Male.— Similar to female except frons
strongly convex on upper half, strongly, trans-
versely impressed just below middle, lower mar-
gins of impressed area elevated laterally, ele-
vations continue to a weak median subcarinate
elevation dividing impression; upper area of
52
BlllCHAM "loUNG Un|V|:HS1TY ScIENCK Bui.l-ETIN
frons coarsely, deeply punctured, \estiture
sparse except for cpistomal brusli.
Type Locality.— X'olcan de Agua, I'^sfiuiTitla,
Guatemala.
Type Mateiual.— The fi'iiiale holotspe, male
allotype, and 56 paratypcs were taken at the
type locality on 19-\'-64, 1000 m, Nos. 608
(type), 609, 612, from Mcnis))criua sp., bv S. L.
Wood.
The holotype, allotype, and paratspcs arc
in my collection.
Araplus fioiilalis. n. sp.
This species is distiuiimshcd troni trcpidus
Wood hv the larger size, bv the punctured de-
clivital intersfriae 2, and by the different fions
in both sexes.
Female.— Length 2.7 nun (paratypes 2.4-2.7
mm), 2.8 times as long as wide; color dark
reddish brown.
Frons broadly flattened from epistoma to
vertex, weakly concave on small median area
above eyes, weakly convex on small, median,
impimctate area on lower half; surface shining,
finely, closely, deeply pmictured; \-estiture of
moderately abundant, fine, long hair. .Antenna
as in trcpidus.
Pnjnotum LI times as long as wide; essen-
tially as in trcpidus.
Elytral outline and disc as in Ircpidu.'; ex-
cept interstriae near declivity witli pimctures.
Declivity about as in trcpidus and refertus ex-
cept interstriae 2 with a row of punctures as
coarse as those of striae; lateral margins without
granules, ^'estiture confined to decli\itv, of fine,
sparse hair.
Male.— Similar to female except frons deep-
ly, transversely conca\i' on upper half of area
below \ipper level of e\ es on median three-
fourths, lateral and lower margins of concavity
strongly, acuteb' carinate except carina inter-
rupted at median line, vestiture inconspicuous
except on epistoma; anterior margin of prono-
tuni rather coarsely serrate; declivitx' more
strongly impressed.
Type Locality.— N'olcan Zunil, Quezalten
ango, Guatemala.
Type Material.— The female hcjlotype, male
allotvpe, and five paratypes were taken at the
type locality on 27-\'-fi4; 1000 m, Nos. 625, 626.
from a composite shrub, by S. I^. ^Vood.
The holotype, allotype, and paratypes ari'
in my collection.
:\raplus niij^rcllu.s, n. sp.
This and the following three form a dis-
tincti\e transitional group between species pre-
\iously placed in Ncodrt/acoetcs Eggers and
SpJicroceros Schedl. The\ have the antennal
club large, with finely marked, strongly pro-
curved sutures, the costal apex of the elytra as-
cends slighth', the declivity is convex, and the
elytral \'estiture tends to be confined to the de-
clivity, closely spaced and more or less scale-
like. This species is distinguished from others in
the group b\' the more slender body, bv the
slender interstrial l)rist]es, and by the subgla-
brous female frons.
I'^kmalk. —Length 1.9 mm (paratypes 1.7-2.0
mm), 2.7 times as long as wide; color black.
Frons convex, a weak, transxcrse impression
above epistoma; surface strongly reticulate-sub-
granulate above eyes, almost smooth and with
;i few small punctures below. Antennal club
rather large, oval, sutures strongly arcuate, 1
septate.
I'ronotuTH 1.1 limes as long as wide; widest
on basal third, siiles moderately arcuate, con-
verging to\\;u(l nanowb' rounded serrate an-
terior margin; summit at middle; asperities
moderately coarse, confused; posterior areas
shining, partly sul)reticulate, with low, longi-
tudinal subasperate crenulations continiu'ng al-
most to base, pimctures obscure, associated with
erenu];itions. Cllabrous.
Elytra 1.7 times as long as wide, 1.7 times
as long as pronotum; sid(\s straight and par-
allel on slightly less than basal two-thirds;
tapered, then ratlier broadly rounded; striae
not impressed except 1 wi'akly, punctiues small,
deep, spaced within row by diameter of a punc-
ture; interstriae four times as wide as striae,
almost smooth, with obscme, minute points,
impunetate. Decli\it\ moderately steep, con\'e\;
striae obsciu'elv impressed, punctures smaller
and less distinct than on disc; interstriae each
with a row of very fine granules. X'estiture con-
fined to ilecli\it\', of rows of narrowly spatu-
late interstrial bristles, each bristle as long as
distance betwcH'ii rows, more closely spaced
within a row.
Male— Similar to female in all respects.
Type Locality— Ten km SE Cartago, Car-
tago, Costa Rica.
Type Material. —The female holotype, male
allot\pe, and 67 parat\pes were taken at the
type locality on 24-L\-63, No. 205 (type), and
.3-VII-6:3, Nos. L3 and 15, 1800 m, from Mtjrica
ptdjcscens. by S. L. Wood,
Biological Series, Vol. 19, No. 1 New Species of American Bark Beetles
53
The liolotype, allotype, and paratypcs are
in my collection.
Araptus vinnulus, n. sp.
This species is distinguished from nigrclhis
Wood by the smaller size, b\' the stouter form,
by the more widely spaced, stouter interstrial
bristles, by the almost obsolete strial punctures,
and bv the frons.
Female.— Length 1.3 mm (paratypes 1.2-1.5
mm), 2.4 times as long as wide; color very
dark brown, almost black.
Frons broadly convex, lower third with
broad, smooth, shining, median line; surface
densely, coarsely, deeply punctured; upper two-
thirds on median two-thirds with moderatel)'
abundant, fine, long hair. Antenna essentially
as in ninrellus.
Pronotum 1.0.3 times as long as wide; as in
ni<i,rclhts except less strongly tapered anteriorh-,
moderately rounded in front, posterior areas
dull. Glabrous.
Elytra 1.44 times as long as wide, 1.44 times
as long as pronotum; sides straight and parallel
on basal half, tapered then rather narrowly
rounded behind; striae not impressed, punctures
minute, sliallow, many almost obsoU>te; inter-
striae shining, with many very minute im-
pressed points and shallow, irregular lines, im-
punctate. Declivity rather steep, convex; as on
disc except interstriae each with a row of small,
rounded granules. Vestiture confined to de-
clivity, of rather stout, spatulate bristles, each
slightly shorter than distance between rows,
similarly spaced within a row.
Male.— Similar to female except frons rather
strongly convex, surface strongh' reticulate, sul)-
glabrous.
Type Locality.— San Ignacio de Acosta, San
Jose, Costa Rica.
Type Materl^l.- The female holotvpe, male
allotype, and 18 paratypes were taken at the
type locality on 5-VII-63, 1500 m, No. 31, from
small branches of Roupala complicafo, bv S. L.
Wood.
Araptus ftiivus. n. sp.
This species is distinguished from vinntihis
Wood by the larger size, bv the more closeh
spaced interstrial setae, by the presence of
stiial hair, and by the longer more abundant
setae on the female frons.
Female.— Length 2.1 mm (paratypes 1.7-2.2
mm), 2.4 times as long as wide; color almost
black.
Frons convex, strongly reticulate, punctures
rather small, sparse; upper two-thirds with
rather dense marginal fringe of long hair, longest
setae on vertex extend aliout two-thirds dis-
tance to upper level of eyes. Antenna as in
vinnulus.
Pronotum 1.03 times as long as wide; as in
vinnulus except more distinctly constricted on
anterior half.
Elytra 1.5 times as long as wide, 1.6 times
as long as pronotum; as in vinnulus except
strial punctures on basal half of disc more dis-
tinct; decli\ital strial punctures obsolete. \'es-
titure confined to decli\'it\', of rows of minute,
fine, strial hair, and rows of longer, spatulate,
interstrial bristles, each bristle slightly shorter
than distance between rows, more closely
spaced within a row, each bristle very slender
on its basal half, flattened on its apical half.
Male.— Similar to female except frons bioad-
Iv convex, glabrous.
Type Locality.— Corro Punta (labeled \'ol-
cau Chiritjui), Chiritjui, Panama.
Type Material.— The female holotype, male
allotype, and 45 paratypes were taken at the
typt>' locality- on 11-L64, 1800 m, No. 422, from
broken Roupala sp. limbs, by S. L. \\'ood.
The h()lot\pe, allotype, and paratypes are
in m\' collection.
Araptus furvescens. n. sp.
This species is distinguished from furvus
Wood b\ the smaller average size, by the feebly
impressed declivital striae, with minute strial
punctures indicated, and by the longer female
frontal pubescence, with pubescent area ex-
tending higher on the vertex.
Female.— Length 1.9 mm (paratypes 1.5-1.9
mm); as in furvus Wood except female frontal
pubescent area extending higher on \ertex, dis-
tance from upper level of eyes to upper limits
of puliescent area 1.5 times distance from epis-
toma to iipprr level of e\es, longest setae on
vertex almost reach epistomal margin, pubescent
area extending \'entrad in lateral areas almost
to epistoma; declivital striae weaklv impressed,
punctures very small but usually visible; de-
clivital interstrial bristles more nearly scalelikc,
each flattened on two-thirds or more of its
length.
Male.— Similar to female except frons more
broadly convex, glabrous.
54
BhIOUAM VoUNC. UNIVtJlMTY SCIENCE Bui.LETIN
Type Locality.— Volcan Pacaya, Estjuintla,
Guatemala.
Type Material.— Tlie female holot)pe, male
allotype, and 40 paratvpes were taken at the
type locality on l-\'I-'64, 1300 m, Nos. f.52,
657, 658, and cut Roupahi limbs, bv S. L.
Wood.
The holotype, allotype, and paratypes are in
my collection.
Arapfus lepidus, n. ,sp.
This species is distinguished from the above
three .species in this group by the presence of
discal interstrial punctures and by the stouter
bodv form. It is distinguished from aztccus
Wood b)' the narrowly rounded, serrate anterior
margin of the female pronotum and by the very
different female frons.
Female.— Length LS mm (paratypes L7-2.0
mm), 2.26 times as long as wide; color dark
brown.
Frons con\e.\, median arini strongly im-
pressed from slightly above upper level of eyes
to just abo\e epistoma, lower lateral margins of
impressed area moderately elevated; surface
shining, smooth, punctures rather fine, deep,
sparse; glabrous except near epistoma. Antennal
club very large, about as in nigreUus Wood, onl\'
median half of suture 1 septate.
Pronotum L05 times as long as wide; es-
sentially as in vinnuhis Wood except crenula-
tions or ridges on posterior half almost obso-
lete, posterior areas smooth, sliining, punctures
very small, shallow, rather sparse. Glabrous.
Elytra L.3 times as long as wide, L2 times
as long as pronotum; sides almost straight and
parallel on basal half, ratlier broadh' rounded
behind; striae not impressed, punctures small,
shallow, rows irregular on anterior two-thirds;
interstriae smooth, shining, about four times
as wide as striae, punctures uniseriate, similar
to those of striae. Declivity steep, convex, sculp-
ture essentially as on disc except punctures
slightly smaller, deeper. Vestiture largely con-
fined to posterior half, consisting of interstrial
rows of slender sealelike bristles, each slightly
shorter than distance bc^twcen rows, similarly
spaced within a row.
Type Locality.— \'olcan, Puntarenas, Costa
Rica.
Type Material.— The female holotype and
three female paratvpes were taken at the type
locality on ll-Xn-63, 1000 m. No. 304, from a
broken branch, by S. L. Wood.
The holotype and paratypes are in my col-
lection.
Araptus mcndictis. n. sp.
This species is distinguished from medialis
\\'ood bv the smooth, polished male frons, with
much finer, less abundant punctures, by the
finely reticulate, much more distinctly punc-
tured female frons, with the pubescence slight-
ly less abundant, finer, and shorter, and by the
larger size.
Female.— Length l.S mm (paratypes L7-L9
mm), 2.8 times as long as wide; color rather
dark reddish brown.
Frons broadh', evenly convex, reticulate and
finch', distinctly, closely punctured on lower
two-thirds, almost smooth and shining in small
area above eyes; vestiture fine, moderately long,
rather abundant, distinctly longer on margins,
particularly abo\e, shining area above eyes sub-
glabrous. Antennal club about as in medialis
except slightly wider.
Pronotum and elytra as in medialis.
Male.— Similar to male medialis except frons
glabrous, smooth, polished, shagreened, punc-
tures much smaller, less abundant.
Type Locality.— Cartago, Cartago, Costa
Rica.
Type Material.- Tiie female holotype, male
allotype, and 22 paratypes were taken at the
t\pe loe;ility XII-1965, from Avocado (Persea
amcricana ) branches, by N. L. H. Krauss.
Tlie holotype, ;illot\pc, and paratypes are
in m\' collection.
Arapliis n'tnulus, n. sp.
This species evidently is not closely allied
to an\' known species. It is distinguished by the
small size, by the simpli', unadorned frons in
both sexes, by the greatly reduced, almost obso-
lete strial punctures, by the regular, almost
scalelike interstrial bristles, and bv other char-
acters.
Female.— Length 1.2 mm (paratypes 1.1-1.3
mm), 2.6 times as long as wide; color dark
reddish brown.
Frons convex, feebly impressed just above
epistoma in lateral areas; surface shining, rather
coarsely, sparsely punctured and with more
numerous minute points; vestiture fine, sparse,
inconspicuous. Antennal club oval, as long as
scape, sutures almost straight, suture 1 finely
septate.
Pronotmn 1.06 times as long as wide; widest
near base, sides weakly arcuate on basal half,
Biological Series, Vol. 19. No. 1 New Species of Americ-^n B.\rk Beetles
55
feebly constricted on anterior half, rather nar-
rowly ronnded in front; anterior margin serrate,
about 12 indistinct serrations; summit indistinct;
Granulations on anterior slope rather coarse,
continued as low, irregular transverse costae to
well behind summit; posterior areas shining,
with numerous impressed points, appearing al-
most subgranulate in lateral areas. Vestiture
erect, moderately abundant, somewhat similar to
elytral setae.
Elytra 1.6 times as long as wide, 1.5 times
as long as pronotum; striae not impressed, mi-
nute punctures almost obsolete; interstriae shin-
ing, setiferous punctures confused with numer-
ous impressed points of almost equal size. De-
clivity rather narrowly con\'e.\; interstriae 1
shallowly impressed; surface sculpture as on
disc. N'estiture of interstrial rows of suberect
apically flattened bristles; each bristle about as
long as distince between rows, somewhat more
closely spaced witliin a row.
Male.— Similar to female in all respects.
Type Locality.— Tampico, Tamaulipas, Mex-
ico.
Type Material.- The female holotype, male
allotype, and 12 paratypes were taken at the
type locality on 14-\TII-72, from Disholcapsis
galls on Quercits vir^inkina, by G. F. Frankie.
The holotype, allotype, and paratypes are
in my collection.
Pseudopitijophthonis festivus, n. sp.
This species is distinguished from pulvereus
Blackman by the planoconcave male frons, by
the more strongly impressed elytral declivitv, by
the more coarsely punctured elytral surface, and
by the very different ehtral vestiture.
Male.— Length 1.8 mm (paratvpes 1.6-1.8
mm), 2.8 times as long as \\ide; color dark
brown.
Frons broadly planoconca\'e from epistoma
to vertex; surface smooth, shining in central
area, marginal are;is finely punctured and bear-
ing a dense tuft of long, \ellow hair. .Antenna as
in pulvereus.
Pronotum 1.2 times as long as wide; as in
pulvereus.
Elytra 1.6 times as long as wide, 1.3 times as
long as pronotum; outline as in pulvereus; sur-
face smooth, shining, punctures moderately
coarse, shallow, confused, very close, width of
interspaces about equal to diameter of punc-
tures. Declivity steep flattened; moderately ele-
vated at suture, a moderately strong sulcus in
area of interstriae 2, lateral convexities rounded;
punctures minute. Ground vestiture of fine,
short, stout, reclining setae; erect scales in in-
terstrial rows except obsolete on lower half of 2
on declivity, each scale about as long as dis-
tance between rows, similarly spaced within a
row, each about four to eight times as long as
wide.
Female.— Similar to male except frons plano-
convex, pubescence sparse, fine, much shorter.
Type Locality.— Eleven km or 7 miles NE
Copala, Sinaloa, Mexico.
Type Material.— The male holotype, female
allot\pe, and two parat>pes were taken ;it the
t\'pe locality on 22-VII-53, from a {hiercus
branch, by S. L. Wood.
The liolotype, allotype, and paratypes are
in mv collection.
Gnathotrichus ohsctirus, n. sp.
This and primus (Bright) are the only spe-
cies in the genus ha\ing confused punctures on
the eltyral disc. It is distinguished from primus
by the minutely reticulate el\ tral surface, by the
smaller size, by the steeper declivity, and by
other features.
Female.— Length 2.5 mm, 3.0 times as long
as wide; color reddish brown.
Frons broad, broadh' convex; surface shining,
finely, convergently aciculate on lower two-
thirds, rather finely, sparsely punctured above,
a strongly developed rather low median carina
from middle to upper level of eyes ( concealed
by pronotum above e\es). Vestiture of fine,
rather sparse hair.
Pronotum 1.24 times as long as wide; as in
dentatus Wood except posterior areas reticulate,
finch', closely, rather deeply punctured.
Elytra 1.7 times as long as wide; sides
straight and parallel on basal two-thirds, narrow-
ly rounded behind; striae obsolete; surface mi-
nutely reticulate, punctures minute, obscure,
confused, a few granules on interstriae 3 near de-
elixitw Deeli\it\' steep, broadly eonxex; \'ery
feebly sulcate on upper half, strial punctures
not e\'ident, interstriae marked by rows of very
small granules, slightly larger on 3; costal mar-
gin at apex rather strongly elevated. Vestiture of
fine hair, on disc of short, moderately abundant
hair and interstrial rows of very long fine hair;
all hair on declivity long, moderately abundant.
Type Locality.— Nine km NE Tezuitlan,
Puebla, Mexico.
Type Material.— The female holotype was
taken at the type localit\ on 2-VII-fi7.' IPOO m.
m
Bnic;iiA.\i VouNc Univehsitv Science Bulletin
No. 152, from a (^tienits log 15 cm in diameter,
by S. L. Wood.
The holotype i.s in mv collection.
Gnathotrichus omissus, n. sp.
This species is distinguished from sulcafti.s
(LeConte) by the minute to obsolete strial
punctures on the disc, by the confused punctures
on the elytral declivity, by the feebh' impressed
elytral declivity, by the much less stronijly acicu-
late female frons, and by the hosts.
Female.— Length 3.5 mm (paratvpes 3.3-3.6
mm), 3.2 times as long as wide; color dark
brown, usualiv witli pale areas on anterior pro-
notum and basal and dccli\'ital areas of elytra.
Frons convex, median line obscurely ele-
vated, surface smootli, shining, sparselv, finely
punctured, median third below upper li'\el of
eyes finelv acicniate; \estiture fine, sparse, in-
conspicuous.
Pronotum 1.3 times as long as wide; essen-
tiallv as in matcriaritis (Fitch) except serra-
tions on anterior margin coarse.
Elytra 1.8 times as long as wide, 1.5 times
as long as pronotum; outline as in matcriaritis:
striae not impressed, punctiues small, in slighth"
irregular rows; interstriae smooth, shining, im-
pressed lines and points moderateh abundant.
Declivity rather steep, convex; weakly, ratluM
narrowlv sulcate; strial punctures obsok'te or
nearly so, e\identlv confused; lateral con\exities
distinctly higher than suture, summit armed by
two or three small granules. X'estiture confined
to declivity, consisting of sparse hair.
Male.— Similar to fcMiKile I'xcept frontal acic-
ulation much stronger and distributed almost
from eve to oyc to well abo\'e eyes; antenna
without long hair; anterolateral angles of prono-
tum bearing a small tuft of hair.
Type Locality.— Volcan Irazu, Cartago, Cos-
ta Rica.
Type Material.— The female holotype, male
allotype, and 38 paratvpes \\ere taken at the
t)pe'locality on 26-1X-63, 2300 m elexation. No.
207, from a broken limb of Orcopanax mihi<ien-
tis, by S. L. \\'ood. One other specimen, not
included in the type series is from \'olcan Paca-
ya, Guatemala.
The holotype, allotype, and paratypes are in
my collection.
Gnathotrupcs dihitus. n. sp.
This species is distinguished from crccentis
Wood by the broadly convex unarmed elytral
declivity, with longer declivital pubescence, by
the larger frontal punctures, and by the smaller
size.
Male.— Length 1.7 mm (paratypes 1.6-1.7
mm), .3.2 times as long as wide; color dark
brown.
Frons broadK' convex, shining, weakly reticu-
late; punctures sparse, moderately coarse, shal-
low; epistomal area distinctly impressed except
on median line; xestiture sparse, hairlike, incon-
spicuous. Antennal club subcircular, three
strongl\- arcuate sutures marked by grooves and
rows of setae, segment 1 very small.
Pronotum 1.2 times as long as wide; sides
feebh' arcuate, subparallel on more than basal
half, broadly rounded in front; anterior margin
armed by 14 serrations; indefinite summit well
in front of middle; anterior slope rather abrupt,
rather coarsch' asperate; posterior areas subre-
tieulate (mostly wavy lines), punctures very
minute, sparse. Glabrous.
Elytra 1.9 times as long as wide, 1.6 times as
long as pronotum; sides almost straight and
parallel on more than basal three-fourths, very
broadK- rounded behind, striae not impressed;
surface" shining, almost smooth, punctures mi-
nute, almost obsolete, confused. D("cli\'ity con-
fined to less than posterior fourth, very steep,
broadh- coinex; punctures larger and deeper
than on disc, confused. Vestiture confined to
declivity, of rather abundant, fine, short, and
rather long hair.
Type Locality.— Tapanti, Cartago, Costa
Rica.
Type Material. —The male holot)pe and
two male parat\pes were taken at the type
locality on 17-I.\-63, 1.300 m. No. 184, from
Miconiu ((iiidata. by S. L. Wood.
The holotype and paratypes are in my col-
lection.
Gnafhotrupes crecenttis. n. sp.
This species is distinguished from dihitus
Wood by the steep, almost flat elytral declivity
which is armed by two pairs of moderately
large tubercles on interstriae 3, by the very short
declivital vestiture, by tlie much finer frontal
punctures, and by the smaller size.
Female.— Length 2.0 mm (paratypes 1.9-2.0
mm), 3.0 times as long as wide; color dark
brown.
Frons as in dihitus except punctures very
fine; sparse setae yer\ long. Pronotum as in
dihitus.
Elytra! outline as in (hhitus, strial punctures
very small, in definite rows, declivity steeper and
Biological Series, Vol. 19. No. 1 New Species of American Bark Beetles
57
much flatter than in (liliitus, punctures larger,
confused, interstriae 3 with two pointed denti-
cles one-third and two-thirds declivital length
from base. \'estiture mosth very short, a few
longer hairs.
Male.— Similar to female e.\cept lower half
of frons more distinctly, transversely impressed,
setae short; strial punctures minute, confused;
declivital denticles slightly larger, punctures
much smaller, vestiture shorter.
Type Locality.— Volcan, Puntarenas, Costa
Rica.
Type Materlal.— The female holotype, male
allotype, and eight paratvpes were taken at the
type locality on ll-XII-63, 1000 m, No. 30S. from
a recenth cut tree limb, b\ S. L. Wood.
The holotype, allotype, and paratypes are in
my collection.
Tricolus siDipIici.s. n. sp.
In this species the frons is rather coarsely
aciculate except for a small, median, raised, cen-
tral granular area, and declivital spines 2 and 3
are essentially absent.
Female.— Length 1.8 mm, 2.8 times as long as
wide; color light brown.
Frons convex, shining, coarsely, con\ergently
aciculate except on small, median, subcircular,
raised granular area at center; \estiture incon-
spicuous. Antennal club broadly obovate; su-
tures 1 and 2 distinetb procurved, obtuse!) sub-
angulate, finely septate except at middle, with-
out long setae on posterior face.
Pronotum 1. 3 times as long as wide; sides
straight and subparallil on basal half, narrowly
rounded in front; summit indefinite, anterior to
middle; anterior slope finely, closely asperate;
anterior margin a rather strongly, acutely ele-
vated continuous costa; posterior areas reticu-
late, punctures obscure, very shallow, moder-
ately close. Glabrous.
Elvtra 1.7 times as long as wide, 1.4 times
as long as pronotum; disc smooth, shining, punc-
tures small, s]iallo\\', some obsolete, evidently at
least slightl)- confused. Declivity occupying pos-
terior third, excavated as in other members of
this genus; face smooth, shining, indistinct punc-
tures small, confused; lateral margins rather
strongly, continuously elevated on heart-shaped
pattern from suture to moderately near posteri-
or margin, gradually increasing in height pos-
teriorly, ending precipitously on lower, median
angle; spine 1 on interstriae 1 at base very small,
pointed, 2 feebly indicated, 3 rounded, not pro-
jecting. Glabrous.
Type Locality.— Palin, Esquintla, Guatema-
la.
Type Materl\l.— The unique female holo-
type was taken at the tvpe locality on 19-V-64,
at 300 m. No. 685, in a Ficus t\vig, by S. L.
Wood.
The holotype is in my collection.
Tricolus inonwtus, n. sp.
This species is distinguished from simplicis
Wood by the much more slender body, by the
narrower declivital excavation, and by the ab-
sence of declivital spine 1.
Female.- Length 1.8 mm, 3.2 times as long
as wide; color verv dark brown.
Frons evidently as in simplicis, upper half
concealed. Antennal club subcircular, sutures al-
most straight; posterior face with two or three
long hairs.
Pronotum 1.4 times as long as wide; essen-
tiallv as in simplicis except punctures on pos-
terior areas deeper, distinct, a slight irregularity
at base anterior to scutellum. Glabrous.
Elytra 2.0 times as long as wide; disc as in
simplicis. Declivity similar to simplicis except ex-
cavated area much narrower, spine 1 represented
only b\' a small callus, 2 represented by a dis-
tinct callus, 3 more broadly rounded, not project-
ing. Glabrous.
Type LocALiT^-.-Fifteen km SE Cartago,
Cartago, Costa Rica.
Type Material.— Tlie uniqui- female holo-
type was taken at the type locality on 24-IX-6.3,
at 2800 m, No. 248, from a branch of Sipanina
nicara<s.uaensis, by S. L. Wood.
Tlic holotvpe is in m\ collection.
Tricolus inaffcctus, n. sp.
This species is distinguished from inornatus
Wood by the larger, stouter body, by the im-
pressed lower, lateral areas of tlu' frons, by the
pronotum, and by slightly wider elytral declivity.
Female.— Length 2.6 mm, 2.9 times as long
as widt'; color black except elytral disc and an-
tennae yellowish brown.
Frons as in simplicis Wood except granular
area more strongly elevated, lateral areas im-
pressed just abo\'e epistoma; mandibles appar-
ently elongate. Anti'iinal club broadly oval, su-
tures straight; posterior face ornamented by more
than 20 long hairs.
Pronotum 1.2 times as long as wide; outline
about as in simplicis except sides on slightly
more than anterior half distinctly ccmstricted,
58
BlllOHAM VOUNG UNl\EHSnY SciENCK Bui.I.ETIN
anterior costa more lu-arly serrate; posterior
areas reticulate, punctures moderately fine,
rather deep. Glabrous.
Elytra 1.8 times as long as wide, 1.6 times as
long as pronotuni; as in inoriwttis except excava-
ted decli\ital area distiiicth wider, calluses of
spines 1 and 2 less conspicuous. Glabrous.
Type Locality.— Volcan Poas, Heredia, Costa
Rica.
Type Material.— The unique female holo-
typc was taken at the type locality on 19-XI-6.3.
2600 m, by S. L. Wood. '
The holotype is in my collection.
Tricolus cccropu. n. sp.
This species is distinguished by the moder-
ately stout body, by the poorly developed de-
clivital spines, bv the strongly produced, acute,
strongly serrate anterior margin of the prono-
tuni, by the large, subrenifcjnn granular area on
the frons, and by the habits.
FE^rALE.— Length 2.4 nun (parat\pes 2.2-
2.5 mm), 2.6 times as long as wide; color dark
brown.
I'rons sonii'what flattened on median three-
fourths from epistoma to upper level of e\cs,
most of flattened area occupied by a raised,
granular area of subreniforni shape. Antenna!
club obscurely subtriangular, sutures weakly
procurved; very feu long setat' on posterior
face.
Pronotum 1..3 times as long as wide; widest
near base, sides weakly converging on basal t\\ o-
thirds, anterior margin xcry strongly, subacutely
produced and coarsely, closely serrate, 14-16
serrations; summit indefinite, near middle; an-
terior slope armed by rather large, low asperi-
ties; posterior area reticulate, punctures sparse,
fine, shallow. Glabrous.
Elytra 1.5 times as long as wide, 1.2 times
as long as pronotum; disc smooth, punctures
small, shallow, confused. Declivity moderately
steep, excavated area narrower and not as deep
as in other species; spines 1 and 2 small, pointed
tubercles, 1 slighth' larger, .3 represented by a
low, poorh developed fold; subapical margin
subacute, not produced. Subglabrous.
Male.— Similar to female.
Type Locality.— Turrialba, Cartago, Costa
Rica.
Type Matehial.— The female holotype, male
allotype, and eight parat)'pcs were taken at the
type locality on 5-VH-63,'at 700 m, No. 19, from
fallen Ccciopia peltata leaf petioles, by S. L.
Wood.
The holotype, allotype, and paratypcs are
in my collection.
Tricolus intrusus, n. sp.
This species is distinguished from the almost
identical cecropii Wood by the much more
acutely produced anterior margin of the prono-
tum, by the coarser punctures on the pronotal
disc, and by the finer, less numerous punctures
in the excavated area of the elytral declivity.
Female.- Length 2.4 nnn (paratypcs 2.1-2.2
mm), 2.7 times as long as wide; color dark
brown.
As in cecropii, except anterior margin of pro-
notum much more acutely produced, tridentate
at its apex (a total of nine serrations on entire
margin); punctures on discal area of pronotum
distincth' larger, e\idently more numerous; punc-
tures on elytral disc evidently slightly smaller;
punctures in excavated face of declivity much
smaller, much less numerous.
Type Locality.-E1 Laurel, 12 km SW Ca-
racas, Venezuela.
Type Matehial.— The female liolotype and
two female paratypcs were taken at the type
localitN' on l-\'-76, at 1300 m. No. 475, from
petioles of fallen Cccropid leaves, by S. L. Wood.
The lioli)t\pe and paratypcs are in my col-
lection.
Tricolus ardis. n. sp.
This spi'cies is distinguished by the acutely
angulati'. strongly serrate anterior margin of the
pronotum. bv the aciculate frons which also has
a small, elexated granulate area, and 1)\' the ely-
tral declivity.
Female.— Length 2.3 mm (paratypcs 2.1-2.5
mm), 3.3 times as long as wide; head, prothor-
;ix, and elytral declivit)' dark brown, basal two-
thirds of elytra light brown.
Frons convex, convergcntly aciculate except
for a small, trans\ersely oval granular area at
center occup\'ing median third. Antennal club
broadly obovate, sutures moderately arcuate;
posterior face without long hair.
Pronotum 1.5 times as long as wide; sides al-
most straight and parallel on basal half, arcu-
ately con\'erging to subacute median point in
front; anterior margin coarsely serrate; summit
indefinite; anterior slope finely, closely asperate;
posterior areas reticulate, punctures fine, shal-
low, not close, some of those at base connected
by transverse lines. Glabrous.
Biological Series, Vol. 19, No. 1 New Species of American Bark Beetles
59
Elytra (including spines) 2.1 times as long
as wide; sides straight and parallel to base of
declivity, arcuately converging to apex of spine
3, apices of spine 3 separated by a distance
equal to width of elytra, posterior emargination
broadly, somewhat U-shaped; surface almost
smooth, shining, punctures shallow, small, large-
ly confused, their centers apparently reticulate.
Declivity moderately steep, excavated in a nar-
rowly heart-shaped area; lateral areas abruptly,
rather strongly elevated, spine 1 at base on in-
terstriae 1 rather small, sharply pointed, 2 one-
third declivital length from base, twice as long
as wide, 3 projecting posteriorly, conspicuously
longer than wide (as seen in lateral aspect).
Glabrous except for an occasional hairlike seta
near declivity.
Male.— Similar to female except for apical
terga of abdomen.
Tyi'e Locality.— Escasu, San Jose, Costa
Rica.
Tvi'E Mateiual.— The female holotype, male
allotype, and 22 paratypes were taken at the
tyix-' locality on 2-X-63, at 1300 m, Nos. 215, 218,
from a tree seedling, by S. L. Wood. Two para-
types are from Cerro Punta near Volcan Chiri-
qui, Chiritiui, Panama, 11-1-64, 1300 m, Nos.
380, 401, tree branches, S. L. \\'ood.
The holotvpe, allotype, and paratypes are in
my collection.
Tricohis parsus, n. sp.
This species is distinguished from inaffec-
tus Wood by the more narrowly rounded ante-
rioi* margin of the pronotum, b\- the acutely ele-
vated, subapical margin of the declivity, and by
the very different declivital spine 3.
Male(?).— Length 2.3 mm, 2.6 times as long
as wide; color almost black.
Frons largely concealed, evidently not acicu-
late, finely granulate, raised granular area as in
ardis Wood. Antcnnal club broadly subquadrate,
sutures feeblv procurved.
Pronotum 1.3 times as long as wide; about
as in ardis except anterior margin narrowly
rounded, not angulate, serrations with their
bases contiguous. Glabrous.
Ehtra 1.6 times as long as wide, 1.3 times as
long as pronotum; sides straight and parallel on
basal two-thirds, gradualK- narrowed, rather
broadly rounded behind, with a conspicuous,
rather narrow emargination at suture; surface
almost smooth, shining, punctures largely con-
fused, small, moderatel) deep. Declivity much
as in inaffecttis except slightly wider, margins
slightly higher, spine more strongly developed
but not pointed, spine 3 more distinctly converg-
ing mesad, its crest not as high, its apex round-
ed, distinctly, weakly projecting; subapical mar-
gin acutely elevated, projecting moderately to
produce sides of median emargination. Glabrous
except for an occasional hairlike seta on sides.
Type Locality.— Fifteen km SE Cartago, Car-
tago, Costa Rica.
Type Materlal.— The unicjue holotype, pre-
sumablv a male, was taken at the type k)cality
on 24-l'X-63, at 1800 m, No. 196, from a woody
vine 5 mm in diameter, by S. L. Wood.
The holotype is in mv collection.
Tricohis rufithorax. n. sp.
This species is distinguished from parsus
Wood by the larger size, by the well-de\'eloped
decli\ital spines, by the subserrate anterior mar-
gin of the pronotum and by other characters.
Female.— Length 2.7 mm (male 3.1 mm),
2.4 times as long as wide; color reddish brown
except elytra black.
Frons convex, reticulate-granulate above,
somewhat punctate laterally, subcircular raised,
granular area occupying slightly more than me-
dian third to upper level of eyes; glabrous except
on epistoma. Antennal club moderately large, al-
most subtriangular, sutures weakly procurved;
posterior face bearing two long hairs.
Pronotum 1.2 times as long as wide; (jutline
as in parsus except anterior margin less narrow-
1\- rounded, its elevated costa weakly serrate; pos-
terior area strongly reticulate, very low, strongly
transverse, asperities continuing from summit to
base. Glabrous.
Elvtra 1.4 times as long as wide, 1.2 times as
long as pronotum; outline about as in parsus ex-
cept sutural emargination smaller and spine 3
projecting very slightly; disc smooth, shining,
punctures fine, confused. Declivity with con-
tours much as in parsus except spine 1 conical,
slightly larger, 2 cylindrical, projecting slightly
more than its basal width, 3 low, costate, ending
abruptb-, not projecting, directed toward sutural
apex but ending remote from it, subapical mar-
gin acutely elevated but less strongly than in
parsus. Glabrous except for a few setae on sides.
Male.— Similar to female except without long
hair on posterior face of antennal club; declivital
spine 2 abnormal, apparently chewed off on both
sides.
60
BnicHAM VouNC University Science Bulletin
Type Locality.— Finca Gromaco on Rio Goto
Brus, Puntarcnas, Costa Rica.
Type Material.— The female holotype and
male allotype were taken at the type locality on
14-\'II-63,' at 500 ni. No. 65, from' a woody '\ine
(liana) less than 1 cm in diameter, by S. L.
Wood.
The holotype and allotype are in my collec-
tion.
Tricohis hadiiis, n. sp.
This species is distinguished from luxlifer
Blandford by the smaller size, by the reddish
brown color, and l)y the more widely spaced
third spines which are very close to the subapi-
cal margin.
Female.— Length 2.S mm (paratypes 2.3-2.8
mm), 2.6 times as long as wide; color reddish
brown.
Frons as in nifitJiorux Wood except raised
granular area occup\ing one-fourth space be-
tween eyes. Antennal club subcircular, sutures
rather weakh- procur\ed; posterior face with up
to a dozen long hairlike setae.
Pronotum L2 times as long as wide; as in
nifithorax.
Elytra L5 times ;is long as wide; outline as in
nifithonix except spine 3 projecting more strong-
ly; disc smooth, shining, with \'ery fine, irregu-
lar lines, pimcturcs fine, shallow, confused. De-
clivity as in nifithorax except spine 2 more slen-
der, twice as long as wide; 3 close to costal mar-
gin, separated bv about lialf its basal width,
subapical margin reaching its posterior margin,
spine 3 projecting, subtriangular, almost as high
as its basal width; sulieostal margin acutely, not
strongly elevated. Sul)gIabrous.
Male.— Similar to female except antennal
club with fewer long setae.
Type Locality.- Pandora, Limon, Gosta Ri-
ca.
Type Material.— The female holotype, male
allotype, and 16 paratvpes were taken at the
type locality on 23-\'III-63, 50 m, Nos. 147, 150,
from a tree branch, by S. L. Wood. One para-
type is from Peralta, Gartago, Gosta Rica, lO-III-
64, 500 m. No. 462, woody vine, one paratypc
is from Gerro Punta near \'olcan Ghiriqui, Chir-
iqui, Panama, 11-1-64, 1800 m. No. 376, tree
brancli; 13 parat\pes are from Ft. Glayton,
Ganal Zone, Panama, 22-Xn-63, from various
collections; all taken by me.
The holotvp(\ allotype, and paratypes are in
my collection.
Tricohis partilis, n. sp.
This species is distinguished from nodifer
Blandford by the smaller size, by the brown
color, bv the more slender body form, by the
more slender spine 3 on the elytral declivity, and
by other characters.
Male.— Length l.S mm (paratypes 1.7-1.8
mm), 2.8 times as long as wide; color brown.
Frons as in nodifer except granular area dis-
tinctly elevated; antennal club subcircular.
Pronotum 1.3 times as long as wide; as in
nodifer except transverse crenulations in discal
area reduced to a few transverse lines.
I'^lytra 1.7 times as long as wide, 1.3 times
as long as pronotum; essentialh' as in nodifer
except spines 1, 2, and 3 more slender, 3 dis-
tinctly longer; lower declivity less broadly flat-
tened; subapical margin more strongly elevated.
Female.— Similar to male except posterior
face of antennal club with several long hairlike
setae.
Type Locality.— X'olcan. Funtarenas, Gosta
Rica.
Type Material.— Tlie male liolotvpe, female
;dlot\pe, and one parat\pe were taken at the
type' locality on ll-XII-63, at 1000 m, No. 304,
from a tree branch, by S. L. Wood. One paratype
is from Finca Gromaco on Rio Goto Brus, Pun-
tarcnas, 14-\'II-63, 500 m. No. 59, tree branch,
S. L. Wood.
The holot\pe. alIot\'pe, and paratypes are in
ni\ collection.
Tricohis fcnoris, n. sp.
This species is distinguished from partihs
Wood h\ the larger size, by the more slender
body form, and by the longer, more slender de-
clivital spines.
Male.— Length 2.1 mm, 3.0 times as long as
wide; color very dark brown.
Frons, antenna, and pronotmn as in partilis;
pronotum 1.3 times as long as wide.
EKtra 1.7 times as long as wide, 1.3 times as
long as pronotum; as in purtiUs except spine 2
three times as long as wide, spine 3 about one
and one-half times as long as wide; subapical
margin more strongly produced.
Type Locality.— Fifteen km SF, Gartago, Gar-
tago, Gosta Rica.
Type Material.— The unique male holotype
was taken at the type locality on 24-IX-63, at
1800 m. No. 196, from a woo'dv \'ine, bv S. L.
Wood.
Biological Series, Vol. 19, No. 1 New Species of American Bark Beetles
61
The holotype is in my collection.
Tricolus frontalis, n. sp.
This species is distinguished by the aciculate
frons, with the granular area absent, and In' the
slender declivital spine 3.
Female.— Length 2.4 nun, .3.1 times as long
as wide; color dark brown.
Frons convex, aciculate, granulate area ab-
sent.
Pronotum 1.3 times as long as wide; as in
fenoris Wood except anterior margin more
broadly rounded.
Elytra 1.9 times as long as wide, 1.5 times as
long as pronotum; about as in fenoris except de-
clivital spine 3 longer, almost three times as
long as wide, spine 3 similar in shape to 2 but
larger, about two and one-half times as long as
wide; subapical margin rather poorly elevated.
Type Locality.— Eight km NE Teziutlan,
Puebla, Mexico.
Type Material.- The unique male holotNpe
was taken at the t\pe locality on 27-\'I-53, at
1600 m, from a tree branch, by S. L. Wood.
The holotype is in my collection.
Tricolus capitalis, n. sp.
This species is distinguished from ruficollis
( Fabricius ) bv the slightly smaller size and bv
the much less strongly elevated subapical mar-
gin of the ehtral declivitv (thereby eliminating
the sutural notch).
Female.— Length 2.1 mm, 2.7 times as long
as wide; color rather dark reddish brown.
Frons con\ex, finely rugose-reticulate; gran-
ular area distinctly elevated, subcircular, occu-
p\ing median third to upper le\'el of eyes; gla-
brous. Antennal club ratlier broadh' obovate,
sutures moderately procurved.
Pronotum 1.3 times as long as wide; sides
straight and parallel on basal half, rather nar-
rowly rounded in front; anterior margin anncd
by about I'ight coarse teeth (bases of some con-
tiguous); summit indefinite, about one-third
pronotum length from anterior margin; anterior
slope rather coarsely, closely asperate; posterior
areas reticulate, punctures fine, those near base
with transverse line or minute shining elevation.
Glabrous.
Elytra 1.6 times as long as uide, 1.2 times as
long as pronotum; disc smooth, shining, punc-
tures rather fine, shallow, confused. Decli\it\'
excavated basically as in other members of the
genus; spine 3 stout, subtriangular, projecting
slightly, as in ruficollis, separated from sub-
apical margin by width of spine; lower declivity
flattened below spine 3 to lateral margin of spine
3; subapical margin almost straight, rather weak-
ly elevated, without sutural emargination, not at
all produced as in ruficollis.
Type LocALiri'.— Thirteen km S El Hato del
Volcan, Chiriqui, Panama.
Type Material.- The female holotype and
one female paratype were taken at the type lo-
cality on 7-1-64, at 1000 m, No. 371, from' a tree
seedling, by S. L. Wood.
The holotype and paratype are in my collec-
tion.
Tricolus naevus, n. sp.
This species is distinguished from ovicollis
Blandford by the slightly larger size, by the very
broad subrenifonn granular area on the frons,
by the narrowh- flattened area lielow declivital
spine 3, and by the much stouter declivital spine
3.
Female.— Length 2.9-3.3 mm, 2.7 times as
long as wide; color almost black.
Frons broadly convex, with lateral areas just
above epistoma conspicuoush' impressed; sculp-
ture of lower area irregular, smooth and shining
laterally, becoming subreticulate abo\e; granu-
lar area \ery broad, occupying median two-
thirds of area between eyes at least three times
longer than wide, its lower margin almost
straight, stronglv cmarginate on its upper mar-
gin. Antennal club subtriangular but narrower
than in ovicollis.
Pronotum 1.3 times as long as wide; as in
ovicollis except anterior margin more narrowly
rounded and more coarsely serrate.
Elytra 1.6 times as long as wide, 1.2 times as
long as pronotum; as in ovicollis except declivital
area below spine 3 flattened only to median mar-
gin of spine 3; spine 3 stout, subtriangular, its
length aliout e(|ua] to its basal widtli. Subgla-
lirous.
Locality.— Tapanti, Cartago, Costa
Type
Rica.
Type Material.— The female holotype and
two female parat\pes were taken at the type
locality on 24-X-64, 1300 m. No. 241, from a
Phoebe mcxicana branch, by S. L. Wood. One
female paratvpe bears identical data except it
is No. 244, from a woody vine, and one with
identical data is 26-XI-64, No. 271, Werklia
ii}sip,nata.
The holotype and paratypes are in my col-
lection.
62
Bhigham Vounc Univeiisitv Sc:iknce Bih.letin
Tricolus scitulus, n. sp.
This species is distinguished from ovicoUis
Blandford by the smaller size, by the uniform
brown color, and by the flatter lower declivity
below spine 3.
Female.— Length 1.9-2.3 mm, 2.6 times as
long as wide; color dark brown, young adults
uniformly brown.
Frons and pronotum as in ovicoUis except an-
terior margin of pronotiun more narrowly
rounded. Antennal club not quite as broad.
Elytra as in ovicoUis except area below spine
3 flatter.
Male.— Similar to female except antennal
club slightly smaller, with fewer long setae on
posterior face.
Type Locality.— Tapanti, Cartago, Costa
Rica.
Type Material.— The female holotspe, male
allotype, and nine paratypes were taken at the
type locality on 26-XL63, at 1300 m, Nos. 241,
265, from Phoclie mexicana l^ranches, by S. L.
Wood. One paratype is from Turrialba, Cartago,
Costa Rica, 5-VII-63, 700 m. No. 23, Crotohria.
S. L. Wood. One paratype is from Cerro Punta
near Volcan Chiricjui, Chiriqui, Panama, ll-I-
64, 1800 m. No. 406, tree branch, S, L. Wood.
A series from La Carbonera Experimental Forest,
50 km NW Merida, Merida, Venezuela, was not
included in the tvpe series.
The holotype, allotype, and paratypes are in
my collection.
Tricolus peUatus, n. sp.
This species is distinguished from the similar
but remoteh' related nodifcr Blandford by the
subapical tuljercle on deelivital spine 3, by the
less strongly elevated subapical margin on the
declivity, by the longer deelivital spine 3, and
by other characters.
Female.— Length 3.0 mm (paratypes 2.7-3.1
mm), 3.0 times as long as wide; color very dark
brown.
Frons, pronotum, and elytral disc as in nodi-
fer, except granular area on frons slightly larger.
Pronotum 1.3 times as long as wide.
Elytral declivity with basic contoius as in
nodifer, except spine 1 slightly larger, spine 2
more slender; spine 3 slightly longer than wide,
more erect than in nodifer, with a conspicuous
tubercle on dorsomedian subapical margin; sub-
apical margin less strongly produced and less
strongly elevated than in nodifer.
Male.— Similar to female except posterior
face of antennal club without long hair.
Type Locality.— Cerro Punta near Volcan
Chiriqui, Chiriqui, Panama.
Type Materlvl.- The female holotype, male
allotype, and two paratypes were taken at the
type locality on 11-1-64, 1800 m. No. .376 (holo-
type), 392 (allotype), 372 and 385 (paratypes),
from tree branches and a woody vine (No. 385),
S. L. Wood. Three paratypes are from \'olcan
Poas, Heredia, Costa Rica, 19-XL63, 2,500 m. No.
260, tree seedling, S. L. Wood.
The holotype, allotype, and paratypes are
in my collection.
Tricolus aciculatus, n. sp.
This species is distinguished from pcUatus
Wood by the smaller size, by the more slender
fonn, bv the absence of a raised granular area
on the frons, and by the presence of a \ery small
gramde in each puncture in the excavated area
of the declivity.
Female.— Length 2.7 mm (paratypes 2.2-2.7
mm), .3.1 times as long as wide; color very dark
brown.
Frons convex, rather weakly impressed above
epistoma in lateral areas; surfaet' sliining, con-
spicuously aciculate on lateral thirds, obscurely
on median third, granular area found in other
species entirely absent; median area above epis-
toma with a broad, low, elevated area somewliat
similar to allied species. Antennal club large,
broadh', asymmetrically oval, sutures almost
straight, obscure, indistinctly septate on central
third; a very few long setae on posterior face.
Pronotum 1.3 times as long as wide; pronot-
um as in nodifcr Blandford except anterior mar-
gin finely subserrate.
Elytra 1.7 times as long as wide, 1.4 times as
long as pronotum; elytra as in pelfatus except
more slender, discal punctures slightly larger,
piincturi'S on deelivital face each with a fine
granule; deelivital spines 2 and 3 more slender
than in peltatus, 3 distinctly longer, bearing a
subapical tuli-.TcIe as in peltatus; subapical mar-
gin of declivity as in peltatus except from dorsal
aspect median fifth abrupth', shallowly emargin-
ate. Subglabrous.
Male.— Similar to female except' posterior
face of antennal club without long hair.
Type Locality.- Ten km or 6 miles SE Te-
zuitlan, Puehla, Mexico.
Type Mati-.uial.- The female holotype, male
allotype, and 46 paratypes were taken at the
Biological Series, Vol. 19, No. 1 New Species of American Bark Beetles
63
t\pc locality on 2-\'II-67, at 1600 in, Nos. 137
and 145, from small Miconki branches, No. 142
from Rtibus, No. 149 host unknown, bv S. L.
Wood.
The holotype, allotype, and paratypcs are in
my collection.
Tricolus l)icflIor
u. ,sp.
This species is distinguished by the color pat-
tern, by the more dorsal po.sition of the frontal
granular area, and by tlu' uniijue decli\it\ .
Female.— Length 3.4 mm (paratype 3.1 mm),
2.7 times as long as wide; color of dorsal sur-
face yellowish brown except pronotal disc from
summit to base and clytral declivity black, ven-
tral surface black except legs light brown.
Frons largeh' concealed, e\iclentl\' eoinex
above, shallowly impressed on lateral thirds of
lower half, median area distinctly elevated abo\e
epistoma; granular area more dorsal in position
than normal, largely above upper le\el of e\es.
apparently wider than long. Antennal club
slightly asymmetrical, sutures weakly procurved.
Pronotum 1.1 times as long as wide; sides
feebly arcuate and subparallel on basal half,
rather strongly converging anteriorlv to rather
narrowly rounded, serrate anterior margin;
sculpture about as other Tricolus.
Elytra 1.7 times as long as wide, 1.5 times
as long as pronotum; sides straight and par-
allel to level of sutural apex, truncate behind
except for projecting spine- 3 and small sutural
emarginatioii; disc and upper decli\it\' about
as in nodifer Blandford except declivital exca\a-
tion more broadly impressed; spine 3 on ex-
treme posterolateral margin, somewhat laterally
compressed, appearing (juadrate from lateral as-
pect, distinctl}' longer than wide, with a con-
spicuous tubercle on dorsomedian apical angle,
its niesal surface somewhat eoncax'e or grooved
toward its apex; subapical margin acutelv,
\\eakly elevated, a shallow sutural emargina-
tion.
Type Locality.— Lower slopes of \'olcan
Poas, 24 km N^V Heredia, Costa Rica.
Type Materml.— The female holot\pc and
one female paratvpe were taken at tlie t^'pe
locality on 14-\TI-63, 1500 m. No. 44. from a
small branch, by S. L. Wood.
The holotype and paratype are in my col-
lection.
Tricolus amplus, n. sp.
This species is distinguished b\- the large
size, by the black color, by the displaced de-
clivital spine 2, by the subtriangular granular
area on the frons, and by other characters.
Female.— Length 3.7 mm (paratypes 3.6-3.9
mm), 2.5 times as long as wide; color black.
Frons convex above, lateral areas above epis-
toma moderately impressed, lateral areas below
upper level of eyes distinctly aciculate; median
granular area largely above eyes, wider than
long, subtriangular, its apex directed orad; ves-
titure of short, sparse hair. Antennal club
slightly asymmetrical, obscurely subtriangular,
sutures weaklv procurved.
Pronotum 1.13 times as long as wide; about
as in hicolor except basal area of disc more
coarsely, more extensively asperate.
Elytra 1.4 times as long as wide, 1.3 times
as long as pronotum; outline as in hicolor ex-
cept posterior margin Insinuate; disc subshining,
almost smooth, punctures \'ery fine, shallow,
confused. Declivit)' much as in allied species
except lateral margin less abruptly, less strongly
elevated; spine 1 in usual position at base on
interstriae 1; spine 2 larger and stouter than
usual and in position of interstriae 2, its base
beginning at level of apex of spine 1 (much
more anterior in position than all other specii'S ) ;
spine 3 strongh', lateralh' compressed, on lateral
margin, quadrate, distinctlv wider than long,
a conspicuous tubercle on its dorsomesal apical
angle; subapical margin acute, distinctly, not
strongly produced, extending to base of spine 3.
Subglabrous.
Male.— Similar to female, including long
hair on posterior face of antennal club.
Type Locality.— Ten km or 6 miles NE
Teziutlan, Puebla, Mexico.
Type Material.— The female holotype, male
allotype, and six parat\pes were taken at the
type locality on 2-\TI-67, at 1600 m. No. 140,
from a limb 5-15 cm in diameter, bv S. L.
Wood.
The holotvpe, allot\pe, and paratypes are in
mv collection.
Amphicranus mirandus, n. sp.
This species is distinguished from hijbridus
Blandford by the pronounced sexual dimor-
phism, by the cleft female epistoma, by the
more slender antennal club, by the absence of
an elevated line on the lateral margins of the
pronotum, and b\' the verv different dccli\ity.
Female.— Length 2.5 mm (paratypes; fe-
males 2.5-2.8 mm, males 2.8-3.5 mm), 3.3 (male
3.9) times as long as wide; color yellowish
64
BiiicHAM Vf)UNG University Science Bulletin
brown except anterior third of pronotiirn and
elytral declivity darker Ijruwn.
Frons convex above, weaklv, traii,s\-er.sel\-
impressed just below upper level of eyes, epis-
tomal area protuberant and sharph elcft on
median line from epistomal margin to trans-
verse impression; surface subrugose-reticulate,
a few fine punctures obscurely indicated; gla-
brous except along epistomal margin and on
sides of head near eyes. Antennal funicle 2-seg-
mented; club 1.6 times as long as wide, oval,
sutures weakly procurved, posterior face with
some long hair.
Pronotum 1.4 times as long as wide; widist
at base, sides straight distinctly con\eiging on
ijasal three-fourths, \'ery broadly rounded in
front; anterior margin unarmed; summit on an-
terior fifth; anterior slope very steep, asperities
rather sparse; posterior areas finely reticulate,
punctures minute, obscure. Glabrous.
Elytra 1.9 times as long as wide, 1.4 times
as long as pronotum; sides almost straight and
parallel on basal three-fourths, broadlv, ratlur
dei-plv U-shaped behind between lateral pro-
cesses; decli\it\- confined to posterior two-fifths;
disc as in hijhridtts. I^celixity ])asieallv as in
litjl>richis; interstriae 1 at base with a niiniitc
tubercle, 2 with a slender si)ine about three
times as long as its basal width; lateral margin
following same course as in Jii/hridus but only
weakly elevated and not serrate; lateral pro-
cesses about as long as their bas;il widths,
separated by slightly gre;iter distance than
deptli of emargination between them; mesal
margin of terminal processes bearing a strongly
elevated, laterally compressed, longitudinal pro-
cess, its upper margin horizontal, its posterior
margin subx'ertieal. Disc glabrous, rather sparse,
fine hair on declivity.
Male.— Similar to female except frons
broadly convex, epistomal area normal; ante-
rior margin of pronotum vertical, appearing
shallowly, broadly emarginate from above; de-
elivital spines and lateral processes distinctly
longer; lateral decli\ital processes with margins
directed mesad (not dorsad) thereby forming
a pronounced constriction in area of emargina-
tion just anterior to its middle; face of decli\'it\'
glabrous.
Tyi'e Locality.— Rincon de Osa, Puntarenas.
Costa Rica.
Type Materlal.— The female holotvpe, male
allotype, and 30 parat\'pes were taken at the
type locality on ll-\ 111-66, at 30 in. No, 85.
from a tree limb 5-15 cm in diameter, bv S. L.
Wood.
The holotype, allot\ pe, and paratypes are in
my collection.
Auiphicrtiuus lurnatilis, n. sp.
This species is distinguished from specta-
hilis (Wood) by the larger size, by the color,
by the coarsely serrate, more narrowly rounded
anterior margin of the pronotum, and b\- the
more gradual, more strongly explanate elytral
declivity.
FrNrALE.-Length 2.7 mm (paratypes 2.5-2.8
mm), 3.5 times as long as wide; color yellowish
brown with head, anterior half of pronotum,
and posterior half of elvtra black.
Frons as in spcctahilis.
Pronotum 1.6 times as long as wide; as in
spcctahilis except anterior margin more nar-
rowly rounded and armed by nine coarse serra-
tions, median one prominent, and posterior
areas less strongly reticulate, more shining.
Elytra 1.9 times as long as wide, 1.3 times
as long as pronotum; ;is in spcctahilis except
disc very smooth, brightly sliining, declivity not
as steep, posterior margin more strongK' explan-
ate, and upper slope of lower major proimnence
on margin of declivity horizontal (as in specio-
siis Schedl), not ascending. Glabrous.
Type Locality.— Six km S San \'ito, Punta-
renas, Costa Rica.
Type Material.- The female holotype and
seven paratypes, apparently all females, were
taken at tlie type locality (in 19-21-111-67, from
a dead tree, bv D. D. Sliwa.
The holotype and paratvpes are in my
collection.
Amphicranus macclltis. n. sp.
This species is distinguished from halteattts
Blandford bv the much smaller size, bv the
color, by the absence of a granular area on the
frons, bv details of the declixital sculpture, and
by other characters.
Male.— Length 1.9 mm (parat\pes 1.8-1.9
mm), 4.0 times as long as wide; color yellowish
brown.
Frons uniformlv convex; surface evidently
minuteh' rugose-reticulate. Antennal funicle 3-
segmented; club 1.3 times as long as wide,
Iiroadlv oval, sutures rather weakly procurved.
Pronotum 1.8 times as long as wide; .sides
straight and almost parallel on basal three-
fourths, rather narrowly rounded in front; an-
terior margin acuteh' ele\ated, siibserrate ex-
cept median serration slightly larger and basallv
separate from marginal eosta; indistinct summit
Biological Series, Vol. 19. No. 1 New Species of American Bark Beetles
65
on anterior fourth; anterior slope steep, conve.x.
most conspicuous asperities arranged in a sub-
transverse row; surface sul)reticulate, punctiues
small, very shallow. Glabrous.
Elytra 2.4 times as long as wide, 1.4 times
as long as pronotum; outline as in bciltcdtus
except apical emargination wider, l^-shaped;
disc smooth, strial punctures fine, shallow, in
rows. Declivity broadly, deeply excavated much
as in balteatus except steeper and somewhat
less strongly explanate; basal margin with
spines on interstriae 1 and 2 minute, spine on
3 moderately large, pointed, lateral margin
acutely, strongly elevated to small denticle on
mesal side at level of sutural apex, margin
oblicjueh' descending from denticle to apex of
lateral process on a straight line; sutural emar-
gination narrowly U-shaped, 1.5 times as deep
as wide; declivital face reticulate, shining,
punctures very obscureb indicated. Glabrous.
Female.— Similar to male except decli\it)
less strongb' explanate, its lateral margins less
stronglv elevated; sutural emargination as long
as wide.
Type Locality.— Rincon de Osa, Puntarenas,
Costa Rica.
Tyi'e Material.— The male holotvpe, female
allotype, and two male parat\'pes were taken at
the type locality on ll-\'III-66, at 30 m. No.
85, from a tree limb, bv S. L. W'ood.
The holot\pe, allotype, and paratypes are in
my collection.
Amphicranus spinescens. n. sp.
This species is distinguished b\ the absence
of a fine, raised line on the lateral margins of
the pronotum, by the very coarse, slender ser-
rations on the anterior margin of the pronotum.
and by the different annature on tlie elytral
declivity.
Female.— Length 2.4 mm (paratypes 2.0-2.4
mm), 4.4 times as long as wide; color yellowish
brown, anterior half of pronotum and posterior
third of elytra a darker brown.
Frons trans\'erselv concave, lower third
weakly, transversely impressed, surface rugose-
reticulate except subgranular on oval area on
median half at upper level of eyes; vestiture
confined to epistomal margin. Antennal funicle
3-segmented; club o\al 1.4 times as long as
wide.
Pronotum 1.8 times as long as wide; sides
straight and paralk'l on basal two-thirds, nar-
rowly, strongly produced in front; anterior mar-
gin armed b\' 10-12 \'eiy coarse, basally sepa-
rate serrations, median ones slender; summit
on anterior third; anterior slope moderately
steep, convex, anterior asperities coarse, smaller
toward summit; posterior areas finely reticulate,
punctures minute, almost obsolete. Glabrous.
Elytra 2.4 times as long as wide, 1.4 times
as long as pronotum; sides straight and parallel
to declivity, broadly emarginate behind, pos-
terolateral angles produced into long forceps-
like processes; disc smooth, subshining, strial
punctures almost obsolete. Declivity steep, ex-
cavated; basal margin subacutely elevated,
armed by three small, pointed denticles one
each at fnterstriae 1, 2, and 3; lateral margin
subacutelv elevated, bearing just below middle
of declivity a yery large, cylindrical, blunt spine
3.5 times as long as its basal width, directed
caudad, below this spine lateral margin con-
tinues to apex of ventrolateral process; \entro-
lateral process subcylindrical, curving slightly
mesad, its length measured from apex of
suture equal to more than half width of el\tra;
posterior emargination \ery broad, deep,
broadly U-shaped, very slightly deeper than
wide.
Male.— Similar to female except declivital
spines slightly longer.
Type Locality.— Rincon de Osa, Puntarenas,
Costa Rica.
Type Material.— The female holotype, male
allot\pe, and 18 paratypes were taken at the
tj^pe locality on ll-MII-66, at 30 m. No. 85, tree
limb, by S. L. Wood.
The holotype, allot\pe, and parat\pcs are
in my collection.
Ainphicramis spinosus, n. sp.
This species is distinguished from spinescens
Wood by the larger size, by the finely punc-
tured pronotal disc, and by the more brightly
shining elytra.
Female.— Length 2.7 mm (male paratypes
2.8-3.0 mm), 4.0 times as long as wide; color
brown.
As in spitiescens except posterior areas of
pronotal disc reticulate, with moderately abun-
dant, fine, distinctly impressed punctures in
both discal and lateral areas and elytra much
more brightly shining.
Male.— Similar to female except declivital
spines shghtlv longer.
Type Locality.— Rincon de Osa, Puntarenas,
Costa Rica.
Type Material.— The female holotype, male
allotype, and two male paratypes were taken
66
BiiiGiiAM ^OuNG Univehsiiv Sciknce Bulletin
at the type locality on ll-VIII-66, at 30 m. No.
85, from a tree limb, by S. L. Wood.
The holotype, allotype, and paratypes are
in mv collection.
Amphicianus mucronaltis, n. sp.
This .species is distinguished from /«,sf/L;/«?(/.s
Blandford by the smaller size, liy the \'ery dif-
ferent female frons, and by the narrower pos-
terolateral elytral processes of the male on
which the spine at the level of the sutural ape.x
is positioned ('((uidistant from the lateral and
mesal margins (not on tlie lateral margin as in
fastigiatiis).
Female.— Length 2.7 mm, 4.0 times as long
as wide; color light brown, pronotuni and
elytral declivity darker.
Frons con\e,\, reticulate ])eIow upper le\el
of eves, shining abo\e, punctures fine, moder-
ately abundant, obscure (.)n reticulate area: me-
dian fourth from epistoma more tlian half dis-
tance to upper level of eves with a shaiph'
elevated, low, oval, granulate area; \estiture
inconspicuous. Antennal club transverselv o\al.
slightly wider than long.
Pronotum 1.7 times as long as wide; as in
fastiiiiatiis except submarginal groo\i's and
callus on anterior slope mueli more poorh'
developed.
Elytra as in fastiiiiatus except discal punc-
tures slightly smaller.
Male.— Length .3.0-.3..'3 mm; similar to male
fastigiatiis except antennal club slightly more
broadly o\al; anterior margin of pronotum more
strongly, narrowly acuminate, process on its
ventral surface almost obsolete; elytral declivity
slightly less strongly explanate, lateral margins
much less strongly elevated, denticles on basal
margin much smaller, denticle at le\el of sutural
apex in center of floor of lateral process ( not
on inner surface of lateral margin as in
fastii^ialus).
Type Locality.— Cerro Ptmta near \olcan
Chiriqin', Panama.
Type Material— The female holotype, male
allotype, and one male paratvpe were taken at
the type locality on 11-L64, at 1800 m, from a
tree branch, by S. L. Wood.
The holotype, allotype, and parat\pe are
in my collection.
Amphicianus aciis. n. sp.
This species is distinguished from fastifiiafns
Blandford by the smaller size, bv the different
female frons, by the more slender male pronotal
process, and b\' the male elytral declivity.
Female.— Length 2.5 mm (female paratype
2.5 mm), 4.0 times as long as widt'; color light
reddish brown, anterior parts of pronotum and
elytral declivity sometimes darker.
Frons about as in female fastiiiiattis except
reticulate to vertex, median area on lower half
rather strongly elevated but not shaiply defined,
its summit nanowly flattened and granular. An-
tennal club obscureh' snbtriangular, 1.3 times
as wide as long.
Pionotum 1.8 times as long as wide; as in
fasti'fi,iotits except grooves and submarginal
ridges on anterior slope more pronounced.
Elytra as in mucronatus Wood.
Male— Length 2.5-2.8 mm; as in male
mucronatus except granular area on frons nar-
rower, exti-nding to upper le\-el of eyes; lower
surface of pronotal mucronate process with a
rather strongly developed, conical tubercle di-
rected ventrad; lateral margin of declivity
slightly higher, a coarse denticle displaced just
mesad of lateral margin at level of sutural apex,
posterodorsal angle rounded and cntireh' devoid
of a tubercle, sutural emargination much nar-
rower.
Type Locality.— El Laurel Experiment Sta-
tion, 12 km SW Caracas, Venezuela.
Type NLvtehial.— The female holotype, male
allotype, and three paratypes were taken at the
type' locality on l-\'-70, 1800 m, Nos. 457, 458,
from a tree bole 13 em in diameter. No. 513
in flight, bv S. L. Wood. One broken male
paratype is from Merida, Merida, Venezuela,
ll-IX'-69, 2000 m. No. 4, Croton bole, S. L.
Wood.
The liolot\pe, allot\pe, and paratvp(>s are
in my collection.
Paracorthi/Ius mtitilus. n. sp.
Tiiis species is distinguished from concisus
Wood b\- the smaller size, by the shallowly
divaricate sutural ;ipex of the elytra, and b\' the
distinctive eh tral declivity.
Female.— Length 2.0 mm (parat\pes 1.9-2.0
mm), 2.7 times as long as wide; color reddish
brown.
Frons as in velutimis Wood except surface
reticulate abo\e shining epistomal margin." An-
tennal club 1.4 times as long as wide, apical
segment only slightly modified.
Pronotum 1.1 times as long as wide; as in
velutimis except sinface reticulate, punctures
very fine, much less closeh- spaced.
Biological Series, Vol. 19, No. 1 New Species of American Bahk Beetles
67
Elytra 1.4 times as long as wide, 1.2 times
as long as pronotum; as in veltitituis except de-
clivity. Declivity subvertical, suljtriincate, shal-
lovvlv concave; lateral margiiLS distinctl\ c]e\at-
ed, rounded, armed on middle third on median
margin by two pointed tubercles, upper one
slightly larger; sutural interstriae weakly elevat-
ed; posterolateral margin subacute at apex,
suture weakly emarginate; surface rugose-retic-
ulate; punctures obscure. \'er\' minute setae
moderately abundant on declivity, similar but
less abundant on disc.
Male.— Similar to female except apical seg-
ment of antennal club slightlv shorter and
more broadly rounded; anterior margin amied
by eight serrations of moderate size; decli\it\-
more evenly concave, its margins weakh', sub-
acutely elex'ated on more than lower three-
fourths, upper margin armed b\' three pairs
of small teeth on interstriae 1-3, lateral pair
conspicuouslv larger, sutural apex more dis-
tinctly emarginate, sutural interstriae more dis-
tinctly elevated.
Type Locality.— Fort Shennan, Canal Zone,
Panama.
Type Material.- The female holotype, male
allotype, and two female paratvpes were taken
at the type locality on lO-XI-57, at light.
The holotype, allotype, and paratvpes are
in my collection.
Paracorthi/lus concisus, n. sp.
This species is distinguished from vehitinus
Wood by the larger size, by the subconcave
elytral declivity, with different armature in the
female, by the sparse pubescence, and by other
characters.
Female.— Length 2.6 mm (paratvpes 2..3-2.6
mm), 2.6 times as long as wide; color reddish
brown.
Frons as in velutimts except obscurely reticu-
late, shining, median carina extending from
epistomal margin to median callus on lower
third. Antennal club 1.9 times as long as wide,
aliout intermediate in shape between velutinus
and mutilus.
Pronotum 1.1 times as long as wide; as in
mtitilus.
Elytra 1.4 times as long as wide, 1.2 times
as long as pronotum; as in mutilus- except
tubercles slightly larger and apex of suture
entire.
Male.— Similar to female except antennal
club less elongate, its apex more broadly
rounded; anterior margin of pronotum armed
by eight serrations; declivity much more
broadly, evenlv impressed, margins on lower
half more distinctly, more narrowly elevated
but not angulate, sutural apex entire and bent
slightly dorsad, two pair of tubercles on inter-
striae 3 displaced mesad from lateral margin,
lower one at middle, upper one on basal fourth.
Type Locality.— Moravia, Cartago, Costa
Rica.
Type Materl\l.— The female holotype, male
allotype, and 14 paratypes were taken at the
type' locality on 11-III-64, 500 m. No. 473, from
a cut tree 35 cm in diameter, by S. L. Wood.
The holotype, allotype, and paratypes are
in my collection.
S'/U/?- FJrpya
Brigham Young University
*'''^ubrTry°°'" Science Bulletin
MAY271P74
HARVARD
UNIVERSI
TAXONOMY OF THE GENUS
CHLORIS (GRAMINEAE)
by
Dennis E. Anderson
BIOLOGICAL SERIES — VOLUME XIX, NUMBER 2
MARCH 1974 /ISSN 0068-1024
BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN
BIOLOGICAL SERIES
Editor: Stanley L. Welsh, Department of Botany,
Brigham Young University, Provo, Utah
Acting Editor: Vernon J. Tipton, Zoology
Members of the Editorial Board:
Ferron L. Andersen, Zoology
Joseph R. Murdock, Botany
WiLMER W. Tanner, Zoology
Ex officio Members:
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Sciences
Ernest L. Olson, Director, Brigham Young University Press
The Brigham Young University Science Bulletin, Biological Series, publishes
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Brigham Young University
Science Bulletin
TAXONOMY OF THE GENUS
CHLORIS (GRAMINEAE)
by
Dennis E. Anderson
BIOLOGICAL SERIES — VOLUME XIX, NUMBER 2
MARCH 1974 / ISSN 0068-1024
TABLE OF CONTENTS
ABSTRACT 1
ACKNOWLEDGMENTS 1
INTRODUCTION 1
NOMENCLATURAL HISTORY OF THE GENUS 2
TAXONOMIC RELATIONSHIPS OF CHLORIS 2
Cytology 3
Leaf Epidermis 5
Anatomy, Embryology, and Seedling Morphology 6
Relationships of Chloris and Other Genera 9
Relationships within Chloris 13
Chloris Swartz 15
Key to the species of Chloris 16
1. Chloris roxlmrghiana Schultes 22
2. Chloris puniculata Scribner in Robinson 24
3. Chloris herroi Arechavaleta 25
4. Chloris ciliata Swartz 27
5. Chloris lamproparia Stapf in Chevalier 30
6a. Chloris canterai Arechavaleta var. canterai 30
6b. Chloris canterai Arechavaleta var. grandiflora ( Rosengurtt & Izaguirre de Artucio) Anderson 32
7. Chloris dantUjiina C. D. Adams 34
8. Chloris cruciata (Linnaeus) Swartz 37
9. Chloris filiformis (Vahl) Poiret in Lamarck 39
10. Chloris suringari Hitchcock in Urban 39
11. Chloris scariosa F. von Mueller 40
12. Chloris prieurii Kunth 42
13. Chloris rohusta Stapf in Chevalier 43
14. Chloris mollis (Nees) Swallen 45
15. Chloris longiaristata Napper 46
16. Chloris quinque.sctica Bhide 48
17. Chloris wightiana Nees ex Steudel 48
18. Chloris montana Roxburgh 51
19. Chloris bournci Rangachariar & Tadulingam 53
20. Chloris inflata Link 53
21. Chloris formosana (Honda) Keng 57
22. Chloris pilosa Schumacher 58
23. Chloris virgata Swartz 60
24. Chloris gai/aiu/ Kiuith 65
25. Chloris castilloniatui Lillo & Parodi 69
26. Chloris orthonoton Doell in Martins 69
27. Chloris aristata (Cervantes) Swallen 70
28. Chloris hrandegei (Vasey) Swallen 72
29. Chloris chloridca ( Presl) Hitchcock 74
30. Chloris ventricosa R. Brown 76
31. Chloris truncate R. Brown 78
32. Chloris pumilio R. Brown 80
33. Chloris lobata Lazarides 82
34. Chloris divaricata R. Brown 83
35. Chloris pcctinata Bentham 86
36. Chloris crinita Lagasca 87
37. Chloris pluriflora (Foumier) Cla)'ton 89
38. Chloris cticullata Bischoff 91
39. Chloris verticillata Nuttall 93
40. Chloris andropogonoidcs Foumier 95
41. Chloris texensis Nash 103
42. Chloris sesqiiiflora Burkart 104
43. Chloris humhcrtiana A. Camus 104
44. Chloris amethi/stca Hoclistt-tter 106
45. Chloris subniutica Humboldt, Bonplantl. Kunth 106
46. Chloris somalewis Rendle 109
47. Chloris cleimntis Merrill 109
48. Chloris radiata (Linnaeus) Swartz 110
49. Chloris pycnotlirix Trinius 114
50. Chloris halophila Parodi 1 17
.51. Chloris ekmanii Hitchcock 119
52. Chloris arcmiria Hitchcock & Ekman in Hitchcock 119
53. Chloris sagnwuna Achille Richard in .Sagra 121
54. Chloris cubensis Hitchcock & Ekman in Hitchcock 123
55. Chloris mossambicensis Schumann 123
56. Chloris hurrnetuis Ander.son 125
UNIDENTIFIED, REJECTED, OR REASSIGNED NAMES 126
LITERATURE CITED 131
TAXONOMY OF THE GENUS CHLORIS (GRAMINEAE)
by
Dennis E. Anderson'
ABSTRACT
This study is a worldwide taxononiic treat-
ment utilizing endomorpliic characters such as
cytology, histology, and ciiihrNology, in addition
to the traditional cxoniorphic features. New
chromosome numbers and histological informa-
tion are reported. Relationships between Chloris
and other genera in the Chlorideae are dis-
cussed. A total of r6 species is recognized in the
genus; each is described and illustrated. Com-
plete synonymies are included as well as a list
of unidentified or rejected names. Distributions
and maps are given for each species, along with
lists of specimens examined dining studies at
various iu-rbaria. Reports of field studies in-
volving introgression among certain species are
also included. A new species, Chloris hurmen-
sis, is described from Bumia, and a new com-
bination, Chloris canterai Arech. var. graiuliflora
( Rosengurtt & Izaguirre de Artucio ) Anderson,
is also proposed.
ACKNOWLEDGMENTS
These studies were begun in the summers
of 1963 and 1964 as part of a National Science
Foundation program in l\esearch Participation
for College Teachers sponsored b\' the Depart-
ment of Botan\-, University of Texas. Additional
financial support was provided b\' this program
through grants ( NSF GY-3002, NSF GE-69:8).
The assistance of Dr. Harold Bold, director of
the summer program, is also gratefully acknowl-
edged. Initialh', Dr. Walter V. Bro\\'n pointed
out to me many of the biosystematie problems
of Chloris in Texas, and I am grateful to him
for his continued encouragement.
.additional direct assistance was received
from the National Science Foundation in 1969
(NSF GB-7235), enabling me to visit a number
of major overseas herbaria and to do additional
field work. The curators were most helpful and
allowed me to borrow man\- specimens. Their
cooperation is much appreciated.
A special debt is owed to Dr. W. D. Clayton
of the Royal Botanic Gardens, Kew, England,
who provided advice, cooperation and friend-
ship, and thus made my stay at Kew most pro-
ductive.
California State University, Humboldt, has
generously provided released time and leaves in
support of these studies, and the continued as-
sistance of President Cornelius Siemens is ac-
knowledged with many thanks.
I would also like to thank Mr. Rupert C.
Barneb\- for his aid in the preparation of the
Latin description of Chloris btirmsiisis.
Michael Lazarides kindly let me examine his
manuscript treatment of Chloris in Queensland,
prior to its publication.
I am grateful to my wife, Nancy, for the end-
less hours she contributed in filing and organiza-
tion, in establishing and verifying geographical
locations, and, finally, in cartography.
INTRODUCTION
With its delicate, digitate clusters of spikes,
Chloris must certainly be among the most strik-
ing of grass genera. The distribution spans
the warmer regions of most of the continents,
from the pampas of Argentina to the sandhills
of Nebraska, from the Cape of Good Hope to
the sands of the Sahara, from Afghanistan to
the island archipelagoes of the South Pacific,
and from the subcontinental islands of Ne.v
Guinea and Borneo through the continent of
^Department of Biology, Humboldt State College, Areata, California 95521.
Bricham Young University' Science Bulletin
Australia. Thus, it is likely that taxonomists in
these areas have seen (me or more of the taxa
in the genus. Though these attributes would
seem to insiue popularity among agrostologists,
Chloris nevertheless has not been subjected to
any systematic review for nearly a centuiy. This
has been an era characterized by extensive ex-
ploration and collection of hitherto unknown
areas, as well as the development of revolu-
tionary systematic thought.
My interest in Chloris began with studies in
hybridizing species in Texas; this interest spread
later to similar studies in the Southeast; and ulti-
mately it encompassed more classical investiga-
tions in manv of the major herbaria of the
United States, Great Britain, and Europe. Ini-
tially, my interests centered in experimental
studies but the continuing frustration of not
being able to name species with confidence led
me to conclude that a classical studv was needed
and that experimental studies could then be
made on a sounder foundation.
Well over 15,000 herbarium specimens and
many mass collections from hybridizing popula-
tions were studied. The numbers of specimens
available van' widely. Some taxa are repre-
sented only by the holot\'pe. Others are known
from such large numbers of specimens that
population descriptions may be advanced with
confidence. Still others ha\'e been collected so
thoroughly that the enormous complexit)' of the
population has revealed the necessity for further
intensive investigation.
This study, then, attempts to bring the tax-
onomv of the genus CliJoris up to date. It is
largely morpho-geographical in nature, with oc-
casional incursions into biosystematics as the
nature and availability of materials allow.
NOMENCLATURAL HISTORY OF THE GENUS
Swartz first described Chloris in 1788, in-
cluding two new species, C. ciliata and C. pc-
traea, and transferred into the genus three Lin-
naean species: Agrostis criiciafa, Agrostis radiata,
and Andropogon poh/dactylon.
With the description of the genus Eitstachijs
by Desvaux (1810) and the transfer to it of
Chloris petraea Swartz, the basic nomenclatural
pattern at the generic level was established,
though many authors now consider Eustachijs
as a subgenus of Chloris.
Other genera, based on various species of
Chloris, have been proposed: Rabdochloa Beau-
vois (1812), including C. cruciata and several
species from other genera; Schtdtesia Sprengel
(1815) containing C. petraea only; and Phacel-
laria Willdenow ex Steudel ( 1840), including C.
suhmutica. None of these genera have been ac-
cepted by recent authors.
More recently, Camus proposed the subgen-
era Monanthochloris, including within it C. per-
rieri (1949), and Pterochloris, which contains C.
hitmhertiana (1950). Subsequently, Camus
(1957) elevated the latter subgenus to the ge-
neric level.
Generic conspeeti of Chloris have been writ-
ten spor;idicallv, Iiut all are of the nature of
catalogs rather than re\isions or monographs.
Of these, the treatment by Steudel ( 1854) is the
most complete. In it he recognized 69 taxa and
included a s\noptic key as well as an enumera-
tion of species, each with a short description.
Later listings by Regel ( 1862, 1863), based upon
the specimens at Leningrad, included a total of
only 21 taxa.
Regional revisionary studies restricted to
Chloris are few, the most notable being those
of the North American species by Nash (1898),
the treatment by Swallen (1939) in l^orth
American Flora, and the publications bv Everist
( 1935a,b, 19.37, 1938) on the Queensland species.
SubsequentK , Lazarides (1972) treated Chloris
in his stud\' of the tribe Chlorideae in Austra-
lia.
TAXONOMIC RELATIONSHIPS OF CHLORIS
Most recent classifications of the Gramineae
are based heavily upon cytological and ana-
tomical characteristics; in contrast, the older sys-
tems were constructed primarily on the basis of
spikelet morphology. The higher categories rec-
ognized in the older classifications (especially
subfamilies) may be quite artificial. This is
reflected most strikingly in the modern systems
b\- the redefinition of the subfamily Festucoi-
deae, which has had many of its genera distrib-
uted to other subfamilies.
All recent treatments have placed Chloris
and most of the rest of the genera in the tribe
Chlorideae in a separate subfamily, which is
usualh' called the Eragrostoidcae or Chlori-
doideae. Tlie number of tribes included in this
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chi.obis (GnAMiNEAE)
subfamily has varied considerably; for example,
Gould ( 196Sa ) recognized seven.
Throughout the development of the more
modem classifications, the group of genera fonn-
ing the Chlorideae has remained relatively
stable, including such familiar genera as Bou-
telotia. Biichlol'. Ctcnium. Cijnodon, Entcropo-
gon, Eustacliys, Gynmopogon, and Tetrapogon,
among others.
The taxonomic parameters of these genera
have been widel)' accepted; yet intensive study
of certain complexes immediately reveals the
artificialit\' of man\' of the genera. Most inves-
tigations of generic relationships have been con-
fined to the study of Narious species of selected
genera; seldom have attempts been made to ex-
amine the majorit\' of species from a number
of viewpoints, especially for endomorphic char-
acters.
A report of such an attempt for ChJoris fol-
lows. The limited a\'ailabilit\' of material for
manv taxa has frustrated certain phases of stud\',
most notabh' those requiring living materials.
Areas and species deserving further investiga-
tion will be immediateK' apparent.
The discussion is divided into three areas:
cvtological, histological, and morphological evi-
dence. An attempt to s\nthesize these lines of
evidence as a basis of classification of the spe-
cies in the genus follows.
Cytology
No single, extensive attempt to determine
chromosome numbers in Chloris has been made,
and the counts reported in Table 1 are largely
taken from incidental reports of chromosome
numbers of a variety of grasses. In some cases
I have been able to verify the identification of
the plant b\' examining the voucher specimen;
for these, the herbarium containing the voucher
is indicated in parentheses after the counter's
name. Chromosome numbers followed by my
name are reported for the first time. Vouchers
for these are deposited in the California State
University Herbarium at Humboldt.
From Table 1 it can be seen that chromo-
some numbers have been listed for fewer than
half of the species in Chloris, and thus, only
limited conclusions can be drawn. Most spe-
cies have a base number of x=10 and onl\- in-
frequently have other numbers been noted. Rel-
atively few diploids have been identified; most
species are at various polyploid levels, with the
decaploid number of 2n = 100 being the highest
known in the genus. Several species have vary-
ing chromosome numbers. Cltloris gatjana is
noteworthy in this respect, with 2n = 20, .30, and
40.
There are few studies of reproductive mecha-
nisms. .Again, the best-known species is Chloris
gaijana. Moffett (1944) suggested that the spe-
Table 1. Cliromosome numbers in the genus Chloris.
Species
Geographic origin
Counted by
Chlom andropogonoides
C. aristata
C. Iicrroi
C. bournei
C. canterai
C. chloridea
C. ciliata
40
100
40
40
50
72
40
80
40
C. crinita
40
Te.vas (without precise locality)
Texas: Nueces Co. (TAESI); San
Patricio Co. (TAES!); Starr Co.
(TAES! TEX! UC!)
Texas: Hays Co.
Mexico: Oaxaca (USI)
Me.xico: Hidalgo (UC! US!)
Mexico: Queretaro
Mexico: Chiapas
Texas : Brazos Co. ( as a weed in a
grass nursery)
India ( without precise locality )
Without locahty
Texas: Milam Co. (TAES! UC! US!)
Mexico: Chiapas
Mexico: San Luis Potosi (US!)
Texas: Wharton Co. (TEX!)
Texas: Walker Co.
Botanical Garden, Lyon (cultivated)
Dominican Republic: Santiago
Texas: Bee Co. (TAESI)
Texas: Cameron Co.
Texas (without precise locality)
Argentina : Cafayate
Brown (1950)
Gould (1960)
Anderson 3,282 (HSC)
Tateoka (1962a) (as C. rufescens)
Gould (1965) (as C. rufescens)
Gould ( 1966) (as C. rtifescens)
Gould & Soderstrom (1970) (as C.
rufescens )
Gould (1960)
Janaki-Ammal in Darlington & Janaki-
Amma! (1945)
Krishnaswamy ( 1940 )
Gould (1958) (as C. pohjdactyla)
Gould & Soderstrom (1970)
Gould (1965)
Brown (1950)
Gould (1958)
Singh & Godward ( 1960)
Gould & Soderstrom (1967)
Gould (1968b)
Anderson 4,546 ( HSC )
Brown (1951)
Bowden & Senn (1962)
Bricham Young University Science Bulletin
Table 1. (continued)
C. cucullata
40
C. gaijana
C. halophila
C. injlata
C. vilosa
C. plurifhra
C. pyciwthrix
40
20
30
40
80
20
40
ca 50
20
30
80
30
36
40
C. radiata
40
C. Toxburghiana
20
C. scariosa
40
C. submutica
ca 65
80
Without locality
Texas: WiUacy Co. (TEX!)
Mexico: Coahuila
Texas: Brooks Co. (TAES!); Kerr
Co. (TAES!); Kimble Co. (TAESI);
Maverick Co. (TAES!); Terrell Co.
(TAES! TEX! UC!); Val Verde Co.
(TAES!)
Texas: Hays Co.
Texas: Travis Co. (3 locations)
England (cultivated)
Ethiopa; Kenya (Nzoia); South Afri-
ca; Southern Rhodesia ( Katambora ) ;
Sudan; Uganda
Southern Rhodesia
Texas ( without precise locality )
central Africa
Kenya: Kitale
Kenya ( cultivated )
Te.xas: Brazos Co. (as a weed in a
grass nursery) (TAES!); Kenedy Co.
Kenya ( cultivated )
Africa
Southern Rhodesia
central Africa
Congo: Kenya (Alengo); Southern
Rhodesia; Tanzania; Uganda
Kenya: Kitale
Tanzania: Kisarawe
Kenya: Sierra Leone (cultivated)
Bolivia: Cochabamba
Without locality
Without locality
Without locality
Taiwan
Dominican Republic: Santiago
Mexico: Yucatan
Texas: Cameron Co.
Pakistan: Karachi, Nazimabad
Nigeria (seed from USPI,
.386)
Sudan
no. 212-
Texas: San Patricio Co. (TAESI);
Willacy Co. (TAES!) (Label on
voucher indicates that n = 42)
South Africa
tropical Africa ( without precise lo-
cality )
Transvaal: Irene
Kenya: Nairobi
Botanical Garden, Lyon ( cultivated )
Kenya
Australia: Queensland (K!)
Without locality
Mexico (without precise locality)
Mexico: Durango
Botanical Garden, Berlin (cultivated)
Mexico: Michoacan ( UC! )
Mexico: Chihuahua
Mexico: (seed from USPI, no.
216-386)
Avdulov (1928)
Brown (1950)
Gould (1966)
Gould (1968b)
Anderson 3,279
Amlcrwii 2,609; 2,612; 2,614; 2,654;
3,274; 3,347; 3,356 (all HSC)
Hunter ( 1934 )
Moffett (1944)
Moffett & Hurcombe ( 1949)
Brown (1950)
Brown & Emery ( 1958 )
Singh & Godward (1963)
Pritchard & Gould ( 1964 )
Gould (1968)
Pritchard & Gould (1964)
Moffett (1944)
Moffett & Hurcombe ( 1949)
Brown & Emery ( 1958 )
Hutton (1961)
Singh & Godward (1963)
Tateoka (1965a)
Pritchard & Gould (1964)
Bowden & Senn (1962)
Avdulov (1928) (as C. barbata)
Krishnaswamy (1940)
Janaki-Ammal in Darlington &
Janaki-Animal (1945) (as C. barbata)
Chen & Hsu (1962) (as C. barbata)
Gould & Soderstrom (1967) (as C.
barbata )
Gould & Soderstrom ( 1970)
Anderson 4,545 (HSC)
Baquar and Saeed (1969) (as C
barbata )
Anderson 3,435 (HSC)
Thomas in Darlington & Wylie
(1955)
Gould (1968b)
de Wet (1954)
Thomas in Darlington & Wylie ( 1955)
Moffett & Hurcombe ( 1949)
Tateoka (1965a)
Singh & Godward ( 1960)
Pritchard & Gould (1964)
Singh & Godward ( 1960)
Krishnaswamy ( 1940 )
Avdulov (1931)
Gould (1960)
Singh & Godward ( 1960)
Gould (1965)
Reeder (1971)
Anderson (HSC)
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gr.\mineae)
Table 1. (continued)
C. tnincata 40
Australia ( without precise locality )
Without locality
Australia (seed from USPI, no.
212-389)
Avdulov (1928)
Krishnaswamy (1940)
Anderson (HSC)
C. verticillata n =
= ca 28
Texas, Mason Co.
Gould (1960)
2n = 40
Texas: Gray Co.
Texas: Archer Co.; Bosque
Brazos Co.; Eastland Co.; Jack
Wise Co.
Co.;
Co.;
Brown (1950)
Gould (1968b)
ca 40
Kansas: Shawnee Co.
Anderson 2,768; 2,769 ( HSC )
40
Nebraska; Dundy Co.
Texas: Sterling Co.
Texas: Travis Co.
Anderson 2,774; 2,776 (HSC)
Anderson 3,002 (HSC)
Anderson 3,338; 3,347; 3,356;
3,361; (HSC)
63
Texas: Montague Co.
Gould (1968b)
C. vir'^ata
20
South Africa: Cape Provinces, Mafe-
king (K!)
Moffett & Hurcombe (1949)
Te.\as ( without precise locality )
Brown (1950, 1951)
Botanical Garden, Bedin (cultiva
ited)
Singh & Godward (1960)
Mexico: Baja California Sur;
Coahuila; Qucrctaro
Gould (1966)
Australia (without precise locality)
Pritchard & Gould (1964)
Texas: Castro Co. (TAES!)
Gould (1968b)
India: Gorakhpur
Gupta (1969)
Texas: Brewster Co.
Anderson 2,579 (HSC)
Madagascar (seed from USPI. no.
219-956)
Anderson (HSC)
26
Without locality
Thomas in Darlington & Wylie ( 1955)
30
Without locality
Krishnaswamy (1940)
40
Without locality
Krishnaswamy (1940) (as C. caudata)
C/iZarts-probable hybrids
n =
= ca 20
Texas: Brazos Co. (TAES!)
Gould (1960) (as C. subdolicho-
stachya )
n =
= ca 20
Texas: San Patricio Co.
Gould (1960) (as C. hitisifutimea)
n =
= ca 31
Texas: Willacy Co. (TAES!)
Gould (1960) (as C. latisquamea)
n =
= ca 32
Texas: Brazos Co. (TAES! )
Gould (1958) (as C. latisquamea)
n =
= ca 33
Texas: Llano Co. (TAES!)
Gould (1960) (as C. suhdolicho-
stachija )
n =
= ca 36
Te.\as: Lampasas Co. (TAES!)
Gould (1958) (as C. latisquamea)
n=42
Texas: Llano Co.
Gould (1958) (as C. latisquamea)
2n =
= ca 56
Te.\as: Mason Co.
Gould (1968b) (as C. latisquamea)
2n =
= ca 68
Texas: Williamson Co.
Gould (1968b) (as C. latisquamea)
'Tlie n number is fjiven vvlieie the roimt uiisinally reported is not logically convertible to the '2n number.
cies was an aponiict. Brown and Emery ( 1958)
reported the occurrence of four-nucleate enibrvo
sacs, a character associated witli apomicts. Hut-
ton (1961), working with different material, re-
ported eight-nucleate, apparently normal, em-
bryo sacs that also showed delayed embryo and
endosperm formation (an unusual feature).
These studies were carried out primarily on
tetraploids. Bogdan (1961) reported both dip-
loid and tetraploid forms to be sexual. This
same conclusion was borne out by circumstantial
evidence in the study bv Pritchard and Gould
(1964),
My own field observations in the Chloris
andropogonoides-C . cucuUata-C. verticillata hy-
brid complex show high seedset in plants with
low pollen stainability, circumstantially suggest-
ing apomixis in this group.
Leaf Epidermis
The epidermal studies in Chloris which are
reported here were initiated by Rogers ( 1967 ) ,
who examined 36 species in this and allied gen-
era. These studies were later expanded by my-
self, and a total of .55 species has now been ex-
amined.
Rogers studied man)' epidermal features of
a large number of specimens of the wide-rang-
ing species, Chloris virgata, in order to estab-
lish a basis for the selection of epidermal char-
acters, to detemiine the spectrum of variation,
and to assess the relationship of geographic ori-
gin to character expression. Table 2 gives the
origin ot these sjiecimt'ns and the epidermal
characters surveyed.
Rogers's study indicated that the density of
prickles varied randomly with respect to geog-
Brigham Young Univehsity Science Bulletin
Oo o OoOopgoOO
80888SRRSSOO
0,0,0,8,^,^,0,0,0, 0,0.0,
8§Mf 9 99" #tii
ogggo 88g§ooq
8919998 8 @99
n n '^ ^ ( — in C-J d —
©eeoo V
OQ R Qf^
On n 9 Q Sy
vv
O 8 Oo
WW XX n^-^ZZ AAA^BI
Fig. 1. Microhairs (upper figure of each set) and
costal silica cells (lower figure) of Austrochlom,
Chloric, Enleropogon, Eustachi/s, and Tetrapogon.
(A) Austrochloris dichanthioides (Everist) Laza-
rides; (B) Chloris brandegei (Vasey) Swallen;
(C) C. divaricata R. Brown; (D) C. submutica
Humboldt-Bonpland-Kunth; ( E ) Eustachys petraea
Swartz; (F) Chloris bournei Rangaehariar and
Tadulingam; (G) Euatachijs distichnplujUa La-
gasca; ( H ) Chloris scariosa F. von Mueller; ( I )
Eustachys retusa Lagasca; (J) Chloris halophila
Parodi; (K) C. ventricosa R. Brown; (L) C. mos.^-
flni/;iVt'ti.siv Schumann; (M) Tetrapogon macrnnlhns
(Jaubert and Spach) Bentham; (N) Chloris pec-
tinaia Bentham; (O) C. robusta Stapf in Chevalier;
(P) C paniculata Scribner in Robinson; (Q) C.
roxburghiana Schultes; (R) Tetrapogon spathaccus
( Hochstetter) Hackel ex Durand and Schinz; (S)
Chlorii ciliata Swartz; (T) C. prieurii Kunth; (U)
C sagracana A. Richard; (V) C. suriiigari Hitch-
cock in Urban; (W) Tetrapogon tcncllus (Rox-
burgh) Chiovenda; (X) Chloris chloridea (Presl)
Hitchcock; (Y) C. somalensis Rendle; (Z) Entero-
pogon dolichostachya ( Lagasca ) Lazarides; ( AA )
Chloris dementis Merrill; (BB) C. radiata (Lin-
naeus) Swartz; (CC) C. texensii Nash; (DD) C.
mollis (Nees) Swallen; ( EE ) Tetrapogon villosus
Desfontaines; (FF) Eustachys paspaloidet (Vahl)
Lanza and Mattei; (GG) E. glauca Chapman;
(HH) Chloris ekmanii Hitchcock; (U) C. pumilio
R. Brown; (JJ) C cuhensii Hitchcock and Ekman;
( KK ) C. ilandi/ana C. D. Adams; ( LL ) Eiitero-
pogon acicularls (Lindley) Lazarides; (MM)
Eustachys neglecla Nash; ( NN ) Chloris truncata
R. Brown; ( OO ) C andropogonoides Fournier;
(FF) C. berroi Arechavaleta; (QQ) C. cruciate
(Linnaeus) Swartz; (RR) C canterai Arechava-
leta; (SS) C. amethystea Hoclistetter; (TT) C.
lobata Lazarides; (UU) Eustachys tenera (Presl)
A. Camus; (VV) Chloris pycnothrix Trinius; (WW)
C. aristata (Cervantes) Swallen; (XX) C. virgata
Swartz; (YY) C. pilosa Schumacher; (ZZ) C. gay-
ana Kunth; (AAA) C. inflata Link; (BBB) C. or-
thonoton Doell in Martins. Arrangement is by over-
all similarity of microhairs. X 475.
raphy, from ab.sont to moderately abundant.
Because of this random variabilit)', prickles were
not emphasized as a taxonomic character.
Of the otlier epidemial characters studied,
Rogers concluded that the most stable were
shape, size, and type of microhair and the shape
of the costal silica cells. Typical microhairs and
costal silica cells are illustrated ( Fig. 1 ) for
40 species of Cliloris and for 14 other species
which were formerly placed in Chloris, but
which are now placed in other genera.
Two types of microhairs, unicellular and bi-
cellular, were observed in the taxa studied.
Table 3 lists the species showing these two
microhair t)'pes. One species complex (consist-
ing of C. aristata, C. gaijana, C. inflata, C. ortho-
noton, C. pilosa. C. jx/rnothrix, and C. virgata)
possesses vmieelliilar microhairs. The spikelets
and inflorescences of these species are similar to
one another, yet they are not unique in the
genus, and several strongly resemble particular
species with bicellular liairs. The remaining
species all have bicellular microhairs. While
several different shapes and sizes of hairs are
recognizable, these cannot be correlated with
an\' macroscopic features.
\'irtuall\' all of the taxa examined have
costal silica cells var\ing from cuboidal to
saddle shaped (Fig. 1; Table 3). Only one spe-
cies differs from this pattern: C cruciata has
costal silica cells which tend to be more elon-
gated and to ha\'e more deeply indented end
walls than the remaining species.
Anatomy. Embryology, and Seedling
Morphology
Anatomical, embr\ological, and seedling
studies on a selected group of 28 taxa in Chloris
and a]Ii(>d genera were completed by Jensen
( 1969 ) . The species were selected to be repre-
sentati\'e of as man\' different sections of the
genus as possible in terms of spikelet morphol-
og\'. Characteristics studied included leaf anat-
omy, embryo stnicture, root-hair position and
aspect, shape of the first seedling leaf, and t\'pe
of starch grains. Individual discussions of these
features follow.
Jensen examined several anatomical features
of leaf cross sections, including the shape of the
Biological Series, Vol. 19, No. 2 Ta.xonomy of the Genus Chloris (Ghamineae)
00
CO
.9^
S
nr
o
«
u -w
"â– a
<
c
â– s -a
H
p to
CJ
P c
6
-.^
l-H
(U
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3:5
o
U
â– 5
c
•C a.
so
s
3
2
1— 1 1-H
4 ei
i-H 1—1
in
CO
1
1
1
i-H
1
â– ^ -"T
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CO
CO
.—I
CO
o
ol
rH ■— (
3
^H
â– ^
oi
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g g
8
3
o
CD
§
05 Tf
g
g
i d> i i
^ O Tt* "^
â– *j* Tj< CO d â– *** '^
ome
dome
dome
dome
1
o
-t3
1 i
-high d
-triang.
-triang.
-triang.
bi)
C
a
â– c
-triang.
-triang.
dome
dome
-triang.
Tj -0 -a -a
tu 0) 0) 0;
e S E E
1
-0 IS -0
dj OJ 1;
E S E
-0 TJ
E 1
21-25
14-24
19-22
20-22
OI
^ CO — «
Cvl Ol Ol
d> i ^
rt fM — (
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f-H I-H
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3-15
21-27
.5-18
6-20
05
3
CO r- lo CD CD
^H ^H ^H ^H ^H
â– 4< -3" O â– "t â– *
oj a> oj u
t« l-< Ui Im
c4 c4 c4 cd
3 3 3 3
cr tr cr IT
o o c o
X !< X X
rt « ctf rt
CO TT in CD
3 3^4
3 5- 3
CO VI
_g 2 o
rt cu rt c3 rt ?3
M" CO -^ c-l O Tt
I I I J I T
-H rt — I O 00 "
t~ in in -^
4 4
in oi oi oi o oi
-H -H o c3> 00 d>
4-» ■•H
o
—I m
7o^
o
U
•3
&
feo
c
«
1
Z
3
Q
3
Q
o
>h3
n-.S
C5
c
o
CJ
o c
<;
C/1
X
1>
â– %&
53 >
D
s
.^
s
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I I J iM O O
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tS
S - .C
e
X t:
^ o
-< Z
Table .3. Epideniud characteristics of selected species
of Austrochloris, Chloris, Enteropogon, Eustachys, and
Tetrapogon.
Costal silica
Figure 1
Species
cell shape
reference
Chloris species
with hicellular microhairs
C. amethystea
cuboidal
SS
C. andropogonoides
saddle
oo
C. herroi
cuboid-saddle
pp
C. hournei
cuboid-saddle
F
C. hrandegei
short saddle
B
C. canterai
cuboidal
RR
C. eld or idea
saddle
X
C. ciliata
cuboidal
s
C. dementis
cuboid-saddle
AA
C. cruciata
very short saddle
to cross shaped
QQ
C. cuhcusis
short saddle
JJ
C. dandyana
saddle
KK
C. divaricatd
narrow saddle
c
C. ekmanii
cuboidal
HH
C. halophila
cuboid-saddle
J
C. lohata
cuboidal
TT
C. mollis
saddle
DD
C. mossamhicensis
cuboid-saddle
L
C. paniculata
round-cuboid
P
C. pcctinata
sho t saddle
N
C. pricurii
saddle
T
C. pumilio
cuboid-saddle
II
C. radiata
cuboid-saddle
BB
C. robtista
saddle
O
C. roxhurghiana
short saddle
Q
C. sagraeana
short saddle
U
C. scariosa
cuboid-saddle
H
C. somalensis
cuboid-saddle
Y
C. sidimtitica
short saddle
D
C. suringari
saddle
V
C. texensUi
short saddle
CC
C. truncata
cuboid-saddle
NN
C. ventricosa
cuboid-saddle
K
Chloris species
tfif/i unicellular microha
irs
C. aristata
cuboid-saddle
WW
C. gai/ana
cuboid-saddle
zz
C. inflata
cuboid-saddle
AAA
C. orthonoton
cuboid-saddle
BBB
C. pilosa
cuboid-saddle
YY
C. pycnothrix
saddle
VV
C. virgata
short saddle
XX
Species formerly i» Chloris, now excluded and placed
in related genera, all with hicellular microhairs
AtLiirochloris dichanthioid
es culxjid-saddle
A
Enteropogon acicularis
short saddle
LL
E. dolichostaclnjs
culx>idal
Z
Eustachys distichophylla
culx)id-saddle
G
E. glauca
cuboid-saddle
GG
E. neglecta
.short-saddle
MM
E. paspaloides
cuboidal
FF
E. petraea
culx)idal
E
E. retusa
saddle
I
E. tenera
cuboidal
UU
Tetrapogon macranthus
cuboidal
M
T. spathaceus
cuboid-saddle
R
T. tenellus
saddle
W
T. villosus
cuboidal
EE
Brigham Young University Science Bulletin
keel, the disposition of vascular bundles, the tex-
ture of the adaxial surface (ribbed or smooth),
the extent of the sclerenchyma girder, the dis-
tribution of bullifomi cells, and the nature of
the vascular bundle sheath. While individual
species may show stable and recognizable pat-
terns, few correlations between leaf anatomy
and spikelet morphology could be made. Like-
wise, there were no consistent correlations of
anatomical features with either microhair or
costal silica cell types.
The classical study of Rceder (1957) in
comparing anatomical features of embryos of
various genera of grasses is well known. Reeder
separated the grass genera studied into essential-
ly two groups, panicoid and festucoid, on the
basis of four embiTO characteristics: ( 1) whether
the lower portion of the scutellum is separate
from the rest of the embr)'0 (panicoid) or fused
(festucoid); (2) the presence of an epiblast
(festucoid) or its absence (panicoid): (3)
whether both vascular traces diverge from about
the same point (festucoid) or are separated by
an elongated area of vascular tissue between
points of divergence (panicoid); and (4) wheth-
er the margins of the embrvonic leaf overlap
(panicoid) or just meet (festucoid).
Jensen (1969) studied three of these charac-
teristics for 25 selected species in C^'Inris and
certain related segregate genera (Table 4).
Typically, all species studied had the scutellum
separate from the rest of embryo (panicoid), an
epiblast present (festucoid), and vascular traces
separated by an elongated area of vascular tis-
sue (panicoid).
Tlie fourth feature, whether or not the mar-
gins of the embr\'onie leaf overlap, was studied
on 19 of Jensen's 2S taxa (Table 5). While
both festucoid and panicoid types are repre-
sented among these species, consistent correla-
tions with other anatomical features or with
Table 4. Ta.xa in CMoris and related segregate genera
having panicoid scutellum type, an epiblast present
(festucoid), and a panicoid arrangement of vascu-
lar tissue.
CMoris aristata
C. herroi
C. hournei
C. prieurii
C. ptjcnothrix
C. radiata
C. canterai
C. rohii.tta
C. cMoridea
C. sidfmutica
C. cruciata
C. truncata
C. dandyana
C. divaricata
C. virgata
Enteropogon acicularis
C. gaijana
C. halopMla
C. inflata
C. mollis
C. ortlionoton
Eustachys distichopliylla
E. glauca
E. neglecta
E. petraea
Table 5. Comparison of embryonic leaf types in se-
lected species of CMoris and related genera.
Species with panicoid embryonic leaves
Chloris herroi C. prieurii
C. chloridea
Species v\ith festucoid embryonic leaves
Chloris artktata C. polydactyhi
C. hournei C. pycnothrix
C. canterai C. radiata
C. cruciata C. rohusta
C. divaricata C. virgata
C. halophila Eusiachys distichophylla
C. mollis E. glauca
C. ortlionoton E. neglecta
gross spikelet morphology are absent. Those
species with festucoid embryonic leaves, for ex-
ample, represent diverse forms with respect to
microhairs and costal silica cells, as well as
spikelet characteristics.
Tateoka ( 1962b) recognized four starch-grain
types— simple, compound, or some variation of
each— as being characteristic of various groups
of grasses. Jensen reported that compound
starch grains were widespread in the species
studied, with one notable exception, Enteropo-
gon acicularis, which had simple starch grains.
This species had been placed in Chloris until re-
centh', wlien it was removed b\' Lazarides
(1972). Jensen found that the following species
all have compound starch grains: Chloris aristata,
C. herroi, C. hournei, C. canterai, C. chloridea,
C. cruciata. C. dandijana, C. divaricata, C. gay-
ana, C. haIo])hihi, C. inflata, C. mollis, C. ortho-
noton, C. prieurii. C. ptjcnothrix, C. radiata,
C. rohttsta, C. suhmutica, C. truncata, C. vir-
gata, Eustachi/s distichopht/Ua, E. glauca, E. ne-
glecta. and E. petraea. I ha\e examined the
starch grains of C. crinita. C. pluriflora, and C.
scariosa; they are all compound.
Differences in root hair development in
grasses were first shown b\' Sinnott ( 19.39 ) and
Sinnott and Bloch ( 1939). Later, the taxonomic
significance was pointed out bv Reeder and von
Maltzalin (19.53) and Row and Reeder (1957).
These authors described one type in which the
root hair piojeets forward at about a 45" angle
from the smaller of two epidermal cells. A
second kind has the root hair developing near
the middle of either sister epidermal cell and
projecting outuard at an angle of about 90°.
A third t\'pe has sister cells of the epidennis
slightly different in size, with the root hair pro-
jecting at an angle of about 60° near one end of
the cell, though Row and Reeder suggest that
tlie variability' in size of the sister cells is more
reliable than the angle of projection. Because
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Cramineae)
of the limited availability of viable seeds, Jensen
was able to examine onh' ten species for this
characteristic, all of which had the root hair pro-
jecting at an angle of about 60° near one end of
the cell. The species examined were: Chloris
canterai, C. gayana, C. inflata, C. pilosa, C.
pijcnothrix. C. stil)mutica. C. truncata, Eiistaclnjs
caribaca, E. distichopluiUa. E. petraca, and E.
rettisa.
The first seedling leaf in panicoid grasses is
relatively wide and curled; that of festucoids is
long and narrow ( Stebbins, 1956 ) . The species
listed above all had panicoid seedling leaves, ac-
cording to Jensen.
Relationships of Chloris and Other Genera
As with manv genera in the Gramineae,
generic delimitations in the Chlorideae may be
difficult to assess; and they, of necessity, may
be somewhat arbitrar\' and artificial. The com-
plex that includes Chloris is no exception. While
most of the species have long been included
within the genus and most of the characteristics
recognized, some species have been placed in
adjacent genera in the past, a few more recentK'
so. A graphical conspectus of Cliloris and re-
lated genera is presented in Fig. 2. The number
of taxa commonly accepted in each is indicated
by the relative size of the circle. Also given
arc those species in ClUoris that resemble the
satellite genera, as well as those in the satellite
genera that have been included in Chloris.
There is little infonnation, aside from spike-
let moiphology, that is useful in generic delimi-
tation within this group. Chromosome counts
are mostly of the same basic number; leaf epi-
dermal and anatomical characteristics do not
appear uniciue to particular genera; genetic in-
fonnation is \irtuall\- nonexistent. Consequently,
speculation on relationships, both intergeneric
and interspecific, rests heavilv upon infomiation
gleaned primarily from the structure of the in-
florescence and spikelet and, secondarily, upon
data concerning vegetative features. However,
the genera in this complex have been long
recognized and have remained relatively un-
changed. The description of each of these, as
commonly accepted toda}-, is given in Table 6,
and their relationships to Chloris follow.
Eustachi/s. This genus was first described
in 1810 by Desvaux, but it has been treated
subsequently as a subgenus of Chloris bv the
EOSTAPFIELLA
NEGLECTA
PASPALOIDES
PETRAEA
EU ST ACHYS
RETUSA
)
A U S T R C H L R I S
TENERA
ULIGINOSA
DISTICHOPHYLLA
/
FLORIDANA
y
iSLAUCA
DELICATULA
G YMNOPOGO
UNISPICEUS
E N T E R P G N
ACICULARIS
DOLICHOSTACHYA
;
D A K N P H L I S
X'CYNOCHLOR I S
Fig. 2. Relationships of Chloris and allied genera.
10 Brigham Young University Science Bulletin
Table 6. Comparison of various characteristics of Chloris and related genera.
Genus
Vegetative characteristics
Inflorescence type
Glumes
Number
Chloris
Cyrtodon
Daknopholis
Enteropogon
tufted, rhizomatous or sto-
loniferous; sheaths usually
overlapping at the base,
blades narrow to broad, not
distichous
rhizomatous to stoloniferous;
sheaths usually overlapping
at the base, blades generally
narrow, acute, distichous
prostrate, repent to stolonif-
erous; leaves basal, sheaths
overlapping, blades small,
subelliptic, obtuse
erect, tufted, sheaths usual-
ly not markedly overlapping
at the base, blades narrow,
flat or convolute
usually a single radiate se-
ries of 4-10 spikes, occasion-
ally 2-more approximate or
only slightly separated series
of 10-20 spikes; rarely an
indefinite number of termi-
nal spikes or as few as 1-2
spikes
a single radiate series of
4-9 spikes
a single radiate series of
2-4 spikes
usually a single terminal
spike, sometimes several to
many in a radiate or sub-
verticillate series
both glumes simi-
lar, narrowly ovate
to lanceolate; acute
to acuminate; awn-
usually 1,
rarely 2 and
then in the
same inflor-
less or only awn-
tipped
escence as
spikelets
with
1
both glumes nar-
rowly lanceolate
to ovate
1
first glume ovate;
second glume tnm-
cate, apex marked-
ly truncate and
1
erose
both glumes nar-
rowly lanceolate to
ovate
1
Eustachys
Gymnopogon
tufted, rhizomatous or sto-
loniferous; sheaths strongly
overlapping, largely basal,
equitant; blades generally
broad, obtuse, occasionally
acute
tufted, sheaths strongly over-
lapping, largely basal;
blades markedly distichous,
stiff, acute to acuminate
usually a single radiate sc-
ries of 4-30 spikes, rarely
only 1-2
a single series of 6-20
spikes, generally borne sing-
ly at a node
first glume ovate-
lanceolate, acute,
awn-tipped or
awnless; second
glume linear to
linear - lanceolate,
retuse, short awned
between the lobes
both glumes nar-
rowly lanceolate,
acuminate
usually 1,
sometimes 2
Neostapfiella erect, tufted; sheaths some-
what overlapping; blades
narrow
Tetrapogon tufted to short stoloniferous
or rhizomatous; sheaths usu-
ally overlapping at the base,
blades narrow to broad, not
distichous
2-3 spikes
both glumes nar-
rowly lanceolate,
acuminate
1 or 2
a single radiate
1-4 spikes
series
of
both glumes simi-
lar, ovate to lance-
olate
usually 2 or
3
majority of authors, the most notable exception
being Nash (1898). EttstacJnjs is obviously
closely related to Chloris, though distinct in
several features. Vegetatively, the strongly flat-
tened, folded, equitant leaf sheaths set Eu-
stachys apart from most species in Chloris. In
addition, the second glume in Eustachys is bi-
lobed at the apex, with a short awn arising be-
tween the lobes; in Chloris the second glume is
acute to acuminate and is rarely awned. The
fertile and sterile florets of Eustachys are com-
monly pale to dark brown to almost black and
are awnless or short-awned. Florets in Chloris
are usually pale to tawny, seldom darker, and
Biological Series,
Vol. 19, No. 2
Taxonomy
OF THE Genus Ciilohis
( Gramineae)
11
Spikelet characters
Fertile floret(s)
Sterile
Awn
Pubescence
: floret(s)
Shape and color
Keel
Margins
Callus
Number
Size and shape
elliptic to hinceo-
usually well
glabrous to
usually
bearded
usually 1 —
usually markedly
late to ovate; often
developed.
jilose to
ciliate or
sometimes
dissimilar from fer-
acute, sometimes
rarely ab-
ong-ciliate
appressed
2 — more
tile floret; usually
obtuse or acumi-
sent
pubescent
rarely
well developed,
nate, rarely bifid;
several
rarely reduced
usually laterally
compressed; tawny
to light brown.
rarely black or
dark brown
ovate to lanceo-
awnless
short
short
short-
to 1,
absent, or, if pres-
late; prominently
ciUate
cUiate
pubescent
highly
ent, reduced and
laterally compres-
reduced
forming only a knob
sed; tawny
at ape.x of rachilla
joint
lanceolate to ellip-
short awned
scabrous
scabrous
bearded
obsolete
tic; prominently
to nnicronate
laterally compress-
ed; tawny
dorsally compress-
usually well
scabrous
scabrous
bearded
1
similar to fertile
ed to nearly latte-
developed
floret, but much
rete, but not lat-
smaller
erally compressed.
linear in side view.
elliptic to lanceo-
late in dorsal view;
tawny to greenish
elliptic, lanceolate
short or ab-
usuallv glab-
conimonlv
bearded
usually 1,
markedly dissimilar
or ovate, acute to
sent
rous, some-
appressed
sometimes 2
from fertile floret;
shortly acuminate;
times pilose
pubescent
usually well devel-
strongly laterally
oped, rarely re-
compressed; often
duced
dark brown or
black, rarely light
brown
usually narrowly
usualh' well
glabrous.
sparsely to
bearded
1-2
highly reduced to
elliptic; nearlv te-
de\eloped
scabrous, or
densely
obsolete
rete to slightly dor-
pilose
ciliate
sally or laterally
compressed; tau ny.
brown or purplish
ovate to lanceo-
well devel-
glabrous to
glabrous
densely
1
similar to fertile
late; laterally com-
oped
pilose
to sparsely
long
floret but slightly
pressed; tawny
ciliate
bearded
smaller
elliptic to broadly
usually well
densely
densely
densely
usually 2-4,
similar to fertile
lanceolate; lateral-
developed
pubescent
pubescent
bearded
sometimes 1
floret but smaller;
ly compressed; us-
lower well devel-
ually tawny
oped, upper small-
er
are usualh' prominenth' awned. Cliloris sub-
mutica is perhaps most similar to Ettstacliys, pri-
maril\ because of the lack of prominent awns.
Sanchez ( 1971 ) enumerated several anatomi-
cal features bv which the leaves of Cliloris and
Eustaclii/s differ. Sanchez reported that Eu-
staclujs has, for the most part, relatively thick-
walled, uniformh- sized bulliform cells; a promi-
nent epidemial band of these lies adaxial to the
midrib. By contrast, Chloris has unequally sized,
thin-walled bulliform cells and lacks a concen-
tration of these in the midrib region. While
none of these characters appear to be exclusive
to either genus, the trend shown clearly re-
12
Bricham Young University Science Bulletin
inforces the exomoiphic differences between
them.
Tetrapogon. This Old World genus is best
separated from Chloris on the basis of having
two ( occasionally three ) fertile florets per spike-
let. Chloris regularly has a single fertile floret,
though very occasionally individuals of certain
species may produce a spikelet with two or more
fertile florets. In addition, both glumes in Tetra-
pogon are thin, papery, or parchmentlike and
(like the spikelets) are relatively large. The
glumes in species of Clihris are not thin and
papery, but are finner and relatively smaller.
Several species in Chloris strongly resemble
Tetrapogon. Chloris inossamhicetisis, an African
species, is similar in many respects, but the bulk
of material which this author has examined has
spikelets with one fertile floret; only a few plants
have spikelets with two or more florets. Chloris
bournei, C. tcightiana, and, to a lesser extent,
C. quinquesetica—aW from India— are similar to
several Tetrapogon species, primarily in size of
spikelets. However, they have just a single fer-
tile floret and are best retained in Chloris.
The strikingly different Chloris scariosa from
Australia could easily be given generic status,
for its spikelets (with their several flabellate
sterile florets) arc unlike any other species in
Chloris. Chloris scariosa has relatively large,
papery glumes, in this respect strongly resem-
bling Tetrapogon. I am maintaining this spe-
cies in Chloris largely because of its single fer-
tile floret. If Chloris scariosa were to be given
generic rank, then a number of other somewhat
unusual species are also deser\ing of such con-
sideration—a process that could hopelessly frag-
mentize the genera in this complex.
Gymnopogon. The most striking difference
between this genus and Cliloris is vegetative.
Qymnopogon is characterized by thick, sharp-
pointed, distichous blades, the sheaths of which
are more or less cquitant and basal. Further-
more, many species of Gymnopogon have the
spikes well separated and spreading, making the
inflorescence quite open. Chloris generally has
broader, more lax blades and sheaths that only
partly overlap. While a few species have distant
spikes, they are mostly borne in one or more
radiating series.
Most species of Gymnopogon either lack
sterile florets or they are very greatly reduced.
In Chloris the sterile floret is generally well
developed.
Overall, Chloris mollis is probably most simi-
lar to Gymnopogon, a fact reflected in its no-
menclatural histoiy. While the leaf blades of
C. mollis are relatively short and sharp pointed.
the sheaths are not as strongly overlapping. The
spikes are somewhat distant, but not as markedly
so as in Gymnopogon. Sterile florets in C. mollis
are well developed; in this respect they are un-
like those in Gymnopogon.
Chloris dementis, C. ekmanii, C. pycno-
thrix, and C. radiata all have mucli reduced
sterile florets; but in all other respects they fit
Chloris well.
Cynodon. Clifford and Everist (1964) de-
scribed a sterile intcrgeneric h\brid, which they
named Cynochloris macivorii, which was found
growing with Cynodon dactylon and Chloris
divaricata on a lawn bowling rink at Ipswich,
Queensland. The identification of this plant
rests upon its intermediacv between the two
putati\e parents with respect to a number of
spikelet, habit, and behavioral characteristics,
since experimental recreation of the hybrid has
not been attempted. On the basis of the data,
there is little doubt as to the correctness of Clif-
ford and Everist's inteipretation, however.
The existence of this bigeneric hybrid at-
tests to the genetic proximity of these two gen-
era; \et the two can ordinarily be separated
readily by morphological features. Most species
of Cynodon have obsolete or verv poorly de-
veloped sterile florets, while in Chloris they are
usually well developed. Likewise, in Chloris
the callus is prominentK' short bearded, whereas
in Cynodon it is less obviously pubescent.
Daknopholis. Described in 1967 by Clayton,
this genus was erected to contain several anom-
alous species of Chloris: C. hoivinii, C. perrieri,
and C. ramosissima. Two features will separate
the new genus from Chloris: ( 1 ) the lack of
a well-defined sterile floret and (2) the presence
of a truncate and erose second glume. In over-
all appearance Daknopholis is more suggestive
of Cynodon than Chloris. The relationships
were discussed in some detail by Clayton
(1967).
Entcropogon. Tliese Old World trooical pe-
rennials (as treated b\' Clayton [19 17], for ex-
ample) are easily separated from Chloris in hav-
ing a single spike or, rarely, two; most Chloris
species have three or more .spikes. In addition,
the fertile lemmas of Entcropogon are strongly,
dorsally compressed and have a raised midnerve.
I do not believe that this difference in num-
ber of spikes accurately reflects the relationships.
Two species with many spikes that are often
treated in Chloris— C. acicularis from Australia
and C. dolichostachya from southeastern Asia-
have strongly dorsally compressed spikelets with
a raised midnerve on the lemma. The similarity
to Enteropogon is such that one would be hard
Biological Series, Vol. 19, No. 2 T.\.\onomv of the Genus Chloris (Cramineae)
13
pressed to separate these two taxa of Chloris
and various species of Enteropogon on the basis
of spikelets alone. They could, of course, be
separated readily on the number of spikes. But
the differences do not end here. Tateoka ( 1962b )
reported that starch grains were simple and an-
gular in Enteropogon, while Chloris has only
compound grains. Both CJdoris acicularis and
C. dolichostaduja have Enteropogon-ty^e starch
grains, further reinforcing the removal of these
species from Chloris. Accordingly, I have fol-
lowed Lazarides (1972) in removing these two
species from Chloris and referring them to En-
teropogon.
Trichloris. This genus has been character-
ized as having three prominent awns on the
fertile and sterile lemmas: one is an extension
of the midnerve and the other two are of lateral
nerves. The relative length of these awns varies
between species.
The prominently threc-awned lemmas of
Trichloris will distinguish it readily from virtual-
ly all species of Chloris in the Western Hemis-
phere. Chloris chloridca and C. halophila have
somewhat bilobed lemma apices, but these are
mucronate, at most.
This same awn difference serves also for the
Eastern Hemisphere, except in Australia. There
are several Australian species, too, that have
lateral nerves extended into awns (prominently
so in C. lohata and C. punulio; less obvious in
C. ilivaricata and C. pectinata.) In all likelihood,
of course, this resemblance is a result of parallel
evolution and docs not reflect a close genetic
relationship. Nevertheless, it is ambiguous to
continue to recognize Trichloris in the New
World, while maintaining the three-awned spe-
cies from Australia in Chloris. Clayton (1967)
recognized this ambiguity and treated the two
species of Trichloris as members of Chloris.
More recently, Sanchez (1971) resurrected
Trichloris on the basis of anatomical studies of
leaves. According to Sanchez, the bulliform
cells of most Argentine species of Chloris have
achloroph\llous cells hing immediatelv below
them in the mesophyll. Such achlorophvllous
cells are onlv rarely found in Trichloris.
While there are recognizable tendencies in
the patterns of variation in these anatomical
features, there is no reinforcement from spike-
let morphology. Consequently, I am following
the lead of Clayton ( 1967 ) in treating the two
species of Trichloris as members of Chloris.
Neostapfiella. This genus from Madagascar
is relatively poorly represented in most herbaria.
The limited material available, however, shows
the spikelets with either two basal fertile florets
or with the single terminal sterile floret identical
to, or strongly resembling, the basal fertile
floret. Additionally, the number of spikes (one
to three) is fewer than that generally found in
Chloris. Neostapfiella is probably more closely
related to Tetrapogon than to any species in
Chloris.
Relationships within Chloris
This synthesis is based primarily upon the
gross morphologv of the inflorescence and spike-
let; but studies of the leaf epidermis, already
discussed, provided important additional infor-
mation. Chromosome number and structure have
contributed virtually nothing, for the genus is
essentially monobasic and the chromosomes very
similar.
Some species complexes are well defined; i.e.,
there is a strong correlation of morphology, an-
atomv, and geographv. Other species may re-
semble one another, but they are geographically
isolated in such a wav as to preclude a direct
cvolutionarv- connection. In still other complexes
there may be little or no outward resemblance;
vet the formation of hybrid swarms recombines
those highly divergent characters and indicates
strong cvolutionan' ties. Still other taxa are
isolated entities, displaying no obvious relation-
ships with any other species or complexes.
Tlie formal recognition of a series of sub-
genera would be folly, not that some complexes
are unworthy of it, but because the naming of
these as subgenera will necessarily create a resi-
due of species that cannot be grouped. The in-
dividual species in this remnant cannot be
treated as a single subgenus, for such would
be blatantly artificial and misleading. The other
extreme would be to treat each as the single
member of its own subgenus, a practice leading
to rather hopeless fragmentation. I have tried
to compromise in the following discussion. Those
groups containing obviously closely related spe-
cies—whether the basis is morphological, geo-
graphical, genetic, or all three— are treated as
species complexes and discussed as such. The
isolated species are discussed individually wher-
ever this author feels that they have importance.
In addition, discussions of relationships, espe-
cially within the various complexes, follow the
descriptions provided later. A graphical synop-
sis of relationships within the genus is provided
in Fig. 3.
The Chloris inflate complex.
Members of this complex are characterized
by single-celled microhairs, by typical Chloris-
type silica cells, and, in most, by a prominent
14
Brigham Young Univebsity Science Bulletin
burmensts
scavvosa
robusta
anethystea
mossambicensis
somalensis
filiformis
Qubmutica
paniaulata
Fig. 3. Relationships of species and species complexes in the genus Chloris.
tuft of hairs on the upper margins of the fer-
tile lemma. Included here arc five very widely
distributed species {Chloris gmjana, C. inflata,
C. pilosa, C. pi/cnothrix, and C. virgata), as well
as several more restricted species (C. aristata,
C. castilloniana, C. jormosana, and C. orthono-
ton). Of these, Chloris jn/cnothrix is the most
aberrant; if it were not for possession of unicel-
lular microhairs, it would be placed confidently
in the following complex.
The Chloris radiata complex.
Six similar species (C. arenaria, C. ctibensis,
C. ekmanii, C. halophila, C. radiata, and C. sa-
graeana) make up this complex. All have com-
paratively narrow spikelets and reduced sterile
florets; relatively long awns are also a common
feature. The cpidennal pattern includes bicel-
lular microhairs and cuboidal silica cells. While
most of the species have inconspicuously ciliate
upper lemma margins, C cuhcnsis may approach
the C inflata complex in having more prominent
pubescence. Though Chloris cruciata is not in-
cluded here because of its somewhat unique
short cells, it is similar in overall spikelet struc-
ture.
The Chloris ciliata complex.
While this group presents more interspecific
taxonomic problems than any other group, it
is well defined as a whole. All species (Chloris
berroi, C. canterai, C. ciliata, C. dandijana, and
C lamproparia, are characterized bv having rel-
atively short-awned, elliptic fertile lemmas that
are prominently spreading-ciliate along most of
the margin. Epidermal patterns are typical of
Chloris. Chloris sesquiflora is similar, but the
marginal pubescence is strongly appressed.
The Clitoris verticillata complex.
Were it not for the extensive introgression
between Chloris andropogonoides, C. cucullata,
and C. verticillata, they would probably not be
included in a single complex. Moi-phologically,
there is considerable divergence, especially in C.
cucullata, in which the broad fertile and sterile
lemmas are distinctive in the genus. While in-
trogression involving the very rare C. texensis
has not been demonstrated, in spikelet and in-
florescence morphology it is obviousl)' similar to
C. andropogonoides.
In Chloris chloridea and C hrandegei there
is a strong tendency toward dorsal compression
of the spikelet. In other respects the plants are
similar to C. halophila, though considerably
more robust. The development of underground
spikelets in Chloris chloridea is unique in the
genus.
Four Australian species have bilobed fertile
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chlouis (Ghamineae)
15
and sterile lemmas. In Chloris pumilio and C.
lohata, this lobing is very prominent; the other
two, C. pectinata and C. cUvaricata, are less so.
The outward similaritv' to C. plurifhra and C.
crinita, both formerly in Trichloris, an American
genus, is probabK- the result of parallel evolu-
tion.
Another pair of Australian species, Chloris
triincata and C. ventricosa, is notable for having
ver\' truncate or ventricose fertile lemmas. Chlo-
ris triincata is strikingK similar to the North
American C. verticillata; again, the similarity
seems best explained by parallel evolution.
Chloris h>ngiaristata and C. prieurii, both
primariK- African in distribution, have several
sterile florets and relati\ely narrow, long-aw ned
lemmas. Tlie glandular strip along the side of
the lemma in C. prieurii is unusual in Chloris.
Clitoris hournei. C. (piincptcsetica, and C.
wightiana all have relatively large spikelets with
obtuse lemmas, coarsely pubescent lemma mar-
gins, and several sterile florets. Thev bear a
strong resemblance to species in Tetrapogon
but ha\e just a single ferHlc floret, whereas
Tetrapogon has more than one.
The above discussion leaves unmentioned a
rather large number of species which are quite
isolated moiphologicalh' ( Fig. 3 ) . Unquestion-
ably, these are best retained as a part of Chloris,
for thev have all of the features characteristic of
the genus; and yet, the elaboration of some in-
di\idual structural feature causes them to stand
alone. None is trulv widelv distributed, and
several are narrow endemics; this suggests adap-
tation to a particular climatic or edaphic regime.
Chloris Swartz
Chloris Swartz, Prodr. Veg. Ind. Occ. 25.
1788.
Rabclochloa Beauvois, Ess. Agrost. 84, 158,
176. 1812.
Phacellaria Willdenow ex Steudel, Nom.
Bot. ed. 2. 1:353. 1840. Herbarium name, given
as a svnonym.
Heterolepis Ehrenberg ex Boissier, Flora
Orientalis 5:554. 1884. Herbarium name, given
as a synonym of Chloris.
Trichloris Fournier, Mex. PI. 2:142. 1886.
Chloropsis Kuntze, Rev. Gen. PI. 2:771. 1891.
Leptochloris Munro ex Kuntze, Rev. Gen. PI.
2:771. 1891.
Pterochloris Camus, Bull. Mus. Hist. Nat.
(Paris). Ser. 2. 29:349. 1957.
Fibrous-rooted annuals or rhizomatous,
stoloniferous, or cespitose perennials, ranging in
^The inflorescence branches of the various species bear spikelets that range from virtually sessile to obviously pedicellate, and thus may
be called fpikes or racemes, depending upon emphasis. In this study the main inflorescence branch is arbitrarily called a spike.
size from only a few cm to over 2 m tall; epi-
dermal cells of root tips slightly different in
size, giving rise to root hairs which project
forward at about 60°; sheaths glabrous, scab-
rous, or villous; hgule often a ciliate crown,
sometimes absent; blades narrow to wide, flat
or rolled, deeply keeled and V-shaped, or with
rounded keels and with several furrows, scleren-
chyma girders complete or only on abaxial side,
bulliform cells generally large and penetraring
the mesophvll, bundle sheaths complete or in-
complete; costal silica cells usually cuboidal-
saddle- or axehead-shaped, very rarely some-
what cruciate, stomatal apparatus medium- to
triangular-dome shaped, bicellular microhairs
usually present, with relatively broad and short
terminal cells, or, less commonly, single-celled;
spikes' usually 5 to 25 (occasionally as few as
1 or 2, rareb' more than 25), frequentlv digi-
tateh arranged, occasionally in several verticils,
sometimes with a few isolated single spikes;
spikelets borne in two staggered rows on one
side of the rachis, sessile to pedicellate, com-
monly rather densely imbricate and appressed,
sometimes sparsely so; glumes usually unequal,
lanceolate, often glabrous with scabrous nerves,
mostly acute, rarely acuminate, shorter than
the florets; fertile lemmas one (very rarely
two), lanceolate, occasionally elliptic, three-
nerved, callus bearded, margins shortly to
prominently pubescent, intemerves usually
glabrous, sometimes scabrous, rarely pilose,
midnerv'cs scabrous or glabrous, rarely long-
pubescent, midnerve usually extended as an
awn, either from the tip or from betv\'een two
teeth, rarely only mucronate; palea shorter than
the fertile lemma, with two scabrous nerves,
otherwise glabrous; stamens three, anthers rela-
tively small; pistil one, with two stigmas; lodi-
cules two; sterile floret usually one (rarely two
or three, mostly barren, very rarely staminate
or perfect, varying widely in size and shape
from highly reduced to well developed and
similar to the fertile floret, rudimentary to
cylindrical to obovoid, acute to truncate, awned
or unawned; caryopsis ovoid, elliptic to obo-
void, round to trigonous in cross section, the
embryo relatively small, lower portions of
scutellum separate from the rest of the embryo,
epiblast present, vascular traces separated by
an elongated area of vascular tissue, embryonic
leaf margins usually not overlapping, very
rarely overlapping; starch grains of endosperm
compound; basic chromosome number x^9
(rarely), 10 (usually), the chromosomes rela-
tively small.
16
Brigham Young University Science Bulletin
Widely distributed in tropical, subtropical, spheres, both north and south of the equator
or warm temperate regions of both hcmi- (Fig. 4).
Fig. 4. Worldwide distribution of the genus Chloris. { This map is a composite of the distribution maps of all
of the species.)
Key to the species of Chloris
1. Inflorescence of many (30 or more) short spikes (6 cm or less) arranged along a cen-
tral axis which is at least 1 cm in length, the spikes not in verticils 2
Inflorescence of less than 30 spikes, the spikes arranged in one or more verticils 3
2. Sterile florets two or more; widely distributed in Africa, occasional in
India C. rox])ur<i}mina, p. 22
Sterile floret one; endemic to Cocos Island, off the west coast of Cen-
tral America C. paniculata, p. 24
3. Spikes two to four, forming a cylindrical, spikelike inflorescence; the spikes attached to
one another at least at the base and often for the full length, separable only with dif-
ficulty; Argentina, Uruguay C. herroi, p. 25
Spikes neither attached to one another nor forming a cylindrical, spikelike inflorescence 4
4. Spikes one to three 5
Spikes four or more 14
5. Sterile florets two or three 6
Sterile floret one (See also Chloris suringari, which has a highly reduced upper sterile
floret.) - .' 10
6. Fertile lemmas 1.8 to 2.8 mm long 7
Fertile lemmas 2.7 to 4.2 mm long 8
7. Inflorescence a single spike; lemmas awned from about the middle of the keel; Cuba,
Cura9ao) C. stirinp.ciri, p. 39
Inflorescence of more than one spike; lemmas awned subapically; southcentral United
States, Caribbean Islands, Mexico, Central and South America C. ciliata, p. 27
Biological Series, Vol. 19, No. 2 T.jv.xonomv of the Genus Chloris (Gr.^mineae) 17
8. Combined length of fertile- and sterile -floret awns 6.5 to 9 mm, usually more than
7 mm; annuals; Africa C. lamproparia, p. 30
Combined length of fertile- and sterile-floret awns 4 to 8 mm, usually less than 7
mm; perennials; South America, Texas 9
9. Plants densely tufted, with basal, narrow, rolled, curved leaves; spikes 3 to
5 cm long C. canterai var. grandiflora, p. 32
Plants not densely tufted; leaves cauline, up to 5 mm wide, not curled or curved; spikes
4 to 14 cm long C. canterai var. canterai, p. 30
10. Spikelets diverging from the rachis at nearly right angles; spikelets obviously
pedicellate; Culja, Haiti, Jamaica C. cruciata, p. 37
Spikelets appressed or cmly slightly divergent from the rachis; spikelets sessile or
nearly so 11
11. Awn of fertile lemma less than 1.5 mm long; internerves appressed-pubescent; Mau-
ritius C. filiformis, p. 39
Awn of fertile lemma more than 2 mm long; margins and keel spreading-pubescent; in-
ternerves glabrous or with inflated hairs 12
12. Awn of fertile lemma longer than 5 mm; Cuba, Jamaica, Hispaniola
_ C. ekmanii, p. 119
Awn of fertile lemma less than 4.5 mm long 13
13. Internc'i-ves glabrous; Cuba, Curasao C. surin^,ari, p. .39
Internerves densely covered with inflated hairs; Malagasy Republic
C. humbertiana, p. 104
14. Sterile florets two or more, the uppermost floret sometimes represented only by
the clavate rachilla joint 15
Sterile floret one 41
15. Sterile florets greatly modified, commonly o\er 3 mm wide, flabellate, margins scarious,
more or less inrolled and enclosing one another; Australia C. scariosa, p. 40
Sterile florets neither greatly modified, flabellate, nor involute, seldom more than 1.5
mm wide 16
16. Fertile lemmas with a row of glands along the sides; Africa, India —
C. prieurii. p. 42
Fertile lemmas without a row of glands, though sometimes appressed-pubescent
with nonglandular hairs 17
17. Fertile lemmas with a short mucro (less than 1 mm long); Africa C. pilosa, p. 58
Fertile lemma with an awn more than 1 mm long .- 18
18. Sterile floret pubescent, either with a prominently appressed-pubescent callus, or
densely pubescent on the midncrve, upper margins, or apex 19
Sterile floret glabrous or, at most, with a few ( 10 or so ) appressed hairs on the back 26
19. Callus of sterile floret bearded 20
Callus of sterile floret not bearded 21
20. Sterile floret unawned, acuminate; rachilla joint not obvious; culms stout, bam-
boolike; Africa C. rohusta, p. 43
Sterile floret obviously awned; rachilla joint prominent; culms not stout and bam-
boolike; New World C. mollis, p. 45
21. Leaf sheath, blade, and spike rachis densely pustulose-pilose; sides of fertile and sterile
lemma prominently and densely pustulose; India C. wip.htiana, p. 48
Leaf sheath, blade, and spike rachis glabrous , scabrous, or sparsely pilose, but never pus-
tulose; sides of lemma glabrous or occasionally appressed-pubescent, but not pustulose.. 22
18 Bricham Young University Science Bulletin
22. Culms densely appressed-pubescent for several mm just below the points of attach-
ment of the spikes 23
Culms glabrous or scabrous below the points of attachment of the spikes 24
23. Upper margins of lowermost sterile floret ciliate; sterile florets usually four ( occasion-
ally 3); uppermost sterile floret spherical and inflated; India ...C. quinqtiesetica, p. 4S
Upper margins of lowermost sterile floret usually not ciliate, occasionally sparsely pubes-
cent; sterile florets usually three (rarely two or four); upper sterile floret resembling
the lower, neither spherical nor inflated; Ceylon, India, Thailand C. montana, p. 51
24. Awn of fertile lemma 2 mm long or less; Burma C. btinnensis, p. 125
Awn of fertile lemma more than 3.5 mm long 25
25. Midnerve of lowest sterile floret densely and prominently appressed-pubescent; fer-
tile lemma longer than 3 mm; lowest sterile floret 1.5 mm or longer, its awn usually
shorter than 4 mm; stoloniferous perennial; southern India C. hournei. p. 53
Midnerve of lowest sterile floret sparsely appressed-pubescent; fertile lemma less than 3
mm long; lowest sterile floret less than 1.5 mm long, its awn over 5 mm; annual or
weak perennial, sometimes rooting at the lower nodes; pantropical C. infhita. p. 53
26. Awn of fertile lemma longer than 30 mm; Africa C. longiaristata, p. 46
Awn of fertile lemma less than 15 mm long 27
27. Sterile florets two or three, only the lowermost awned, the others awnless 28
Sterile florets two or three; all are awned 34
28. Fertile lemmas prominently ciliate along most of the margin, the cilia spreading
at nearly right angles; lemmas broadly elliptic 29
Fertile lemma margins glabrous or with .strongly spreading hairs near the apex,
but never along tlie lower margins 33
29. Fertile lemmas 1.5 to 2.8 mm long 30
Fertile lemmas 2.9 to 4.2 mm long 31
30. Spikes three to five (rarely six or seven), usually less than 8 cm long; Texas,
Mexico, Central America, southern Brazil, Argentina C. ciliata, p. 27
Spikes 7 to 40 (usually 10 to 20), usually more than 8 cm long, occasionally as
short as 5 cm; Florida, West Indies, South America C. dandijanci, p. .34
31. Spikes one to four (often two); combined length of awns of fertile and sterile floret 6.5
to 9 mm (usually about 7 mm); annuals; Afric;i C. himpwparia, p. .30
Spikes two to nine (usually three or more); combined length of awns of fertile and ster-
ile floret 4 to 8 mm (usually less than 7 mm); perennials; South America, Texas .32
32. Plants densely tufted with basal, narrow, rolled, curved leaves; spikes 3 to 5 cm
long C. canterai \ar. grandijUmi, p. 32
Plants not densely tufted; leaves cauline, up to 5 mm wide, not curled; spikes 4
to 14 cm long C. canterai var. canterai. p. .30
33. Annual plants arising from a fibrous root system, occasionally rooting at the lower nodes
if culms are decumbent; Africa C. pilosa, p. 58
Perennial plants, usually with prominent, aggressive stolons, occasionally tufted; widely
distributed in the warmer parts of the world C. gaijana, p. 65
(NOTE: Occasional specimens of C/iZoris uenfn'ro.sr; having two sterile florets will -
key to this point. )
34. Awn of fertile lemma less than 2 mm long; Burma C. burmensis, p. 125
Awn of fertile lemma more than 3.5 mm long 35
Biological Series, Vol. 19, No. 2 T.^.xonomv or the Genus Ciii.oris (Cramineae) 19
35. Culms densely appressed-pubesceiit for several mm just below the points of attach-
ment of the spikes; Ceylon, India, Thailand C. montana, p. 51
Culms scabrous or glabrous just below the points of attachment of the spikes 36
36. Upper sterile floret inflated, nearh spherical; widespread in wanri re-
gions C. inflata, p. 53
Upper sterile floret cylindrical or very narrowly turbinate 37
37. Rachis pilose; eastern China, Hong Kong, North Vietnam, Taiwan ...C. formosana, p. 57
Rachis scabrous 38
38. Fertile lemma with three awns, the lateral about equal to the central, or shorter.... 39
Fertile lemma with only a single awn, lateral awns absent 40
39. Awns subequal, the lateral only slightly shorter than the central C. crinita, p. 87
Awns unequal, the lateral about one-tenth the length of the central ..C. pluriflora. p. 89
40. Fertile lemmas acute at the ape.x; margins not inrolled, densely ciliatc, especially
on the upper margins; spikelets tawny: robust, stoloniferous perennials more
than 50 cm tall; widespread in warm regions C. gmjana, p. 65
(NOTE: On rare occasions, specimens of C/i/o;/.s HKWsa7ni»!censis, from southern
Africa, and C. sa'^raeana, from tlie Caribbean region, both rarely haying two
sterile florets, will key to this point. )
Fertile lemmas obtuse to rounded at the apex; margins inrolled and appressed-
pubescent; spikelets dark brown to black at maturity; relatively short, tufted
perennials; Australia C. truncata. p. 7S
41. Sterile lemma bilobed for upper one-third to nearly all of its length, the lateral lobes
usually awned 42
Sterile lemma not prominently bilobed 47
42. Fertile lemma margins pubescent to ciliate on upper half, the hairs usually prom-
inent and spreading, especially toward the apex 43
Fertile lemma margins glabrous, scabrous, or strongly appressed-pubescent, but
upper margins without prominent spreading hairs 44
43. Fertile lemma margins pubescent mostly on upper half; sterile floret bilobed nearly to
the base, the lobes usually widely divergent at maturity; sterile lemma lobes gradu-
ally tapering into an awn; fertile lemma awns subeciual; Australia C lohata, p. 82
Fertile lemma margins pubescent along most of length (the hairs spreading apically,
appressed basally); sterile floret bilobed to about the middle, very occasionally to the
base, the lobes not widely di\ergent at maturity; sterile lemma lobes usually trun-
cate to somewhat rounded, the awn arising laterally; median awn of fertile lemma
usually about twice the length of the lateral awn C. piimUio, p. 80
44. Lateral and central awns of fertile lem ma subequal to equal; southwestern United
States, Mexico, and South America C. crinita, p. 87
Lateral awns of fertile lemma about one-tenth the length of the central awns 45
45. Spikelets densely pectinate, crowded, diverging at wide angles from the rachis; Aus-
tralia C. pectinata, p. 86
Spikelets not densely pectinate, only slightly imbricate, appressed to the rachis 46
46. Spikes in two or more verticils; South America C. halophila, p. 117
Spikes in a single radiate series; Australia C. divaricata, p. 83
47. Fertile lemma awnless or with a short awn, generally less than 2 mm long 48
Fertile lemma prominently awned, the awn usually more than 3 mm long 52
20 Bricham Young University Science Bulletin
48. Sterile floret very narrowly cylindrical, usually about 0.2 mm wide, usually
sparsely pilose near the apex; Baja California, Mexico C hrandegeu p. 72
Sterile floret broader, usually more than 0.3 mm wide; at most merely scabrous
at the apex 49
49. Sterile floret inflated-spherical, about 1 mm wide, usually as wide as long; Texas, New
Mexico, Oklahoma, northeast Mexico C. cticullata, p. 91
Sterile floret not inflated-spherical, less than 5 mm wide, usually at least twice as long
as broad 50
50. Fertile lemma margin glabrous, scabrous, or appresscd-pilose for no more than
one-half of its length; keel usually glabrous, occasionally scabrous near the
apex, rarely sparsely appressed-pubescent; Mexico, Colombia
C. sul>imitica, p. 106
Fertile lemma margin and keel appressed-pubescent for most of length 51
51. Sterile floret elliptical, flattened, about 1 mm long; fertile lemma prominently spread-
ing-pilose on margins and keel; South America C. sesquiflora, p. 104
Sterile floret cylindrical to naiTowlv turbinate, inflated, ca 2 mm long; fertile lemma
margins and keel appressed-pubescent; Africa C. ametht/stca, p. 106
52. Fertile lemma broadly elliptic, prominently long-ciliate along most of the mar-
gin, the cilia spreading at nearly right angles 53
Fertile lemma margins glabrous or with appressed to widely spreading hairs near
the apex, but never along the lower margins 54
53. Fertile lemma more than 3.5 mm long, gibbous; awn definitely subapical; Ar-
gentina C. castilloniana. p. 69
Fertile lemma less than 3 mm long, elliptic, not gibbous; arising only slightly subapi-
cally; Florida, Bahama Islands, Jamaica, South America . C. chmdyana, p. .34
54. Plants producing cleistogamous underground spikelets at the ends of thin rhi-
zomes; southwestern United States, Mexico, Central America, and V'cne-
zuela C. chloridea, p. 74
Plants not producing cleistogamous underground spikelets 55
55. Spikes naked 2 to 5 cm from the liase; endemic to eastern Texas; rare, possi-
bly extinct C. texemis, p. 103
Spikes floriferous to near the base 56
56. Spikes borne in two or more verticils, these usually well separated. (See also C.
virgata which may have two verticils, closely inserted.) 57
Spikes borne in a single terminal whorl 61
57. Sterile floret greatly reduced, usually less than 1 mm long; annuals; widely distributed
in American tropics; Hawaii C. racliata, p. 110
Sterile floret longer than 1 mm, well developed; perennials, tufted or stoloniferous 58
58. Upper margins of fertile lemma with a prominent tuft of spreading white hairs,
these usually longer than 1 mm; side of fertile lemma grooved, the groove
usually with many glandular hairs ( though these may be somewhat obscure or,
rarely, absent); Brazil Ascension Island C. orthonoton, p. 69
Upper margin of fertile lemma scabrous or appressed short-pilose; side of fertile
lemma without a groove or glandular hairs 59
59. Fertile lemma 2.0 to 3.5 mm long; widely distributed in central and southcentral United
States C. verticiUuta, p. 93
(NOTE: Various introgressants of Chloris verticillata, C. cticuUata, and C. ondropof^o-
noides may key to this point. See full discussion of this problem in the text.)
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 21
Fertile lemma longer than 3.8 mm 60
60. A\\'n of fertile lemma longer than 9 mm long; South America C. halophila, p. 117
Awn of fertile lemma shorter than 5 mm long; Baja California, Mex-
ico C. hrandeiiei, p. 72
61. Fertile lemma with a dense tuft of spreading hairs on the upper margins, the longest
usually more than 1.5 mm long (see also C. aristata and C. cubensis, which occasion-
ally have rather long hairs near the fertile lemma apex) ^ 62
Fertile lemma without a dense tuft of long spreading hairs on the upper margins,
though often with shorter (generall)' less than 1 mm), appressed hairs, occasionally
scabrous or glabrous 64
62. Fertile lemma with a shallow lateral groove 63
Fertile lemma without a lateral groove; southern Africa C. mossamhicensis, p. 123
63. Annual plants; fertile lemma usually prominently carinate, gibbous; lateral groove
of fertile lemma glabrous or appressed-pilose, not glandular; widely distribu-
ted C. virgata, p. 60
Perennial, stoloniferous plants; fertile lemm.x rounded on the back, neither carinate nor
gibbous; lateral groove of fertile lemma usually with many glandular hairs
(though these are sometimes obscure, or, rarely, absent); Brazil, Ascension
Island C. orthonoton, p. 69
64. Spikelets bright green to olive green; Ethiopia, Somali Republic
-. C. soinalensis, p. 109
Spikelets pale brown, tawny to black, but not greenish, at least at maturity 65
65. Fertile lemma usually less than 2 mm long; Phillippini' Islands C. dementis, p. 109
Fertile lemma more than 2 mm long 66
66. Spikes naked 2 to 5 cm from the base; endemic to eastern Texas; rare, possibly
extinct C. texensis, p. 103
Spikes floriferous to near the base 67
67. Sterile floret more than 0.6 mm wide, often nearly 1 mm wide 68
Sterile floret less than 0.6 mm wide 70
68. Upper fertile lemma margins densely ciliate or spreading-pilose, the margins not
inroUed; Central America . C. aristata, p. 70
Upper fertile lemma margins glabrous, scabrous, or appressed-pilose, the margins
inrolled, especially below 69
69. Margins of fertile lemma glabrous or sparsely pilose; fertile lemma apex generally acute,
occasionally somewhat rounded; fertile lemma tawny to purple tinged, seldom dark
brown or black; Australia C. ventricosa. p. 76
Margins of fertile lemma appressed-pilose; fertile lemma apex broadly obtuse
to rounded; fertile lemma dark bro\vn to black at maturity. Australia
C. truncata, p. 78
70. Awn of fertile floret usually less than 5.5 mm long; southern Texas, northeastern
Mexico C. andropogonoides, p. 95
Awn of fertile floret usually more than 6 mm long 71
71. Sterile floret less than 1 mm long 72
Sterile floret more than 1 mm long ; 74
72. Culms slender, wiry; blades filiform or narrow, less than 1 mm wide (frequently
less than 0.5 mm wide); Cuba, Jamaica, Hispaniola C. ekmanii, p. 119
Culms stout, not wiry; blades (at least the wider) 5 to 10 mm wide 73
22 Bricham Young University Science Bulletin
73. Awn of fertile lemma 6 to 13 mm long; fertile lemma margins sparsely pilose;
leaf blade apex acute to subacute; widely distributed in American tropics;
Hawaii C. racliata, p. 110
Awn of fertile lemma 10 to 45 mm long; fertile lemma margin scabrous, rarely sparsely
pilose; leaf blade apex obtuse, rarely acute; South America, Africa
C. pipiothrix, p. 114
74. Culms slender, wiry; blades filiform or very narrow, usually less than 1 mm wide,
frequently less than 0.5 mm wide; Cuba, Hispaniola, Jamaica
C. ekmanii, p. 119
Culms stout, not wiry; blades, at least the larger, generally broader than 1.5 mm .-. 75
75. Blades usually densely pubescent on both upper and lower surfaces; awn of fertile lemma
20 to 24 mm long; sterile floret narrowly cylindrical, 1.6 to 1.9 mm long, 0.2 to 0.3
mm wide; Cuba C. arenaria, p. 119
Blades usually glabrous or scabrous, occasionally sparsely pilose near the base; awn of
fertile lemma less than 13 mm long; sterile floret comparatively broader, 0.3 to 0.6
mm wide, 0.7 to 1.9 mm long 76
76. Spikes strongly divergent from the vertical axis, becoming horizontally spreading
or even somewhat reflexed at maturity; keel of fertile lemma glabrous, occa-
sionally very sparsely appressed-pubescent; longest callus hairs 0.3 to 0.8 mm
long; marginal hairs of fertile lemma strongly appressed, shorter than 0.5 mm
long; Caribbean Islands, Cuba, Haiti, Jamaica C. sagracaiui, p. 121
Spikes not divergent more than 45° from the vertical axis; keel of fertile lemma
usually prominently appressed-pubescent, occasionally sparsely so, rarely gla-
brous; longest hairs of the callus I.O to 1.2 mm long; marginal hairs of the lemma
somewhat spreading, especially toward the apex, the longest usually 1 mm or
longer; Caribbean Islands, Cuba, Haiti, Jamaica .C. cuhemis, p. 123
I. CHLORIS ROXBURGHIANA Schultes, spikelets densely imbricate, ca 17 per cm of the
Mantissa 2:3.39. 1824. (Based on Chloris rachis length; glumes narrowly lanceolate, gla-
pohjstachija Roxburgh. See explanatory note brous except for the slightly scabrous midrib;
in text.) Fig. 5, A-E. first glume 0.9 to 1.0 mm long, ca 0.1 mm wide;
^, , . , , _, , , -- second glume 1.9 to 2.2 mm long, ca 0.2 mm
Chlons pohjstochya Roxburgh Hortus ^^j^^. ^^^j,^. ,^^^,^^^^ ^^ ^^ 2.I mm long, ca 0.4
Bengalensis 82. 1814. Nomen nudum. Non ^^^ ^^.^^ ^^^^^^^ ^^ ^^^.^^.^ ^,^^„^^ ,,^,.^^^f^^, ^^,^^_
Lagasca, IHlb. ^^^^ glabrous to sparsely ciliate, especially
Chloris pohjstacluja Roxburgh, Flora In- above; sides sparsely appressed-pilose to gla-
dica 1:332. 1820. Non Lagasca, 1816. (See b^ous, apex acute, awn 10 to 15 mm long; sterile
explanatory note in text. ) florets usually two (occasionally three) gla-
Chloris myriostachija Hochstetter, Flora brous, shaped much like the fertile lemma, lower
38:204. 1855. sterile floret 0.7 to 1.0 mm long, 0.2 to 0.3 mm
(HOLOTYPE: "Hb. abyss. Buch. nr. 1416," wide, upper floret(s) progressively smaller, awn
not seen, but description is of this taxon. ) of lower sterile floret ca 10 mm long; caryopsis
Chloris myriostachija viu. minor Chiovenda ca 1.1 mm long, 0.4 mm wide, ellipsoidal, trigo-
in Pirotta, Annuario Reale 1st. Bot. Roma nous; chromosome number 2n = 20.
8:54. 1903. (Description is of a small var- Only C/i/on* paniculafa \ms as large a num-
iant of the species ) ^^^ °^ spikes as C. roxburghiana. These two are
most easily differentiated by the number of
Perennial to 125 cm tall, arising from a sterile florets, C. roxburghiana always having at
strongly stoloniferous, woody base; sheaths least two, while C. paniculata always has one.
glabrous below, often pilose above; ligules Chh)ris roxburghiana is a stoloniferous perennial
pilose; blades up to 30 cm long, 5 to 6 mm found in Africa and India. Chloris paniculata
wide, glabrous or scabrous; spikes many (usu- has a unique growth habit, with well-developed,
ally 50 or more) attached along a short axis at short, vertical rhizomes bearing the frayed rem-
the culm apex and fonning a dense inflores- nants of previous years' growth as well as many
eence up to 15 cm long and 4 to 6 cm wide; long, arching, basal leaves; it is endemic to
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
23
Fig. 5. C/iforis roxhurghiana and C. paniculata. (A-E) C. roxburghiana. (A) habit, x 1/5; (B) spikelet, partly
dissected, x 15; (C,D) lower and middle sterile florets, respectively, x 10; (E) caryopsis, x 10. (F,G)
C. paniculata. (F) habit, x 1/5; (G) spikelet, partly dissected, x 15.
24
Bricham Young University Science Bulletin
Cocos Island, which is off the western coast
of Panama.
Chloris mijriostachya has been used as a
name for this species for many years, especially
in African floras. Part of this erroneous usage
can be traced to the difficulty of typifying the
Roxburgh names. William Roxburgh first used
C. pohjstachija in 1814 (Hortus Bengalensis
82.), publishing it without a description. Iden-
tification of this name rests with its subsequent
publication by Roxburgh (Flora Indica 1:332.
1820.), who included a fairly complete descrip-
tion. By that date, however, Lagasca y Segura
(1816) had published the same name for a
Mexican species (sec under C. suhimiticu).
Shortly afterward, Schultes (Mantissa 2:. 3.39.
1824.) published a new name, C. roxhurghkina.
based upon C pohjstaclu/a Roxburgh. No plant
specimen that mav be connected positively with
Roxburgh's name can be located; however, im-
portant clues may be obtained from the Rox-
burgh drawings at Kew and from the descrip-
tions given b>' Roxburgh, both those of the
published Flora Indica as well as the copy of
the manuscript of that work in the Kew Li-
brary. The drawings, labelled "C. pohjstachija,"
show an inflorescence of sixteen spikes arranged
along a short common axis, and a series of dis-
sected spikelets showing broadly ovate-elliptic
fertile lemmas and two similar sterile florets.
While the inflorescence drawing does not
portray accurately the more t\pieal specimens
of this taxon from Africa ( though Indian col-
lections bearing a strong resemblance are com-
mon) the spikelet drawings leave no question
as to the identity of Roxburgh's Chloris pohj-
stachija and thus, indirectly, C. roxhurghiana
Schultes.
Chloris roxburghiana is common (sometimes
dominant) in grasslands, brush, and disturbed
areas, often on sandy loams from low to me-
dium elevations (Fig. 6). Associates include
Cenchrus sp., Pennisettim sp., Sporobohis sp..
Acacia mellifera, Blepharis sp.. Commiphora sp.,
and Euphorbia spinescetu.
Representative specimens examined: AN-
GOLA: Mossamedes Dtr., Camucuio, Azancot
de Menezes 363 (K). BOTSWANA: Maklautsi-
Shashi Rivers, Palapye, de Beer, s.n., 9 May
1957 (K). ETHIOPIA: Harrar Prov., 40° 39' E,
10° 10' N, Burger 2,899 (K); Sidamo Prov., 20
mi N of Moyale (Kenya), Moonei/ 7,421 (K).
INDIA: Nilgiri Dtr., ' Kitagin Ghat, Fischer
2,086 (K). KENYA: Kibwezi Plains, Balhj 1,329,
8,090 (K); 25 mi SSW of Kitui, Bogdan AB
5,126 (K,US); Lugard's Falls Road, Tsavo Na-
tional Park, Greenwatj and Kanuri 12,636 (K);
Fig. 6. Distribution of Cliloris mxl>urnhiana. Inset
A: India.
Wei Wei 50 mi N of Kapenguria, Trelawnij AB
4,325 (K,UC). MALAGASY REPUBLIC: Mom-
bas, anon. (K). MALAWI: Lilongwe Dtr., Salu-
beni 428 (K). MOZAMBIQUE: Louren^o
Marques Dtr., between Moambas and Sabie,
Torre 2,237 (K). REPUBLIC OF THE
CONGO: Plaine du Lac Edouard, Pare Nation-
ale Albert, Louis 4,787 (K, US). SOMALI RE-
PUBLIC: Boundaiy Pillar 93, 8°37'N, 45°9'E,
Gillett 4.197 (K, US). SOUTHERN RHODE-
SIA: 30-60 mi S of Ft. Victoria, Rodin 4,251 (K,
UC, US); Sabi River Valley, Melsetter, Whellan
1,021 (K). SUDAN: Equatoria Prov., Kidepo
resthouse, Mijers 11,236 (K). TANZANIA:
Tanga Dtr., Kange Estate, Faulkner 832 (K);
SW of Umba River, Kivingo, Greenuaij 1,996
(BM, K); Lake Manyara National Park, Green-
wai/ and Kirrika 11,089 (K, US). UGANDA:
K;inamugit, Eggeling E 2.946 ( K, US ) ; Turkana
Dtr., Oropoi Valley, Liebenberg 1,776 (K).
UNION OF SOUTH AFRICA: Kruger Natiimal
Park, between Punda Maria and Pafuri (Mo-
zamljique), Godfrey SH-1.729 ( K, US); be-
tween Beauty and Ellisras, Werdemann and
Oberdicck 1,801 (K).
2. CHLORIS PANICULATA Scribncr in Rob-
inson, Proc. Amer. Acad. Arts Sci. 38:262.
1902. (HOLOTYPE: "Nuez Isl . . . an islet
near Cocos Isl.: Snodgrass and Heller, no.
968;" in GH. Not seen, but description is
clear.) Fig. 5, F and G.
Perennial to 70 cm tall, arising from an
upright, stout, underground stem bearing many
rootlets and shreds of leaf sheaths; sheaths
Biological Seiiies, Vol. 19, No. 2 Taxonomy of the Genus Chlohis (Gramineae)
25
glabrous; ligule lacking; blades very long and
narrow, up to 50 cm long and 5 mm wide, arch-
ing, glabrous except for the scabrous margins;
inflorescence paniclelike, made up of at least
50 spikes racemosely arranged on the upper 10
to 12 cm of the culm; spikes about 5 cm long
at lower part of inflorescence, becoming pro-
gressively shorter near the tip; glumes narrow
to broadly lanceolate, glabrous except for the
scabrous midnerve; first glume ca 1.2 mm long,
0.2 mm wide; second glume ca .3 mm long, 0.4
mm wide; fertile lemma ca 2.6 mm long, 0.6
mm wide, broadly lanceolate, glabrous except
for the prominently bearded callus and scab-
rous keel, apex ± obtuse, awn 2.5 to 2.8 mm
long; sterile floret one, ca 1.5 mm long, 0.3 mm
wide, glabrous, awn 1.5 to 1.8 mm long.
Only Chloris roxhurghiana has as many
spikes as C. panicithta. Differences between
these two species have been discussed under C
roxlmrghiana.
Chloris paniculata is an endemic known
only from Cocos Island and its associated islets
—an archipelago lying about 300 miles south-
west of Costa Rica, to whom it belongs. Rob-
inson (1902) and Stewart (1912) both reported
it as being abundant on rocky cliffs near the
coast, both on the main island and on the islets.
Most of the vegetation of the island area is
made up of common Central American species
or pantropical weeds. Only eight endemics in
a total of about a hundred species were re-
ported by Stewart. Fosberg and Klawe (1966)
also report C. paniculata from Cocos Island.
I cannot suggest a species in Chloris. nor
for that matter anv other genus in the Chlori-
deae, that might be said to be closely related
to C paniculata. The previously discussed re-
semblance to C. roxhurghiana is superficial, for
there are manv spikelet details in which they
differ. Certain species of Leptochloa have a
similar aspect because of the large number of
spikes. Leptochloa, however, has spikelets which
unifonnh' have a larger number of fertile flor-
ets, and the sterile floret is not as elaborated in
structure and size as is generally the case in
Chloris.
Specimens examined: COCOS ISLAND: ex-
posed rocky cliffs near the ocean, Stewart 260
(F,MO,NY,US).
3. CHLORIS BERROI Arechavaleta, Anales
Mus. Nac. Montevideo 5:388. 1896.
(HOLOTYPE: "Berro, Uruguay, Estancia
de Soriano, Estancia de Vera." This speci-
men was not seen, though specimens at K!
and US! are labeled essentiallv the same
and are the species as commonlv under-
stood. The description and illustration ac-
companying the original description leave
no (juestion as to the identity of the taxon,
however. ) Fig. 7, A-D.
Chloris accumbens Hackel ex Arechavaleta,
Anales Mus. Nac. Montevideo 5:391. 1896.
Nomen nudum, pro sijn. C. berroi.
Perennial from a fibrous root system, tufted,
15 to 80 cm tall; sheaths glabrous; ligule ciliate;
blades narrow, 3 to 15 cm long, 1.5 to 2.0 mm
wide, glabrous or sparsely pilose near the base;
spikes two to four, 3 to 12 cm long, tightly
appressed, adherent, and forming a narrow,
cylindrical, spikelike inflorescence; spikelets
densely imbricate, 9 to 12 per cm of the scab-
rous-hispid rachis; glumes glabrous, lanceolate;
first ghnne 1.5 to 1.6 mm long, ca 0.3 mm wide;
second glume 2.1 to 2.6 mm long, 0.3 to 0.6 mm
wide; fertile lemma ovate, 2.7 to 3.5 mm long,
0.8 to 1.2 mm wide, margins and keel ciliate
with hairs up to 2 mm long, awn 2.7 to 3.4 mm
long; sterile florets usually three, up to 1.9 mm
long, lowennost awned, upper unawned, gla-
brous; caryopsis 1.2 to 1.8 mm long, 0.5 to 0.7
mm wide, obovoid, trigonous; chromosome
number 2n^40.
Chloris berroi can be distinguished from
all other species of the genus by its cylindrical
inflorescence; this results from the interweaving
of the spikelets on adjacent spikes and, more
especially, from the hispid hairs of the rachises.
The spikes remain interconnected past maturity,
except for the very tips, and may be separated
from one another only by actually tearing them
apart.
Individual spikelets of Chloris berroi are
very similar to the other species in which the
lemnias have ciliate margins, especially those of
C. ciliata and C. clanchjana. Other than the
inflorescrnce differences previously mentioned,
there seem to be no other characteristics sepa-
rating them.
Chloris berroi is restricted to the Rio de la
Plata region of Argentina and Uruguay (Fig.
8), where Parodi ( 1919) reported it to be abun-
dant in the campo.
Specimens examined: ARGENTINA: Prov.
Buenos Aires; La Plata, Spegarrini 1,433 (NY);
Pergamino, Niedfeld 38 (US). Prov. Entre Rios:
camino de Puerto Constanza a Gualeguaychu,
Burkart 10,529 (US); Dep. Federacion: Estan-
cia "Buena Esperanza," Pedersen 6,254 (US).
URUGUAY: Dep. Canelones: Montevideo,
Sello s.n. (MO). Dep. Florida: Arroyo Mansa-
viUagra, Gallinal 5,790 (MO). Dep. Soriano,
26
Bricham Young UNivERSiTi' Science Bulletin
Fig. 7, Chloris berroi and C. ciliata. (A-D) C. benoi. (A) h;ibit.. x 1/5; (B) spikelet, x 10; (C) lower sterile
floret, X 15; (D) upper sterile floret, x 15. (E-I) C. ciliata. (E) habit, x 1/4; (F) spikelet, x 10; (G)
sterile florets, x 15; (H) upper sterile floret, x 15; (I) caryopsis, x 10.
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gbamineae)
27
Fig. 8. DistribuKon of Chloric herroi (southern South
America ) .
Berw 6,385 (US). Dtp. Tacuarembo, Herter
587a (MO).
4. CHLORIS CILIATA Swartz, Prodr. Veg.
Ind. Occ. 25. 17S8. (HOLOTYPE: Not
seen. Swartz's original description in 1788 is
clear; his amplification in 1797 leaves no
doubt.) Fig. 7, E-I.
Anchopoiion ptihescens Aiton, Hortns Ke-
wensis 3:423. 1789. (Based on Chloris cil-
iata Swartz.)
Ctinodon ciliatus (Swartz) Raspail, Ann.
Sci. Nat. Bot. (Paris) 5:303. 1825. (Based
on Chloris ciliata Swartz. )
Chloris propinqua Steudel, Syn. PI. Glum.
1:204. 1854. (HOLOTYPE: "Duchai.ssing
legit in Guadeloupe." A fragment in US!,
from the Steudel specimen in P, is badly
shattered but would appear to be Chloris
ciliata. The description seems to fit Chloris
ciliata Swartz.)
Chloris ciliata var. texana Vasev, U. S. Dept.
Agr. Div. Bot. Bull. 121. pi. 30. 1890.
(HOLOTYPE: "Near Brownsville" US!)
Chloris texana (Vasey) Nash, Bull. Torrev
Bot. Club 25:441. 1898. Based on Chloris
ciliata var. texana Vasey.)
Chloris nashii Heller, Muhlenbergia 5:120.
1909. (Based on Chloris ciliata var. texana
Vasey. )
Perennial 25 to 60 cm tall, tufted, erect;
sheaths glabrous, hgule absent or reduced to
a short ciliate crown (Northern Hemisphere) or
densely and conspicuously pilose (Southern
Hemisphere); blades 10 to 20 cm long, ca 5
mm wide, long-acuminate, glabrous or scabrous;
spikes three to five (rarely si.x or seven), 3.5
to 6.0 cm long (occasionally up to 8 cm) some-
what flexuous and spreading, glumes narrowly
lanceolate, glabrous except for the scabrous
midrib, becoming hyaline near the margins;
first glume 1.3 to 1.7 mm long, 0.2 to 0.4 mm
wide; second glume 2.0 to 2.5 mm long, 0.2 to
0.4 mm wide; fertile lemma 1.8 to 2.8 mm long,
0.8 to 1.1 mm wide, strongly flattened, elliptic,
margins and keel strongly ciliate, the cilia 0.5
to 1.5 mm long, awn 0.9 to 2.7 mm long; sterile
florets two, lowermost enclosing the upper;
lower sterile floret 1.3 to 1.8 mm long, 0.8 to
1.8 mm wide, truncate, glabrous, awn 0.9 to 1.4
mm long, upper floret similar but smaller, 0.8 to
1.1 mm long, 0.9 to 1.2 mm wide, membranous,
awnless; caryopsis ca 1.4 mm long, ca 0.7 mm
wide, obovoid to ellipsoid; chromosome number
2n=40.
The species in this complex (Chloris herroi,
C. canterai, C. ciliata, C. dandijana, and C.
lamproparia) are characterized by elliptic to
lanceolate lemmas, the margins of which are
densely ciliate, usually along the entire length.
Chloris herroi is easily distinguished, for it has
spikes so interconnected by hispid hairs on the
rachises that it appears to have but a single
spike. The remaining species, however, present
a more complex pattern. While most collections
will fit a given species quite well, certain geo-
graphic areas (most often in South America)
may have individuals which are somewhat inter-
mediate with respect to the characteristics. Fig-
ure 9 illustrates selected features of a large
number of specimens chosen at random from
throughout the ranges of the species.
While this diagram reveals no distinct sub-
populations, certain variation trends may be
discerned. One trend is characterized by indi-
viduals with relatively short lemmas, short
marginal cilia on the lemmas, short awns,
and' few spikes (six or less). These are the
plants commonly called C. ciliata and are repre-
sented bv the open circles situated roughly in
the lower left quadrant of the diagram. A sec-
ond trend, represented mostly by open circles
with an upward line, is shown in the upper
left quadrant. These collections had longer
28
Bricham Young University Science Bulletin
CO
<
n:
<
LlI
en
<
Ll
O
n
I—
CD
z
UJ
3.5-
3.0-
2.5-
2.0
1.5
1.0-
0.5-
46 A
6 4
44 4 4
44 44 4 4 o
4 o 4
44 44444 po 44 oo i
444444 A ^
o4 444 44 A ooo
A A 4444 44 4 o o
A 44444 44
4 4 4 44AA44o 4
4 4
4 44o44 44 44
44o4444 4 o o
4 44 4 o
44 44oA4 4 o o o o
4 o
4 o o o oo
o oooooo o ooo
o ooo oo o o
o oooo ooo o oo o o
o ooo o oo
o o ooo
o oooo
t
9 f
t
O NEW WORLD
• OLD WORLD
6 4 SPIKES 7 OR MORE
O • SPIKES 6 OR LESS
COMBINED LENGTHS OF AWNS OF
STERILE AND FERTILE FLORETS:
O • MORE THAN 7.5 MM
O •LESS THAN 7.4 MM
1.5
2.0
2.5 3.0 3.5 4.0
LEMMA LENGTH IN MM.
4.5
Fig. 9. Comparison of various morphological and geographical characteristics of Clitoris canterai var. canterai
and var. grandiflora, C. ciliata, C. dandyana, and C. lamproparia.
lemmas and marginal hairs, a larger number of
spikes (often more than 20), but short awns;
these are representative of plants commonly
called C. damhjana. Extremes of these popula-
tions are easily separated, though individual
plants may diverge considerably from the
"average" member of each trend.
The third and fourth trends overlap more
with respect to the morphological characteris-
tics analyzed and are most easily separated
geographically. These two groups are inter-
mingled in the upper right quadrant and differ
from both of the pre\ious trends in length of
the lemma and of its marginal cilia. Within this
group, the New World collections are perennials
with relatively short lemmas, marginal cilia, and
awns. These are the plants commonlv named
C. canterai. The remaining specimens are Old
World annuals and have, generally, longer
lemmas, marginal cilia, and awns; these are
commonly called C lamproparia.
Well-defined discontinuities between the
various populations have not been demon-
strated, at least on the basis of the characteris-
tics employed. However, it does not seem that
merging all of these into a single species is
justified, for discernible variation trends are
evident. Thus four species are recognized, each
centering in one of the four trends discussed
above.
Certain other characteristics were of value
in differentiating elements within these taxa, but
did not clarify the relationships between them;
these are discussed individually under the ap-
propriate species. All populations are deserving
of further study, especially in areas of sympatry.
Mass collections as well as cytological exam-
ination and genetic analysis ^\â– ill be especially
valuable.
Tabic 7 compares all of the species in this
complex witli respect to a variety of characters.
Figures 7, 11, 1.3, and 15 should also be used
for comparison.
There aie two recognizable geographic vari-
ants within Chloris ciliata: (1) a North Ameri-
can population with nonciliate ligules and (2)
a South American group with prominently cil-
iate ligules. No other differences could be
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (CfRamineae)
29
Table 7. Comparison of Chloris berroi, C. canterai, C. ciliata, C. dandyana, and C. lamproparia.
Annual
or
Species
perennial
C. berroi
C. canterai
perennial
perennial
C. ciliata
perennial
C. dandyana
perennial
Fertile
lemma
Spike
Culm
type
Number
Length
( in cm )
Length
( in mm)
Marginal
hair
length
( in mm )
Awn
length
( in mm )
C lamproparia annual
erect, 15-80 cm tall
erect, to 1 in tall (ex-
cept shorter and ces-
pitose in var grandi-
flora)
erect, tufted, 25-60
cm tall
erect, usually tufted,
rarely stoloniferous,
up to 1.35 cm tall
erect to decumbent,
10-60 cm tall
2-4
2-9, often
3-6
3-5, rarely
6-7
4-28, u.sual-
ly more
than 8
1^, often
2
3-12 2.7-3.5
4-14 2.7-3.7
3.5-6 1.8-2.8
7-20 1.6-2.8
ca 4
3.2^4.2
up to 2.0
1.5-3.0
0.5-1.5
1.0-3.0
2.0-3.0
2.7-3.4
2.4-5.5
0.9^2.7
1.4^.8
3.5-6.0
found, and the two are not givt>n taxononiic
statu.s.
Chloris ciliata is widely distributed in open
grasslands in the south central United States,
Mexico, Central America, the Caribbean region,
and in southern South America (Fig. 10).
Representative specimens examined: BA-
HAMA ISLANDS: Na,ssau, Curtiss 72 ( F, MO,
NY, US). ARGENTINA: Prov. Cordoba: Bell
Ville, Paradi 3,055 (US); Prov. Entre Rios:
Dep. Federacion, Estancia "Buena Esperanza,"
Pedersen 6,197 (US); Prov. Formosa: ]or<ien-
sen 3,328 (US); Prov. Jujuy: camino a San
Pedro, Parodi 9,796 (US); Prov. Mendoza:
Santa Rosa, Jensen-IIaanip s.n., 1904-5 (MO,
US); Prov. Salta: N of Salta City, Stephens and
Hartley SH 57 (US). CUBA: Prov. Habana:
Marianao, Ekrnan 1,192 (US); Prov. Pinar del
Rio: Bay of Mariel, Britton and Earle 7,570
( NY ) ; Prov. Santa Clara Lomas de Banao, Luna
888 (NY). HAITI: Port-de-Paix, Ekrnan 4,324
(US); Massif de la Pelle, Port-au-Prince, Ek-
rnan N. //. 8,145 (US); Gonave Island: vicinity
of Etroite, Leonard 3,371 (US); Tortue Island:
Basse Terre, Leonard and Leonard 12,444 (NY,
US); vicinity of Jean Rabel, Leonard and
Fig. 10. Distribution of Chloris ciliata. (A) Southern South America,
Islands. (C) Central America and Caribbean Islands.
( B ) Puerto Rico, Leeward, and Windward
30
Bricham Young University Science Bulletin
Leonard 12,934 (US). JAMAICA: Middlesex
Co.: Clarendon Parish, Inverness, Harris 12,755
(F, MO, NY, US); Manchester Parish, Harris
12,689 (F, MO, NY, US); Cornwall Co., St.
James Parish, Montego Bay, Hitchcock 9,671
(US). LEEWARD ISLANDS: Antigua: Dead-
sands Bay, Box 31 (US); Guadeloupe Island,
Duss 31,587 (NY, US); St. Barthelemy, Questel
337 (US). MEXICO: Coahuila: Castaiios,
Reeder, Reeder and Soderstrom 3,270 (US);
Nuevo Leon: Monterey, Kenoijer 315 (MO);
Oaxaca: Toniellin, Hitchcock 6,220 (US); San
Luis Potosi, 7.5 mi N of El Naranjo, McGregor
et al. 813 (US); Tamaulipas: 22 km S of Vic-
toria, Martinez and Lmjando F-2,283 (TEX,
US); Yucatan: Chichen-Itza, Swallen 2,409
(US); Tizimin, Swallen 2,491 (US); Uxmal,
Swallen 2,612 (US); Peto, Swallen 2,672 (US);
Merida, Souza-Novelo 15 (US). PUERTO
RICO: Ponce, A. Chase 6,495 (NY, US).
SANTO DOMINGO: Prov. Santo Domingo:
Llano Costero, Ekman 12,486 (US); Barahona,
Fuertes 308 (US). UNITED STATES: New
York: Yonkers, mill yard, Nash s.n., 24 July 1898
(NY). Texas: Aransas Co., Rockport, Swallen
10,297 (US); Bee Co., Pettus, Tharp 43,071
(TEX); Bexar Co., 20 mi S of San Antonio,
Silveus 137 (TEX); Calhoun Co., 2 mi SW of
Port Lavaca, Rowell and Webster 2,117 (TEX);
Cameron Co., Laguna Atascosa Refuge, Fleet-
wood 3,800 (TEX); De Witt Co., near Edgar,
Owens 155 (US); jim Wells Co., 6 mi W of Alice,
Gotdd and Kapadia 8,854 (UC); Karnes Co.,
5 mi S of Kennedy, Emery 773 (TEX); Kleberg
Co., King Ranch, M. C. Johnston 541,669 (TEX);
Matagordo Co., near Palacios, Rogers 6,607
(TEX); San Patricio Co., 1 mi SE of Sinton,
Gould 9,903 (TEX); Starr Co., Fort Ringgold,
E of Rio Grande City, Tharp and York 51-84
(TEX); Travis Co., Zilker Park, W. V. Brown
s.n., 6 Oct 1957 (TEX); Wharton Co., 12 mi
N of El Campo, W. V. Brown 3,265 (TEX);
Willacy Co., Sauz Ranch, M. C. Johnston and
Davis, 5,319.84 (TEX). URUGUAY: Dep. Mon-
tevideo, Herter 78,127 (US). VIRGIN IS-
LANDS: St. Croix: Bassin Yard, Ricksecker 42
(F, MO, NY, UC, US). WINDWARD IS-
LANDS: Barbados, Hitchcock 16,514 (US);
Grenada: Richmond Hill, Broadway 1.869
(NY); Martinique, Du.ss 1,273 (MO, NY. US).
5. CHLORIS LAMPROPARIA Stapf in Che-
valier, Bull. Bot. Soc. France Mem. IV.
8:220. 1912. (HOLOTYPE: Chad, "Bag-
uimii, emplacement de la ville de Massenia,
25-31 Aout 1903, no. 9633 bis." P! ISO-
TYPE: K!) Fig. 11.
Annuals 10 to 60 cm tall, erect to decum-
bent; stems usually branched at the base, these
sometimes rooting at the lower nodes; sheaths
glabrous; ligule a short ciliate crown; blades
5 to 20 cm long, 3 to 5 mm wide, sparsely pus-
tulose-pilose above, especially near the base,
glabrous to scabrous below; spikes one to four
(often two) ca 4 cm long, appressed or only
slightly divergent; spikelets densely inserted oh
the rachis, usually divergent, ca 10 per cm of
the long scabrous-hispidulous rachis; glumes
lanceolate-ovate, thin, brown, glabrous (except
for the scabrous midnerve); first glume 3.9 to
4.4 mm long, 0.6 to 0.9 mm wide, apex acute;
second glume 6.5 to 8.0 mm long, ca 1 mm wide,
apex acute to acuminate; fertile lemma 3.2 to
4.2 mm long, 0.8 to 1.7 mm wide, broadly ovate
to elliptic, dark brown, keel appressed-pubes-
cent, margins with spreading ciliate hairs, 2 to
3 mm long, internerves glabrous, apex acumin-
ate, awn 3.5 to 6.0 mm long; sterile florets two
or three; lowest sterile floret 1.3 to 1.6 mm
long, 1.0 to 1.5 mm wide, triangular, apex trun-
cate, awn 2.3 to 4.7 mm long, upper florets
similar in shape, but progressively smaller, un-
awned; caryopsis 1.5 to 2.0 mm long, 0.8 to 1.0
mm wide, ovoid to obovoid.
Chloris lamproparia is the only species in
the group with ciliate margins on the lemma
that is distributed in the Old World, and it is
also the only annual. It is most similar to C.
canterai, differing from it in having larger spike-
lets, though occasional specimens are inter-
mediate in spikelet size and can be assigned
most easily by their geographic origin and
habit.
Chloris lamproparia is distributed across cen-
tral Africa (Fig. 12), where it is reported to be
common in wet grasslands and savannas.
Representative specimens examined: CAM-
EROUN: Tchabaol, 45 km ENE of Maroua,
Letouzeij 6,473 (K). MALI: Bore, Dcmange
14/1.957 (K). NIGERIA: Domaturu, De Leeiiw
1,097a (K); Zaria Dtr„ Tat/lor H. 2,624/32 (K).
SUDAN: Darfur Prov.: 3 km NE of Kes, Jack-
son 4,040 (K); Jcbel Marra, Zalingei, Wickens
2,304 (K); Equatoria Prov., 13 mi SW of Ka-
poeta. Peers s.n., 29 Aug 1953 (K); Kordofan
Prov.: El Muglad, Andretos 3,063 (K); near
Hamadi, on El Obeid-Dilling road, Harrison
901 (K). TANZANIA: Tumba Rukwa, Michel-
more Cl,557 (K). UGANDA: Moroto, Wilson
15 (K). UPPER VOLTA: Ouagadougou, Chev-
alier 24,674 (K).
6a. CHLORIS CANTERAI Arechavaleta var.
CANTERAI. Anales Mus. Nac. Montevideo
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chlohis (Gbamineae)
31
r~\
Fig. 11. Chloris lamproparia. (A) habit, x 1/4; (B) spilcelet, partly dissected; (C) lemma with one side re-
moved, showing palea and sterile floret within; (D) sterile florets; (E-F) upper sterile florets; (G) cary-
opsis. (B-G) \ 10.
5:385. 1896. (HOLOTYPE: "Anderson . . .
Montevideo." Not seen, but original descrip-
tion clearly identifies the plant.) Fig. 13,
A, C-J.
Chloris parodiana Rosengurtt and Izaguirre,
Bol. Soc. Argentina Bot. 12:128. 1968.
(HOLOTYPE: "Paraguay, Paraguari . . .
Rosengurtt B-5,979, MVFA." Not seen, but
description and illustration are clear. )
Perennial up to 1 m tall, tufted and erect;
sheaths glabrous; ligule long-ciliate, with cilia
up to 7 mm long; blades narrow, often rolled
and appearing filiform, 1 to 5 mm wide, up to
25 cm long, glabrous to slightly scabrous, occa-
sionally sparsely long-pilose; spikes two to nine
(often three to six), 4 to 14 cm long, usually
more than 7 cm long, erect to curving; spike-
lets densely imbricate, appressed, light to me-
diimi brown, ca 11 per cm of the scabrous
rachis; first glume 1.6 to 2.4 mm long, 0.5 to
0.6 mm wide, narrowly lanceolate, markedly
inequilateral; second glume 2.3 to 3.8 mm long,
0.4 to 0.6 mm wide, narrowly lanceolate, equi-
32
Bbicham Younc University Science Bulletin
Fig. 12. Distribution of Chloris lamproparia.
lateral or inequilateral; fertile lemma narrowly
ovate, 2.7 to 3.7 mm long, usually more than 3
mm, 0.6 to 1.1 mm wide, ovate, callus bearded,
densely ciliate on keel and margins, marginal
cilia 1.5 to 3.0 mm long, intcmerves glabrous,
awn 2.4 to 5.5 mm long; sterile florets two
(sometimes three), similar, turbinate-flattened;
lower sterile floret 1.1 to 1.8 mm long, about
half as wide, glabrous; awn 1.5 to 3.5 mm long;
upper sterile floret(s) similar to lower in shape,
but smaller, about 0.5 mm long, awnless; car-
yopsis ovoid-ellipsoid, 1.3 to 2.0 mm long, 0.8 to
0.9 mm wide; chromosome number 2n=36.
6b. CHLORIS CANTERAI Arechavaleta var.
GRANDIFLORA (Rosengurtt & Izaguirre
de Artucio) Anderson, comb. nov. Fig. 13,
B-J. Chloris <:,ran(UfIora Rosengurtt and Iza-
guirre de Artucio, Bol. Soc. Argentina Bot.
12:127. 1968. (HOLOTYPE; "Uruguay,
dcp. Florida . . . Gallinal et Rosengurtt, PE-
5,790 . . . MUFA." Not seen, but original
description and illustration are clear.)
Perennial, densely cespitose, 5 to 30 cm tall
(often about 20 cm) arising from a dense
cluster of very narrow, often curved and rolled
leaves; spikes three to five, ca 3 to 6 cm long;
other characteristics as in var. canterai, except
chromosome number unknown.
The most marked difference within Chloris
canterai is a vegetative one. A group of speci-
mens, primarily from Argentina ( Fig. 14, open
circles) is shorter than usual and has a well-
defined, cespitose growth habit. No spikelet
distinctions could be found. A few of these
have been collected in the same precise loca-
tion as specimens of var. canterai; but the
collection data indicate that they are found in
dry, rocky areas as opposed to the lower, wetter
sites in which var. canterai is typically found.
While one immediately suspects that an environ-
mental factor may be influential, or that the
cespitose individuals are merely more heavily
grazed, the morphological disccmtinuity is so
well defined that it is felt that the population
is worthy of being recognized taxonomically.
Rosengurtt and Izaguirre de Artucio (1968)
have given this population the name Chloris
grandiflora. I feel that this overemphasizes the
differences, however, and I have consequently
reduced it to a varietal rank.
In addition, Rosengurtt segregated a second
species from this variable population, naming
it Chloris parodiana. It is based largely upon
the relative length of lemma cilia. I do not feel
that the differences emphasized are stable, at
least on the basis of the data summarized in
Fig. 9; and I, consequently, have treated it as a
synonym under var. canterai. Further studies
of a more experimental nature will be needed
to detemiine the validity of all of these taxa.
While some specimens of Chloris canterai
and C. lamproparia may be difficult to separate,
there are ample differences. Chloris canterai is
a tufted. South American perennial (Fig. 13)
with relatively smaller and narrower spikelets
and more spikes per inflorescence than C.
lamproparia, which is an African annual (Fig.
11).
Chloris canterai var. canterai is restricted to
northeastern Argentina, extreme southern Brazil,
Paraguay, and Uruguay ( Fig. 13, shaded cir-
cles), where it is reported as occurring on moist
soils on the campo. Chloris canterai var. grandi-
flora, on the other hand, is found on dry sites,
often rocky areas, most commonly in Argentina.
A single specimen from BM! (collected by
Gardner in 1839) is labeled "Banks of the Rio
Gurgueia, Piaui," an area of Brazil far to the
north of any existing collections and probably
represents a labeling error. All North American
plants arc naturalized introductions.
Representative specimens examined: CHLO-
RIS CANTERAI var. CANTERAI. AR-
GENTINA: Prov. Chaco: Tintina, Parodi 812
(US); Prov. Corrientes: Mercedes, Parodi 6,242
(US); Prov. Santa Fe: bajos del Rio Salado,
Burkart 9,077 (US); Prov. Formosa: Pilcomayo,
Morel, 6.569 (US). BRAZIL: Rio Grande do
Sul, Sao Gabriel, Rambo 25,567 (US); Uru-
guaiana, Poste Zootecnico, Stcallen 7,617 (US).
PARAGUAY: Dep. Paraguari: Caapucu, Rosen-
gurtt 5,405 (US); Dep. Presidentc Hayes: S of
Villa Hayes, Rosengurtt 5,631 (US). UNITED
STATES: Texas: Atascosa Co., 4.5 mi SE of
Jourdanton, /. C. Johnson 2,044 (TEX); Jack-
son Co., 4 mi E of Edna, Gould 6,781 (TEX,
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
33
Fig. 13. Chloris canterai var. cantcrai and var. grandiflora. (A) habit, C. canterai var. canterai, x 1/6; (B)
habit, C. cantcrai var. grandiflora, x 1/3; (C) spikelet, x 10; ( D.E ) sterile florets, partially dissected, x 15;
(F) uppermost sterile floret, x 15; (G) caryopsis, x 10; { H-J ) outlines of spikelets, showing variation, x 10.
34
Bricham Young University Science Bulletin
• =C, CANTERAI VAR. CAMERA 1
0=C. CANTERAI VAR, GRANDIFLORA
Fig. 14. Distribution of Chloris canterai var. canterai
(dots) and var. grandiflora (open circles) in South
America and Te.\as (Inset A).
UC); 14 mi NE of Port Lavaca, Gould and
Reeves 8,208 (TEX, UC); La Ward, Tharp et
al. 51-1,603 (TEX); 4 mi SE of Morales, Turner
3,276 and Tharp 53-430 (TEX); Milam Co.,
Buckholts, Gould 7,692a (TEX); Wharton Co.,
6 mi S of El Campo, Anderson 3,288 (HSC).
URUGUAY: Dep. Cancloncs: Pando, Montaro
709 (K); Dep. Cerro Largo: Rio Negro, Gal-
linal et al. PE-5,531 (MO, UC, US); Dep. Flor-
ida: Arroyo Mansavillagra, Rosengurtt 5,852
( US ) ; Dep. Paysandu, without preci.sc locality,
Rosengurtt B-909 (US); Dep. Rocha: Camino
de las Indios, Rosengurtt B-6,938 (K); Dep.
Salto: without precise location, Del Puerto s.n.
in LS74 (US). CHLORLS CANTERAI var.
GRANDIFLORA. ARGENTINA: Prov. Buenos
Aires: Sierra de la Ventana, Bartlett 20,029
(US); San Nicolas, P«ror// 1,023 (US). Prov.
Corrientes: Dep. Mercedes, Estancia "Ita
caabo," Parocli 6,118 (US). BRAZIL: Rio
Grande do Sul, Uruguaiana, Poste Zootccnico,
Swallen 7,656, 7,697 (US). URUGUAY: Dep.
Colonia: San Juan, Herter 18,789 (US); Dep.
Florida: Arroyo Mansavillagra, Rosengurtt 5,790
(US).
CHLORIS DANDYANA CD. Adams, Phv-
tologia 21:408. 1971. (See discussion of this
name below. ) Fig. 15.
Andropogon harhatum Linnaeus, Svst. Nat.
ed. 10. 2:1305. 1759. Non A. harhatum Lin-
naeus, 1771. (See discussion of this name
below. )
Andropogon polydactijlon Linnaeus, Sp. PI.
ed. 2. 2:1483. 1763. (See discussion of this
name below.)
Chloris pohjdactijla (Linnaeus) Swartz,
Prodr. Veg. Occ. 26. 1788. (Based on An-
dropogon polydactijlon Linnaeus. )
Chloris consanguinea Kunth, Rev. Gram.
1:89. 1829, pubhshed in key; Enum. Pi.
1:264. 1833, republished with full descrip-
tion. (HOLOTYPE: "Rio Janeiro," not seen,
but description is clear. )
Chloris elata Desvaux, Opuscules 73. 1831.
(HOLOTYPE: "Brasilio," not seen, but de-
scription is clear. )
Chloris arundinacea Nees ex Steudel, Syn.
PI. Glum. 1:207. 1854. (HOLOTYPE:
"Sello. Brasil." Fragment of type in US!
from B.)
Chloris harhata (Linnaeus) Nash, Bull.
Torrey Bot. Club 25:443. 1898. Non C. har-
hata ( Linnaeus ) Swartz, 1797. ( Based on
Andropogon harhatum Linnaeus, 1759. )
Chloris pohjdactijla f. stolonifera Parodi,
Rcvista Argentina Agron. 20:24. 1953.
(PARATYPE: US! A stoloniferous form.)
Perennial to 135 cm tall, usually tufted,
rarely stoloniferous, erect, arising from a thick
fibrous root system; sheaths glabrous, becoming
long-pilose at the ligule; blades up to 45 cm
long, 1.5 cm wide, usually glabrous with scab-
rous margins, occasionally scabrous on both
surfaces; spikes four to twenty-eight (usually
more than eight), flexuous, to 20 cm long, usu-
ally more or less .spreading, sometimes even
drooping; spikelets densely imbricate, averaging
12 per cm of the scabrous to hispid-ciliate
rachis; glumes pale, glabrous, with scabrous to
short-hispid midnerves; first glume 1.0 to 2.5
mm long, 0.2 to 0.4 mm wide, ovate-lanceolate;
second glume 1.9 to 3.5 mm long, 0.2 to 0.4
mm wide, narrowly lanceolate; fertile lemma
1.6 to 2 8 mm long, 0.5 to 0.9 mm wide,
elliptic, prominently flattened, usually pale
brown, margins prominently ciliate with hairs
at least 1 mm long and often up to 3 mm long,
keels densely appressed-pilose, awn 1.4 to 4.8
mm long; sterile florets usually two, the upper
hidden in the lower; lower sterile floret cvlin-
drical to narrowly turbinate, 1.0 to 1.6 mm long,
glabrous, awn 1.7 to 4.0 mm long; upper sterile
floret flattened and thin, 0.5 to 0.9 mm long,
0.3 to 0.7 mm wide, unawned; caryopsis ca 1
mm long, 0.5 mm wide, ellipsoidal.
Because of a complicated nomenclatural
problem involving Chloris dandijana and C
inflata, it is necessary that a complete review
of the history of these taxa be made.
BiOLOGiCAi, Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
35
M\miViM
4 ^!;:l^^»»»-s
Fig. 15. Chloris dandyana. (A) Inflorescence, x 1/3; (B) spikelet from specimen with long leimna margin
hairs; (C) spikelet from specimen with short lemma margin hairs; (D) sterile florets; (E) caryopsis; (F-J)
outline of series of spikelets, showing variation. ( B-] ) x 10.
In 1759 Linnaeus described Andropogon
barhatum (Syst. Nat. ed. 10. 2:1305), providing
a brief description and a reference to "Sloane.
hist. t. 65. f. 2." The protologue is inconclusive.
The plate in Sloane is likewise not confirming,
but the description of a Jamaican plant accom-
panying the plate (p. Ill) would seem to fit
best the plant described above. Reference will
be made later in this discussion to a specimen
labeled Aiidropogon barhatum in the Linnaean
Herbarium.
Linnaeus again described Andropogon bar-
batum in 1760 (Amoen. Acad., p. 412), chang-
ing and amplifying the description and drop-
ping the reference to Sloane, citing instead
"Brown, jam. 364." The description again is not
conclusive, but it would still appear to be the
above-described species.
In 1763 the above names, references, and
descriptions were tied together (Sp. PL, ed. 2.
2 ; 1483 ) ; but the name of the plant was changed
to Andropogon polijdactijlon, no reason being
given for the change. There is no question of
Linnaeus's intent, for he copied directly the
description accompanying the 1759 publication
of Andropogon barbatum and referred to Brown
and Sloane, this time adding a direct reference
to "p. Ill" of the latter work. Reference was
also made to the Amoenitates (1760).
In addition to the changes of names in the
written descriptions, corroborative evidence is
obtained from the Linnaean Herbarium, for
there (in the Andropogon folder) is a specimen
of the plant described above with three epithets
written on it: "barbatum," "fasciculatum," and
"polydactylon." The first two names are both
lined through; the last is not. Nonetheless, An-
dropogon polydactylon is clearly a superfluous
name and is illegitimate.
In 1771 Linnaeus again used the name An-
dropogon barbatum (Mantissa, p. 302), this
time providing a clear description of a plant
from the East Indies. There is no question
about this description; it obviously refers to the
36
Brigham Young Univebsity Science Bulletin
pantropical weed with inflated, truncate sterile
florets, commonly called either Chloris barbata,
C. paraguaijensis, or (as in this study) C. inflata.
There are two specimens— one immature, the
other mature— of this ta.xon in the Linnaean
Herbarium. One is labeled "barbatum"; the
name is not lined through. Again, there is no
explanation for Linnaeus's second and different
usage of Andropogon barbatum. The protologue
is a description only; there are no references to
previous publications. There is little question
then about the species referred to; it is cer-
tainly not the same ta.xon which was given this
name previously.
The transfer of Andropogon barbatum Lin-
naeus (1771) by Swartz cannot stand, for the
basionym is a later homonym. It did, however,
occupy barbata in Chloris, so that Andropogon
barbatum Linnaeus (1759)-the first name ap-
plied to the species with ciliate margins on
the lemmas described above— cannot be used
as was proposed by Nash ( 1898 ) .
Thus, all of the names used for this species
are unacceptable and the new name Chloris
dandtjana Adams is adopted.
Chloris dandijana is distinguished from
other members of this group primarily in spike-
let size and lemma pubescence— characters in
which it may approach C. ciliata, at least in
South America. It is distinguished from C. can-
terai in usually having a relatively large num-
ber of flexuous spikes. There are, however,
some specimens that differ from this pattern—
these plants usually having relatively short hairs
on the lemma margins and sometimes only
seven to ten short spikes. In an attempt to
learn more about this pattern of variation, many
specimens were scored for these and other
characters and various correlations were at-
tempted. Fig. 16 shows the geographic origin
of some of these specimens and their character-
istics. No sharply defined patterns emerged,
except that individuals with combinations of
short lemma hairs and short spikes occurred
randomly in South America but were neither
found in the West Indies nor in Florida. In
South America, however, these plants have no
distinctive distribution pattern. If these variants
were always found in areas of sympatry, a good
case could be made for introgression with an-
other species; but the presence of aberrant
plants in Venezuela, for instance, where the
nearest sympatric populations are in southern
Brazil, virtually rules out all but the most
ancient or fast-moving gene flows. Further
studies, especially attempts at hybridizing var-
ious portions of the populations, may shed light
• LEMMA MARGIN QUA MORE THAN
2.2 MM LONG
•-LEMMA MARGIN CILIA LESS THAN
2.1 MM LONG
• LEMMA LENGTH MORE THAN 22 MM
-• LEMMA LENGTH LESS THAN 21 MM
• SPIKES EIGHT OR MORE
4 SPIKES SEVEN OR FEWER
• SPIKES MORE THAN 7.0 CM LONG
f SPIKES LESS THAN 6.9 CM LONG
Fig. 16. Geographic distribution and morphological
characteristics of Chloris dandtjana.
on this problem. Since there are no sharp dis-
continuities in the variation pattern and only
slight correlations of the characteristics with
geographical distribution, I do not feel justified
in naming the variants.
Representative specimens examined: AR-
GENTINA: Prov. Catamarca: Dep. Capital,
Choya, Castillon 42,882 (US); Prov. Chaco,
without precise location. Asp 21 (US); Prov.
Cordoba, Rio Tercero, Stuckert 13,689 (US);
Prov. Corrientes: Isla Corrientes en el Rio Uru-
guay, Cerca Monte Caseros, Rosengurtt B-3,679
(US); Prov. Formosa: Costa Alegre, N de la
Primavera, Morel 7,039 (US); Prov. Jujuy: 15
km SE of San Pedro, Eyerdam and Beetle
22,320 (UC, US); Prov. Misiones: San Javier,
Obcra, Schwindt 4.779 (US); Prov. Salta: Dep.
Oran, Maldonado 790 (F, US); Dep. Cande-
laria, Ruiz de los Llanos, Montenegro 322. 325
(US); Rosario de la Frontera, Los Banos, Ven-
turi 7,974 (US); Prov. Santiago del Estero: be-
tween Forres and Robles, Bartlett 17,944, 19,744
(US); Beltran, Maldonado 510 (US); Prov.
Tucuman: Dep. Leales, Tres Pozos, Descole-
Schreiter s.n.. 5 Dec 19.38 (NY, US); Rio Chico,
Escaba, Monetti 42,931 (US); Cruz Alta,
Schreiter 3,987 ( US ) ; Burruvacu, Venturi 2,537,
2,661 (US), 7,536 (F, US). BAHAMA IS-
BiOLOciCAi. Series, Vol. 19, No. 2 Taxonomy of the Genu.s Chloris (Gramineae)
37
LANDS: Cat Island, Orange Creek, Britton and
Milkpaugh 5J33 (F, NY); Eleuthera I.sland,
Governor's Harbor, Hitchcock s.n., 14 Nov 1S90
(F, MO); Long Island, vicinity of Clarence
Town, Britton and Millspaugh 6,245 (F, NY);
New Providence, Blue Hills, Wihon 8,247 (F,
NY); Hog Island, near New Providence, Wilson
8,292 (F, MO, NY). BOLIVIA: Dep. Santa
Cruz: among bushes in savanna, Cardenas 5,362
(US); Dep. La Paz: Coripata, Hitchcock 22,689
(US); Dep. Cochabamba: without precise loca-
tion, Spiag,ai 26 (US). BRAZIL: Bahia: Cruz
das Almas, Pinto 746 (UC, US); Matto Grosso:
Corumba, A. Chase 11,136 (US); Aquidauana,
A. Chase 11,046 (US); between Campo Grande
and Dourados, A. Chase 10,915 (F, NY, US);
Minas Geraes: Buritys, near Rio Sao Francisco,
A. Chase 10,460 (US); Priapora, Dorsett and
Popenoe 3,436 (US); Pernambuco: vicinity of
Pernambuco, A. Chase 7,757 ( NY, US ) ; Rio de
Janeiro: San Pedro, NW of Gabo Frio, A. Chase
10,158 (US); vicinity of Sao Leopoldo, Leite
471 (NY); Geno Largo, near San Luiz, Ramho
53,116 (US); Santa Vitoria de Palmar, SwuIIen
7,416 (US). BRITISH GUIANA: Essequibo Co.,
Rupunini River, Loxton s.n., 26 Sep 1955 (K).
COLOMBIA: Dep. Antioquia: 1 km E of Puente
de Occidente, Pelaez and Barkleij 595 (US);
Dep. Bolivar: Tierrabomba Island, Cartagena
Bay, Killip and Smith 14,136 (NY, US): Dep.
Torrecillo: near Turbaco, Killip and Smith
14,639 (F, MO, NY, US); Div. Boyaca, Soata,
Cuatrecasas 1,094 (F, US); Dep. Gundinimarca:
Route de Honda a Guaduas, Humbert 27,097
(US); Girardot, Pennel 168 (NY); Dep. Huila:
3°23'N, 74°49'W, Fosherg 19,589 (US); Dep.
Valle: Hova del Rio Sanjunifiuin, Cnafreca.^as
15,366 (US); Dep. Magdalena: E of Los Ve-
nadas, 10°N, 73°43'W, ^ Ditgand 5,829 (US);
Dep. Santander: 12 km from Bucaramanga, Lati-
genheim 3,198 (UC, US). ECUADOR: Prov.
Imbabura: cntre Paramb;as y Pajon, Acosta-
Solis 12,667 (F, US). GRAND CAYMAN:
Hitchcock s.n., 19 Jan 1891 (MO). JAMAICA:
Cornwall Co.: Santa Cruz Mts., Britton 1.172
(NY); Middlesex Co.: Lower Clarendon Parish,
Inverness, Harris 12.743 ( F, MO, NY, US ) ; Lee-
ward Islands: Antigua, road to Hermitage Bay,
Box 26 (US). PARAGUAY: Dep. Boqueron:
Puerto Casado, Hartley SH 141 (US); Puerto
Sastre, Ramirez 5, 19 (US); Mariscal Estigar-
ribia, Rosengurtt B-5,813 (US); Dep. Goncep-
cion: Estancia "Fanciere," Ramirez 193 (US);
Dep. Paraguari: Caapucu, Anderson 1155A
(US). PERU: Dep. Cajamarca: Jaen, Hutchin-
son, 1,422 (UC, US); Dep. San Martin: Puca-
yacu, 11 km S of Tarapoto, Ferretjra 7,725 (US).
URUGUAY: Dep. Artigas: Santa Rosa del
Guareim, Herter 1,048 (US). UNITED
STATES: Florida: Dade Co., Miami, Tracy
8,857 (F, MO, NY, TEX, US); Monroe Co.":
Upper Matccumbe Key, A. Chase 3,914 (US);
Long Key, Silveus 5,329 (TEX); Key Vaca,
Sicallen 5.189 (US); Plantation Key. Swollen
5,210 (US); Big Pine Key, Swallen 14,461 (US);
Ke\- Largo, Weatherwa.x 61,009 (UC). VENE-
ZLTELA: Carabobo, Valencia, Carreno 8,236
( US ) ; Distrito Federal, vicinity of Caracas,
Bailey and Bailey 200, 554 (US); Merida, Tovar,
Fendler 1.854.5 (US); Miranda, between Los
Tequcs and El Garrizal, Pittier 12,973 (NY, US);
Tachira, La Mulera, Tamayo 2.296 (US);
Trujillo, San Jacinto, Tamayo 1,698 (US); Zulia,
savannas of Lagunillas, Jahn 658 (US).
8. CHLORIS CRUCIATA (Linnaeus) Swartz,
Prodr. Veg. Ind. Occ. 25. 1788. (Based
on Agrostis cruciata Linnaeus. ) Fig. 17,
A-E.
Agrostis cruciata Linnaeus, Svst. Nat., ed.
10. 2:87.3. 1759. ("Sloan, jam. t. 69. f. I."
cited by Linnaeus. The illustration cited is
a good likeness of the species treated here.
A specimen, in LINN!, labeled Agrostis
cruciata is somewhat immature and does
not ha\'e the spreading spikelets character-
istic of the species. In all other respects
the specimen fits the usual circumscrip-
tion. )
Rahdochloa cruciata (Linnaeus) Beauvois,
Ess. Agrost. 84, 158, 176. 1812. (Based on
Chloris cruciata [Linnaeus] Swartz.)
Cynodon cruciafus (Linnaeus) Raspail,
Ann. Sci. Nat. Bot. (Paris) 5:.30.3. 1825.
(Based on Chloris cruciata [Linnaeus]
Swartz. )
Chloris humholdtiana Steudel, Syn. PI.
Glum. 1:205. 1854. ("C. cruciata H. B. [vix
Swartz.] Ins. Cuba" cited. Description
clearly of C. cruciata as commonly under-
stood. )
Chloris hrevightma Wright, Anales Acad.
Ciencias Habana 8:200." 1871. (HOLO-
TYPE: "1848. Ganuela. . . ." is cited in pro-
tologue. "1848" is evidently an error for
"1548" for a photograph, in US!, of the
type in GH has the latter number. Isotypes,
also with the number "1548" are in BM!
and Gi.)
Tufted perennial up to 35 cm tall; sheaths
glabrous; ligule shortly pilose; blades very nar-
row, usually less than 0.5 mm wide, up to 6
38
Brighaai Young University Science Bulletin
$m^
Fig. 17. Chloris cniciata, C. filifoTmis, and C. suringari. (A-E) C. cruciata. (A) habit, x 1/6; (B) portion
of single spike, x 2; (C) spikelet, partly dissected, x 10; (D) sterile floret, x 10; (E) caryopsis, x 10.
(F-J) C. suringari. (F) habit, x 1/5; (G) portion of spike showing rachis and attached spikelets, x 10;
(H) spikelet, partly di.ssected to show lowest sterile floret, x 10; (I) upper two sterile florets, x 10; (J)
caryopsis, x 10. (K-L) C. filifonnis. (K) habit, x 1/3; (L) spikelet, partly dissected, x 10.
cm long, involute; inflorescence of two or three
radiate branches, spreading horizontally; spikes
1.0 to 2.5 cm long; spikelets somewhat ap-
pressed when immature, eventually spreading
at nearly right angles to the rachis ; spikelets
only slightly ( if at all ) imbricate, averaging
five or six per cm of the rachis length; first
glume ovate-lanceolate, 0.7 to 1.3 mm long,
0.1 to 0.3 mm wide, glabrous, midrib sometimes
slightly scabrous; second glume narrowly lan-
ceolate, 1.6 to 2.2 mm long, 0.1 to 0.3 mm wide,
glabrous, midrib occasionally scabrous; fertile
lemma pale, narrowly lanceolate, 2.2 to 2.8 mm
long, 0.3 to 0.5 mm wide, glabrous, occasionally
the inroUed hyaline margins with a few short
hairs, callus bearded, awn 7.5 to 17.0 mm long;
sterile floret one, 0.5 to 1.4 mm long, 0.2 to 0.3
mm wide, narrowly truncate, borne on a fairh'
long rachilla joint, awn 3.7 to 7.0 mm long;
caryopsis narrowly cylindrical, 1.4 to 1.5 mm
long, ca 2.0 mm wide.
Chloris cniciata is the only species in the
genus with a tufted habit, very narrow leaves,
two or three horizontally spreading spikes, and
spikelets that are divergent from the rachis at
right angles (Fig. 17). Other species from the
Caribbean (especially C. sagraeana and C. ek-
inanii) have similar, though appressed, spike-
lets; they also lack the other features.
Epidermal studies of this species ( Rogers,
1967, and those reported earlier in this paper)
revealed the presence of somewhat cruciate
short cells, a feature not found elsewhere in
Chloris. All inflorescence and spikelet character-
istics are typical of Chloris, however, and I do
not feel that ta.xonomic recognition of this spe-
cies as a distinct genus is warranted.
Chloris cruciata has been collected in stony
Biological Series, Vol. 19, No.
Taxonomy of the Genus Chloris (Ghamineae)
39
soils (often of serpentine origin) from Cuba,
Haiti, and Jamaica (Fig. 18).
Representative specimens examined: CUBA:
Prov. Habana: Guanabacoa, Lomas de Jatas,
Ekman 279 (US); San Miguel de Casuora,
Loma de la Vita, Leon 11,753 (US); Prov. Las
Villas: 10 km S of Santa Clara, Howard et al.
100 (UC); vicinity of Sancti Spiritus, Leon
1,497 (US); Oriente Prov.: Bazate, banks of
Rio Canto, Ekman 3,016 (K, NY, US). HAITI:
Massif du Nord, between Cerca-Carvijal and
Bois-Charles, Ekman H607 (US). JAMAICA:
Cornwall Co., Troy, Harris 12,647 (F, K, MO,
NY, US); St. Elizabeth Parish, between Ipswich
and Black River, Hitchcock 9,597 (US); Middle-
sex Co., vicinity of Mandeville, Britton 1,029
(NY); SurrcN' Co., St. Andrew Parish, Stonv
Hill, Harris 'll,341 (F, K, NY, US); Clifton,
Harris 11,375 (F, NY, US); near Kingston, along
Stonv Hill Road, Hitchcock 9,276 (F, NY, UC,
US); Blue Mountains, Yallahs Valley, Perkins
1,481 (US).
9.
CHLORIS FILIFORMIS (Vahl) Poiret in
Lam. Encycl. Method. Bot. Suppl. 2:238.
1811. (Perhaps based on Ctinosurus fili-
formis Vahl.) Non Chloris filifortnis Poiret.
Fig. 17, K-L.
Ci/nosurus filiformis Vahl, Symbolae Bot.
2:20. 1790. (HOLOTYPE: Photograph, in
US!, of type from Vahl Herbarium, C. )
Chloris filiformis Poiret, in Lam. Encvcl.
Meth. Bot. Suppl. 2:237. 1811. (TYPE
MATERIAL: P! There are several speci-
mens at P, in the Herbarium Richard,
labeled "Chloris filiformis Poiret. He. de
France." While Poiret did not designate a
ts'pe as such, they probably represent au-
thentic material. Tlie specimens fit the de-
scription closelv; and this is, unquestion-
ably, the same taxon first described by
\'ahl under Cynosurtis, though Vahl is not
cited by Poiret in the protologue of this
name. )
Chloris distachya Kunth, Rev. Gram. 1:291.
pi. 57. 1830. "(HOLOTYPE: "Chloris dis-
tachya Kunth, Insula franciae. Ex Museo
Fig. 18. Distribution of Chloris crucUita.
Paris." Not seen, but description and ac-
companying plate are clear. )
Stoloniferous or tufted perennial, 10 to 30
cm tall; sheaths glabrous to sparsely pilose,
compressed, keeled; ligule a very short ciliate
crown, often with a few longer hairs near the
margins; blades narrow, 1 to 6 mm long, about
1 mm wide, folded, often curved, sparsely pilose
near the base, becoming apically glabrous,
acute; spikes one or two, 1 to 3 cm long, erect;
spikelets rather densely imbricate, ca 12 per cm
of rachis length, more or less appressed; glumes
lanceolate, glabrous; first glume 1.7 to 2.0 mm
long, ca 0.4 mm wide; second glume 2.5 to 2.8
mm long, ca 0.4 mm wide; fertile lemma 2.6 to
2.8 mm long, 0.5 to 0.6 mm wide, lanceolate-
elliptic, apex acute, callus bearded, internerves
densely appressed-pubescent, awn 1.0 to 1.2
mm long; sterile floret 1.3 to 2.0 mm long, 0.2
to 0.5 mm wide, narrowly lanceolate to linear,
acute, sparsely appressed-pubescent, awn 0.5 to
1.1 mm long; caryopsis not seen.
Known only from Mauritius and nearb\- is-
lets, Chloris filiformis is easily distinguished
from the similar C. .suring,ari (from Cuba and
Cura^'ao) by having considerably shorter awns
and lemmas with appressed pubescence on the
internerves rather than spreading pubescence
on the margins and keels (Fig. 17, K-L, F-J,
respectively ) .
Specimens examined: MAURITIUS: Mon-
tague Longue, dry regions, Botiton s.n., without
date (K);"R()und' Island, rocky hillside, 520 ft.,
//. //. Johnston s.n., 28 Nov 1889 (K); Round
Island, Wiene s.n., Nov 1937 (K). Hubbard and
Vaughn (1940) reported this species also from
the islets of Gunners Quoin and Le Morne.
10. CHLORIS SURINGARI Hitchcock in Ur-
ban, Svmb. AntiU. 7:167. 1912. (HOLO-
TYPE: "Curasao ad Brakkeput . . . W. F. R.
Suringar, Jan 14, 1885" Type fragment, in
US!, from Krug and Urban Herbarium.)
Fig. 17, F-J.
Densely tufted, very shortly stoloniferous,
diminutive perennial up to 15 cm tall; sheaths
very short, about 1 cm or less, striate, sparsely
to densely pilose below, becoming glabrous
above on some culms; ligule a short tuft of
hairs; blades highly reduced, less than 0.5 mm
wide and 2 cm long, usually somewhat curved;
apex blunt to acute, glabrous to sparsely pilose;
inflorescence a single spike 1.0 to 2.5 cm long;
spikelets imbricate, about 15 per cm of the
rachis; first glume 1.0 to 1.5 mm long, 0.4 to
0.5 mm wide, lanceolate, appressed to the
40
Brigham Young University Science Bulletin
rachis, one-nerved, glabrous, marginal areas thin
and hyaline; second glume very narrowly lan-
ceolate, diverging from the rachis at about 70°,
2.3 to 3.0 mm long, ca 0.3 to 0.4 mm wide, one-
nerved, glabrous, margins hyaline; fertile lemma
ovate-lanceolate, 1.7 to 2.5 mm long, 0.4 to 0.6
mm wide, densely pubescent on the callus,
margins and keel with hairs about 0.7 mm long,
avvned from about the middle of the keel, the
awn 2 to 3 mm long, straight or slightly genicu-
late; sterile florets three, the lowest about 1 mm
long, 0.3 to 0.4 mm wide, avvned from about
the middle, the awn 1.5 to 2.0 mm long; upper
sterile florets much reduced, awned; caryopsis
ca 1.2 mm long, 0.5 mm wide, ovoid, flattened
to slightly trigonous.
This unique species, endemic to Cuba and
Cura9ao, may be immediately separated from
the similar species, C. filiformis, in having rela-
tively long awns and a single spike ( Fig. 17,
F-J). Additional notes on the relationships of
these species have been presented under C.
filiformis.
Collection labels indicate that Chloris sur-
ingari grows on limestone benches and rocks
not far from the seashore. Only four specimens
were seen in all herbaria consulted.
Specimens e.xamined: CUBA: Oriente Prov.,
El Guanal, Leon 11,564, Jan 1940 (US). CURA-
SAO: without precise location, Bohlinp^h 9.281.
1909-1910 (US); limestone bench near Willem-
stad, Britton and Shafcr 3,058, 20-27 Mar 1913
(US); Kleine Berg, Mordant 251, 23 Nov 1966
(P)-
11. CHLORIS SCARIOSA F. von Mueller,
Fragm. Phyt. Australiae. 6:85. 1867.
(HOLOTYPE: MEL, not seen. There are
two specimens at Kew, collected by von
Mueller in 1879, both of this species as
commonly understood. One is labeled
"Chloris scariosa F. Muell. Gracemere.
Bowman. Herb. F. Mueller. 1879." The
other is labeled "Chloris lativalvis Muell.
Sturt's Creek. M. Gregory." Lazarides
[1972] says that the latter specimen is an
isotype of the holotype at MEL. The pro-
tologue. which is very short, refers to
"glumis latis scariosis insignis." Since von
Mueller, as evidenced by other descriptions
on the same page, was familiar with the
species of Chloris in Australia, there is no
doubt as to the taxon to which he referred
and Lazarides 's designation of the Gregory
specimens as the type is reasonable. Chloris
lativalvis was apparently never published.)
Fig. 19, A-D.
Erect cespitose or stoloniferous perennial,
25 to 50 cm tall; sheaths glabrous to very
sparsely spreading pubescent; ligule a short
ciliate crown; blades up to 20 cm long, 5 to 8
mm wide, glabrous to scabrous or very sparsely
pilose; spikes four to six, 3 to 5 cm long, virgate
to somewhat spreading; spikelets imbricate, ca
7 per cm of the scabrous rachis; glumes un-
equal, purplish, membranous, glabrous except
for the scabrous midrib; first glume elliptic to
linear, 4.0 to 4.7 mm long, ca 0.5 mm wide,
rounded at the apex; second glume 5 to 7 mm
long, ca 1 mm wide, narrowly oblanceolate to
obovate or elliptical, apex bifid; fertile lemma
highly modified and differing from any other
species in Chloris, callus narrow and elongated
at the base of the lemma, 1.0 to 2.5 mm long,
densely bearded, sharp-pointed, body of lemma
ovate to ellipsoid, 1.0 to 2.5 mm long, 0.5 to
1.2 mm wide, margins membranous, prom-
inently winged, glabrous below, becoming
densely spreading ciHate above, keel sparsely
to densely pilose, especially below, awn 5.8 to
8.5 mm long; sterile florets three to five, broad,
membranous margined, with prominent veins;
lowermost floret broadly cordate or even am-
plexicaulous at the base and partly surrounding
the corky rachilla, tapering to a narrow tip
above, ca 4.5 mm long, ca 5 mm wide, awn 5
mm long, upper sterile florets progressively re-
duced; caryopsis 1.3 to 1.5 mm long, 0.8 to 1.0
mm wide, broadly obovoid-ellipsoid, trigonous;
chromosome number 2n=^40.
No other species in the genus has such
highly modified spikelet parts as Chloris scar-
iosa. The broadly winged fertile lemmas and
sterile florets at once set this species apart from
all others. This unique character was recognized
by Thellung (1919), \\ho erected the section
Hackelochloris to include it.
Furthei-more, there is a strong overall re-
semblance between C. scariosa and various spe-
cies of Tetrapogon. Certainly, the two genera
approach closely at this point and must be
rather arbitrarily separated on the basis of fer-
tile floret number. Both epidermal and anatomi-
cal features have been studied; no significant
differences between the two genera can be
demonstrated. On these bases it does not seem
desirable to erect a new genus for this single
species.
Chloris scariosa has been collected from a
variety of soils, most commonly in arid regions
of Queensland and Northern Territory, Aus-
tralia (Fig. 20).
Representative specimens examined: AUS-
TRALIA: Northern Territory: 5 mi S of Yam-
BiOLOCicAi, Series, Vol. 19, No. 2 Taxonomy of the Genus Cm,onis (Gramineae)
41
Fig. 19. Chloris scariosa and C. prieurii. (A-D) C. scariosa. (A) habit, x 1/4; (B) spikelet, partly dis-
sected, X 5; (C) sterile florets, partly dissected, x 5; (D) caryopsis, x 10. (E-H) C. prieurii. (E) habit,
X 1/4; (F) spikelet, partly dissected, x 10; (G) sterile florets, partly dissected, x 15; (H) caryopsis, x 10.
42
Bricham Young Universit-i- Science Bulletin
Fig. 20. Distribution of Chloris scariosa.
bah Station, Perry 3,368 (K); near coast, 60 mi
N of Wollogorong Station, Pern/ 1,218 (K, US);
8 mi N of Tennant Creek, Perry 551 (K); Burt-
well, Cleland 349 (K). Queensland: Burke Dtr.,
Cloncurry, Allan 20 (K); Forest Home Station,
Brass 1,790 (K); near Normantovvn Township,
flood flats of Norman River, Lazarides 4,248
(K, US); Charters Towers, Michael 1,299 (K);
Antil Plains, near Townsville, White 8,909 (K,
NY, US); Warrego Dtr., claypans near Cun-
namulla. White 11,566 (K, US). South Austra-
lia: Cordillo (Downs), Clehnul s.n., 5-24 (Ma\-,
1924?). Western Australia: between Gasco\ne
and Fortescue River, King s.n. 1885 (K).
SWITZERLAND: Derendingen, Kammgarn
Fabrik, Prob.st s.n., 25 Oct 1919, adventive (K).
12. CHLORIS PRIEURH Kunth, Rev. Cram.
1:89. 1829. (HOLOTYPE: "... He St.
Louis [Senegal] Roger Dedit, Major 1S25."
P! ISOTYPE: K! Fig. 19, E-H.)
Chloris cryptostachya Steudel ex J. A.
Schmidt, Beitr. Flora Cap Verdischen In-
seln 148. 1852, (HOLOTYPE: " . . . C.
punctulata Hochstetter hrbr. u. it. iiubic. nr.
23 Cordofan." P! ISOTYPE: K!)
Chloris punctulata Hochstetter ex Steudel,
Syn. PI. Glum. 1:205. 1854. (HOLO-
TYPE: "Hrbr. un. it. nubic. nr. 2.3." P!
ISOTYPE: K!)
Chloris subtriflora Steudel, Syn. PI. Glum.
1:208. 1854. (HOLOTYPE: "LaPrieur in
Senegambia." No collection that can be
identified positively as the holotype can be
found, though there are specimens in the
Herbarium Richard in P! that were col-
lected by Le Prieur in Senegal in 1829.
These are not labeled as Chloris triflora,
and there are no other data on the labels.
Dr. W. D. Clayton, in a personal commu-
nication, says that the type cannot be found
at CN. Under these circumstances, it must
be presumed lost. The description, while
not detailed enough for positive identifica-
tion, suggests strongly that the plant re-
ferred to is C. prieurii. )
Chloris multiradiata Hochstetter, Flora
38:204. 1855. (The original description
seems to refer to C. prieurii. There are
possible isotypes in K! and P! The K speci-
men is C. virgata; the P specimen, C.
prieurii. )
Stoloniferous perennial up to 80 cm tall;
sheaths glabrous, occasionally becoming pilose
near the ligule; ligule variously short-ciliate to
long-pilose; blades narrow, 10 to 14 cm long, up
t(j 5 mm wide; lower surface glabrous, upper
surface scabrous to pilose; spikes appressed to
slightly di\'erging, three to seven, 6 to 11 cm
long, spikelets imbricate, 8 to 11 per cm of
the scabrous-hispidulous rachis; glumes thin
and delicate, narrowly lanceolate, glabrous with
slightly scabrous midnei-ves; first glume 2.1 to
2.2 mm long, 0.2 to 0.3 mm wide; second glume
3.7 to 4.0 mm long, 0.2 to 0.3 mm wide; fertile
lemma nairowly elliptic, 3.3 to 4.7 mm long,
0.4 to 0.7 mm wide, with a row of clavate
glandular hairs along the middle of the lemma
side, awn 10 to 18 mm long; sterile florets
four or five, the lower successively enclosing
the upper and becoming progressively more
reduced; lowemiost sterile floret cylindrical, 1.5
to 2.5 mm long, 0.2 to 0.4 mm wide, awn 8 to
17 mm long, uppermost sterile floret more or
less flabellate, ca about 0.3 mm long, 0.1 mm
wide, awn 1.5 to 2.0 mm long, intermediate
sterile florets gradualh' changing from cylin-
drical to flabellate; caryopsis 2.0 to 2.5 mm
long, 0.5 mm wide, ellipsoid, strongly flattened,
light brown.
(July two species of Chloris have a row of
glandular hairs on the side of the lemma— C.
prieurii and C orthonoton. The two are easily
separated. C. prieurii is limited to the Old
\\'orId ( Fig. 21 ) and has four or five sterile
florets (Fig. 19, E-H); C. orthonoton is, on
the other hand, from South America and has
a single sterile floret (Figs. .33, A-D; 39).
Representative specimens examined: BOTS-
WANA: between Kapupahedi and Tamso, Oka-
vango National Territory, anon. (K). CHAD:
Bilthie, Ro.ssetti 39 (BM). CAPE VERDE IS-
LANDS: Boa Vista, Chevalier 44,423 (K).
ETHIOPIA: Nario Dikeno, Schimper 1,607
(MO). GHANA: Accra, Irvine 3,023 (MO).
Biological Series, Vol. 19, No. 2 Taxonomy ok the Genus Chlohis (Gramineae)
43
Fig. 21. Distribution of Chloris prieurii. Inset A:
India.
INDIA; Rajasthan Prov., Jaipur, Raizada 2/53
(K). MALI: San.sancling, Chevalier 24,958 (P);
Nara, Chudeau sm., 26 Aug 1918 (P); Tim-
buktu, Hagerup 239 (US). NIGERIA: Bomu
Prov., 48 mi N of Damaturu, De Leeuw 1,150
(K). SENEGAL: Kaolack, Berhaut 456 (P);
Mbambev, Chevalier 33,795 (P); Pointe Louis,
Roberfi/' 149.16.2 (K). SUDAN: Blue Nile
Prov., Bagein, 50 mi S of Khartoum, Jackson
2,336 (K); Darfur Prov., ]chv\ Mana, Blair 140
(K); Khartoum Prov., Khartoum, Schweinftirth
813 (US); Kordofan Prov.: El Muglad, An-
drews 3,062 (K); El Obeid, Harrison 76 (K).
UNITED STATES (adventive): Alabama: Mo-
bile, Mobile and Ohio wharf, Mohr 44 (US);
North Carolina; Wilmington, McCarthi/ 90
(US). UPPER VOLTA; Gourma, Rossetti 110
(K).
13. CHLORIS ROBUSTA Stapf in Chevalier,
Bull. Soc. Bot. France Mem. IV 8:221.
1912. (DUPLICATES OF SYNTYPES;
"Haut-Chari, Ndelle, 20-2.5 Dec 1902, Chcx-
alicr, 6.991; Nicreria du Nord, Borgu. Bar-
ter, S7S." K! ) Fig. 22. A-D.
Robu.st perennial to .3 m tall, aquatic to
subaquatic with very strong, heavy culm bases,
and often dense rooting at the lower nodes,
stoloniferous with stolons up to 7 m long;
sheaths glabrous; ligule a minute naked crown;
blades up to 40 cm long and 20 mm wide,
glabrous and glaucous or scabrous; spikes seven
to twenty-eight, 6 to 10 cm long, erect, some-
what flexous; spikelets imbricate, ca nine per
cm of rachis length; first glume ovate-lanceolate,
glabrous except for the slightly scabrous mid-
nerve, 1.6 to 2.5 mm long, ca 0.2 to 0.3 mm
wide; second glume linear-lanceolate, arcuate,
glabrous except for the scabrous midnerve, 4.0
to 5.5 mm long, 0.4 to 0.5 mm wide; fertile
lemma 3 to 4 mm long, 0.4 to 0.5 mm wide,
narrowly elliptic, apex long-acuminate, tapering
into the awn, upper % of the margin with long
hairs, these up to 2.5 mm long; callus bearded,
awn ca 0.5 to 1.0 mm long; sterile florets two;
lower sterile floret similar to the fertile floret
in shape, staminate, often with a rudimentary
pistil, 2.5 to 3.5 mm long, ca 0.3 mm wide,
margins with cilia up to 2.5 mm long, callus
bearded, upper lemma sides sparsely pubescent,
palea present, about as long as the sterile
lemma; upper sterile floret clavate, less than 1
mm long, pilose. Caryopsis not seen.
Chloris rohusta is unique in many respects,
including the bamboolike growth habit, pref-
erence for aquatic to subaquatic habitats, and
spikelets with sterile florets that are regularly
staminate. These features, together with the
overall morphological aspect of the spikelets,
suggest that this species should be separated
from the rest of Chloris. Epidennal and ana-
tomical characters, however, are those typically
found in other species of Chloris; and, there-
fore, there seems to be little justification for
segregating it.
This species has been collected throughout
equatorial Africa (Fig. 23). Its tall, bamboo-
like, elastic culms form large colonies along
and in rivers and streams. Associated grasses
include Phragmites communis and species of
Andropogon.
Representative specimens examined; CAM-
EROUN; riverain forest of Sanaga River, near
Goyoum, 20 km W of Deng Deng, Breteler
980 (K). DAHOMEY: entre de Lac Ozri et
Zagnando, Chevalier 23.054 (K). GHANA; 3
mi from Yendi, Kufapindi River, Adams and
Akpahla 4,101 (K); 30 mi from Navrango,
Tumu Road, Adams and Akpabla 4,361 (K);
Sakogu-Shishe Road, old Morago River bridge,
Ankrah 20.507 (K); Daka River near Yendi,
Brand 444 (US); Weija, near Accra, Hall 2,552
(K); \\'enchi, between Banda and Menji, Mor-
ton GC 25,256 (K); near Daboya, Thorold 288
(K). I\'ORY COAST: Marabadiassa, along
Bandama Blanc River, As.si 7.178 (K); entre
Sanlo and Kalepui, Assi 9,291 (K); bords du
Bandama River, Dume, Roherty s.n. (MO
1640174). NIGERIA: between Oyo and Iseyin,
44
Brigham Young University Science Bulletin
Fig. 22. Chloris robusta and C. Mollis. (A-D) C. rohmta. (A) lower and middle portions of stem, \ 1/3;
(B) upper stem and inflorescence, x 1/4; (C) spikelet, partly dissected, x 10; (D) sterile florets, x 15.
(E-J) C. mollis. (E) habit, x 1/4; (F) spikelet from type of C. mollis, partly dissected, x 10; (G)
floret from type of C. anlsopoda, x 10; (H) sterile florets from spikelet with two sterile florets, x 20; (I)
sterile floret from spikelet with single sterile floret, x 15; (J) caryopsis, x 5.
Biological Series, Vol. 19, No. 2 T.^.xonomv of the Genus Chlobis ( Gr.a.mineae)
45
Fig. 23. Distribution of Chloris robusta (equatorial
Africa).
Brenan and Keaij, 8,959 (K); Yola, Dalziel 269
(K); Lagos, Foster 2 (K); Jebba, on Niger
River, Uagerup 692 (K, US); Onitsha Prov.,
near Nzam Anambra Forest Reser\'e, Oramili
Creek, Jones 6,888 (K); Oyo Prov., Dtr. Ibadan,
Awba Hills Forest Reserve, Onochie and Jones
14,666 (K); bv Chinchaga River, near Muina,
Meikle 736 (K); Kaiama, Ward 35 (K, US).
REPUBLIC OF THE CONGO; Prov. Equa-
teur, Bangui, Ubangi River, MeruiUan 209 (K).
SIERRA LEONE: Moa River, Mafindo Falls,
near Kailahun, Deifihton 4,004 (K, US). SU-
DAN; Equatoria Prov., 40 mi NW of Yei, Myers
7,865, 8,496 (K); Bahr Chazal Prov., Mvolo.
Schwcinfurfh 2,843 (US). UGANDA: Kitgum,
Cluia. Thomas 4,346 (K).
14. CHLORIS MOLLIS (Nees) Swallen, North
AiiKT. Flora 17:596. 19.39. (Based on Ct/w-
nopo^on mollis Nees.) Fig. 22, E-J; 24.
Gi/mnopogon mollis Nees, Agrost. Bras.
427. IS.3,3. (FRAGMENT OF TYPE?: US!
Labeled "G\ninop(>gon mollis Nees. Type.
Campi Mimoso dicit. Prov. Piauhy." The
name of the herbarium from which the
fragment was obtained is illegible. While
the infomiation on the label is sketchy, it
does match exactly the collection location
noted by Nees. This, together with the
very complete original description, leave
no question as to the typification of this
name.)
Gijmnopogon rupestre Ridley, Jour. Linn.
Soc. Bot. (London) 27:73. 1891. (HOLO-
TYPE: BM! ISOTYPE: US! "Fernando do
Noronha. Cliffs on Portuguese Bay. Ridley,
Ramage, Sept. 14,1887.")
Chloris luetzeUmrgii Hitchcock, Proc. Biol.
Soc. Washington 36:197. 192.3. (HOLO-
TYPE: "Serra do Borborema, State of Para-
hvba do Norte, Brazil, April, Lutzelburg,
No. 12451" US!)
Chhris anisopoda Scribner in Robinson,
Proc. Amer. Acad. Arts Sci. 38:118. 1902.
ISOSYNTYPES: [Galapagos Islands]
"Charles Isl., Baur, 333; Snodgrass and
Heller, 459" US! SYNTYPES; GH, not
seen. )
Chloris angustiflora Arcsehoug, Kongl. Sven-
ska Fregatten Eugenics Bot. III. 118. 1910.
(TYPE: not seen, but description of speci-
men, collected on the island of Puna, by
Andersson is conclusive.)
Chloris leptantha Hitchcock e,\ Urban,
Symb. Antill. 7:166. 1912. (HOLOTYPE:
"Hab. in insulis orae \'enezuelensi adjacen-
tibus Bonaire, 19 Feb 1885. W. F. R. Sur-
ingar." US!)
Chloris rupestris (Ridlev) Hitchcock, U. S.
Dopt. Agric. Misc. Publ. 243:126. 1936.
(Based on Gijmnopogon rupestre Ridley.)
Annuiil 40 to 60 cm tall from a fibrous,
rather poorlv developed root system; sheaths
usually denseh- appressed- to spreading-pilose,
especialK near the apex, occasionally i-nclosing
cleistogamous spikelets; ligule long-pilose;
blades variable in size, ranging from short and
verv narrow up to -30 cm long and 1 cm wide,
appressed-pilose on both upper and lower sur-
faces; inflorescence of three to seven spikes, 6
to 9 cm long, mostly radiate at the apex of the
culm, sometimes with one or two single spikes
inserted separately just below; spikelets imbri-
cate, seven to ten per cm of rachis length;
glumes thin, transparent, narrowly lanceolate,
glabrous except for the scabrous midnerve; first
glume 3 to 4 mm long, 0.2 to 0.3 mm wide,
Fig. 24. Distribution of Chloris mollis. Insets: (A)
northern South America. (B) eastern Cuba, Ja-
maica, Hispaniola. (C) Galapagos Islands.
46
Brigham Young University Science Bulletin
glabrous; second glume 3.1 to 4.6 mm long, ca
0.3 mm wide, glabrous; fertile lemma very
narrow, 2.5 to 6.8 mm long, ca 0.3 mm wide,
glabrous e.xcept for the prominently bearded
callus and the short-ciliate upper margins, awn
4.0 to 7.5 mm long; sterile floret usually one
(occasionally two), and then the lowermost
bearing rudimentary flower parts; lowermost
sterile floret cylindrical, 1.2 to 1.6 mm long,
0.1 to 0.2 mm wide, awn 3.7 to 5.8 mm long,
callus prominently pubescent with spreading
hairs, margin ciliate above; upper sterile floret,
when present, ca 1 mm long, 0.1 mm wide,
callus bearded, awn ca 2.5 mm long; caryopsis
ca 3.3 mm long, ca 0.5 mm wide, narrowly
cylindrical.
ChloTts mollis is unique in the genus in hav-
ing sterile florets with a prominently pubescent
callus and long awns.
As may be seen from the synonymy, this
species has been placed in both Gymnopogon
and in Chloris. In some respects it does have
characters of both genera. Many species of
Gymnopogon are characterized b\' having short,
stiff, sharp-pointed leaf blades, which appear
crowded at the base of the culm because of
their short, overlapping sheaths. Chloris mollis
has these characters, except that the leaves are
not crowded at the base. Gt/mnopogon tends
to have inflorescence branches somewhat stag-
gered at the culm apex rather than in the ver-
ticillate arrangement so common to Chloris. In
this respect, Cliloris mollis is intemiediate be-
tween the two genera, with the upper spikes
more or less radiate but occasionally with a few
remote lower spikes. In Gi/mnopogon the sterile
floret is usually much reduced, often being rep-
resented only by a swollen area at the junction
of rachilla and awn. One species, G. foliosus,
however, does have a reasonably well-developed
sterile floret and bears a remarkable resem-
blance to Chloris mollis. Essentially then, we
are faced with the problem of which characters
to accept. I believe that by emphasizing pri-
marily the rather unusual vegetative character-
istics and inflorescence of such species of Gt/m-
nopogon as G. floridamts, G. cliapmanniamis.
etc., that the highest degree of homogeneit\'
may be established in the resulting group. In
such a system, Chloris mollis is still essentially
unclaimed, for its outward vegetative appear-
ance and inflorescence type do not fit well with
Gymnopogon. However, the well-developed
sterile florets of C mollis are certainly more
like the majority of Chloris species rather than
most of the Gymnopogon taxa.
Representative specimens examined: BRA-
ZIL: Bahia: Juazeiro, Dorsett and Popenoe
411h (US); Ceara: Crateiis, Swollen 4,481
(US); Iguatu, Swollen 4,399 (K. US); Fernando
de Noronha: Morro Francez, Simmons 9,263
(US); Maranhao: Municipio de Loreta, be-
tween Balsas and Paniaiba Rivers, Eiten and
Eiten 4,503 (K, US); Paraiba: Pombal, Picket
3,804 (US); Soledade, Pickel 3,851 (US); Per-
nambuco: Serra do Genipapo, A. Chase 7,686
(US); Pcsqueira, Pickel 1,662, 3,675 (NY, US);
Piaui: Fazenda Nacional, Sivallen 4,179 (NY,
US), 4,296 (K, US); Picos. Sivallen 4,228 (US);
Rio Grande do Norte: Santa Cruz, Swollen
4,558 (US); Angicos, Swollen 4,714 (K, US).
COLOMBIA: Bolivar Div.: Cartagena, Hitch-
cock 9,906 (US); Magdalena Div.: Santa Marta,
H. II. Smith 2,752 (F, K, MO, NY, US). COS-
TA RICA: Boruca, Pittier 3,675 (BR). CUBA:
Oriente Prov.: Caimenera Naval Station, Hiorom
and Ramsden s.n., 6 Feb 1919 (US). CU-
RASAO: near Willemstad, Britton and Shafer
2,906 (US). DOMINICAN REPUBLIC: Prov.
Barahona: Las Salinas, Fuertes 1,921 (US).
ECUADOR: Prov. Guayas: Chanduy, in littore
Maris Pacific!, Spruce 6,436, Apr 1862 (K, NY,
US); between Guayaciuil and Salinas, Hitch-
cock 20.056 (US);' Chongon, A.^plund 7,671
(US). GALAPAGOS ISLANDS: Charles Is-
land: near Black Beach, Howell .9,.3.55 (NY,
US), Svenwn 184 (US); Post Office Bay,
Howell 8,801 (US). GUATEMALA: Dep. Za-
capa: I mi W of Tcculutan, Stei/ermark 29,192
(US). HAITI: Anses-a-Pitre, Ekman 6,999 (G,
K, US); jean Rabel, Leonard and Leonard
12,581 (US). JAMAICA: Gordontown, Harris
11,515 (F, K, MO, US); near Kingston, Harris
12,212 (K, US). LESSER ANTILLES: Bon-
aire Island, Boldingh 7,208B ( US ) ; Aruba, Sur-
ingar 12 (US). PERU: Prov. Lambayeque: en-
tre Motupe y Olmos, Lopez 1,238, 2,463 (US);
Prov. Tumbes: entre Pocitos v Una de Gato,
Fcrreyra et al. 10,6.34 (US).' VENEZUELA:
Dep. Federal, Gran Roque, Gines 4,615 (US);
Anzoatequi, 14 km S of San Mateo, Garroni 53
(US).
15. CHLORIS LONGIARISTATA Napper, Kir
kia 3:117. 1963. (ISOTYPE: "Tangain ika,
Ngudu. R. R. Staples. 382. 1/4/1935."' K!)
Fig. 25, E-H.
Stoloniferous perennial 30 to 70 cm tall;
siieaths somewhat compressed, glabrous; ligule
a short ciliate cro\vn; blades up to 15 cm long,
3 to 6 mm wide, acute, glabrous below, sparsel)'
long-pilose above and on the margins, especially
near the base; spikes three to five. 3.5 to 12.0
cm long, appressed to slightly divergent; spike-
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Ciiloris (Gramineae)
47
Fig. 25. ChloTis quinquesetica and C. longiaristata. ( A-D ) C. quinquesetica. (A) habit, x 1/4; (B) spikelet,
partly dissected, x 10; (C) sterile florets, x 10; (D) caryopsis, x 10. (E-H) C. longiaristata . (E) habit, x
1/4; (F) spikelet, partly dissected, x 5; (G) sterile florets, x 5; ( H) caryopsis, x 5.
48
Bricham Young UNrv'EBsrrv Science Bulletin
lets slightly divergent, ca four per cm of the
scabrous rachis; glumes very narrowlv lance-
olate, pale, membranous, tapering to a delicate
point or a short awn; first glume 3.7 to 4.9
mm long, 0.2 to 0.4 mm wide; second glume
5.5 to 6.7 mm long, 0.3 to 0.5 mm wide; fertile
lemma dorsally compressed, 7.5 to 8.1 mm long.
0.8 to 0.9 mm thick, 1.0 to 1.5 mm wide, nar-
rowly lanceolate to elliptic, callus well devel-
oped, tapering, shaip pointed, densely bearded,
margins, sides, and keel glabrous except coarse-
ly scabrous near the apex, awn .30 to .35 mm
long, subtended by two short teeth; sterile
florets two or three, lowermost sterile floret
narrowly cylindrical, 2.0 to 2.7 mm long, 0.3
to 0.4 mm wide, glabrous below, sparsely
scabrous above, gradually narrowing into the
awn, awn 20 to 35 mm long, subtended by a
pair of deltoid teeth, rachilla joint well de-
veloped; upper sterile florets progressively re-
duced; car\()psis ca 3 mm long, ca 0.7 mm
wide, dorsal!)- flattened, shallowly trigonous.
Chloris longiaristata is most closely related
to C. prieiirii, from which it mav be separated
by its ven,- long awns, longer than those on
any other species of Chloris, and lack of ciliate
hairs on the upper lemma margins.
Specimens examined: TANZANIA: Mwanza,
Lloyd 4 (K).
16. CHLORIS QUINQUESETICA Bhide, Jour.
Proc. Asiatic Soc. Bengal n.s. 8:311. pi. 24.
1912. (HOLOTYPE: "Papadi Bassein,
growing on the bounds of rice fields in
semisalt lands. R. W. Bhide. .30-8-191 1."
K! ) Fig. 25, A-D; ,30.
Perennial, culms eri'ct to strongh stolonif-
erous and rooting at the lower nodes, to .50
cm tall; sheaths glabrous; ligule a tuft of hairs;
blades to 15 cm long, 2 to 4 mm wide; upper
surface glabrous to scabrous, lower surface
glabrous to loosely pilose; spikes three to eight
(rarely two, occasionally up to eighteen, as
indicated in original description), only slightK'
divergent, 1.5 to 5.0 cm long, decunent at the
base and united into a common axis bearing
a few spikelets; culms villous for 5 to 6 mm
below the spikes; spikelets densely imbricate,
averaging ca 20 per cm on the prominenth
hirsute rachis; glumes linear-lanceolate to ovate,
membranous, glabrous except for the scabrous
midnei-ve; first glume 1.5 to 1.8 mm long, 0.2
to 0.3 mm wide; second glume 2.2 to 2. .5 mm
long, 0.5 to 0.6 mm wide; fertile lemma 2.0
to 3.1 mm long, 0.8 to 0.9 mm wide, ovate,
keel and lateral nerves usually appressed-pubes-
cent, sometimes glabrous, margins densely
spreading-pubescent with hairs about 1 mm
long, callus densely bearded, awn 4 to 8 mm
long; sterile florets three or four; lowest sterile
floret tiuncate, 1.3 to 1.6 mm long, 0.5 to 0.6
mm wide, glabrous below but with a prominent
fringe of hairs along the upper margin, awn
3.5 to 6.0 mm long; upper florets enclosed by
the lower and progressively reduced, becoming
more inflated; carvopsis 1.4 to 1.6 mm long,
0.5 to 0.7 mm wide, ellipsoid.
Onlv two species of Chloris— C. (fuitvjuese-
tica and C. montana—\\Ave a dense fcltlike pu-
bescence just below the points of attachment
of the spikes. Chloris quinquesetica usuallv has
four sterile florets (occasionally three); the
lowermost sterile floret is ciliate along the
upper margin, and the upper sterile floret is
spherical and inflated (Fig. 25, A-D). On the
other hand, Chloris inontana usually has three
sterile florets. These are conunonK' all glabrous,
though occasionally sparsely pilose; and the
upper sterile floret resembles the lowest and
is not spherical and inflated (Fig. 27, F-I). A
comparison of C. qiiinqiiesetica and similar
species is shown in Table 8.
Specimens examined: INDIA: Kistna Dtr. :
Vijayarion, Barber 8,243 (K); Kannefalli,
Bourne s.n., without date, (K); Juhi, near Cawn-
porc, Diithie 7,7.59 (K); Cawnpore. Makldom-
pur, Mohbs s.n., Aug 19.33 (K); Sindi, Hooper
s.n., without date (K); Madh Island, Santapau
21,284 (K); Andra Pradesh, Secunderabad,
without collector or date (K).
17. CHLORIS WIGHTIANA Nees ex Steudel,
Syn. PI. Glum. 1:206. 1854. (ISOTYPES:
There are three sheets with apparentK' four
different collections mounted upon them at
K!, among which are undoubtedly dupli-
cates of the synt\pes cited bv Nees and,
subsequently, Steudel. Among these is a
specimen labeled "Herb. Wight. 1766.
Chloris Nxightiana NE. Wall s.n. .3809." The
label corresponds in many respects to the
specimen cited in the original description. )
Fig. 26.
Chloris incompleta Wight ex Steudel, Svn.
PI. Glum. 1:206. 1854. Pro sijn. Non'C.
incompleta RotJi, 1821.
Tufted to somewhat stoloniferous perennial,
10 to 30 cm tall; sheaths densely pustulose-
pilose; ligule a densely ciliate crown; blades up
to 4 cm long, ca 4 to 6 mm wide, densely pustu-
lose-pilose on both surfaces; spikes three or four,
1.0 to 2.5 em long, appressed or onlv slightly
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris ( Cr.^mine.\e)
49
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50
Brigham Young University Science Bulletin
Fig. 26. Chlom wightianii. (A) habit, \ 1/4; (B) leaf (left) and enlargement of blade (right) showing
pustulose-ba.sed hairs; (C) spilcelet, partly dissected, x 10; (D) sterile florets, x 15; (E) caryopsis, dorsal
view (left) and lateral view (right), x 10.
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Ciiloris (Gramineae)
51
divergent; spikeli't.s clen.sel\- imbricate, .spread-
ing widely from the pii.stulose-pilose rachis, ca
12 per cm of rachis length; glumes broadly
lanceolate, thin, membranous, glabrous except
for the midnei-ve, which is sometimes scabrous;
first glume 2.0 to 2.5 mm long, 0.2 to 0.3 mm
wide; second glume 2 to 3 mm long, 0.3 to 0.4
mm wide; fertile lemma 2.7 to 3.1 mm long,
0.9 to 1.0 mm wide, elliptic, callus, margins,
and keel denseh' and prominently spreading-
pilose, lemma sides pustulose, awn 7.5 to 10.0
mm long; sterile florets three or four, lowest
sterile floret broadly triangular, 1.4 to 1.5 mm
long, 1.9 to 2.0 mm wide, pustulose, midrib
densely pilose, awn 6 to 7 mm long, upper
sterile florets progressively reduced; caryopsis
ca 1.3 mm long, ca 0.9 mm wide, ca 0.5 mm
thick, ovoid-elliptic, strongly dorsally flattened.
Ciiloris wigJitiami may be easily separated
from all other species in the genus by its pustu-
lose pubescence; this is especially noticeable on
the sheaths and blades. Only Ciiloris arcnaria
is as pubescent, but it is not pustulose. A com-
parison with other similar species is presented
in Table S.
Only two collections in addition to the iso-
types mentioned above were studied in the
herbaria. One label indicated the plant was
collected on alkaline soil.
Specimens examined: INDIA; Peninsula In-
diae Orientalis, Wig/if without precise localit)',
date, or collection number (K); Tinnevelly Dtr.,
Tamparaparani Ri\er delta, Lumlararaj v.n., 28
Dec 1943 (K).
IS. CHLORIS MONTANA Roxburgh, Flora
Indica ed. Care>- 1:331. 1820. (T\pe not
found. There is, however, a colored plate
at K!, number 882 in the Roxburgh Flora
Indica drawings labeled "Chloris montana
Roxb." This plate clearly shows a villous
upper culm and a good spikelet dissection,
leaving no question about the plant to which
Roxburgh was referring.) Fig. 27, F-I, 28.
Andropo'^on tefru-aristiitiis Roxburgh, ex
Hooker, ' Flora British India 7:292.' 1896.
Pro si/n.
Chloris montana Roxburgh var. glatica
Hooker f. in Trimen, Flora Ceylon 5:276
1900. (ISOTYPE?: "from the herbarium.
Ceylon. Jaffna, abundant. H. Trimen 22. 2
ii 90." US! The collection data correspond
with those given in the original description,
but no collection number or date was gi\en.
The plant is small, possibly only a seasonal
dwarf; no spikelet differences were noted.)
Stoloniferous, occasionally tufted, perennial
from 20-50 cm tall; sheaths glabrous; ligule a
short ciliate crown; blades up to 12 cm long,
2-4 mm wide, glabrous below, glabrous to scab-
rous above; spikes two to five, 1.5 to 6.5 cm
long, slightly divergent; upper culms puberu-
lent; spikelets densely inserted, appressed, av-
eraging ca 15 per cm of the scabrous to sparsely
pilose rachis; glumes narrowly lanceolate, pale,
translucent, glabrous except for the scabrous
midnerve; first glume 1.5 to 1.7 mm long, ca
0.2 mm wide; second glume 2.2 to 2.7 mm long,
0.2 to 0.4 mm wide; fertile lemma 2.1 to 3.5
mm long, 0.5 to 0.7 mm wide, elliptic, upper
margins ciliate with hairs up to 1 mm long,
keel glabrous, sides glabrous to sparsely ap-
pressed-pilose, callus bearded, awn 4.2 to 9.5 mm
long; sterile florets usually three (rarely two or
four); lowest sterile floret 0.9 to 1.7 mm long,
0.4 to 0.6 mm wide, truncate, usually glabrous,
rarely with a few appressed hairs in the mid-
nerve region, awn 3.9 to 9.5 mm long; upper
sterile florets progressively reduced, temiinal
sterile floret elongated to subspherical, usually
only slightly inflated; caryopsis 1.7 to 2.0 mm
long, ca 0.3 to 0.4 mm wide, ellipsoid, trigonous.
Some specimens of Chloris montana have the
sterile lemma apex quite ciliate, though not as
densely as C. quinqttesetica, which they may
resemble in other respects (Figs. 27, F-I; 25,
AD).
Occasional specimens may approach C.
inflata. particularly in the number of sterile flo-
rets. Chloris montana usually has three sterile
florets, though sometimes only two. Likewise,
(.'. inflata usualh has only two sterile florets,
though some plants from the South Pacific
island archipelagoes regularly have three. How-
ever, C. montana is a stoloniferous perennial with
puberulous upper culms and without inflated
upper sterile florets (Fig. 27, F-I). Chloris in-
flata is usualh' a tufted annual though oceasion-
alh stoloniferous, and has glabrous upper
euims and inflated upper sterile florets ( Fig. .30,
A-M). Additional differences between these
and other similar species are presented in
Table 8.
Representative specimens examined: INDIA:
Nellore Dtr., Tada, Bourne 2,471 (K); Kistna
Dtr., Manilipatain, Madras, Gamble 17,396 (K);
Madras Dtr.: Dugarazpatam, Gamble 20,389
(K); Adzar, Gamble 20,784 (K); Chingleput
Dtr., Aradi, anon. IS Sept 1917 (K); Chitoor
Dtr., Panappakkam, without collector or date
(K); Pondicherry, Perrottet s.n., 1839 (K).
MAURITIUS: Round Island, Barkley?, without
52
BniGHAM Young University Science Bulletin
17. Chloris bournei and C. montana. ( A-E ) C. bouniei. (A) habit, x 1/4; (B) spikelet, partly dissected,
10; (C,D) sterile florets, showing variation, x 10; (E) caryopsis, x 10. ( F-I ) C. montana. (Â¥) habit,
1/4; (G) spikelet, partly dissected, x 10; (H) sterile florets', dissected, x 10; (1) caryopsis, x 15.
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chlohis (Gramineae)
53
Fig. 28. Distribution of Chloris montana. Inset A:
Mauritius, Reunion, and Round Island.
collection number or date (K). THAILAND:
HuaHin, Kerr 13,477 (K).
19.
CHLORIS BOURNEI Raiigachariar and
Tadulingam, Jour. Indian Hot. Soc. 2:189.
1921. (TYPE: MH, not seen. Original
description and accompanying illustration
Fig. 27, A-E.
arc clear.
Stoloniferous perennial up to 1 m tall, with
geniculate culms ascending from a tufted l)ase;
sheaths glabrous, ciliate near the membranous
ligule; blades up to 10 cm long, 5 mm wide,
glabrous, scabrous, or sparsely pilose above,
pilose on lower surface, especially at base; spikes
three to nine, 6 to 9 cm long; spikelets imbri-
cate, ca six per cm of the scabrous rachis;
glumes broadly lanceolate to ovate, glabrous
and membranous except for the scabrous to
hispid midnerve; first glume 1.8 to 2.5 mm long,
0.3 to 0.4 mm wide; second glume 3.2 to 3.7 mm
long, 0.4 to 0.5 mm wide; fertile lemma elliptic
to obovate, 3.0 to 3.7 mm long, 0.8 to 1.3 mm
wide, densely appressed-pubescent along callus,
margins, and keel, awn 3.5 to 7.0 mm long;
sterile florets two or three; lowest sterile floret
obovate, 1.5 to 2.8 mm long, ca 1 mm wide,
glabrous to sparsely pubescent on the midnerve,
somewhat inflated, often containing rudiments
of the palea, stamens, and pistil, occasionally
apparently fertile, somewhat inflated, awn 3 to
4 mm long; caryopsis ca 1.7 mm long, 0.7 mm
wide, ellipsoid-ovoid; chromosome number 2n
=40.
ChloTis bournei and C. inflata are similar,
though C. bournei has much larger spikelets
(Table 8; Fig. 27, A-E). Also, Chloris bournei
is a stoloniferous perennial, while C. inflata is
a tufted or stoloniferous annual.
Representative specimens examined: INDIA:
Coimbatore Dtr.: Talamalai Forest Reserve,
Jacob 100 (K); Coimbatore Botanical Garden,
Janaki-Ammal 1,172 (K); Bombay, Hubb, Koelz
19,901 (US); Arcot Dtr.: Ammapettai, banks of
Uppanera, Narayanaswami 4,171 (K); Tanjore
Dtr.: Ponnakani Meda, Narayanaswami 5,141
(K); Bellary Dtr.: Tornagallu, Rangachari s.n.,
II Aug 1901 (K); Madras Presidency: near Ah-
mednagar, Shattuck s.n., (K, US 879426); prope
Palamaltah, Wight 3,316 (US); Godaveri Dtr.:
Samalkot, anon., 19 Jan 1918 (K).
20. CHLORIS INFL.\TA Link, Enum. PI. 1:105.
1821. (FRAGMENT OF TYPE?: "Chloris
inflata Link. Hb. Link. Sem. ex California."
US! It is uncertain whether this is a frag-
ment of the type, but the original descrip-
tion does refer to seed having been sent by
Dr. Chamisso and that the origin was Cah-
fomia. The locality must be an error. For-
tunately, the original description clearly ap-
plies to the species described below.) Fig.
30, A-M.
Andropogon barJiatum Linnaeus, Mantissa
302. 1771. Noil A. barbatum Linnaetis Syst.
Nat. ed. 10. 2:1305. 1759. (For a full ex-
planation, see below and also under Chloris
(landi/ana.)
• = BOURNEI
A=QUINQUESETICA
Fig. 29. Distribution of Chloris bournei (circles I
C. quinquesetica (triangles).
and
54
Bricham Young University Science Bulletin
Fig. 30. Chloris inflata and C. formosana. (A-M) C. inflata. (A) hubit, x 1/3; (B) spikelet, partly dissected,
X 10; (C-G) fertile florets, showing variation, x 10; (H-L) sterile florets, showing variation, x 10; (M)
caryopsis, x 10. (N-Y) C. formosana. (N) spikelet. partly dissected, x 10; (O-S) fertile florets, showing varia-
tion, X 10; (T-X) sterile florets, showing variation, x 10; (Y) caryopsis, x 15.
Biological Series, Vol. 19, No. 2 Ta.xonomv of the Genus Chloris (Gramineae)
55
Chloris harbata (Linnaeus) Swartz, Flora
Indiae Occ. 1:200. 1797. ( Basrcl on An-
dropogon harhatum Linnaeus, 1771, but
not Andropogon harhatum Linnaeus, 1759. )
f^on Chloris barhata ( Linnaeus ) Nash, 1898.
Chloris longifolia Steudel, Syn. PI. Glum.
1:205. 1854. (HOLOTYPE:' 'Hrhi. Cum-
ing, nr. 696." P! FRAGMENT OF TYPE:
US!)
Chloris paraguuiensis Steudel, Syn. PI.
Glum. 1:204. 1854. (TYPE: "Rengger
legit in Paraguay." Not seen. Original de-
scription clearly fits the above species.)
Chloris harbata Swartz var. divaricata
Kuntze, Rev. Gen. PI. 2:771. 1891. (ISO-
TYPE: "Turong. Anam. Com. 2/1889." K!)
Chloris rufescens Steudel, Syn. Pi. Glum.
1:206. 1854. (Ti'PE: "Un'me legit in Ins.
Maurit." Not seen. The original descrip-
tion is not conclusive, but it seems to fit
this species best.) Non Chloris rufescens
Lagasca, 1805.
Annual 15 to 95 cm tall, erect to decumbent
and rooting at the lower nodes; sheatlis gla-
brous; ligule short, pilose, occasionally mcreh-
shortly cihate; blades flat, up to 15 cm long,
0.3 to 0.6 mm wide, glabrous, occasionally pi-
lose at the base; spikes seven to fifteen, 3 to 8
cm long, digitate, ± erect, often somewhat
fk'xuous and purplish; spikelets densely imbri-
cate, about 14 per cm of the scabrous rachis;
glumes lanceolate to narrowly ovate, hyaline,
glabrous except for the scabrous midnerve;
first glume 1.2 to 2.1 mm long, 0.1 to 0.3 mm
wide; second glume 2.3 to 2.7 mm long, 0.2 to
0.3 mm wide; fertile lemma 2.0 to 2.7 mm long,
0.4 to 0.7 mm wide, oxate to elliptic, callus and
upper margins spreading to appressed-pilose,
with hairs up to 1 mm long, keel glabrous to
pilose, sides of lemma occasionallv sparsel\ ap-
pressed-pilose, awn 4.0 to 7.7 mm long; sterile
florets usuall)' two (rarel\- three), lowest sterile
floret 0.9 to 1.3 mm long, 0.4 to 0.9 mm wide,
narrowly turbinate, tnmcate, inflated, usually
glabrous, occasionalh' sparsely appressed-pilose
on back, awn 5 to 7 mm long; upper sterile floret
obovoid to subspherical, markedly inflated,
about % the lengtli and width of the lower;
caiyopsis 1.1 to 1.4 mm long, 0.4 to 0.5 mm
wide, ellipsoid to obovoid; chromosome num-
ber 2n = 20, 40, ca 50.
Chloris inflata varies greatly in several char-
acters, including size, growth habit, spikelet
pubescence, and number of sterile florets. Ni'i-
ther discernible geographic patterns nor con-
sistent correlation of characters can be dis-
tin<Tuished, except that specimens from the island
archipelagoes of the southwest Pacific show a
high frequency of plants with three sterile flo-
rets. One collection with three sterile florets
was named C. harbata var. divaricata by Otto
Kuntze (1891), though he mentioned only that
the awn of the sterile floret was divaricate to
reflexed. The plant is essentially identical to
other populations of C. inflata in all other
respects.
Chloris inflata is similar to C. formosana
(Fig. 30, N-Y), C. montana (Fig. 21, F-I), and
C. quinquesetica (Fig. 25, A-D). It is readily
separated from the last two by being annual
and lacking a felt)' pubescence at the apex of
the culm just below the point of attachment
of the spikes. In addition, it has more spikes
(7-15) than Chloris montana (2-5). C. quin-
quesetica usually has four sterile florets; C.
inflata, two or three.
Detailed differences and a short key, sepa-
rating Chloris inflata and C. montana, are given
under the latter species. Table 8 presents a
comparison of this and other related species.
Considerable confusion has been evident as
to whether Chloris harbata (Linnaeus) Swartz,
C. inflata Link, or C. paraguatjensis Steudel
should be the accepted binomial. Certainly, the
last name is predated by the other two and must
be rejected. A full discussion of the relationship
of Andropogon harhatum Linnaeus (both 1759
and 1771), Chloris polydactyla (Linnaeus)
Swartz, and Chloris harbata (Linnaeus) Swartz,
has been presented in the treatment of C
dandt/ana and should be consulted (p. 34).
Essentially, I am rejecting C. harbata (Lin-
naeus) Swartz, for the basionym (Andropogon
barbatum Linnaeus, 1771) is a later homonym.
Swartz clearly referred to Andropogon barbatum
Linnaeus, 1771, when making the transfer; and
thus, C. harbata cannot be accepted as a new
name as proposed by previous authors. The
next available name is Chloris inflata.
Chloris inflata has been collected in a wide
variety of habitats throughout the warm temper-
ate, subtropical, and tropical regions of the
world (Fig. 31). Chloris inflata is often a
\veed in waste areas, but it may be found also
in cultivated fields, along beaches, etc. It ap-
pears to be very salt-tolerant: one gathering
from La Salinas, Dominican Republic {Howard
and Howard 8,411, NY! US!), was growing on
pure salt rock. Still other collections are from
sand dunes, loams, or limestone derivatives.
Representative specimens examined: (Note:
Chloris inflata is very well represented in major
herbaria and over 700 individual specimens
56
Brigham Young Universiti- Science Bulletin
Fig. 31. Distribution of Chloris inflata. Inset A; Hawaiian Islands
South Kennedy
Station, Adams
LANDS: Giaiul
were examined. The following list represents
only a small fraction of these collections. A com-
plete list is available from the author in mimeo-
graphed form.) ADMIRALTY LSLANDS: Bis-
marck Arch., Goodenough Island, Milne Bay
Dtr., Brass 24,419 (K).' ANTIGUA: near St.
John, Rose et al. .3,377 (NY, US). ARGEN-
TINA: Buenos Aires, Isla Manel, Parodi 740
(US). AUSTRALIA: Northern Territory: near
Darwin, Black 1:270.200 (K); Queensland:
Dtr.: 1.3 mi SE of Eaglefield
L079 (K). BAHAMA IS-
Turk Island, Millsixiii^h and
Millspaugh 9,002 (F, NY); Inagua, Hitchcock
s.n., 3 Dec 1890 (F, MO). BARBADOS: Bridge-
ton, F. W. Jolnison 1,194 (NY). BORNEO:
Balikpapan, Frost i (K). BRAZIL: Bahia: 46
km VVSW of Joazeiro, A. Chase 7.939 ( F, US);
Pernambuco: Bello Jardim, Serra do Genipapo,
A. Chase 7.698 (F, MO, US). BRITISH
GUIANA: Georgetown, Graham 345 (K). BUR-
MA: Mandalav, White 132-237 (US); Sagaing,
Huk s.n., 5 July 1890 (K). CANTON ISLAND:
without precise location, Degener and Hathaway
21,251 (MO, NY). CELEBES ISLAND: Mana-
do, Egvjan 1.732 ( K, US). CEYLON; Trincoma-
lee coast, Kingdon-Ward 23,064 (K). COCOS
KEELING ISLAND: St. John 26,465A (K).
COLOMBIA: Dep. Atlantico: entre Palmar de
Varelay Pondera, Dttgand 4.42S (US). CUBA:
Havana Prov.: without precise location. Baker
1,441 (F, US); Oriente Prov.: Guanta-
namo Bay, Britlon 2.0.32 (NY, US); Isla
de Pinos, Curtiss s.n., Apr. 1904 (NY).
CURASAO: near Willemstad, Curran and I la-
man 63 (NY, US). DOMINICA ISLAND: near
Morne Daniel, Hodge 3.893 (NY). DOMINI-
CAN REPUBLIC: Prov. Barahona: Neiba Val-
ley, Howard and Howard 8,360 (NY, US); Prov.
Monte Cristo: Guayubin, Abbott 876, 878
(US); Prov. Santo Domingo: vicinity of Ciudad
Trujillo, A//«rf/ 14,626 (NY). FERNANDO DE
NORONHA: Quixaba, Cutjler 9,395 (US). FIJI
ISLANDS: Viti Levu Island, Greenwood 1,213
1.308 (US). FRENCH GUIANA: Cayenne,
lloock s.n., 2 Jun 1962 (NY). GHANA': near
Labadi, Adams 3,835 (K). GRAND CAYMAN:
without precise locatitm, Moggs 61 (K). GREN-
ADINES: Petit Martinicjue, Howard 10.917
(NY). GUADELOUPE: Basse Terre, Dms
3,158 (NY, US). HAITI: Gonave Island, vicinity
of Etroite, Leonard 3,339a {Â¥, NY, US); Tortue
Island, vicinity of LaVallec, Leonard and
Leonard 11.262 (UC). HONDURAS: Dep.
Atlantida, Standleij 56,587 ( F, US). INDIA:
Madura Dtr.: Pulnev Hills, Dindigal. Anglade
1.109 (K); Chingleput Dtr.: Madnis, Gamble
16,415 (K); Calcutta, Heifer s.n., 18.36-.38 (BR,
NY); Coimbatore Dtr.: Anai Katti, Rapi and
Naganathan 4,802 (K). IVORY COAST:
Abidjan Plateau, Adjanohoun 428A (K).
JAMAICA: between Portland Point and
Rocky Point, limestone plain, Britton 1.903
(NY); Kingston, Harris 9,049 (F, K, NY, US).
BiOLOcicAi. Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
57
JAVA: Ambarawa, Brooks s.n., Mar 1909 (BM).
KENYA: Voi Dtr., Tsavo National Park, E of
Lugard Fall.s, Grceiucai/ ami Kamiri 12.S74
(K); Kwale, SW of Mombasa, Bogdan AB
3,194 (K). LINE ISLANDS: Christmas Island,
Pearson P22 (K). LIBERIA: Cape Palmas,
Hale 157 (US). MACAU: vacant lot, Hitchcock
19,153 (US). MALAGASY REPUBLIC: No,ssi
Be, Hildchramit 2.939 (US), 2,932 (K). MA-
LAYSIA: Singapore, Bor S61 (K); Selonger
State, Port Swettenham, BuckiU 12,839 (K).
MARSHALL ISLANDS: Guam, Fosberg 25,334,
31,223 (US); Pagan Island, Bonham 33 (US);
Saipan Island, Fosberg 31,287 (US); Tinian Is-
land, Fosberg 24,798 ( US ) ; Bikini Atoll, Taiilor
1,500 (UC); Eniwctok Atoll, Fosberg 24.407
(US); Kwajalein Atoll, Fos]>erg 31,202 (US);
Majura Atoll, Fosl>rrg 31.193 (US). MAR-
TINIQUE: St. Anne, S of Dunkertiue-
Catherini' mangrove, Ei^lcr 39—73 (NY).
MAURITIUS: Battc-rie Dauphine, Saner 2.701
(F). ME.\IC:0: Tamaulipas: N of Ciudad
Mante, Beetle M-521 (UC); Yucatan: Me-
rida, Swallen s.n.. Aug 1932 (US). MOLLUCA
ISLANDS (HALMAHERA): Morotai, Main
and Aden 1.477 (US). NEW CALEDONIA:
Anse Vata, McKee 4.063 (K). NEW PROVI-
DENCE: Nassau, Matthews 61 (K). NIGERIA:
Lagos, Ward 9 (K). PALAU ISLAND: Ara-
kabesan Island, Ilosaka 3.354 (US); Peleliu Is-
land, Fosberg 31,952 (US). PANAMA: near
Miraflores, Locks, Allen 1,752 (F, K, MO, NY).
PAPUA: Daru Island, Brass 6.403 ( BM, US);
Fairfax, 9°55'S, 147°20'E, Gillison NGF 22,211
(K); SW base of Mt. Lawes, Eucalyptus savan-
nah, Sclwdde 2.644 ( K, US). PEMBA IS-
LAND: Vaughn 252 (BM). PHILIPPINE IS-
LANDS: Luzon: Manila, Clemens s.n., Sep 1924
(UC, US); Corregidor, Cunung 696 (K, MO).
PUERTO RICO: Mona Island! Sardinera, Brit-
ton et al. 1.831 (NY, US); Isla de Culebra,
Brown and Wheeler 141 (NY, US); Guanica
Bav, A. Chase 6,529 (US). RYUKYU ISLANDS:
Okinawa, Amano 6,800 (US). ST. KITTS:
Hitchcock 16,369 (US). ST. LUCIA: Brooks 20
(US). SAMOA EAST: Tutuila Island, Collarnio
508 (US). SERAM (CERAM) ISLAND: Am-
boina, Robinson 1,647 (BM). SEYCHELLES
ISLANDS: Mahe Island, Boivin s.n., 17 Nov
1855 (K). TAIWAN: Chiayi, Devol 7,095 (UC).
TANZANIA: Dar Es Salaam, Hitchcock 24.424
(US); Tanga Pro\-.: Sakura Sec: Panxani
Dtr., Tanner 2,027 (K, UC). THAILAND:
Bangkok, Kerr 3,761, 7.853 (BM); Wangka,
Kwae Noi River basin, Wichian 330 (K)."tI-
MOR: Kupang, R. Brown s.n.. Apr 1S0.3 (BM).
TOBAGO: Spexside, Purseglove P. 6,358 (K).
TRINIDAD: Woodbrook, Broadwai/ 7.177 ( F,
MO). UNITED STATES: Hawaii: Oahu, N of
Koko Head, Degener 10,835 (NY); Texas:
Hidalgo Co., McAllen, roadside, Anderson
3,875 \USC). VENEZUELA: Lara, Palmasola,
Pittier 6,385 (US). VIRGIN ISLANDS: St.
Croix, Thompson, 5 Oct 1921 (NY, US); St.
Thomas, Hitchcock 16.320 (US); Tortola, Velez
3.123 (K). WAKE ISLAND: Peale Islet, Fos-
berg 34.928 (US). WEST IRIAN (IRIAN
BARAT): Merauke River, Kelapi Lima, McKee
1.705 (K). ZANZIBAR: Cooke s.n.. 1861 (MO).
21. CHLORIS FORMOSANA (Honda) Keng,
Claves Gen. Sp. Gramineamm Primarum
Sinicanun 197. 1957. (Based on Chloris
barbata Swartz var. jormosana Honda.) Fig.
30, N-Y.
Chloris harhata Swartz \ar. formosana
Honda, Bot. Mag. Toho 40:4.37. 1926.
(SYNTYPES: "Taiwan: 'Takao, G. Naka-
hara, no. 6.35, anno 1905." "Baehu, B. Ha-
yata anno 1919." Not seen. Description re-
fers to species as commonl)' understood.)
Perennial often stoloniferous, 30-75 cm tall,
erect to decumbent; sheaths glabrous, rarely
sparsely pilose; ligule a very short ciliate fringe;
i)lades up to 20 em long, usuaii\' folded, gla-
brous, acute; spikes five to nine, 4 to 8 cm long,
erect to somewhat lax; spikelets densely imbri-
cate, ca fourteen per cm of the pilose rachis;
glumes lanceolate, glabrous, except for the
scabrous midnerve; first glume 1.0 to 1.5 mm
long, ca 0.2 mm wide, acute; second glume
2.2 to 2.5 mm long, 0.2 to 0.3 mm wide, apex
obtuse, mucronate; fertile lemma 2.3 to 3.0 mm
long, 0.5 to 0.7 mm wide, with a lateral groove,
which is occasionally appressed-pilose, upper
margins denseh' pilose with hairs up to ca 1
mm long, keel glabrous, slightly gibbous, awn
2.0 to 5.7 mm long; sterile florets two; lower
sterile floret oblanceolate to truncate, 1.6 to
1.9 mm long, 0.6 to 0.7 mm wide, flattened or
only slightl\- inflated, awn 2.3 to 5.0 mm long;
upper sterile floret of same shape as lower, but
smaller, 0.5 to 0.7 mm long, 0.4 to 0.5 mm wide,
glabrous, awn 1.5 to 2.5 mm long; car\'opsis
1.2 to 1.6 mm long, 0.3 to 0.4 mm wide, ellip-
soid.
Chloris formosana is most easily confused
with C. inflata. The following key will separate
the two:
1. Lower sterile floret 1.6 to 1.9 mm long,
flattened or only slightb' inflated; upper
sterile floret about one-third the size of
58
Brigham Young University Science Bulletin
the lower, scarcely inflated; upper
sheaths, ligules, and Iowit portions of
leaves glabrous, rarely sparsely pilose;
perennials.
C. fonnosana (Fig. 30, N-Y)
1. Lower sterile floret 0.9 to 1.3 mm long,
usually inflated; upper sterile floret two-
thirds to aljout the same size as the
lower, usually inflated and spherical; up-
per sheaths, ligule, and lower portions of
the leaves usuallv loosely pilose, though
sometimes sparsely so; annuals, occasion-
ally rooting at the lower nodes.
C. inflata (Fig. 30, A-M)
Other species that might be confused with
C. formosana are compared in Table 8.
Collected from sandy or gravelly soils, often
near the ocean in China, Hong Kong, Taiwan,
and Vietnam (Fig. 32).
Representative specimens examined: CHINA:
Fukien Prov.: Amoy, Chung 6,055 (US), Price
1,363 (K); Kwangtung Prov.: Taio, Chun 3,-
114B, 3,123 (NY); Pakhoi, Hitchcock 19,255
(US); Hainan Island: Hoihow, Hitchcock
19,547 (US), McChire 10,451 (BM, K), Tsan^i
986 (NY). HONG KONG: Bor H.K. 76 (K),
Ford s.n., 25 Aug 1893 (K, NY), Hance 1.943
(BM); Quarry "Bay, Sampson 851 (BM).
NORTH VIETNAM: Tonkin, route de Haiphong
a Doson, bords des chemins, Petelot 425 (NY,
US). TAIWAN: Takao, Henri/ s.n., 1S93-4
(BM), Henry 1,023 (K), Plaijfair' 53 (K); with-
out precise location, Tonaka 10.352 (BM NY
UC).
22. CHLORIS PILOSA Schumacher, Beskr.
Guinciske Planter 75. 1827. (PHOTO-
GRAPHS OF HOLOTYPE: K! US! Type
specimen, in Isert and Thonning Herbarium,
C, has mucronate lemmas.) Fig. .33, E-K.
Fig. 32. Distribution of Chlaris formosana (southeast
China, Hainan, and Taiwan).
Chloris breviseta Bentham in Hooker f.,
Niger Flora 566. 1S49. (SYNTYPES: "Cape
Coast, Don" BM! "Cape Coast, Vogel" K!)
Chloris nigra Hackel. Bol. Soc. Broteriana
21:179. 1908. (TYPE: "Insula S. Jacobi
Prom Viridis, pr. Trinidade, leg. A. Bar-
jona" not seen, but description is clear.)
Chloris virgata Swartz var. breviseta (Ben-
tham) Pilg(>r e.x Peter, Beih. Repert. Sp.
Nov. 40:262. 1931. (Based on Cliloris
breviseta Bentham. )
Annual often weakly rooted, sometimes
shortly stoloniferous, erect or in stoloniferous
specimens somewhat decumbent, usually 30 to
70 cm tall, occasionalK' to 2 in; sheaths gla-
brous to denscK' pilose; ligule densely and prom-
inently pilose; blades 2 to 10 mm wide, up to
.30 cm long, scabrous; spikes five to nine, 3
to "^j cm long; spikelets loosely imbricate, ca five
to seven per cm of the scabrous rachis; glumes
membranous, une(jual; first glume 1.1 to 1.6
mm long, 0.2 to 0.3 mm wide, narrowly lanceo-
late, glabrous except for the scabrous mid-
nerve; second glume 1.9 to 2.3 mm long, 0.3 to
0.4 mm wide, glabrous with scabrous midnerve,
rounded at the apex and bearing an awn ca 0.3
mm long; fi'rtile lemma 2.3 to 3.5 mm long,
0.7 to 1.5 mm wide, broadly ovate or elliptic
in overall outline, prominently gibbous on the
keel; var\ing from pale to dark gra\ or black,
often mottled when mature, callus bearded,
sides with a prominent glabrous or appressed-
pilose furrow, apex acute to blunt, mucronate,
or with an awn up to 6 mm long; steiile florets
two (very rarely one) glabrous, becoming
scabrous at the apex, lower sterile floret 1.5 to
2.2 mm long, 0.7 to 1.1 mm wide, narrowed at
the base, becoming inflated at the truncate
apex, glabrous below, apex scabrous, mucronate,
or \\ ith an awn up to 3.0 mm long, upper sterile
floret a triangular inflated rudiment less than
1 mm long, ca 1 mm wide, usually hidden in
the lower sterile floret, awnless; anthers ca 0.4
to 0.5 mm long; caryopsis 1.3 to 1.5 mm long,
ca 0.5 to 0.6 mm wide, obovoid, trigonous;
chromosome number 2n = 20, 30.
Awn length in this species varies widely.
Most specimens fall in one of two groups, those
with awns less than 1 nun long, generally only
mucronate (represented on the map in Fig.
34 by unshaded circles) and those with longer
awns, usually from 2 to 6 mm (indicated on
the map by shaded circles). No strong correla-
tions of awn length can be made with geo-
graphic origin, nor with other morphological
characters. While the variation in awn length
BiOLOCiCAi. Series. Vol. 19. No.
Taxonomy ci- the Genus Ciilohis (Gramineae)
59
Fig. 33. Chloris cnstilloniuud and C. pilosa. (A-D) C. castilloniami. (A) habit, x 1/5; (B) spikelet, partly dis-
sected, .\ 10; (C) sterile floret, \ 15; (D) caryopsis, \ 10. (E-K) C. pilosa. (E) habit, .x 1/3; (F) spikelet
of awned variant, partly dissected, x 10; (G) spikelet of unawned variant, x 10; (H) sterile florets of a\vned
variant, x 15; (I) sterile florets of unawned variant, x 15; (J) upper sterile floret, x 15; (K) caryopsis, x 10.
60
Bbicham Vounc University Science Bulletin
produces plants with somewhat diverse appear-
ances, tlieir nomenclatural recognition is not
warranted.
Chloris pilosa most closely resembles C. vir-
gata, from which it may be separated in having
prominently pilose ligules and broader lemmas
which have shorter marginal cilia.
Widely distributed in equatorial Africa,
Chloris pilosa has been collected in weedy areas
and as a native in savannahs. It is occasionalh'
cultivated as a forage grass. The natural distri-
bution is shown in Fig. 34.
Uepresentative specimens examined: BEL-
GIUM (adventive): Dunquenjue, anon. (US).
CAMER(HTN: Boro, 35 km ENE of Maroua,
Bounougou 13 (K); Darga, 30 km ESE of Ma-
roua, Koechlin s.n., May 1965 (K); Victoria Dtr.;
roadsides, weedy, Maitland 9 (K). CAPE
VERDE ISLANDS: San Nicalao, Loive s.n., 22
Feb. 1864 (BM); Sao Tiago, Grandvaux-Bar-
hosa, 5,755 (K). CONGO: LeopoldviHc, Fas-
seaux 1,019 (K). DAHOMEY: Parakou, Ris-
opoulos 1,197 (K). ETHIOPIA: Serilia, Ghattas.
^chucinfurth 2,283 ( K, US). FERNANDO PO;
Mann 118 (K). GAMBIA: Sankuli-Kunda,
Pirie s.n.. Sept 1933 (K). GHANA: Nungua.
Accra Plains, 5°40'N, 0°06'W Ankrah 20,150
(K); 2.5 mi SW of Kpetchu Ferr\ . 7°55'N.
0°22'E, Ankrah 20,387 (K); Baikpa, 6°51'N,
0°26'E, Rose-Innes 31,189 (K); Axim, Vigne 395
(NY, US). GUINEA: Nzerekore, Bald-
win 9,722 (K, US); Kouroussa, Pobequin 546
(K). IVORY COAST: Kouiblv, Adjanolwun
413A (K). LIBERIA: Grand Ba.ssa Co.: Timbo,
Baldivin 11,223 (K); 10 mi inland from River
Cess, Baldwin 11.228 (K); Sinoe Co.: Sasstown,
Baldwin 11,603 (K). MAURITANIA: Dahr,
16°30'N, 7°W, Rossetti 61/172 (K). NIGERIA:
• SPIKELETS AWNFD f:; " • -C v.^ '"Af
O SPIKELETS AWNLESS/ A- '\) °»T^
ORMUCRONATE \ ii<.„_ <V ft- ■■■>
Fig. 34. Distribution of Chloris pilosa. Dots represent
specimens with awned lemmas; open circles repre-
sent specimens with unawned or merely mucronate
lemmas.
Prov. Oyo; Ibadan, Brenan 9,601 (K, US);
Prov. Zaria: Samaru, Freeman S. 116 (K); Jebba,
llagerup 739 ( K, US ) ; Jos Plateau, along De-
limi River, Naraguta, Laivlor and Hall FHl
46,540 (K); Toro, 20 mi S of Jos. Semple 176
(US). REPUBLIC OF THE CONGO: Musa,
De Giorgi 1,299 (K); Eala, Germain 1,695
(K); Kasenye, Johnston 1,042 A (K); Bukama,
Shantz .565 '(K). SUDAN: Cordofan Prov.: El
Obeid, Harrison 75 (K); Darfur Prov.: Jebel
Marra, Wickens 2,279 (K); Equatoria Prov.:
Lado Dtr., Cartwright 27 (K); Upper Nile
Prov.: Zerat Island, Harrison 1,005 (K). SIER-
RA LEONE: Mano, Deighton 674 (K); Musaia,
Deighton 5,386 (K); Freetown, Deighton 2,029
(K). SENEGAL: Dakar, Baldwin 5,714 (K).
TANZANIA: Tanga Dtr.: Kange Estate, Faulk-
ner 808 (K); Rufiji Dtr.: Utete boma, Milne-
Redhead and Taylor 7,537 (K); Morogoro Dtr.:
Duturi, Nicholson 29 (K). TOGO: Palime,
Stage 73 (K). UGANDA: Buganda Dtr.,
Hitchcock 24,934 (K); Lango Dtr.: Amugo,
]ohn.ston 1,037 (K); Ankole Dtr.: Kiruhara,
Thornton 39 (K); near Mt. Elgon, Snowden
1,207 (K).
23. CHLORIS MRGATA Swartz, Flora Indiae
Oce. 1:203. 1797. ( Holotype not .seen, but
description clearly refers to the species as
commonh' understood.) Fig. 35.
? Chloris pubescens Lagasca, Var. Cienc.
Lit. Artes (Madrid) 4:143. 1805. Descrip-
tion recopicd in Lagasca, Gen. Sp. Pi. 5
Nov. 1816. (A specimen, at G!, labeled
"Chloris pubescens Lagasca, ex Peruvia, La-
gasca 1807" is Chloris virgata. The original
description is not conclusive, and this name
is included here provisionall)'. )
Rahdochloa virgata (Swartz) Beauvois, Ess.
Agrost, 84, 158, 176. 1812. (Based on
chloris virgata Swartz, but Rahdochloa not
x'ahdly published.)
Chloris elegans Humboldt, Bonpland,
Kunth, Nov. Gen. .Sp. PI. 1:166. /)/. 49.
1816. (HOLOTYPE: "Mexico, Queretaro.
M..\. Bonpland n. 4194." P! Fragment in
US! )
C/(/,)r;.s nlba Prc-sl, Rel. Ilaenke 1:289. 1830.
( Description clearly refers to Chloris vir-
gata. Scribner, 1899, examined and de-
scribed the presumed type in the Bemhardi
Herbarium, saving it was the same as C
elegans. )
Chloris caudata Trinius ex Bunge, Mem.
Savans Etr. Acad. Petersburg 2:144. 1835.
BiOLOCiCAi. Sehies, Vol. 19, No. 2 T.•\xONo^I^ (U- the Genus Chlouis (Gramineae)
61
Fig. 35. Chloris virgata. (A) habit, .x 1/4; (B) spikelet. partly dissected; (C-F) fertile lemmas, showing
variations; (G-1) lower sterile florets, showing variation; (J) rudimentary palea of lower sterile floret (left)
and attached upper sterile floret (right); (K) caryopsis, lateral view; (L-N) caryopses, dorsal views, show-
ing variations. (B-N) x 10.
62
Brigham Young Universiti' Science Bulletin
(Description, combined with geographical
citation, "Hab. propre Pekinum," seems
to refer to C. virgata. A specimen in the
Cosson Hb., at P!, collected by Bunge in
1835, and labeled "Chloris caudata," per-
haps in Bunge script, is C. virgata.)
Chloris penicillata Willdenow ex Steudel,
Nom. Bot. ed. 2. 1:353. 1840. Pro si/n.
Chloris decora Nees ex Steudel, Syn. PI.
Glum. 1:205. 1854. First published as
novien nudum by Nees in Royle, Illust.
Bot. Himalayan Mts. and Flora Cashmere
1:416. 1840.' (HOLOTYPE: "Rovle Hrbr.
nr. 87" K! )
Chloris meccana Hochstetter ex Steudel,
Syn. PI. Glum. 1:205. 1854. (ISOTYPES:
"802. Chloris meccana Hochst. ex Steudel
... d. 11 Febr. 36, leg. W. Schimper" G!
US!)
Chloris iiwdagascariensis Steudel, Syn. PI.
Glum. 1:203. 1854. (HOLOTYPE: ". . . Le
Jolis legit 1849 . . ." P! )
Chloris notocoma Hochstetter, Flora 38:204.
1855. (ISOTYPES: "Hb. abyss. Hohenacker
nr. 2125" G! P! )
Chloris nmltiradiata Hochstetter, Flora 38:
204. 1855. (Description is of C. virgata.
Type, "Hb. abyss. Buch. nr. 486" in P! is
C. prietirii, a duplicate at K! is C. virgata. )
Chloris alba Presl var. aristulata Torrey,
Rept. Expl. Survey Mississippi River Pacific
4:155. 1857. ( A ' short-awned variant.)
Chloris hrachijstacltiis Andersson in PetiTs,
Naturwiss. Reise Mossambique 556. 1864.
(Based on Chloris alba Presl.)
Agrostomia barl)ata Cervantes, Naturaleza
(Mexico City) 1:.346. 1870. (TYPE: Cuer-
na\aca, Mexico, not seen, but description
fits C. virgata. )
Chloris barbata var. decora (Nees) Ben-
tham. Flora Australiensis 7:613. 1878.
(Based on C. decora Nees.)
Chloris alberti Regel, Acta Horti Petropoli-
tani 7:6,50. 1881. (TYPE: "Mongolia occi-
dentali prope Takiansi, A. Regel," not seen.
Geographic origin and description seem to
suggest C. virgata.)
Chloris tetrastachtjs Hackel ex Hooker f.,
Flora British India 7:291. 1896. Pro syn
Chloris virgata var. elegans (Humboldt
Bonpland Kunth) Stapf in Thistleton-Dver,
Flora Capensis 7:642. 1900. (Based on C.
elegans. )
Chloris pohjdactijla Swartz ssp. nmltiradi-
ata (Hochstetter) Chiovenda in Pirotta,
Annuario Rc-ale 1st. Bot. Roma 8:54. 1903.
(Based on C. multiradiata Hochstetter).
Chloris gahriclae Domin, Biblioth. Bot. 85:
.368, Fig. 83. 1915. (HOLOTYPE: "Queens-
land; am Flinders River bei Hughenden,
haufig, Domin, II. 1910." K!, on extended
loan from PR.)
Chloris rogeoni Chevalier, Rev. Bot. Appl.
Agric. Trop. 14:127. 19,34. (SYNTYPES:
"Dire. 43,871, El Oualadjii 4,3,937 et 42,-
,357." All except the last at P! Fragments
of last two at K! )
Chloris tibestica Quezel, Bull. Soc. Hist.
Afrique Nord 48:84. 19,57. (TYPE: "Emi
Koussi," not seen. Description and subse-
quent illustration in Quezel, 1958, leave no
doubt as to identity.)
Annual, extremely variable in size, ranging
from a few centimeters to over a meter; culms
usually tufted, several arising from one root
system, occasionalU' stoloniferous; sheaths usu-
ally glabrous, occasionalK' rather densely pilose
toward the apex; ligule glabrous, or with a cili-
ate fringe of hairs up to 4 mm long; blades up
to 30 cm long and 1.5 cm wide, usually gla-
brous witli scabrous margins, occasionally pilose;
spikes four to twenty, 5 to 10 cm long, usu-
ally more or less erect; spikelets densely imbri-
cate, averaging 10 per cm of the scabrous or
hispid-ciliatc rachis; glumes pale brown, lanceo-
late, glabrous with scabrous midnerves; first
glume 1.5 to 2.5 mm long, 0.2 to 0.4 mm wide;
second glume 2.5 to 4.3 mm long, 0.3 to 0.5
mm wide; fertile lemma 2.5 to 4.2 mm long, 0.7
to 1.3 mm wide, pale brown, occasionally
puiplish or blotched with dark brown spots,
keel usually prominently gibbous, keel, mid-
ner\'es, and lower margins glabrous to promi-
nently pilose, margins long-ciliate, with spread-
ing hairs near the apex, awn 2.5 to 15.0 mm
long, usualh' more than 5 mm long; sterile flor-
et one, on robust specimens, occasionally two;
lower sterile floret 1.4 to 2.9 mm long, 0.4 to
0.8 mm wide, awn ,3.0 to 9.5 mm long; upper
sterile floret, when present, usually borne on a
short rachilla and greatU' reduced, sometimes
absent, leaving only the naked rachilla; caryop-
sis 1.5 to 2.0 mm long, ca 0.5 mm wide, ellip-
soid-trigonous; chromosome number 2n=20,
26. .30, 40.
Without doubt, Chloris virgata is the most
variable of all of the annual species in the
genus. Great variation is shown in many traits,
including such vegetative features as height, leaf
size, tomentum, and habit, as well as spikelet
Biological Series, Vol. 19, No. 2 T.axonomy of the Genus Chloris (Gramineae)
63
characters (especially various pubescence on
the lemma, such as presence or absence of hairs
on the keels, midnerves, and margins). In addi-
tion, the lemmas themselves ma\ be variously
gibbous, keeled, or merely folded at the mid-
nerve. Only two attributes seem constantly cor-
related: (1) the conspicuous tuft ot spreading
hairs on the upper lemma margins and (2) the
appressed to erect or only slightly divergent
spikes.
No well-marked patterns of variation can be
identified, though several somewhat cr)'ptic
populations are noticeable. Most specimens
from Australia and some from Asia and Africa
have relativel)' narrow lemmas that are not con-
spicuously gibbous, though other collections
from the same general area may show the
typical prominently gibbous keel. Some of the
populations, especially those from Tibet and
Nepal, ma\' eventually prove to be distinct; but
presently, so few specimens are available for
study that it seems advisable to treat the entire
variable population as a single specie's. By con-
trast, western hemisphere material seems to be
quite uniformly gibbous, usualh' conspicuously
so.
Occasionalh' novelties, such as a specimen
collected by Dr. T. L. Bancroft at Falm Island,
Queensland (K!), may be very different. This
particular specimen was unique in having dense-
ly pilose glumes. In other respects it is similar
to typical specimens of C. virgata and is consid-
ered here as merely an aberrant individual.
Chloris virgata closely resembles C. gaijana
and C. pilosa in spikelet characters. It may be
easil)- separated from the former in being an
annual and from the latter in having relatively
long awns.
CIdorif; virgata lias Ijcen widel\- collected
from many habitats, primarily in warm temper-
ate to tropical areas, but extending well into
temperate areas where hot summers are com-
mon (Fig. 36). Over 1,300 different specimens
were examined in this study. The following list
represents only a small fraction of the total.
Representative specimens examined: AF-
GHANISTAN: Helmand River Valley, irrigated
field. Long 326 (US). ANGOLA: Kaoko veld,
N of Ohopoho, ca 1S°E x 14°S, de Winter and
Leistncr 5,177 (K). ARGENTINA: Prov.
Buenos Aires: Pergamino, Parodi 834 (US);
Piov. Gatamarca: 6 mi SW of Andalgala, Bartlett
20,226 (US); Prov. Cordoba: Dep. dc Santa
Maria, Ihmziker 1.294 (MO); Prov. La Rioja:
Patguia, Laliitte ij Castro 2 (US); Prov. Men-
doza; Santa Rosa, Jen^en-Haartip s.n., 1904-05
(US); Prov. Salta: Dep. Rosario de Lerma, Las
Tres Acquias, Montenegro 455 (US); Prov. Tu-
cuman: Dep. Trancas, Rio Tipainago, Venturi
4,354 (NY, US). AUSTRALIA: Queensland:
Darling Downs, Ashcroft s.n., May 1916 (K);
Petrie, Blake 170 (K); Western Australia: Wan-
Fig. 36. Distribution of Chloris virgata. Inset A: Hawaiian Islands
64
Brigham Younc University Science Bulletin
dagee, Minilya River, Gardner 3,234 (K). BO-
LIVIA: Cocliabamba Dep.: without precise lo-
cation, Btichtien 2,512 (US); Dep. Tarija, near
Entre Rios, West S,250 (MO, US). i«)TSWA-
NA: Kgalagedi, without precise location, Leist-
ner 3,103 (K); Ngwato, Bokalaka area, Francis-
town, McClintock s.n., 18 Dec 1966 (K). BRA-
ZIL: Bahia: near Rio Sao Francisco, Joazeiro,
A. Chase 7,946 (F, US); Ceara: Campo Salles,
Swallen 4,2HH (US, Piaui: between Floriano and
Oeiras, Swallen 4,171 (US); Rio Grande do
Norte: Santa Cruz, SuaUcn 4,849 (US). BUR-
MA: Mingaladon, Pokhant 1,375 (K). CHINA:
Honan Prov.: Cheng-Chou, Chiao 18,500 (F);
Hopeh Prov.: near Peking, Chcn^ 2.038 (US);
Inner Mongolia: Ordos, Tapiichan, Hsiu II 3,799
(US); Kansu Prov.: Yao Kai, near Lichen, Ching
258 (US); Kiangsu Prov.: Nanking, Chiao
12,916 (K, NY, UC); Manchuria: Darien, /»//«;!«
â– 59a (NY); Harbin, Dorsett and Dorsett s.n..
(US); Shungari River, Komarov 167 (BM, K);
Shansi Prov.: Sui Guan Dtr., I Ian Ngan 32
(UC); Shantung Prov.: Cliingtao, Li Chuan,
Chiao 2,864 (F, K, NY, UC, US); Shensi Prov.:
Chou-chih-hsien, Knng K3,734 (US); Yunnan
Prov. La-Kov, Mairc Apr 1914 (BM, F). CU-
RAgAO: Sovonet, Suringar 9 (US). ECUA-
DOR: Prov. Guavas: W of Cluavacjuil, .\sphin'J
15,369 (K); Puntilla, Salinas, ' 2 IS'S, 81 W,
Svenson s.n.. 1 Mar 1941 ( F, MO); Prov. Loja:
between La Tonia and Loja, Hitchcock 21,403
(US). EL SALVADOR: vicinity of San Salva-
dor, StancUcij 19.234 (NY). GALAPAGOS IS-
LANDS: Charles Island, Post Office Ba\ , lloucll
8,838 (US); Chatham Island: Wreck Bav, Ilou-
eU 8..'i95 (US), Stewart 1,254 (K, MO); Indefati-
gable Island, Howell 9,917 ( US ) ; North Sc'smour
Island, Howell 9.967 ( US ) ; South Se\'mour Is-
land, Howell 9,924 (US). GUATEMALA: near
Jalapa, Kellerman 7,975 (F, NY); vicinitv of
Zacapa, Standlei/ 74.606 (F). HONDURAS: 5
km N of Talaiiga, Johanncssen 892 (UC); Dep.
Morazan: vicinitv of El Zainorano, Standlei/ s.n..
27 June 1949 (F). INDIA: Allahabad, Dudgeon
s.n., 1 Nov 1919 (MO); Bombay, Gamble 21,-
028 (K); Central Pro%'.: Nagpur, Haines 3.609
(K); Assam, 21 mi marker on Ledo road, 27°
20'12"N, 96°2'55"E, Juan L8C93 (US); Bihar
Prov.: Ranka Koelz 18,971 (K, US); Shi-
moga, Mysore, Meehold 10.495 (K); Gwalior
St.: Wisner 35 (US). KENYA: Esageri
Station, Nakuru to Eldama Ravine Road,
Bogdan AB2,014 (K); 12 mi N of Magadi,
Clayton DC56 (K); 20 mi E of Isiolo, Stewart
474 (K); Nairobi, Webster K33 (K). KOREA:
in campis Chinampo, Faurie 1,244 (BM); Pyeng-
yang. Smith s.n., 5 Sept 1935 (US). LESOTHO:
Lvr\be, Dieterlen 6,326 (K). MEXICO: Aguas-
calientes: 2 mi S of Aguascalientes-Zacatecas
line, Emery 182 (TEX); Baja Cahfornia: Magda-
lena Island, Brandegee s.n., 21 Jan 1889 (UC);
57 km NW of San Ignacio, Carter, Alexander,
and Kellogg 1,950 (K, UC, US); Chihuahua: 2
mi N of Lucero, Weber and Charette 11,604
(UC); Coahuila: W of Puerto de las Monjas,
I.M. Johnston 8,640 (MO, US); Colima: Alzada,
Hitchcock 7.054 (US); Distrito Federal: Ixtapa-
lapan, Matuda 25,685 (MO); Durango: 3 mi N
of Donato Guerra, Emery 337 (TEX); Jahsco:
Guadalajara, La Barranca, M.E. Jones s.n., 19
Nov 1930 (MO, UC); Guanajuato: Acambaro,
Hitchcock 6.932 (US); Guerrero: near Iguala,
Canon de la Mano Negra, Rose 9,385 (NY, US);
Michoacan: 26 mi SE of Sahuayo, Pratt 699
(TEX); Morelos: Cuernavaca, Hitchcock 6,835
(US); Navarit: Tepic, Acaponeta, Rose 14,248
(NY, US)'; Oaxaca: I.xtepec, Fisher 35,311 (F,
MO, NY); Puebla: Tehuacan, Hitchcock 6,073
(US); Queretaro: 5 mi from Queretaro, Pratt
774 (TEX); San Luis Potosi: 6 mi S of Mate-
huala, McGregor et al. 516 (US); Sinaloa: San
Ignacio, Ortega 4.468 (US); Sonora: 25 mi W
of Angostura, Santos 1,831 (F, K); Tamaulipas:
near Aldoma, Martinez and Luyando F-2,177
(TEX); Veracruz: 6 to 7 km N of Tierra Blanca,
Santos 3,313 (NY); Yucatan: Izamal, Gaumer
1,085 (F, MO, NY, US); Zacatecas: 30 mi N of
Zacatecas, Emery 307 (TEX). MADEIRA IS-
LANDS: near Funchal, Sledge s.n., 14 Apr
1949 (BM). MALAWI: Zomba, Cormack 176
(K); Domasi, between Zomba and Ncheu,
Jackson 132 (K). MARIANAS ISLANDS: Sai-
pan, Kagman Peninsula, Fosberg 31.288 (NY,
US). MAURITANIA: Atar Parielle, Adam
21.799-1 (K). MOgAMBIQUE: Prov. Mo9am-
bique: Mocuba, Namozoa, Faulkner 29 (K);
Lourenyo Marques Dtr.: near Costa do Sul,
Gomes e Sowm 3. 438 (K). MONGOLIA: Sha-
harakh Usu, Chancy 407, 537 (NY, UC, US);
llOmiNWofHaHin, EnTcsson 478 (US). NE-
PAL: Rohagan, Suli Gad, Polunin. Si/kcs and Wil-
liams 3,37-3. 3.376 (BM). NICARAGUA: road to
Granada, Hitchcock 8,706 (US). NIGERIA:
Bornu, Gudumbali, Rains s.n., Oct 1961 (K).
PAPUA: 9 mi NW of Ore Bay, Reeder 831
(US). PARAGUAY: chaco boreal, Ro/os 7,724
(US). PERU: Prov. Apurima: 20 km N of
Abancay, Stork et al. 10,539 (UC, US); Prov.
Lima: Huarochiri, Weberbauer 5,287 (F); Prov.
Avacucho: entire Pucjuio v Nazca, Fcrreyra
.5,52.3 (K, US); Prov. Piura:'l km W of Talara,
Beetle 26,197 (F, K, UC, US); Prov. Tumbes:
Contramirante Villar, Ferreyra 12,214 (US).
REPUBLIC OF THE CONGO: Kasenye,
Biological Series, Vol. 19, No. 2 Taxonomy ov the Genus Chloris (Gramineae)
65
Johnston 1.042, 1,084a (K); Katanga, Tumbwe,
Stjmoens 8,382 (K). SAUDI ARABIA: SO to
90 km N of Aden, Wahah 1,904 (US). SENE-
GAL: without precise location, Adanson 74
(BM). SEYCHELLES ISLANDS: Long I.sland,
Sqxiihbs Gil (K). SOCOTRA ISLAND: Bal-
four 341 (K). SOMALI REPUBLIC: Ca'ad
Forest Reserve, 7 mi W of El Afwein, Boaler
B 130 (K); Erigavo Dtr.: Hubcra, McKinnon
S249 (K, US). SOUTHERN RHODESIA: Shan-
gani Dtr.: Gwampa Forest Reserve, Goldsmith
48,828 (K); Lower Sabi Dtr.: Rattraij 1,230
(K); Miami Dtr.: termite mound, K.34 Experi-
mental Farm, Wild 1,773 (K). SOUTHWEST
AFRICA: Avis Dam, Windhoek, Liehenberg
4,446 (UC, US); Grootfontein, Schoenf elder
88 S494 (K). SUDAN: Darfur Prov.: Nyorlete,
12°5S'N, 24°4'E, Blair 11 (K); Wadi Toro,
13°6'N, 23°56'E, Blair 144 (K); Jebel Mara,
Zalingei, Wickens 1,786 (K); Kassala Prov.: be-
tween Berber and Suakin, Schiieinfurth 399,
572 (US). TANZANIA: Tumba, Bullock 2,415
(US); Mwanza Township, Cannidiael 1,160
(K); Mbulu Dtr.: near Mdala River, Lake Man-
yara National Park, Greenuaij and Kanuri 11,-
239 (K); Dar-es-Salaam, Marshall 50A (K);
Monik Plateau, above Lake Natron, Newbould
6,206 (K): 7 mi NE of Old Shimanga, Welch
62 (K). TIBET: Lhasa, Richardson s.n., autumn
1946 (K). UGANDA: Jinja, Hitchcock 24,954
(K, US); Karamoja, 16 mi W of Moroto, Lang-
dale-Brown 1,586 (K). UNION OF SOUTH
AFRICA: Sunday River, N of Monke\' Ford,
Burchell 2,862 (K); Zoutpansberg, Kruger Na-
tional Park, Codd 5,407 (K); 4 mi N of Kroon-
stad, 27°3.5'S, 27°5'E, Schecpers 1.266 (K); IS
mi E of Pietersburg, van Viewen 1,612 (K).
UNITED ARAB REPUBLIC: Jebel Elba, Wadi
Kasisrob, Shabetai 6 (K). UNITED STATES:
Alabama: Cliambers Co., Langsdale, Banks
306 (US); Arizona: Chiricahua Mts., Paradise,
Blumer 1,715 (K, MO, NY); Sulphur Spring Val-
ley, Forbes 1,634, 1,643 (NY, US); Toro'" Can-
yon, Baboquivari Mts., Gilman 9 (NY); Pima
Co., Wilmont Road, Tucson, Ginter s.n., 22
Sept 1943 (UC); Santa Cruz Co., Patagonia,
Hitchcock 3,645, 3,664 (US); Ajo, M.E. Jones
24,795 (MO, NY); California: Fresno Co., along
Hwy 180, 5 mi W of Sanger, Bacigahipi 2,487
(UC); Merced Co., S of Newman, Mason and
Smith 8,223 ( UC ) ; Riverside Co., Ford Dry Lake,
24 mi W of Blythe, Grinncll 1.081a (UC); Yolo
Co., 3 mi N of Davis, along HWY 99 W, Crainp-
ton 3,140 (UC). Colorado: Powers Co., road-
side near Holly, Harrington 740 (UC, US);
Hawaii: Oahu, Honolulu, Kaimuki, A. Chase
12,677 (US); Kansas: Stevens Co., E of Hu.go-
ton, McGregor 17,016 (KANU); Louisiana:
beach. Lake Ponchartrain, C.A. Brown 2,387
(US); Maine: North Berwick, wool waste, Par-
lin 1.516 (US); Maryland: Canton, chrome ore
piles. Reed 32,714 (US); Massachusetts: Mill-
burv, wool waste. Gates 31,785 (US); Missouri:
Courtney, Bush 9,733 (MO); Nevada: Lincoln
Co., between Crystal Springs and Ash Springs,
Train 2,415 ( NY)'; New Mexico : Grant Co., Man-
gas Canyon, 16 mi WNW of Silver City, Barkletj
14,732 (NY); Sierra Co., Animas Creek, Met-
calfe, s.n. (MO, NY); New York: Yonkers
wool mill, Bickncll. .v.n., 4 Sept 1898 (NY);
North Carolina: Guilford Co., W of Greens-
boro, Blomquist 1,962 (F); North Dakota: Nor-
ton Co., Mandan, Stevens 2,649 (UC, US);
Oklahoma: Payne Co., Stillwater, Henson 343
(UC); South Carolina: Myrtle Beach, Blom-
quist s.n., 22 Aug 1930 (US); Texas: Brewster
Co., 9 mi S of Marathon, Ferris and Duncan
2,841 (MO); Hemphill Co., 5 mi E of Canadian,
Rouell, Jr. 4,266 (TEX); Jeff Davis Co., 8 mi
S of Ft. Davis, W.V. Brown 3,269 (TEX); Utah:
0.5 mi N of south entrance, Zion National Park,
Harrison 11,092 (UC); Whipple Exped., Camp
60, Ft. Smith to Rio Grande, Bigelow s.n.,
1853-4 (NY); Wright Exped., western Texas
to El Pa.so, Wright 762 (MO, NY, US). VENE-
ZUELA: vicinity of Caracas, Bailey and Bailey
201 (NY, US).' ZAMBIA: Zambra, Namuala-
Pemb road, Astle 2,107 (K); Abercorn, Siame
608 (K), Trapnell 1,735 (K); Monze, Lochinvar
Ranch, van Rensburg 1,238 (K); 10 mi NE of
Serenje, Vasey- Fitzgerald 2,956 (MO). ZANZI-
BAR: Hitchcock 24,460 (US).
24. CHLORIS GAYANA Kunth, Rev. Gram.
1:89. 1829, in key; redescribed 2:293, /;/.
.58. 1830. ( Holotypc not seen, but Kunth's
complete description and plate are of the
species as commonly understood.) Fig. .37,
38.
C'hloris ahi/ssinica Hochstetter ex Achille
Ricliard, Tentamen Florae Abyssinicae
2:406. 1850. (DUPLICATE OF SYN-
TYPE; "Schimperi iter Abyssinicum, sectio
• tertia, ISOO . . . =nr. 79. Sect. I, prope
Djeladcranne in vallibus. V.i. 1844." BM!
K!)
Chloris glahrata Andersson in Peters, Nat-
urwiss. Reise Mossambiquc Bot. 2:557.
1864. ( Holotype not seen, but description
clear. )
Chloris gayana ssp. Chi. [sic] oligostachys
Barratte and Murbeck in Murbeck, Acta
66
BiucnAM Young Univehsity Science Bulletin
Fig. 37. Chloris gatjana (A) habit, x 1/6; (B) spikelet, partly dissected; (C-F) florets, showing variation in
size and shape of fertile lemma and size, shape, number, and sexual condition of sterile florets; (G) lower-
most sterile floret; (H) middle sterile floret; (I) uppermost sterile floret; (J) caryopsis. (B-J) x 10.
Univ. Lund II. Sect. 2. 36:8. pL 13. 1900.
(The authors clearly refer to this as a new
subspecies, yet they interjected an abbre-
viation of the generic name, thereby chang-
ing the relative order of taxa. The descrip-
tion and plate are of a variant witii small
spikclets.)
Eustacliys gayana Mundy, Rhodesia Agric.
Jour. 14:142. 1922. (Presumably based on
Chloris gaijami Kunth, but basionym not
cited. )
Chloris gayana f. oligostachys (Murbeck)
Maire and Weiller, Flore Afrique Nord
2:204. 1953. (Based on Chloris gayana
ssp. oligostachys Murbeck.)
Clitoris gayana f. genuina Maire and Weil-
ler, Flore Afrique Nord 2:204. 1953.
( Based on Chloris gayana Kunth. )
Biological Series, Vol. 19, No. 2 Ta.xonomy of the Genus Chloris (Gramineae)
67
Perennial up to 3 m t;xll, usually stolon-
iferous, sometimes tufted, erect; sheaths gla-
brous to scabrous, often ciliate apicallv; ligule
ciliate; blades up to .30 cm long and 1.5 cm
broad, scabrous; spikes nine to thirty, 8 to 15
cm long, usualh' somewhat divaricate; spikelets
pale, rather densely imbricate, ca 10 per cm
of the scabrous rachis; glumes lanceolate to
narrowly ovate, scabrous, especially on the
nerves; first glume 1.4 to 2.8 mm long, 0.3 to
0.4 mm wide, acute to awn-tipped; second
glume 2.2 to 3.5 mm long, 0.3 to 0.6 mm wide,
acute to awn-tipped; fertile lemma 2.5 to 4.2
mm long, 0.7 to 1.0 mm wide, ovate to obovate
to elliptic, pubescence variable, margins usually
with a prominent tuft of hairs near the apex,
occasionally appressed-pilose for full length, in-
terner\es usually glabrous, sometimes scabrous
or appressed-pilose, lateral nerves usually gla-
brous, occasionally appressed-pilose, somewhat
gibbous, callus bearded, awn 1.5 to 6.5 mm
long; sterile florets two to four (rarely one).
the lowest 2.2 to 3.2 mm long, 0.5 to 0.8 mm
wide, similar to the lemma, but more cylindri-
cal, staminatc or barren, awn 0.8 to 3.2 mm
long; upper florits awnless or awn-tippcxl,
progressi\'elv smaller \\ith the terminal reduced
to a turbinate cup; earvopsis 1.0 to 1.5 mm
long, 0.5 mm \\'ide, ellipsoidal; chromosome
number 2n = 20, 30, 40.
This species is the most variable of the
perennial species in the genus. Both stolonifer-
ous and nonstoloniferous plants are found, with
the former much more common. Individuals
vary greatly in size, though plants less than
0.5 m are uncommon. Spikelet moipholog)' is
inconstant. Lemma pubescence varies from
glabrous to densely pilose. The number of ster-
ile florets ranges from two (veiy rarely one)
to four, with the lowest "sterile" floret usually
staminatc, occasionally sterile. The second low-
est floret is usually sterile; upper florets are
always so. Twenty-one spikelets randomly
chosen from one collection (Wickens 2,715)
showed five different combinations of floret
number (two or three) and sexual condition
( male or barren).
A number of variants have been named. Of
these Chloris ahijssinica Hochstetter ex Achille
Richard has been mostly widely used, though
geneialK not recenth'. Duplicates of a syntype
were examined (BNI, MO, K); these did not
differ markedly from the variation spectrum
exhibited by the remaining specimens that were
examined. Stapf (19.31) provided a short key
to Chloris aht/ssinica and C. gaijona, separating
them on the presence or absence of stolons and
the length, sexuality, and number of sterile
florets. These characters were studied on a rep-
resentatixe number of specimens collected
throughout Africa, but neither discontinuities
in measurements nor major patterns could be
distinguished.
There is a possibilit\ of a very casual cor-
Fig. 38. Distribution of Chloris gaijana. Inset A; Hawaiian Islands.
68
Bricham Young University Science Bulletin
relation with ecological conditions, for tnfted
plants were frecjuently, but not alwa\s, col-
lected troni salt pans.
Representative specimens examined: (A
total of over 800 different specimens of this
species were studied. The following list repre-
sents only a small portion of this group.) AN-
GOLA: Huila Dtr. : Namaculungo, Teixera
1,407 (BM). ARGENTINA: Prov. Cordoba:
Cruz del Eje, ViUafarie 2,413 (US); Prov. Cor-
rientes: Dep. Lavalle, Estancia "La Pastrol"
Pedersen 3,864 (NY, UC, US); Prov. Salta:
Metan, Cabrera 3,089 (NY); Prov. Tucuman:
Trancas, Venturi 6,480 (US). AUSTRALIA:
New South Wales: Kenebri, 11 mi N of Bara-
dine, Constable 15,978 (UC); Northern Terri-
tory: near Darwin, Allen 67 (K); Queensland:
Conjuboy, Blake ami Lazarides s.n., 26 Feb
1954 (US); Kenmore, Clemens s.n., 18 Ma\
1943 (US); Western Australia: Latham, Moore
K197 (K). BOTSWANA: Koudoum Valley, 30
km E of Southwest Africa border. Wild and
Drummond 7,056 (BM). BRAZIL; Minas
Geraes: Lavras, A. Chase 8,827 (F, VIO, NY,
US); Rio Grande do Sul: Mono Santa Ana,
Rambo 41,982 (US). COLOMBIA: Pasto,
Medcllin, Rivera 131 ( K ) ; vicinity of Medellin,
Toro 299 (NY, US). COSTA RICA: Turrialba,
Leon 286 (MO). CUBA: Prov. Ilerradura:
Pinar del Rio, Ekwan 14,108 (NY). EASTER
ISLAND: Mataveri, waste land, Chaplin 1,041
(NY). ECUADOR: Prov. Guayas: Milagro,
Acosta-Solis 70,693 (US); Prov. Pichincha:
Valle del Guaillabamba River, Acosta-Solis
21,259 (US), A.'iplund 7.036 (K). ETHIOPIA:
Ocule Casai, Pappi 2,081 (MO, NY, UC); 10
mi N of Assale, Se/nple 2 ( US ) ; Metahara.
Semple 10 (US). GUATEMALA: near Chimal-
tenango, Standlei/ 80,483 (F, K). HAITI: plaine
du Nord, Ekman H. 9,890 (K, US). HONDU-
RAS: Dep. Morazan: Zamorano, Rodri'^nez 103
(F), 3,277 (F, MO, US); Standlei/ 1,767a,
13,035, 21,041 (F), 18,979 (F, NY, US); Wil-
liams and Molina 12,216 (F). INDIA: Mysore,
Bangalore, Koelz 19,898 (UC); Salem Dtr.;
Hosen Cattle Station, Naratjanaswami 3.028
(K). JAMAICA: St. Andrew Parish, Richmond
Park, Harris 12,707 (F, K, MO, NY). KENYA:
Klboko, Bogdan AB 2,239 (UC); Longonot,
Hitchcock 24,828 (K, US); Nakuru, Hitchcock
25,075, 25,082 (K, US); between Oljoro-o-Nyon
River and Lake Naivasha, Mearns 636 ( US ) ;
between Narok River and Lake Naivasha,
Mearns 628 (NY, US). LORD HOWE IS-
LAND: Binoleij 25 (K, UC). MALI: El Oual-
adji. Chevalier 43,927 (K); Tombouctou, Olaf-
sen 205 (US). MALAWI: Southern Nvasa Dtr.:
Dambo, Namwera Road, Ballard C.h.7 (K);
Chikwawa Dtr.: Bra.^s 17,996 (MO, NY, US);
Kasita River near Ekwendeni, ]ack.son 366 (K);
Port Herald Dtr.: Phipps 2,606 (MO). MAU-
RITIUS: Curepipe, Vaughan 1,844 (K). MEX-
ICO: Morelos: Cuenievaca, Lijonnet 2,807
(US). MOgAMBIQUE: Nya.ssa Dtr.: Ama-
ramba, 22 km de Nova Freixo, Torre and Paiva
10,624 (K). NEW ZEALAND: near Tauranga,
Hodgkins 70,536 (US). NIGERIA: Prov.
Bornu: Lake Chad, Daggash 24,970 (K); Ngala
Dtr.: Wulgo, Daveij 2 (K); Rann Dtr., Daveij
s.n., 17 Apr 1947 (K); River Kalkala, SW corner
of Lake Chad, Golding 76 (K). REPUBLIC
OF THE CONGO: Kasenve, Johmton 1,057
(US); Albertville, Under s.n., 23 Jan 1927 (K).
RWANDA: Pare National Kagera, Lcbnm 9.482
(K); Biumba, Mutara, colline Nyaka Kanga,
Troupin 8,813 (K). SIKKIM: Gangtok, Dr.
Bors collector s.n.. 15 Jul 1945 (K)."sOMALI
REPUBLIC: Farquahorson s.n.. Sept 1931 (K).
SOUTHERN RHODESIA; Katasi Reserve,
Bikita, Cleghorn 177 (BM). SOUTHWEST
AFRICA: Omuramba Khaudum, ca 18M°S,
20'i°E. de Winter and Marais 4.667 (K); Oka-
vango, at Bagani pontoon, Kaprivi, de Winter
and Wiss 4345 (K). SUDAN: Zalingei, Robert-
son 10 (BM); Dafur Prov.: Nyertiti, Robert-
son 22 (BM); Jebel Marra, Dariba Crater,
Robertson 140 (BM, K); Equatoria Prov.: Nim-
ule, Shantz 907 (US). SYRIA: Ali al Gharbi,
Graham 360 (BM). TANZANIA: Mbeye Dtr.:
Rujewa, Anderson 1,117 (K); Musoma, Em.<ion
232 ( US ) ; Ngari Roiigi plain, western slopes of
Mt. Kilimanjaro, Grecnwatj 6,741 (K); Ai-usha
Region, Ol Doingo Sambu, Greenwaij 4,413
(K); Moshi Dtr.: Sanya River, Haarer 1,097
(K); Rukwa, Lea LR8 (K); Banagi, Serengeti
National Park, Lcippert S603 (K);' 45-65 mi S
of Dodoma, Lijnes 76a (US); Selous Game Re-
serve, Nicholson 48 (K); Ngorongoro Crater,
Paulo 317 (K), Piemiesel and Kepliart 506
(US); Kilwa, Rifiji, Schlieben 2.541 (K); Lindi
Dtr.: Lutamba, Schlieben 5,880 (K); Monduli,
near Arusha, Semple s.n., Mav 1945 (US);
Olduvai, Williams 681 (K). THAILAND: Hua
Hin, lones s.n., June 1931 (BM). TONGA-
TAPU ISLAND GROUP: 'Eua, Houma, Set-
chell and Parks 15,431 ( UC, US). UNITED
STATES: Arizona: Yuma Valley, Fochtman 70
(NY); California: Merced Co., N of Los Banos,
Mason and Smith 8,226 (UC); Florida: Her-
nando Co., Brooksville, McFarlin 6,072 (US);
Hawaii- Kiholo, A. Chase 12.688 (US); Mis-
sissippi: Lumberton, Real s.n., 10 Jun 1920
(US); Texas: Port Lavaca, Allen 17 (US); Kle-
berg Co., .5.5 mi S of Kingsville, Corij 51.303
(NY, UC); Hidalgo Co., 0..5 mi N of Edinburg,
Ferris and Duncan 3.064 (MO, NY, US); Rock-
Biological Series, Vol. 19, No. 2 Ta.xonomy of tiie Genus Chloris (Gramineae)
69
port, Fisher 41.145 (US); Kleberg Co., King
Ranch, Lundell 14.877 (US); Cameron Co., Port
Isabel Road, Rimi/on 4,138 (MO, NY).
UGANDA: Old Entebbe, Darker 257 (UC);
Luzinga, Ditmmer 4,351 (US); Mbulamuti,
Hitchcock 24,975 (US); Tororo, Swamp, Hitch-
cock 24,980 (US); vicinity of Kabula Miiliro,
Mearns 2,532 (US); Mbalc, near Mt. Elgon,
Smusden 1,098 (US); S of Namasagali to Ka-
muli Road, Bugabula City, Wood 948 (MO).
UNION OF SOUTH AFRICA: between Sellen-
dam and Riversdale, Godfreti SI I 1,312 (VC.
US); near Pietermaritzburg, thornveld, Godfreij
SH 1,912 (US); Mposa, Godfreti and Baijer SH
1,463 (US). URUGUAY: Dep. Montevideo:
Osten 19.923 (US). VENEZUELA: Merida, La
Mesa, Uanhurti-Tracij 16 (K). ZAMBIA:
Mweru Wantipa swamp, 8°50'S x 29°40'E,
Tijrer 206 (BM).
25. CHLORIS CASTILLONIANA Lillo and
Parodi e.\ Parodi, Pli\sis 4:176. 191S.
(TYPE: not seen. OriginalK collected by
Castillon in the Calchaqui Valley, Argen-
tina. The original description
panving illustration positi\'el\
plant.) Fig. 33, A-D.
Caespitose perennial up to 1 m tall; she
glabrous; ligule short-ciliate; blades 10-20
long, 4 to 5 mm wide, glabrous; spikes eight to
fifteen, 9 to 11 cm long, apprcssed to strongly
ascending; spikelets densely imbricate, ap-
pressed, about eight per cm of the scabrous
rachis; glumes lanceolate, thin, membranous,
glabrous except for the scabrous midnerve;
first glume 1.6 to 2.0 mm long, 0.3 to 0.4 mm
wide; second glume 2.9 to 3.9 mm long, 0.4 to
0.6 mm wide, sometimes deHcately mucronate;
fertile lemma 3.5 to 4.7 mm long, 1.0 to 1.3 mm
wide, lanceolate-elliptic, rather prominently gib-
bous, callus flattened and elongate, densely
bearded, lemma margins densely ciliate with
hairs 1 to 2 mm long, keel glabrous, occasionally
an appressed-pilose groove present along the
lemma side, awn 2.2 to 2.5 mm long, arising
from well below the acute apex; sterile floret
one, 1.5 to 2.0 mm long, ca 0.4 mm wide, gla-
brous, narrowly obovoid, the apex somewhat
truncate, awn arising subapically, ciu-ving up-
ward, awn 1 to 2 mm long; caryopsis ca 2 mm
long, 0.5 mm wide, ellipsoidal, somewhat tri-
gonous.
Chloris castilloniana is most closely related
to C virgata, from which it may be separated
by its perennial habit as well as in having promi-
nent cilia along most of the lemma margin ( Fig.
and aeeom-
id(ntif\' the
iths
cm
33, A-D; 35). Chloris virgata is an annual with
the cilia restricted to the upper portion of the
lemma margin.
The few specimens that have been collected
are from sandy savannas in Argentina (Fig.
39, triangles).
Specimens examined: ARGENTINA: For-
mosa Territory: orillos del arroyo Ferreyra,
Asp 20 (US); Las Lomita, Parodi 8,455 (NY,
US); Prov. Catamarca: Mollecito, 20 km S of
Andalgala, Bartlett 20,239 (US); Dep. Belen,
Las Faldas, Lillo 45,625 (NY, US); Prov. La
Rioja: Patquia, Lahitte y Castro 3 (US).
26. CHLORIS ORTHONOTON Doell in Mar-
tins, Flora Brasiliensis 23:64. 1878. (ISO-
TYPES: "Glaziou n. 3610" BR! K! US!)
Fig. 39; 40, A-E.
Chloris guaranitica Parodi, Revista Argen-
tina Agron. 28:106. Fig. 2. 1961. (ISO-
TYPE: Argentina: Corrientes, Dep. Mbu-
rucuya, Estancia Santa Maria . . . Pederson
no. 3742." K! )
Stoloniferous, erect perennial to 70 cm tall;
sheaths glabrous, occasionally pilose near the
ligule; ligule prominently pilose; blades 0.5 to
A<ASTILLONIANA
• =ORTHONOTON
Fig. 39. Distribution of Chloris orthonoton (dots) and
C. castilloniana (triangles).
70
Brigham Young University Science Bulletin
0.7 mm wide, up to 25 cm long, pilose near the
base, otherwise glabrous to scabrous; spikes
four to eleven, 4 to 10 cm long, in several verti-
cils near the culm apex; spikelets eight to ten
per cm of the scabrous rachis, densely imbricate,
appressed; glumes lanceolate, acuminate, gla-
brous except for the scabrous midnerve; first
glume 1.6 to 2.4 mm long, 0.2 to 0.3 mm wide;
second glume 2.3 to 3.6 mm long, 0.3 to 0.5 mm
wide; fertile lemma 3.0 to 4.5 mm long, 0.7 to 1.6
mm wide, narrowly lanceolate to elliptic, mar-
gins long-ciliate, especially near the apex, keel
glabrous to appressed-pubescent, internerves
with a longitudinal groove bearing one or more
rows of dark glandular hairs, callus bearded,
awn 7.5 to 20.0 cm long; sterile floret one, 1.5
to 2.2 mm long, 0.3 to 0.5 mm wide, cylindric-
turbinate, glabrous, awn 3.5 to 9.0 mm long;
anthers ca 0.4 mm long; caryopsis ca 1.7 mm
long, 0.4 mm wide, narrowly ellipsoid, trigonous.
Chloris orthonoton is closely related to C.
aristata. Table 9 and Fig. 40 show the main
differences between the two.
Occasional specimens of Chloris orthonoton
may resemble C. vir^ata. The latter, however,
is a fibrous-rooted annual, lacks glandular hairs
in the lateral groove of the lemma, and has a
prominent tuft of cilia only along the upper por-
tion of the lemma margin.
A variant of this species, lacking pubescence
on the midnerves of the lemmas and having
somewhat coarser spikes, has bei'ii named
Chloris isiiaranitica Parodi (1961). In all other
respects the plant is identical with typical C.
orthonoton, and I do not feel it wortlu of
nomenclatiual recognition.
Representative specimens examined: AS-
CENSION ISLAND: roadside near airfield,
Duffey 1/83 (US). ARGENTINA: Prov. Cor-
rientes: Dep. Mburucuya, Parodi 3,742 (BR).
BRAZIL: Prov. Bahia: Baliia, A. Chase 7.871.
8,033, 8,051 (US); Feira de Santana, A. Chase
8,060 (US); Prov. Ceara: Acude, Municipio de
Maranguape, Drouet 2,180 (F, US); Camocim
to Granja, Swallen 4,634 (US); Martinopolis,
Swallen 4,641 (US); Minas Geraes: Lavras,
Black 887B ( US ) ; Juiz de Fora, A. Chase 8,625
(US); Sao Miguel, NW of Fomiiga, A. Chase
10,544 (US); between Sucupira and Omega, A.
Chase 11,162 (US); Paraiba: Soledade, Cesan
692 (US), Pickel 3,827 (US); Pernambuco: Bel-
lo Jardim, A. Chase 7,700 (F, US); vicinity of
Recife, A. Chase 7,720 (F, NY, US); Tapera,
Pickel 81 (F); Rio de Janeiro, NW of Cabo
Frio, A. Chase 10,122 (US); Niteroi, A. Chase
10,037 (US); Rio Grande do Norte: Estremoz
to Natal, Swallen 4,773 (US); Nova Cruz to
Montanhas, Stcallen 4,822 (US); Santa Cruz,
Sirallen 4,861 (US); Rio de Janeiro, C.laziou
17,363, 1888 (NY, US).
27. CHLORIS ARISTATA (Cervantes) Swallen,
North Aim-r. Flora 17:,596. 19.39. (Based
on Agrostomia aristata Cervantes. ) Fig. 40,
F-J.
Agrostomia aristata Cervantes, Naturaleza
(Mexico City) 1:345. 1870. (TYPE: "San
Angel. San Agustin de las Cuevas." Not
seen, but description is clear.)
Stoloniferous perennial with erect culms to
80 cm tall; sheaths glabrous; ligule a short-
ciliate crown; blades up to 15 cm long, 5 mm
wide, usualK glabrous or scabrous, occasionallx'
the upper surface pilose; spikes four to eight,
4.0 to 8.5 cm long, erect, somewhat appressed,
radiate; spikelets appressed, densely imbricate,
ca five to eleven per cm of scabrous rachis;
glumes narrowly lanceolate, glabrous t-xcept for
the scabrous midnerve; first glume 1.8 to 2.3 mm
long, 0.2 to 0.4 mm wide; second glume 3.0 to
4.1 mm long, 0.2 to 0.5 mm wide; fertile lemma
3.4 to 4.5 mm long, 0.8 to 1.0 mm wide, lanceo-
late to elliptic, apex acute, margins appressed-
pilose below, the hairs slightly spreading above,
keel usualK' glabrous, occasionally with a few
appressed hairs, midnerves glabrous, callus
bearded, awn 6.0 to 12.0 mm long; sterile floret
one, 1.6 to 24 mm long, 0.6 to 0.7 mm wide,
awn 5 to 7 mm long; caryopsis 1.5 to 1.9 mm
Table 9. Comparison of characteristics of Chloris orthonoton and C. aristata.
Species
Ligule
Lemma margin pubescence
Lemma internerves
C. orthonoton
C. aristata
long-ciliate ( ca 2
mm long)
short-ciliate ( less
than 1 mm long)
cilia more than 1.5 mm long; ap-
pressed basally, becoming spreading
apically
cilia less than 1 mm long; appressed
most of length or only slightly spread-
ing apically
lateral groove glandular-pubes-
cent; otherwise glabrous
lateral groove and internerves
glabrous, occasii)iuillv ap-
pressed-pubescent, but not
glandular
Biological Seiuks, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gbamineae)
71
Fig. 40. Chloria orthonoton and C. aristiita. (A-E) C. orthonoton. (A) habit, .x 1/4; (B) spikelet, x 5; (C)
florets, ,x 5; (D) sterile floret, x 5; (E) caryopsis, x 7.5. ( F-J ) C. arhtata. (F) habit, x 1/4; (G) spikelet,
X 5; (H) florets, x 5; (I) sterile floret, x 5; (J) caryopsis, x 7.5.
long, ca 0.5 nini wide, ellipsoid, trigonou.s;
chromosome number 2n = 100.
Differences between Chloris aristata and C.
orthonoton have been discussed previously.
This species has occasionally been called
Chloris nifescens Lagasca, a name which can-
not be typified and the original description of
which is not complete enough for positive iden-
tification. Accordingly, this name is rejected
( see under excluded names. )
Chloris aristata has been widely collected
from prairies and grasslands throughout much
of Central America (Fig. 41).
Representative specimens examined: COSTA
RICA: ccrca de San Ramon, Brenes s.n., 3
Jan 1935 (F); San Jose, Cooper 5,997, Hitch-
72
Bbicham Younc University Science Bulletin
Fig. 41. Distribution of Chloris aristata.
cock 8,469 (US), Valerio i04 (F, K); San Rafael
de Cartago, Pittier 7,109 (US); between San
Pedro de Montes de Oca and Curridibat,
Standley 32,817 (US). GUATEMALA: moun-
tains E of Quiche, Grout 640 (US); Guatemala
City, Hitchcock 9,039 (US); La Aurora, Ritana
699 (US); Dep. Alta Verapaz, near San Juan
Chamelco, StamUcy 92,435 (US); 3 km S of
Huehuetenango, Williams et al. 22,109 (F, US).
HAITL W of Palmiste, Leonard and Leonard
12,538 (NY). HONDURAS: Dep. Choluteca:
vicinity of San Marcos de Colon, Standley
15,765 (F); Dep. Morazan: vicinit\- of El Zanio-
rano, Standlcii 1501. 26,327 (F); Sicallen
10,832 (US). MEXICO: Aguascalientes, Hitch-
cock 7,454 (US); Chiapas: Monserrate, Pur-
pus 388 (F); Coahuila: Parras de la Fuentes,
Palmer 450 (F, MO, NY); Saltillo, Hitchcock
5,620 (US); Durango: Duiango, Hitchcock 7,657
(US); Guanajuato: 2.5 km N of San Fran-
cisco del Rincon, Sohns 355 (US); Irapuato,
Hitchcock 7,412 ( US ) ; Acamboro, Hitchcock
6,927 (US); Hidalgo: Jaeala, V. //. Chase 7,098
(US); La Placita, 10 mi S of Jaeala, Gould
10,389 (US); Jalisco: 15.5 mi E of Guadalajara,
Leavenworth 1,852 (US); San Nicolas, Hitch-
cock 7.193 (US); Michoacan: vicinity of Mo-
relia, Arsenc 3,134 (MO, NY, US); 'Uiuapan,
Hitchcock 6,965 ( US ) ; Nuevo Leon, Monterrey,
Hitchcock 1,263 (MO, NY); Galeana, V. H.
Chase 7,738 (US); Oaxaca: Cuicatlan, San Juan
Bautista, Conzatti 4,024.5 (US); Cd. Oaxaca,
Hitchcock 6,110, 6,129, 6,170 (US); Puebla:
Tehuacan, Hitchcock 6.033 (US); Queretaro:
Queretaro, Hitchcock 5,804 (US); San Luis
Potosi: Cardenas, Hitchcock 5,749 (US); Ve-
nado, Whiting 624. 711 (US); Veracruz: Jalapa,
Hitchcock 6,692 (US); Orizaba, Hitchcock
6,336 (US).
28. CHLORIS BRANDEGEI (Vasev) Swallen,
Amer. Jour. Rot. 22:41. 19.35. (Based on
Diplachne hrandegei Vasey ex Brandegee. )
Fig. 42.
Diplachne hrandegei Vasey ex Brandegee,
Proc. California Acad. Sci. H. 2:213. 1889.
(HOLOTYPE: "Magdalena Island. T. S.
Brandegee 18 January 1889." US! )
Leptochloa hrandegei (Vasey ex Brande-
gee) Hitchcock, U.S. Dept. Agric. Bur.
Plant IndustiN Bull. 33:21. 1903. ( Ba.sed
on Diplachne hrandegei Vasey ex Brande-
gee.)
Gouinia hrandegei (Vasey ex Brandegee)
Hitchcock, U.S. Dept. Agric. Bur. Plant In-
dustiy Bull. 33:21. 1903. (Based on
Diplachne hrandegei Vase>- ex Brandegee.)
Perennial to 1 m tall, densely tufted from a
coarse, fibrous root s\stem; lower sheaths tloc-
cose, becoming glabrous above; ligules pilose,
the lower more so than the upper; blades scab-
rous; inflorescence with up to 20 spikes ar-
ranged in a series of somewhat distant sub-
verticils, occasionally the spikes single at a
node; spikes 5 to 10 cm long; spikelets distant,
ca three or four per cm of the spike length,
strongly appressed; glimies markedly unequal,
narrowly lanceolate, thin, hyaline, glabrous ex-
cept for the slightly scabrous midnerve; first
glume 1 5 to 1.8 mm long, ca 0.3 mm wide;
second glume 3.3 to 4.0 mm long, ca 0.5 mm
wide; fertile lemma 4.7 to 5.5 mm long, 0.5 to
0.8 mm widi', strongly dorsally compressed,
iiarrowK' elliptic to linear in side view, more
broadh' elliptic in lace view, apex acute to
shortly acuminate, margins, keels, and inter-
nerves scabrous to hirsute, callus bearded, awn
1.0 to 4.5 mm long; sterile floret one, ca 1.5 mm
long. 0.3 mm wide, ver\- narrowly c\'1indrieal to
ellipsoid, scabrous to sparsely appressed-pilose,
awn 0.7 tn 1.9 mm long; earyopsis 2.5 to 3.0 mm
long, ca 0.7 mm wide, ca 0..3 mm thick, strongly
dorsally compressed, ellipsoid, slightly trigonous.
Chloris l)randegei is similar to and probably
closely related to C. chloridea, though distin-
guished from it by veiy short awns on the lem-
mas and a lack of underground spikelets.
Chloris hrandegei is endemic to Baja Cali-
fornia Sur, Mexico (Fig. 43, triangles), where
it grows on rocky slopes and in arroyos.
Specimens examined: MEXICO: Baja Cali-
fornia Sur: La Purisima, Brandegee L91 (US);
San Jose del Cabo, Brandegee s.n., 6 Oct 1890
(NY); Sierra de la Giganta, 25°58'N,
111°.32/'2'W, on road from Loreto to San Javier,
Carter 5.014 (UC); Cerro Teombo, 25"51'N,
111°25'W, N of Fortezuelo de Gabilan, 1100 m.
BiOLOGiCAi, Series, Vol. 19, No. 2 T.\xonomv of the Genus Chloris (Gramineae)
73
Fig. 42. Chloris hmndegei (A) habit, x 1/4; ( B,C,E ) .spikelet, partly dissected; ( D,F,G ) florets, showing varia-
tion; (H) caryopsis. (B-H) x 5.
74
Brigham Young University Science Bulletin
Fig. 43. Distribution of Chloris brandegei (triangles)
and Chloris suhmutica (dots).
Carter 5,064 (UC); 37 km SE of San Ignacio,
among lava rocks in giavcllv aiTO\o, Carter et al.
1,977\\JC, US); Siena de la Giganta, Arroyo
del Cajon de Tecomaja, SW of Puerto Escon-
dido, 25°43'N, 111°20'W, Carter and Kellogg
2,897 (UC, US); El Triunfo, A/. E. Jones 27,658
(UC); Carmen Island, Palmer 8,621, 1 to 7 Nov
1920 (US); Punta San Lorenzo, ca 25 mi E of
La Paz, Wiggins 15,638 (UC).
29. CHLORIS CHLORIDEA (Presl) Hitchcock,
Proc. Biol. Soc. Washington 41:162. 1928.
(Based on Dincha chloridea Presl.) Fig. 44.
Dineha chloridea Presl, Hel. Haenke 1:291.
1830. (TYPE: Not seen, but there is a frag-
ment in US!, from Haenke Herbarium, PR,
labeled: "Mexico . . . Presl." This and the
description clearly identify the name.)
Etitriana chloridea (Presl) Kunth, Rev.
Gram. Supplenda, p. 23. 18.34. (Based on
Dineha cidoridea Presl. )
Gymnopogon virletii Fournier, Mex. PI.
2:144. 1886. First published as nomen
nudem in Hemslev, Biol. Centr. Amer. Bot.
3:560. 1855. (TYPE: "San Luis de Potosi.
Virl. n. 1441." There is a fragment in US!,
from P, labeled "Mexi(jue. Prov. de San
Luis . . . \'irlet d'Aoust 1851," which is
C. cidoridea. )
Gymnojwgon longifolius Fournier, Mex. PI.
2:144. 1886. (FRAGMENT OF TYPE:
US!, originally from P, labeled ". . . Mex-
ique prov. \'era Cruz. Gouin. 1867. 52.")
Chloris longijolia (Fournier) Vasey, Contr.
U.S. Nat. Herb. 1:284. /;/. 7.9. 1893. Non
Steudel, 1854. (Based on Gymnopogon
longijolia Fournier. )
Chloris clandestina Scribner and Merrill,
U.S. Dept. Agric. Div. Agrost. Bull. 24:25.
1901. ( Based on Gymnopogon longifolius
Fournier. )
Perennial to 1 m tall, tufted, erect, bearing
cleistogamous spikelets at the tips of long, scaly,
branched rhizomes; each cleistogamous spike-
let consists of a series of three to five over-
lapping bracts enclosing a saclike structure,
which in turn encloses a pistil and three stamens;
mature cleistogamous fruit broadly ovoid-ellip-
soid, up to 4 mm long and 2.5 mm wide,
brown; sheaths mostly glabrous, occasionally be-
coming sparsely long-pilose near the ligule; low-
er ligules with a prominent tuft of hairs, upper
ligules often reduced to a low crown; blades
up to 30 cm long and 1 cm wide, scabrous, oc-
casionally pilose, innovations often sparsely pi-
lose; spikes three to fifteen ( usually less than
10), usually 6 to 10 cm long, in several well-
separated verticils, occasionally borne singly at
a node; spikelets rather distant, ca four per cm
of the scabrous rachis, appressed except for the
divergent awns; glumes niarkedh' unequal, gla-
brous except for the scabrous inidnerve; first
glume 1 to 2 mm long, ca 0.4 nun wide, lanceo-
late; second glume 2.0 to 3.5 mm long, ca 0.3
mm wide, narrowly lanceolate; fertile lemma 4.5
to 7.5 mm long, ca 1 mm wide, eu 0.5 mm
thick, dorsally compressed, linear to narrowly
lanceolate in side view, lanceolate in top view,
apex acuminate, callus densely bearded, mar-
gin ciliate, lemma otherwise glabrous, awn 6.5
to 15.0 mm long, often arising from between two
setae; sterile floret one, 1.4 to 3.0 mm long, less
than 0.3 mm uide, often somewhat flattened,
scabrous to short-pilose, awn 2 to 8 mm long;
car\opsis ca 4.5 mm long, 0.8 mm wide, some-
what (lattened, long-acuminate; chromosome
number 2n 40, SO.
The imderground cleistogamous spikelets of
Chloris chloridea make it one of the most un-
usual grassi's. These spikelets are borne on a
rather intricately branched rhizome system ( Fig.
44, A), the branches of which are brittle and
often are not collected. The cleistogamous spike-
lets on the specimens studied set a high pro-
portion of seed, while most of the above ground
spikelets were sterile. Whether this is due to
the immaturity of the specimens examined, en-
vironmental factors, or is an indication of a more
complicated reproductive pattern, must await
further field studies.
Chloris chloridea is quite variable in spike-
let size, and some plants may approach C. bran-
degei in lemma length, but C chloridea will al-
BiOLOciCAi. Series, Vol. 19, No. 2 Taxonomy ok the Genus Chloris (Gramineae)
75
Fig. 44. Chloris cMoridea. (A) habit, x 1/3; (B) spikfli t. partly dissected. .\ 7. .5; (C,D) florets, showing
variation, .\ 7. .5; (E) carvopsis from aboveground spikelet, x .5; (F) underground spikelet, x 5; (G) flower
of underground spikelet, x 15; (H) caryopsis from underground spikelet, x 5.
76
Bricham Younc University Science Bulletin
ways have relatively long awns and, of course,
the underground spikelets.
This species is widely distributed in the
southwestern United States, Mexico, Honduras,
and Venezuela and has been collected in grass-
lands, brushy areas, and old fields (Fig. 45).
Fig. 45. Distribution of Chloris chloridea.
Representative specimens examined: HON-
DURAS: Dep. Morazan; El Zamorano, Rod-
riguez 3,516 (F, US), StaiuUeij 13,076 (F,
NY, US), 14,454 (F), 14,H96 ("f, UC, US),
16,073 (F), 3,9.36 (US), Suallen 11,156 (US).
MEXICO: Baja California Sur: El Triunfo, M. E.
Jones s.n., 6 Oct 1930 (US); Guanajuato: 6 km S
of Ltoii, Soltm 384 (MO, US); Jalisco: Tu,\pan,
A/<?.T(« 1,046 (F, MO, UC, US); Michoacan;
Monte Leon, Pringle 3,994 (US); Morelos: near
Cuernavaca, Pringle s.n., 23 Oct 1902 (F, NY);
Nue\() Leon: Colonia del \'alle, Monterrex , F. F.
Smitli M267 (TEX); Oa.xaca: Tomellin,' //(7r/(-
cock 6,226 (US); San Luis Potosi: Las Canoas,
Hitchcock 5,766 (US); Sinaloa: Topolobampo,
Palmer 2.38, Sept 1897 (K, US); Sonora: Hermo-
sillo, Hitchcock 3,572 (US); Taniaulipas: 3 mi N
of Santo la Marina on road to Jimenez, Crtitch-
fiehl and Joltn.ston 4,972 (TEX); Veracruz: 5 mi
E of San Luis Potosi border, M. C. Johnston
6,106 (TEX, UC, US). UNITED STATES: Ari-
zona: Pima Co., W of Topawa, Coodding 194-45
(NY, US); Texas: Cameron Co., Laguna Atas-
cosa Refuge, Fleetwood 3,857 through 3,859,
3,861 (TEX); Brownsville Municipal Airport,
Rum/on 2,928 (US); Brownsville, Sdvcus 379 ( K,
US, TEX); 1 mi N of Combes, Silveus 1.583.
1,586 (MO); 0.5 mi N of Conches, Silveus 6.781
(TEX); 5 mi S of San Benito, Silveus 7,432
(TEX); Kleberg Co., Kingsville, Diaz 1,951
(US); 4 mi S of Kingsville, Meyer s.n., 24 Oct
1958 (US); Nueces Co., ca 5 mi S of Banquete,
M. C. Johnston 542,273 (TEX); San Patricio
Co., Odem, Silveus 2,408 (TEX). VENEZUE-
LA: Estado Lara, La Ruczga, cerea dc Bar-
quisimeto, Tanuiijo 4,230 (US).
30. CHLORIS VENTRICOSA Robert Brown,
Prodr. Flora Nov. Holl. 186. ISIO. (HOLO-
TYPE: "1. Chloris ventricosa Richmond"
BM!) Fig. 46.
Chloris sclerantha Lindley in Mitchell, Jour.
E.xped. Interior Australia '31. 1848. (TYPE:
"New South Wales; near Muddal banks of
Bogan River, Mitchell" CGE. Not seen.
Fragment of type at K! )
Chloris ventricosa var. tenuis Bentham,
Flora Australiensis 7:613. 1878. (SYNTYPE:
"Rockhampton, O'Shanesy. Herb. F. Muel-
ler 1879 [?]" K!)
Perennial to 1 m tall, stoloniferous, erect;
sheaths glabrous, scabrous or sparsely pilose;
ligule area scabrous to prominently pilose;
blades glabrous, scabrous, or sparsely pilose;
spikes two to nine, 5 to 11 cm long; spikelets
densely imbricate, ;i\x'raging 10 per cm of the
scabrous racliis; glumes scarious, narrowK lan-
ceolate to ovate, acute to acuminate, glaI)rous ex-
cept for the scabrous midnerves; first glume 1.2
to 2.3 mm long, ca 0.5 mm wide; second glume
2.5 to 4.1 mm long, ca 0.8 mm wide; fertile
lemma 2.0 to 5.4 mm long, 0.6 to 1.2 mm
wide, elliptic to obovate, sometimes ventricose,
acute to obtuse, usuallv glabrous except for the
scabrous midncrve, occasionally scabrous on the
interner\es and with a fvw scattered, appressed
hairs along the scabrous inrolled margin, es-
pecially near the apex, callus bearded, awn 1
to 11 mm long; sterile floret one (very rarely
two), 1.0 to 2.6 mm long, 0.3 to 1.0 mm wide,
cylindric to narrowly turbinate, apex obtuse to
truncate, awn 0.5 to 7.5 mm long; anthers 0.7
to 1.4 mm long; caryopsis 1.5 to 2.1 mm long,
ca 0.4 mm wide, narrowly obovoid, ± trigonous.
This species is quite variable, mostly with
respect to the height and apparent vigor of the
plant, the size of spikelet parts, and the shape
of the sterile florets. Attempts at correlating
various features of about 75 specimens were
made, but no conspicuous patterns could be de-
tected. This corroborates an earlier study by
Everist (193.5b), in which he found many in-
termediate plants between various extremes.
Certain of these have received names. Chloris
ventricosa \ar. tenuis Bentham is based on a
small plant with relatively small spikelets, while
a jjomewhat larger variant was called C. scle-
rantha Lindley.
Chloris ventricosa and C. truncata are closely
allied. Most specimens can be differentiated by
the length of the spikes, but some specimens of
C. truncata have abnormally short spikes. The
following key will separate most specimens:
Biological Series, Vol. 19, No. 2 T.\.\onomv of the Genus Chloris (Gramineae)
77
1. Lemma margin.s apprcssod-pilosc, rarely
sparsely so; awn of sterile floret ( the lower,
if two are present) 3.1 to 16.0 mm long,
usuallv 4 mm or more; spikes 5 to 23 cm,
usually 10 cm or more in length; lemmas
0.7 mm wide or less, usually ca 0.5 mm
wide; spikelets usually ver\' dark, black or
nearly so. C. tnmcata. Fig. 4S.
1. Lemma margins usually glabrous or
scabrous, not appressed-pilose, rarely veiT
sparsely pilose; awn of sterile floret (the
lower, if two are present) 0.5 to 7.5 mm
long, usuallv about 2 mm long; spikes gen-
erally short, 5 to 11 cm long, usually less
than 8 cm long; lemmas 0.6 mm wide or
more, usually about 0.8 to 1.0 mm; spike-
lets tawny, rarely dark brown.
C. ventricosa, Fig. 46.
Chloris ventricosa is widespread in eastern
Queensland and Ni'w South Wales ( Fig. 47,
B ) , where Everist ( 1935b ) reported that it ap-
pears to prefer heavier and richer soils.
Representative specimens examined: AUS-
TRALIA: New South Wales: Gravesend, Break-
well s.n., May, 1913 (F); Warialda, Hadleij
s.n., Apr 1908 (F); Hunter Valley, 9 mi NW of
Scone, Story 6,932 (K); beween Windsor and
Kingswood, Vickenj and Constable 16,121 (K,
Fig. 46. Chloris tculricosa. (A) hahit, x 1/6; (B) spikelet, partly
variation, .\ 15; (G) sterile floret, x 15; (H) caryopsis, .\ 10.
dissected, x 15; (C-F) florets, showing
78
Bricham Young University Science Bulletin
Fig. 47. Distribution of Chlork truncata (.M and C. ventricosa (B).
MO, US); Svdnev, Wilkes Exped s.n., 1832-42
(US); Queensland: Chinchilla, Beasleij 128 A
(K); near Wandoan, Behon s.n., 1930 (K);
Miles, Belson s.n., 1930 (K); Wallumbilla, Bel-
son s.n., 1930 (K); Downfall Creek, near Gilgul-
gil, Behon s.n., 1930; near Petrie, Blake 24 (K);
Allora, Darling Downs, Blake 294 (K); Dray-
ton, Blake 5,177 (K); Mundubbera, Blonsome
19, 54 (K); Laidley, Clemens s.n., 11 May 1943
(US); Kenmore, Clemens s.n., 18 May 1943
(US); Crow's Nest, Clemens s.n., 2A Feb 1944
(US); Yanaman Forest Reserve, Clemens s.n..
2 Aug 1944 (NY, UC, US); Moreton Dtr., Mt.
Coolon, Clemens s.n.. Mar 1945 (K); Jericho
vicinity, Clemens s.n.. Mar- Apr 1946 (US); Cal-
lide Cotton Research Station, Biloela, Coudrtj
55 (K); Maranoa Dtr., Cumming 20 (NY);
Rockhampton, Davies s.n., 23 Apr 1932 (K);
Gatton, Davies s.n.. May 1932 (K); Maranoa
Dtr., Boatman Station, Everist 2,779 (K, US);
Darling Downs Dtr., Benandri, 35 mi SE of
Texas, Everist 7,137 (US); Mt. Gravatt, Jack-
son s.n.. Mar 1931 (K); near Mt. Sturgeon Sta-
tion, Lazarides 3,644 (K, US); Leichhardt Dtr.,
2 mi NE of Moura Township, Lazarides 6,910
(K); Moonie River, Roe 822 (US); Burnett,
Kingaroy, Smith 3,042 (K); Toowoomba, White
6,642; Wondai, White 7,214 (K); Dalbv, White
s.n., Apr 1916 (K).
31. CHLORIS TRUNCATA Robert Brown,
Prodr. Flora Nov. Roll. 186. 1810. (HOLO-
TYPE: "Port Jackson . . . R. Brown" BM!
ISOTYPE: K!) Fig. 48.
Chloris elon^ata Poiret in Lamarck, Encycl.
Suppl. Meth. Bot. 2:236. 1811. (HOLO-
TYPE: Photograph and fragment, from
Desfontaines Herbarium at FI, in US! )
Chloris metcast acluja Sehrad. ex Schultes,
Mantissa 2:.340. 1824. (TYPE: "In Nova
Hollandia." Not seen, but description is
clear. )
Chloris truncata R. Br. f. ahhreviata Thel-
lung, Viertelijahr. Naturforsch. Gesell.
Zurich. 64:706. 1919. (ISOTYPE: Switzer-
land: Derendingen, Probst 27 Sept 1917.
K!)
Perennial, 30 to 50 cm tall, stoloniferous;
sheaths glabrous, iigules short-ciliate; blades 9
to 17 cm long, 0.2 to 0.3 mm wide, glabrous to
scabrous; spikes five to thirteen, 5 to 23 cm
long, radiate; spikelets appressed, rather distant,
ca six per cm of the scabrous rachis; glumes
narrowlv lanceolate, thin, hsaline, glabrous ex-
cept for the scabrous midrib; first glume 1.4 to
2..3 mm long, 0.2 to 0..3 mm wide; second glume
2.8 to 4.2 mm long, 0.2 to 0.4 mm wide; fertile
lemma l.S to 4.5 mm long, 0.2 to 0.7 mm wide.
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
79
Fig. 48. Chloris tnincata. (A) habit, x 1/3; (B) spikelet, partly dissected; (C-E) florets, showing variation;
(F) sterile floret; (G) caryopsis. ( B-G ) \ 10.
80
BniGHAM YouNO Univkhsity Science Bulletin
narrowly elliptic, glabrous except for tlie ap-
pressed-pilose margins, often becoming black at
maturity, apex truncate, callus bearded, awn 3.1
to 16.0 mm long; sterile floret one, rarely two,
1.3 to 3.5 mm long, 0.5 to 0.9 mm wide, apex
truncate, awn 3.1 to 12.5 mm long; anthers ca
0.6 mm long; cai-yopsis 1.7 to 2.2 mm long.
0.3 to 0.5 mm wide, ellipsoid to narrowh' obo-
vate, trigonous; chromosome number 2n=40.
Chloris truncata is most closely related to
C. ventricosa. Differences between the two have
been discussed under C. ventricosa.
Everist (1935b) reported that this species
seems to favor "black soil open downs, edge of
red soil country, and the edges of brigalow and
balah country." Fig. 47— A illustrates the dis-
tribution in Australia.
Representative specimens examined: AUS-
TRALIA: Capital Terr.: Canberra, McKee 7,712
(US), 8,931 (K), Evans 2,608 (K); New South
Wales: Nyngan, Baker s.n.. Dec. 1899 (US);
Hastings River, Berkler s.n. (K); Tocumwal,
Black m (US); Ballina, Black s.n.. Feb 1S94
(US); Singleton, Boorman s.n.. Nov 1914 (K);
Menindee Dtr., Quondang Tank, Constable
4,716 (K, US); Warialda, Hadlet/ s.n., Apr 1908
(F); Narromine, Helms 31,321 (K, NY); Guym,
McKee 646 (K); Stephen's Creek, Morris 489
(K); near Sydney, Vickenj 88 (K); Warren to
Collie Road, Vickenj K26 (K); Coonamble,
Vickenj K27 (K); Windsor to Kingswood,
Vickertj and Constable 16,08.5 (K); Belltrees
via Scone, White 8 (US); Queensland: Brisbane
River, Bick and White s.n.. May 1916 (K);
Petrie, near Young's Crossing, Blake 48 (K);
Wallangarra, Blake 4.422 (K); Cambooya,
Cashmore 808 (US); Dalveen, Clark s.n.. Mar
1916 (K); Darling Downs Dtr., Ballandean,
Clemens 44,480 (K); Brisbane, Turbot Street
pavement, Clemens s.n., 18 Mar 1942 (US);
Stanthorpe, Clemens s.n., 13 Nov 1943 (US);
Bald Mt., Clemens .s.n, 9 Nov 1944 (US); 35
mi SE of Texas, Everi.st 7,140 (K, US); Cun-
namulla, Gilnith Plains, Hartlei/ A18.5 (US);
Roma, White 9,573 (K); Darling Downs, be-
tween Inglewood and Milmenan, White 9,778
(K); Dalby, White s.n., Apr 1916 (K); Noondee
Station near Warwick, Young 12 (K); South
Australia: Koonamore Vegetation Reser\-e, Pal-
tridge 22 (K); Victoria: 2 mi S of Bacchus
Marsh, Aston 282 (US); Rochester, Davies s.n.,
28 Jan 1932 (K); Melbourne University ground.
Sonenberg s.n., Jan 19.35 (K); Western Aus-
tralia: Merredin," Allbani 11.48 (US): Carnar-
von, red muddy soil, Gardner G 3,003 (K);
Yundamindera Run, 120 mi NW of Kalgoorlie,
Pearse 1,931 (K). FIJI ISLANDS: Suva, Ti/r-
bet 42 (K). UNITED STATES: Hawaii, Ft.
Shafter, Hitchcock 13,849 (F).
32. CHLORIS PUMILIO Robert Brown. Prodr.
Flora Nov. Holl. IS6. 1810. (LECTO-
TYPE: BM! Additional specimens K! BM!
See discussion of nomenclature below. ) Fig.
49, J-P; 50, B.
Chloris pallida Hackel, Bot. jahrb. 6:244.
1885. (HOLOTYPE: "In Australia boreali-
occidentali Ig. Pemberton-Walcott." W,
Fragment at K! )
Chloris ptimilio (Robert Brown) Kuntze,
Rev. Gen. PI. 2:771. 1891. (Based on C.
pumilio Robert BrowiL )
Chloris ruderalis Domin, Biblioth. Bot.
85:365. 1915. (SYNTYPES: "Queensland:
Prope opp. Atherton legi II. 1910." ". . . in
xerodrymio prope opp. Mareeba legi II.
1910." ". . . prope opp. Charters Towers
legi II. 1910." All at K!, on extended loan
from PR.)
Chloris ruderalis f. biaristulata Domin,
Biblioth. Bot. 85:365. 1915. (HOLO-
TYPE: A specimen at K!, on extended loan
from PR, lalieled: ". . . Queensland apud
Mungana prope opp Chillagoe legi II. 1910"
consists of se\eral plants, one of which
fits the description in having two ver\' short
lateral teeth at the base of the sterile floret
awn and is possiblx the plant upon which
this fonn is based. )
Chloris ruderalis f. rolmsta Domin, Bib-
lioth. Bot. 85:.365. 1915. (TYPE: Ap-
paientlv one of the specimens on the sheet
discussed above, but no particular plant
can be assigned positively to this name. )
Annual 60 to 90 cm tall, tufted, or rooting
at the lower nodes, from a fibrous root svstem;
sheaths glabrous except pilose near the apex;
ligule a short-ciliate crown with a few long
hairs; blades 7 to 12 cm long, ca 3 mm wide,
scabrous abo\e with a few long hairs near the
base, glabrous below except for the scabrous
midrib; spikes fom' to twelve, 4 to 8 cm long.
erect to slightlv divergent; spikelets densely
inserted. a%eraging 9 to 16 per cm of the scab-
rous rachis; glumes narrowK lanceolate, gla-
brous except for the scabrous midnerve, mem-
branous; first glume 1.5 to 2.5 mm long, ca
0.3 to 0.4 mm wide; second glume 2.5 to 5.0
mm long, with a short hairlike awn; fertile
lemma .'3.7 to lO.O mm long ( including the
lateral awns). 0.4 to 0.8 mm wide, narrowly
lanceolate-elliptic, callus bearded, margins in-
BioLOGiCAi, Series, Vol. 19, No. 2 Ta.xonomv of the Genus Chloris (Gr.^mineae)
81
Fig. 49. Chloris lohata imd C. pumilio. (A-I) C. lohata. (A) habit, x 1/4; (B) spikelet, partly dissected, x
5; (C-E) florets, showing variation, X 5; (F) sterile floret, ventral view, x 5; (G) sterile floret, seniilateral
view, X 5; (H) group of sterile florets, showing variation, x 5; (I) caryopsis, x 10. (J-P) Chloris pumilio.
(J) spikelet, partlv dissected, x 5; (K) group of fertile florets, showing variation, x 5; (L) lower sterile
floret, lateral view', x 5; (M) upper sterile floret (not always present), x 5; (N) sterile floret, ventral view, x
5; ( O ) group of sterile florets, showing variation, x 5; ( P ) caryopsis, x 5.
82
Brigham Young University Science Bulletin
rolled, appressed short-pilose below, becoming
spreading long-pilose above, otherwise miniitelv
scabrous, apex bifid, the lobes gradually taper-
ing into awns up to 7 mm long, midnerve ex-
tending as an awn 7 to 16 mm long, almost
alwa)'s at least twice the length of the lateral
awns; sterile floret one (rarely two), 1.3 to 4.0
mm long (including the lateral awns), 0.3 to
0.7 mm wide, glabrous to slightly scabrous, apex
minutely ciliate, divided to as much as halfway
into two lobes, the lobes asymmetrical and with
lateral awns 1.0 to 3.5 mm long, central awn
3 to 15 mm long, apex minutely ciliate; ear\ opsis
1.2 to 3.3 mm long, ca 0.5 mm wide, ellipsoid,
trigonous.
Chloris piiinilio and C. hihata are very simi-
lar. The best characters to use to distinguish
C. pumilio are: lemma margins pilose for full
length, sterile floret bilobed onl\ halfwa\ to
the base, and the asymmetrically awned lobes
of the sterile floret.
Robert Brown (ISIO) described Chloris
pumilio on the b;isis of specimens collected on
islands off the northern coast of Australia. The
description is as follows:
"4. C. pumilio, spicis 2-4 erectiusculis,
glumis bifloris: valvuhs acuminato-aristatis,
perianthiis eiliatis lanceolatis, triaristatus.
aristis lateralibus \alvula brevioribus: inter-
media elongata. ( T. ) v. v."
This description was compared with the original
Brown manuscript notes at the librar\ of the
British Museum (Natural History). While many
of the words in this manuscript description are
illegibly written, it seems to matcli Brown's
published description in the important points.
The phrase ". . . arista lateralibus valvula
brevioribus . . ." suggested something (juite dif-
ferent from the taxon that has been called C.
pumilio for many years— the plant treated under
C. lohata in this paper. The latter species has
lateral awns that are clearly equal or nearly
equal to the central awn and much longer than
the lemmas. As with many of Brown's speci-
mens, typification is not straightforward, since
many were distributed and confused after his
death. In the case of this plant, however, there
can be little doubt; for in the British Museum
(and also at Kew), there are several sheets of
Brown's specimens containing many plants— all
fitting the above description. Most of these have
the typical blue label of the Bennett distribution
of Brown's specimens and, as such, are not
necessarily strictly authentic. One sheet, how-
ever, has on the back the handwritten nota-
tion "Nova Hollandia Ora Scptentrionalis. R.
Brown." The latter sheet is taken here to be
the lectotype, for it is the particular sheet se-
lected and annotated by Brown. All of the
specimens of the Bennett distribution at the
British Museum and Kew match the lectotype
as well.
The totalit) of this evidence leads to only
one conclusion: the plant (commonly called
Chloris rudcralis for many years) must now be
called C. pumilio, leaving the plant commonly
called C. pumilio without a name. This was
correctl)' recognized by Lazarides ( 1972 ) , who
provided the new name C. lohata for it.
Specimens examined: AUSTRALIA: North-
ern Territory: Katherine River levee, 14°20-2rS,
E of 132"E,' Black 17,414 (K, US); Queensland:
Stannary Hills, Bancroft 1,90S: Forest Home
Station,' Br«.9.v 1,821 (K); Gilbert River, Cum-
berland, Brass 8,857 (K); Gilbert River, with-
out precise location. White 1,489 (K); Stanley,
near Townsville, Chisholm-Ellis 16 (K); Towns-
ville. White s.n., 11 Feb. 1918 (K); Thursday
Island, Ilockings s.n., Apr 1931 (K); 2 mi S
of Nonnanton Township, Lazarides 4,272 (K);
Hayman Island, White 10.222 (K); 'Western
Australia: De Grey River, Mulgee Station, An-
derson 1,898 (K); North Kiniberle\- Dtr., May
River, Ba.sedow 75 (K); Broome, ' Bf/^es 7, .902
(K); Holmes 1270.210 (US); Wandagee, Mouri-
bandy, Gooch s.n., 16 Jul 1937 (K); Kimmaya,
Holmes s.n., Apr 1942 (US); Whim Creek,
Mitchell, without precise location or date (K);
Muhie, Roebounie, Morrison s.n.. Mar 1899
(K)'.
33. CHLORIS LOBATA Lazarides, Australian
Jour. Bot. Supplement 5. 20. 1972. ( HOLO-
TYPE: "Queensland: Burke District; 4
miles east of Nonnanton Township. 6. iii.
1954. Lazarides 4.289." CANB. ISOTYPES:
K! US! ) Fig. 49, A-I.
Chloris pumilio auct. non Robert Brown.
Annual 15 to 45 em tall, ert-et or geniculate,
often branching from thi' lower nodes; sheaths
glabrous, often sparseK' pilose toward the sum-
mit; ligule a densely ciliate crown with a few
long hairs intenningled ni'ar the edges; blades
up to 15 cm long, 0.2 to 0.6 mm wide, glabrous
to scabrous above ;md below; spikes three to
nine, erect to somewhat divergent. 2.5 to 6.0
cm long; spikelets appressed and imbricate, ca
8 to 15 per cm of the scabrous raehis; glumes
narrowly lanceolate, acuminate, membranous,
glabrous except for the scabrous midnen-e; first
glume 1.6 to 2.5 mm long, 0.2 to 0.4 mm wide;
second glume 2.6 to 3.9 mm long, 0.2 to 0.3 mm
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
83
wide; fertile lemma narrowly elliptic, 5.5 to 11.0
mm long (including the lateral but not the
middle awn), 0.3 to 0.6 mm wide, callus white
bearded, margins glabrous to scabrous below,
rather prominently long-ciliate near tin- apex,
remainder of lemma prominently scabrous antl
punctate, lateral nerves of lemma prolonged
into two awns 2 to 6 mm long, midnerve pro-
longed into an awn 3.0 to 10.5 mm long; sterile
floret one, 3.3 to 7.5 mm long (including tlu'
lateral awns), 0.4 to 0.5 mm wide, divided
nearly to the base into two flattened lanceolate
lobes, each with an awn ca 2 to 4 mm long,
middle awn 3 to 8 mm long; car)'()psis 1.7 to
2.4 mm long, ca 0.4 mm wide, ellipsoid,
trigonous.
As has been discussed under the previous
species, this taxon has for many years been
mistakenly called Chloris pumilio.
Collected on heavy soils on banks, water-
holes, and alluvial flats, mostly in Queensland,
Australia, but also occasionally found in North-
ern Territorx and Western Australia. ( Fig.
.50, C).
Representative specimens examined; .A,US-
TRALIA: Northern Territory, on levee of Roper
Fig. 50. Distribution of Chloris pectinata
pumilio (B), and C. lobata (C).
(A)
River, Else\- Station, E of Mataranka, Blake
17,517 (K, US): Kathenne, Tijack-Bake, May
1943 (UC). Queensland: Forest Home Sta-
tion, Gilbert River, Brass 1.704, 1,787 (K); Gil-
bert River, without precise location. White 1.492
(K); Nonnan Ri\'er, Gulliver, without precise
collection number and date ( K, US); near Nor-
manton, Lazarides 4,243 (K, US); Mt Molloy,
McKee 9.121 (K); 39 mi E of Lawn Hill Sta-
tion, Pernj 1.115 (K, US); between Townsville
and Rollingstone, White 8,895 ( K, US). West-
em Australia: 10 mi SE of Mt. House Station,
Lazarides 5,157 (K, US).
34. CHLORIS DIVARICATA Robert Rrown,
Prodr. Flora Nov. Holl. L186. 1810.
(LECTOTYPE: â– 3. Chloris divaricata, Port
I. Keppel Ra\. Whorlwater Ray . . ." BMl
See additional nomenclature note below.)
Fig. 51, K-R.
Chloris ct/nodontoidcs Balansa, Bull. Soc.
Rot. France 19:318. 1872. (HOLOTYPE:
"Bords des Chemins, Nouvelle Cale-
donie . . . Paneher, 18-70" P! )
Perennial 20 to 50 em tali, tufted to shortly
stoloniferous, sometimes apparently niat-fonn-
ing; sheaths glabrous; ligule a short-ciliate
crown; blades up to 15 cm long, generally 5
to 9 em, 1.0 to 1.5 mm wide, glabrous to scab-
rous; spikes three to nine, 4 to 17 cm long,
divaricate, becoming horizontal; spikelets sparse-
1\- inserted, about three to seven per cm of
tlie scabrous rachis, appressed, only the awns
divergent; glumes very narrowly lanceolate,
thin and membranous, glabrous except for the
scabrous midnei-ve; first glume 0.9 to 1.8 mm
long, ca 0.3 mm wide; second glume 2.0 to 2.9
mm long, ca 0.3 mm wide; fertile lemma linear
to narrowly lanceolate, 2.9 to 4.0 mm long, in-
eluding the teeth at the apex, callus bearded,
margins glabrous, scabrous or with a few ap-
pressed siiort hairs near the apex, keel scabrous,
internei-ves scabrous, awn 7.5 to 17.0 mm long,
sterile floret one, 1.2 to 1.9 mm long, ca 0.3
mm wide, narrowly elliptic, glabrous, apex
acute, bilobed K to 'A of its length; awn 4.5 to
9.5 mm long; car\'opsis ca 2.2 mm long, 0.4 mm
wide, narrowl)' ellipsoid.
Brown's original description listed the sym-
bol â– 'T.v.\-." as the place of collection. Accord-
ing to studies of original expedition charts by
Burbidge ( 1956), this is taken to be Cape Shield
near Groot Eylandt in the Gulf of Caqjcntaria.
I could not find specimens labeled as such at
the British Museum, but there were five speci-
mens labeled "Chloris jmrpuram. East Coast,"
84
Bnir.HAM Young University Science Bulletin
G *^H "I y\
Fie. 51. Chloris pectimta and C. divaricata. (A-J) C. pcclinala. (A) habit, x 1/4. (B) spikelet, partially dis-
^ sected; (CD) florets, sliowiiii; ^.,,rIati,„I; (K-1) sterile florets, dorsal view, partia ly flattened, ;<h°^"ng van-
ation; (J) caryopsis. (K-R) C. divaricata. (K) habit, x 1/4; (L) spikelet, partially d'ssected; (M-Q) sterile
florets, dorsal view, partially flattened, showing variation; (R) caryopsis. ( B-J ) and (M-H) x lu.
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
85
a sixth of the Bennett distribution of Brown's
specimens, with the usual blue label, and a
seventh, labeled "3. Chloris divaricata. Port I.
Keppel Bay. Whorlwater Bav. Broadsound . . ."
The label of the last specimen is handwritten
and has two slits cut in it— a feature thought
to indicate that this is the specimen selected
by Brown from the bulk of the collections and
studied b\' him on his journey home from Aus-
tralia. Fortunately, all specimens are of the
same taxon.
The two collection sites mentioned, the one
in the description and the second on the vari-
ous Brown specimens at the British Museum,
are widely separated— Cape Shield being on the
west shore of the Gulf of Carpentaria and "Port
I. Keppel Bay . . ." on the east coast of Queens-
land. The latter specimen was obviousl) seen
and studied by Brown; and even though its col-
lection site is at variance with that cited in the
description, it seems best to designate it as the
leetotype of the species.
Cliloris divaricata from Australia and C.
cijnodontoidcs from Fiji, Tonga, and New Cale-
donia have commonly been treated as distinct
species, though, more recently, Lazarides
( 1972 ) treated them as varieties of C. divari-
cata. The priman' differences emphasized are
that C. divaricata is not stoloniferous and has
ielati\'el\- long spikes, while C. cijnodontoidcs
is stoloniferous and has short spikes. Table 10
presents the results of measurements of 56 dif-
ferent specimens from all of the geographic
ariMS represented.
As can be seen, onlv minor statistical dif-
ferences exist between the two populations.
While average spike number and length mav
differ, the ranges overlap greatly. Likewise, the
presence or absence of stolons is not sufficienth
exclusive to be used in separating the two
groups.
Lazarides (1972) has mapped the two vari-
eties, showing var. ci/nodontoides in inland
areas of southern Queensland and coastal onl\'
in the north. Collections of var. divaricata may
also be noted from these regions but are much
more commonly found coastward and south-
ward, extending well into New South Wales.
The specimens I have seen do not support
such a clear-cut separation, for stoloniferous
plants with relatively large numbers of long
spikes are intermingled with tufted plants with
fewer, shorter spikes. Plants showing different
combinations of these features or showing inter-
mediate values may also be found. Until addi-
tional field studies are made, I feel that recog-
nition of distinct varieties is not warranted.
CIdoris divaricata most closely resembles C.
pcctinata, though it differs in having sterile
florets 1.2 to 1.9 mm long (as opposed to 1.7
to 2.9) and only slightly imbricate spikelets.
The spikelets of C. pectinata are strongly
imbricate.
Everist ( 1935a ) reported that this species
occurs in "practicalK' all types of soil, from
heavy black soils to light sandy loams." It has
been collected in eastern Queensland and New
South Wales, Australia; Fiji Islands; and Tonga
Islands (Fig. 52). Specimens from Hawaii,
Texas, and France are adventives, as are prob-
ably those from New Caledonia.
Representative specimens examined. AUS-
TRALIA: New South Wales: 6 mi NW of Mus-
wellbrook, Storij 7.079 (K); Warialda, Vickenj
18.028 (K). Queensland: Brisbane River, on
lidges, Badey without collection number or
date (K); Brisbane, Bailey without collection
number or date (US 879,412); Dalby, Beiers
47 (K); near Wallumbilla, Belson s.n., 1930
(K); Petrie, near Young's Crossing, Blake 49
(K); Dravton, Blake 5,176 (K); Mundubbera,
Blonwme'41 (K); Mt. Molloy, Brass 2,405 (K);
South Brisbane Park, Cdemens s.n., 20 Feb 1943
(US); near Mt. Gravatt, Clemens s.n., 11 Mar
1943 (US); Kenmore, Clemens s.n., 18 May
1943 (US); Rosewood, Clemens s.n., 18 Jan
1944 (US); Bunya Mts., Clemens s.n.. Mar 1944
(US); Bauple, Wide Bay Dtr., Clemens s.n.,
10-20 Jun 1945 ( K ) ; Charleville, Warrego Dtr.,
Clemens s.n., 9 Nov. 1945 (K); Mt. Fox, Clemens
s.n., Sept-Dec 1949 (K); Callide Cotton Re-
search Station, Biloela, Cowdrij 1,930 (K);
Table 10. Comparison of spike length, spike number, and growth habit of plants called C. divaricata and C.
cijnodontoidcs.
Geographic
origin of
plant
Number of spikes
Range Average
Length of spikes
Range Average
Stolons
Number of Number of
plants having plants lacking
stolons stolons
Australia 2^9 5.7 4.5-16
Fiji, Tonga,
and New CiJedonia 3-9 5.2 4-8
11.1
6.3
16
11
27
86
Bricham Young University Science Bulletin
^D
.-f*"
,o
<i>,
^S:>
D
^
E
Fig. 52. Distribution of Chloris clivaricata. (A) east-
em Australia. ( B ) Fiji Islands. ( C ) New Caledonia.
(D) Hawaiian Islands. (E) Tonga Islands.
Rosedale, Dovey 439 (K); Boatman Station,
Everist 2,778 (K), 2,931 (K, US); Benandri, 35
mi SE of Texas, Everist 7,139 (K, US); Bishop
Island, mouth of Brisbane River, Everist and
Blake 367 (K); Macalister, Harris 5.505 (K);
10 mi S of Miles, Johnson 719 (K); Rcedsdale,
5 mi NW of B\ mount, Johnson 2,251 (K);
Gympie, Kenny without collection number or
date (US 1,386,466); Conjubov Station, Laza-
rides 4,137 (K); 1 mi NE of Niall Station,
Lazarides 4,630 (K, US); Moreton Dtr., Water-
ford, Mead 7 (K); Jandowae, NE of Dalby,
King's Creek, E of Bowman,
1870 (K); Mitchell Dtr., Brixton,
13 Nov 1930 (K); Holcombe, via
s.n., 21 Jun 1930 (K);
28 mi VVNW of Nebe
Moore 36 (K);
Mueller s.n.
Ratcson s.n.,
Tara, Salter
Elphinstone.
near Lake
Township,
White 7S9
(K, US);
Murgon,
Story and Yapp 85 ( K ) ; St. George.
(US); Rockhampton, White 3,375
Clermont, White 3,416 (K, US)
White 7,230 (K); Rolleston, White 8,830 (K,,
Darling Downs, between Inglewood and Mil-
merran. White 9,790 (K); Wellington, White
s.n., 17 Mar 1916 (K); Mareeba, White s.n.,
Jan 1918 (K); Laidley, White s.n., 17 Mar 1921
(K). FIJI ISLAND: Suva, Greenwood 226A
(K); Viti Levu, Nandronga, Singatoka, Green-
wood 226C (K, NY, US); ViH Levu, Nafasiri,
near Nasinu, Greenwood 226D ( US ) ; Viti Leon,
near Casey, Suva, TothiU 43 (K); Totoya Is-
land, Tothill 201 (K); Matuka Island, Tothill
291 (K); Kondavu Island, Turbet 8 (K).
FRANCE: Dunquerque, port area, anon. (K).
NEW CALEDONIA: Noumea, Bernier 812 (K);
foot of Ouen Toro, McKee 11,499 (K. US).
TONGA ISLANDS: Tongatapu, Kolomotuia,
Sokai 843 (K). UNITED STATES: Hawaii:
Honolulu, near Bishop Museum, Neal s.n., 7
Sept 1934 (K); Niihau, St. John 23,623 (K);
University of Hawaii campus, St. John 24,832
(K); WjWer .s.n., 19 Jan 1931 (K). Texas: Bexar
Co., U.S. Experiment Station, San Antonio, Sil-
veus 2,564 (TAES); Nueces Co., near Tuloso-
Midway High School. Hwv. 9, between Corpus
Christi and Calallen, W. V.' Brown 755 (TAES);
Refugio Co., 2 mi SE of Woodsboro, along
Road 1360, Waller et al. 2,118 (TAES).
35. CHLORIS PECTINATA Bentham. Flora
Australiensis 7:612. 1878. (SYNTYPE:
"Charlotte Waters. Giles" K!) Fig. 51,
A-J.
Chloris divaricata var. muelleri Domin, Re-
pert. Sp. Nov 9:553. 1911. (ISOTYPE:
"Sturts Creek. Feb 56 . . . Mueller." K!)
Chloris j)ectin(ita var. tt/pica Domin, Bib-
lioth. Bot. 85:366. 1015." (Based on C. pec-
tinata Bentham.)
Chloris pectinata var. fallax Domin, Bib-
lioth. Bot. 85:366. 1915. (HOLOTYPE:
"Queensland . . . Rolling Downs . . . Domin
II. 1910." K!, on extended loan from PR.)
Cldoris divaricata var. minor Black, Flora
South Australia Pt. 1. 83. pi. 4. 1922.
(ISOTYPE: "Oodnatta . . . Miss Staer . . ."
K!)
Annual 20 to 75 cm tall, erect, often branched
above, especially on taller specimens; sheaths
glabrous; ligule a dense short-ciliate crown, oc-
casionally with a few longer hairs intermixed;
blades up to 15 cm long, 2 to 5 mm wide, gla-
brous to scabrous; spikes four to thirteen, 5 to
11 cm long, erect at first, later becoming widely
divaricate to horizontally spreading at maturity;
spikelets densely inserted on the rachis, spread-
ing, ca 10 to 14 per cm of the scabrous rachis;
glumes narrowlv lanceolate, membranous and
glabrous except for the scabrous midner\('; first
glume 1.4 to 2.5 mm long, 0.1 to 0.3 mm wide;
second glume 2.9 to 4.3 mm long, 0.2 to 0.3" mm
wide; fertile lemma 3.0 to 6.2 mm long (includ-
ing lateral teeth), 0.4 to 0.6 mm wide, linear to
narrowly lanceolate, becoming dark gray to
black at maturity, callus bearded, margins
Biological Series, Vol. 19, No. 2 T.v.xonomy of the Genus Chloris (Gramineae)
87
glabrous, scabrous, or appressed short-pilose
(the hairs less than 0.2 mm long), sides of
lemmas sparsely to densely glandular and scab-
rous, occasionally glabrous, lemma teeth 0.5 to
10.0 mm long, awn 6 to 37 mm long; sterile
floret one, 1.7 to 2.9 mm long, 0.2 to 0.3 mm
wide, flattened, bilobed no more than halfwav
to the base, glabrous, awn 4 to 10 mm long;
anthers ca 0.6 mm long; caryopsis ca 2.3 mm
long, 0.3 mm wide, narrowly ellipsoid, trigonous.
Chloris pectiimta is most closely related to
C. divaricata. Differences have been discussed
under the latter taxon.
This species is widely distributed in central
and western Queensland and also in Northern
Territory and South Australia (Fig. 50, A).
Everist (193S) reported that it occurs in a
variety of habitats but develops best in low-
lying clay soils.
Representative specimens examined: AUS-
TRALIA: Northern Territoiy: 26 mi NNW of
Brunette Downs, Blakc 17,819 (K, US); Mac-
Donald Downs, Chalmers 32 (K); 22 mi NE
of Argadargada Station, Lazarides 5,258 (K,
US); Barkl)' Highway, James River crossing,
Pernj 710 (K); Queensland: Eulo, near Cunna-
mulla, Allan 10.948 (US); Forest Home Sta-
Hon, Gilbert River, Brass 1,822 (K); Charleville
Dtr., Warrego, Clemens s.n., 7 Nov 1945 (K);
Jericho, Clemens s.n., 6 Mar 1946 (US);
Mitchell Dtr., Springvale, Clemens s.n., 2 Apr
1946 (US); Maranoa Dtr., Boatman Station,
Everist 2,778 (US); Cook Dtr., road from Taba-
cum to Springmount, Goodall s.n., 2 Apr 1961
(US); Hughenden, Hawthorn s.n., Jun 1919
(K); 27 mi NW of Cloncuru', Lazarides 4.308
(K, US); 3 mi SW of Malbon Township.
Lazarides 4,401 (K, US); Toorak Experiment
Station, Lazarides 4,446 (K, US); Adovale,
MacGiUivraij 1,029 (K); Cunnamulla, McKee
s.n., 11 Apr 1963 (K); 25 mi NNE of Cam-
ooweal. Perry 955 (K, US); Blackall, Ranking,
13 (K); Longreach Dtr., Thomas s.n.. Mar 1927
(K); Clermont, White 3,422 (K); Antil Plains
near Townsville, White 8,893 ( K, US); War-
rego Dtr., Carbeen, near Cunnamulla, White
11,556 (K); Mitchell Dtr., Tower Hill, White
s.n., Apr 1919 (K); Quilpie, White s.n.. 20 Mav
1928 (K); South Australia: Cordillo Downs,
Cleland s.n.. May 1924 (K); Oodnadatta, Staer
s.n., Jan 1913 (K); Western Australia: Wan-
dagee, Minilya River, Gardner 3,236 (K); Wan-
dagee Station, near Caniar\'on, Meadh/ A/77
(K); Warrambic, Roebourne, Meares 345 (K).
36. CHLORIS CRINITA Lagasca, Var. Cienc.
Lit. Arts (Madrid) 4; 143. 1805. NoHien
nudem; Nov. Gen. et Sp. 5. 1816. The
first cited publication has not been located
in any United States library, but Mrs. Ag-
nes Chase found the series in the British
Museum and placed copies of Lagasca's
protologues on file at US! The name was
apparently first used without a description;
later Lagasca provided a description in the
second work mentioned above. Parodi
( 1947 ) states that no specimen with the
appropriate number could be found among
the types at Madrid, but that a figure bear-
ing Lagasca's handwritten annotation is of
the species as commonly understood. Fig.
53, D-F.
Chloris mendociim Philippi, Anales Univ.
Chile 36:208. 1870. The original descrip-
tion cites a collection by Mendoza. A speci-
men in P! labeled "Phillipe pi. chilense
Mendocina. Chloris mendocina Phil. Ipse
Mendoza" may be the type. It and the
description are Chloris crinita.
Trichloris fasciculata Fournier v\ Bentham,
Jour. Linn. Soc. Bot. (London) 19:103.
1881. Nomen nudem; Fournier, Mex. PI.
2:142. 1886. (HOLOTYPE: "San Luis de
Potosi. Virl. n. 1440" P! )
Trichloris hlanchardiana Fournier ex Scrib-
ncr, Bull. Torrev Bot. Club 9:146. 1882.
(HOLOTYPE: "Santa Cruz Valley near
Tucson . . . Pringle, May 19" US!)
Chloropsis hlanchardiana (Fournier)
Kuntze. Rev. Gen. PI. 2:771. 1891. (Based
on Trichloris hlanchardiana Fournier.)
Chloropsis crinita (Lagasca) Kuntze, Rev.
Gen. PI. 2:771. 1891. (Based on Chloris
crinita Lagasca.)
CIdoropsis fascictdata (Fournier ex Ben-
tham) Kuntze, Rev. Gen. PI. 2:771. 1891.
( Based on Trichloris fasciculata Fournier
ex Bentham. )
Leptochloris crinita (Lagasca) Munro ex
Kuntze, Rev. Gen. PI. 2:771. 1891. (Based
on Chloris crinita Lagasca.)
Trichloris verticillata Fournier ex Vasey,
U.S. Dept. Agric. Div. Bot. Bull. 12. Pt.
II: pi. 25. 1890. "Chloropsis hlanchardiana
Gav in Herb." is cited; description and illus-
tration are of C. crinita.
Chloropsis inendocina (Phihppi) Kuntze,
Rev. Gen. PI. 3:.348. 1898. (Based on
Chloris mendocina Philippi.)
Trichloris mendocina f. hlanchardiana
(Fournier ex Scribner) Kurtz, Bol. Acad.
Nac. Ciencias Cordoba (Rep. Argentina)
88
Bricham Young University Science Bulletin
Fig. 53. Chloris phiriflora and C. crinita. (A-C) C. jih.trijlora. (A) habit, x l/."); (B) .spikelet, partly di.s-
sected, x 10; (C) caryopsis, x 10. (D-F) C. crinita. (D) habit, x 1/5; (E) spikelet, partly dissected, x 10;
(F) caryopsis, x 10.
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
89
16:270. 1900. (Based on Trichloris blanch-
arkina Fournicr c.x Scribner. )
Chloris trichodcs Lagasca ex Parodi, Revis-
ta Argentina Agron." 14:62. 1947. Citation
of an herbarium name.
Tricltloris crinita (Lagasca) Parodi, Re-
vista Argentina Agron. 6.3. 1947. (Based
on Chloris crinita Lagasca.)
Stoloniferous or tufted perennial up to 1
m tall; sheaths glabrous to sparsely hirsute;
ligule ciliate with cilia up to 3 mm long; blades
up to 20 cm long, 5 to 10 mm wide, scabrous
above and below; spikes six to twenty, up to
15 cm long, erect, arising in several close ver-
ticils, spikelets densely arranged on the rachis,
averaging ca seven to nine per cm of the scab-
rous rachis; glumes lanceolate to ovate, acumi-
nate to awned, glabrous except for the scabrous
midrib; first glume 0.8 to 1.1 mm long, ca 0.3
mm wide, linear to narrowly lanceolate; second
glume 2.0 to 2.5 mm long, ovate, with an awn
up to 2 mm long; fertile lemma 2.4 to 3.S mm
long, ca 0.5 mm wide, dorsally flattened, nar-
rowly lanceolate to elliptic, scabrous, especialh
above the middle, apex three-awncd, central
awn S to 12 mm long, the lateral usually some-
what shorter; sterile floret usually one (occa-
sionally two), greatly reduced, cylindrical, the
floret proper ca 1.0 to 1.5 mm long, gradualh'
narrowing at the apex into three subequal awns,
the awns ca 5 to 7 mm long; second sterile
floret (when present) similar to the first but
smaller; caryopsis 1.7 to 2.3 mm long, ca 0.6 mm
wide, ca 0.3 mm thick, strongly dorsally flat-
tened; chromosome number 2n^40.
Onh three species in Chloris have three long
awns on the feitile and sterile florets— C. crinita,
C. lobata. and C. pumilio. Chloris crinita is a
New World tufted perennial with the lemma
apex tapering abraptly to the lateral awns.
Both Chloris lobata and C. pumilio are Aus-
tralian annuals in which the upper lemma is
split, the lobes gradually narrowing into the
awns. There are manv additional, subtle, quali-
tative differences; and I believe the resemblance
is largelv superficial.
Chloris crinita is widelv distributed in the
south central to southwestern United States,
northern Mexico, and, rarely, has been collected
in South America (Fig. 54).
Representative specimens examined: AR-
GENTINA: Prov. Buenos Aires: Dep. Villarino.
60 km E of Rio Colorado, Eijerdam et al. 2.3,526
(US); Prov. Catamarca: Santa Maria, Castillon
3,290 (US); Prov. Chaco: Siena Blanca, Jor-
^ensen 2,878 (US); Prov. Cordoba: Reinedias,
Niedfeld s.n., 17 Apr 1920 (US); Prov. Corrien-
tes: Dep. Mburucuya, Estancia Santa Teresa,
Pedersen 63 (US); Prov. Fonnosa: Guayculea,
Jorgensen 3,330 (US); Prov. Jujuy: Laguna de
la Brea, Fries 115 (US); Prov. Mendoza: Vallee
de Rio Atuel, St. Raphael, Wilczek 526 (US);
Prov. Salta: Coronel Moldes, Htinziker 1,110
(US); Prov. San Juan: Colonia Fiscal, Sar-
iniento, Bartlett 20,522 (US); Prov. Santiago del
Estero: Rio Huayco Hondo, Bartlett 19,754
(US); Prov. Tucuman: Trancas, Tapia, Venturi
2,345 (US). BOLIVIA: Mataral, anon., 1804
(US). MEXICO; Chihuahua: 100 km W of
Cuarto Cienegas, Harvey 1,255 (US); Durango:
Torreon, Hitchcock 7.724 (US); San Luis Potosi:
20 km NE of Rio Verde, Rzedowski 5,157 (US).
PARAGUAY: Dep. Chaco: Loma Para, Rojas
2,751 (US). UNITED STATES: Arizona: Ben-
son, Griffiths 1,969 (US); New Mexico: Organ
Mts., W of Dona Ana, Wooton and Standletj s.n..
Sept 1907 (US); Texas: vicinity of El Paso,
Chase 5,899 (US); Fort Worth, Tracy 8,208
(US); Pecos, Silveus309 (US).
37. CHLORIS PLURIFLORA (Fournier) Clay-
ton, Kew Bull. 21:102. 1967. (Based on
Trichloris pluriflora.) Fig. 53, A-C.
Trichloris pluriflora Fournier in Hemsley,
Biol. Centr. Amer. Bot. 3:560. 1885.
Fig. 54. Distribution of Chloris crinita. Inset A: South-
ern United States and Mexico.
90
Bricham Young University Science Bulletin
Nomen nudem; Fournier Mcx. Pi. 2:142.
1886, with description. (HOLOTYPE:
"Karw. in Herb. Petrop." Fra^nent of tvpe
in US!)
Chloropsis pluriflora (Fournier) Kuntze,
Rev. Gen. PL 2:771. 1891. (Based on
Trichloris pluriflora Fournier.)
Trichloris pluriflora f. macra Hackcl in
Stuck., Anales Mus. Nac. Hist. Nat. Buenos
Aires 14:116. 1911. Notnen nudem.
Though this name is without a description,
a dupHcate of a specimen in US! from the
Hackel Herbarium is t)'pical Chloris pluri-
flora.
Trichlori.<i lilloi Parodi, Physis 9:256. 1928.
(HOLOTYPE: "Argentina, Tucuman, Es-
quina Grande, Departamento de Cliicligasta
. . . Jorgenscn, no. 1396." Not seen, but
description and accompanying illustration
clearly refer to Trichloris pluriflora.)
Stoloniferous or tufted perennial up to 1.5
m tall; sheaths glabrous to sparsely hirsute;
ligule ciliate, cilia up to 3 mm long; blades up
to 30 cm long, up to 1 cm wide, scabrous to
sparsely hirsute above and below; spikes seven
to twenty, up to 20 cm long, erect, in few,
rather distant verticils; spikelets densely ar-
ranged on the rachis, averaging ca seven to nine
per cm of the scabrous rachis; glumes lanceolate
to narrowly lanceolate, glabrous except for the
scabrous midnerve; first glume 2 to 3 mm long,
ca 0.2 to 0.3 mm wide; second glume 3 to 5 mm
long, ea 0.5 mm wide; fertile florets one or two,
occasionally a third with rudimentar)' pistils
and stamens, narrowly lanceolate, 3 to 5 mm
long, margins short-ciliate near the middle, up-
per portions sparsely scabrous, otherwise gla-
brous, central awn 8 to 12 mm long, lateral awns
0.5 to 1.5 mm long; sterile florets two or three
(very rarely only one), the lower often with
rudimentary pistils and stamens, narrowly lan-
ceolate; lowermost sterile floret 1.5 to 3.0 mm
long, ea 0.3 mm wide; upper sterile florets pro-
gressively reduced, margins sometimes short-cil-
iate near the middle, othenvise glabrous, central
awn up to 8 mm long, lateral awns ca 0.2 to
1.0 mm long; car\opsis 1.8 to 2.2 mm long, ca
0.5 mm wide, ca 0.2 mm thick, strongly dorsally
flattened; chromosome number 2n=80.
Chloris pluriflora is most similar to C. utollis
(a Central and South American species) and
to C. pumilio and C. lobata (both from Aus-
traha). It may be separated from all three in
having usually two or more sterile florets. In
addition, the presence of the lateral awns will
differentiate it from C. mollis. Neither of the
Austi-alian species has the sterile floret with a
bearded callus.
The presence of the lateral awns in Chloris
pluriflora and in some Australian species would
suggest a close relationship, but there are many
features that are not shared; and I believe the
similarits' is superficial.
Chloris mollis is very similar phenetically,
lacking only the well-developed lateral awns;
I believe that it and C. pluriflora are closely
related.
For many years Chloris pluriflora has been
placed in Trichloris, along with C. crinita. Ar-
guments for the merger of the two genera were
presented by Clayton (1967). Earlier in the
present study, I discussed the difficulty of main-
taining Trichloris as a separate genus.
Chloris pluriflora is found in the south-
western United States, northern Mexico, and
South America (Fig. 55).
Representative specimens examined; AR-
GENTINA: Prov. Buenos Aires: Palermo,
Burkart 4,658 (US); Prov. Catamarca; An-
dalgala, Jorgensen 1,396 (US); Prov. Cordoba:
without location, Stuckert 8,959 (US); Prov.
Jujuv: Ceoro Moreno, Vcnturi 8,191 (US);
Prov. Salta: Coronel Moldes, Bartlett 19,646
(US); Prov. Santiago del Estero: C. Pelligrini,
Venturi 5,716 (US); Prov. Tucuman: Barranca
Colorado, Venturi 810 (US). BOLIVIA: Villa-
Fig. 55. Distribution ot Chloris pluriflora. Inset A:
Southern United States, Mexico, Central America,
and Caribbean Islands.
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Cramineae)
91
niontes, anon., K. Pflanz no. 2,097 (US).
CUBA: Prov. Habana: Casa Blanta, Ekman
16,879 (US). ECUADOR: Prov. Loja: benvccn
La Toma and Loja, Hitchcock 21,400 (US).
GUATEMALA: Peten, Tikal National Park,
Lundell 16,066 (US). MEXICO: Coahuila:
Sabina.s, Nelson 6,827 (US); Neuvo Leon: 12
mi S of Montcrrev, Mueller 375 (US); Oaxaca:
Canon df Santa Catarina, Conzatti 3,985 ( US ) ;
Tamaulipas: Podilla, Swallen 1,734 (US).
PARAGUAY: Asuncion, Puerto Casado, Rajas
2,307 (US). PERU: Cuzco, Gunther 66.5
(US). UNITED STATES: Texas: Cameron
Co., Resaca del Rancho Viejo, 6 mi NE of
Brownsville, Cory 51,411 (US); Duval Co., 2 mi
SW of Benavides, Harvey 895 (US); Kleberg
Co., W of Kingsville, Sicallen 10.258 (US);
Webb Co., Laredo, Griffiths 6,542 (US); Wil-
lacy Co., E of Ravmondville, Swallcn 10,100
(US).
38. CHLORIS CUCULLATA Bischoff, Ann.
Sci. Nat. Bot. (Paris) III. I9:.3.57. 1853.
(TYPE: "Provincia Tamaulipas prope Mat-
amoros . . . Engelmann, 1849". Not seen,
but description elcarl\- refers to this taxon. )
Fig. 56, A-K.
Perennial, tufted, erect. 15 to 60 em tall;
sheaths glabrous; ligule a crown of very short
hairs; blades 2 to 4 mm wide, up to 20 cm
long, often greatly reduced in size upward on
the culm, glabrous to scabrous; spikes ten to
twenty, 2 to 5 cm long, borne in several close,
radiating \erticils at the apex of the culm,
flexuous or curved; spikelets densely inserted on
the scabrous rachis, averaging 14 to 18 per
cm of rachis, spreading at nearh' right angles,
straw-colored at first, later becoming fuscous;
glumes lanceolate to obovate, glabrous except
for th(> scabrous midnei-ve, membranous; first
glume 0.5 to 0.7 mm long, 0.4 to 0.5 mm wide,
ovate to broadly lanceolate; second glume 1.0
to 1.5 mm long, 0.4 to 0.6 mm wide, obovate;
fertile lemma 1.5 to 2.0 mm long, 0.7 to 1.0
mm wide, broadly elliptic, apex obtuse, callus
short-bearded, keel and marginal nerves ap-
pressed-pilose, otherwise glabrous, awn 0.3 to
L5 mm long; sterile floret one, LO to 1.5 mm
long, 1.0 to 1.5 mm wide, markedly inflated
with the upper margins inrolled, unawned or
with a short awn up to 1.5 mm long; canopsis
0.9 to 1.2 mm long, ca 0.5 mm wide, obovoid,
rounded to trigonous; chromosome number
2n=:40.
Chloris cucullata is quite distinct from most
other species in the genus. It hybridizes readily
with C. andropogonoides and C. verticillata, but.
in pure form, it differs markedly from these two
species (see Table 11 and Fig. 56, A-K, L-U;
58, D-K). A complete discussion of the hybrid-
ization and introgression between these species
follows the description and discussion of C.
andropogonoides (p. 95).
Chloris cucullata is widely distributed
throughout much of Texas and is also found
in adjacent areas of New Mexico and Mexico
( Fig. 57, shaded circles ) . It is a common road-
side or waste area weed in many areas.
Representative specimens examined: NOTE:
Additional citations may be found in the
list of hybrid populations studied following
the discussion of introgression and hybridization
of C. cucullata, C. verticillata, and C. andro-
pogonoides.) MEXICO: Nuevo Lchju: below
La Boca Canyon, R. F. Smith M596 (TEX).
Tamaulipas: .34 mi S of Matamoras, Crutch-
field and Johnston 5,475 (TEX); 5 km S of
Nuevo Laredo, Dominguez 53 (TEX); 24 km
S of Nuevo Laredo, IDominguez and McCart
8,208 (TEX). UNITED STATES: New Mex-
ico: Chaves Co., 21 m E of Roswell along Hwy.
.380, Anderson 4,740 (HSC); Dona Ana Co., Las
Cruces, Plank sji., July 1894 (NY); Eddy Co.,
21 mi NW of Carlsbad, Gould and Kapadia
9,543 (TEX). Texas: Andrews Co., 8 mi NW
of Andrews, Rowell 5,879 (TEX); Armstrong
Co., Palo Duro Canyon, Goodman 5,Z7<S' (TEX);
Atascosa Co., 1 mi N of Pleasanton, Tharj) and
Johnston 541,745 (TEX); Bastrop Co., 2 mi NE
of Cedar Creek, McCart 5,725 (TEX); Baylor
Co., Sevmour, Reverd.on 3,440 (US); Bexar Co.,
San Antonio, near Lake Mitchell, Hitchcock
1,269 (UC); Brooks Co., 2.5 mi E of Falfurrias,
Gould, Reeves, and Morrow 6,603 (TEX, UC);
4 mi SE of Encino Division, King Ranch, Gould
and Morroic 6,720 (TEX, UC),' Burnet Co., 3
mi E of Buchanan Dam, Gould, Brown, and
Celarier 5,476 (UC); Caldwell Co., 8 mi E of
Luling, M. C. Johnston 6,215 (TEX); Cam-
eron Co., Laguna Atascosa Wildlife Refuge,
Fleetwood 3,472 ( TEX ) ; Colorado Co., Colum-
bus, Anderson 3,282a (HSC); Comal Co., E of
New Braunfels, near Mission Valley Mills, An-
derson 3,888 (HSC); Duval Co., E of Crestonia,
Barkley 13,003 (TEX); Ector Co., 18 mi W
of Odessa, Warnock 7,902 (TEX); Frio Co., 4
mi E of Zavala Co. line on Hwy. 76, M. C.
Joluuton 6,192a (TEX); Garza Co., E of Post,
Tharp and Gindnede 51-1,562 (TEX); Gon-
zales Co., Athens, Tharp and Higdon s.n., 23
Apr 1938 (TEX); Hall Co., near Estelline,
Gould 7,721 (UC); Hidalgo Co.: McAllen,
vacant lot, Anderson 3,876 (HSC); Santa Ana
National Wildlife Refuge, M. C. Johnston
92
Bbicham Young University Science Bulletin
Fig. 56. Ckloris cucutlata and C. .erticiUan.. (A-K) C. c^:M.a. , A) h.hit x 1 /4 ^^^^^l^^^ :^^'fZ
^ (left) and lower side (right) x 4; (C) spikelet, partly chssecte^^^^^ lW-V)C. verticillata. (L) habit,
above, X 10; (F-J) florets, showing variation, x 10; (K) ^^^^P'"'.!/, '^ ^m) spikelet partly dissected,
X 1/4- (M) portion of spike, upper side left and lower side (>'ght , x 4. ( I\ ) .spiKeier, pa y
X 10 (O) sterile floret, x l6; (P-T) florets, showing variation, x 10; (U) caryopsis. x 10.
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chlohls (Ghamineae)
93
o=C VERTICILLATA
•=C.CUCULLATA
o =C. ANDROPOGONOIDES
Fig. 57. Distribution of Chloris cucultata (shaded
circles), C. verticillata (dotted circles), and C.
andropogonoides (open circles).
.5.3-674 (TEX); Jim Hogg Co., 30 mi S of Heb-
bronville, Tharp 5,912 (TEX); Jim Wells Co.,
Romarsid Ranch, Freeborn ami Freeborn 524
(TEX); Karnes Co., Karnes Citv, ]â– C. Johnson
813 (TEX); Kenedy Co., 2 mi 'S of Sarita, An-
derson 3,305 (HSC); King Ranch, Norias Divi-
sion, M. C. Johnston 5.319.77 (TEX); Kleberg
Co., King Ranch, Santa Gertrudis Division,
M. C. Johnston 5.319.76 (TEX); King Ranch,
Laureles Division, M. C. Johnston 5.319.81
(TEX); 2 mi N of Kingsville, Swallen 10,085
(US); La Salle Co., ,3 mi S of Dille\-, Painter
and Barkley 14,310 (TEX); Live Oak Co.,
Tharp and Gimbrede 52-5.52 (TEX); Llano
Co., Enchanted Rock, Web.ster 47,261 (TEX);
McMullen Co., 10 mi S of Tilden, Alvarez et al.
7,714 (TEX); Mason Co., 18 mi N of Fredricks-
bm-g, Emert/ 814 (TEX); Midland Co., Mid-
land, Tracy 7.962 (TEX); Nolan Co., 10 mi
S of jet. of Hms. 126 and 53, Anderson 4,651
(HSC); Nueces Co., ca 1 mi N of Kingsville,
Anderson -3.302 ( HSC ) ; Corpus Christi. ^Heller
s.n., 14-21 Mar 1894 (UC, US); Mustang Is-
land, /4 mi S of Port Aransas, Anderson 3,293
(HSC); Pecos Co., 6 mi N of Ft. Stockton,
Warnock T464 (TEX); Randall Co., Palo Duro
Canyon, Reed -3,984 (TEX); San Patricio Co.,
Welder Wildlife Refuge, near Sinton, Gould
and Hijcka 8,020 (UC); Starr Co., 6 mi NW
of La Gloria, Sanchez et al. 8,3-33 (TEX);
Stonewall Co., Double Mt., Tharp and Gim-
brede 51-1,551 (KANU, TEX); Tarrant Co.,
Ft. Worth, McMullen s.n., Aug-Oct 1927
(TEX); Taylor Co., Abilene State Park, Hen-
derson 6.3-899 (TEX); Terrell Co., 2.5 mi S of
Sheffield, Gould 9,699 (TEX, UC); Travis Co.:
Camp Mabry, Austin, Tharp s./i., 21 Apr 1938
(TEX); near Bee Caves, jet. of Hwys. 620 and
71, Anderson 3,273 (HSC); east end of Mans-
field Dam, Anderson 3,274 (HSC); Val Verde
Co.: 9 mi S of Del Rio, along Hvvy. 277, An-
derson 4,466 (HSC); 35 mi N of Del Rio, near
Loma Alta, Warnock and McBryde 15,154
(TEX); Ward Co., 1 mi NE of Sand Hills,
Rose-Innes and Moon 183 (TEX); Webb Co.:
Laredo, Martinez and Trevino 3 (TEX); 40
mi NW of Laredo, Emery 668 (TEX); Wichita
Co.: Wichita Falls, McKce s.n.. Sept 1927
(TEX); Red River, above Burkburnett, Tharp
s.n., 20 Julv 1921 (TEX); Willacv Co.: 2.5 mi
S of Ytturia, Anderson 3,311 (HSC); Redfish
Bay, Tharp s.n., 5 Mar 19.34 (TEX); 7 mi N
of Ravmondville, Tharp and Brown 3,2-32
(TEX)'; Wilson Co., Parks -s.n., 25 Dec 1946
(TEX); Zapata Co., 19 mi S of Laredo, Mar-
tinez and Trevino 37 (TEX).
39. CHLORIS VERTICILLATA Nuttall, Trans.
Amer. Philos. Soc, n.s. 5:150. LS37.
(HOLOTYPE; ". . . Arkansas. Wm. Nuttall"
Photograph of tvpe, from PH, in US! ISO-
TYPE: K! ) Fig. 56, L-U.
Perennial 14 to 40 cm tall, tufted, erect or
decumbent, sometimes rooting at the lower
nodes; sheaths glabrous; ligule a ciliate crown
with some hairs up to 2 mm long; blades to
15 cm long, 2 to 3 mm wide, glabrous to scab-
rous; spikes ten to sixteen, 5 to 15 cm long,
usuallv in several verticils, these separated by
5 mm or more, inflorescence terminating in a
vertical spike; spikelets sparsely inserted, aver-
aging ca four to seven per cm of the scabrous
rachis, appressed; glumes lanceolate, membra-
nous, glabrous except for the scabrous midnei-ve;
first glume 2 to 3 mm long, ca 0.3 mm wide;
second glume 2.8 to 3.5 mm long, 0.3 to 0.4
mm wide; fertile lemma 2.0 to 3.5 mm long,
1.5 to 1.9 mm wide, elliptic to lanceolate, callus
bearded, keel glabrous to appressed-pilose,
margins appressed-pubescent, apex acute to
somewhat obtuse, awn 4.8 to 9.0 mm long;
sterile floret one, 1.1 to 2.3 mm long, 0.5 to
1.0 mm wide, oblong, somewhat inflated, trun-
94
Brigham Young University Science Bulletin
cate, glabrous, avvn 3.2 to 7.0 mm long; caiyop-
sis 1.3 to 1.5 mm long, ca 0.5 mm wide, ellip-
soid, rounded to trigonous; chromosome num-
ber 2n = ca 28, 40, 63.
Chloris verticillata resembles the Austrahan
C. trtincata; it differs in having a single sterile
floret (rarely two). Chloris verticillata also has
several somewhat remote verticils of spikes
terminated by a single erect spike, while C.
truncata has a single verticil and lacks a ter-
minal spike.
Chloris verticillata, C. ondropogonoides, and
C. cucullata introgress in Texas. A discussion of
this follows the treatment of C. ondropogo-
noides.
Chloris verticillata (commonly called "wind-
mill grass") is widely distributed in the central
United States, where it is commonly found as
a weed of roadside, lawn, or waste areas ( Fig.
57, dotted circles). Disturbance, cultivation, and
overgrazing have largely eliminated much of
the native vegetation in this area, but earl\
records (Barkley, 1965) indicated that windmill
grass was found in generally low areas of the
native prairie.
Representative specimens examined: UNIT-
ED STATES: Arizona: Gila Co., 7 mi W
of Young, anon. (UC). Colorado: Baca Co..
Sand Canvon, 25 mi S of Pritchett, C. L. Por-
ter 4,272 (TEX); Che\enne Co., near Cheyenne
Wells, Harrington and Smith 384 (UC); Crow-
ley Co., Rocky Ford, Harrington 685 (UC);
Powers Co., 10 mi W of Lamar, Harrington 724
(UC). Iowa: Story Co., Ames, /. P. Smith 2,597
(HSC). Kansas: Anderson Co., 3 mi S of Gar-
nett, Horr and McGregor 3,591 (KANU, TE.X);
Barber Co., 1 mi W of Medicine Lodge, Mc-
Gregor 14,029 (KANU); Bourbon Co.,1 mi S
of Uniontown, Ungar 111 (KANU); Chase Co.,
4 mi E of Marion, Horr 3.497 ( KANU, TEX ) :
Chautauqua Co., roadside grassland, Horr 1,540
(KANU); Clav Co., waste ground, Imler s.n.,
14 Jun 1929 (KANU); Coffee Co., Burlington,
Ander.wn 2,765 (HSC); Dickinson Co., without
precise location, Imlcr s.n., 15 Jun 1929
(KANU); Ellis Co., 6 to 7 mi SVV of Hays,
Rijdberg and Imler 1,248 (KANU); Ellsworth
Co., 1 mi E of Terra Cotta, McGregor 9,212
(KANU); Finney Co., Garden City, Kellerman
s.n., 20 Aug 1882 (KANU); Ford Co., 1 mi
NW of Ford, Hulhert 3.207 (KANU); Gear)-
Co., waste land, Imler s.n., 15 Jun 1929
(KANU); Grant Co., Thompson 55 (UC);
Greenwood Co., 3.5 mi W or Woodson Co. line
on Hwy. 54, Lathrop 3.186 (KANU); Harvev
Co., 8.7 mi E of Newton, Harms 2,192 (KANU,
UC); Kiowa Co., 7 mi SE of Haviland, Horr
and McGregor 3,785 (KANU, UC); Meade Co.,
County Park, Horr 3,272 (KANU); Montgom-
ery Co., 1 mi N of Independence, along Hwy.
7.5, Anderson 2,761 (HSC); Morris Co., 4 mi
SE of Council Grove, Richards 1,788 (KANU);
Morton Co., 7.3 mi N of Elkhart, Richards
2,492 (KANU); Neosho Co., 5 mi E of Erie,
Holland 488 (KANU); Osage Co., Lyndon,
Anderson 2,766, 2,767 (HSC); Pottawatomie
Co., without precise locality, Imler s.n., 13 Jun
1929 (KANU); Republic Co., T4S, R2W, sec.
22, Morley 1,227 (KANU); Riley Co., without
precise location, Gates 12,801 (UC); Shawnee
Co., near Wakarusa, Anderson 2,768, 2,769
(HSC); Stafford Co., 8 mi NE of Hudson, Mc-
Gregor 10.515 (KANU); Wichita Co., grass-
land, Agrelius .'i.n., 12 Aug 1912 (KANU);
Wilson Co., 8 mi N of Neodesha, along Hwy.
75, Anderson 2,762, 2,763 (HSC); Woodson
Co., T24S, R14E, sec 21, Lathrop 471 (KANU).
Missouri: Atherton, Bush 7,352B (TEX).
Nebraska: Cherry Co., Valentine, Tolstead
4-463 (NEB); Dundy Co., 3 mi S of Benkel-
man, Andcr.wn 2,774, 2,776 (HSC); Hitchcock
Co., Swanson Reservoir, Anderson 2,772 (HSC);
Nemaha Co., Auburn, Andenson 2,770 (HSC);
Nickolls Co.. Superior, Bates 6,635 (NEB);
Redwillow Co., Indianola, Anderson 2,771
(HSC). New Mexico: Cuny Co., 14 mi W of
Melrose, along Hwy. 60, Anderson 4.663 (HSC);
San Miguel Co., Las Vegas, Whitehouse s.n.,
3 Sept 1929 (UC). Oklahoma: Cimarron Co.,
6 mi E of Kenton, Rogers 5,933 (TEX); Cotton
Co., 5 mi N of Burkbumett, Rose-Innes and
Moon 973 (TEX); Muskogee Co., rockv soil,
Behh s.n.. H Sept 1940 (TEX); Payne Co., W
of Stillwater, E.stes 57 (TEX). Texas: Ander-
son Co., Engeling Wildlife Management area,
Calley and Mar.^i 13 (TEX); Archer Co., 7
mi S of Windthorst, Anderson 3.921 (HSC);
Bell Co., old Tennessee Valley community site,
York and York 54,447 (TEX); Brewster Co.,
24 mi E of Alpine, along Hwy. 90, Ander.wn
4.447 (I ISC); Altuda Pa.ss, Glass Mts., Warnock
W362 (TEX); Burnet Co., Longhoni Caverns
State Park, near Marble Falls, Silvetis 7.642,
7,643 (TEX); Clay Co., Prospect Hill, Silveus
117 (TEX); Comanche Co., DeLeon, Ander.wn
3,919 (HSC); Culberson Co., McKittrick Can-
von, Guadalupe Mts., Fischer s.n., 20 Jul 1950
(KANU); Dallam Co., 21 mi NW of Dalhart,
Rowell 5,388a (TEX); Dallas Co., Dallas, com-
mon on prairie, Bii.^li 1.164 (US); Garza Co.,
3 mi E of Post, Tharp and Gimhrede 51-1,558
(TEX); Gray Co., 5 mi W of Pampa, Brown
3,382 and Tharp GR 239 (TEX); Hays Co., W
of Wimberly, /. C. Johnson 258 (TEX); Hemp-
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
95
hill Co., 5 mi E of Canadian, Rowell 4.078,
4,174 (TEX); Howard Co., Big Spring, Hitch-
cock 13,366 (US); Hudspeth Co., 4 mi NW of
Sierra Blanca, Anderson 4,420 (HSC); Jones
Co., eastern part of county, Tharp and Ginihrede
51-1,556 (TEX); Lampasas Co., N of Lampasas,
Silveus 7,540 (TEX); Menard Co., Ft. Mc-
Cavett, Tharp 5,224 (TEX); Mills Co., Priddy,
Anderson 3,918 (HSC); Nolan Co., 10 mi S of
jet. Hvv)'s. 126 and 53, along .53, Anderson
4,650 (HSC); Palo Pinto Co., 0.5 mi N of
Stravvn, Anderson 3,920 (HSC); Parker Co..
SW of Weatherford, Silveus 7,543 (TEX);
Pecos Co., 5 mi E of Brewster-Pecos Co. line,
along Hwy. 90, Anderson 4,460 (HSC); Potter
Co., N of Amarillo, Brenckle 48,302 (US);
Randall Co., near Palo Duro Canvon, young
s.n.. Sept 1917 (TEX); Runnells Co., 0.5 mi
N of Ballinger, Anderson 4,641 (HSC); San
Saba Co., Richland Springs, Anderson 3,333
(HSC); 2.1 mi SE of San Saba, Cory 58,249
(UC); Sherman Co., 18 mi S of Stratford,
Weaver and McLaughlin, without number or
date (TEX); Sterling Co., Sterling Citv, park,
Anderson 3,002 (HSC); Tarrant Co., near Ft.
Worth, Ruth 302 (NY, UC); Taylor Co., Camp
Berkeley, Tolstead 7,445 (UC); Travis Co., Bee
Caves, Anderson 3,272 (HSC); Wichita Co.,
Burkburnett, Anderson 3.923 (HSC); Wichita
Falls, McKee s.n., June 1927 (TEX); William-
son Co., N of Leander, Johnston and McCart
5,279 (TEX).
40. CHLORIS ANDROPOGONOIDES Four-
nier, Mex. PI. 2:143. 1886. (HOLOTYPE:
"Mexique: Prov. de San Luis. Virlet d'Aoust.
1851. 1462." P! The specimen is depau-
perate, probably from overgrazing, but the
spikelets are those of the species as com-
monly understood.) Fig. 58, D-K.
Chloris tcmiispica Nash, Bull. Torrev Bot.
Club 25:436. 1898. (HOLOTYPE:
". . . Nealley . . . 1889." US! )
Perennial 10 to 40 em tall, tufted to shortb
stolon if erous; sheaths glabrous; ligule a short,
naked crown; blades ca 1 mm wide, to 1.5 cm
long, glabrous to scabrous except basally sparse-
ly pilose; spikes six to thirteen, 4 to 15 cm long,
usually radiate in a single series, occasionally
a second poorh' de\eIoped verticil produced
above; spikelets distant, ca four to seven per
cm of the scabrous rachis, appressed; glumes
narrowly lanceolate, acute, thin, glabrous ex-
cept for the scabrous midnei-ve; first glume 2.0
to 2.3 mm long, ca 0.2 mm wide; second glume
3.0 to 3.3 mm long, ca 0.3 mm wide; fertile
lemma 1.9 to 2.7 mm long, 0.5 to 0.6 mm wide,
narrowly lanceolate to elliptic, apex acute, callus
bearded, margins and keel appressed-pilose,
otherwise glabrous, awn 1.9 to 5.2 mm long;
sterile floret one, 0.9 to 1.7 mm long, 0.25 to
0.5 mm wide, narrowly cylindrical, apex obtuse,
glabrous, awn 2.5 to 3.5 mm long; caryopsis
1.3 to 1.4 mm long, ca 0.4 mm wide, ellipsoid,
rounded to trigonous; chromosome number
2n=40.
Chloris andropogonoides is similar to C. vct-
ticillata and C. texensis. It may be separated
from the former by its single verticil of spikes,
generally shorter awns, and narrower sterile
florets (Figs. 56, L-U; 58, A-C, D-K). Addi-
tional differences are given in Table 11.
Chhris andropogonoides is generally smaller
in spikelet dimensions than C. texensis but dif-
fers more obviously in having spikes that are
floriferous to near the base, while the spikes
of C. texensis are naked for several cm above
the base.
As discussed in the following section, Chloris
andropogonoides, C. cucullata, and C. verticil-
lata are involved in extensive h\bridization and
introgiession in Texas.
Chloris andropogonoides is restricted primar-
ilv to grassy roadsides and prairie relicts of the
coastal plain of southern Texas and northeastern
Mexico (Fig. 57, open circles). Occasional very
wideh' separated populations may be found far-
ther inland, as at Rock Springs, Texas (Fig. 64).
Representative specimens examined: MEX-
ICO: Coahuila, Musquiz Palm Canyon, Marsh
984 (TEX); Tamaulipas: 1 mi E of Ejido de
San Lazaro, 24°.35'N, 99°13'W, Graham and
Johmton 4,291 (TEX); El Canelo Ranch, 24 mi
N of San Fernando, M. C. Johnston 4,880
(TEX). UNITED STATES: Texas: Aransas
Co., Rockport, A. Chase 6,067 (US); Atascosa
Co., 3 mi S of Hindcs, M. C. Johnston 6,197
(TEX); Bee Co., 2.6 mi N of Skidmore, An-
derson 2,699 (HSC); Bexar Co., 7 mi N of San
Antonio, M. C. Johnston 2,409 (TEX); Cald-
well Co., near Luling, Silveus 4,170- A (TEX);
Cameron Co.: 10 mi E of Brownsville on Hwy.
4, Anderson 3,318 (HSC); Laguna Atascosa Na-
tional Wildlife Refuge, Fleetwood 3.374 (TEX);
levee of Resaca de la Cringa, M. C. Johnston
542.186 (TEX); Comal Co., E of New Braun-
fels, near Mission Valley Mills, Anderson 3,889
(HSC); Edwards Co., substation no. 14, Cory
52.467 (US); Hays Co., 2 mi W of San Marcos,
Emery 835 (TEX); Hidalgo Co., Santa Ana
National Wildlife Refuge, Fleetwood 3,161,
3,373 (TEX); Jim Wells Co., King Ranch, Santa
Gertrudis Division, M. C. Johnston 542,104
96
Bbicham Young University Science Bulletin
Fig. 58. Chloris texemis and C. andropogonoides. (A-C) C. 'texcnsis. (A) habit, x 1/4; (B) spikelet, partly
dissected; (C) sterile floret. (D-K) C. andropogonoides. (D) habit, k 1/3; (E) spikelet, partly dissected;
(F-H) fertile lemmas, showing variation; (I) sterile floret; (J) group of sterile florets, showing variation;
(K) caryopsis. (B, C, E-K) .x 10.
BiOLOCiCAi. Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
97
Table 11. Characteristics differentiating Chloris cucullata, C. verticillata, and C. andropogonoides.
Characteristic
C. cucu
llata
C. verticillata
C. andropogonoides
Height
15-60 cm
14-40 cm
10-40 cm
Length of spikes
2-5 cm
5-15 cm
4-15 cm
Number of verticils
2 or 3
2-5 cm
usually 1
Relative distance
between verticils
very close together
, ap-
usually well separated,
pearing to
arise
from
5 mm or more apart
same point
Number of spikelets per
cm length of spike
14-18
4-7
4-7
Lemma length
1.5-2.0 mm
2.0-3.5 mm
1.9-2.7 mm
Lemma awn length
0.3-1.5 mm
4.8-9.0 mm
1.9-5.2 mm
Sterile floret
Length
1.0-1.5 mm
1.1-2.3 mm
0.9-1.7 mm
Width
1.0-1.5 mm
0.5-1.0 mm
0.25-0.5 mm
Length/Width
0.6-1.5
1.6-3.0
2.7-4.8
Length of awn
0-1.5 mm
3.2-7.0 mm
2.5-3.5 mm
Method of
seed dispersal
spikelets falling
from
tumbleweed, culm de-
tumbleweed, culm dehisces
spike at maturity
hisces at uppermost
node, allowing inflores-
cence to break from
plant at maturity and
roll
at uppermost node, allow-
ing inflorescence to break
from plant at maturity and
roll
(TEX); Alice, /. G. Smitli s.n., 7 June 1897
(US); Kenedy Co., King Ranch, Norias Divi-
sion, Swallen 10,211 (US); Kerr Co., Kerrvillc,
Hitchcock .5,26.5 (UC); Lacey's Ranch, Palmer
11,230 (UC); Kleberg Co., King Ranch, Laure-
les Division, M. c/]ohnMon 541,14" (TEX);
between Kingsville and Ricardo, Swollen 10,091
(US); Nueces Co., near Agua Dulce Creek,
10 mi W of Chapman Ranch, Anderson 3.297
(HSC); San Patricio Co., Portland, Silveus 50
(US); Starr Co., 1 mi S of Rio Grande City,
Anderson 4,524 (HSC); N of Roma, Tharp
47,438 (TEX); Travis Co., Barton Springs,
Higdon 5 (TEX); Willacy Co., E of Ra\mond-
ville, Silveus 7,313 (TEX); Lyford, ' T/iar/)
7,644 (TEX, US).
Hybridization and Introgression
Involving Chloris cucullata, C. verticillata,
and C. amlropogonoides
The identification of specimens of Chloris
from central and southern Texas often presents
a difficult problem as characteristics of C. cucul-
lata, C. verticillata, and C. andropogonoides
seem to blend and rccombine in various ways.
"Pure" populations are well defined and may
be keyed with few problems. The characteris-
tics of such populations are presented in Table
11 and Figure 59.
Of these characters, the length of the lemma
awn and the width of the sterile floret gave the
best separation of all three species; these fea-
tures were chosen as the main axes of the
scatter diagrams to follow. Of the remaining
characters, the best for distinguishing all three
species are the length of the spikes, the number
of spikelets per unit length of the spike, the
distance between successive verticils, and lemma
shape. These are indicated on the scatter
diagrams by radiating lines attached to the
gl\ph.
Fig. .59 shows the disposition of selected
specimens of Chloris cticullata. C. verticillata,
and C. andropogonoides plotted b)- using the
characters described above. Each sample en-
tered into Fig. 59 is a single specimen chosen
; 1 GO
75-
O
_i
y 50-
U
(/I 25-
• zC. CUCULLATA
©tC verticillata
O.C ANDROPOGONOIDES
SPIKELETS PER CM OF SPIKE
o
14-16
<? 9
9-13 4-8
6 6
LENGTH OF SPIKES (IN CM )
o
3-5
-\
[IN CM)
LEMMA LENSTh/wiDTH O
11-21 2 2-32
-o -o
13-1,7 18-22
10 20 30 40 50 60 70 80
LEMMA AWN LENGTH ( IN MM)
Fig. 59. Scatter diagram of selected specimens repre-
senting pure populations of C. cucullata (upper
left), C. verticillata (right middle), and C. andro-
pogonoides (lower middle).
Brigham Young University Science Bulletin
from populations containing only one of the
three species. In the case of C. verticillata, this
presented httle problem as the natural range
of this species extends far to the north of the
ranges of the other two (Fig. 57). Tlius, the
samples of C. verticillata used in Fig. 59 are
representative of populations from Oklahoma,
Kansas, Nebraska, and Colorado. The examples
of C. cucuUata and C. ondropogonoides chosen
for Fig. 59 were selected from those popula-
tions in Texas that had plants of only one
species present and in which the other species
were not present nearby.
From Fig. 59 it may be seen that the three
species are well defined. Chloris cucullata
stands well apart from the other two taxa on
the basis of all six characters used. Chloris ver-
ticillata and C. ondropogonoides are somewhat
closer but are easily separated by the simultane-
ous use of sterile floret width and lemma
awTi length (see Fig. 56, L-U, 58, D-K). In
addition, C. andropogoiwides usually has longer
and narrower lemmas than C. verticillata. The
dashed lines drawn around each of the three
species groupings in Fig. 59 represent the
approximate range of variation of lemma ami
length and sterile floret width on the specimens
studied; these will be reproduced as frames of
reference in the scatter diagrams to follow.
Chromosome numbers of 2n = 40 have been
rejTorted several times for all three taxa. A
chromosome number of 2n^ca 2S has been
reported once by Gould ( 1960).
The collection sites sampled for cytok)gicaI
and mass collections of h\'brid populations are
shown on the map in Fig. 60 and listed in Table
12. For convenience of discussion, these areas
are given the names of nearby landmarks or
population centers.
In the illustrations of individual populations
following, unshaded squares ( [U ) represent
• KASON CO
ZiLKER PWK^PATTEBSm PARK
# EDWARDS CO •*
•EASTROP ST PARK
• SAN P*RCOS
RIO GRANDE C|T>
• SKtCrDRE
Fig. 60. Location of collecting areas sampled for in-
trogression studies.
those plants in which virtually all of the pollen
mother cells examined showed normal meiotic
figures. FulK shaded s([uares (â– ) depict those
plants in which nearly all of the pollen mother
cells or their products showed meiotic irregu-
larity, including lagging chromosomes or
chromosome fragments, micronuclei, multiva-
lents, or excluded chromosomes. Half-shaded
scjuares ([J) exemplif}- those plants in which
less than 20 percent of the PMC's or their
products showed meiotic aberrations. At least
25 PMC's were examined in detemiining the
relative amount of meiotic regularity or irregu-
larity of each plant plotted in the diagrams.
The following discussion of populations rep-
resents the different combinations of the three
species involved: the three possible pairs of
species and the combination involving aJll three
species.
Table 12. Populations utilized in introgression studies {all from Texas)
Name of population
Species involved
Fig.
Bastrop State Park
Edwards Co. ( Rock Springs )
Kinney Co.
Mason Co.
Nolan Co.
Patterson Park
Pawelekxille
Pettus
Raymondville
Richland Springs
Rio Grande City
San Marcos
Skidmore
Zilker Park
anomalous
C. ondropogonoides x C. verticillata
anomalous
C. cucuUata x C. verticillata
C. cucullata x C. verticillata
C. cucullata x C. verticillata
C. cucullata x C. andropogonoides
C. cucullata x C. andropogonoides
C. cucullata x C. andropogonoides
C. cucullata x C. verticillata
C. cucullata x C. andropogonoides
anomalous
anomalous
C. cucullata x C. andropogonoides x C. vcrticiUata
66
64
68
61
63
67
65
Biological Series, Vol. 19, No.
Taxonomy of the Cenu.s Chloris (Gramineae)
99
Chloris cuctiUata x C. verticiUata
This Inbiid t\pe is well represented in Pat-
terson Park, a city park in the northeast section
of Austin, Texas (Fig. 60). In this area Chloris
cucuUata and C. verticiUata and their introgres-
sants fonn extensive colonies in distinbed areas,
along roadwa)'s, paths, and in neglected sec-
tions of the park. This population is illustrated
diagrammatically in the scatter diagram shown
in Fig. 61 and in the photograph in Fig. 62.
The scatter diagram of the Patterson Park
population shows a number of plants corre-
sponding in all major characters with the "pure"
populations of C. cucuUata and C. verticiUata.
In addition, there are man\- intennediate forms
that variously recombine the characters of these
two species. In those specimens studied cyto-
logicallv, the plants lying within the range of
expected variation of the pure species have nor-
mal meiotic behavior, while those studied from
the intermediate group show, for the most part,
high percentages of irregular meiotic figures.
The Patterson Park population is just one of
several showing this pattern of introgression.
Others with similar characteristics, when repre-
sented in a graph, include Mason Co., Nolan
Co., and Richland Springs ( Fig. 60).
Chloris cucuUata x C. andropogonoides
This population, represented graphical K in
Fig. 63, is found in a relatively stable grassland
community along a railroad right-of-wa\- near
Raymond\ille, in southern Texas (Fig. 60).
Many individuals of Chloris andropogonoides
and C. cucuUata, as well as intermediate plants,
are found in this area. Two other species of
Chloris, C. gatjana and C ciliata, are also com-
M4^^:^4^
i 1 I 1 ) â– t ' f
Fig. 62. Silhouette photograph of representative in-
florescences of Chloris populations from Patterson
Park, Travis Co., Te.xas.
mon in this area but are not involved in the
parentage of any of the hybrids.
Again, the intermediate plants show a high
frequency of meiotic irregularity. Unfortunate-
ly, preserved anthers of the parental species
were too young for meiotic study. Several of
the intermediate plants studied, however,
showed higher chromosome numbiTs than the
2n— 40 reported for the parental species. Dip-
loid counts of 2n = ca 60 were most frequent,
though the high degree of irregularity made
exact counts impossible. Plants with 2n = ca 60
could ha\e resulted from the combination of
reduced and unreduced gametes.
In this population the intennediates are
well represented in the lower left quadrant
of the graph. By comparison, the Patterson
Park diagram ( Fig. 61 ) shows a poor represen-
tation of intennediates in this section. This shift
is largely influenced by variation in awn length
and is to be expected in view of the differences
OF CELLS EXAMItCO Q PLl
5 1,00
<? 9 :
I ALMOST ALL CELLS E
NTS NOT STUDIED CYTOLOGfCAL LV
SPLKELETS PER CM OF SPIKE Q Q
(AMINEO HmEOSiS NORMAL BmEOTIC IRREGULARITY B MECTC iRREGULAR.T>
LENGTH OF SPIKES (I
CMJ
o
DISTANCE BETWEEN UPPeRMOST O
AND LOWERMOST VERTICIL O 144
(IN CM.)
LEfvWA LENGTm/wiDTM ^^
^^<?-9^â– ^-<f-
-f^^f^^+
20 30 40 50 60
LEMMA AWN LENGTH (IN MM)
Fig. 61. Scatter diagram of population of Chloris citnil-
lata X C. verticiUata from Patterson Park, Austin,
Te.xas.
3l 25
glOO
OPLANTS NOT STUOED criOLOGICALLT
SPWELETS PER CM OF SPIKE Q
ALMOST ALLCELLS EXAMINED
LEN3TM OF SPIKES t
o
â– 16 9-13 â–
6 i
o :
coo! <> Y
oo; <>-i
-â– -â– -'o^<X) ,
DISTANCE BETWEEN LiPPERmOST O Q- .O—
AND LOWERMOST VERTICIL O-lO U-2t 22-32
(IN CM.)
,8'^2 13^.7 l.8"^2
LEMMA length/width
^O.^-^
T-
'A
20 30 40 50 60
LEMMA AWN LENGTH (IN MM)
Fig. 63. Scatter diagram of populations of Chloris
cucuUata x C. andropogonoides from near Ray-
mondville, Texas.
100
Bricham Young University Science Bulletin
in awn length of the putative parents (see
Table 12).
Several additional collection sites had popu-
lations with intiogression patterns similar to
that of Raymondville, including Pavvelekville,
Pettus, and Rio Grande City (Fig. 60).
Chloris andropogonoides x C. verticillata
The collection site for this population is in
Edwards County, Texas, on the grounds of the
Texas A and M Experimental Station about 25
miles northwest of Rock Springs (Fig. 60).
While "good" Cliloris andropogonoides and C.
verticillata arc present in abundance, C. ciicul-
lata was not observed in the area. The mass
collection of plants made in this area is repre-
sented in Fig. 64.
Unlike previous scatter diagrams, the inter-
mediate populations shown in Fig. 64 are not
of obvious parentage. On the basis of the
species present, it would be expected that the
recombinants would fill the areas between C.
verticillata and C. andropogonoides in the
graph, though certainly more long-awned inter-
mediates similar to C. andropogonoides are
present than in either of the previous two popu-
lations discussed. The unexpected position of
the remaining intennediate plants might be the
result of the presence at some time in the past
of C. cucuUata in the area, still showing its in-
fluence on the remaining population. A more
likely possibilit\', however, is that the inter-
mediates merely represent recombinants of
some of the more extreme morphological \'ar-
iants of C. andropogonoides and C. verticillata.
This population was collected when the
plants were in seed and cytological studies were
not possible.
I have been unable to find additional popu-
lations that I feel are representative of this
species combination. Chloris cucullata is nearly
ubiquitous in areas in which C. andropogonoides
and C. verticillata arc sympatric and usually
enters into the introgression pattern when
present.
Chloris cucidlata x C. andropogonoides x
C. verticillata
The population represented in Fig. 65 is
from a grassy field utilized intensively as a
recreation area in Zilker Park, a citv park in
the southwest part of Au.stin, Texas. Plants
that can be referred without hesitation to C.
cucullata are common in the area. Others are
clearly close to C. verticillata and C. andro-
pogonoides but are somewhat taller than ex-
pected. An abundant population of inter-
mediates is also present.
175
2,50-
2
5 1254
glOO-
l-
u
O 75
-I
Li.
^ 50H
SP«ELf TS PEn CM OF SPIKE O O O
M-16 9 '3 4-8
LENGTH OF SPKESdNCM) O 6 O
3-^ 6.8 9-11
DISTANCE BETWEEN UPPERMOST O O O-
ANO LOWERMOSTVERTlCiL 040 11-21 ??02
(IN CM)
LEMMA LENGTh/wCTH O "O — O
' 8-12 tJ-17 18-22
2 30 40 50 60
LEMMA AWN LENGTH (IN MM)
Fig. 64. Scatter diagram of population of Chloris
andropogonoides x C. verticillata from the Texas
A and M Experiment Station near Rock Springs,
Texas.
In comparing Fig. 65 with the diagrams
of Patterson Park and RaMiiondville ( Fig. 61 and
63), one can plainly see a number of mutual
features. The Zilker Park population shows
many plants in the lower left and left center
portions of the diagram, in this respect similar
to the Raymondville population, and indicates
a strong influence of C. andropogonoides. It
is also strongly expressed in the upper right-
hand portion of the recombinant cluster, a fea-
tiu-e it shares with the Patterson Park popula-
tion. By contrast, the Raymondville population
is not well expressed in the upper right-hand
portion of the diagram, nor is the Patterson
Park population well represented in the lower
left. Thus, the Zilker Park plants seem to repre-
sent various recombinations of all three species.
Dmeosis normal
in all cells
examined
2
2
2 125
HMEiOTIC iRREGULARITT â– MEIOTIC IRREGULARITY IN ALMOST ALL CELLS EXAMINED
IN ^cr^ or less „
OF CELLS EXAMINED PLANTS NOT STUDIED CYIQLOGKTAllt
SPKELETS PER CM OF SPIKE O Q O
9 13
o
â– -0 O '
"r-,0;
-ft- -
a
a
LENGTH OF SPKESdNCMl O 6 6
3-5 6-8 9 11
DISTANCE BETWEEN UPPERMOST O O O-
ANO LOWERMOST VERTICIL O-IO 11-21 22-32
LEMMA length/width O O — O
e-12 13-17 18-22
„f^^
VO 2.0 30 40 50 60
LEMMA AWN LENGTH (IN MM)
Fig. 65. Scatter diagram of population of Chloris
cucuUata x C. andropogonoides x C. verticillata
from Zilker Park, Austin, Texas.
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
101
As with the otlier species combinations ex-
amined cvtologically, the intemiediate popula-
tions in Zilker Park show a high degree of
meiotic irregularity. Only those plants assigned
to C. cucullata show little or no meiotic aber-
rancy.
Anomalous popidations
Not all populations in the areas of .synipatry
are as clearly understood as those just dis-
cussed, for many colonies of heterogeneous
intemiediate plants in southern Texas coexist
with only one of the putative parents, the other
probable parent having been completely
swamped by introgression and existing only in
a much diluted fonn. In these collections it is,
most frequenth', Chloris androj)og,onoi(h's and
C. vcrticillata that are missing; C. cucullata of-
ten persists in pure form. I have chosen four
examples from among many stiidied as repre-
sentative of these anomalous populations. These
are: Bastrop State Park, Skidmorc, Kinney Co.,
and San Marcos.
The Bastrop State Park population (Fig.
66) is found along a grassy roadside just out-
side the west entrance of the park ( near Bas-
trop, in Bastrop Co. ) . On the basis of other
mass collections examined, this appears to be
the remains of a three-way cross involving C.
cucullata, C. andropogonoides, and C. vcrticil-
lata, with the latter two species swamped.
Chloris cucullata. while still present, is no
longer well defined, for a number of plants
are similar to "pure" C. cucullata but lack cer-
tain character combinations.
The Skidmore population ( Fig. 67 ) was
collected in a low roadside ditch in Bee Co.,
Texas (Fig. 60). It is similar to the one at
5 1.50
2
z
- 125
I
o
S 100
u
ir
O 75-
y
X
50
25
SPiKELETS PER CIV
I op
' O
1 S <â– V <?
OF SPIKE o Q O
14-18 9-13 4-6
LENGTH OF SPIKES (IN CM) O O O
3-5 6-a 9-11
DISTANCE BETWEEN UPPERMDST O O O-
ANO LCWERMOST VERTICl o-lO 1 »-2I 23-32
--^-;
10 2.0 30 40 50 60
LEMMA AWN LENGTH (IN MM)
Fig. 67. Scatter diagram of anomalous Chloris popula-
tion from near Skidmore, Bee Co., Texas.
Bastrop State Park except that only one plant
typical of C. andropogonoidcs was found. The
Skidmore site is out of the natural range of
C. vcrticillata, and the presence of this species
is not to be expected. Chloris cucullata is well
represented, as is a series of intermediates. In
this instance the intermediate plants do not
closely approach C. cucullata morphologically.
The Kinney Co. collection (Fig. 68) is from
open bnishland, associated with such shrubs as
Prosopis and Lcucophijllum. In this population
the intermediates are still more isolated from
C. cucullata, and no plants even approaching
C. andropogonoidcs or C. vcrticillata were
found.
The San iVIarcos collection (Fig. 69) is from
a dr)', grassy roadside ditch near the Blanco
River bridge northeast of San Marcos, Texas,
and about two miles east of the eastern edge
of the Edwards Plateau. This population shows
well-defined Chloris cucullata and C. andro-
QMEOSrs
E>AMl^
NORMAL
I^LLS
ED
JMEOTIC «WDGULAf*<TY BmEIOTIC
IN 20V. OR LESS „
r.r CELLS EXAMINED O PLANTS
IRREGULARITY IN ALMOST ALL CELLS EXAMINED
NOT STUDIED CVTOLOGICALLY
1.75
SPIKELETS PER CM OF SPIKE Q Q Q
M-IB 9-13 d-8
175
5PIKELET5 PER CM OF SPiKE
O
4-18
9
9-13 4-8
6 i
6-8 9-11
5 1,50
LENGTH OF SRKESdNCM.) O O O
2 1.50
r----.
LENGTH OF SPIKES (IN CMJ
o
3-5
2
z
1
3-!. 6-8 9-11
DISTANCE BETWEEN UPPERMOST O Q—
2
z
' 1
DSTANCE BETWEEN UPPERMOST O
AND LOWERMOST VERTICl. O-IO
o- o-
11 -2J 2-2-12
AND LOWERMOST VERTICIL 0-10 \ 12122-3?
(IN CM.)
^1.25
W
<?
(IN CMJ
LEMMA LENGTH /WIDTH O O — O
8-12 13-17 18-22
^125
I 1
LEMMA lENGTm/wiDTM
.b9.
o -o
13-17 13-Z2
o
5 1.00
UJ
1 ,><?
<? Q
o
G
^100
UJ
.75
b.
-0
-0
t
§75
u.
^-^"^
UJ
UJ
i so
UJ
1-
-6
=! 50
UJ
t-
I
1
1
1
.2 5
25
1
1
1
1
1
10
20
30 40
50 60 70 80
10 20 30
4.0
50 60 70
60
LEh
^MA AWN LE
-JGTHdN MM)
LEMMA
AWN
LENGTH (IN MM)
Fig. 66. Scatter diagram of anomalous Chloris popula-
tion from near Bastrop State Park, Texas.
Fig. 68. Scatter diagram of anomalous Chloris popula-
tion from Kinney Co., Texas.
102
Brigham Young University Science Bulletin
DmEOSIS normal flMEfOTC IRREGU.AR1TV
111 Ai I .-CI I c lu on-L no I F^^
â– mcOTIC iRREBULARITT IN ALMOST ALL CELLS EXAMINED
O PLANTS NOT STUDIED CYTOLOGICAlly
2 150
2
Sf^KELETS PER CM OF SPIKE
,0
io
'O
LENGTH OF SPIKFi {IN CM 1 O
3-5
aSTANCe BETWEEN UPPERMOST O
AND LOWEFJMOST VERTICIL 0-TO
(IN CM )
LEMMA length/width O
9 9
9-13 -1-8
6 i
6 - B 9 â– ' 1
o o-
1.1-21 2 2-3.2
:t
^?m^
20 30 40 50 60
LEMMA AWN LENGTH (IN MM)
Fig. 69. Scatter diagnim of anomalous Chloric popula-
tion from near San Marcos, Texas.
pogoiwides, as well as many heterogeneous,
cytologically aberrant intermediates. The posi-
tion of the intermediate plants is similar to the
Zilker Park population (Fig. 65), suggesting
that all three putative parents were present at
one time. I searched the area many times over
a period of several seasons and could not find
well-defined C. verticillata, although plants
similar to that species were found about a mile
north of the site. Since this collection area is
close to the limestone-rich Edwards Plateau,
the seemingly preferred habitat of C. verticil-
lata, one would expect the latter species nearby.
Discussion.
Ecologically, populations with large num-
bers of intennediate plants are most frequently
associated with highly disturbed areas. The
best collecting sites, therefore, are roadside
ditches, waste places, and city parks. None
of the species involved will persist in cultivated
or grazed areas, nor will they compete well
with an established native flora or such ag-
gressive weeds as Johnson grass. Before the
excessive disturbance caused by man, it is likely
that the three species involved were only oc-
casionally found, and then only in such areas
as blowouts, slipouts, and banks and alluvial
areas along creeks and streams. There is little
information about the ecology of these species
before the rampant habitat destruction by man;
however, a report of a grass-collecting trip
through the Kansas prairies by J. B. Norton
in 1898 (Barkley, 1965) indicates that Chloris
verticillata grew along the sides of sloughs and
on low ground but did not compete well with
upland grasses.
Cytological examination of the intennediate
plants shows them to be highly aberrant in
meiosis, producing few stainable pollen grains.
It would be expected on this evidence that the
rate of seedset would be relatively low. This,
however, is not always the case. Plants from
the intermediate populations may actually have
good seedset— in some cases, a high percentage.
Such plants are frequently associated with
stands that are quite uniform morphologically.
The combination of highly aberrant meiotic be-
havior, good seedset, and extreme uniformity
suggests that an apomictic mechanism may be
in operation.
Nomenclature
The wide array of variable intermediate
plants in this complex would thus seem to be
the product of extensive hybridization and in-
trogression, possibly still further complicated
by apomixis. The intermediate plants will key
to Chloris suhdolichostachtja and C. latisqua-
mea in many standard grass floras but frequent-
ly will not fit the descriptions of these taxa
precisely. It seems likelv that these names, in
reality, have been applied to segments of the
much larger hybrid populations and do not
represent biological species in the usual sense.
The extreme variabilit\' of the intennediate
populations presents a difficult noinenclatural
problem. The wide variety of morphological
types represented certainly cannot be individu-
ally named, for this would saturate the litera-
ture with names that carry little biological
meaning and might apply to plants of low
genetic stability. The problems become addi-
tionallv compounded in that there is no way
of telling parts of the hybrid population of C.
cucuUata x C. andropogonoides from parts of
the C. cucuUata x C. verticillata populations
(compare Fig. 61 and 63) and from the popu-
lations involving all three species (Fig. 65).
To use the existing names C. latisquamea and
C. suhdolichostachtja for the intermediates is
unsatisfactory, for there is no way of deter-
mining which, or how many, of the three species
are involved in the ancestry of their types. In
short, these names cannot be precisely defined.
Likewise, the usage of a single name for a com-
plex of morphological forms obscures their bio-
logical nature.
I feel that the only solution is to designate
the intermediate plants by a formula based upon
their probable parentage. Thus, a population
suspected to be the result of C. cucuUata intro-
gressing with C. andropogonoides should have
its intermediate plants designated by the for-
mula name C. cucuUata x C. andropogonoides.
While this will not satisfy those who want a
particular name for a particular plant, it does
Biological Sehies, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
103
satisfy what is, to me, a more important aspect
of taxonomy that is seldom reflected in existing
nomenclature: it helps to define the reasons for
the existence of a taxonomic problem.
The original descriptions of the following
names seem to fit the characteristics of hybrid
plants.
Chloris vei-ticiUata var. aristidata Torrey and
Gray in Whipple, U.S. Kept. Expl. Pacific
2:176. 1857.
Chloris siihdolichostachtja C. Mueller, Bot.
Zeitung (Berlin) 19:341. 1S61.
Chloris verticillata var. intermedia Vascy in
Coulter, Contr. U.S. Nat. Herb. 2:528. 1894.
Chloris hrevispica Nash, Bull. Torrev Bot.
Club 25:4.38. 1898.
Chloris latisquaniea Nash, Bull. Torrev Bot.
Club 25:439. 1898.
Specimens and populations examined (all at
HSC): C. cucullata x C. verticillata. TRAVIS
CO.: Austin, Patterson Park, along Aiiport
Blvd., 0.5 mi SE of its junction with Hwy. 35,
Anderson 3,336, 12 July 1964 (mass collection),
3,338-3.361, 12 Julv 1964 ( cy tovouchers ) .
MASON CO.: along Hw)'. 29, 2.5 mi E of the
junction of Hwys. 29 and 377, 3 mi E of Grit,
Anderson 4,633, 2 Julv 1968 (mass collections).
NOLAN CO.: along' Hwy. 53, 9 mi S of the
junction of Hwys. 126 and 53, Anderson 4,653.
2 July 1968 (mass collecHons). SAN SABA
CO.: Richland Springs, along Hw>'. 190, road-
side ditch near cit)' water tower, Anderson
3,333, 9 Jul\- 1964 (mass collections).
C. cucullata x C. andropogonoides. WIL-
LACY CO.: grassy roadside, 1 mi N of Ray-
mondville, along Hwy. 77, Anderson 3,308.
3,316, 1 July 1964 (mass collections), 3,.m9-
3,315, 1 July 1964, 4,564-4,572, 28 June 1968
(cytovouchers). KARNES CO.: 6 mi S of the
junction of Hwys. 123 and 889 and the village
of Pawelekville, Anderson 4,611, 30 June 1968
(mass collection), 4,595-4,610, 30 June 1968
(cytovouchers). BEE CO.: 3.3 mi N of Pettus,
1.6 mi N of the junction of Hwys. 181 and 798,
Anderson 2,781-2,789, 13 Julv 1964 (mass col-
lections). STARR CO.: 8 mi W of Rio Grande
City, along Hwy. 83, Anderson 3,330. 2 Julv
1964 (mass collection), 3,323-3,329 (cytovouch-
ers); 1 mi S of Rio Grande City, Anderson 4.524.
27 June 1968 (mass collection); ca 1 mi SE
of Arrovo Los Olomos bridge, ca 3 mi SE of
Rio Grande City, Anderson 4,542, 27 June 1968
(mass collection), 4,525-4,541 (cytovouchers).
C. andropogonoides x C. verticillata. ED-
WARDS CO.: Texas A and M Field Station.
25 mi NW of Rock Springs, Anderson 3,903, 12
July 1965 ( mass collection ) .
C. cucullata x C. andropogonoides x C. ver-
ticillata. TRAVIS CO.: Austin, Zilker Park
playing field, Anderson 3,011-3.090, 6 June 1964
(mass collection): 2,603-2,628, 28 June 1963;
2,650-2,664, 3 July 1963; .3,262-3,268, 25 June
1964 (all cytovouchers).
Anomalous populations. BASTROP CO.:
just outside W entrance of Bastrop State Park,
Anderson 3.128-3.138, 10 June 1964 (mass col-
lections), 3,116-3,127, 10 June 1964 (cytovouch-
ers). KINNEY CO.: along Hwy. 693, 1-3 mi
E of its junction with Hwy. 277, Anderson
4,479, 26 June 1968 (mass collection), 4,470-
4,478, 26 June 1968 (cytovouchers). HAYS
CO.: near bridge over Blanco River, 1 mi NE
of San Marcos on Hwy. 21-80, Anderson 3,276,
28 June 1964 (mass collection), 3,278-3.282. 28
June 1964, 4,617-4,619, 30 June 1968 (all cyto-
vouchers). BEE CO.: 2.6 mi N of Skidmore,
Anderson 2.698-2.699. 13 July 1963 (mass col-
lections), 2.691-2.697, 13 July 1963 (cyto-
x'ouchers ) .
4L CHLORIS TEXENSIS Nash, Bull. Torrey
Bot. Club 23:151. 1896. (HOLOTYPE:
"Texas. G. C. Nealley." US! ) Fig. 58, A-C.
Chloris nealleyi Nash, Bull. Torrey Bot.
Club. 25:435. 1898. (Based on Chloris
texensis Nash.)
Perennial 30 to 45 cm tall, cespitose; sheaths
glabrous to sparsely pilose; ligule a low mem-
branous crown; blades narrow, ca 4 mm wide,
up to 15 cm long, scabrous; spikes eight to ten,
up to 20 cm long, radiate, occasionally in two
verticils, bearing loosely imbricate spikelets only
on distal half, naked at base; spikelets ca three
or four per cm of the scabrous rachis; glumes
lanceolate, glabrous except for the scabrous mid-
nei-ve; first glume 2.7 to 3.0 mm long, 0.4
mm wide; second glume 3.5 to 3.8 mm long,
0.5 mm wide; fertile floret 3.7 to 4.3 mm long,
0.7 to 0.8 mm wide, lanceolate to narrowly
ovate, callus bearded, upper margins sparsely
appressed-ciliate, otherwise glabrous, apex
acute, awn 7 to 11 mm long; sterile floret
2.0 to 2.5 mm long, 0.5 mm wide, narrowly
elliptic, apex acute, glabrous, awn 4.5 to 6.5
mm long; caryopsis ca 2.3 mm long, 0.5 mm
wide, ellipsoid, trigonous.
Chloris texensis is most closely allied to C.
verticillata and C. andropogonoides (Fig 56,
L-U; 58, D-K). It differs from C. verticillata in
usually having a single verticil of spikes and
relatively narrow sterile florets. It may be sepa-
rated from C. andropogonoides by the rela-
tively larger spikelets and longer spikes. Chloris
104
Bricham Voung University Science Bulletin
texensis lacks spikelets on the lower portion of
the spikes, whereas both C. verticillata and C.
andropogonoides are floriferous to the base.
Chloris divaricata, a similar Austrahan spe-
cies, is separable in having deeply lobed sterile
florets, scabrous fertile lemmas, and spikes that
are floriferous to the base (Fig. 51, K-R).
Chloris texensis has sterile florets that are mere-
ly acute, not deeply lobed, glabrous fertile
lemmas, and spikes that are naked at the base
(Fig. 58, A-C). Chloris divaricata is apparently
an escape in Texas and has been confused with
C. texensis.
Only six specimens of the species, including
the type, have been found in herbaria con-
sulted. All are from Brazoria, Brazos, or Harris
County, Texas, except the type, for which a
precise collection locality was not provided.
On two separate occasions I searched the
area of two collections, Silveus 423 and 427,
from the A and M Field Station near Angleton.
Most of the land in the area is either culti-
vated, heavily grazed, or in stages of old-field
succession; and I could not locate either C.
texensis or any native vegetation that might
support it. Because neither the plant nor native
vegetation could be found, and because few
specimens have been collected since 1931, I
feel that it is quite likely that the species is
in danger of becoming extinct.
Specimens examined: UNITED STATES:
Texas: Brazoria Co., Columbia, Bush 1,361 (MO,
NY); experimental farm near Angleton, Sil-
veus 423 (TEX); open sandy woods and
swamps, Angleton, Silveus 427 (TEX); Brazos
Co., without precise location, Malone s.n., Oc-
tober 1938 (TAES); HaiTis Co., exposed clay
lowland, 3 mi S of Alvin, Parks 117 (TAES).
42. CHLORIS SESQUIFLORA Burkart, Bol.
Soc. Argentina Bot. 12:287. 1968. (TYPE:
"Dep. Concep. del Unig., bajos cerea del
rio Cualeguaychu, vecindades del Parque
Unzue, A. Burkart. 25,676 . . . .' SI, not
seen. ISOTYPE: HSC!) Fig. 70, AC; 71,
triangles.
Stoloniferous perennial up to 45 em tall;
sheaths glabrous; ligule a short-eiliate crown;
blades up to 20 cm long, 4 to 5 mm wide,
glabrous to scabrous; spikes four or five, 6.0
to 6.5 em long, ascending, somewhat arcuate;
spikelets rather densely inserted, appressed,
imbricate, averaging about 11 per cm of the
scabrous racliis; glumes lanceolate to narrowly
ovate, acute to acuminate, glabrous except for
the scabrous midnerve; first glume 2.1 to 2.2
mm long, 0.3 to 0.5 mm wide; second glume
2.7 to 2.9 mm long, 0.4 to 0.5 mm wide; fertile
lemma 2.1 to 2.9 mm long, 0.9 to 1.1 mm
wide, ovate to elliptic, apex acute, callus short-
bearded, keel and margins densely appressed-
pilose, awn 0.7 to 1.0 mm long; sterile floret
one, ca 1 mm long, 0.3 mm wide, strongly
flattened, membranous, elliptic, awn 0.3 to 0.5
mm long; anthers ca 0.7 mm long.
Only two specimens of Chloris sesquiflora,
including the type, were seen. While it re-
sembles outwardly some of the species with
ciliate lemmas (especially C. ciliata), the flat-
tened, poorly developed single sterile floret and
the appressed pubescence of lemma margins
and keels separate it easily. It is probably not
closely related to C. ciliata.
Specimens examined: PARACUAY: Plaine
de Paraguari, plaques desseckles, dans les ar-
giles impermeables, Balansa 264 (BR).
43. CHLORIS HUMBERTIANA A. Camus,
Bull. Soc. Bot. France 91:63. 1944.
(HOLOTYPE: "Madagascar . . . Mananara
. . . pentes orientales du Vohipaly . . . Hum-
bert, no. 141.50." P! ISOTYPE: K!) Fig.
70, D-F.
Pterochloris humhertiana (A. Camus) A.
Camus, Bull. Mus. Hist. Nat. (Paris) Ser.
2. 29:.349. (Based on Chloris humhertiana
A. Camus.)
Perennial 20 to 30 cm tall, stoloniferous,
with erect culms arising at intei"vals from strong
stolons; leaves basal; sheaths flattened, equitant,
glabrous; ligule a very short-ciliate crown;
blades 1.5 to 2.0 em long, 3 to 5 mm wide,
glabrous, folded, slightly arcuate, apex obtuse;
spikes two or three, 3 to 4 cm long, partially
enclosed in upper sheaths when young, be-
coming exserted at maturity; spikelets imbri-
cate, ca five to seven per cm of the glabrous
laehis; glumes laneeolate-ovate, thin and mem-
branous, glabrous except for the scabrous mid-
nei-ve; first glume 1.0 to 1.5 mm long, ca
0.3 mm wide; second glume ca 1.5 mm long,
0.2 to 0.3 mm wide; fertile lemma 2.3 to 2.5
mm long, 0.7 to 1.2 mm wide, body of lemma
elliptic, with a promini'iit wing at the apex,
making the lemma appear narrowly tnnicate,
callus bearded, margins scabious below, gla-
brous above, internerves densely pubescent vvith
prominent, inflated hairs, awn 2.4 to 3.5 mm
long; sterile floret ca 1 mm long, ca 0.5 mm
wide, elliptic, flattened, apex acute, awn 3.0
to 3.8 mm long, arising below the apex.
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Cm.onis (Gramineae)
105
Fig. 70. Chloric se.squiflora and C. humbertiana. (A-C) C. sesquiflora. (A) habit, .x 1/3; (B) spikelet, partly
dissected, x 10; (C) sterile floret, x 10. (D-F) C. humbertiana. (D) habit, x 1/3; (E) spikelet, partly dis-
sected, X 10; ( F ) sterile floret, x 10.
106
Bhigham Young University Science Bulletin
Chloris humbertiana is a very distinct spe-
cies; its strongly winged lemmas, with inflated
hairs on the intemerves, are unique in the
genus.
Emphasizing these characters, Camus ( 1950 )
proposed the subgenus Pterochloris to include
Chloris humhertitina. Later (1957), Camus ele-
vated Pterochloris to generic rank, retaining C.
humbertiana as its single species.
While Chloris humbertiana is unusual in
Chloris, it is no more so than a number of other
species and I am maintaining it in Chloris.
Only four specimens of this species were
seen in the herbaria consulted and all are from
xerophytic bushland in the Malagasy Republic
(Madagascar).
Specimens examined: MALAGASY REPUB-
LIC: Pateau Mahafetz, Bosser 15,698 (P); Cap
Sainte-Marie, Humbert and Capuron 29,264
(P); d'Amparihy a rembouchuie de la Mcna-
randra, Humbert and Capuron 26,347 (P);
environs d'Ampandrandava, Setjrig 809 (P).
44. CHLORIS AMETHYSTEA Hochstetter,
Flora 38:205. 1855. (ISOTYPE: "... Abys-
sinia leg. Schiinper recu 1855. 1267 . . ."
G!) Fig. 72, A-K.
• = HALOPHILA
A = SESQUIFLORA
Fig. 71. Distribution of Chloric halophila (dots) and
C. sesquiflora (triangles).
Stoloniferous perennial up to 80 cm tall;
sheaths glabrous below, becoming loosely pilose
toward the ligule, especially on the margins;
ligulc a short-ciliate crown, often with longer
hairs near the edges; blades up to 10 cm long,
ca 5 to 6 mm wide, glabrous with scabrous mar-
gins and midnerves; spikes four to six, 5 to 9 cm
long, ascending when young, becoming more
loosely spreading at maturity, usually radiate,
sometimes in two verticils; spikelets appressed,
ca 7 per cm of the scabrous rachis; glumes un-
equal, glabrous except for the scabrous mid-
nerve; first glume 1.5 to 2.5 mm long, 0.2 to 0.5
mm wide, lanceolate, apex acute; second glume
2.5 to 3.8 mm long, ca 0.4 mm wide, linear,
apex acute to mucronate, sometimes somewhat
lacerate, glabrous except for the scabious mid-
nerve; fertile lemma 3.5 to 4.5 mm long, 0.7
to 1.2 mm wide, elliptic to obovate, often pur-
plish tinted, callus densely bearded, margins
densely appressed-pilose, keel sparsely to dense-
ly appressed-pilose, apex acute to obtuse, usu-
ally awnless, occasionally with a short awn less
than 1.7 mm long; sterile floret one, 1.4 to 2.5
mm long, 0.6 to 1.2 mm wide, narrowly turbi-
nate, apex truncate, somewhat inflated, awn-
less; caryopsis ca 2.2 mm long, 0.6 mm wide,
trigonous, ellipsoidal.
Chloris amethijstea seems most closely re-
lated to C. i^aijana and C. mossambiccnsis, dif-
fering from both in having short awns. While
relatively few specimens are available, C.
amethijstea appears to be restricted to Ethiopia
and Kenya (Fig. 73, triangles), occurring in
montane scrub and grasslands at the forest
edge on white or yellowish sandy loams and at
elevations of 3,000-6,000 feet. Associated plants
include Ficus, Acacia, Comhretum, and Com-
miphora.
Specimens examined: ETHIOPIA: Territory
Agow, Dschascha, Hohenacker 2,247 (K, US);
between Assala and Awash Rivers, Semple 19
(US). KENYA: Mt. Marsabit, Adam.wn E. A. H.
11,666 (K, US), Bo<^dan A. B. 4,774 (K);
Stony Athi River, 20 mi SE of Nairobi, Bogclan
AB .3,1.52 (K); Moyale, 3°32'N, .39°03'E, Gillett
12,852,14,065 (K).
45. CHLORIS SUBMUTICA Humboldt, Bon-
pland, Kunth, Nov. Gen. Sp. 1:167. pi. 50.
1816. (There is a fragment, in US!, from
P, that may be from the tv'pe. It is labeled
"Chloris submutica HBK mihi n. "4182.
Mexique." The original description gives
"Tezcucensi" as the collection site, but no
other p:.'rtinent information is given. The
detailed original description and accom-
Biological Series, Vol. 19, No. 2 Taxonomy „i. thf. Genus Chloius (Chamineae)
^' dl^sJ'd" 75 IPvT, '"'^ "^^ .'"'""f^"- (A-K) C. arncthystea. (A) habit, .x 1/6; (B) spikelet, partly
S^n V 7'f' Kf^ ^P'^'''}/h?r"S ^^,"«""" i" ^'^vn length, .x 10; (F-J) sterile florets, showing
™°"' -^ 7.5; W caryopsis, .x 5. (L-V) C. sub,„uUca. (L) habit, x 1/4; ( M ) spikelet, partly dissectecf
X /..b, (N-1) sterile florets, showing variation, x 7.5; (U) caryopsis, x 5. f > l- /
108
Brigham Young University Science Bulletin
Fig. 73. Distribution of Chloris amethystea (triangles)
and C. somalensis (circles).
panying illustration leave no doubt as to
the taxon described.) Fig. 72, L-V.
Chloris pohjstacluja Lagasca, Elenchus
Plantarum 59. 1816. (A fragment of a
specimen, in US! from "Hb. Madrid"
labeled "Chloris polystachya Lag. Nee' iter
Hb Madrid" may be from the type. The
original description is inconclusive. )
Eustachijs suhmutica ( Humboldt, Bonpland,
Kunth) Roemer and Schultes, S)St. 2:614.
1817. (Based on Chloris suhmutica Hum-
boldt, Bonpland, Kunth.)
Chloris inermis Trinius, Gram. Uniflorus
232. 1824. (Based on Chloris pohjstachija
Lagasca. )
Eustachijs polystachya (Lagasca) Kunth,
Rev. Gram. 1:88. 1829. (Ba'scd on C/i/orf.?
polystachya Lagasca. )
Phacellaria panicea VVilldenow ex Steudel,
Noin. Bot. ed. 2. 1:353. 1840, pro syn.
Chloris suhhijlora Steudel, S\n. PI. Glum.
1:206. 1854. (HOLOTYPE: P! A fragment
of an additional Steudel specimen, labeled
by him as C. suhhijlora, is in US!, originally
from CN.)
Chloris pratensis Lesible ex Steudel, Syn.
PI. Glum. 207. 1854. Pro syn., C. suhhi-
jlora.
Perennial 30 to 75 cm tall, erect, usually
tufted, occasionally short-stoloniferous; sheaths
glabrous below, becoming ciliate at the ligule
with hairs up to 2 mm long; blades up to 20
cm long, ca 5 mm wide, the surfaces and mar-
gins scabrous, occasionally long-pilose at base
of blade; spikes 5 to 17, up to 7 cm long, usu-
ally erect when young, becoming somewhat re-
flexed at maturity; spikelets rather densely im-
bricate, averaging about twelve per cm of the
scabrous rachis; glumes lanceolate, acute, pale,
glabrous with scabrous midnerves; first glume
1.5 to 3.2 mm long, 0.2 to 0.9 mm wide; second
glume 2.5 to 3.4 mm long, 0.3 to 0.7 mm wide;
fertile lemma 2.8 to 3.7 mm long, 0.6 to 1.1 mm
wide, pale, broadly linear to elliptic, callus
bearded, rest of lemma glabrous except for the
appressed-pilosc margin, apex obtuse, unawned
or with a very short mucro; sterile floret one,
1.4 to 2.2 mm long, 0.3 to 0.9 mm wide, trun-
cate, awnlcss or occasionally with a very short
mucro; anthers ca 0.5 to 0.9 mm long; caryopsis
1.7 to 2.3 mm long, 0.5 to 0.6 mm wide and
thick, ellipsoid, trigonous; chromosome number
2n=ca 65, 80.
Chloris suhmutica outwardly resembles
species in Eustachys, primarily because of the
appressed, awnless spikelets. Tlie second glume,
however, lacks the rounded apical lobes sub-
tending the short awn, a characteristic of Eu-
stachys. Also, C. suhmutica lacks the conspicuous
equitant leaf sheaths of Eustachi/s.
Chloris suhmutica Humboldt, Bonpland,
Kunth and C. polystachi/a Lagasca were both
published in 1S16. According to Stafleu (1967),
the particular section of the Nova Genera et
Species Plantarum, containing the original de-
scription of C. suhmutica, was published in
Mav 1816; while C. poh/staclu/a Lagasca, pub-
lished in Elenchus Plantarum, was probably
published in June or Jul\' 1816. In addition to
the slightly earlier date of publication, C. .suh-
mutica is clearly identifiable and has been
widely accepted, far more so than C. poly-
stachya. For these reasons C. suhmutica is
maintained here.
Cliloris suhmutica has been extensively col-
lected in Mexico, occasionally in Columbia,
Guatemala, New Mexico, and Venezuela ( Fig.
43, circles ) . Most specimens are from eleva-
tions of 3,000 to 6,000 feet and on a variety
of soil types.
Representative specimens examined: CO-
LOMBIA: Dep. Cundinamarca: Sabana de
Bogota, cerca de Madrid, Tarragon et al. IS Cu
104 (US). GUATEMALA: Quezeltenango,
Chiquilaga, de Koninck 11 (US). MEXICO:
Aguascalientes : Aguascalientes, Hitchcock 7,453
(US); Chiapas: N of San Cristobal de Las
Casas, Breedlove 6,759 (F); Chihuahua: Ma-
jalca, Pilares, 2,080 m, Harvey 1,483 (US); San
Juanito, Knohloch 5,425 (F); Coahuila: Saltillo,
5,300 ft, Fisher 57 (US), Hitchcock 1,261 (NY,
UC) 5,58.5 (US); Durango: Tepehuanes,. F(.s/ier
44,221 (MO), Palmer ^262, 4-25 June 1906
(MO, NY, UC, US); Otinapa, Pahner 341, 25
June-5 Aug 1906 (MO, NY, UC, US); Dls-
trito Federal: Lomas de Mixcoac, Lyonnet 2,701
Biological Sehies, Vol. 19, No.
Taxonomy of the Genus Chloris (Gramineae)
109
( US ) ; pyramid of Cuicuiico Tlalpan, McDcin-
ieh 47 (F); Guanajuato: 6 km S of San Frlipe,
Sohns 405 (US). Hidalgo: Pachua, 8,000 ft,
Hitchcock 6,748 (US). Jali.sco: Zapotlan,
Hitchcock 7,129 (US); Michoacan: 5 mi N of
Patzcuaro, Webster 2,700 (US); Navarit: Tepic,
Santa Teresa, Rose 2,143 (US); Nuevo Leon:
Galeana, Hacienda Pablillo, Tmjhr 127 (F,
MO); Puebla: vicinit\' of Puebla, Arscne 330
(MO, NY, US); Queretaro: Cd. Queretaro,
Hitchcock 5,818 (US); Sonora: Rio de Bavispe,
El Picaco de Pilar, Santos 2,168 (US); San Luis
Potosi: Cd. San Luis Potosi, Schaffner 130 (NY,
US); Tamaulipas: 6 km W of Miquihuana. 2 940
m, Stanford et al. 737 (MO, US); Tlaxcala: be-
tween San Cristobal and Calpulalpan, Sohns 580
(US); Zaeatecas: 15 km W of Concepcion del
Oro, near Zacatecas-Coahuila border, Stanford et
al. .5.58 (MO, NY, UC, US). UNITED STATES:
New Mexico: Dona Ana Co., State College,
Norris 1,026 (US), Parker 459 (US). VENE-
ZUELA: Estado de Merida: between Mucaruba
and Mueliachies, Rudd 541 (US).
46. CHLORIS SOMALENSIS Rendle, Jour.
Bot. (London) 37:66. 1899. (IIOLO-
TYPE: "Wagga Mountain, Somaliland.
Mrs. E. Lort' Phillips. 1897." BM!) Fig.
74, F-L.
Perennial to 70 cm tall, cespitose, erect;
sheaths glabrous, overlapping; ligule a short-
pilose crown; blades 15 to 30 cm long, to 7
mm wide, glabrous; spikes three or four, ca
8.5 cm long, erect when young, later spreading;
spikelets appressed, dense, ca ten per cm of
the scabrous rachis; glumes unequal, linear-
lanceolate, thin and glabrous to slightly scab-
rous, especially on the midnerse; first glume
1.4 to 3.0 mm long, 0.2 to 0.4 mm wide; second
glume 3.2 to 5.0 mm long, ca 1.0 mm wide;
iFertile lemmas 3.5 to 5.2 mm long, 0.8 to 1.0
mm wide, lanceolate, olive-green, callus beard-
ed, margins glabrous below, becoming spar-
ingly appressed-pilose near the apex, otherwise
glabrous, awn 7 to 12 mm long; sterile floret
one, 0.5 to 2.0 mm long, 0.1 to 0.5 mm wide,
usually greatl\- flattened and reduced, some-
times slighth' inflated, scabrous, awn 1.5 to 3.5
mm long: anthers 1.0 to 1.5 mm long; car\'opsis
2.0 to 2.2 mm long, 0.5 to 0.6 mm wide.' ellip-
soid, trigonous.
In general the olive-colored spikelets and
greatly reduced sterile florets readily separate
Chloris somalensis from related species. Occa-
sional specimens have exceptionally large rudi-
ments and mav eventually prove distinct as
better series of specimens become available.
Included here would be Glover and Gilliland
755 (BM!).
Onl\- six specimens were available in all
herbaria consulted. Tlicse were collected in
Ethiopia and the Somali Republic ( Fig. 73,
circles ) in open, grassy savannahs with Jiini-
periis and Dodonaea. Soils were partly lava with
black clay and partly sandstone with red, sandy
soil in areas with moisture derived largely from
low clouds.
Specimens examined: ETHIOPIA: Hamasen,
Nefarit, Baldrati s.n., 7 Mar 1916 (K); Sidamo,
Mega, 4°5'N, 38°20'E, ca 2,200 m, GiUett 14,296
(K, US), Moonetj 5,565 (K). SOMALI RE-
PUBLIC: Golis Range, Baldostie Codie, Drake-
Brockman 480 (K); Iva Makarai, 9°56'N, 45°E,
Drake-Brockman 481 (K).
47. CHLORIS CLEMENTIS Merrill, Philippine
Jour. Sci. 40:181. 1929. (HOLOTYPE:
"Luzon: Pangasinan Province, Calasiao . . .
Clemens 17,267." UC! ISOTYPES: BR! F!
NY! US!) Fig. 74, A-E.
Perennial 20 to .30 cm tall, with delicate,
erect culms arising from the prominent stolons;
sheaths very short, glabrous below, becoming
somewhat ciliate along upper margins; ligule a
short membranous ring; blades 1.0 to 2.5 cm
long, 2 to 4 mm wide, obtuse, slightly apiculate,
glabrous; spikes two to five, 2 to 3 cm long,
radiate; spikelets appressed, imbricate, ca five
per cm of the scabrous rachis; glumes lanceolate
to ovate, membranous, glabrous except for the
scabrous midnerve; first glume 0.5 to 0.6 mm
long, ca 0.2 mm wide; second glume 0.7 to 0.8
mm long, ca 0.2 mm wide; fertile lemma 1.5 to
1.8 mm long. 0.4 to 0..5 mm wide, lanceolate
to elliptic, callus bearded, margins sparsely
ciliate apically, scabrous below, keel sparsely
scabrous, otherwise glabrous, awn 5.5 to 6.5 mm
long; sterile floret one, ca 0.3 mm long, 0.1
mm wide, much reduced, flattened, often con-
cealed within the lemma, awn ca 1 mm long;
car\opsis ca 1.2 mm long, 0.3 mm wide, nar-
rowly ellipsoid.
Chloris dementis resembles several other
species in gross appearance, but differs mark-
edly in detail. Its strongly stoloniferous habit
is similar to that of C. divaricata as well as
to EustachijS tenera and to various species of
Cijnodon. It differs from Chloris divaricata in
having much smaller sterile florets ( Figs. 73,
A-E; 51, K-R). Sterile florets in Cynodon are
either greatly reduced or absent. Etisfachijs is
characterized b\' having bilobed second glumes.
Other species (such as Chloris ekmanii, C.
pijcnothrix, C. radiata, and C. sagraeana) may
no
Bbigham Young University Science Bulletin
Fig. 74. Chlorii dementis and C. somalensis. (A-E) C. dementis. (A) habit, x 1/3; (B) spilcelet, partly dissected,
X 10; (CD) sterile florets, showing variation, x 30; (E) caryopsis, x 10. (F-L) C. somalensis. (F) habit,
X 1/3; (G) spikelet, partly dissected, x 10; (H-K) sterile florets, showing variation, x 10; (L) caryopsis, x 7.5.
have similar spikelets but are tufted rather than
stoloniferous.
No specimens were seen other than the
holotype and its isotypes.
48. CHLORIS RADIATA (Linnaeus) Swartz,
Prodr. Veg. Indiam Occ. 26. 1788. (Based
on AgTOstis radiata Linnaeus.) Fig. 75, F-J.
Agrostis radiata Linnaeus, Syst. Nat. ed. 10.
2:873. 1759. (See nomenchitural note
below. )
Ct/nosurus scopariiis Lamarck, Encycl.
Meth. 2:188. 1786. (HOLOTYPE: Not
positively identified, but there is a frag-
ment from P, in US!, labeled "Cynosurus
scoparius de St. Dominique Lam.")
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chlohis (Gramineae)
111
Chloris scoparius (Lamarck) Desfontaines,
Tabl. ficole Bot. Mus. Hist. Nat. 14. 1804.
( Based on Ct/nosurus scoparius Lamarck. )
Cliloris i^Iaucescens Steudcl, Ssn. PI. Glum.
1:206. 1854. (HOLOTYPE: "Guadeloupe.
Duchaissing" P! Fragment in US! )
Chloris tacncnsis Steudel in Lecliler Ber-
berides Ainer. Australis 56. 1857. Nomen
niulein, but a specimen "St. 1.569" is cited.
This specimen, in P!, labeled C. tacnensis
in Steudel's script is C. radiata (Linnaeus)
Swartz.
Chloris biflora Kegel, Annotationes Bot.
Index Seminum 28. 1863. Pro syn.. C.
radiata.
Chloris scoparius Kegel, Annotationes Bot.
Index Seminum 28. 1863. Pro syn.. C.
radiata.
Chloris pallescens Kegel, Annotationes Bot.
Index Seminum 28. 1863. Pro syn., C.
radiata.
Chloris crinita Salzman ex Doell in Martins
Flora Brasiliensis 23; 64. 1878. Pro .syn.,
C. radiata. Non Lagasca, 1805.
Gymnopogon radiata (Linnaeus) Parodi,
Ph\sis 4:180. 1918. (Based on Agrostis
radiata Linnaeus. )
Annual usually with dense fibrous roots, oc-
casionally rooting at lower nodes, 30 to 60 cm
tall; sheaths glabrous, occasionally pilose; ligule
a short-ciliate crown, sometimes with a few
longer hairs intennixed; blades 10 to 30 cm
long, up to 10 mm wide, usually pilose, occa-
sionally glabrous or scabrous, apex usualh'
acute; spikes five to fifteen, 5 to 8 cm long in
one, two, or sometimes three ascending verti-
cils; spikelets densely imbricate, ca 11 to 15
per cm of the scabrous rachis; glumes \ery
narrowly lanceolate, glabrous except for the
scabrous midner\'e; first glume 0.7 to 1.6 mm
long, 0.1 to 0.2 mm wide; second glume 2.0
to 2.7 mm long. 0.1 to 0.2 mm wide; fertile
lemma 2.8 to 3.3 mm long, 0.4 to 0.6 mm wide,
lanceolate to elliptic, glabrous, except upper
margins short-ciliate, callus bearded, awn 6 to
13 mm long; sterile floret one, 0.4 to 0.7 mm
long, ca 0.1 mm wide, greatK' reduced, borne on
a long rachilla joint, awn 3 to 5 mm long; car\-
opsis 1.4 to 1.5 mm long, 0.3 to 0.4 mm wide,
trigonous; chromosome number 2n^40.
The correct name of this plant cannot be
established conclusively. Linnaeus' original de-
scription refers to a plate and description in
Sloane ("Sloane jam. t. 68. f. 3"). Neither the
text nor the plate are clear enough for posi-
tive identification; both could fit the species
described above or possibly another Caribbean
species, for example, C sagraeana. In the Lin-
naean herbarium there is a specimen of Pat-
rick Browne's which Linnaeus might have seen;
it is fragmented but seems to be more similar
to C. sagraeana. No other information can be
obtained either to substantiate or refute the
name. While most of the evidence in this prob-
lem is either nonsubstantive, or even somewhat
negative, I do not wish to reject circumstantially
a name that has been used commonly for this
species for nearly two centuries. Until additional
information can be found, I propose continued
acceptance of Chloris radiata (Linnaeus)
Swartz for the plant described above.
Chloris radiata most closely resembles C.
pycnothrix. Differences have been discussed
under the latter species. Other Caribbean spe-
cies are similar in spikelet and inflorescence
features but have larger sterile florets (see
Table 13 for a further comparison. )
Chloris radiata is widely distributed through-
out the eastern Caribbean, northern South
Africa, and Central America, where it is a
common weed in waste areas, along paths,
and roadways, etc. (Fig. 76).
Kepresentative specimens examined: BA-
HAMA ISLANDS: Nassau, Hitchcock s.n., Nov
1890 (F, MO). BOLIVIA: Coripati, Yungas,
Bang 2.173 ( F, MO, NY, US). CAKOLINE
ISLANDS: Palau group, Koror Island, Fosberg
32,327 (NY, US). COLOMBIA: Div. Antio-
quia: Fradonia, Archer 503 (US); Div. Cauca:
El Valle, Triana 345 (US); Div. Nariiio, be-
tween Dos Kios and La Union, Archer 3,394
(US); Div. Santander: Boca de Rosario, Kio
Magdalena, Pennell 3,904 (MO, NY, US).
COSTA KICA: Port: Limon, Hitchcock 8,417,
8,42S (US); Prov. San Jose: between San
Pedros Montes de Oca and Curridabat, Stand-
ley 41.283 (US). CUBA: Prov. Pinar del Rio:
Mariel, Ekman 12.939 (F, NY, US); Prov.
Habana: Guatao, Leon s.n., Oct. 1923 (US).
DOMINICAN REPUBLIC: Prov. Pacificador:
Pimentel, Abbott 691 (US); Prov. Seibo: vi-
cinit\' of Hato Mayor, Allard 13,643 (US); Prov.
Santiago: vicinit)' of Santiago, Allard 14,553
( US );'^ Prov. Duarte: Valle de Cibao, Ekman
13,259 (US); Prov. Puerto Plata: Puerto Plata,
Ekman 14.448 (US). ECUADOR: Prov. Chim-
borazo: Alausi, Hitchcock 20,695 (US); Prov.
Esmeraldas: Rio Santiago, Acosta-Solis 19,710
(US); Prov. Manabi: Bahia, Acosta-Solis 12,002
(F); Prov. Pichincha: between Santo Domingo
and Quininde, Acosta-Solis 13,289 (US).
FRENCH GUIANA: vicinity of Cayenne,
Broadway 44 (NY, US). GALAPAGOS IS-
112
Bbigham Young University Science Bulletin
Fig
75. ChloTis pycnothrix and C. radiata. (A-E) C. pycnothrix. (A) habit, x 1/5; (B) spikelet partly dissected,
X 7.5- (C) group of fertile florets, showing variation, x 10; (D) group of sterile florets, showing vanation
X 10- (E) caryopsis, x 10. ( F-J ) C. rajiata. (F) habit, x 1/5; (G) spikelet. partly dissected, x 7.5; (H
group of fertile spikelets, showing variation, x 7.5; (I) group of sterile florets, showing variation, x 10; (J)
caryopsis, x 10.
Biological Seiues, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
113
-^^"^^^^
.K^
«*> I *v
/ c
D
X
E
^h>-
d?
â– â– J
» A
^«
<*
•
.^
C\
Fig. 76. Distribution of Chlofia radiata. In.sets: (A) Galapagos Islands. (13) Hawaiian Islands. (C) Marianas
Islands. (D) Okinawa and adjacent islets. (E) Palau Islands.
LANDS: Albemarle Lsland, Villaiiiil, IIowcIl
8,954 (US). GUATEMALA: Dep. Izabel: vi-
cinity of Puerto Barrios, Standley 24,761 (US);
Dep. Jutiapa; between Jutiapa and La Cclera,
Standley 76.021 (US). GUYANA: New Am-
sterdam, Hitchcock 16,833 (F, MO, NY, UC,
US). HAITL Tortue Island, La Vallee, Leon-
ard and Leonard 11,585 (NY, UC, US); vi-
cinitN' of Jean Rabel, Leonard and Leonard
13,752 (US). HONDURAS: Dep. Morazan:
vicinity of El Zamorano, Standley 24,665 (F).
JAMAICA: Cornwall Co., Ipswich, Hitchcock
9,600 (US); Middlesex Co., Ewarton to Lin-
stead, Hitchcock 9,437 (US); Surrey Co., Blue
Mtns., Robertsfield, Harris 11,942 ( F, MO, NY,
US). LEEWARD ISLANDS: Antigua: Ben-
dais Road, Box 4 (US); Guadeloupe: Point a
Pitre, F. W. Johnson 1,093 (NY); St. Kitts.
Hitchcock 16,538 (US). MARIANAS IS-
LANDS: Rota Island, Fosherg 3,027 (US);
Saipan, Mt. Tapotchau, Fosherg 25,181 (NY,
US). MEXICO: Oaxaca: vicinit)' of Cafetal
Concordia, Morton and Makrinus 2',598 (F, US).
NICARAGUA: San Juan del Sur, Hitchcock
8,592 (US); vicinity of Jinotepe, Standley 8,535
(F). PANAMA: Bocas de Toro, Laguna de
Chiriqui, Hart 66 (US); Taboga Island, Hitch-
cock 8,073 (US); Barro Colorado Island,
Standley 41,125 (US). PERU: Dep. Caja-
marca: Chilete, Anderson 744 (US); Dep. Lima:
La Molina, Ferreyra 11,118 (US); Dep. Lor-
eto: Aquaitia, Rio Yurac Yacu, Seihert 2,076
(MO, US). PUERTO RICO: Monte Mesa,
vicinity of Mayaguez, A. Chase 6,179 (US);
vicinity of San Juan, SW of Bayamon, A. Chase
6,388 ( US ) ; Viequis Island, vicinity of Isabel
Segunda, Shafer 2,474 (NY, US). RYUKYU
ISLANDS: Daitoshima group, Rasa Island,
Oct 1919 (US); Okinawa, Boro-
Island, Amano 5,756 (US). EL SALVA-
DOR: Dep. Sonsonate: Acajutla, Standley
21,908 (US); Dep. de la Libertad: vicinity of
Ateos, Standley 23,343 (MO, US). SURINAM:
Lower Saramacca River, Lanjouw 263 (US).
TOBAGO: Orange Hill, Broadway 4,649 (US).
TRINIDAD: Santa Cruz, Cangrehal, Broadway
2,279 (MO, US). UNITED STATES: Hawaiian
Islands, Oahu, Haunaman Bay, A. Chase 12,681
Kawagoe s.n
114 Brigham Young University Science Bulletin
Table 13. Comparison of Chloris arenaria, C. cuben^is, C. ekmanii, C. halophila, C. pycnothrix, C. radiata, and
Culms
Mature inflorescence
shape
Leaf Blade
Species
Width
(in mm)
Pubescence
C. arenaria
stout; GO-
TO cm tall
spikes horizontally
spreading or
somewhat deflexed
1.5-3.0
scabrous to densely
pilose above;
densely pilose below
C. cubensis
stout; 50-
60 cm tall
spikes erect, diverg-
ing no more than
45° from vertical axis
about
1.5 mm
glabrous, occa-
sionally short
pilose
C. ekmanii
slender,
wiry; 30-
45 cm tall
spikes horizontally
spreading or
deflexed
less
than 1.0
mm, often
less than 0.5
mm
sparsely to densely
pilose, occasionally
glabrous
C. halophila
stout; 15-
50 cm tall
spikes in several
verticils, hori-
zontally spreading
2-3
glabrous or scabrous,
sparsely pilose
above the base
C. pijcnothrix
stout
spikes ascending
when young, diverg-
ing at maturity
3-5
glabrous to
scabrous
C. radiata
stout
spikes ascending
5-10
usually pilose,
occasionally gla-
brous or scabrous
C. sagraeana
stout
spikes horizontally
spreading to
somewhat deflexed
1.3-3.0
glabrous
(US); Oregon: Linnton, on ballast, /. C. Nel-
son 487, 9 Oct 1915 (US). VENEZUELA:
Periveca, Capacho, Tachira, Tamaijo 2,297
(UC, US). VIRGIN ISLANDS: St. Croix,
Britton and Cowell 50 (NY); Tortola, Britton
and Shafer 755 (NY, US). WINDWARD IS-
LANDS: Barbadoes, Maxwell coast, Allan 317
(K); Dominica, Hitchcock 16,440 (US); Gre-
nada, Richmond Hill, Broadway 978 (US); St.
George's, Broaduay s.n., 12 Nov 1905 (US);
Martinique, Halm 696 (NY, US); St. Lucia,
Hitchcock 16,485 (US); St. Vincent, Smith and
Smith 581 (K).
49. CHLORIS PYCNOTHRIX Trinius, Gram.
Unifloris 234. 1824. (HOLOTYPE: "Chloris
pycnothrix . . . Rhadd pycnothrix mi Sta.
Cathar am Chamisso." Fragment, in US!,
from Trinius Herbarium.) Fig. 74, A-E.
Chloris beyrichiana Kunth, Rev. Gram. 1:89.
1829; 2:289. pi. 56. 18.30. (TYPE: ". . .
Brasiliae graminosis, prope Rio-Janeiro . . ."
Not seen, but description and plate clearly
refer to this species. )
Chloris humilis Kunth, Rev. Gram. 1:89.
1829; 2:531, pi. 180. 1832. (TYPE: "prope
Rio-Janeiro." Not seen, but description and
plate are definitive. )
Chloris ohtnsifolia Desvaux, Opuscules 73.
1831. (Type not seen, but description is
clear.) Non Trinius, 1824.
Chloris intermedia Achille Richard, Tent.
Florae Abyssinicae 2:407. 1850. (HOLO-
TYPE: "Abyssinia, Chire, Leg. Quartin Dil-
lon and Petit" P! )
Chloris leptostachya Hochstetter ex AchiUe
Richard, Tent. Florae Abyssinicae 2:407.
1850. (ISOTYPES: "Abyssinica, ad agrorum
margines et in incultis prope Adoam.
Schimper, Sectio secunda, 951." BM! G! K!
P!)
Chloris salzmanii Steudel, Syn. Pi. Glum.
1:206. 1854. (HOLOTYPE: "Brazil, Bahia,
in argillosis aridis . . . Salzmann 664." G! )
Chloris leptostachya var. intermedia
(Achille Richard) Durand and Schinz, Con-
spectus Florae Africae 5:861. 189.5. (Based
on Chloris intermedia Richard. )
Chloris radiata var. beyrichiana (Kunth)
Hackel in Stuckert, Anales Mus. Nac. Hist.
Nat. Buenos Aires 6:489. 1906. (Based on
Chloris beyrichiana Kunth; see also Chloris
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae) 115
C. sagraeana.
Fertile lemma
Awn
length
Sterile floret
Spikelets per Length Width length Length Width
cm of rachis (in mm) (in mm) (in mm) Margin pubescence (in mm) (in mm)
5-7
3.5-3.7
ca 0.5
20-24
appressed-ciliate; 1.6-1.9
hairs short
0.2r0.3
5-9
6-10
3.1-3.6
2.0-3.0
0.6-0.7
0.4-0.5
9-12
5.5-10
densely ciliate, at
least some hairs
more than 1 mm
long
1.4-1.9
0.4-0.6
sparsely to densely
appressed-pilose on
upper margins;
hairs short
0.4-1.1
less than
0.2 mm
4-7
3.8-5.5
0.6-0.8
9.0-26
appressed-pilose 1.5-2.5
0.2-0.4
12-17
11-15
4-9
2.5-3.0 0.4-0.5
2.8-3.3 0.4-0.6
2.6-3.5 05-0.7
10^5 glabrous, or u ith 0.3-1.6
few scattered hairs
near apex
6-13 upper margins short- 0.4-0.7
pilose with ap-
pressed hairs
7-13 densely appressed- 0.7-1.6
pilose wim hairs
less than 0.5 mm long
0.2 mm
or less
ca 0.1
0.2-0.5
radiata var. beyrichiana Hackel in Kurtz
under "Unidentified Names.")
Gijmnopof^on radiatus var. beyrichiana
(Kunth) Parodi, Phvsis 4:173. 1918.
( Based on Clitoris beyrichiana Kunth. )
Gymnopogon haumani Parodi, Physis 4:1S3.
1918. (Type not seen, but description and
illustration are conclusive.)
Gymnopogon beyrichianus (Kunth) Parodi,
Revista Centr. Estud. Agron. Vet. Univ.
Buenos Aires 18:148. Fig. .5.5. 1925. (Based
on Chloris beyrichiana Kunth, possibly
misapplied. )
Annuals with erect culms arising from short
stolons, up to 4.5 cm tall; sheaths glabrous,
occasionally becoming long-pilose near the
collar; ligule ciliate, occasional!)' long-pilose;
leaves 4 to 9 cm long, 3 to 5 mm wide, glabrous
or scabrous, apex usually obtuse; spikes four to
eleven (often three to nine), 3 to 8 cm long,
usually borne in a single verticil, though oc-
casionalh' with two; spikes ascending when
young, becoming divergent at maturit)'; spike-
lets densely imbricate and tightly appressed to
the inflorescence axis, averaging 12 to 17 per
cm of the scabrous rachis; glumes thin, nar-
rowly lanceolate, glabrous except for the scab-
rous-hispid midnerve and the somewhat scab-
rous apex, rarely pilose; first glume 1.3 to 1.7
mm long, 0.1 to 0.3 mm wide; second glume
2.5 to 3.5 mm long, 0.2 to 0.3 mm wide; fertile
lemma narrowly elliptic, 2.5 to 3.0 mm long,
0.4 to 0.5 mm wide, glabrous except occasion-
ally a few scattered hairs near the apex, callus
bearded, awn 10 to 45 mm long, usually 10 to
25; sterile floret greatly reduced, usually less
than 1 mm long, occasionally up to 1.6 mm
long, 0.2 mm or less wide, awn 4.5 to 11.0 mm
long, caryopsis 1.2 to 1.4 mm long, 0.3 to 0.4
mm wide, narrowly ellipsoid; anthers ca 0.5
mm long; chromosome numbers 2n=30, .36, 40.
Chloris pycnothrix most closely resembles
C. radiata and, somewhat less so, C arenaria,
C. cubensis, C. ekmanii, and C. sagraeana.
Table 13 presents the more important character-
istics differentiating these species.
Plants combining various characteristics of
Chloris pycnothrix and C. radiata are occasion-
ally found in areas that are unquestionably
within the range of C. pycnothrix. For exam-
ple, collections from the Galapagos Islands
have long awns, a characteristic of C pycno-
thrix, but otherwise fit C. radiata. Specimens
116
Brigham Young University Science Bulletin
with the short-ciliate lemma margins of C.
radiata, from parts of Venezuela and North
Africa, are referred to C. pycnothrix only be-
cause the preponderance of their characters
belong to that species. Individual specimens
from these areas may be difficult, if not im-
possible, to refer to one species or another on
the basis of only a few key characters. These
aberrant specimens must represent casual, ran-
dom variation in spikelet features, for these
plants otherwise maintain the distinctive epi-
dermal features of the species (see Table 13;
Fig. 1).
Chloris pycnothrix is widely distributed
through Africa and northern South America
( Fig. 77 ) . Collection data indicate that these
plants are frequently found as weeds in dis-
turbed areas. While usually described as an
annual, greenhouse specimens have persisted
for over two years before dying, suggesting
that this species might behave as a weak pe-
rennial under favorable conditions.
Representative specimens examined: AN-
GOLA: Huila Dtr.: Posta Zootechnica, Teixeira
Fig. 77. Distribution of Chloris pycnothrix. Insets:
(A) Galapagos Islands. (B) Reunion and Mauritius.
637 (K); Golungo Alto, Welwitsch 2,985b (K).
ARGENTINA: Prov. Corrientes: Empedrado
Dap., Estancia "Las Tres Marias" Pedersen
3,053 (US); Prov. Misioncs: Posades, Parodi
4,017 (US); Prov. Formosa: without precise
location, Jorgensen 43,053 (US). BRAZIL:
Prov. Bahia: 39 km NE of Bahia, A. Chase
8,016 (F, MO, NY, US); Prov. Maranhao: Caro-
lina to San Antonio dc Balsas, Stcallen 3,951
(US); Prov. Matto Grosso: Tres Lagoas, A.
Chase 10,759 (US); Prov. Parana: Guaratuba,
Morro do Brejatuba, Hatschhach 4,469 (US);
Prov. Rio Grande do Sul: vicinity of Guaiba,
Beetle 1,498 (UC); Prov. Catarina; Campo Ere,
Fazenda Sao Vicente, Castellanos 24,742 (K).
BURUNDI: environs de Karuzi, Vander Ben
2,075 (K). CAMEROONS: Bertoua, Bahlwin
13,954 (US); Wakwa, 10 km S of N'Gaoundere,
Breteler 488 (K); Nkambe Div.: Bamcnda
Highlands, 6°35'N, 10''40'E, Hepper and Char-
ter 1,938 (K). CENTRAL AFRICAN RE-
PUBLIC: 35 mi from Bambari, Kt/rle FAIO
(K). ETHIOPIA: Bagemedcr Prov.: Lake
Lana, Bahudur Gorge, Grahham 116/21 (K);
Harar Prov.: NE of Lake Alemaya, 15 km NW
of Harar, Burger 1,079 (K). GALAPAGOS
ISLANDS: Santa Cruz Island, Academy Bay,
Schimpff 39 (NY, US). GHANA: Venchi,
7°49'N, 2°0TW, Rose-hmes 30,519 (K); Nzere-
kore, Baldwin 9,709 (US), .9,725 (K). GUINEA:
Mamou, Chevalier 34.887 (K). IVORY
COAST: Pakobo Nord, Adjoushoun 384A (K).
KENYA: S mi N of Tsavo Falls, 6,300 ft, Bar-
ney 1,123 (K); Nairobi, Bogdan B44 (UC);
Amboseli National Park, 50 mi E of Namanga,
Bogdan AB 5,500 (K); S Lake Elmenteita,
6,000 ft, Clayton DC 27 (K); Mt. Suswa,
Glover 3,785 (K). MALAGASY REPUBLIC:
Mt. Belamboany, Perrier de la Bathie 14 (K).
MALAWI: Shiri Highlands, Buchanan 55 (K);
Mlanje, Burtt-Davy ^1962/29 (K); near Zomba,
Cormack 165 (K). MARTINIQUE: roadside,
Velez 3,447 (US). MAURITIUS: Curepipe.
Vacoa, Vaughan A-29 (K). MOQAMBIQUE:
Mocuba Dtr.: Namagoa, Faulkner 18 (K), 145
(K, US); 26 km W of Lourenyo Marcjues,
Gomes e Sousa 3,247 (K). NIGERIA: Samam,
Zaria, Clayton DC 1,332 (K); Bauchi Plateau,
Lely P769 (MO); Toro, 20 mi E of Jos, Scmple
173 (US). PARAGUAY: Dep. Guaira: Villarica,
Jorgensen 3,532, 3,533 (US); Dep. Misiones:
Florida, Campal s.n., 15 Apr 1938 (US); Dep.
San Pedro: Primavera, Woolston 6.40 .(K).
PERU: Dep. Apurimac: Abancay, Stordt, with-
out collection number and date (K). REPUB-
LIC OF THE CONGO: Marungu, Dubois 1..363
(K); Eala, Lebrun 454 (MO, US); Yangambi,
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
117
Louis 16,007 (K); Katanga, 25 km N of Luhum-
bashi (Elizabethville), Schmitz 7,384 (K). RE-
UNION: St. Denis, Rivels, without collection
number and date (K). SOMALI REPUBLIC:
Hargeisa, 9°33'N, 44°E, GiUett 3.917 (K);
Medishe, Glover and GiUihnd 941 (K, US).
SOUTHERN RHODESIA: Sali.sbuiy, Brain 440,
1,541 (MO); Stapleford Forest Reserve, Gilli-
land 237 (K); Kariba Gorge, Goldsmith 73/60
(K); Urungvve National Reserve, Zwipani area,
Goodier 513 (K); Victoria Falls, wet spray
areas, Parker 4,202 (K); near Sinoia, Rodin
4,376 (F, MO, UC, US). SUDAN: Dai fur
Prov.: Gur Lambong, Jebel Marra, Wickens
2,593 (K); Equatoria" Prov.: Yd, Harrison 488
(K); Khartoum Prov.: Khartoum, Jackson 4,186
(K); Northern Prov.: Wadi Haifa, Pettet 90
(K). TANZANIA: Kondoa Dtr.: near Sam-
bala, Ritrtt 2,603 (K); Ngorongoro Crater,
Burtt 4,338 (K); Musoma, Emson 241 (K);
Bukoba Dtr.: Ngono Bridge, Gillman 67 (K);
Kilimanjaro, Lyamungo, Greenwaij 3,133 (K);
Seronera, Serengeti, Greenwaij 9,874 ( K ) ; Zan-
zibar, Hitchcock 24,445 (US). UGANDA:
Nabilatuk, near Emonagabem, Dtjson-Hudson
55 (K); Entebbe, Hitchcock 24,874 (US); 7 mi
W of Moroto Karamoja, Lan<^dak'-Broum 1,572
(K); Busoga, Nawendwa, Maitland 1,406 (K);
Mbale, near Mt. Elgon, Snowden 1,090 A (K).
UNION OF SOUTH AFRICA: Cape Provinces,
East London, McCaUum-Wehster N94 (K);
Natal, SW of Mapumulo, Umtavi Valley, Moll
1,843 (K); Transvaal, Kruger National Park,
de Winter and Codd 600 (US). VENEZUELA:
E of La Guiana, Ctirran and Hainan 902
(NY). ZAMBIA: Chilonga, Mt. Makula Re-
search Station, Hinds 6 (K); Mpika Dtr.:
Luangwa Game Reserve, Mfuwe, Mitchell 2,945
(K); Kasama Dtr.: Mungwi, Robinson 4,080
(K); Mazabuka Dtr.: Mochipapa to Mabwin-
gombe Hills, White 7,523 (K).
50. CHLORIS HALOPHILA Parodi, Rev. Ar-
gentina Agron. 12:45. Fig. 1. 1945. (ISO-
TYPE: "Argentina, Buenos Aires, Perga-
mino . . . Parodi 2769" US! ) Fig. 78.
Perennial 15 to 50 cm tall, sometimes up
to 80 cm, weakly stoloniferous; sheaths glabrous,
sometimes sparsely pilose above; ligule ciliate
or a short, membranous crown; blades glabrous
or scabrous, sometimes becoming sparsely pilose
on upper surface near the base; spikes four to
thirteen, 3.5 to 12.0 cm long (usually 7 to 8
cm), arranged in several verticils; spikelets ap-
pressed, somewhat distant, ca four to seven
per cm of the scabrous rachis; glumes narrowly
lanceolate, glabrous except for the scabrous mid-
nerves; first glume 1.5 to 3.0 mm long, 0.2 to
0.3 mm wide; second glume 3.7 to 5.3 mm long,
0.3 to 0.4 mm wide; fertile lemmas 3.8 to 5.5
mm long, 0.6 to 0.8 mm wide, narrowly lance-
olate or elliptic, apex acute to acuminate,
callus bearded, margins appressed-pilose, awn 9
to 26 mm long; sterile floret one, 1.5 to 2.5 mm
long, 0.2 to 0.4 mm wide, lanceolate to elliptic,
apex acute to acuminate, glabrous to somewhat
scabrous, awn 7 to 13 mm long; anthers ca 0.7
mm long; caryopsis 1.9 to 2.4 mm long, 0.4 to
0.6 mm wide, ellipsoid, trigonous; chromosome
number 2n = 80.
Chloris halophila is similar to C radiata and
C pijcnothrix in outward appearance but has a
coarser inflorescence and larger spikelets ( com-
pare Fig. 75 and 78). Additional differences
are given in Table 13.
For many years the names Chloris beyrich-
iana, C. radiata var. beijrichiana. and Gtjmno-
pogon beijrichianaus were misapplied to this
taxon, especially by Parodi. Parodi later (1945)
recognized these names as being properly ap-
plied to C. radiata and provided a new name,
C. halophila. for the plant described above.
Chloris halophila has been collected through
a great altitudinal range from ca 2,000 feet
elevation in Tucuman Province, Argentina, to
over 11.000 feet in Bolivia (Fig. 71, circles).
It occurs in grasslands, especially in saline
conditions.
Representative specimens examined: AR-
GENTINA: Prov. Catamarca: Dep. Ambato,
Camino al Rodeo, Cristobal 406 (UC); Prov.
Cordoba: S of de Mar Chiquita, Burkart 20,457
(UC); Dep. Punilla La Cocha, Stuckert 20.191
(G); Prov. Tucuman: valle de Tafi, Bruch s.n.,
1908 (NY); Dep. Burruyacu: Sal Morado, Dias
799 (NY); Rio de Nio, Monetti 43,048 (US);
Tapia, Rodriguez 182 (US); Leales, Chamar
Pojo, Venturi 711 (K, US); Prov. Salta: Pasaje
del Rio Juaramendo, Lorentz 344 (F); Rosario
de Lerma, Meyer 3,464 (NY); Dep. Agua Cal-
iente: Cavelaria, Venturi 3,773 (MO, US).
BOLIVIA: Dep. Cochabamba: Cochabamba,
Cutter 7,381 (US); Mizque, Eyerdam 24,987
(F, G, K, MO, UC); Rio Tapacari. Kuntz s.n.,
1892 (NY); Dep. La Paz: vicinity of La Paz,
ca 10,000 ft. Bang 82 (BM, G, MO, NY, US).
ECUADOR: Prov. Carchi: Pistu a la PalesHna,
Acosta-Solis 16.557 (US); Prov. Loja: between
Las Lomas and Loja, Hitchcock 21,409 (US);
Prov. Pichincha: descenso del Quincha a Guayl-
labamba, Acosta-Solis 10,329 (F, US); Caja-
marca Prov.: Cajamarca, 2,600 m, Anderson 630
(UC, US). PERU: Dep. Cuzco: Cuzco, Hitch-
cock 22,492 (US); 5 km S of Ollantaytambe,
118
Brigham Young University Science Bulletin
3H
Fig. 78. Chloris hahphila. (A) habit, x 1/5; (B) spikelet, partly dissected, x 7.5; (C-^H) fertile florets, show-
ing variation, x 5; (I-N) sterile florets, showing variation, x 7.5; (O) caryopsis, x 7.5.
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Cramineae)
119
Hitchcock 22,552 (K, NY, US); Dep. Hauncave-
lica, Izcuchaca, entre Cucnca y La Mejorada,
3,100-3,150 m, Tovar 2,448 (US); Prov. Huan-
cavo: Mantero, N of Huancavo, 3,300 m, An-
derson 683 (UC, US).
51. CHLORIS EKMANII Hitchcock, U.S. Dept.
Agrie. Misc. Publ. 243:129. 19.36. (HOLO-
TYPE: "Bal)iijal, Jagiicy Grande, prov. Ma-
tanzas, Cuba . . . Ekman . . . 16594." US! )
Fig. 79, A-F.
Perennial 30 to 45 cm tall, occa.sionally up to
60 cm tall, tufted to short-stoloniferous; culm.s
slender, wir\', somewhat geniculate at the base;
sheaths compressed-keeled; blades up to 10 cm
long, usually less than 1 mm wide, sparsely to
densely pilose on both sides, occasionally gla-
brous; spikes three to six, 1.5 to 5.0 cm long, slen-
der, flexuous, erect to appressed when young, at
maturity spreading horizontally or becoming
somewhat reflexed; spikelets imbricate, ap-
pressed when young, somewhat divergent at
maturits', ca 6 to 10 per cm of the scabrous
rachis; glumes lanceolate, glabrous except for
the scabrous midnerve; first glume 1.1 to 2.0 mm
long, 0.1 to 0.3 mm wide; second glume 2.3 to
3.6 mm long, 0.1 to 0.3 mm wide; fertile lemma
2 to 3 mm long, 0.4 to 0.5 mm wide, narrowh
lanceolate to elhptic, callus bearded, sparsely to
densely appresscd-ciliate on upper margins,
otherwise glabrous, awn 5.5 to 10.0 mm long;
sterile floret one, 0.4 to 1.1 mm long, less than
0.2 mm wide, usually included in the fertile
lemma, awn 3.3 to 6.0 mm long; carj'opsis ca
1.4 mm long, ca 0.3 mm wide, ellipsoid, trigo-
nous.
Chloris arenaria, C. cul)ensis, C. ekmanii, C.
racliata, and C. sagraeana form a complex of
species in the Caribbean region, the area to
which all except C. racliata are restricted. While
most specimens can be named with reasonable
confidence, some either have intermediate fea-
tures or else recombine characteristics of two
species. These are often found in, but are not
restricted to, areas of sympatr)'. Field studies
are needed, especially in Cuba; and precise def-
initions of the taxa invoh'cd and evaluation of
their biological significance must await further
infomiation. The treatment presented here will
enable one to name most specimens. A further
discussion of variation patterns is presented un-
der the species most closely concerned. Table 13
presents a comparison of the characteristics of
these species.
Chloris ekmanii is most similar to C. sagrae-
arm but is a more delicate plant with wiry stems
and, usually, very narrow leaves. The first
glume, less than 1.5 mm long, and the sterile
floret are both smaller than any other species in
this complex. Occasional specimens have gla-
brous rather than hain' leaves, but this character
does not correlate with either geographic origin
of the specimen or with other features.
This species is apparently widely distributed
in Cuba, Jamaica, and Hispaniola ( Fig. 80, cir-
cles) in a variety of ecological sites, such as
along railways, roadways, and rivers, shrubby
savannas, pastures, and pinclands. It is often
collected in bright red clay soils, though other
collections have been made on rocky or even
serpentine soils.
Representative specimens examined: CUBA:
Prov. Camagiiey: Camagiiey to Santayana, Brit-
ton 2,428 (NY); San Serapio, N of Camaguey
City, Leon 16,927 (US); Prov. Havana: near
Almandares River, Leon 287 (US); San Miguel,
Eknuin 10,912 (US); La Pitanill, San Miguel de
Casanova, Leon 12,454, 19,517 (US); Guanaba-
coa, Ekman 13,645 (F, NY, US), Hitchcock
23,299 (K, US), 23,240 (US); Jata Hills, Ekman
856 (US); Isla de Pifios, Nueva Gerona, along
Rio Los Casas, Ekman 11,682 (NY); Prov. Las
Villas ( Santa Clara ) : Trinidad, road to San Juan
de Letran, Ekman 13,975 ( US ) ; Gara, Leon
1,480 (NY); vicinity of Sancti Spiritus, Serguis
2,569 (US); Pitajones to Ciegos de Ponciano,
Shafer 12,225 (NY, US); Prov. Matanzas: Ja-
giiey Grande, Ekman 16,954 (F); Prov. Oriente:
Guamo, Hitchcock 23,412, 23,427 (US); Hol-
guin, Eknmn 3,229 (F); Prov. Pinar del Rio: El
Sabalo, Finca Sabanalamar, Killip 32,294 (US);
Loma Pelada de Buenovista, Cavajabos, Leon
13,226 (US); Bahia Honda, Wilson 9,421 (NY).
DOMINICAN REPUBLIC: Prov. Montecristi:
Guayabin, Abbot 916 (US); Santiago Prov.:
Licev al Medo, near Peiia, Jiminez 3,910 (US).
JAMAICA: Cornwall Co., Montego Bay, Hitch-
cock 9,695 (US); Middlesex Co.: Lititz, Man-
chester, Harris 12,434 (F, K, NY, US), Harris
12,697 (K, NY, US); Inverness, Lower Clar-
endon, Harris 12,727 (K, NY, US); between
Ewarton and Linstead, Hitchcock 652 ( F, K,
MO, NY, UC, US); New Forest, Southern Man-
chester, Hitchcock 9,823. 9,847 (US).
52. CHLORIS ARENARIA Hitchcock & Ek-
man ex Hitchcock, U.S. Dept. Agric. Misc.
Publ. 243:131. 1936. (HOLOTYPE: "Cuba,
La Grifa, prov. Pinar del Rio. Ekman
11253." US! ISOTYPES: F! K!) Fig. 79,
G-K.
Chloris eleusinoides Grisebach var. vestita
Greenman ex Combs, Trans. Acad. Sci. St.
Louis 7:477, pi. 39. 1897. (Fragment of
120
Bricham Young University Science Bulletin
, , C. ckmatiii. (A) habit, x 1/3;
sterile florets, showinj
(B) ligiilc region, x 5; (C,D)
Fig. 79. Chloris ekmanii and C. arenaria. (A-F) _ _ ,.-, - , . , „ „
spikelets, partly dissected, x 7.5; (E) sterile florets, showing variation, x 10; (F) earyopsis, x 10. (G-L)
C. arenarUi. (G) habit, x 1/4; (H,I) ligule region, x 5; (J) spikelet, x 7.5; (K, L) sterile florets, show-
ing variation, x 15.
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
121
Fig. 80. Distribution of Chloris cubensis (squares),
C. chmanii (circles), and C. sagraeana (triangles).
Inset A: Puerto Rico and the Lesser Antilles.
type: "Rio Damuji, Rodas Cuba . . . Combs
631" US! Origin of fragment unknown. )
Perennial 60 to 70 cm tall; culms cespitose,
erect to somewhat decumbent at base; sheaths
compressed-keeled, puberulent. becoming dense-
ly so near the apex; ligule a dense tuft of
hairs; lower leaves 5 to 10 cm long, 1.5 to 3.0
mm wide, becoming shorter upward on the
culm; blades scabrous to densely pilose above,
denseh' pilose below; spikes five to six, 4 to 6
cm long, digitate, ascending when young, be-
coming horizontal or somewhat reflexed at ma-
turit)'; spikelets rather denselv imbricate, ap-
pressed, ca five to seven per cm of the scabrous
rachis; glumes narrowly lanceolate, acuminate,
glabrous except for the scabrous midnerve; first
glume 2.2 to 2.5 mm long, 0.2 to 0.3 mm wide;
second glume 3.3 to 3.5 mm long, 0.2 to 0.4 mm
wide; fertile lemma 3.5 to 3.7 mm long, ca 0.5
mm wide, lanceolate to elliptic, callus bearded,
otherwise glabrous except for the short, ap-
pressed-ciliate margins, awn 20 to 24 mm long;
sterile floret one, 1.6 to 1.9 mm long, 0.2 to 0.3
mm wide, narrowly c\lindrical, glabrous, awn
15 to 20 mm long; caryopsis not seen.
Chloris arenaria mav' be separated from other
species in this complex by its densely villous
leaf blades and sheaths and bv its long awns on
the lemmas ( 20 to 24 mm ) .
Only one of the two specimens studied had
collection infonnation. This plant ( Eknuin
11,253) was collected in pineland on white sand.
Specimens examined: CUBA: without pre-
cise location, Wright 3,819 (K, NY, US).
53. CHLORIS SAGRAEANA Achille Richard
in Sagra, Hist. Fisica Politica Natural Isla
Cuba XI. Botanica 315. 1850. (HOLO-
TTPE: "Chloris sagraeana nob. no. v. fol.
obtusa Cuba. . . ." P! Fragment of type:
US! ) Fig. 81, I-T.
Chloris morales-coelloi Leon ex Britton,
Bull. Torrey Bot. Club 53:4.58. 1926. (ISO-
TYPE: ". . . Cajobabo . . . not far from
mouth of the Jojo River, Oriente . . . Leon
. . . 12.320." US! )
Perennial 15 to 80 cm tall, cespitose, erect or
ascending from a geniculate base; sheaths gla-
brous, ligule short-ciliate; blades up to 15 cm
long, 1.3 to 3.0 mm wide, glabrous; spikes three
to six, .3.5 to 15.0 cm long (usually ca 4 to 9 cm
long), becoming widely divergent or somewhat
reflexed at maturity'; glumes narrowly lanceolate,
glabrous except for the scabrous midrib; spike-
lets appressed, ca four to nine per cm of the
scabrous rachis; first glume 1.6 to 2.2 mm long,
0.2 to 0.4 mm wide; second glume 2.4 to 3.5 mm
long, 0.5 to 0.7 mm wide; fertile lemma 2.6 to
3.5 mm long, 0.5 to 0.7 mm wide, elliptic, callus
bearded, with hairs less than 0.8 mm long, mar-
gins strongly appressed-pilose, the hairs usually
less than 0.5 mm long, keels glabrous, occasion-
ally very sparsely pilose, awn 7 to 13 mm long;
sterile floret one (occasionally two), 0.7 to 1.6
mm long, 0.2 to 0.5 mm wide, cvlindrical, gla-
brous, apex acute, awn 3.3 to 9.0 mm long; cary-
opsis 1.6 to 1.7 mm long, 0.4 to 0.5 mm wide,
ellipsoid, trigonous.
Chloris sagraeana is the most variable species
within the complex that includes C. arenaria, C.
cubensis, C. ekmanii, and C. radiata. This vari-
ation is especially noticeable with respect to
the size of the fertile and sterile lemmas. Fre-
quently specimens with exceptionally long lem-
mas will produce two sterile florets; and on oc-
casion, the lowennost "sterile" floret will pro-
duce a caryopsis. though it otherwise resembles
the upper sterile floret in nK)rphology. This vari-
ation in stmcture mav or mav not be correlated
with size. A specimen with two sterile florets,
coincidentally very robust, was named Chloris
morales-coelloi by Leon (in Britton, 1926).
Chloris sagraeana may be separated from C.
ekmanii by its more robust stature and broader
leaves and from C. cubensis in having shorter,
appressed hairs on the lemma margins (Table
13).
This species has been collected from a vari-
ety of habitats, including pinelands, serpentine
outcrops, coral rocks, and pastures ( Fig. 80,
triangles ) .
Representative specimens examined: BAHA-
MA ISLANDS: Fortune Island, Hitchcock s.n.,
Nov 1890 (F, MO); Inagua Island, Hitchcock
122
Brigham Young UNivEHsrri- Science Bulletin
Fig. 81. Chloris ciihensis and C. sagraeana. (A-H) C. ciibensis. (A) habit, .\ 1/3; (B) ligule region, x 3; (C)
spikelet, partly dissected, x7.5; (D) florets, x 7.5; (E-G) sterile florets, showing variation, x 10; (H) caryop-
sis, X 10. (I-T) C. sagraeana. ( I,J ) habit, x 1/3; (K) ligule region, x 3; (L) spikelet, partly dissected, x 7.5;
(M) spikelet with two sterile florets, partly dissected, x 7.5; (N) spikelet from type, partly dissected, .\ 7.5;
(0,P) additional florets, showing variation, x 7. .5; ( Q-S ) sterile florets, showing variation, x 10; (T) caryop-
sis, X 10.
Biological Series, Vol. 19, No.
Taxonomy of the Genu,s Chloris (Gramineae)
123
s.n., 3 Dec 1S90 ( F, MO, NY); Nassau, Britton
and Brace 596 (F, NY), Curtiss 81 (US). BAR-
BADOS: Allen s.n., Jan 1942 (K, NY). BARBU-
DA: S of Codrington, Danfortit 3 (US). CUBA:
Prov. Havana: vicinit\' of Havana, Ansouin 779
(US); between Morro and Cojimar, Ekmun 389
(US); Guanabacoa, Lomas de los Jatos, Ekman
10,902a (US); Near Marianas, Leon 1,965 (US);
near Luzano, Leon 4,807 (US); Santa Fe, Leon
13,096 (US); Cuabal de Jesus Maria Minas,
Leon and Hioram 4,783, 4,787 (US); Prov. Ma-
tanzas: Babiyal, Ekman 16,954 (US); Prov. Pi-
nar del Rio: La Grifa, Ekman 11,252 (US);
Mariel, coral reef E of town, Ekman 12.837
(US); Prov. Oriente: pinelands, crest of Sierra
Nipe, Leonard and Leonard 3,183 (US). DO-
MINICAN REPUBLIC: Prov. Montecriste:
Valle de Cibao, Villa Vasquez, Ekman 13,128
(K, US); Prov. de Santiago: Mao, Abbott s.n.,
23 Feb 1921 (US). GUADELOUPE: without
precise localit)', Diiss 4,110 (F, NY, US); Les
Pointes, Quesial 1,740 (US). HAITI: Pignon to
San Rafael, Cook et al. 138 (US); Port-au-
Prince, Ekman 2.107 ( K, US ) ; Tortue Island l)c-
tween La Vallee and Ravine-Rosetiere, Ekman
4,058 (K, US); Corail, Dutreuil, Ekman H
10,748 (K. US); Grande Cayemite, Pointe
Macon, Eyerdam 336 ( F, NY, US); vicinit\' of
Petionville, Leonard 4,984 ( NY, US ) ; vicinit)- of
Gros Morne, Dep. Artibonite, Leonard 9,839
(US); Port-de-Paix, Leonard and Leonard
11.051 (US); Baie des Moustiques, Leonard and
Leonard 12,067 (US); vicinitv of Jean Rabcl,
Leoiuird and Leonard 12,552 (K, MO, NY, US);
vicinitv of Mole St. Nicolas to Bombardopolis,
Leonard and Leonard 13,225 (UC, US).
JAMAICA: Cornwall Co., Savanna-la-Mar,
Hitchcock 9,890 (F, MO, NY). Middlesex Co.,
Ashlev Hall Savannah, Lower Clarendon, Harris
12,738 (F, NY, US).
54. CHLORIS CUBENSIS Hitchcock & Ek-
man c\ Hitchcock, U.S. Dept. Agric. Misc.
Publ. 243:131. 1936. (HOLOTYPE: " . . .
Cuba, prov. Oriente, Gamboa . . . Ekman
. . . 14960." US!) Fig. 81, A-H.
Perennial usualh' 50 to 60 cm tall, occasion-
ally up to 90 cm, tufted to stoloniferous; sheaths
glabrous, ligule a short-ciliate or naked crown
up to 0.5 mm tall; leaves 10 to 15 cm long, ca
1..5 mm wide, often folded, scabrous to sparseh'
pilose above, glabrous below; spikes three to
seven, 5.5 to 8.5 cm long, radiate, ascending;
spikelets appressed, imbricate, ca five to nine
per cm of the scabrous rachis; glumes narrowly
lanceolate, membranous, glabrous except for tlie
scabrous midnerve; first glume 1.7 to 2.4 mm
long, 0.2 to 0.3 mm wide; second glume 2.9 to
3.9 mm long, 0.2 to 0.4 mm wide; fertile" lemma
3.1 to 3.6 mm long, 0.6 to 0.7 mm wide, lanceo-
late, callus long-bearded, upper 2 3 of margins
long-ciliate with hairs up to 1 mm long, keel and
sides sparseh' pilose, lemma awn 9 to 12 mm
long; sterile floret one, 1.4 to 1.9 mm long, 0.4 to
0.6 mm wide, cylindrical, apex obtuse, awn 6 to
9 mm long; caryopsis ca 1.3 mm long, ca 0.3 to
0.4 mm wide, ellipsoid.
Chloris cubensis is similar to C. saeraeana,
from which it may be separated by its generally
larger size, less divergent mature spikes, and by
the long hairs on the lemma margins and, usu-
ally, on the lemma keel. (Fig. 81; Table 13).
Specimens have been collected on roadsides
and open dr)- areas (Fig. SO, squares).
Representative specimens examined: AN-
TIGUA: Nonsuch Bav, Facey Creek, Box 556
(US). BARBADOS: 'without precise location,
Hitchcock 16.499 (US). CUBA: Prov. Cama-
gii(>y: Minas, Ekman 15,368 ( F, NY, US); Prov.
Havana: Loma de Tierra, Ekman 13,469 (K,
US); Vibora, Leon 859 (US); Calvarco, Leon
860 (US); Prov. Las Villas: Loma de Belen,
Leon 15,627 (US); Prov. Oriente: Holguin, Ek-
man 1,014 (F, K, MO, NY, UC, US), Ekman
15,715 (F, NY, US). HAITI: Tortue Island, vi-
cinitv of Basse Terre, Leonard and Leonard 12,-
5.:5,S'(US). JAMAICA: Hanover Park, Bailev's
Point, Adams 7,964 ( BM ) ; Cockpit, Vere, Har-
ris 12.462 (F, MO, NY, US); Spanish Lower
Road, Harris 12,476 (K). PUERTO RICO: Rio
Piednxs, Alberts 273 (US).
55. CHLORIS MOSSAMBICENSIS Schumann,
Notizbl. Bot. Gart. Berlin 1:104. 1895. (ISO-
TYPE: "Mozambik, Cabareino Grande . . .
No. 88. leg. Prelado." BM!) Fig. 82.
Tetrapogon mossambicensis ( Schumann )
Chippcndahl ex B.S. Fisher in Meredith,
Grasses and Pastures of South Africa 198.
1955. (Based on Chloris mossambicensis
Schumann.)
Perennial, tufted to aggressively stolonifer-
ous, 20 to 100 cm tall (usually 40 to 80 cm);
sheaths glabrous to scabrous; ligule a short-cili-
ate crov.'n, often with a few long hairs inter-
spersed, especially near the margins; blades up
to 30 cm long, ca 8 mm wide, obtuse to acumi-
nate; spikes three to six (often four or five), 3
to 10 cm long, divaricate to horizontally spread-
ing; spikelets sparsely inserted, imbricate, diver-
gent, ca eight or nine per cm of the scabrous
rachis; glumes narrowly lanceolate, thin, mem-
branous, glabrous except for the scabrous mid-
124
Brigham Young University Science Bulletin
Fig. 82. ChloTis mossambicensis. (A) habit, x 1/5; (B) spikelet, partly dissected, x 5; (C) sterile florets, x
7.5; (D) partly dissected "sterile" florets, showing upper sterile floret (u.s.f. ), palea of lower "sterile" floret
(pa.), lemma of lower "sterile" floret (1.), and flower, x 20; (E-J) florets, showing variation, x 5; (K)
caryopsis, x 7.5.
nerve; first glume 1.6 to 2.6 mm long, ca 0.2 mm
wide; second glume 3.0 to 4.4 mm long, 0.3 to
0.4 mm wide; fertile lemma 3.0 tO' 4.7 mm long,
0.8 to 1.2 mm wide, elliptic to ovate, apex acute
to obtuse, callus densely bearded, margins ap-
pressed-pilose below, becoming ciliate near the
apex with hairs ca 1 mm long, keel appressed-
pilose, internerves glabrous, awn 3.5 to 12.0 mm
long; sterile floret usually one (occasionally
two), 1.5 to 3.3 mm long, 0.7 to 1.5 mm wide,
narrowly to broadh' triangular, usually barren,
occasionally fertile and setting seed, apex trun-
cate, usually glabrous, very occasionally sparsely
pilose, awn 2 to 9 mm long; caryopsis 1.2 to 1.3
mm long, 0.3 to 0.5 mm wide, ovoid to obovoid,
rounded to trigonous.
Tliis species has been treated variously as a
member of Chloris or of Tetrapogon. As brought
out in i-arlier discussion, the two genera are very
closely related and must be somewhat arbitrari-
ly separated. Essentially, Tetrapogon regularly
has more than one basal fertile floret, vyhile
Chloris has just one. A sampling of nineteen
specimens, from different locations in southern
Africa, showed seventeen with a single fertile
floret and two \\ath two fertile florets. Because
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
125
Fig. 83. Distribution of Chloris mossambicensis (south-
em Africa).
of this and an overall rc,sein1)laco to tlie prepond-
erance of Chloris species, I am treating this
species as a member of the genus CJdoris.
Chloris mossambicensis is wideK' distributed
in soutliern Africa (Fig. 83), where it lias l)een
collected in a variety of habitats. A more de-
tailed and expanded distribution map may be
found in Chippendall ( 1955).
Representative specimens examined; KEN-
YA: 14 Falls, Athi River, near Doinyo Sapuk,
Bogdan AB 1,590 (K); Nairobi to Mombasa
Road, near Samburu Station, Bogdan AB 3,616
(K); Mariakani, Bogdan AB 5,429 (K); Kwale
Dtr., near Tani, bc-twetn Samburu and MacKin-
non Road, Drummond and llensleij 4,189 (K).
MOgAMBIQUE; Sul do Save, Maputo River,
entre Changalane e Catuane, Mijre and Carval-
hao 1,440 (K); between Moamba and Boane,
Schwcickcrdt 1,919 A (K); Louren90 Marques.
Rio Limpopo, entre Mianga e Guija, Torre 6,614
(K). SWAZILAND: 11 mi W of Stegi, Codd
and Dyer 2,863 (K). TANZANIA: Tanga Prov..
Mkwaja, Pangani, banks of Bohomaia River, van
Renshurg 516 (K). UNION OF SOUTH AFRI-
CA: Natal: 8 mi N of Mkuze on Candover Road,
Edwards 3,257 (K); Zululand, lower Umfolozi
region. Ward 2,442 (K); Transvaal: 16.5 mi SE
of Pretoria Kop, Acocks 16,713 (K); Barberton
Dtr., near Krokodipoort siding, Liehenherg 2,-
406 (K); Kruger National Park, 4 mi S of Sku-
kuza, de WinU'r and Codd 546 (K).
56. CHLORIS BURMENSIS sp. nov. Fig. 84.
Culmi inferiores cum foliis igiioti. Foliorum
vaginae glabrae; ligulae breves ± truncatae,
scabrae vel brcviterciliatae; laminae 10-18 cm
longae, 5-16 mm latae, acuminatae, inferne
glabrae, superne scabrae. Spicae 7-9, adscen-
dcntes, 6-18 cm longae, dilute virides. Spiculae
appressae, secus rhachin scabram ± 9-10 per
utnimque cm dispositae. Glumae lanceolatae
tenues membranaceae, praeter carinam scabram
glabrae. Cluma prima 1.7-2.2 mm longa, 0.2
-0..3 mm lata. Gluma secunda 2.5-3.2 mm
longa. :+: 0.5 mm lata. Lemma fertile ellipticum,
apiee acutum, 2.5-2.8 mm longum, 0..8-0.9
mm latum, callo angusto barbato, secus margines
carinamque dense appressepilosum; arista 1-2
mm longa. Flosculi steriles 2, inferior obovoideus
obtusus vel acutus, 1.5-1.7 mm longus, 0.6-
0.7 mm latus, praeter carinam sparsim appresse-
pilosam, glabcr, arista 0.7 to 1.6 mm longa, su-
perior ob()\oideus apicc ± tmncatus 1.0-1.5
mm longus, 0.5-0.8 mm latus, versus apicem
scaber ceterum glaber, arista rb 0.5 mm longa.
Caryopsis ignota.
Lower stems and leaves unknown. Leaf
sheaths glabrous, ligule a densely scabrous or
short-eiliate, tnincate crown; upper leaf blades
10 to 18 cm long, 5 to 16 mm wide, scabrous
above, glabrous below, acuminate; spikes seven
to nine, 6 to 18 cm long, ascending, pale green;
spikelets appressed, ca nine or ten per cm of
the scabrous rachis; glumes lanceolate to nar-
rowly ovate, thin and mtMubranous, glabrous ex-
cept for the scabrous midnervc; first glume 1.7 to
2.2 mm long, 0.2 to 0.3 mm wide; second glume
2.5 to 3.2 mm long, ca 0.5 nmi wide; fertile lem-
ma 2.5 to 2.8 mm long, 0.8 to 0.9 mm wide, el-
liptic, callus nanow, bearded, margins and keel
densel\- appressed-pilose, apex acute, awn 1 to 2
mm long; sterile florets two; lower sterile floret
1.5 to 1.7 mm long, 0.6 to 0.7 mm wide, obo-
\oid, keel lightly appressed-pilose, otherwise
glabrous, apex acute to obtuse, awn 0.7 to 1.6
mm long, upper sterile floret I.O to 1.5 mm long,
0.5 to 0.8 mm wide, broadlv obovoid, glabrous
below, scabrous at the tmncate to rounded apex,
awn ca 0.5 mm long; anthers ca 0.4 mm long;
car\opsis unknown.
HOLOTYPE: Burma, roadside between
Meiktila and Pvawbye. On calcareous stiff clay.
V. Them Lain 311.' 23 Oct 1947. K!
Chloris biirmcnsis is, \\ithout question, dis-
tinct from all other species in Chloris. The com-
bination of awned sterile florets in pairs and the
densely appressed-pubescent, elliptic fertile
florets (Fig. 84) separates this taxon from all
other species. The onl\- specimen available— the
holotype— is unfortunately without any basal
parts; so infonnation on height, habit, or longev-
ity is not available. Superficially, C. hurmensis
resembles some species of Tetrapogon, but de-
tails of its spikelet structure place it definitely in
Chloris.
126
Brigham Young University Science Bulletin
K VL VM
Fig. 84. Chloris burmensis. (A) inflorescence and upper culm, x 1/3; (B) ventral view of portion of rachis,
X 3; (C) dorsal view of portion of rachis, x 3; (D) spikelet, partly dissected, x 10; (E) fertile floret with one
half removed, showing palea and flower, x 15; (F) palea, ventral view, x 15; (G-I) lower sterile florets,
showing variation, x 10; (J-M) upper sterile floret, showing variation, x 10.
UNIDENTIFIED, REJECTED, OR REASSIGNED NAMES
Tlie following alphabetical list includes all
names used in Chloris that are presently uniden-
tifiable, not located, or assigned to other genera.
Chloris acicularis Lindley ex Mitchell, Jour.
Exped. Interior Trop. Australia 33. 1848. Basis
of Enteropogon acicularis (Lindley) Lazarides.
Chloris acicularis var. queenslandiae Domin,
Biblioth. Bot. 85:368. 1915. = Enteropogon
acicularis (Lindley) Lazarides, on basis of the
description.
Chloris acuminata Trinius, Species Gram.
Icon. 3:/)/. 305. 1831. =Eustachi/s disticho-
phtjlla (Lagasca) Nees.
Chloris affinis Caro and Sanchez, Kurtziana
6:224. 1971. Fig. 2. Specimens of this plant
were not available in time to be considered
in this publication. On the basis of the original
description and accompanying illustration, it
would seem to be most closely related to
Chloris pijcnothrix Trin.
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
127
Chloris agowensis Hochstetter ex Kegel, An-
notationes Bot. Index Seminum 26. 1862.
Nomen nudum, synonym of Chloris spathacea
Hochstetter.
Chloris anoplia (Nees) Foumier ex Hem-
sley, Biol. Cent.-Amer. Bot. 3:558. 1885. =?,
possibly Leptocldoa sp. Based on Leptochloa
anoplia Nees.
Chloris argentina (Hackel) Lillo and Parodi,
Physis 4:180. 1918. Based on Chloris clis-
tichophijlhi var. argentina Hackel which — Eu-
stacht/s retiisa (Lagasca) Kunth.
Chloris argentinensis Parodi, Rev. Fac. Ag-
ron. and Vet. Univ. Nac. Buenos Aires 2:28.3.
1919. Error for Chloris argentina?
Chloris aristida Salzmann ex Steudel, Syn.
PI. Glum. 1:218. 1854. Nomen nudum, as s\n-
onyin of Gijmnopogon foliosus Nees.
Chloris hahiensis Steudel, S\n. Pi. Glum.
1:208. 1854. =Eitstachi)s caribaea (Sprongcl)
Herter.
Chloris hahiensis f. glabrescens Hackel, Re-
pert. Sp. Nov. 8:46. 1910. =Eustac}it/s cari-
baea (Spri'iigel) Herter.
Chloris hoivinii A. Camus, Bull. Soc. Bot.
France 79:845. 1933. Basis of Daknopholis
hoivinii (A. Camus) Clayton.
Chloris brachi/athcra (Hackel) Herter, Re-
vista Sudamer. Bot. 6:146. 1940. Based on
Chloris ciliata Swartz var. brachi/athera Hackel
in Kurtz, which see.
Chloris brevipila Rosengurtt and Iza<ji;uirre
de Artucio, Bol. Soc. Argentina Bot. 12:120. Fig.
2. 1968. ^Eustachtjs sp., on basis of descrip-
tion and illustration.
Chloris calvesccns Hackel in Chodat and
Hassler, Bull. Herb. Boissier II. 4:279. 1904.
= Eustaclii/s sp., on basis of description.
Chloris campulodes Trinius ex Steudel, Noin.
Bot. ed. 2. 1:353. 1840. Nomen nudum.
Chloris capensis (Houttyn) Thellung, Re-
pert. Sp. Nov. 10:289. 1912. Based on Andro-
pogon capensis Houttyn, which is based on
Andropogon muticum Linnaeus, a superfluous
name. Andropogon muticum Linnaeus is uniden-
tifiable.
Chloris capensis var. hahiensis (Steudel)
Parodi, Revista Argentina Agron. 20:26. 1953.
— Eustachys caribaea ( Sprengel ) Herter.
Chloris capensis var. glabrescens (Hackel)
Parodi, Revista Argentina Agron. 20:20. 1953.
= E«.sfflt/i(/s caribaea (Sprengel) Herter.
Chloris card)aea Sprengel, Syst. Veg. 1:295.
1825. Basis of Eustachys caribaea (Sprengel)
Herter.
Chloris castilloniaiui Lillo and Parodi var.
pubescens Caro and Sanchez, Kurtziana 6:230.
1971. Specimens of this plant were not avail-
able in time to be considered in this publica-
tion. On the basis of the original description
it would seem to be part of the variation pat-
tern of Chloris castilloniaija Lillo and Parodi.
Chloris cenchriformis (Achille Richard)
Baillon ex Schweinfurth and Ascherson, Beitr.
Flora Aethiopiens 310. 1867. Based on Lepi-
dopironia cenchriformis Achille Richard. =Tet-
rapogon cenchriformis (Achille Richard) Pilger.
Chloris cheesmanii Hackel ex Cheesman,
Trans. Linnean Soc. Bot. 6:305. 1903. =En-
teropogon unispiceus (F. Mueller) Clayton on
basis of description.
Chloris ciliata Swartz var. brachyathera
Hackel ex Kurtz, Bol. Acad. Ciencias Cordoba
(Rep. Argentina) 16:257. 1900. Without de-
scription. As described bv Parodi ( Rev. Fac.
.\gron. and Vet. Univ. Nac. Buenos Aires 2:271.
1919) = Chloris ciliata Swartz.
Chloris compressa De Candollc, Cat. Horti
VIonspeliensis 94. 1813. =? Specimens P!,
labeled as this species, of uncertain origin, are
C. radiuta Swartz. A photo, in US!, of a speci-
men in Montpellier labeled as C compressa,
perhaps in de Candolle's script, is also C
radiata. The protologue might applv equally to
C. radiata or C. virgata Swartz.
Chloris confertifolia Trinius in Sprengel,
Neue Entdeckungen 2:74. 1821. =Eustaclnjs
disticiu^plnjlla (Lagasca) Nees.
Chloris ctenioides Steudel, Syn. PI. Glum.
1:423. 1854. =Dactyloctenium sp., on basis of
fragment of type ( US! ) from CN.
Chloris curtipendula Michaux, Flora Bor.
Amer. 1:59. 1803. Basis of Bouteloua curti-
pendula (Michaux) Torrev in Emory.
Chloris cynodoidea Lagasca, Elenchus Plan-
tarum 5. 1816. Nomen nudem.
Chloris cynodon (Linnaeus) Trinius, Gram.
Unifloris 229. 1824. Based on Panicum dactylon
Linnaeus —Cynodon dactylon (Linnaeus) Per-
soon.
Chloris delicatula Clarke in Hooker, f. Flora
British India 7:290. 1896. Basis of Gymnopogon
delicatula (Clarke) Bor.
Chloris dichanthioides Everist, Queensland
Agric. Jour. 49:4.32. /)/. 155, 19.38. Basis of
Austrochloris dichanthioides (Everist) Lazar-
ides.
Chloris digitaria Humboldt, Bonpland,
Kunth, Nov. Gen. and Sp. PI. 1:168. 1816.
= Leptochloa sp. on basis of description.
Chloris digitata (Roxburgh) Steudel, Syn.
PI. Glum. 1:207. 1854. Based on Melica digi-
tata Roxburgh which = Enteropogon dolicho-
stachya (Lagasca) Lazarides.
128
Bricham Young University Science Bulletin
Chloris (listacluja Bojer, Hortus Mauritianus
371. 1837. = Chloris filiformis (Vahl) Poiret?
An unnamed Bojer specimen from Mauritius
(in G!) is C. filiformis. The protologue is in-
complete.
Chloris distichophijlla Lagasca, Gen. Sp.
Nov. 4. 1816. Basis of Eustachys distichophijlla
(Lagasca) Nees.
Chloris distichophijlla var. acuminata (Trin-
ius) Hackel, Bull. Herb. Boissier II. 4:279.
1903. = Eustachys distichophijlla (Lagasca)
Nees.
Chloris distichophijlla var. argentina Hackel
in Stuckcrt, Anales Mus. Nac. Hist. Nat. Buenos
Aires 11:113. 1904. = Eustachys disticlwphylla
(Lagasca) Nees.
Chloris distichophijlla var. f^emdna Hackel
in Stuckert, Anales Mus. Nac. Hist. Nat. Buenos
Aires 11:113. 1904. = Eustachys distichophijlla
(Lagasca) Nees.
Chloris dolichostachya Lagasca, Gen. Sp.
Nov. 5. 816. Basis of Enteropogon dolicho-
stachya (Lagasca) Lazarides.
Chloris duhia Humboldt, Bonpland, Kunth,
Nov. Gen. and Sp. PI. 1:169. 1816; /;/. 694.
1825. Basis of Leptochloa duhia (Humboldt,
Bonpland, Kunth) Net>s.
Chloris durandiana Schultes, Mantissa 2:.341.
1824. Based on Chloris gracilis Durand, which
may equal Chloris radiata (Linnaeus) Swartz.
See also C gracilis Durand in this list.
Chloris dusenii Ekman, Arkiv for Botanik
10:26. 1911. = Eustachys uligino.sus (Hackel)
Herter; isotype in US!
Chloris elegans (Kunth) Roberty, Petite
Flore Ouest.-Afrique 387. 1954. Basionym not
cited.
Chloris elegans (Kunth) Roberty, Bull. Inst.
Fran9aLse Afrique Noire Ser. A. 17:51. 1955.
Based on Ctenium elegans Kunth.
Chloris eleusinoides Grisebach, Flora British
West Indian Islands 539. 1864. Unidentified.
Type not seen, but description suggests Chloris
sagraeana A. Rich.
Chloris emarginata Beauvois, Ess. Agrost. 79,
158. 1812. Nomen midem.
Chloris ecpdtans Trinius, Sp. Gram. Icon. 3.
pi. 309. 1831. =Eustachijs paspaloides (Vahl)
Lanza and Mattei.
Chloris falcata Swartz, Ges. Naturf. Freunde
Berlin Neuc Schrift 3:160. /;/. 1. Fig. 1. 1801.
= Harpechloa capensis Kunth.
Chloris fasciculata Schrader in Schultes,
Mantissa 2;.3.39. 1824. =Eustachys disticho-
phijlla (Lagasca) Nees.
Chloris fasciculata (Linnaeus) Thellung,
Repert. Sp. Nov. 10:289. 1912. Non Chloris
fasciculata Schrader in Schultes. Based on An-
dropogon fa.yciculatum Linnaeus =Microstegium
sp. on basis of holotype ( LINN! ).
Chloris floccifolia ( Forskal ) Pcjiret in
Lamarck, Encycl. Method. Bot. Suppl. 2:238.
1811. Based on Cynosurus floccifolius Forskal
= Eleusine floccifolia (Forskal) Sprengel.
Chloris floridona (Chapman) Wood, Amer.
Bot. and Florist Pt. 2. 407. 1871. = Eustachys
floridana Chapman.
Chloris foliosa Willdenovv, Sp. PI. 4:924.
1806. Basis of Gymnopogon foliosus (Will-
denovv) Nees. Fragment of t\pe, from B, in
US!
Chloris foliosa Lagasca, Gen. Sp. PI. Nov. 5.
1816. Non Chloris foliosa Willdenovv. Uniden-
tified. A fragment, in US!, of a Lagasca speci-
men dated 1820. from M, is Gymnojwgon sp.
Chloris geminata Hochstetter, Flora 38:205.
1855. =Tetrapogon sp. on basis of description.
Chloris glaiica (Chapman) Wood, Amer.
Bf>t. and Florist Pt. 2. 407. 1871. = Eustachys
glauca Chapman.
Chloris glahrescens (Hackel) Rosengurtt and
Izaguirre, Bol. Soc. Argentina Bot. 12:123.
1968. Based on Chloris hahicnsis var. glabres-
cens Hackel = Eustachys caribaea (Sprengel)
Herter.
Chloris gracilis Durand, Chloridis Speciebus
10. 1808. =? Tvpe not seen, description may
be of Chloris radiata (Linnaeus) Swartz. Non
Chloris gracilis Humboldt, Bonpland, Kunth,
1816.
Chloris gracilis Humboldt, Bonpland, Kunth,
Nov. Gen. Sp. PI. 1:168. 1816. Non Chloris
gracilis Durand. = Leptochloa sp. on basis of
description.
Chloris gryllus (Linnaeus) Honckeny, Svn.
PI. Gennaniae 1:437. 1792. =:Chry.wpogon
gryllus (Linnaeus) Trinius on basis of descrip-
tion. Based on Andropogon gryllus Linnaeus.
Chloris "uineensis Schumacher, Beskr. Guin-
eiske Planter 75. 1827. =Dactijlocteniuin sp.
on basis of photograph of type ( in US! ) in the
Isert and Thonning Herbarium.
Chloris hlspida Durand, Chloridis Speciebus
17. 1808. =Bouteloua sp. on basis of
description.
Chloris imherhis Desfontaines, Cat. PI. Pa-
risiensis ed. 3. 15. 1829. Nomen nudem.
Chloris incomplcta Roth ex Roemer and
Schultes, Syst. Veg. 2:607. 1817. = Entero-
pogon dolichostachya (Lagasca) Lazarides on
basis of photograph of tvpe in K!
Chloris lepidopironia Desveaux ex Regel,
Annotationes Bot. Index Seminum 26. 1862.
Biological Series, Vol. 19, No. 2 Taxonomy of the Genus Chloris (Gramineae)
129
Nomen mulcm. lierbarium name published as a
synonym.
Chloris longiharbis Michaux ex Beauvois,
Ess. Agrost. 79, 158. 1812. Nomen luidcm.
Chloris nmcrantha Desveaux, Opuscules 73.
1831. =Tetrapogon sp. on basis of description.
Chloris macrantha Jaubcrt and Spach, lllustr.
PI. Orient. 4:42. /;/. 328. 1851. Non Chloris
macrantha Desveaux. —Tetrapogon sp. on basis
of description.
Chloris macrostachija Hochstetter ex Achille
Richard, Tent. Florae Abxssinicae 2:408. 1850.
Basis of Enteropogon macrostachyus (Hoch-
stetter ex Achille Richard) Munro ex Bentham.
Chloris maritimiis (Kunth) Trinius, Gram.
Unifloris 235. 1824. Based on Ct/nodon mari-
timus Kunth. =- Cijnodon sp.
Chloris mearnsii Merrill, Philippine Jour.
Sci. 3:220. 1908. = Enteropogon sp. on basis
of description.
Chloris monostacJiya Michaux, Flora Bor.
Anier. 1:59. 1S03. —Ctenium sp. on basis of
description.
Chloris monostachijo (Vahl) Poiret in
Lamarck, Enc\cl. Method. Bot. Suppl. 2:238.
1811. Non Chloris monostachi/a Michaux. Basis
of Enteropogon monostachi/os (\'ahl) K. Schu-
mann ex Engl. ( Based on Ct/nosurus niono-
stachijus X'ahl. )
Chloris moorci F. Mueller, Linnaea 25:444.
1852. Unidentified. A specimen at K!, collected
and labeled as C. moorei by von Mueller,
though not the type, is Enteropogon acicuhiris
( Lindlev ) Lazarides.
Chloris mucronata Michaux, Flora Bor.
Amer. 1:59. 1803. =Dactijloctenium sp. on
basis of description.
Chloris multiradiata Hochstetter \'ar. ragazzii
Pirotta, Anales 1st. Reale Bot. Roma 6:157.
1896. Unidentific>d, possibh' a \'ariant of Chloris
virgata Swartz.
Chloris neglecta Nash, Bull. Tone\' Bot. Club
22:423. 1895. Basis of Eustachijs neglecta
(Nash) Nash.
Chloris ohttisifolia Trinius, Gram. Unifloris
233. 1824. Unidentified, description suggests
Eustaclu/s tener ( Presl ) A. Camus.
Chloris ohttisifolia Balansa, Jour, de Bot.
(Morot) 4:166. 1890. Non Chloris ohttisifolia
Trinius 1824. =Etistachys tener (Presl) A.
Camus.
Chloris pallida Willdenow, Sp. PI. 4:293.
1806. Based on Andropogon provinciale Retz. =
?. The letter name unidentified, though descrip-
tion is not of a Chloris.
Chloris pallida ( Edgeworth ) Hooker, f..
Flora British India 7:289. 1896. Non Chloris
pallida Willdenow. =Schoenfeldia pallida
Edgeworth, on basis of description.
Chloris panicea Willdenow, Sp. PI. 4:923.
1806. ^Enteropogon dolichostachya (Lagasca)
Lazarides. Type not seen, but description
strongly suggests this species.
Chloris parva G. Mimeur, Bull. Mus. Hist.
Nat. (Paris) 22:128. 1950. =Chloris prieiirii
Kunth? Mimeur cites a specimen sent in 1880,
Talmy 86, from Senegal and deposited in P.
This cannot be located, but another specimen
sent by Talmy, also from Senegal, but in 1882,
is Chloris prictirii. The original description like-
wise strongly suggests this species.
Chloris parvispictila Caro and Sanchez,
Kuitziana 6:227. 1971. Fig. 3. Specimens of
this plant were not available in time to be
considered in this publication. On the basis
of the original description and accompanying
illustration it would seem to lie most closely
related to Chloris dandyana.
Chloris paspaloidcs Hochstetter, Flora 38:206.
1885. =Etistachys paspaloidcs (Vahl) Lanza
and Mattei.
Chloris paiili-diicis Schweinfurth, Beitr.
Flora Aethiopiensis 298. 1867. Nomen inidtim.
Chloris paijtensis Steudel, S\ii. PI. Glum.
1:207. 1854. =Cynodon dactijlon on basis of
fragment of type (in US!) from Urville
Herbarium.
Chloris pedicellata Steudel, Flora 33:232.
1850. Nomen ntidttm, as s\ nonvm of Schellingia
tenera Steudel.
Chloris pendtda Salzmann ex Steudel, Syn.
PI. Glum. 1:208, 1854. Nomen nudum, as
svnonym of Chloris hahiensis Steudel.
Chloris penicellata (Vahl) Persoon, Syn. PI.
1:87. 1805. =Tetrapogon sp.? on basis of de-
scription of Cynostirtis penicillatus Vahl, upon
which it is based.
Chloris penicillata Jan. ex Trinius, Gram.
Uniflores 231. 1824. Non Chloris penicillata
(Vahl) Persoon. Nomen ntidtim. as synonym
of Chloris compressa De Candolle.
Chloris perrieri A. Camus, Bull. Soc. Bot.
France 96:93. 1949. =Daknopholis hoivinii
(A. Camus) Clayton on basis of holot\pe in P!
Chloris peregrina Durand, Chlorides Specie-
bus 11. 1808. Unidentified. Type not seen,
description suggests Chloris inflata Link.
Chloris petraea Swartz, Prodr. Veg. Ind. Occ.
25. 1788. Basis of Eustachys petraea (Swartz)
Desfontaines.
Chloris piperita Michaux ex Steudel, Nom.
Bot. ed. 2. 1:353. 1840. Nomen ntidum, as
synonym of Campulosus rnonostachytis Beauvois.
Chloris poaeformis Humboldt, Bonpland,
130
Brigham Young University Science Bijlletin
Kunth, Nov. Gen. and Sp. PL 1:169. 1816.
= LeptochIoa sp. on basis of description.
Chloris poltjdacttjla Swartz var. hreviaristata
Hackel in Kurtz, Bol. Acad. Ciencias, Cordoba
(Rep. Argentina) 16:257. 1899. Unidentified.
Type specimen not seen, but description sug-
gests Chloris ciliata Swartz.
Chloris pohjdactijla Swartz f. major Kegel,
Annotationes Bot. Index Seminum 25. 1862.
Unidentified. Type not seen. Description incon-
clusive.
Chloris polydactijla Swartz f. media Kegel,
Annotationes Bot. Index Seminum. 25. 1862.
Nomen nudum.
Chloris pohjdactijla Swartz f. minor Kegel,
Annotationes Bot. Index Seminum 25. 1862.
Based on Chloris elegans Humboldt, Bonpland,
Kunth, which =Chloris vir^ata Swartz. Prob-
ably misapplied to Chloris pohjdactijla Swartz.
Chloris polijstachia Lagasca, Elenchus Plan-
tarum 5. 1816. Nomen nudum.
Chloris procumhens Durand, Chlorides Spe-
ciebus 16. 1808. =Bouteloua sp. on basis of
description.
Chloris prostrata ( Willdenow ) Poiret in
Lamarck, Encycl. Method. Bot. Suppl. 2:239.
1811. Based on Dacttjloctenium prostratum
Willdenow. = Dacttjloctenium sp.
Chloris pubescens Peyritsch in Wawra and
Peyritsch, Akad. Wiss. Wien Sitzungsber. Math.
-Naturwiss. Klasse 38:385. 1860. Non C.
pubescens Lagasca, 1805. —Eustachys sp. on
basis of description.
Chloris pulchra Schumacher, Beskr. Guinei-
ske Planter 76. 1827. =Ctenium sp. Photo-
graph of type, in US!, from Isert and Thonning
Herbarium.
Chloris queenslandiac Domin, Biblioth. Bot.
85:368. 1915. Nomen nudum, as synonym of
Chloris acicularis var. queenslandiac Domin.
Chloris radiata Heyne ex Both, Nov. PI.
Indiae Orientalis 61. 1821. Non Chloris radiata
Swartz. Nomen nudum, as synonym of Chloris
incompleta Roth, which = Enteropogon dolicho-
stachtja (Lagasca) Lazarides.
Chloris radiata Swartz var. beijrichiana
(Hackel) Kurtz in Rio and Achaval, Geog. Prov.
Cordoba 1:331. 1904. Attributed incorrectly to
Hackel by Kurtz, presumably based on Chloris
betjrichiana Kunth, but basionym not cited.
Chloris ramosissima A. Camus, Not. Syst.
Mus. Hist. Nat. Paris 12:155. 1946. =Dakno-
pholis boivinii ( Camus ) Clayton on basis of
holotype in P!
Chloris repens Steudel, Nom. Bot. ed. 2.
1:353. 1840. Nomen nudum, as synonym of
Eleusine indica (Linnaeus) Gaertner.
Chloris retusa Lagasca, Elenchus Plantarum
5. 1816. = Eustachys retusa (Lagasca) Kunth
on basis of fragment and photograph of type
in US!
Chloris rhachitricha Steudel, Syn. PI. Glum.
1:205. 1854. =Eulalia ? on basis of type speci-
men in P!
Chloris ridleyi Hackel, Oesterr. Bot. Zeitschr.
52:237. 1902. = Eustachys tener (Presl) A.
Camus. Isotype at K!
CIdoris roxhurghii Edgeworth, Jour. Asiatic
Soc. Bengal 21:183. 1851. Based on Melica
digitata Roxburgh which = Enteropogon doli-
chostachi/a (Lagasca) Lazarides.
Chloris rufescens Lagasca, Var. Ciencias Lit.
and Artes (Madrid) 4:143. 1805. Type not
located. Original description too meager for
recognition of plant.
Chloris savatieri Baillon, Bull. Soc. Linn.
(Paris) 2:1070. 1893. Nomen nudum, given as
an herbarium name in a discussion of Tetra-
pogon sp.
Chloris scariosa Beauvois, Ess. Agrost. 79,
158. 1812. Non Chloris scariosa F. Mueller.
Nomen nudum.
Chloris secundus (Pursh) A. Eaton, Manual
Bot. North Amer. ed. 5. 173. 1829. Based on
Ctpwsurus secundus Pursh. =Bouteloua sp. on
basis of description.
Clitoris septentrionalis C. Mueller, Bot.
Zeitung (BerHn) 19:340. 1861. =Eu.stachys
petraea (Swartz) Desveaux on basis of descrip-
tion.
Chloris simplex Schumacher, Beskr. Guinei-
ske Planter 74. 1827. = Enteropogon sp. on
basis of photograph (in K! ) of type speci-
men in Isert and Thonning Herbarium.
Chloris spathacea Hochstetter ex Steudel,
Syn. PI. Glum. 1:204. 1854. =Tetrapogon
cenchriformis on basis of isotype at K!
Chloris spathacea Baillon, Bull. Soc. Linn.
(Paris) 2:1070. 1893. Non Chloris spathacea
Hochstetter, 1854. =Tetrapogon sp. on basis
of tvpe in P!
Chloris subaequigluma Rendle, Cat. African
Plants collected by Dr. F. Welwitsch 2:222.
1899. Basis of Bracteola .subaequigluma
(Rendle) C. E. Hubbard.
Chloris swartziana Doell in Martins, Flora
Brasiliensis 23:68. 1878. —Eustachys petraea
(Swaitz) Desveaux, on basis of description.
Chloris swartzii C. Mueller, Bot. Zeitung
(Berlin) 19:341. 1861. = Eustachys petraea
(Swartz) Desveaux. Based on Chloris petraea
Swartz.
Chloris tenella Koen ex Roxburgh. Hortus
Bengalensis 82. 1814. Nomen nudum. Chloris
Biological Series, Vol. 19, No. 2 T.\xonomy of the Genus Chloris (Gramineae)
131
tenella Roxburgh, Flora Indica ed. Carey, 1:330.
1820. =Tetrapogon sp. on basis of description.
Chloris tenera (Prcsl) Scribncr, Ann. Rcpt.
Missouri Bot. Card. 10:41. /)/. 40. 1899.
= Eustachys tener (Prcsl) A. Camus.
Chloris temiior Poiret ex Schultes, Mantissa
2:321. 1824. Nomen nii(him, as svnon\'ni of
Leptcliloa cynosuroicles Rocmcr and Schultes.
Chloris tetrameris Trinius, Gram. L'nitlorcs
235. 1824. =Enterof)ogon. doliclwstadnju
(Lagasca) Lazarides on basis of description.
Chloris transiens Pilger, Bot. Jahrb. 51:418.
1914. Basis of Schocnefehlia transiens (Pilger)
Chiovenda.
Chloris triangiihita Hochstctter ex Achille
Richard, Tent. Florae Ab\ssinicae 2:409. 1850.
= Tetrapogon sp. on basis of description.
Chloris uliginosa Hackel, Repert. Sp. Nov.
7:320. 1909. Basis of Eustachijs nligiiwsus
(Hackel) Herter.
Chloris unispicea F. Mueller, Fragm.
Phytogr. Australiae 7:118. 1870. Basis of En-
teropogon unispicea (Mueller) Clayton.
Chloris villosa ( Desfontaines ) Persoon, Syn.
PI. 1:87. 1805. Based on Tetrapogon villosits
Desfontaines. = Tetrapogon sp.
Chloris villosa var. sinaica Decaisne, Ann.
Sci. Nat. Bot. (Paris) II. 2:12. 1834. = Tetra-
pogon sp. on basis of description.
Chloris virgattts Nees ex Steudel, Syn. PI.
Glum. 1:213. 1854. Nan Chloris virgata
Swartz. Unidentified. Description ma\- be of
Leptochloa sp.
Leptochloris greggii Munro ex Merrill, U.S.
Dept. Agric. Div. Agrost. Circ. 32:7. 1901. Not
located.
Trichloris latifolia Vasey, U.S. Dept. Agric.
Spec. Rept. 63:32. 1883. Not located.
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S-(^^' ^ls°''l
Mus ccMp zcRfL'flham Young University
LIBRARY ^ . D II X-
Science Bulletin
HARVARD
UNIVERSITY
A COMPARISON OF METEOROLOGIC
MEASUREMENTS FROM IRRIGATED
AND NON-IRRIGATED PLOTS,
PROVO, UTAH, 1970-1972
by
Ferron L. Andersen
Phil D. Wright
and
J. Carl Fox
BIOLOGICAL SERIES — VOLUME XIX, NUMBER 3
APRIL 1974/ISSN 0068-1024
BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN
BIOLOGICAL SERIES
Editor: Stanley L. Welsh, Department of Botany,
Brigham Young University, Provo, Utah
Acting Editor: Vernon J. Tipton, Zoology
Members of the Editorial Board:
Ferron L. Andersen, Zoology
Joseph R. Murdock, Botany
WiLMER W. Tanner, Zoology
Ex officio Members:
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Sciences
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Brigham Young University
Science Bulletin
A COMPARISON OF METEOROLOGIC
MEASUREMENTS FROM IRRIGATED
AND NON-IRRIGATED PLOTS,
PROVO, UTAH, 1970-1972
by
Ferron L. Andersen
Phil D. Wright
and
J. Carl Fox
BIOLOGICAL SERIES — VOLUME XIX, NUMBER 3
APRIL 1974/ISSN 0068-1024
TABLE OF CONTENTS
ABSTRACT 1
INTRODUCTION 1
MATERIALS AND METHODS 3
RESULTS AND DISCUSSION 5
CONCLUSIONS 34
ACKNOWLEDGMENTS 36
LITERATURE CITED 36
This publication is supported in part bv BVU Faculty Research Grant 115-77-724 and in part by Public
Health Service Grant AI 10588.
A COMPARISON OF METEOROLOGIC MEASUREMENTS
FROM IRRIGATED AND NON-IRRIGATED PLOTS,
PROVO, UTAH, 1970-1972
by
Ferron L. Andersen\ Phil D. Wright-, and J. Carl Fox'
ABSTRACT
A comparative study of micrometeorologic
conditions on irrigated and non-irrigated pas-
ture plots was conducted at Provo, Utah, from
1970 to 1972. Daily measurements were taken
of the following: precipitation cither as rain or
snow, new snowfall and total snow depth dur-
ing the winter; relative humidity in a standard
weather shelter; number of hours at maximum
relative humidity; cloud cover each morning;
potential evaporation; total wind 1 m above
ground level; temperature extremes in a standard
weather shelter; and temperatures both on ir-
rigated and non-irrigated plots with sensing de-
vices located 5 cm beneath soil surface under
grass cover, at soil surface under grass cover, and
at soil surface on bare ground. During the
pasture months of May through October, week-
ly soil moisture measurements from irrigated
and non-irrigated plots were gravimetrically de-
termined. All data were entered on columnized
work sheets, kev punched, and subsequentlv as-
similated and tabulated by a FORTRAN IV
program with the IBM .360/65 computer. Graph-
ical representations of all daily measurements
for the three vears, as uell as others depicting
the effect of irrigation and snow cover for se-
lected weeks, were completed.
The total precipitation for 1970, 1971, and
1972 was 491.0, 726.4, and 390.1 mm, respectively,
and the average monthly mean temperatures in
a standard weather shelter for the three years
were 9.2, S.8, and 9.4°C. Other representative
yearly values for 1970, 1971, and 1972, respec-
tively, were total snowfall: 1165.9, 1877.1, and
909.3 mm; average maximum and minimum rela-
tive humidity: 97 and 43, 97 and 46, and 97 and
46 percent; average hours at maximum relative
humidity each day: 7, 8, and 8 hr; average cloud
cover each morning: 3-, 3-, and 3-tenths of cov-
ered sky; total potential evaporation and daily
average for May through October: 983.8 and
5.3, 1030.7 and 5.6, and 1274.2 and 6.9 mm;
average monthl)' soil moisture content for both
irrigated and non-irrigated plots for May through
October: 16.1 and 8.4, 16.3 and 5.8, and 8.1 and
3.8 percent; wind totals and daily averages:
19,315 and 52, 22,691 and 61, and 22,255 and
60 km; average annual temperatures 5 cm be-
neath soil surface on irrigated and non-irrigated
plots for 1971 and 1972: 8.4 and 9.0 and 8.2
and 10.4°C; average annual temperatures at soil
surface under grass cover on irrigated and non-
irrigated plots for 1971 and 1972: 9.7 and 10.6
and 10.9 and 12.8°C; and average annual tem-
peratures at soil surface on bare ground on ir-
rigated and non-irrigated plots for 1971 and
1972: 12.9 and 16.5 and 14.9 and 15.0°C. Tem-
perature extremes on irrigated plots during the
pasture season were moderated considerably
from those recorded on the non-irrigated plots,
and irrigation, when done routinely, provided
adequate soil moisture throughout the entire pas-
ture season. These findings are especially ob-
vious when data other than yearly and monthly
means are examined. Irrigation is undoubtedly
the major factor in providing optimum moisture
and temperature conditions for the development
and survival of all types of pastureland bio-
logical organisms in creating optimum microen-
vironments in otherwise desolate, arid regions.
INTRODUCTION
Micrometeorology, the study of atmospheric of a uniform definition of the zone in question,
conditions in microclimates, entails numerous Reference can be made to both horizontal and
difficulties, not the least of which is the lack vertical measurements, which may vary from
'Department of Zoology, Brigham Young University. Provo. Utah 84602
'Department of Agriculture, Salt I^ike City, Utah 84114
^Veterinary Research Laboratory, Montana State University, Rn/eman, Montana 59715
Bricham Young University Science Bulletin
a few millimeters to many meters in both direc-
tions. Sutton (1964) stated that micrometeorology
deals with atmospheric phenomena 'in the first
few hundred feet above the earth's surface,"
whereas Holmes and Dingle ( 1965 ) stated that
the microclimatic zone consists generally of the
layer of air which is markedly altered by the sur-
face of the earth or other surfaces thereon,
and that the microclimate of a tree, therefore, is
that area extending several meters out from the
branches which acquires unique properties be-
cause of the tree. Whitman and Wolters ( 1965 )
defined microclimate as that zone beneath the
level of a standard weather shelter (about 1.5
m ) . Andersen, Levine, and Boatman ( 1970 )
stated that the microclimate of the free-living
stages of ruminant nematodes is the layer of
space between the ground surface and the top
of the pasture vegetation. Since the project
described herein dealt with measurements on
irrigated and non-irrigated pasture plots, the
definition used is that given by the latter authors.
The practice of irrigation is the single most
important process which permits maximum uti-
lization of agricultural lands in arid regions of
the earth. In the United States, of a total of
2,.322,016,000 acres, 340,998,000 are cultivated,
of which a3,022,000, or approximately 10 per-
cent are irrigated (Israelsen and Hansen, 1967).
In Utah, however, of 52,721,550 total acres,
2,155,186 are cultivated and 1,348,627, or about
63 percent, of that amount are irrigated. In
Utah County, 76 percent of all cropland is ir-
rigated (Utah Agricultural Statistics, 1972).
Thus, we see that irrigation is far more impor-
tant to agriculture in Utah and other western
arid states than in areas where sufficient pre-
cipitation is accumulated through rain or snow
only.
While irrigation is the most important factor
toward creating cultivatable lands in arid re-
gions, indiscriminate usage of available water
often leads to soil erosion, alkalization, or high
water tables, with resultant reduction in plant
growth and productivity. In addition to these un-
desirable effects, irrigation may create suitable
microhabitats for the development and survival
of detrimental biological organisms, many of
which could not exist or at least could not reach
such high population levels on otherwise arid
lands. The creation of desert bridges from one
irrigation field to another may allow harmful
insects to traverse broad expanses of arid re-
gions (Rainy and Hess, 1967), and faulty irri-
gation may fill borrow pits or other depressions
for long periods of time, thereby creating favor-
able breeding grounds for numerous arthropod
or molluscan vectors of disease. Mosquitoes,
for example, are by far the most important of
all arthropods of public health significance,
since several species are vectors of malaria and
encephalitis. In spite of extensive eradication
campaigns, malaria is still the most important
infectious disease in the world; however, arbo-
viral encephalitis is far more significant in the
United States proper. In 1970, onlv sixteen per-
sons acquired malarial infections in the United
States (Center for Disease Control, Malaria
Surveillance, 1971), whereas during the same
year, 110 cases of arboviral encephalitis were re-
ported ( Center for Disease Control, Neurotropic
Viral Diseases Surveillance, 1972). Anopheles
freeborni, the primary vector for malaria in the
western part of the United States, and Citlex
tarsalis, (the "irrigation mosquito"), the primary
vector for western and St. Louis encephalitis,
both breed in water impoundments and seepage
areas, such as those associated with faulty or
inefficient irrigation (Rainy and Hess, 1967).
The breeding habitats of both species generally
decrease each summer with the recession of sur-
face waters, except where irrigation waters main-
tain their breeding sites. Surtees (1970) showed
a 70-fold increase in malaria vectors in an ir-
rigated area in Kenya, and Reeves and Hammon
( 1962) demonstrated that human infections with
encephalitis were definitely associated with work
on irrigated agricultural lands in California.
Snails, which serve as intermediate hosts
for the blood flukes that cause schistosomiasis
(bilharziasis) in humans in many tropical and
subtropical regions, have reproduced extensively
in newly created habitats associated with mas-
sive irrigation projects. The first report by the
stud\- group on schistosomiasis in Africa ( World
Health Organization, 1950) stated that: "The
introduction or development of irrigation
schemes, as well as the change from basin to
perennial irrigation, has always resulted in a
considerable increase in the incidence and inten-
sity of bilharziasis wherever that infection ex-
isted or was introduced by outside laborers. The
severity of the infection may be such as to
cause the abandonment of an irrigation scheme
created at considerable expense."
Irrigation also creates fa\'orable microen-
vironments in pastures for the free-living stages
of manv parasitic nematodes which otherwise
could not exist in overall arid zones. As early
as 1944, Furman pointed out that irrigational
practices in the dry interior Sacramento Valley
regions of California resulted in increased popu-
lations of nematode larvae parasitic to sheep.
Honess and Bergstrom (1966) showed compa-
rable results for populations of nematode larvae
parasitic to cattle in Wyoming. Surveys in cen-
Biological Series, Vol. 19, No. 3 Meteobologic Mej\slirements
tral Utah have shown tliat 68 to 71 percent of
all cattle and 90 to 96 percent of all sheep
raised on irrigated pastures harbor parasitic
helminths of at least one species ( Fox, Ander-
sen, and Hoopes, 1970; and Wright and Ander-
sen, 1972). During the past decade numerous
workers have shown that there is a positive cor-
relation between optimum micrometeorologic
conditions and the development and survival of
many parasites ( See Crofton, 1963; Levine, 1963;
Kates, 1965; Williams and Mayhew, 1967; An-
dersen, Levine, and Boatman, 1970; Williams
and Bilkovich, 1971; Gibson, 1973; and Levine
and Andersen, 1973).
Because of the importance of irrigation to
agriculture and animal husbandry in central
Utah and the demonstrated high prevalence of
parasites in domestic animals, a study was un-
dertaken to measure microenvironmental con-
ditions on irrigated and non-irrigated pasture
plots in that area.
MATERIALS AND METHODS
A 9in X 9m experimental weather station de-
signed to yield measurements from irrigated and
non-irrigated plots was established at the Brig-
ham Young University Animal Science Farm lo-
cated at Provo, Utah (Fig. 1). Coordinates for the
station are 111° 39' 1" W. longitude and 40°
16' 1" N. latitude. Elevation at the site is 4,653
feet (1163 m) above sea level. Meteorologic
measurements from the area were either re-
corded or manually observed each dav for
1970, 1971, and 1972. The plot was fenced with
5-foot ( 150 cm ) chain link fence from the cen-
ter of a 2-acre (.8-hectare) sheep pasture at
the fami. The existing vegetation at the site con-
sisted predominantlv of meadow fescue ( Festuca
elatoir), alfalfa {Medicago sativa), clover {Tri-
folitnn repens), and dandelion (Leontodon
taraxacum). Tlie vegetative cover was mowed
weekly to an approximate height of 3 to 4 in
(7.5 to 10 cm) and the clippings discarded. The
soil was characterized as the Keigley series ( 1
percent slope, dark brown, deep, well-drained.
r? ,^^::S^s^^-»'':F
Fig. 1. Weather station for measurement of meteoro-
logic conditions on irrigated and non-irrigated pas-
ture plots at BYU Animal Science Farm, Provo,
Utah.
Fig.
2. Non-irrigated plot separated by a 5* x 20' metal
frame.
silty clay loam) (Soil Survey of Utah County,
Utah, 1972). The average volumetric moisture
content at field capacity (Vsrd bar) was 2.2 in
(5.5 cm) of water per foot (30 cm) of soil, and
at the permanent wilting point ( 15 bars) was 1.6
in (4.0 cm) per foot (30 cm) of soil (Soil Con-
servation Service, Utah, 1972).
A non-irrigated section near the center of
the plot was established by placing a 5 ft by 20
ft ( 150 by 600 cm ) rectangular galvanized metal
frame into the ground to an approximate depth
of 3 ft 8 in (HO cm), with about 4 in (10 cm)
protruding above the ground (Fig. 2) in order
to keep the water from that part of the plot dur-
ing periods of irrigation. The pasture and the
irrigated portion of the weather station were
usually irrigated every 8 to 10 days during the
warmer part of each pasture season; however,
irrigation could only be performed by personnel
not associated with the research project and the
intervals between irrigations were not always
consistent. During 1972, the timing was further
disrupted until mid-June by road and ditch con-
Bricham Young University Science Bulletin
Fig. 3. Standard non-recording rain gauge.
struction in areas directly adjacent to our experi-
mental plot. In spite of these limitations, irriga-
tion times are indicated on the precipitation
and soil moisture graphs for each of the three
years of the study (Fig. 12, 13, and 14) and
hopefully give some indication of the effects
produced.
Precipitation as rain was measured in a stan-
dard non-recording rain gauge (Fig. 3) in hun-
dredths of an inch. During the winter, the
total snow cover was determined with an ordi-
nary ruler by measuring in inches the total depth
of snow to the soil surface at several representa-
tive sites, and the daily snowfall was likewise
determined by measuring the depth to the pre-
vious day's crust. Precipitation in the daily snow-
fall was calculated as one-tenth that of the
measured new snow depth, since that is the
average reported snow depth to water equivalent
ratio (Mcintosh and Thorn, 1969).
During the six months from 1 May through
31 October, soil moisture content from both ir-
rigated and non-irrigated plots was determined
gravimetrically by taking 6-in (15-cm) core
samples each Monday with a 1-in (2.5-cm) di-
ameter soil auger. The samples were immedi-
ately taken to the laboratory, where they were
weighed, dried at 105°C for four days, and then
reweighed.
Relative humidity (RH) was measured in a
standard U.S. Weather Bureau shelter (Steven-
Fig. 4. Hygrothermograph for daily measurement of
temperature (top graph) and relative humidity
( bottom graph ) recorded in a standard weather
shelter.
son screen) located approximately 5 feet ( 1.5 m)
above ground level with a standard hygrother-
mograph which gave a dual record of both
temperature and relative humidity on the same
curvilinear chart (Fig. 4). The humidity element
consisted of multiple strands of specially treated
hygroscopic hairs which expanded over the 0-
100 percent scale range with 3 percent accuracy
between 20 to SO percent RH and 5 percent
beyond those points. The instniment was cali-
brated periodically by adjusting the recording
arm to 100 percent RH during rainy periods, and
it was checked occasionally with a sling psy-
chrometer or Assman spring-driven psychro-
meter. Tlie number of hours each day when
the relative humidity was at maximum (98-
100 percent) was also noted and recorded.
Potential evaporation over the irrigated plot
was measured by an evaporimeter consisting of a
metal pan with a 250-sq cm evaporating sur-
face resting over a spring-loaded device and re-
cording arm (Fig. 5). Since the clock revolved
Fig. 5. Evaporimeter for daily measurement of poten-
tial evaporation.
Biological Series, Vol. 19, No. 3 Meteorologic Measurements
Fig. 6. Non-recording anemometer for measurement of
daily wind total.
daily, the recorded spiral pattern was difficult
to read once the chart had been removed;
hence, the evaporation pan was filled daily and
the potential evaporation read from the scale
each morning.
Total wind movement at the site was read
daily from a non-recording totalizing anemo-
meter positioned 1 m above the vegetative sur-
face (Fig. 6). Cloud cover for each da\- was esti-
mated visually in tenths of sky covered so as
to cast a shadow from the sun at the time the
instruments were read.
Daily temperature measurements were taken
with standard maximum and minimum ther-
mometers in' the weather shelter (Fig. 7) and
also recorded by the hygrothermo graph at that
Fig. 7. Maximum and minimum thermometers lo-
cated in a standard weather shelter,
location as described above. Temperatures were
also recorded on the irrigated and non-irrigated
plots by three-lead distance thermographs ( Fig.
8) with mercury bulb sensors located 2 in (5
cm) beneath soil surface under 3 to 4 in (7.5
to 10 cm) of grass cover, at soil surface under
^rass cover, and at soil surface on bare ground
(Fig. 9).
All meteorologic observations were taken
daily at approximately 8:00 a.m. M.S.T., and
charts for the recording instruments were
changed each Monday morning. All data were
entered on specially designed worksheets ( Fig.
10 and 11) columnized for a FORTRAN IV pro-
gram, kcvpunched on 80-column IBM cards,
and then transferred to tape for storage and
subsequent assimilation and tabulation of daily,
monthh', and yearU- summaries with the IBM
360/65 computer. All mensural data not already
in the metric system were so transposed by ap-
propriate subroutines. Daily measurements of
most of the moisture and temperature totals or
extremes recorded over the three-vear period
were graphed manually and photographed for
reproduction herein.
RESULTS AND DISCUSSION
Meteorologic measurements of all data col-
lected during the three-year study period as
described above are tabulated as dailv, month-
ly, and yearly summaries. However, since the
daily summaries for any one month require three
computer sheets per month, or 108 total pages,
they are not included in this paper. Copies of
these computer tabulations and the FORTRAN
IV program are available upon request for
the cost of reproduction. Monthly and year-
ly totals and means are included herein as
Tables 1 to 3, 4 to 6, and 7 to 9 for 1970,
1971, and 1972, respectively. Tables 10 and 11
give the monthly precipitation and the average
monthh' mean temperatures, respectively, as re-
corded for these three years at the official
weather station for Provo ( KOVO radio station ) ,
located approximately 6 km south of our station.
Brighaai Young University Science Bulletin
Fig. 8. Three-lead distance thermograph located in a
standard weather shelter.
For additional comparisons herein, Tables 12
and 13 give average monthly values during
1970-72 for potential evaporation and total wind
movement, respectively, when recorded for May
through October at the Utah Lake weather sta-
tion (approximately 25 km northwest of our
station) near Lehi, Utah. Table 14 hsts average
monthly soil temperatures for 1970-72 recorded
Fig. 9. Mercury-filled sensing device located at soil
surface on bare ground.
4 in (10 cm) beneath barren, sandy-loam soil
at Salt Lake City, Utah ( Climatological Data,
Utah. 1970, 1971, and 1972).
Graphical representations for all daily data
measured in the study period (except daily
cloud cover) are shown in Fig. 12 to 50. Fig.
51 depicts the marked impact of irrigation dur-
ing one selected week as reflected in the tem-
peratures recorded with the three-lead distance
thermograph with sensors located at the three
ground sites described above. Fig. 52 and 53
contrast the daily temperatures for two dif-
ferent weeks during winter periods with and
without snow cover on the ground.
The yearly precipitation recorded in this
study for 1970, 1971, and 1972 was 491.0, 726.4,
and 390.1 mm, respectively (Tables 1, 4, and
MONTH
YEAR
METEOROLOGIC DATA
PARASITOLOGY B Y U
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Page 2
TEMPERATURES
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Biological Series, Vol. 19, No. 3 Meteobologic Measurements
j30
PRECIPITATION AND SOIL MOISTURE
PROVO, UTAH. 1970
®
.SOIL MOISTURE - IRRIGATED
•denotes time Of IRRIGATION
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PRECIPITATION AND SOIL MOISTURE
PROVO, UTAH, 1971
®
SO/L MOISTUKE - IKKIGATED
16 I 15 I 15
JANUARY FEBRUARY MARCH
I 15 I
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PRECIPITATION AND SOIL MOISTURE
PROVO. UTAH, 1972
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SO(L MOISTURE - IRKIGiTED
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^DENOTES TIME OF IRRIGATION
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16
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111
tIiiNII. I
15 I 15 I 15 I 15 I 15
AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER ,
_J I I 1 1 1
Bbigham Young UNrvERSiTY Science Bulletin
SNOW COVER
PROVO. UTAH, 1970
®
â– M
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JANUARY FEBRUARY MARCH
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APRIL
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MAY
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MARCH
IS
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IS I IS I IS I IS I IS I 15 I IS I IS
MAY JUNE JULY AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
Biological Sehies, Vol. 19, No. 3 Meteohologic Measurements
RELATIVE HUMIDITY IN WEATHER SHELTER
PROVO, UTAH, 1970
MAX
MIN
nrr^^^^
ii ! . :â– f
— I 1 1 1 1
1 15 I 15 I 15
JANUARY FEBRUARY MARCH APRIL MAY JUNE JULY AUGUST SEPTEMBER CXTOBER NOVEMBER DECEMBER
I I I I I I I I I I I I I
RELATIVE 'HUMIDITY IN WEATHER SHELTER
PROVO, UTAH, 1971
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MAX
MIN
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~i 1 1 1 1 1 1 1 1 1 1 1 1 r
I 15 I <5 I IS I IS I IS I IS I IS I
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I 15 I IS I IS I 15
JANUARY , FEBRUARY^ MARCH ^ APRIL ^ MAY ^ JUNE ^ JULY ^ AUGUST ^SEPTEMBER^ OCTOBER ^ NOVEMBER, DECEMBER ,
RELATIVE HUMIDITY IN WEATHER SHELTER
PROVO, UTAH, 1972
n
(20)
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MAX
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1 I 1 I I
I I I I I I I I 1 1 1
15 I IS I 15 I 15 I IS I 15
JULY AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
1 I I I I I
10
Bricham Young University Science Bulletin
15 I 15 I 15 I 15 I 15
AUGUST SEPTEMBER OCTOBER , NOVEMBER DECEMBER.
DAILY DURATION OF RELATIVE HUMIDITY AT 98-100%
PROVO, UTAH, 197 ^
I 15 I 15 I 15
JANUARY FEBRUARY MARCH
I I 1
15
APRIL
15
JUNE
16
JULY
15 I 15 I 15 I 15 I 15
AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
1 1 1 1 I
DAILY DURATION OF RELATIVE HUMIDITY AT 98-100%
PROVO. UTAH, 1972
I 15 i 15 i 15 I 15 I 15 I 15 I 15 1 15 I 15 I 15 I 15 I 15
JANUARY FEBRUARY MARCH APRIL MAY JUNE JULY AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
I I I I I I I I I I I I I
Biological Series, Vol. 19, No. 3 Meteobologic Measurements
11
POTENTIAL EVAPORATION
PROVO, UTAH, 1970
(24)
-05 ?;
15 I 15 I 15
JANUARY FEBRUARY MARCH
15
APRIL
15
MAY
15
JUNE
15
JULY
IS I 15 I 15 I 15 I IS
AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
POTENTI^^ EVAPORATION
PROVO, UTAH, 1971
®
15
JANUARY
I IS I
FEBRUARY
15
MARCH
15
APRIL
15
MAY
IS
JUNE
15
JULY
IS I 15 1 15 I IS I IS
AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
POTENTIAL EVAPORATION
PROVO, UTAH, 1972
(26)
I IS
JANUARY
15 I
FEBRUARY
IS
MARCH
IS
APRIL
15
MAY
15
JUNE
IS
JULY
IS I 15 I IS I IS I IS
AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
12
Bhigham Young University Science Bulletin
15
1 15 1
15
15 1
15
1 15
15
JANUARY
FEBRUARY
MARCH
APRIL
MAY
JUNE
JULY
15 I 15 I 15 I 15 I 15
AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
I I I I I
;j 120
o 120
T 1 1 1 1 1 1 1 r
15 I 15 I IS I 15 I 15
AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
Biological Series, Vol. 19, No. 3 Meteorolocic Measurements
13
15 I 15 I 15 I 15
JANUARY FEBRUARY MARCH APRIL
I I I
15 I 15
MAY JUNE
I
JULY AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
I I I I I I
TEMPERATi'r^E IN WEATHER SMELTER
PROVO, UTAH, 1971
, .-â– â– -/, N A ,^A,^â– ; ,
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I IS I 15 I IS I IS I 15 I IS I IS I IS I IS I IS I 15 I 15
JANUARY FEBRUARY MARCH APRIL MAY JUNE JULY AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
TEMPERATURE IN WEATHER SHELTER
PROVO. UTAH, 1972
-1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 I 1 I I I I I
I 15 1 15 I IS I 15 I 15 I IS I IS I 15 I 15 I IS I IS I 15
JANUARY FEBRUARY MARCH APRIL MAY , JUNE , JULY AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
14
Bricham Young University- Science Bulletin
30
â– c
COMPARISON OF MAXIMUM AND MINIMUM TEMPERATURES
5 CM DEEP IN SOIL UNDER 10 CM GRASS COVER ON
IRRIGATED AND NON-IRRIGATED PLOTS
PROVO. UTAH. 1971
NON-IRRIGATED, 5 CM BENEATH SOIL SURFACE
IRRIGATED. "
T I r
15 I IS I
JANUARY FEBRUARY
IS
MARCH
IS
APRIL
-r
IS
MAY
1
IS
JUNE
IS
JULY
"T
IS
AUGUST
T 1 r
IS I IS
SEPTEMBER OCTOBER
IS I IS
NOVEMBER DECEMBER
I I
COMPARISON OF MAXIMUM AND MINIMUM
S CM DEEP IN SOIL UNDER 10 CM GRASS
TEMPERATURES NON-IRRIGATED , 5 CM BENEATH SOIL SURFACE "*^
COVER ON WIN — -
IRRIGATED AND
NON- IRRIGATED PLOTS
PROVO. UTAH,
1972
""""^^°' M*N ~
~
(35)
i:V'^Vvn^
_
-
10^
-
.' "^" V. ' ' \>if
■V-' »
f^
IS I IS
JANUARY FEBRUARY
15
MARCH
IS
APRIL
IS
MAY
IS
JUNE
15
JULY
140
122
IS I 15 i IS I IS I 15
AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
Biological Seiiies, Vol. 19, No. 3 Meteobologic Measurements
15
COMPARISON OF DAILY MEAN TEMPERATURES 5 CM DEEP IN
SOIL UNDER 10 CM GRASS COVER ON IRRIGATED AND
NON- IRRIGATED PLOTS
PROVO. UTAH, 1970
°C
20
NON- IRRIGATED
IRRIGATED
(36)
15
JANUARY
IS I
FEBRUARY
15
MARCH
15
APRIL
15
MAY
15
JUNE
IS
JULY
IS I IS I IS I IS I 15
AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
COMPARISON OF DAILY MEAN TEMPERATURES 5 CM DEEP
IN SOIL UNDER 10 CM GRASS COVER ON IRRIGATED AND
NON -IRRIGATED PLOTS
PROVO, UTAH, 1971
NON -IRRIGATED
IRRIGATED
I
IS
JULY
\
IS
AUGUST
IS I IS I
JANUARY FEBRUARY
IS
MARCH
IS
APRIL
15
MAY
15
JUNE
COMPARISON OF DAILY MEAN TEMPERATURES 5 CM DEEP
IN SOIL UNDER 10 CM GRASS COVER ON IRRIGATED AND
NON- IRRIGATED PLOTS
PROVO, UTAH, 1972
NON-IRRIGATED
IRRIGATED
I 1 I I I 1 I I I I I 1 I I I 1 I I I
15 I IS I IS I IS I 15 I IS I IS I IS I IS I IS
MARCH APRIL MAY JUNE JULY AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
I I 1 I 1 I I I I
IS I IS I
JANUARY FEBRUARY
16
Brickam Younc University Science Bulletin
COMPARISON OF MAXIMUM AND MINIMUM TEMPERATURES
AT SOIL SURFACE UNDER 10 CM GRASS COVER ON
IRRIGATED AND NON-IRRIGATED PLOTS.
NON- IRRIGATED, AT SOIL SURFACE
IRRIGATED,"
MAX
MIN
"T
I 15 I IS
JANUARY FEBRUARY
1 1 1 1 1 1 r
15 I IS I 15 I 15
SEPTEMBER OCTOBER NOVEMBER DECEMBER
J I I 1 1
Biological Sehies, Vol. 19, No. 3 Meteoholocic Measurements
17
COMPARISON OF DAILY MEAN TEMPERATURES AT SOIL
SURFACE UNDER 10 CM GRASS ON IRRIGATED AND
NON- IRRIGATED PLOTS
PROVO, UTAH. 1970
NON-IRRIGATEO
IRRIGATED
(42)
1 —
15
MARCH
T 1
I 15
, APRIL
1
15
JUNE
1
15
JULY
~1 —
15
15
15 I IS I
JANUARY FEBRUARY
I I I I
15
MAY
AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
J I I I I I
COMPARISON OF DAILY MEAN TEMPERATURES AT SOIL
SURFACE UNDER 10 CM GRASS ON IRRIGATED AND
NON- IRRIGATED PLOTS
PROVO, UTAH. 1971
JANUARY FEBRUARY
I I L
IS
MARCH
IS
APRIL
15
MAY
IS
JUNE
IS
JULY
IS I 15 I IS I IS I IS
AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
1 I I I I
IS i IS i IS I IS I IS
AUGUST SEPTEMBER OCTOBER NOVEMBER DECEMBER
18
Brigham Young University Science Bulletin
COMPARISON OF MAXIMUM AND MINIMUM
TEMPERATURES AT SOIL SURFACE ON BARE
GROUND ON IRRIGATED AND NON- IRRIGATED
PLOTS
COMPARISON OF MAXIMUM AND MINIMUM
TEMPERATURES AT SOIL SURFACE ON BARE
GROUND ON IRRIGATED AND NON -IRRIGATED
PLOTS
Biological Seiues, Vol. 19, No. 3 METEonoLocic Measurements
19
COMPARISON OF DAILY MEAN TEMPERATURES
ON BARE GROUND ON IRRIGATED AND NON-IRRIGATED PLOTS
PROVO, UTAH, 1970
®
NGN- IRRIGATED
IRRIGATED
1 1 1 T
15 I 15 I
JANUARY FEBRUARY
I L
1
15
MARCH
I 15
APRIL
15
MAY
15
JUNE
15
JULY
SEPTEMBER OCTOBER NOVEMBER DECEMBER
J I I I I
COMPARISON OF DAILY MEAN TEMPERATURES
ON BARE GROUND ON IRRIGATED AND NON-IRRIGATED PLOTS
NON- IRRIGATED
IRRIGATED
COMPARISON OF DAILY MEAN TEMPERATURES
ON BARE GROUND ON IRRIGATED AND NON- IRRIGATED PLOTS
PROVO, UTAH, 1972
NON -IRRIGATED
IRRIGATED
20
Bricham Young Univehsity Science Bulletin
MONDAY TUESDAY WEDNESDAY THURSDAY FRIDAY SATURDAY SUNDAY
I \ i ._1 I I I
7) with the highest monthly amount of 125.7
mm received during March 1971 and the lowest
amount of 2.0 mm recorded during May 1972.
The yearly precipitation totals at the official
Provo weather station (KOVO) were 494.8,
323.5, and 274.8 mm, respectively, for the three
years indicated (Table 10). Whereas the totals
for 1970 and 1972 were quite similar for the
two sites, the total amount recorded for 1971
at our station was more than twice that reported
at KOVO. Even though the rainfall may be ir-
regular in distribution for any one storm, such
a discrepancy for an entire year's total is diffi-
cult to explain. The fact that KOVO radio sta-
tion is located more toward the center of Utah
Valley, whereas our station is to the north and
considerably closer to the surrounding moun-
tains, may have contributed significantly to this
difference. Also, since the main differences re-
corded occurred during the months of March,
October, November, and December of that year,
some of the discrepancy may be attributed
to the fact that different methods were em-
ployed at the two stations to detcnnine the
precipitation in snow. As mentioned above, we
measured snowfall with an ordinary njler and
calculated precipitation therein as one-tenth that
of the depth of each snowfall. Official weather
stations either invert the rain gauge cone over
the new snow cover to the last preceding crust
or collect snowfall directly into the rain gauge
at 1 m height. The collected snow is then melted
and the water content therein measured directly
with the rain gauge ruler. Collection of snow
in the rain gauge cannister at 1 m height is more
subject to errors associated with concurrent
wind movements, whereas reliability of our
method suffers from the fact that the amount
of moisture in snow may vary between l/6th
to l/30th of the total snow depth for any one
storm. Since we had no way of conveniently
melting the snow at our station, we felt it best
to use the approximation as described.
In addition to this measured precipitation,
as stated earlier, the amount of water applied
to the station via flood irrigation was not mea-
sured; however, such water was usually applied
for one to two hours during each irrigation
period. Since our station was located at the end
of an irrigation ditch that was used solely for
that pasture, our plot was never excessively ir-
rigated at any time during the study. When_ cul-
tivated lands are located at the ends of main
irrigation ditches where excess water cannot be
transferred or shut down, or when located ad-
jacent to laterals which run continuously dur-
Biological Sehies, Vol. 19, No. 3 Meteobolocic Measurements
21
20
10
-10
10
1 1 1 1
COMPARISON OF DAILY TEMPERATURES UNDER 222-292 MM SNOW COVER
4-10 JAN 1971, PROVO, UTAH
(52)1
5CM UNDER GRASS COVER
^ y ^
SOIL SURFACE UNDER GRASS COVER
-^ \ â– \
-10
SOIL SURFACE WITHOUT GRASS COVER
SAM
6AM
SAM
SAM
SAM
BAM
SAM
MONDAY TUESDAY WEDNESDAY THURSDAY
I I I I L
FRIDAY
SAM
SATURDAY SUNDAY
J I
SAM
SAM
SAM
SAM
SAM
SAM
SAM
SAM
MONDAY
TUESDAY
WEDNESDAY
THURSDAY
FRIDAY
SATURDAY
SUNDAY
ing the irrigation season, such lands can often
receive far more water than is optimum. Holmes
and Watson ( 1967) stated that farmers in certain
areas of southern Australia customarily use too
much water for irrigation and often apph' \va-
ter crest depths of about 20 cm in order to cover
high spots on lands in that area. As a result, as
much as 50 percent of the water applied through
irrigation must be pumped back into the river.
DeVries and Birch ( 1961 ) compared environ-
mental riieasuremcnts in Australia on three irri-
gation pastures to that of an adjacent dry lot
east of Rochester, Victoria, where the frequency
of irrigation was approximately once every 15
days, and found that the drainage from irrigation
lands in that area varied from 3 to 26 percent
of the irrigational inflow. They assumed an
average of 10 percent of irrigational water sup-
plied would eventually leave as drainage. To
our knowledge, the only comparable data for
irrigational lands in central Utah is that by
Israelsen, et al., who showed that in 1944 the
average efficienc)- of flood irrigation was 40
percent for 11 farms they tested in Utah Count)'.
Presumably the overall efficiency has improved
since then, especially in areas in which sprinkler
irrigation is used, but it is not uncommon to
find pasture lands continually flooded with flows
from drain streams during every day of the sum-
mer, and even into winter months in this area.
The impact of the precipitation pattern and
the accompanying irrigation of pasture plots on
the creation of suitable microenvironments for
biological organisms is more apparent if one ex-
amines the precipitation data of this current
study for June, July, August, and September,
since irrigation in Utah is done routinely only
during that approximate period of the year.
During the three-year study period the total pre-
cipitation for those four months and the monthly
averages over that period were 167.3 and 41.8
mm for 1970; 101.9 and 25.5 mm for 1971; and
53.9 and 13.5 mm for 1972. The long-term pre-
cipitation total and the monthly averages for
those four months (based on all data collected
from 1931 through 1960), as reported from the
official Provo weather station, are 70.2 mm total
and 19.5, 14.2, 21.0, and 15.5 mm for June,
July, August, and September, respectively
(Table 10). The normal values for all other
months of the year average more than 25 mm of
precipitation per month, whereas these four
months all average below 25 mm per month.
Parasitic nematodes, which are prevalent in cen-
22
Brigham Young University Science Bulletin
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29
Table 12.
Total monthly potential evaporation* at Utah Lake station, Lehi, Utah.
1970 1971
1972
Month
mm in mm in
mm
in
May
June
July
August
September
October
Total
187.00
206.00
232.25
220.50
115.00
86.75
7.48
8.24
9.29
8.82
6.20
3.47
164.25
229.00
272.00
226.50
171.00
ND
6.57
9.16
10.88
9.06
6.84
ND
1,047.50
43.50
1,062.75"
42.51""'
221.75
225.25
293.00
218.25
151.00
ND
8.87
9.01
11.72
8.73
6.04
ND
1,109.25"" 44.37°
"" Measured from standard 4-ft ( 120-cm ) diameter evaporating pan
""' 5-month total only
ND =^ Not determined
Table 13. Total monthly wind movement' at Utah Lake station, Lehi, Utah.
Month
May
June
July
August
September
October
Total
1970
1971
1972
km
miles
km
miles
km
miles
3,563
2,227
2,990
1,869
3,373
2,108
2,941
1,838
2,850
1,781
2,963
1,852
2,626
1,641
2,397
1,498
2,624
1,640
2,045
1,278
2,338
1,461
1,877
1,173
3.402
2,126
3,438
2,149
2,547
1,592
2,632
1,645
3,291
2,057
2,566
1,604
17,209
10,755
17,304
10,815
15,950
9,969
" Measured with standard wind-totalizing anemometer located 6 inches (15 cm) above standard 4-ft (120-cm)
diameter evaporating pan
Table 14. Average monthly soil temperatures" at Salt Lake City, Utah.
Month
1970
1971
1972
°C
°F
°C
°F
°C
op
1.3
34.4
1.6
34.8
0.0
32.0
3.0
37.4
2.4
36.3
1.7
35.1
5.1
41.1
4.2
39.6
8.3
46.9
7.2
44.9
9.6
49.3
9.3
48.7
13.4
56.2
13.4
56.1
16.1
61.0
18.4
65.2
19.4
66.9
22.5
72.5
21.9
71.4
26.4
79.6
26.5
79.7
23.3
74.0
26.3
79.4
26.3
79.3
16.3
61.3
17.9
64.2
19.6
67.2
10.3
50.5
9.6
49.3
13.2
55.8
5.7
42.2
3.1
37.5
4.8
40.6
2.2
36.0
-0.5
31.1
LI
33.9
10.7
51.2
11.2
52.1
12.4
54.4
27.8
82.0
33.9
93.0
30.0
86.0
-1.7
29.0
-2.8
27.0
-1.7
29.0
January
February
March
April
May
June
July
August
September
October
November
December
Yearly Mean
Yearly Ma.ximum
Yearly Minimum
° Recorded 10 cm deep in barren, level, sandy-loam soil
tral Utah and which are picked up by animals
grazing on pasture, reportedly require a mini-
mum monthly average of 50 mm precipitation
(Levine, 1963) in order for the free-living
stages to develop and survive. Tims, it appears
that irrigation is the factor which supplies the
additional moisture needed to create optimum
microenvironments for these parasites.
The total snowfall for each of the three
years of this study was 1165.9, 1877.1 and 909.3
mm (Tables 1, 4, 7), and the percentage of
total moisture received as snow during those
years was 23.7, 25.8, and 23.3 percent for 1970,
1971, and 1972, respectively. The number of
da)'S with at least 25 mm of snow cover on the
groimd was 38, 62, and 39 days, respectively.
Snowfall during winter months is important to
this gt>ographic area as the major source of water
to be used in irrigation during the following
summer season. The impact of snowfall on bio-
30
Bricham Young University Science Bulletin
logical organisms in microenvironments comes
about through the moderating effect it has on
temperature extremes. This effect can be read-
ily seen by examining Fig. 52 and 53, which
depict temperature fluctuations at soil surface
with and without snow cover. For those para-
sitic organisms with free-living stages outside
their normal hosts, this moderation may be an
important factor in their extended winter sm-
vival. Andersen, Levine, and Boatman (1970)
showed that free-living infective larvae of cer-
tain nematodes parasitic in sheep could survive
twice as long when placed on plots during win-
ter months than during the warmer periods of
the year, which fact may have been due in
part to the moderating effect of snow cover.
The impact of snow cover on temperature at
ground level under snow cover compared to that
in a standard weather shelter was also demon-
strated by Andersen and Levine ( 1967 ) who
recorded a range during one day in January
1965, at Urbana, Illinois, of and -2.2°C under
10 cm snow cover compared to -6.1 and -17.8°C
in a standard weather shelter. In the study here-
in reported, we found that the temperature on
4 January 1971 at soil surface under 22 cm of
snow (Fig. 52) varied onl\' from -3 to -4°C
(mean of -3.5°C), whereas that in the weather
shelter at the same location ranged from -7.8
to -25.6°C (mean of -16.7°C). Three weeks
later on 25 January 1971, with no snow cover
(Fig. 53), the temperature at soil surface on
bare ground varied from 15 to -3°C ( mean of
6°C), whereas that in the weather shelter ranged
from 12.2 to 0.0°C (mean of 6.1°C).
The extremes in relative humidity measured
in the weather shelter fluctuated markedly
throughout the year in the present study, but
the daily maximum recorded was generally high
even during warm summer periods. The average
maximum and minimum humidity percentages
recorded for the three vears were 97 and 43
percent, 97 and 46 percent, and 97 and 46 per-
cent, respectively (Tables 1, 4, and 7). Percent-
ages less than 98 to 100 percent were recorded
only 18 days during each of the three years.
In a comparable study of meteorologic measure-
ments on pasture lands at Urbana, Illinois, An-
dersen, Levine, and Boatman (1970) recorded
average daily maximum and minimum relative
humidities of 96 and 56 percent for 1965 and
94 and 60 percent for 1966.
Whereas in our current study we measured
relative humidity onlv over the irrigated plot,
deVries and Birch ( 1961 ) compared relative
humidity in a Stevenson screen (approximately
150 cm above soil surface) located over weather
stations in three irrigated pastures in Australia
with that measured over a dry lot area approxi-
mately 6 km away. By rotating hygrothermo-
graphs at 4 to 6 week intervals over a two-year
study in order to eliminate systematic errors,
they found that relative humidities were 5 to
10 percent higher in summer in the irrigated
fields than in the non-irrigated region. The
evaporation of moisture and increased transpira-
tion from the higher vegetation in the irrigated
stations undoubtedly contributed to this elevated
reading and would naturally hold for any geo-
graphical region where such comparison could
be made. Since our non-irrigated plot was con-
fined within the irrigated section of the weather
station, we were unable to make such a measure-
ment. A comparison of relative humidities re-
corded within Stevenson screens on adjacent
wet and dr)' plots would be meaningless as an
adequate reflection of environmental influences
emanating from the surface atmosphere, since as
Halstead ( 1958 ) stated, meteorologic conditions
may differ more from the instrument shelter to
the ground below than might exist between
weather stations located 100 miles apart.
Since the cvaporimeter we used could not
be read during freezing weather, data are given
only for the pasture period of May through
October. The totals and the daily averages of
those periods for the three years were: 983.8
and 5.3 mm, 1030.7 and 5.6 mm, and 1274.2 and
6.9 mm, respectively (Tables 1, 4, and 7). Po-
tential evaporation was generallv highest during
the months of July and August, except in 1971
when the greatest amount was recorded in June.
Information on actual evaporation is pre-
ferred to that of potential evaporation, but as
Holmes and Watson (1967) pointed out, the
measurement of actual I'vaporation is one of
the most difficult of all water budget com-
ponents to obtain. Formulae are available for
calculation of potential evapotranspiration—
evaporation and transpiration— (Hamon, 1961)
for indications of the overall water balance at
specific locations, but they are not used herein,
since our project dealt with basic actual mea-
surements only. Levine ( 1963 ) , however, pointed
out that one should carefully differentiate be-
tween actual and potential evapotranspiration.
The actual evapotranspiration amount would
be low indeed over arid desert soils, but the po-
tential there would be extremely high. This
would be realized under conditions of irriga-
tion such as occurred in our study. Burmaa and
Louden (1968) showed that potential evapo-
transpiration was 20 percent greater for irri-
gated ryegrass-alfalfa pastures in Wyoming high-
lands than that for wheatgrass-alfalfa pastures
but did not propose an explanation for this.
Biological Series, Vol. 19, No. 3 NIeteobologic Measurements
31
Potential evaporation recorded from our sta-
tion compared to that at the Utah Lake Station
near Lehi, Utah (Table 12), was 983.8 and
1047.5 mm, respectively, for May through Oc-
tober 1970; 971.7 and 1,062.75 mm for May
through September 1971; and 1217.8 and
1,109.25 mm for May through September 1972.
These comparisons suggest that the measured
potential evaporation is similar throughout Utah
Valley. However, since the wind totals for those
periods at the Utah Lake Station (Table 13)
were nearly twice that measured at our station,
one would expect correspondingly higher poten-
tial evaporation measurements instead of such
comparable levels as recorded. A partial ex-
planation of this apparent discrepancy might
relate to the fact that we used a recording in-
strument with an 18-cm-diameter evaporating
pan, whereas the Utah Lake Station used a
standard non-recording 120-cm-diameter evapo-
rating pan. A reliable comparison between sta-
tions is further complicated in that the water
level in our instrument was routinely within 1
cm of the surface of the pan, whereas it is not
infrequent for water levels in large standard
non-recording evaporimeters to be 10 to 15 cm
below the surface. Such a difference would in-
fluence greatly the effect of wind movements
upon the resultant evaporation recorded.
Daily cloud cover was only rated visually
each morning and thus was the most sub-
jective of all data taken during this study.
Nevertheless, this showed a general pattern of
approximately 40 to 60 percent average daily
cover during the early and later parts of each
year and 10 to 30 percent during the pasture
season (Tables 1, 4, and 7). Tliese data were
accumulated in order to approximate the de-
gree to which cloud cover might affect condi-
tions at microcnvironments, and in turn affect
such factors as temperature and evaporation at
that site. The amount of solar energy reaching
the soil surface would naturally be the same on
either irrigated or non-irrigated plots, providing
the amount and type of vegetation were simi-
lar. Since vegetation is consistently less on non-
irrigated pastures in such a region as central
Utah, and since vegetation moderates the micro-
climate by providing a barrier between the soil
and immediate atmosphere above (Cabom,
1973), one would generallv expect warmer mi-
crocnvironments on non-irrigated plots than on
irrigated ones. The amount of solar energy avail-
able to wami the soil would depend further on
such factors as the reflective properties of the
soil (albedo), soil moisture, and the amount
of water leaving the soil upward through evapo-
transpiration. Andersen, Levine, and Boatman
(1970) measured solar radiation during June
through November with a recording pyrheli-
ometer over pasture lands at Urbana, Illinois,
in 1965 and 1966 and noted this decreased grad-
ually in the fall, naturally coincident with the de-
crease in daily sunlight hours. The average daily
gram calories per cm- measured at their station
for 1965 and 1966 were highest in July for both
years (848 and 542) and lowest in November
(385 and 188) for all montlis during the periods
indicated. To our knowledge, comparable field
data are not available for other sites near cen-
tral Utah. The immediate effect of cloud cover
upon temperatures can readily be seen by exam-
ining Fig. 51 herein. With the exception of the
impact of .3 mm precipitation on Thursday of
that week indicated, all other jagged lines in
the temperature curves are attributed mainly
to changes in cloud cover. While this effect is
most marked in the temperatures recorded at
soil surface on bare ground, the decrease in tem-
perature during heavv' cloud cover is detectable
on the recordings at soil surface under grass
cover as well.
Soil moisture measurements were taken
weekly during May through October of each
year. Tlie yearly averages of soil moisture con-
tent on a dry-weight basis for those months
for the irrigated and non-irrigated plots re-
spectively were 16.1 and 8.4 percent for 1970,
16.3 and 5.8 percent for 1971, and 8.1 and 3.8
percent for 1972 (Tables 1, 4, and 7). In gen-
eral, the percent soil moisture varied in accord-
ance with times of irrigation or rainy periods
(Fig. 12, 13, and 14), with the difference between
irrigated and non-irrigated plots most apparent
during 1970 and 1971. During 1972, road and
ditch construction adjacent to the weather sta-
tion prevented irrigation of the pasture until
mid-June, at which time the moisture level on
the irrigated plot climbed to 15.0 percent from
the preceding week's measurement of 1.4 per-
cent. Irrigation continued sporadically after that
time, and differences in soil moisture content
between the t^vo plots were not marked dur-
ing August and September. Both levels then
climbed simultaneouslv in October, when a
total of 119.6 mm of rain was recorded.
The results obtained from our soil moisture
measurements appeared more erratic during the
three years than anticipated. Tliis may have
been due in part to our small soil sample size
and to the non-homogeneity of the soil. We
routinely collected only 20 to 30 g of soil, where-
as Israelson et al. (1944) used 200 g samples
from the 11 farms they studied in Utah County.
Obviously, samples of that size cannot be re-
32
Bhicham Young University Science Bulletin
moved repeatedly from small plots such as we
had available.
Of interest was the fact that the soil mois-
ture showed an immediate decline within one to
two days after each irrigation. Tliis is undoubt-
edly due to evapotranspiration from the short
grass cover on pasture lands as well as good
soil permeability. Leonard et al. (1971) found
that irrigation of soils once every 15 days under
a red-pine forest in New York kept the soil
moisture level near that measured shortly after
snow melt in the spring. Crops grown on soils
characteristic of Utah County, however, are
commonly irrigated every 8 to 10 days maximum.
Andersen, Levine, and Boatman (1970)
found that the actual soil moisture 2.5 cm be-
low soil surface on pasture grasses at Illinois
did not reach the permanent wilting point dur-
ing 1965, when 1,069.9 mm of precipitation,
somewhat evenly distributed throughout the
year, were recorded; but that soil moisture did
reach that level 27 consecutive days during 1966
when a total of 944.5 mm was recorded. Dur-
ing the second year, however, a five-week period
in August and September received only a total
of 12.2 mm rain at which time the moisture
deficit occurred. Thus, we see that the distri-
bution of precipitation during summer months
is more influential in determining the micro-
environmental conditions than is the total
amount accumulated during any one month.
In our study, the longest period of time without
measureable precipitation during the summer
months was 22 days during June and July in
1970, 35 days during June and July in 1971, and
23 days during May and June in 1972. Irriga-
tion, if done routinelv on well-managed pas-
tiues, compensates for these longer dry spells
in arid regions such as central Utah and will
effectively keep the soil moisture above levels
detrimental to vegetative cover during the sum-
mer season.
Wind patterns measured at 1 m above the
ground during the year were very erratic but
usually totaled between 1,000 and 2,500 km per
month over the three-year study period. The
totals and the daily averages for each year
were 19,315 and 52 km, 22,691 and 61 km, and
22,2.55 and 60 km, respectively (Tables 1, 4, and
7). The highest monthly total of 2,756 km was
recorded during April 1972 and the lowest
monthly total of 1,183 km during November
1972. Total wind movements during May
through October for our station compared to
that at Utah Lake Station near Lehi, Utah
(Table 13). were 9,737 and 17,209 km for 1970,
11,604 and 17,.304 km for 1971, and 10,716
and 15,950 km for 1972. Since the anemometer
at the Utah Lake station was only one-half the
approximate height above ground level as was
ours, one would expect an even greater differ-
ence in wind totals had both measurements been
recorded at the same level. The reduced wind
at our station, situated on the east bench of
Utah Valley near the mountains, is undoubtedly
due to topological features and the location of
numerous buildings nearby. As discussed above,
since the Utah Lake station had considerably
more wind than was measured at our plots, one
would expect correspondingly more potential
evaporation at that site. The fact that this did
not occur, however, is probably explained in
part by the two dissimilar gauges used.
Andersen, Levine, and Boatman (1970) found
that total wind movement at soil surface on
pasture lands was only 14 percent of that mea-
sured 1 m above ground at the same location.
The impact of this phenomenon on pasture
microenvironments would be that of reduced
evaporation therefrom compared to that actually
measured by standard evaporation pans located
above the microclimatic zone of the pasture
vegetation.
DeVries and Birch ( 1961 ) found that wind
velocities at 2 m were generally greater at sta-
tions located in irrigated pastures in Austraha
than on a non-irrigated plot 6 km distant from
irrigated lands, but they felt that the difference
was due largely to different topography in the
two areas and to the roughness of the irrigated
pastures. They pointed out that the effect of ir-
rigated lands on meteorologic conditions of ad-
jacent drylands would become negligible at a
distance equal to the length covered by the
wind over the irrigated area.
Temperature extremes measured in the stan-
dard weather shelter showed that the annual
averages for the monthly maximum, minimum,
and mean temperatures were 16.5, 1.9, and
9.2°C; 16.3, 1.3, and 8.8°C; and 17.3, 1.5, and
9.4°C for the three years, respectively (Tables
2, 5, and 8 ) . The average monthly means agreed
very closely with the corresponding figures of
9.6, 9.1, and 10.0°C recorded for the same years
at the official KOVO weather station (Table 11).
As discussed above, data from weather shel-
ters are a poor indication of conditions at or
near soil surface level; however, such data do
offer the advantage of having been taken under
similar physical conditions as prescribed by
the U.S. Weather Bureau. Presumably geo-
graphical areas with similar macroclimates
would have similar microclimates if measured
under comparable vegetative cover, topography,
soil texture, and soil moisture. Slight differences
in temperatures measured with the maximum
Biological Sehies, Vol. 19, No. 3 Meteobolocic Milasurements
33
and minimum thermometers within the weather
shelter in this project compared to those re-
corded with the hygrothermograph located at
the same site can be explained in part by the
difference in time lag before response of the
different instruments. Mercury- or alcohol-filled
thermometers have a time lag of approximately
3 minutes, whereas hygrothermographs require
up to 30 minutes (Landsberg, 1941). Because
of this, temperature trends recorded by the
hygrothermograph would tend to level out soon-
er and would generally yield lower maximum
and higher minimum temperatures.
Considerably more pertinent to the present
study is the comparison of temperatures re-
corded at or near the soil surface on the irri-
gated part of the station with those on the non-
irrigated plot. Since all leads for the 3-lead dis-
tance thermographs were not positioned until
April 1970, comparative annual statistics for
that year cannot be given. For 1971, the yearly
averages for maximum, minimum, and mean
temperatures recorded 5 cm beneath soil sur-
face under grass cover were 11.3, 5.4, and 8.4°C
in irrigated plots and 12.1, 5.9, and 9.0°C in
non-irrigated plots (Table 5). Comparable fig-
ures for 1972 were 11.4, 5.0, and 8.2°C and
13.9, 6.8, and 10.4°C, respectively (Table 8).
The yearly averages for maximum, minimum,
and mean temperatures recorded at soil surface
under grass cover were 13.8, 5.6, and 9.7°C in
irrigated plots and 16.5, 4.6, and 10.6°C in non-
irrigated plots for 1971 (Table 6). Comparable
figures for 1972 were 15.8, 5.9, and 10.9°C and
20.7, 4.9, and 12.8°C, respectively (Table 9).
The yearly averages for maximum, minimum,
and mean temperatures recorded at soil surface
on bare ground were 24.6, 1.2, and 12.9°C in
irrigated plots and 31.8, 1.2, and 16.5°C in non-
irrigated plots for 1971 (Table 6). Comparable
figures for 1972 were 30.0, -0.2, and 14.9°C and
32.8, -2.7, and 15.0°C, respectively (Table 9).
The annual maximum and annual minimum ex-
tremes for those years for each site where tem-
peratures were measured were: weather shel-
ter: 36.1 and -2.5.6°C for 1971 and 36.7 and
-25.6°C for 1972; 5 cm beneath soil surface
under grass cover: 29.0 and -4.0 (1971) and
28.0 and -7.0°C (1972) for the irrigated plot
and 30.0 and -5.0 (1971) and .37.0 and -6.0°C
(1972) for the non-irrigated plot; at soil sur-
face under grass cover: 34.0 and -4.0 ( 1971 ) and
41.0 and -6.0°C (1972) for the irrigated plot
and 43.0 and -6.0 (1971) and 54.0 and -8.0°C
(1972) for the non-irrigated plot; and at soil
surface on bare ground: 62.0 and -11.0 (1971)
and 63.0 and -15.0°C (1972) for the irrigated
plot and 72.0 and -12.0 (1971) and 68.0 and
-16.0°C (1972) for the non-irrigated plot.
In the study at Illinois, where comparable
measurements on pasture lands were taken dur-
ing 1966 (Andersen, Levine, and Boatman,
1970), the annual maximum, annual minimum,
and average monthly mean for temperatures re-
corded in the weather shelter were 35.6, -24.4,
and 8.0°C, respectively; beneath soil surface
under 10 cm grass cover: 36.0, -10.0, and 12.9°C;
at soil surface under 10 cm grass cover: 46.0,
-10.0, and 14.1°C; and at soil surface on bare
ground: 64.0, -21.5, and 15.3°C. From this com-
parison, it is evident that the annual extremes
and yearly means are similar at both geographi-
cal locations, especially if temperatures recorded
from the non-irrigated plot are compared with
those on the pasture plot at Illinois with no ir-
rigation. The difference in the yearly means for
each site monitored for the last year reported
for each project ( 1966 for Illinois and 1972 for
Utah) were: 1.4°C in the weather shelter,
2.5°C at 5 cm beneath soil surface under grass
cover, 1.3°C at soil surface under grass cover,
and 0.3°C at soil surface on bare ground.
The only temperatures routinely measured
at or near soil surface in Utah of which we are
aware are those recorded 10 cm beneath barren
soil at Salt Lake Cit\- ( Climatological Data,
Utah, 1970, 1971, 1972). The annual maximum,
annual minimum, and average mean at that site
for 1970 were 27.8, -1.7, and 10.7°C, respective-
ly; 33.9, -2.8, and 11.2°C for 1971; and 30.0,
-1.7, and 12.4°C for 1972 (Table 14). Average
annual means at that station compared to the
average annual mean of the temperature 5 cm
beneath soil surface under grass cover at our
non-irrigated plot for the two years, where com-
parisons could be made, thus differed by 2.2°C
for 1971 and 2.0°C for 1972. If compared, how-
ever, to means tabulated from measurements
made at soil surface under grass cover, the dif-
ference was only 0.6°C for 1971 and 0.4°C for
1972.
DeVrics and Birch (1961) noted that tem-
peratures 5 cm beneath the surface in irrigated
stations in Australia were approximately 10°C
cooler in summer than at their non-irrigated plot.
They attributed this partly to the cooling effect
of evaporation and partly to the shading from
the more dense vegetation on the irrigated sta-
tions. Watts (1973) noted that temperatures 5
cm beneath soil surface on bare ground in En-
gland reached a maximum earlier in the day and
then decreased more quickly at night than did
those temperatures measured at comparable
depths under black polythene or glass cover.
Also, the bare ground had to be irrigated six
34
Brigham Young University Science Bulletin
times as often as the covered plots. Of particu-
lar interest was the fact that he was able to
closely correlate growth measurements in plant
cover with mean daily temperatures at 5 cm
depth. This suggests that this level would be a
good one to monitor for all meteorologic pro-
jects directly or indirectly related to research
on plant or animal productivity.
Leonard et al. (1971) noted that the impact
of irrigation on soil temperatures was influenced
considerably by the temperature of the irrigat-
ing water. In their study in New York with pine
forest cover, temperatures of inugation water
ranging from to 5..5°C above that of the soil
surface brought about a subsequent correspond-
ing increase in the soil temperatures. In our
study, the temperature of the irrigation water
was invariably lower than that of the soil sur-
face. The reflection of this finding upon the soil
temperature profile can readily be seen in Fig.
51, which shows a gradual drop of about 5°C in
the temperature recorded 5 cm beneath soil sur-
face under grass cover, an immediate drop of
about 10°C in the temperature at soil surface
under grass cover, and a corresponding decrease
of about .30°C at soil surface on bare ground.
When the temperature data for this project
are examined on a daily basis ( Fig. .33-51 ) , the
effect of irrigation on these temperatures can
readily be seen. Almost without exception, the
maximum daily temperatures on the irrigated
plots during the pasture season were cooler than
on the non-irrigated plots. This difference was
least demonstrable with the measurements taken
5 cm beneath soil surface under grass cover
(Fig. 33, 34, and 35), next most noticeable
on the soil surface measurements under grass
cover (Figs. .39, 40, and 41) and most notice-
able with the recording of the maximum levels
on the bare soil surface plot ( Fig. 45, 46, and
47). The difference in minimum daily tempera-
tures was not nearly so obvious, however, as
noted from the same figures referred to above.
Because of these similarities in minimum tem-
peratures, the compared means of the daily
temperatures on the irrigated and non-irrigated
plots for each of the three sites and for each
of the three years ( Fig. .36, 42, and 48 for 1970;
Fig. 37, 43, and 49 for 1971; and Fig. 38, 44,
and .50 for 1972) are nearlv the same. Although
mean comparisons may be helpful for general
conclusions, the fact that they automatically
eliminate extremes and ranges makes them dif-
ficult to interpret or use meaningfully. They are
included herein only to show how similar these
means were on the irrigated compared to the
non-irrigated plot. Compared means of the three
different sites or compared means for the three
different years were not graphed, although the
monthly averages of these data can be found
in the annual summaries.
Even though daily observations were made
on the presence or absence of dew or frost each
morning in the present study, such measure-
ments were somewhat subjective, especially dur-
ing summer rainy periods or when snow covered
the ground in the winter. Because of these facts,
and since observations were made only for the
irrigated section of the station, the data are
neither graphed nor recorded herein. However,
the mornings with detectable dew or frost are
noted on the daily computer printouts which
are available if desired. Under most circum-
stances dew was present on the pasture grass
whenever the temperature at soil surface under
grass cover was above freezing and when the
relative humidity measured in the weather shel-
ter reached 98 to 100 percent during the night.
Frost naturally occurred with temperatures in
the microclimate below freezing and when rela-
tive humidity was at maximum.
Andersen, Levine, and Boatman ( 1970 )
found that moisture from dew remained on
pasture grass in Illinois for a daily average of
5.1 to 11.8 hours for months between May and
October, when such measurements were re-
corded in their study. Daily totals were high
even during warm summer days with relatively
infrequent precipitation.
Even though the presence of dew on the
pasture grasses would be an additional source of
moisture for all biological organisms in that
microenvironment, such moisture alone cannot
prevent the ultimate drying of the vegetation
later on during the day. Thus, as Andersen, Le-
vine, and Boatman (1970) pointed out, the alter-
nate h\dration and dehydration of such organ-
isms as nematode larvae mav be more detrimen-
tal to their survival in microhabitats than that of
continual desiccation. Additional studies are
needed on tlie impact of moisture deficits upon
biological organisms in microenvironments.
CONCLUSIONS
Meteorologic measurements obtained during
the three-year study showed a marked impact
of irrigation on the moisture and temperature
profiles of experimental pasture plots. Irrigation
not onlv lowers the soil temperature (due to
evaporation and to the fact that the irrigating
Biological Series, Vol. 19, No. 3 Meteorolocic Measurements
35
water used was invariably cooler than the soil
on the plots ) but also cools the air immediately
above the soil surface. It also brings about a
more dense vegetative cover with a resultant
increase of shade, thereby contributing to in-
creased evaporation and transpiration (evapo-
transpiration ) from the additional surface area
of the cover.
The overall moisture balance in the micro-
climate is thus dependent upon the accumulated
precipitation and irrigation, contrasted with that
lost to the niacroclimate through evapotranspira-
tion. This loss is influenced by the combined
effects of the type and amount of vegetative
cover, wind movements within or immediately
above the grass, moisture content of the air, and
the temperature at or near the soil surface zone.
The temperature of the microclimate is de-
pendent upon the solar energy reaching the sur-
face, the albedo or reflective power of the soil
and vegetative cover, the soil texture, and
amount of soil moisture. It is therefore evident
that irrigation influences directly or indirectly
nearly all factors in the pasture microclimate.
These interact in a complex manner to bring
about favorable environments for increased plant
productivit\- and optimum conditions for de-
velopment and survival of biological organisms
which inhabit that zone.
Irrigation was extremelv sporadic in our
study, but, as mentioned above, the timing of
this was not under our control. A much better
method would have been to use sprinkling irri-
gation, so that not only the timing but also the
amount of water added could have been con-
trolled. At the time the project was started,
however, only flood irrigation was available
and was therefore used throughout the study.
Nevertheless, flood irrigation is considerably
more common than sprinkling in this area, and
irrigation intervals under natural conditions are
also frequently sporadic.
Additional conclusions of this study relate to
each of the meteorologic measurements taken.
The precipitation patterns differed markedly
during each year, which is tvpical of any semi-
arid region, but nevertheless showed the gen-
eral dryness associated with the pasture season
in Utah. The fact that the long-term averages
of precipitation for Provo (Table 10) are less
than 25 mm of rainfall during June, July, Au-
gust, and September illustrates the necessity for
irrigation of all cultivated croplands in this area.
Other forms of precipitation, such as snow or
dew, also have an impact upon the pasture
microenvironment; but with respect to plant
productivity or creation of optimum environ-
ments for living organisms, they are not nearly
as important as rain or irrigation of cultivated
croplands or pastures in Utah. Snow cover
moderates temperatures considerably at ground
level in the winter, and dew adds additional
moisture during the pasture season which could
provide the film of water necessary for vertical
migratory movements on the vegetation by such
organisms as nematode larvae. Even though dew
may represent a helpful addition of moisture
during the pasture season for increased plant
productivity, its impact on living organisms in
the microclimatic zone is not clear, since alter-
nating hydration and dehydration of organisms
in that environment have been shown to be
more detrimental to development of organisms
in that microhabitat than that of sustained des-
iccation. Additional research is needed on the
effect of moisture deficits on organisms inhabit-
ing pasture microenvironments.
Relative humidity measurements were gen-
erally near maximum at least once during each
24-hour period even in warm weather and peri-
ods of infrequent rains. Since the small non-irri-
gated plot was established in the center of the
irrigated part of the station, no measurements
of relative humidity over non-irrigated areas
were obtained. Such comparison would only
be possible if measurements could be taken
from large fields, and then one would have dif-
ficulty locating two separate areas with similar
topography, yet far enough removed where
wind movements would not influence the read-
ings. Also, no measurements were obtained on
relative humidity from the pasture microclimate
per se, since, to our knowledge, the only devices
capable of measuring relative humidity from
such an environment cannot withstand satura-
tion and thus cannot be used in extended field
trials such as this one.
Potential evaporation was recorded only from
the irrigated part of our station, since, as dis-
cussed above, the non-irrigated plot was within
the irrigated area and was too small to permit
a valid measurement from that section only. The
level naturally inclined during the early part of
the pasture season, reached its peak during the
warmest months, and declined gradually there-
after. Temperature and wind were the most
influential factors on this measurement. Bury-
ing the recording part of the evaporimeter into
the ground so that the top of the evaporating
pan would be level with the top of the grass
cover would yield a measurement of potential
evaporation more indicative of that occurring in
the microenvironment. Such an arrangement
could be used with sprinkling, but with the ir-
rigation system we had available, the instrument
36
Bricham Young University Science Bulletin
would likely have been flooded each time ir-
rigation occurred.
The percent of cloud cover each morning was
undoubtedly the most subjective of all data col-
lected but nevertheless indicated trends during
the year. The main drawback of this procedure
was that it did not reflect subsequent changes
during the day. Even though a rough indication
of daily cloud cover was noticeable on the
temperature curve recorded for one week on
bare ground (Fig. 51), all other daily tracings
of that measurement are not available to the
reader. A recording pyrheliometer would be ad-
visable for such studies in the future.
Soil moisture samples gave extremely erratic
results, probably relating in part to the small
sample size taken. However, the compared mois-
ture percentages on the irrigated and non-irri-
gated plots gave a good indication of the im-
portance of irrigation to this geographical re-
gion. Gravimetric measurements for determina-
tion of soil moisture are inferior to those avail-
able with more costly equipment but neverthe-
less give a simplified indication of the moisture
present. Larger or duplicate samples would have
helped considerably, as would have conversion
to volumetric water content values to help
smooth out soil non-homogeneity.
Wind measurements were recorded only at
1 m height and gave only a general indication
of air movements in grass. We were unable to
follow the method used by Andersen, Levine,
and Boatman (1970) with an anemometer
placed at soil surface level as well, since with
flood irrigation the totalizing mechanism of the
anemometer would have been flooded each time
irrigation occurred. Such a measurement could
be included, however, with a comparable pro-
ject done with sprinkling irrigation.
Temperatures recorded in the present pro-
ject were the most complete of all measurements
included herein. Weather shelter temperatures
were. included only for comparison of the macro-
climate with the pasture microclimate and for
standardization of this project with such tem-
peratures recorded in other geographical areas.
Since temperatures in weather shelters are taken
in a consistent manner as described by the U.S.
Weather Bureau, measurements taken of macro-
and microclimates at one site should provide a
fairly accurate indication of the microclimate at
a second comparable niacroclimatic site, pro-
viding similar conditions of soil and vegetative
co\'er existed. The average monthly maximum
temperatures recorded in the weather shelter
were gcneralU' higher and the average monthly
minimum temperatures generally lower than
those recorded on the irrigated plot 5 cm be-
neath soil surface or at soil surface under grass
cover. However, the grand means of the monthly
means for those measurements were quite com-
parable. Measurements on bare irrigated ground
showed greater temperature extremes than those
under grass cover or those recorded from the
weather shelter. Also, those temperatures re-
corded on non-irrigated plots on bare ground
showed extremes greater than those on irrigated
plots. This was due in part to the fact that dry
bare soil on non-irrigated surfaces loses more
heat by outgoing radiation at night than those
where the soil is darker due to the additional
moisture from irrigation. Moist soils also have a
higher heat capacity. Temperatures recorded 5
cm under grass cover were dampened in all ef-
fects and were influenced least by those factors
which contribute to the overall heat and mois-
ture balance of the pasture microenvironment.
The computer programs designed for this
project proved to be very helpful in assembling
and tabulating all data collected. Work is now in
progress to develop additional programs for
computerized construction and plotting of
graphs of all daily observations after they are
once columnized and keypunched on IBM cards.
The successful completion of such programs
will greatly facilitate the complete assimilation
of all meteorologic data measured in future
years.
ACKNOWLEDGMENTS
Appreciation is extended to Mr. Raul Marin,
Mr. Paul Roper, Mr. Spade Whittemore, and
Mrs. Barbara Woolf for technical assistance with
this project.
LITERATURE CITED
Andersen, F. L., and N. D. Levine. 1967. Methods
and problems in microenvironmental measurements.
Illinois Vet. 10:10-17.
Andersen, F.L., N. D. Levine, and P. A. Boatman.
1970. Survival of third-stage Trichostrongi/lus
colubriformis larvae on pasture. J. Parasit., 56:209-
232.
BuRMAN, R. D., AND T. L. LouDON. 1968. Evapo-
Biological Series, Vol. 19, No. 3 Meteorolocic Measurements
37
transpiration and microclimate of irrigated pas-
tures and alfalfa under high altitude conditions.
Transactions of the ASAE. pp. 123-128.
Cabohn, J. M. 1973. Microclimates. Endeavor, 32:30-
33.
Center for Disease Control, Malaria Surveii,-
LANCE. 1970 Annual Report, 1971. U.S. Depart-
ment of Health, Education, and Welfare, Atlanta,
Ceorgia. 25p.
Center for Disease Control, Neurotropic Viral
Diseases Surveillance. 1970 Annual Report, 1972.
U.S. Department of Health, Education, and Wel-
fare, Atlanta, Georgia. 26p.
Climatolocical Data, Utah Annual Summaries.
vol. 72 (no. 13), 1970; vol. 73 (no. 13), 1971;
and vol. 74 (no. 13), 1972.
Crofton, H. D. 1963. Nematode parasite population
in sheep and in pasture. Technical Communication
no. 35. Commonweidth Agricultural Bureau, F;u-n-
ham Royal, Bucks, England. 104p.
DeVries, D. a., and J. W. BmCH. 1961. The modifi-
cation of climate near the ground by irrigation for
pastures on the Riverine Plain. Aus. J. of Agr. Res.
12:260-272.
Fox, J. C, F. L. Andersen, and K. H. Hoopes. 1970.
A survey ,.1 the helminth parasites of cattle and
sheep in Utah Valley. The Great Basin Nat., 30:
131-145.
Kurman, D. p. 1944. Effects of environment upon
the free-living stages of Ostertagia ctrcumcincta
(Stadelmann) Trichostrongylidae: II. Field E.xperi-
ments. Am. J. Vet. Res., 5:147-153.
Gibson, T. E. 1973. Recent advances in the epi-
demiology and control of parasitic gastroenteritis
in sheep. Vet. Rec, 92:469-473.
Halstead, M. a. 1958. Panel discussion, whither
weather, climate and agriculture. Proc. 6th Ann.
Counc. Publ. 576, ( 1958) :60-75.
Hamon, W. R. 1961. Estimating potential evapo-
transpiration. Proc. Am. Soc. Civil Engrs., J. Hy-
draulics Div., 87, HY3:Pt. 1, pp. 107-120.
Holmes, J. W., and C. L. Watson. 1967. The wa-
ter budget of irrigated pasture land near Murray
Bridge, South Australia. Agr. Meteor., 4:177-188.
Holmes, R. M., and A. N. Dingle. 1965. The re-
lationship between the macro- and microclimate.
Agr. Meteor., 2:127-133.
Honess, R. F., and R. C. Bergsthom. 1966. Tri-
chostrongylosis of cattle in Wyoming. Science Mono-
graph 2. Agr. Ex. Sta., Univ. of Wyoming, Lara-
mie, Wyoming.
IsRAELSEN, 6. W., AND V. E. HANSEN. 1967. Irriga-
tion principles and practices. John Wiley and
Sons., Inc., New York. 447 p.
ISRAELSEN, O. W., W. D. Criddle, D. K. Fuhhi-
MAN, AND V. E. Hansen. 1944. Water-applica-
tion efficiencies in irrigation. Agr. E.\p. Sta., Utah
State Agr. Coll., Logan, Utah. Bulletin 311, 55 p.
Kates, K. C. 1965. Ecological aspects of hehninth
transmission in domesticated animals. Am. ZooL,
5:95-130.
Landsberg, H. 1941. Physical climatology. Gray
Printing Company, Inc. Dubois, Pa. 283 p.
Leonard, R. E., A. L. Leaf, J. V. Berclund, and
P. J. Craul. 1971. Annual soil moisture-tem-
perature patterns as influenced by irrigation. Soil
Science, 3:220-227.
Levine, N. D. 1963. Weather, climate, and the bio-
nomics of ruminant nematode larvae. Adv. in Vet.
Sci., 8:215-561.
Levine, N. D., and F. L. Andersen. 1973. Develop-
ment and survival of Trichostrongylus colubri-
formls on pasture. J. Parasit. .59:147-165.
McIntosh, D. H., and a. S. Thom. 1969. Essentials
of meteorology. Wykeham Publications ( London )
LTD. London and Winchester. 239 p.
Rainev, M. B., and a. D. Hess. 1967. Public health
problems related to irrigation, pp. 1070-1081. In
R. M. Hagan, H. R. Haise, and T. W. Edrainster
( eds. ) Irrigation of agricultural lands. American
Society of Agronomy, Madison, Wisconsin.
Reeves, W. C, and W. McD. Hammon. 1962. Epi-
demiology of the arthropod-borne encephalides in
Kern County, California, 1943-1952. Univ. Calif.
Publ. Hlth. vol. 4.
Soil Conservation Service, Utah. Central Utah
County Soil Survey, 1972. 161 p.
Soil Survey of Utah County, Utah. 1972. United
States Department of Agriculture, U.S. Govern-
ment Printing Office, Washington, D.C. 16 Ip.
Subtees, G. 1970. Large-scale irrigation and arbo-
virus epidemiology, Kano Plain, Kenya. I. De-
scription of the area and preliminary studies on
the mosquitoes. J. Med. Ent. 7:509-517.
Sutton, G. 1964. Micrometeorology. Sci. Amer., 208:
62-76, 142.
Utah Agricultural Statistics. 1972. Utah State
Department of Agriculture, Salt Lake City, Utah.
129 p.
Watts, W. R. 1973. Soil temperature and leaf ex-
pansion in Zea mays. Expl. Agric, 9:1-8.
Whitman, W. C, and G. Wolters. 1965. Micro-
climatic gradients in mixed grass prairie, pp. 165-
185. In R. H. Shaw (ed.). Ground level clima-
tology. AAAS, Washington, D.C.
Willl^ms, J. C, AND F. R. Bilkovich. 1971. De-
velopment and survival of infective larvae of the
cattle nematode, Ostertagia ostcrtagi. ]. Parasit.
57:327-338.
Williams, J, C. and R. L, Mayhew. 1967. Survival
of infective larvae of the cattle nematodes, Co-
operia punctata, Trichostrongylus axei, and Oeso-
phagostomum radiatum. Am. J. Vet. Res., 28:629-
640.
World Health Organization. 1950. Joint study
group on bilharziasis in Africa. Rep. 1st. Sess.
Tech. Rep. 17:16.
Wright, P. D., and F. L. Andersen. 1972. Para-
sitic helminths of sheep and cattle in Central Utah.
J. Parasit., 58:959.
Brigham Young University
Science Bulletin
MUS. CCMP. ZOOL
LIBRARY
'"^'ATaxonomic study of the
•^n^t^stern collared lizards,
crotaphytus collaris and
cromphytus insularis
by
Nathan M. Smith
and
Wilmer W. Tanner
BIOLOGICAL SERIES ~ VOLUME XIX, NUMBER 4
APRIL 1974/ISSN 0068-1024
INFORMATION FOR CONTRIBUTORS
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versity Press and the library of Brigham Young
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In the preparation of manuscripts, authors are
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To facilitate review by referees, send to the
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sity, Provo, Utah 84602.
Brigham Young University
Science Bulletin
A TAXONOMIC STUDY OF THE
WESTERN COLLARED LIZARDS,
CROTAPHYTUS COLLARIS AND
CROMPHrrus insularis
by
Nathan M. Smith
and
Wilmer W. Tanner
BIOLOGICAL SERIES — VOLUME XIX, NUMBER 4
APRIL 1974/ISSN 0068-1024
TABLE OF CONTENTS
INTRODUCTION 1
REVIEW OF LITERATURE 2
MATERIALS AND METHODS 3
External Morphology 3
Cranial Morphology 7
RESULTS 8
External Morphology 8
Cranial Morphology 9
DISCUSSION 9
External Morphology 12
Cranial Morphology 19
Population Characteristics and Systematics 20
Phyletie Relationships 24
SUMMARY AND CONCLUSIONS 27
ACKNOWLEDGMENTS 27
APPENDIX I , 28
APPENDIX II 28
LITERATURE CITED 28
A Taxonomic Study of the Western Collared Lizards,
Crotaphytus collaris and Crotaphytus insularis
by
Nathan M. Smith' and Wilmer W. Tanner-
INTRODUCTION
The range of the Western Collared Lizard
Crotaphytus collaris baileiji Stejneger (1890)
as formerly construed included northern Mexico,
the Baja California Peninsula, the Islands Angel
de la Guarda and Tiburon in the Gulf of Cali-
fornia, and parts of the seven western states of
Arizona, California, Colorado, Idaho, Nevada,
New Mexico, and Utah ( Stejneger and Barbour,
1917). Within this vast area the geophysical
features vary considerably and may serve as
population barriers.
Although geophysical variation is broad
throughout the range, niche requirements are
narrow. The collared Uzards require boulders
for basking, lookouts, and shelter, and occupy
mountain slopes, canyons, rocky gullies and
boulder-strewn alluvial fans where vegetation
is sparse (Stebbins, 1966). The many mountain
ranges oriented on north-south axes have pro-
vided favorable conditions for extensive distri-
bution of the collared lizard in the Great Basin,
Arizona-Sonoran Desert, and Baja California
Peninsula. Narrow niche requirements cause
collared lizards to form demes with valleys be-
tween the mountain ranges acting as isolating
barriers. The Imperial Valley, Gran Desierto
of southern California and northwestern Mexico,
is an important potential barrier between popu-
lations of the Great Basin and the Baja Cali-
fornia Peninsula. The island populations in the
Gulf of California were probably members of
the mainland population in late Miocene or
early Phocene (Anderson, 1950), with Isla
Tiburon still part of the mainland as recently as
10,000 years ago (Lawlor, 1971). This possible
geophysical arrangement has been discussed
by Tanner ( 1966 ) to e.xplain the distribution
of the night snakes in and around the Gulf of
California. Portions of the Great Basin, which,
according to Wells and Jorgensen ( 1964 ) were
significantly cooler (and perhaps moister) 8,000
to 40,000 years ago, have only become suitable
habitat for the collared lizard since the Wis-
consinian.
Systematists recognizing the potential isolat-
ing factors within the range originally attributed
to C. c. haileiji have described C. c. auriceps
Fitch and Tanner ( 1951 ) from the Upper Colo-
rado River Basin, and C. c. fusciis Ingram and
Tanner ( 1971 ) from the Chihuahuan Desert.
Also, two insular forms, C. iiistihiris Van Den-
burgh and Slevin ( 1921 ) and C. dickersonae
Schmidt ( 1922 ) , although given species rank,
have long been recognized as closely related to
C. collaris Sav ( Burt, 1928; Fitch and Tanner,
1951; Ingran/and Tanner, 1971). Allen (193.3)
referred to the collared lizards from Tiburon
Island as C. c. dickersonae.
Collecting trips were made to determine the
geographic limits of the several populations
and to gather live specimens for electrophoretic
analysis of blood proteins. As a result, two new
subspecies of Crotapht/tus were named (Smith
and Tanner, 1972). The present paper brings
together the basic techniques and data used
with an analysis of the characteristics of each
population of C. collaris and C. insularis.
In this comparative ta.xonomic study multi-
variate analysis techniques are utilized (Ingram
and Tanner, 1971) to study general external
morphology and comparative cranial morpholo-
gy. Disc gel electrophoresis, which was used
to compare populations of C. c. bailey, C. c.
auriceps, and C. c. bicinctores is not included
in this report because of the limited scope of the
material available, but the data are available
from the authors upon request. An analysis is
made of the westem-collaris complex and its
relationship to C. c. auriceps, C. c. bailey, and
C. c. fiiscus. The Great Plains population, C. c.
collaris, is not included, however; a cursory
analysis indicates that it also may be a hetero-
genous group, and a study of its relationship
is planned.
^Graduate Department of Ijbrary and Information Sciences, Brigham Young Universit>', Provo, Utah 84602.
-Department of Zoology, Brigham Young University, Provo, Utah 84602.
Bricham Young UNrvEHSiTY Science Bulletin
REVIEW OF LITERATURE
An adequate review of the literature up to
1971 which is pertinent to this study was pre-
sented by Ingram and Taruier (1971:1-2) and
will not be repeated here.
In 1971, Ingram and Tanner reaffirmed C.
c. auriceps and defined the following distin-
guishing characteristics of C. c. baileiji: (1)
the body is dark green; ( 2 ) yellow, if present on
the head, does not extend posteriorly beyond
the supraorbital semicircles or ventrally to the
throat; (3) it has fewer fused interorbitals and
a greater number of supralabials and loreal-
lorilabials series than C. c. coUaris. Ingram and
Tanner then defined C. c. fuscus as the popu-
lation found in the Chihuahuan Desert. They
indicated C. c. fuscus most closely resembles
C. c. baileyi and is distinguished from it by a
combination of external morphological char-
acters, the sum of which gives good distinc-
tion. In addition, C. c. fuscus has a brown body
dorsum with no trace of green and the head
is a light to cream color, with no trace of yellow.
Smith and Tanner (1972) designated the
Great Basin collared lizard ;is C. c. bicinctores.
It can be distinguished from C. c. baileiji and
other members of the coUaris complex by ( 1 )
fewer scales and fewer spots in the dorsal sep-
aration of the first collar, (2) greater number
of enlarged median internasals, (3) second col-
lar not extending onto the forearm, (4) greater
number of scales from interparietal eye to the
anterior edge of first collar, and (5) more ex-
tensive black pigmentation of the ventral groin.
C. c. bicinctores is easily distinguished from
the Baja California population (C. i. vestigium)
by its collars and dorsal pattern. It is distin-
guished from C. c. dickersonae by its brown
body and dorsal pattern (Fig. 1).
In addition. Smith and Tanner (1972) indi-
cated a close relationship for the populations
from Angel de la Guarda Island, Gulf of Cali-
fornia, Mexico, and the Baja California Penin-
sula. The Angel Island population was desig-
nated C. /. insularis, and the population inhab-
iting the area north and west of Palm Springs,
Riverside County, California, and south from
there along the eastern slopes of the mountains
into Baja California Sur, Mexico, was desig-
nated as C. i. vestigium. Characteristics of C
i. vestigium which distinguish it from C. i.
insularis are fewer subdigital toe lamalae on the
second and fourth toes, a larger second collar/
svl index, and a smaller dorsal separation of
both the first and second collars.
Axtell (1972), after indicating intergradation
between C. c. bicinctores and C. c. baileyi,
chose to designate bicinctores as C. insularis
bicinctores. Axtell's choice of designation is
predicated on the basis his evidence indicates
little genetic introgression. He lists similarities
for bicinctores and c. insularis as a complete
dark ventral collar, similar gular markings, com-
pressed tail, extensive dark abdomino-groin pig-
mentation, small dorsal granules, and no green-
ish coloration.
A brief ta.xonomic history of C. c. baileyi
and the insular collared lizard populations from
Angel de la Guarda and Tiburon islands are
listed with their synonymies as follows:
Crotaphytus coUaris baileyi Stejneger
Crotaphytus baileyi Stejneger, 1890, N. Amer.
Fauna, 3:103 (Type locality: Painted Des-
ert, Desert of the Little Colorado R., Ari-
zona; U.S. Nat. Mus.).
Crotaphijtus coUaris baileyi: Stone and Rehn,
1903, Proc. Acad. Nat. Sci. Phila., 55:30.
Crotaphytus coUaris dickersonae Schmidt
Crotaphytus dickersonae Schmidt, 1922, Bull.
Amer. Mus. Nat. Hist., 46:638 (Type lo-
cality: Tiburon Island, Sonora, Mexico; U.S.
Nat. Mus.); Smith and Taylor, 1950, Bull.
U.S. Nat. Mus., 199:93.
Crotaphytus coUaris dickersonae: Allen, 1933,
Occ. Pap. Mus. Zool. Univ. Mich., 259:7;
Smith and Tanner, 1972, Great Basin Nat.,
32:26.
Crotaphytus coUaris baileyi: Stone and Rehn,
1903, Proc. Acad. Nat. Sci. Phila., 55:30.
Crotaphytus coUaris auriceps Fitch and Tanner
Crotaphytus coUaris auriceps Fitch and Tanner,
195i, Trans. Kans. Acad. Sci., 54(4) :553
(Type locality: three and one-half miles
north-northeast of Dewey Bridge, Grand
Co., Utah; Kans. Mus. Nat. Hist).
Crotaphytus coUaris baileyi: Stone and Rehn,
1903, Proc. Acad. Nat. Sci. Phila., 55:30.
Crotaphytus coUaris fuscus Ingram and Tanner
Crotaphytus coUaris fuscus Ingram and Tanner,
1971, Brigham Young Univ. Sci. Bull. Biol.
Scr., 13(2):23-24 (Type locality: sis and
one-half miles north and one and one-half
miles west of Chihuahua City, Chihauliua,
Mexico; Brigham Young Univ. Herpet.
Mus.).
Biological Series, Vol. 19, No. 4 Western Collared Lizards
Crotapht/tus coUaris haileiji: Stone and Rehn,
1903, Proc. Acad. Nat. Sci. Phila., 55:30.
Crotaphi/tus coUaris hicinctores
Smith and Tanner
Crotaphi/tus coUaris hicinctores Smith and Tan-
ner," 1972, Great Basin Nat., 32:27-29 (Type
locality: Mercury Pass, Nevada Test Site,
Nye Co., Nevada; Brigham Young Univ.
Hcrpet. Mus. ).
Crotaphtjtus coUaris haileiji: Stone and Rehn,
1903, Proc. Acad. Nat. Sci. Phila., 55:30.
Crotaphytus insiilaris hicinctores: Axtell, 1972,
Copeia, No. 4:721.
Crotaphytus imukiris insularis
Van Denburgh and Slevin
Crotapht/tus insularis Van Denburgh and Slevin,
1921, Proc. Calif. Acad. Sci., ser. 4, vol.
11:96 (Type locality: Angel de la Guarda
Island, Baja California, Mexico; Calif. Acad.
Sci.).
Crotaphytus insularis insularis: Smith and Tan-
ner, 1972, Great Basin Nat., 32:27.
Crotaphytus coUaris haileyi: Stone and Rehn,
1903, Proc. Acad. Nat. Sci. Phila., 55:30.
Crotaphytus insularis vestigium
Smith and Tanner
Crotaphytus insularis vestigium Smith and Tan-
ner, 1972, Great Basin Nat., 32:27 (Type
locality: Guadalupe Canyon, Ju;u-ez Moun-
tains, Baja California, Mexico; Brigham
Young Univ. Herpet. Mus.).
Crotapht/tus colUiris haileyi: Stone and Rehn,
1903, Proc. Acad. Nat. Sci. Phila., 55:30.
MATERIALS AND METHODS
External Morphology
Selection and Gathering of Material
Materials examined are from four principle
sources: (1) the collection at the Brigham
Young University Herpetological Museum; (2)
materials examined at the California Academy
of Sciences, Los Angeles County Museum, San
Diego Society of Natural History, California
State College at Long Beach, University of
California at Berkeley, and the University of
Arizona; (3) materials loaned from the insti-
tutions listed in the acknowledgments; and (4)
live specimens collected and color photographed
in the field with 35mm Agfachrome.
Several collecting trips, supported by an
NDEA Educational Allowance, were made be-
tween 1969-1971. The first trip, in June 1969,
was an extensive one, co\'ering two weeks and
4,000 miles. During this period, specimens were
collected from southwestern Colorado, western
New Mexico to the Mexican border, most of
Arizona, southeastern Cahfomia, and south-
western Utah. During the last three weeks in
May 1970, an extensive collecting trip was made
to southern Nevada, southern California to the
Mexican border, and most of Arizona. Speci-
mens were taken from the Oatman-Kingman,
Arizona, area, which is near an area of con-
firmed intergradation for C. c. hicinctores and
C. c. fca/feyi (Axtell, 1972). In July 1970, speci-
mens were collected through central and south-
eastern Utah. Specimens were taken from both
the east and west sides of the Colorado River
in the area of Moab, Utah.
The fourth and final collecting trip was to
Tiburon Island, Gulf of California, Mexico, dur-
ing the first week in April 1971. A number of
collared lizards (14) were caught and photo-
graphed on the island and three were caught
on the mainland opposite the island. All speci-
mens collected during these four trips are de-
posited in the Brigham Young University Her-
petological Museum. A series from the Nevada
Test Site (Research Grant Number AT(ll-l)
1496, between the U.S. Atomic Energy Commis-
sion and Brigham Young University) and from
Pyramid Lake, Nevada, were obtained by the
junior author. Also, two trips were made by
him to the Palm Springs area, and specimens
were taken in Chino Canyon and east of
Thousand Palms, Riverside County, California.
Following the procedure used by Ingram and
Tanner (1971), collared lizards with a snout-
vent length less than 80 mm or that were dam-
aged were not included in the computer analy-
sis portion of this study.
The collared lizards examined in this study
are as follows:
ARIZONA: Apache Co., BYU 497; LACM
16895; UIMNH 7524; USNM 38056. Coconino
Co., CU 30081; SDSNH 12772-76; UIMNH
62445, 74789-92, 6543; USNM 45025-26, 60111,
60113, 60115, 60117-20, 86942; UU 217, 3006-08.
Gila Co., UIMNH 74797-98. Graham Co.,
UIMNH 82348-50. Mohave Co., BYU 32104;
Brigham Young University Science Bulletin
Biological Series, Vol. 19, No. 4 Western Collared Lizards
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Brigham Young UNrvERsm- Science Bulletin
UIMNH 62446, 74785; USNM 86941, 115675-76;
SDSNH 5291, 17302, 31769, 31937, LMK 31768.
Navajo Co., UU 1603. Pima Co., CAS 81420;
LACM 3983, 5899; UAZ 636, 666, 669-670, 672,
1509, 1511, 19986, 28625, 34364, 35539, 35715;
Santa Cruz Co., UAZ 11241; Yavapai Co., BYU
2423; USNM 11860, 15689-90, 15992; UIMNH
74776. Yuma Co., BYU 2437, 32097-98, 32100-03;
CAS-SU 15873, 18299; LACM 16892-93;
SDSNH 16731, 17602, 33301; UAZ 623-27,
643-44, 680-81, 35673, 35676, 35714; UIMNH
74793, 74847.
CALIFORNIA: Imperial Co., BYU 33.321;
CAS-SU 13843; LACM 16898, 37783; SDSNH
11791, 12244-45, 1.3908-09, 29697-98, 38310. Inyo
Co., BYU 409.34-35; CAS 22153-54, 65216,
65221-22, 65224, 65481-82, 65515-17, 65581,
65585, 65636, 65662, 65681, 84241, 89621, 89626-
30, 89633-34; CAS-SU 22072-73; LACM 26824,
36667, 36670; SDSNH 15878-79. Kern Co., BYU
31948; CAS 111006. Los Angeles Co., CSCLB
2747-51; LACM 16870, 2681L 26814, 63187-90;
SDSNH 2834-36, 19481-82. Mono Co., CAS
84962. Riverside Co., BYU 2422, 2429-30, 2432,
2435, 2438, 31983, 32099; CAS-SU 22711; CSCLB
2744, RDK 600430-1, RDK 600611-1, RBL
600327-2, JAN 700510-2, VAP 660513-4; LACM
16873, 16875-76, 16885-89, 26821-22, 52889-90;
SDSNH 20698-99, 39751; UCMVZ 1, 210-11, 231,
243, 74609-11, 76737, 41089, 51850-51, 71723. San
Bernardino Co., BYU 3.3.320; CAS 5245, 36268,
42185, 42785-91; CSCLB 2720-21, 2735-36,
2738-39, RBL 600529-5, RCS 700412-12;
LACM 63179-80, 63182-83, 63188; SDSNH 2490.
5874, 11087, LMK 20888, 29664. 38703, 39874;
UCMVZ 5.368, 316.34, 52486. San Diego Co..
BYU 3994; CAS 62875, 64368; CAS-SU 7930;
SDSNH 1687, 4701, 11088, 11958, 12551, 13250,
18591-92, 22.327.
COLORADO: Los Animas Co., CU 1292,
2939. Mesa Co., BYU 1134. San Miguel Co.,
CU 4448, 4451, 4453, 4456, 4458.
IDAHO: Ada Co., BYU 507; CAS 64173.
Butte Co., BYU 30772. Owyhee Co., BYU 2834;
CAS 55247-48, 55253, 64163-64; NNC 7, 28;
SDSNH 1444-45.
KANSAS: Montgomery Co., BYU 22167.
Wilson Co., KU 41, 48-50, 54.
MEXICO: Isla Angel de la Guarda, CAS
49151, 50873-76, 50878-79, 86754-55, 86783-84;
CAS-SU 21948, 22712; LACM 4001-02, 9854-55;
SDSNH 46001; UCMVZ 96635-37. Baja Califor-
nia, CAS 11545-46, 14000-01, 17048, 18822,
19124-26; CSCLB LWR 670619-1, LWR670619-
2, KAW 700324-6; CU 45855-61; LACM 16994,
16996-99, 63167-73, 63175-78; SDSNH 17052,
19788-92, 24392, 30107-11, 37815, 41612, 52950-
51, 52999; UCMVZ 9590, 51140, 73568. Chihua-
hua, BYU 1.3383-86, 1.3410-11, 13736, 14211,
15184, 16969-76, 17010, 17014; UCMVZ 70704.
Sonora, BYU 2425, 39993, .39995, 409.30-31; CAS
12768-70, 12770, 12772-77, 104467; CU 44201-
03; CSCLB 2730, 2752-55, 2757, 2759-61, 2763-
64, 29.59; LACM 52882, 52886, 88798; SDSNH
CRM117, .35917-19, 43254, 44063; UAZ 694,
697-710, 1513-14, 4634, 9625-26, 10248, 10597,
12114, 16578, 20144, 20255, 28072, 28662, 30226,
31391, 31482, .3.3781; UCMVZ 10163; USNM
26W. Isla del Tiburon, BYU 2426, 2433, 2443,
3163-65, 3167-69, 3172; CAS 14002-12, 5.3263-64;
SDSNH 46003-06, 50665. Zacatecas, CAS 95961.
NEVADA: Clark Co., BYU 461, 40932-33;
CAS-SU 20083-85. Churchill Co., LACM 268.39-
43. Lincoln Co., CAS 37025-.36, .37029-31, 370.33,
37035-36, 37038-39, 37041-42, 37044, 37046,
37058-59. Lyon Co., CAS 20576-80, 20583-85,
22660, 22662-64, 22690, 22705-06; LACM 26838.
Nye Co., BYU 2418, 2424, 17275, 18815-16,
22189-96, 23629-30, 2.3882-83, 30088, 30587; CAS
37514, 37693-95, 37697, 37699-710, 37716, .37721;
CAS-SU 20086-9.3, 20096. Washoe Co., BYU
2401, 2442, 3115; CAS 21487, 21489-90, 21492-94,
21496, 21498-500, 21.505-06; SDSNH .38311,
38676-77.
NEW MEXICO: Chaves Co., LACM 3974-
75. Dona Ana Co., USNM 22268, 25423. Lincoln
Co., LACM 16990. Luna Co., BYU 31940, 31944,
.32121. Quay Co., USNM 44940. Sierra Co.,
LACM 16992.
OKLAHOMA: Carter Co., BYU 500, 1574.
OREGON: Malheur Co., UAZ 21124.
TEXAS: El Paso Co., USNM .59.351; UTEP
.52, 55, 57. Garza Co., CU .32277. Jeff Davis Co.,
UAZ .35145. Randall Co., CU 1.3554-56. Roberts
Co., USNM 32866. Stcphans Co., BYU 13117.
UTAH: Beaver Co., BYU 12715. Emery
Co., BYU 16496, 16774, 20089-90. Garfield Co.,
BYU 11740-41, 12699-701, 31882-83; UU 3329.
Grand Co., BYU 2421, 24,34, 3114, 12854-55,
Kane Co., BYU 2732, 11331, 11.384, 11.386b,
14660, 14894-96, 14898-902, 18921, 21259-60,
21262-63, 40936-.39; UU 1449. Millard Co., BYU
445, 447-48, 8753, 8755, 8883, 12946, 21000-02,
21703; UU 2802. San Juan Co., BYU 2419,
18.335, 31945-46, 31951, 31981, 32088. Tooele
Co., BYU 4.305-06, 14818, 14820-21. Utah Co.,
BYU 552, 1455, 16.30-31, 2844, 3116, 1.3041,
14689. 16602, 22129, .30382; UU 20.38. Washing-
ton Co., BYU 446, 8754, 11.324, 11.386a, 12190,
12875.
SPECIMENS SEEN-NOT USED IN THE
COMPUTER ANALYSIS: CAS 14002-03, 21608,
39114-15, 64966; CAS-SU 19125; LACM 8798-
99, 16993, 36666, 52885-86, 63184; SDSNH
Biological Series, Vol. 19, No. 4 Westebn Collared Lizabds
40074, 40139; UIMNH 74801; UU 122, 124,
33597.
Statistical Methods
After an initial examination of the museum
specimens available at Brigham Young Univer-
sity and a review of the literature with its de-
scriptions of the collared lizards from Tiburon
and Angel de la Guardia Islands, it was hy-
pothesized that there are eight distinct collared
lizard populations in the Great Basin, the Baja
California Peninsula, Tiburon and Angel islands,
the Upper Colorado River Basin, Central Ari-
zona, Chihuahua, Mexico, and the Great Plains.
The methods of analyses employed in this
study, with minor exceptions, are those of In-
gram and Tanner (1971:4). In step 2, we were
limited to 17 and 13 specimens, respectively,
for Angel and Tiburon islands which made a
total sample of 150 individuals. In step 4, eight
clusters were fonned. Also we omitted step 5
and the color and pattern analysis.
Tlie terminologv for taxonomic characters is
that used by Ingram and Tanner (1971:5).
The characters used in the evaluation, with
their numbers, are as follows: (1) tail length/
hind-leg length; (2) tail length/snout-vent
length; (3) snout- vent length; (4) intemasal
scales; (5) number of fused interorbital scales;
(6) fronto-parietal scales; (7) scales from the
union of the posterior canthal and subocular to
the supralabial; (8) supralabial scales; (9) in-
fralabial contacts postmental; (10) gular scale
rows; (11) number of enlarged internasals; (12)
scales from rostral to interparietal; (13) dorsal
scales from interparietal to anterior edge of first
collar; (14) dorsal scales from the anterior edge
of first collar to posterior edge of second coUar;
(15) total dorsal scales; (16) total ventral
scales; (17) dorsal separation of the first collar;
(18) dorsal separation of the second collar;
(19) number of spots within the dorsal separa-
tion of first collar; (20) second collar length/
snout-vent length; (21) subdigital lamellae of
right hind foot, second toe; (22) fourth toe sub-
digital lamellae; (23) fifth toe subdigital lamel-
lae; (24) femoral pores, right side; (25) second
collar extends onto the upper ami; (26) dorsal
light elongate spots present; (27) the pregroin
brown black; (28) the coloration of the pre-
groin fonns spot on the thigh; (29) dorsal
ground color is blue green; (30) small spots
on side of head; (31) large spots on side of
head; and (32) reticulation on side of head
forms patternal bars. Characters 25-32 are
binary (0 ^= characteristic is absent, 1 ^ it is
present ) .
Cranial Morphology
Stejneger (1890:103) distinguished C. baileyi
from C. coUaris stating that in the former the
"head [is] narrower, and the snout [is] longer."
In 1921, \'an Denburgh and Slevin established
C. insularis on the basis of a narrower head and
longer snout than baileyi. Unfortunately, neither
Stejneger nor Van Denburgh and Slevin listed
the data upon which these statements were
based. If their data contained measurements
from juveniles, then it is possible these char-
acters are not significant. To test this possi-
bility, a section on cranial morphology is in-
cluded in this study.
Selection of Material
A total of 126 specimens was selected from
the seven areas used: the Great Basin group
(N^20) from western Arizona, eastern Cali-
fornia, Idaho, Ne\ada, and western Utah; the
Baja group (N=20) from Baja and southern
California; C. /. insularis (N=16) from Angel
Island; C. c. dickersonae (N=20) from Tiburon
Island; C. c. auriceps (N = 10) from the Upper
Colorado River Basin; C. c. baileyi (N = 20)
from central Arizona; and C. c. fuscus (N = 20)
from Chihuahua, Mexico. Only specimens known
to represent these populations were used and all
specimens had a snout-vent length greater than
80 mm. Approximately equal numbers of males
and females were included in each sample.
The following collared lizards were exam-
ined in this portion of the study:
ARIZONA: Coconino Co., BYU 32116;
LACM 28895; UIMNH 6453, 7525, 74786. Mo-
have Co., UIMNH 7524, 74781, 74787, 82349.
Yavapai Co., UIMNH 74767-74, 74776-77. 82354.
CALIFORNIA: Inyo Co., LACM 36666,
36670. Los Angeles Co., LACM 63187. Riverside
Co., LACM 16873, 16875. San Bernardino Co.,
LACM 63179.
MEXICO: Isla Angel de la Guarda, CAS-SU
21948, 22712; CAS 50873-76, 50878-79, 86755,
86783-84; LACM 4001-02, 9854-55; SDSNH
46001. Baja California, BYU 23.337; CU 45855-58,
45860-65; LACM 16993-94, 16996-99, 63171. Chi-
huahua, BYU 13736, 14211, 15184, 15186-87,
16989, 16971-77, 17010, 17014; USNM 2725,
14242, 14307, 14307a. Isla del Tiburon, BYU
3163, 3167-68; CAS 14003-07, 14009, 14011-12,
53263-64; CU 26679-80, 35168; SDSNH 44063,
46005-06, 50665.
NEVADA: Clark Co., BYU 23629, 23883.
Churchill Co., LACM 26843. Lincoln Co., CAS
37033, 37035. Lvon Co., CAS-SU 20576, 20585,
22690, 22705; LACM 26838. Washoe Co., CAS
21499.
8
Brigham Young University Science Bulletin
TEXAS: El Paso Co, UU 493.
UTAH: Garfield Co, BYU 12700-01. Grand
Co., BYU 551, 1625-26, 11737, 12854-55, 31949-
50, 31981, 32111. Millard Co., BYU 21000-02.
Characters Used
A total of 16 characters were analyzed from
six measurements taken from the intact, un-
skinned head. All measm^ements were made to
the nearest 0.1 mm, using a Vernier Caliper. The
measurements were as follows: (1) total skull
length, distance from posterior of quadrate to
anterior of rostral; (2) eve to snout, distance
from the anterior corner of the e^e to anterior
of rostral; (3) head width at widest point, a
point posterior to the eye, in the parietal region;
(4) head width at eyes, distance from jaw to
jaw at a point medial to each eye; (5) head
width at nostrils, distance from each jaw at a
point medial to each nostril; and (6) head depth
at eves, depth of head from a point medial to
supraoculars.
The six measurements made for each skull
were modified to form the following 16 char-
acters which are listed with their character num-
ber: (1) total skull length; (2) eye-to-snout
length; (3) head width at widest point; (4) head
width at eyes; (5) eye-to-snout length/total skull
length; (6) head width at \videst point/total skull
length; (7) head width at eves/total skoill length;
(8) head width at nostrils/total skull length; (9)
head depth at eyes/total skull length; (10) head
width at widest point/eye-to-snout length; (11)
head width at eyes/ eye-to-snout length; (12)
head width at nostrils /eve-to-snout length; (13)
head depth at eyes/eye-to-snout length; (14)
head depth at eyes/head width at widest point;
(15) head depth at eyes/head width at eyes;
and (16) head depth at eyes/head width at
nostrils.
Statistical Analysis
Dixon's BMD07M Computer Program (1968),
the same stepwise discriminant analysis used in
the external moqDhology portion of this study,
was used at this point because the statistical
verification of the seven populations was pre-
viously demonstrated.
The discriminant program calculates the
mean, standard de\iation, and the within-group
correlation coefficient for each character. The
U- and F- statistics which test the null hypothe-
sis of no difference among populations are
listed at each step. The number of correctly
identified specimens is displayed at each step
and the a posteriori probability for an individ-
ual's membership in each of the seven groups is
given.
RESULTS
External Morphology
Cluster Analysis
Cluster analysis was used to form eight
groups of closest morphological similarity (Fig.
2). To test for the independence of the eight
groups formed by clustering from the eight
groups postulated by geographical locality, a
contingency table was formed (Table 1). The
null hypothesis was tested by chi-square with
forty-nine degrees of freedom. The test statistic
is significant at the 0.001 level.
m
X=(l-
,49) =^==1
i=l j = l
(0„-Ei-)VEij
X=(l-oc,49) = 485.7
X=( 0.999,49) = 85.4
Therefore, X^ (l-oc,49) > X= (0.999,49) and
the null hypothesis is rejected.
Specimens from the various locations were
labeled as follows: the Great Basin region of
Idaho, western Utah, parts of western Arizona,
Nevada, and California south and west to the
eastern edge of the Coachella Valley, Riverside
County— the Great Basin group (C. c. bicinc-
tores); Baja and southern Cahfomia, west of
Coachella Valley— the Baja group (C. /. vestigi-
um); Angel Island-C. /. insularis; Tiburon
Islandâ €” C. c. dickersonae; Upper Colorado Riv-
er— C. c. duriceps; Central Arizona— C. c. baileyi;
Chihuahuan Desert— C. c. juscus; and other
specimens from Colorado, New Mexico, Texas,
Oklahoma— C. c. collaris.
Discriminant Analysis
Discriminant analysis, which predicts mem-
bership within a group on the basis of a set of
continuously scaled attributes for the individual
(Cocley and Lohnes, 1971), was used to define
more accurately the relationships seen after
cluster analysis. The BMD07M Computer Pro-
gram of Dixon (1968) was used, with the out-
put consisting of (1) character mean and stan-
dard deviation for each group (Table 2); (2)
the coefficient of the discriminant function for
each group (Table 3); (3) the F-statistic to
show the difference between each group (Table
Biological Series, Vol. 19, No. 4 Western Collared Lizards
4); (4) the number of case.s classified in each
group after each new character enters the
analysis (Table 5); (5) the U-statistic which
tests the difference between groups (Table 6);
and (6) the probability of each specimen's
membership within a group (Table 7).
The output of the BMD07M lists both the
U-statistic and appro.ximate F-statistic. Each
tests the null hypothesis of no difference be-
tween groups, the U-statistic is 0.00006 with
24,6,164 degrees of freedom. The approximate
F-statistic was used because of the availability
of tables. The appro.ximate F-value is 24.90764
with 144,831.93 degrees of freedom. The tabuhu-
F-value is F(. 999,120,120) = 1.76 (Ostle, 1963).
Therefore, F(l-oc, 144,831.93) > F(. 999,120,
120) and the null hypothesis of no difference
between groups is rejected.
The F-statistics to test between each pair of
groups are listed in Tabic 4. The tabular F-
value is F(. 999,24, 120) = 2.40 (Ostle, 1963);
therefore, the null hypothesis of no difference
is rejected for each of tlie groups compared at
the .001 level of significance.
Each postulated group contained lizards
known to be from a gi\ en population. Individual
lizards were compared to the discriminant func-
tions ( Table 3 ) and assigned to one of the seven
possible populations each time a new character
entered the program (Table 6 and 7). To test
the program's ability to classif)' lizards from
known populations, the percentage of correctly
identified were divided by total number in the
geographic sample (Ncorr Ntot). This was
summed over all groups to get the total sample
correctly identified (Table 8). Approximately
94 percent of the sample was identified correct-
ly, which is well within the 75 percent taxo-
nomic rule of Mayr (1969).
Cranial Morphology
Discriminant Analysis
The discriminant analysis of Dixon ( 1968 )
was used to test its ability to recognize, on the
basis of cranial morphology, the seven previously
postulated populations. The mciin and standard
deviation was calculated for each character
(Table 9). Again the approximate F-value was
used to test the null hypothesis of no difference
between groups instead of the U-statistic be-
cause of the availability of F-tables. The ap-
proximate F-value is F(l-oc ,54,570.59) =4.7506.
The tabular F-value is F( 0.999,40, 120) = 1.84
(Ostle, 1963). Therefore, F (1- a: ,54,570.59 ) >
F( 0.999,40,120) and the null hypothesis of no
difference is rejected at the .001 level of signifi-
cance.
An F-matrix of 9,111 degrees of freedom was
calculated to determine if all groups differ sig-
nificantly (Table 10). The tabular F-value is
F( 0.999,9,60) = 3.69 (Ostle, 1963). Tliercfore,
not all groups are distinguishable at the .001
level of significance.
Table 11 lists the number of individuals
classified into each group after nine characters
have entered the analysis. From this informa-
tion, the number of correctly identified speci-
mens was calculated to see how many popula-
tions fit the 75 percent rule (Table 12). Only
the Great Basin population was distinguishable
\\ith 75 percent accuracy using cranial mor-
phology alone.
DISCUSSION
Ingram and Tanner (1972:6) indicated that
the collared lizards of western Utah, Idaho, Ne-
vada, California, and Baja California are ver\'
different from those east of the Colorado River.
The objective of this study has been to dis-
tinguish the collared lizard populations of these
\\ estern states from those which are contiguous
to the east (C. c. haileiji, C. c. auriceps, and
Table 1. A contingency table to test the independence of Ward's clustering method and the proposed groups.
Proposed Groups
Clusters
Great
Baja
Angel
Tiburon
Upper
Central
Chihuahuan
Great
Basin
Calif.
Island
Island
Colorado
Arizona
Desert
Plains
1
4
6
6
2
12
7
3
16
.0
4
2
17
5
9
9
2
4
6
9
5
4
4
7
1
3
12
8
1
3
2
12
10
Brigham Young University Science Bulletin
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Biological Series, Vol. 19, No. 4 Western Collabed Lizards
11
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12
Brigham Young University Science Bulletin
Table 2. A listing of the means and standard deviations for the 32 characters measured on the individuals used
Groups
bicinctores
(N=50)
vestigium {N = 30)
Standard
insularis
{N^21)
Standard
Standard
Character
Mean
Deviation
Mean
Deviation
Mean
Deviation
1
2.2
0.1
2.3
0.5
2.3
0.1
2
1.9
0.1
2.1
0.4
2.1
0.1
3
91.2
7.1
95.0
15.0
97.4
10.5
4
5.9
0.6
5.0
0.5
5.3
0.6
5
0.0
0.2
0.0
0.0
0.2
0.4
6
3.0
0.8
3.3
0.8
3.0
0.6
7
8.3
0.9
7.2
1.2
8.8
0.8
8
14.7
1.2
15.0
1.6
16.0
1.2
9
0.1
0.3
0.6
0.5
0.1
0.4
10
59.2
4.7
63.1
7.2
67.4
5.4
11
1.3
0.9
2.5
1.3
3.0
1.0
12
15.9
1.3
16.1
1.6
17.3
1.5
13
34.8
3.6
36.9
4.4
41.3
3.7
14
29.1
4.5
25.5
3.9
25.1
0.7
15
174.2
11.2
177.6
12.9
188.6
9.7
16
206.3
11.1
216.2
18.4
227.0
11.4
17
5.1
4.1
19.7
4.0
35.4
12.3
18
2.5
3.4
26.6
9.3
44.1
4.1
19
0.0
0.0
0.1
0.4
0.0
0.0
20
0.1
0.0
0.1
0.0
0.0
0.0
-21
19.7
1,2
19.2
1.7
22.0
1.0
22
37.6
2.0
36.1
3.0
41.7
2.0
23
20.1
1.7
27.1
37.1
24.7
1.7
24
17.8
1.3
19.5
2.0
20.2
1.3
25
0.0
0.0
0.0
0.0
0.0
0.0
26
0.0
0.2
1.0
0.2
0.9
0.4
27
0.5
0.5
0.6
0.5
0.4
0.5
28
0.5
0.5
0.6
0.5
0.3
0.5
29
0.3
0.4
0.0
0.0
0.0
0.0
30
0.2
0.4
0.3
0.5
0.6
0.5
31
0.3
0.5
0.6
0.5
0.3
0.5
32
0.9
0.3
0.7
0.5
0.7
0.5
C. c. fiiscus) and to determine their relation-
ships to the previously established species, C.
insularis and C. dickersonae. To delimit their
characteristics, an investigation was made of
their external morphology. Because cranial mor-
phology has been used as a prominent character
to distinguish collared lizard populations ( Stej-
neger, 1890; Van Denburgh and Slevin, 1921),
we have investigated, by discriminant analysis,
its ability to define unilaterally the seven popu-
lations within this study (Table 9).
External Morphology
Initially, eight collared lizard populations
were postulated (Table 1). The clustan pro-
gram of Wishart ( 1968 ) was used to partition
individuals into clusters of closest morphologi-
cal similarity. The program was discontinued
after eight clusters had been fonned and a con-
tingency table constructed to test for the in-
dependence of the postulated groups with those
formed by cluster analysis. The chi-square test
indicated that the groups fonned b\' the two
methods are significantly related.
Once the above populations were established,
the stepwise discriminant analysis program of
Dixon (1968) was used to compile discriminant
functions to define more precisely the character-
istics of each population. The discriminant pro-
gram forms a new character, Z, which is the
value of the set of discriminant fimctions fonned
from linear combinations of the original char-
acters. Characters arc added in a linear stepwise
manner to maximize the difference between each
population. Using this procedure, it is possible
with 24 variables to identify correctly 90 per-
cent of each group in this study (Table 7).
Within the discriminant anahsis, five of the
first seven characters listed dealt with collar
pattern (Table 7). When the mean and one
standard deviation are plotted for the four non-
binary variables of these five collar characteris-
tics (Fig. 3), several population groupings are
Biological Series, Vol. 19, No. 4 Western Collared Lizards
13
in forming the discriminant functions.
Groups
dickersonae
{N = 20)
Standard
baileyi (iV
= 20)
Standard
fllSCUS
{N = 20)
Standard
auriceps
{N = 10)
Standard
Mean
Deviation
Mean
Deviation
Mean
Deviation
Mean
Deviation
2.2
0.1
2.2
0.1
2.1
0.2
2.2
0,3
2.0
0.3
1.9
0.1
1.9
0.2
1.8
0.2
92.6
24.1
92.2
6.0
93.2
7,8
94.3
7.3
5.3
0,5
6.1
0.4
5.8
0,8
5,8
0.8
0.0
0,0
0.2
0.4
0.5
0.8
0,1
0.3
3.6
0,6
2.0
0.6
1.7
0.8
2.3
0.8
7.6
1.1
7.4
1.0
6.5
0.8
8.6
1.5
13.6
0.6
13,0
1.8
14.5
1,3
12.5
0.7
0.2
0.4
1.0
0.2
0.9
0.2
0.9
0.3
60.3
5.3
61.2
6.0
62.4
6,2
66.1
4.5
1.6
0.5
0.5
0.7
0.5
0,6
0.1
0.3
16.5
1.2
14.6
1.6
14.3
1,7
15.7
1.2
40.4
3,0
24.3
6.9
27.6
3,5
24.5
6.1
25.7
1.9
33.5
5.1
27.8
3,3
33.7
6.5
173.0
13.0
161.0
10.0
152.1
9.2
158.1
7.9
208.6
9.4
189.7
10.1
184,3
9,4
195.4
9.8
6.7
2.6
24.1
7.0
23.4
5,7
28.9
4.7
2.6
1.1
4.5
7.0
5.9
5,9
1.4
1.8
0.0
0.0
1.9
0.5
1.9
0,2
1.9
0.3
0.2
0.0
0.2
0.0
0.2
0,0
0,2
0.0
19.3
0.6
19.3
1.6
18.7
1,6
17.3
1.3
34.6
1.3
34.6
2.0
34.6
3,0
33.0
3.9
18.8
1,5
15.5
1.6
15.3
1,7
15.0
1.7
18.6
0.9
17.7
1.6
18.1
1,5
18.6
1.6
0.2
0.4
1,0
0.0
1.0
0.0
0.9
0.3
1.0
0.0
0,0
0.0
0.0
0.2
0.0
0.0
0.7
0.4
0,0
0.0
0.0
0,0
0,0
0.0
0.9
0.3
0,0
0.0
0.0
0,0
0,0
0.0
1.0
0,0
0,9
0.4
0.0
0,0
0,8
0.6
0.8
0,4
0,4
0.5
0.9
0,4
0,1
0.3
0.2
0,4
0.4
0.5
0.0
0,0
0.5
0.5
0.5
0.5
0.2
0.4
0.3
0,5
0.5
0.5
possible. Comparison of the number of unique
spots within the dorsal separation of the first
cohar ( Fig. .3I\' ) shows the existence of a dis-
tinct eastern (auriceps, baileyi, fuscus) and
western complex (dickersonae, bicinctores, ves-
tigium, insularis). When the dorsal separation
of the second collar is compared (Fig. 3III),
C. insularis and C i. vestigium are shaiply de-
fined while C. c. bicinctores and C. c. dicker-
sonae are allied with the coUaris complex. We
note from our data that C. c. dickersonae is
allied with both the eastern and western com-
plexes depending upon the variable studied,
thus supporting its possible intennediarv posi-
tion. The blue to green color pattern of C, c.
dickersonae, as well as its oral pigmentation
(Axtell, 1972), link it with the eastern complex,
but the discriminant analysis program,
BMD07M, confuses it more often with C, c.
bicinctores than with any of the eastern popu-
lations. Moreover, using the clustan program
(Fig. 2), we had difficulty distinguishing C. c.
dickersonae and C. i. vestigium. Therefore, a
strong relationship of C. c. dickersonae to the
western populations is indicated.
The a posteriori technique used by Ingram
and Tanner (1971) was used in this study to
detennine areas of intergradation within the
western complex and between it and the coUaris
complex. To do this, a large number of collared
lizards ( .369 ) from all areas of the western com-
plex were inserted into the discriminant analysis
program as unknowns, and the a posteriori
probability for membership in each of the seven
groups was calculated. Analysis of this portion
of the BMD07M program indicated most of
the populations apparently exhibit little inter-
gradation ( Fig, 4 ) , however, exceptions to this
will be discussed in the succeeding .section on
population characteristics.
.Although we were unable to locate in the
field distinct hybrid zones for C. c. bicinctores
and members of the coUaris complex, we
strongly believed they existed, and this has since
14
Brigham Young University Science Bulletin
Table 3. A listing of the coefficients of the discriminant functions after 24 characters have been entered into
the discriminant analysis.
Groups
Character
bicinctores
vestigium
instdaris
dickersonae
baileyi
ftl-SCtlS
auriceps
3
0.36226
0.44728
0.42848
0.45130
0.43575
0.50236
0.42086
4
14.63895
10.93610
12.74734
12.18521
15.86134
14.90975
13.40478
5
11.35417
12.24668
15.48502
12.77270
14.02354
16.45863
13.60895
6
-2.54088
-0.94370
-2.33630
-0.26514
-5.53481
-5.14219
-5.64448
7
3.23171
1.82818
4.22291
1.55059
4.06717
3.76096
5.55091
8
3.85278
3.45407
3.26712
3.24393
3.52133
3.97844
2.81999
9
18.72758
24.49905
24.25928
17.58437
23.31706
21.23149
23.47716
11
0.23926
2.55489
3.48512
0.17327
0.30980
-0.25982
-0.14585
12
4.85020
5.77467
6.38672
5.32886
5.51403
5.72580
6.58892
13
0.19337
0.19038
0.04178
0.60606
-0.49817
0.03978
-0.39207
15
0.72727
0.62564
0.68925
0.54926
0.82512
0.66800
0.73172
16
0.74275
0.84646
0.73013
0.79176
0.62601
0.66604
0.72710
17
-0.17721
0.13376
0.61486
-0.18022
0.28508
0.20895
0.38181
18
0.52595
1.49761
1.88647
0.82280
0.80442
0.91227
0.67961
19
8.27618
8.79661
4.42210
9.42294
37..53778
39.39021
39.79042
20
464.44434
484.50488
327.03882
581.21729
553.91357
610.33252
.525.34448
21
6.43277
5.13985
7.11056
5.47452
5.72452
5.30954
3.84222
23
-0,21490
-0.18263
-0.27868
-0.17966
-0.22353
-0.20750
-0.19154
24
2.39604
3.14634
2.47195
3.44348
3.56539
3.53492
3.99242
25
-11.70876
-12.66015
-7.65450
-3.18096
40.52048
36.31189
38.14626
26
-10.62225
16.41115
10.58251
17.64139
-9.40444
-8.44764
-9.30175
29
-8.40836
-8.49082
-8.37933
-0.25997
4.76858
-5.80569
3.35294
30
18.44745
16.25493
20.84308
18.84085
17.67749
20.89793
15.69166
32
0.50571
-1.09726
0.24414
-2.22648
-5.08565
-5.09013
-1.87483
constant
-395.93677
-418.50757
-472.26001
-416.04150
-476.39014
-478.31909
-458.56494
Table 4. A summary of the F-statistics which .show the differences between individual groups F (.999,24,120)
= 2.40, at the .001 level of significance (Ostle, 1963).
Groups
Groups
bicinctores
vestigium
instdaris
dickersonae
baileyi
fiisctis
vestigium
insularis
dickersonae
baileyi
fuscus
auriceps
45.05
66..55
17.89
24.81
19.79
47.47
75.37
82.96
93.10
56.11
72.24
76.71
89.32
54.13
5.76
45.20
51.57
62.22
38.66
3.01
6.55
Table 5. Number of cases classified into each group after 24 characters entered into tlie stepwise discriminant
analysis program.
Groups
Groups
bicinctores
vestigium
insularis
dickersonae
baileyi
fuscus
auriceps
bicinctores
49
1
vestigium
29
1
instdari.!
1
20
-•
dickersonae
20
baileyi
18
1
1
fuscus
20
auriceps
1
9
Biological Series, Vol. 19, No. 4 Western Collared Lizards
15
Table 6. Summarv table of the sequential entry of characters into the BMD07M discriminant analysis program
with their F-value to enter or remove along vv-ith the U-statistic at each step.
Variable
F-value
Step
Entered Removed Enter or Remove
U-statistic
1
19
334.6821
0.0755
2
18
178.9013
0.0100
3
26
74.8575
0.0026
4
29
26.4363
0.0013
5
25
2.3.6078
0.0007
6
17
13.0184
0.0005
7
20
9.4747
0.0003
8
21
7.8510
0.0003
9
30
7.1107
0.0002
10
13
5.8263
0.0002
11
7
5.1555
0.0001
12
5
2.4926
0.0001
13
4
2.3347
0.0001
14
9
2.2488
0.0001
15
11
2.2193
0.0001
16
24
2.1778
0.0001
17
6
1.9843
0.0001
18
15
2.1924
0.0001
19
32
1.6604
0.0001
20
16
1.2632
0.0001
21
12
1.2978
0.0001
22
8
1.1968
0.0001
23
23
1.0629
0.0001
24
3
1.0144
0.0001
Table 7. A percentage comparison of correctly identified individuals within groups as new variables were added
to the discriminant functions.
Variable
Added
Groups
Step
hicinctores
vestigium
insularis
dickersonae
haileiji
ftisctis
auiiceps
1
19
100
5
95
2
18
62
77
95
65
5
60
70
3
26
96
73
95
100
5
65
70
4
29
96
83
95
100
35
100
30
5
25
96
87
95
100
85
100
10
6
17
96
90
95
100
60
100
30
7
20
96
93
90
100
65
100
60
8
21
96
93
90
100
65
100
60
9
30
96
93
95
100
60
100
80
10
13
96
93
95
100
70
100
90
11
7
96
97
95
100
75
100
90
12
5
96
97
95
100
75
100
90
13
4
96
97
95
100
70
100
80
14
9
96
97
95
100
70
100
80
15
11
96
97
95
100
70
100
80
16
24
96
97
95
100
75
100
90
17
6
96
97
95
100
75
100
90
20
16
98
97
95
100
85
100
90
24
3
98
97
95
100
90
100
90
Table 8. Percentage of sample identified correctlv using
the discriminant analysis.
Group
Sample Size
Ncorr./Ntot.
Percentage
hicinctores
50
49/50
98.00
vestigium
30
29/30
96.67
insularis
21
20/21
95.25
dickersonae
20
20/20
100.00
haileyi
20
18/20
90.00
fiiscus
20
20/20
100.00
auriceps
10
9/10
90.00
Total
171
165/171
94.15
been verified bv Axtell (1972) in an area north
of Kingman, Arizona. Axtell further cites speci-
mens from Coconino and Maricopa counties,
Arizona, as possible hybrids. Banta (1960) de-
scribed a greenish collared lizard taken west
of Davis Dam, Clark County, Nevada, but in-
dicated it was a color trait seldom seen in
western populations of collared lizards. A pre-
served specimen, LACM 26824, from Inyo
Countv, California, well within the range of C. c.
hicinctores, appears to be green and has a very
16
Bricham Young University Science Bulletin
Table 9. A listing of the means and standard deviations of the 16 cranial morphology characters.
Groups
bicinctores
vestigium
insularis
Mean
Standard
Mean
Standard
Mean
Standard
Character
Deviation
Deviation
Deviation
1
25.61
1.87
27.57
3.32
27.39
3.25
2
11.63
0.77
12.54
1.14
12.88
1.50
3
16.31
1.48
17.37
2.33
16.31
1.90
4
14.06
0.77
15.11
1.90
14.92
1.52
5
0.45
0.02
0.46
0.02
0.47
0.01
6
0.64
0.03
0.63
0.03
0.60
0.02
7
0.55
0.03
0.55
0.02
0.55
0.02
8
0.02
0.00
0.02
0.00
0.02
0.00
9
0.02
0.00
0.02
0.00
0.02
0.00
10
1.40
0.09
1.38
0.10
1.27
0.05
11
1.21
0.07
1.20
0.07
1.16
0.05
12
0.04
0.00
0.04
0.00
0.04
0.00
13
0.05
0.00
0.05
0.00
0.05
0.01
14
0.04
0.00
0.04
0.00
0.04
0.00
15
0.05
0.00
0.04
0.01
0.04
0.00
16
1.40
0.09
1.38
0.10
1.27
0.05
Table 10. A summary of the F-statistics which show the differences beUveen individual groups in the cranial
morphology study. F( .999,9,60) = 3.69, at the .001 level of significance (Ostle, 1963).
Groups
Groups
bicinctores
vestigium
insularis
dickersonae
baileyi
fuscus
vestigium
1.43030
inmlaris
6.97606
4.87994
dickersonae
2.46489
1.71034
7.09683
baileyi
2.78564
4.65735
13.80026
,5.94830
fuscus
6.63592
9.44739
18.35388
9.77859
1.860.56
auriceps
3.67191
3.90008
11.17101
5.49404
1.46455
3.15051
Table 11. Number of cases classified into each group after
analysis program for cranial morphology.
liaracters entered into the stepwise discriminant
Groups
Groups
bicinctores
vestigittm
insularis
dickersonae
baileyi
fuscus
auriceps
bicinctores
15
1
2
2
vestigium
2
9
4
3
2
insularis
1
4
11
dickersonae
5
5
1
7
1
1
baileyi
4
2
9
3
2
fuscus
1
1
6
9
3
auriceps
2
3
1
4
Table 12. Percentage of sample identified correctly
from discriminant analysis of the head characters after
nine characters considered.
Sample
Ncorr. /
Percentage
Group
Size
Ntot.
bicinctores
20
15/20
75.00
vestigium
20
9/20
45.00
insularis
16
11/16
68.75
dickersonae
20
7/20
35.00
baileyi
20
9/20
45.00
fuscus
20
9/20
45.00
auriceps
10
4/10
40.00
Total
126
64/126
50.79
unu-sual dorsal recticiilation. Future investiga-
tion could reveal hybrids in the Moab area of
Grand County, Utah. It is difficult to locate
contact zones because collared lizards seldom
cross the vallevs which separate them from their
preferred mountain slopes. Hence thev are
deniic even within the range of unquestionably
similar populations.
Following the technique of Rao (1953), a
graph of the first two canonical variables was
plotted which represents in two-dimensional
space the relationships of the seven populations
studied ( Fig. 5 ) . The overlapping populations
Biological Series, Vol. 19, No. 4 Western Collared Lizards
17
Groups
dickersonae
baileyi
fuicus
auriceps
Mean
Standard
Mean
Standard
Mean
Standard
Mean
Standard
Deviation
Deviation
Deviation
Deviation
29.16
5.25
26.55
3.01
25.64
3.60
25.37
2.36
12.92
1.82
11.82
1.31
11.37
1.43
11.27
0.86
18.32
3.45
18.07
2.62
17.79
2.82
17.13
2.11
15.43
2.11
15.02
1.66
14.87
1.76
14.78
1.78
0.45
0.02
0.45
0.02
0.44
0.02
0.45
0.03
0.63
0.02
0.68
0.03
0.69
0.03
0.67
0.03
0.53
0.04
0.57
0.03
0.58
0.02
0.58
0.03
0.02
0.00
0.02
0.00
0.02
0.00
0.02
0.00
0.02
0.00
0.03
0.00
0.03
0.00
0.03
0.00
1.41
0.10
1.53
0.10
1..56
0.10
1.52
0.14
1.20
0.07
1.27
0.10
1.31
0.05
1.31
0.13
0.04
0.01
0.04
0.00
0.04
0.00
0.04
0.00
0.05
0.01
0.06
0.00
0.06
0.00
0.06
0.00
0.(H
O.OI
0.04
0.00
0.04
0.00
0.04
0.00
0.04
0.01
0.05
0.00
0.05
0.00
0.05
0.00
1.41
0.10
1.53
0.10
1.56
0.10
1.52
0.14
(I)
lllll
05 1
015 OT^
(III
"When the second collar is absent, the dorsal sep.ira
tion is arbitrarily set at 45.
+
Fig. 3. Means and standard deviations plotted for the
four nonbinary characters that contribute most to
the distinction of the four western-complex popu-
lations from C. c. haileiji, C. c. auriceps, and C. c.
fuscus: (1) second collar length/snout-vent length,
( II ) number of scales within dorsal separation of
first collar, (III) number of scales within dorsal
separation of second collar, ( IV ) number of spots
within dorsal separation of first collar. (C) cen-
tral Arizona, (A) upper Colorado River Basin,
( F ) Chihuahuan Desert, ( T ) Tiburon Island, ( I )
Angel Island, (B) Baja California-southern Cali-
fornia, and ( G ) Great Basin populations.
18
Brigham Young University Science Bulletin
Percent of
Sa m p le
100-
Great Basin
( Idaho, Nevada, western Utah, southwestern Arizona,
eastern Cahfomia)
80-
Percent of
Sample
Sonora, Mexico
(West of Highway 15, Organ Pipe National Monument
area and southern end Tinajas Altas Mountains exclusive
of Tiburon Island)
100-
80-
Fig. 4. Continued on next page.
Biological Series, Vol. 19, No. 4 Western Collared Lizards
19
Percent of
Sample
100-
Baja California, southern California south from the San
Gorgonio Pass
B BXI 1 G GXB U
Fig. 4. A comparison of collared lizard populations from the Great Basin, northwest Sonora, Mexico, and Baja
California-southern California.
A key to the abbreviations used in Fig. 4.
A- — auriceps population from the Upper Colorado River Basin.
B — vestigium from the Baja Cdifomia, southern California population.
C — baileyi population from Central Arizona.
F — fuscus population from Chihuahuan Desert.
G — hicinctores population from Idaho, Nevada, western Utah and Arizona, and eastern California.
I — ii^sttlaris population from Angel Island.
T — dickersonae population from Tiburon Island.
GXT — intergrade population of hicinctores and dickersonae.
CXG — intergrade population of bailctji and hicinctores.
A.XC — intergrade population of auriceps and hicinctores.
FXG — intergrade population of fuscus and hicinctores.
CXF — intergrade population of baileyi and fuscu.^.
BXI — intergrade population of vestigium and insularis.
G.XB — intergrade population of vestigium and hicinctores.
U — Individuals not assignable to any of the above groups.
are Great Ba.sin and Tibviron, witli a few mem-
bers of the Tiburon group shifting toward the
coUaris complex. This is e.xpected, since we con-
sider the Tiburon-Sonoran populations to have
been derived from those populations evolving
to the cast and now designated as C. c. fuscus.
The Baja California group is more closely re-
lated to the Tiburon group in the characters
studied than it is to the Great Basin popula-
tion. This is in agreement with the results of
the cluster analysis. There are two valid reasons
members of the collaris complex did not sep-
arate as well in this plot as the one shown by
Ingram and Tanner (1971:23): (1) a more re-
duced scale was used to plot this graph, and ( 2 )
not all the same characters used by Ingram and
Tanner were used in this study. These two
points should be kept in mind along with the
fact that our statistical tests supported these
eastern populations ( Table 4 ) .
Cranial Morphology
The skull has been shown to be significant
in the definition of subspecific taxa (Avery
and Tanner, 1964, 1971; Presch, 1969; Robison
and Tanner, 1962). Van Denburgh and Slevin
( 1921 ) declared C. imularis to have a longer,
narrower head than C. c. baileyi. In comparing
the cranial morphology of the seven popula-
tions of this study, only the Great Basin group
reached 75 percent distinction (Table 12). The
character which best separates the western and
collaris complexes is head width at widest point/
total skull length (Fig. 6). Within these com-
plexes, the distinctions, except for the Great
Basin group, are not significant.
20
Bricham Young Univebsity Science Bulletin
C
$
C
F C$
F
CA FF$$FC
C$$**F $
AC $
C
F AF C
A C
F
T
T
GG GG
G GGGG
GGG-^GG
G GGGG
G
G G
T
T T
*T XT G
GT
B B
B
B B B
BB B B
B *
BBB I
A key to the symbols used
A - Upper Colorado River Basin population
B - Baja population
C - Central Arizona population
F - Chihuahuan Desert population
G - Great Basin population
I - Angel Island population
T - Tiburon Island population
" - Group mean
$ - Group overlap
B $
I
* I
I I
II
I
Fig. 5. A two-dimensional representation of the external morphological interpopulation relationships formed from
canonical varieties.
Although Van Denburgh's data were appar-
ently linear rather than proportional (1922:116),
his assumption that C. iiisiihiiis is distinguishable
from C. c. baileyi, as defined by us, by cranial
characteristics is correct (Table 11). Additional
research is needed to determine if osseous ele-
ments are responsible for the differences in
cranial proportions.
On the basis of cranial measurements, mem-
bers of the western complex show a closer rela-
tionship to one another than to members of the
collaris complex. This is the same relationship
which evolved in the cluster analysis portion of
this study. Using the a posteriori probabilities
generated by the BMD07M program, it is not
possible to determine distinct areas of intergra-
dation for the Great Basin group and the
collaris complex.
Population Characteristics and Systematics
The characteristics which distinguish mem-
bers of the collaris complex have been sum-
marized by Ingram and Tanner (1971:21-24).
The characters which separate them from" the
western complex are: (1) one or more spots
present in the dorsum of the first collar; (2)
absence of a prominent ventral black groin patch
on the males; (.3) fewer scales occurring
BioLOCiCAi. Series, Vol. 19, No. 4 Western Collared Lizards
21
Trr
Fig. 6. The mean and standard deviation tor head
width at widest point/total .skull length; (K) C. c.
fuscu?, (C) C. c. haileyi, (A) C. c. auriceps, (G)
C. c. hicinctores, (T) C. c. dickersonae, (B) C. i.
oesfigiuni, (1) C. i. insiilaris.
from the interparietal to the anterior border of
the first collar, and (4) the .second collar usu-
ally extending onto the upper arm. Additional-
ly, AxtcU (1972) suggests a rounded tail and
dark oral pigmentation as characteristic of the
coUaris complex.
Tlie four populations of the western complex
are believed to represent two species of Crota-
phtjtus. The Great Basin and Tiburon Island-
Western Sonoran populations are considered
C. coUaris. The southern California, Baja Cali-
fornia, and Angel Island populations are con-
sidered C. insularis. The Great Basin popula-
tion, which we consider C. c. hicinctores (al-
though similar to C. insularis with its brownish
dorsal coloration, ventral collar, gular, and ab-
doniino-lateral pigmentation), is distinctly dif-
ferent in dorsal pattern and collar characteristics.
Upon visual inspection, there is no confusion be-
tween a C. c. hicinctores and C. insularis; this
is true even between females ( Smith and Tanner,
1972: Fig. 1). Such is not the case when com-
paring females of two closely related subspecies
such as C. c. fuscus and C. c. hailet/i. Cluster
analysis ( Fig. 2 ) and canonical analysis ( Fig.
5) sharply distinguish C. c. hicinctores from
C. insularis; however, the distinction between
Baja California and Angel Island populations,
two populations we consider to be subspecifical-
ly related, is not so apparent.
In our earlier paper (Smith and Tanner,
1972), we indicated an area of sympatry for C. c.
hicinctores and C. /. vestigium in the vicinity of
the San Gorgonio Pass, Riverside County, Cali-
fornia. This assumption was based upon a fe-
male hicinctores, CSCLB VAP 660513-4, said
to have been taken from the side of the road
leading from Palm Springs to the aerial tram-
way in an area which is occupied by vestigium.
This specimen, when reconsidered, may repre-
sent the results of human intervention in collared
lizard distribution. Dr. Ernest Tinkum and
James Davis of Indio, Riverside County, Cali-
fornia (pers. comm.), indicate they have not ob-
served hicinctores from this area. Until a
thorough ecological study can be completed
within the vicinity of the San Gorgonio Pass,
we now prefer to believe hicinctores and ves-
tigium represent allopatric species (Fig. 7). Ad-
ditional specimens of interest from this area are
LACM 16SS9, a female hicinctores from Caba-
zon, Riverside County, California; and UCMVZ
1, a female vestigium from Hallgrade, near
Cabazon. The latter specimen was taken by
W. P. Taylor in 1908. Stephen B. Ruth, from
the USMN'Z, has examined the field notes and
reports that Taylor's base camp was I'A miles
south of Cabazon and that on May 9 Taylor
walked up to Hurley Flat (which was south of
his camp) via the Hallgrade. In this case, the
specimen was seemingly taken on the low foot-
hills at the base of die San Jacinto Mountains.
Axtell (1972) has taken the position that
hicinctores should be placed in tlie Crotaphijtus
insularis species complex. His arrangement is
based on some morphological similarities be-
tween these species, and we readily agree that
there are similarities. However, if one is to ac-
cept his h\pothesis that hicinctores is specifically
distinct from C. coUaris because introgression
is very narrow in the Cerbat Mountains where
he has pinpointed intergradation, then by that
reasoning— and on the basis of the information
presented above— we would suggest that hi-
cinctores is also specifically distinct from in-
sularis. In the information to follow, we will
give indications of additional intergradation be-
tween hicinctores and other coUaris suhspecies
in northern Sonora, Mexico. These data seem to
relate hicinctores more closely to haUeiji, fuscus,
and dickersonae, and we, therefore, prefer to
group hicinctores with coUaris at present.
The possibility that hicinctores, which has
been found as far north in the upper Colorado
River Basin as the Potash Road (BYU 3114),
Grand County, Utah, intergrades with C. c.
auriceps has not not yet been demonstrated.
However, Axtell (1972) has demonstrated that
hicinctores does intergrade in Arizona with C.
c. haileyi, which is a close relative of auriceps.
In the southern portion of its range, hi-
cinctores has not been shown to contact or inter-
grade directly with C. c. dickersonae; but it
does intergrade with C. c. fuscus, and C. c.
22
Brigham Young University Science Bulletin
Fig. 7. Range map for Crotaphtjtus collaris and Crotaphtjtus imularis: (1) C. c. bicinctores. (2) C. i. veitigium,
(3) C. i. insularis, (4) C. c. dickersonae, (5) C. c. aunceps, (6) C. c. baileyi, (7) C. c. /iwcus.
Biological Series, Vol. 19, No. 4 Western Collared Lizards
23
fuscus intcrgrades with C. c. dickersonae (See
the following discussion of population character-
istics for dickersonae).
Specimen UAZ 12114 from an area midway
between MacDougal and S\kes Crater in north-
western Sonora, Mexico, is considered by us to
be an intergrade between C. c. bicinctores and
C. c. fusctis. Its appearance is very much like
bicinctores. The a posteriori probability was
1.000 for bicinctores. It has, however, jtiscus
characteristics such as its second collar extend-
ing on to the forearm, the first collar having
an anterior extension that could be interpreted
as the incomplete formation of spots, a rounded
tail (a coUaris character suggested by Axtell,
1972 ) , a ver\- reduced ventral groin patch, minor
pigmentation of the dewlap, postmentals con-
tacting the infralabials, and .34 subdigital lamel-
lae on the fourth toe.
Similarly, UAZ 10248 from 36.7 miles west
of Sonoita has a posteriori probabilities of 0.687
for C. c. aiiriceps and 0..3()7 for C. c. fuscus.
Visual inspection of the color and dorsal pat-
tern reminds one of bicinctores. Additional
bicinctores characteristics include: 3 frontopa-
rietals, 3 scutae in dorsal separation of first
collar, no scutae in dorsal separation of second
collar, and 21 subdigital lamellae on the fifth
toe. It is seemingly an intergrade of fuscus
and bicinctores.
Specimens which we have observed from
Organ Pipe National Monument, Pima County,
Arizona (UAZ 636, 1511; LACM 3983, 5899),
have appeared to be fuscus: but visual inspec-
tion of the first collar pattern suggests it is not
nearly so characteristic as those from the type
locality in Chihuahua, Mexico. When the dorsal
collar pattern is masked and only the pattern on
the back is \isible, it is often difficult to dis-
tinguish these fuscus specimens from bicinctores.
Several specimens from the area of Caborca
and San Luis, Sonora, are especiallv interesting
because of the a posteriori probabilit\' listed for
them by the BMD07M program. On the basis
of this information, one would suspect this to
be an area of intergradation or at least an area
which represents the remnants of genes repre-
resenting bicinctores, dickersonae, fuscus, and
other members of the eastern complex, which
may nou' be geographically or ecologicallv iso-
lated. The specimens, with their localities and
probabilities, are UCMVZ 10163, d", 30 miles W
Caborca, .406 fuscus, .353 bicinctores, .241
dickersonae; CAS 104467, 9 , 9 miles E Caborca,
.529 bicinctores, .435 baileiji, .024 fuscus. Addi-
tional field studies in this area are needed before
complete relationships and distributions can be
ascertained. However, present data indicates
considerable intergradation.
The members of C. c. dickersonae are similar
to C. c. bicinctores in many respects, including
collar pattern, but are distinguished from them
by their blue or green dorsal ground color and
dorsal light elongate spots. These two popula-
tions are so closely related that many of their
morphological characteristics, although different,
show considerable overlap (Table 2). The dor-
sal light elongate spots of dickersonae are sug-
gestive of those found in C. insularis. The ven-
tral pigmentation of male dickersonae is not as
pronounced as that seen in bicinctores or in-
sularis. The range of dickersonae can now be
extended north from near Kino Bay to Puerto
Libertad ( UAZ 20144 ) and to El Desomboque,
Sonora, Mexico (UAZ 704, 705, and 30226). Its
range south from Kino Bav is not completely
known, but two specimens (BYU 409.30-31)
from the hills north of Guaymas show inter-
grading characters between dickersonae and
fuscus.
Both of the above specimens have a
posteriori probabilities of 1.000 for dickersonae.
Msual inspection reveals the male ( BYU 40931 )
to have dorsal light elongate spots, a blue
green cast, 3 frontoparietals, no fused interorbi-
tals, 9 loreal-lorilabials and 2 enlarged inter-
nasals— all of which characterize dickersonae.
Conversely, the second collar extends boldly
onto the upper forearm, the postmentals contact
the infralabials, there are 5 scutae within the
dorsal separation of the first collar, and the tail
is round— all of which characterize fuscits. In
addition, the first collar unites ventrally and
there is bold abdomino-lateral pigmentation,
which is not characteristic of either population.
Axtell (1972) noted this exaggerated effect on
the variables found in his intergrades of bicinc-
tores and l)ailetii. Such intergrade variables
cause one to wonder if bicinctores has not arisen
as a result of hybridization between several of
the subspecies and established its range in the
lo\\'cr Colorado River Basin with its major ex-
tension into the Great Basin. The female of this
pair (BYU 40930) has dickersonae characteris-
tics of 3 frontoparietals, 8 loreal-lorilabials, 7
scutae in the dorsal separation of the first collar,
and dorsal light elongate spots. Her fuscus
characteristics include 1 fused interorbital, 21
scutae from the interparietal to anterior edge of
first collar, 5 scutae in the dorsal separation of
the second collar, and the extension of the sec-
ond collar onto the arm. We consider both of
these specimens to be intergrades between fus-
cus and dickersonae.
24
Brigham Young University Science Bulletin
The following additional specimens from the
Guaymas area are listed with their museum
number and their BMD07M a posteriori proba-
bilities. The probabilities are not consistently in
favor of any given population indicating a
gene pool that has traits for clickersonae, ftiscus,
bicinctores, baileyi, and auriceps. We have not
been able to collect collared lizards in the area
near Guayn as and suggest that this will be es-
sential to understand better the complete rela-
tionship of (Uckersonae and fuscus. The speci-
mens, with their identification and a posteriori
probabilities, are: LACM 52886, d, 9 miles
NW Guaymas. .005 dickersonae, .004 fuscus,
.991 bicinctores; LACM 52882, 9 , 9 miles NW
Guaymas, 1.000 fuscus; UAZ 10597, 9, 29.3
miles S Los Pocitos, .795 fuscus, .205 bailetji;
SDSNH 35918, d, between Guaymas and Em-
palme, .669 auriceps, .268 fuscus, .058 baileyi,
.006 bicinctores; SDSNH .35919, d, between
Guaymas and Empalme, .404 bailetji, ..339
fuscus, .214 bicinctores, .043 auriceps; and
SDSNH CRM 117, d, Playa Miramar SW of
Guaymas, 1.000 fuscus.
Specimen GAS-SU 14002, reported collected
from Tiburon Island, is a perfect example of C.
i. insularis and presents an anomalistic situation.
The specimen was collected 7 May 1952. CAS-
SU 14003-14012 taken by the same party two
days earlier, 5 May 1952, from Tiburon Island,
are typical dickersonae. To us it seems very
possible the GAS-SU 14002 has been mislabeled.
The Baja Califoniia-southem Galifoniia pop-
ulation, C. insularis vesti'^ium, represents a dis-
tinct taxon. It differs from C. c. baileyi in those
characters which distinguish it from C. c. bicinc-
tores. Crotaphytus i. vestigium can be distin-
guished from the Great Basin population by its
reduced first and second collar and the distinct
dorsal transverse bands which separate light
elongate spots (Smith and Tanner, 1972: Fig.
1).
Crotaphytus i. vestigium can be distin-
guished from C. i. insularis by fewer lamellae
on the second and fourth toe, a larger second
collar/s-v ratio, and a smaller dorsal separation
of both the first and second collars (Table 2).
Phyletic Relationships
It is noteworthy that cluster analysis separ-
ated collared lizards into eastern and western
groups, producing four clusters for the western
complexes and four for the eastern. These, we
believe, can be used as indicators of phyletic
relationships (Fig. 2). Within the western
group, the most distinct population is C. i.
insularis from Angel Island, one of the older
islands in the Gulf of California (Anderson,
1950). Female collared lizards from Angel
Island, according to Banta (1961), bear a simi-
larity to the leopard lizard Crotaphytus tois-
lizenii silus. It is probable that the collared
lizards from Angel Island display some of the
genetic variance which gave rise to C. wislizenii
and are a relict population which was dcri\'ed
from a more widespread population, the rem-
nants of which are now found on the island
and Baja California Peninsula. Banta ( 1961 )
believes C. collaris is an older population than
the leopard lizards. This we have not been able
to demonstrate from our data; however, pre\ious
studies have shown that Crotaphytus reticulatus
of eastern Mexico and southcentral Texas, a
fonn which lacks the collar pattern of C. col-
laris, is probably an intennediate between C
collaris and C. ivislizenii (Robison and Tanner,
1962; Montanucci, 1969). l>e "^wo species in the
genus which may be considered the more primi-
tive members of the collared lizard complex are
found on the extreme eastern edge (reticulatus)
and the extreme western edge (insidaris) of
their distribution.
The historical distribution of the collared
lizard in the Gulf of California region is a
matter of speculation at present. Those found
on Angel Island and the Baja California Penin-
sula are more closeh' related to I'ach other
than they are to C. c. dickersonae from Tiburon
Island or bicinctores in the Great Basin ( Fig.
5), indicating a long period of isolation in the
western portion of the Gulf. Tlic Baja Cali-
fornia Peninsula is thought to have formed
some four to ten million years ago in late Plio-
cene by crustal movement ( Moore and Buffing-
ton, 1968). A peculiar feature in this h\pothesis
is the fomiation of the northern portion of the
Gulf of California before the existence of a
southern entrance (Larson, Menard, and Smith,
1968). It is possible, therefore, to hypothesize
at least three modes of distribution of the col-
lared lizard from mainland Mexico to the Baja
California Peninsula: (1) the peninsular popu-
lations were isolated by the peninsular drift
which formed the Gulf of California; (2) the
collared lizard extended its range to the Baja
California Peninsula by a migration south of the
newly forming Gulf of California, later to be
isolated as the peninsula was formed; (3) the
collared lizard migrated aroimd the northern
end of tlie Gulf of California and later liad .its
range restricted to the peninsula, isolating it
from the more recent populations to the north
and east.
The origin of the Angel Island population
from the Baja California population is difficult
Biological Series, Vol. 19, No. 4 Western Collared Lizards
25
to assess because of the absence of a good fossil
record. Angel Island is a deep-water island
formed during the Pliocene (Anderson, 1950).
A possible land bridge between the island and
the peninsula might have pemiittcd unrestricted
distribution of the collared lizard. Later, when
the land bridge was inundated, the insular pop-
ulation became a relict. It is also possible that
the collared lizard was rafted to the island be-
fore the channel became extensive.
Tiburon, a shallow-water island, was un-
doubtedlv separated from Sonora, Mexico, when
the level of the seas rose during the recession of
the last glacial period of 10,000 to 15,000 years
ago (Lawlor, 1971). E\cn today, the channel
between the island and the mainland is very
narrow (ca. 5 miles) and shallow at low tide.
Crotaphi/fus c. dickersonac was described from
Tiburon Island (Schmidt, 1922), but this study
has shown that it is not restricted to the island
and occurs at least along the nearby Sonoran
coast. Moiphologv indicates that dickersonac
has not di\erged to any great extent from the
original stock which gave rise to it and hicinc-
tores of the Great Basin. The canonical analysis
(Fig. 5) indicates that dickersonac diverged
from the coUaris complex from whicli all mem-
bers of the \\c'Stern compk^x of collaris probably
arose at a much later date than the insularis
stock.
The Great Basin, formed in late Tertiary
(Banta, 1961), did not present a favorable en-
vironment for the modern collared lizard until
late Pliocene or early Pleistocene, when many
north-south oriented mountains were formed
(Axlerod, 1948). The nortlu'rn range of the
collared lizard population in the Great Basin
ebbed and enlarged with changing Pleistocene
climatic conditions ( Banta, 1961 ) until the arid
conditions beginning about 11,000 vears ago
(Antevs, 1955; Broecker, 1957) permitted its
present northern extension. The modern collared
lizard apparently did not invade the Great
Basin or the Great Plains before Pleistocene
(Brattstrom, 1954: Harris and Findlex , 1964;
and Gehlbach, 1965a), although Gilmore (1941)
described the genus Aciprion Cope from the
Oligocene deposits of Wyoming as similar to
Crofaphytus.
The Great Basin collared lizard, C. c. bicinc-
tores, is possibly a more xeric form than mem-
bers of the eastern coUaris complex. Miller and
Stebbins (1964) report that collared lizards do
not occur at elevations above 5700 feet in the
moimtains of Joshua Tree National Monument,
San Bernardino County, California; whereas
members of the coUaris complex are found up to
7500 feet elevation in the Zuni Mountains of
northwestern New Mexico (Gehlbach, 1965b).
The preference of a more mesic environment by
members of the coUaris complex might be ex-
plained by natural selection for their green col-
oration; conversely, the brown coloration of
the more xeric western fomi seemingly provides
them an adaptive advantage in their arid en-
vironment.
Tiburon-Sonoran and Great Basin popula-
tions evidently had their genesis in northcentral
Mexico, probably from the line of C. c. fiiscus,
spreading west to the Gulf Coast and the Colo-
rado River and later invading and occupying the
west side of the river as well as the extremely
dry areas of western Arizona. In Utah, the Colo-
rado River has served as an effective barrier,
limiting the range of C. c. hicinctores to the
west side of the riv(^r. A proposed phylogenic
relationship is shown in Fig. 8. Distribution
and relationship is based primarily upon mor-
phological characteristics.
The phylogeny of the genus Crotaphytus ap-
parently had its beginnings in southcentral
United States and northcentral Mexico before
the Pleistocene ( Robinson and Van Devender,
1973). Indications are that the genus, or its
immediate ancestral stock, was well established
and gave rise to a radiation, which resulted
in a division of the genus into the primitive
collared lizard and the leopard lizard branches,
perhaps as early as late Pliocene. Although the
fossil record does not support a Pliocene radi-
ation for the crotaphxtines, if Aciprion Cope is a
member of this iguanid line, its origin may then
be extended back into middle Tertiary. We
surmise that the original stock of the coUaris
group may not have exhibited the collar so typi-
cal of the group as we recognize it today. We,
therefore, suspect that C reticidatiis is a rem-
nant of a much older and originally more wide-
spread population than members of the present-
ly extant collared lizard group. We further
suspect that this primitive stock continued to
evolve and gave rise to the coUaris group which
has been so successful in its invasion of the great
plains, the Rio Grande Valley, the plateaus
and mountains of Sonora, Arizona, New Mexico,
Utah, and the Great Basin since the Pleistocene.
We are not certain as to whether the insularis
group represents a primitive coUaris stock which
has recently been replaced in most of the west-
em distributional areas or if it represents a
modification of the western coUaris group which
has, through isolation, become modified from the
ancestral stock which also gave rise to the So-
noran and Great Basin populations. The re-
26
Bricham Young Univehsity Science Bulletin
C . c. bic i nctores
C. i in su la r is
ANCESTRAL STOCK
Fig. 8. Proposed phylogenetic relationship of Crotaphytus.
duced collar in the insularis populations is so
distinct from the collar pattern found in the
other western subspecies of coUaris that we are
inchned to believe the instilaris group is more
primitive than other collared lizards. The in-
sularis group perliaps preceded the coUaris
groups into the western deserts and at present
represents a reHct which arose shortly after the
coUaris ancestral stock was established in the
more central and southern part of their present
distribution.
The ancestral stock of the coUaris complex
appears to have been derived from a stock simi-
lar to C. c. fiiscus. Tlicre seems to be little
reason to doubt its close relationship to those
forms now occurring from the Great Plains west-
ward to the Colorado River. Although these
populations have been separated into a series of
subspecies, they are all similar in many basic
characteristics. Their distinction from the west-
em coUaris subspecies is undoubtedly a series
of adaptive modifications resulting from the
changing environment and perhaps isolation.
The extensive development of the collars of
hicinctores and dickersonae, plus the fact that
they do not overlap the distribution of insularis.
is additional indication that they are more recent
arrivals than insularis. These circumstances
again suggest a relict distributional pattern. As-
suming this to be the general trend in the past
development of this branch of the genus Crota-
phytus, we could readih' assume that the ebbing
of the Ice Age allowed the then established
coUaris stock to extend its distribution north-
ward and to the west. It occupied extensive
areas in central and western United States and
at the same time was undergoing adaptive
changes which led to the distinct subspecies
populations now extant.
Perhaps the most extensive subspecific group
is that of hicinctores. Even though this group
extends from southern California and southwest-
em Arizona northward into Nevada, Utah, and
southern Idaho, its distribution presumably is
so recent or its genetical makeup so stable that
subspeciation has not occurred. This is in con-
trast to the subspecies cline extending northward
from Chihuahua into southeastem Utah. The
Great Plains populations must still be care'fully
examined, and we expect that they may also
show a similar evolutionary development to that
observed in the baileyi branch.
Biological Series, Vol. 19, No. 4 Western Collared Lizards
SUMMARY AND CONCLUSIONS
27
The heterogeneous nature of the collared
lizards west of the Colorado River prompted
this study. Samples from known populations
of C. c. auriceps, C. c. haileiji, and C. c. juscus
were used as a base for defining the western
populations. External moqihology and cranial
morphology were compared by cluster, canonical,
and discriminant analyses.
Results of these analyses show a western
complex of four populations which are distinct
from the three base populations of the coUaris
complex. Members within the western complex
are distinguishable with at least 90 percent re-
liabilit\' b\' discriminant anahsis. Because of
intergradation patterns seen in western Sonora,
Mexico, Crotaplii/tus dickersonae is considered
to be Crotaphijtus coUaris dickersonae. The
Great Basin population is retained as Crotaphij-
tus coUaris hicinctores. The population from
southern California and Baja California, which
is distinct and has no demonstrated intergrades
with C. c. hicinctores, is considered Crotaphij-
tus inmdaris vestigium, and the population from
Isla Angel de la Guarda is designated Crota-
phijtus insularis insularis.
The range of C. c. dickersonae is shown to
extend from Isla Tiburon onto the Sonoran
coast opposite the island and from Bahia Kino
north to El Desemboque. Crotaphijtus c. hicinc-
tores extends from the Yuma, Arizona, area
north through eastern California, western Ari-
zona, Nevada, central Utah (west of the Colo-
rado River), southern Idaho, and southeastern
Oregon. The western range of C. c. haileyi is
restricted to central Arizona. The range of
C. i. insularis is restricted to Isla Angel de la
Guarda and C. i. vestigium to a narrow strip
extending along the eastern mountain slopes
from central Baja California, Mexico, to the
foothills south of the San Gorgonio Pass, near
Palm Springs, California.
ACKNOWLEDGMENTS
We express our gratitude to Drs. A. Lester
Allen, Arthur O. Chapman, and Stephen L.
Wood for their helpful suggestions in their re-
\'iew of this stud\-. Appreciation is expressed
to Mr. William Ingram III for statistical sug-
gestions and use of his computer program
decks, and to Dr. Wayne A. Larson, Dr. Melvin
Carter, Dr. Clive D. Jorgensen, Mr. Keith Blake,
Mr. John Taylor, and Mr. Jerr\- Hintze for statis-
tical and computer assistance; to Mr. Donald
Karr, Dr. Kenneth Larsen, Mr. Jeffrey Smith,
Mr. William Ingram III, and Mr. Robert Mc-
Morris, who have accompanied us on extensive
collecting trips during the three years from
1969 to 1971, we express a heartfelt thanks.
Materials were supplied bv the following in-
stitutions and indixiduals: Drs. Alan E. Leviton
and Steven C. Anderson of the California
Academy of Sciences (CAS and CAS-SU); Dr.
David B. Wake, University of California at
Berkeley (UCMVZ); Dr. John W. Wright and
Mr. Arden Brame of the Los Angeles County
Museum (LACM); Dr. Richard B. Loomis,
California State College at Long Beach
(CSCLB); Mr. Allan J. Sloan of the San Diego
Society of Natural History (SDSNH); and Drs.
Charles H. Lowe and Michael D. Robinson,
University of Arizona (UAZ). They were very
helpful during our visits to their respective in-
stitutions and in providing specimens from the
museum collections. We also wish to thank Dr.
T. Paul Maslin, University of Colorado (CU);
Dr. William E. Deullman, University of Kansas
(KU); Mrs. D. M. Smith, University of Illinois
Natural History Museum (UIMNH); Drs. James
A. Peters and George R. Zug, U. S. National
Museum (USNM); Dr. Walter R. Quanstrom,
Northwest Nazarene College (NNC); Dr.
Robert G. Webb, University of Texas at El
Paso (UTEP); and Dr. John Legler, University
of LTtah (UU), and Brigham Young LTniversity
(BYU). Mr. James Davis, Indio, California,
provided us live and preserved specimens from
the foothills on both sides of the Coachella
Valley; for this we extend our appreciation. A
National Defense Education Act Title IV Fel-
lowship provided financial support for the
senior author from 1968 to 1970.
28
Brigham Young University Science Bulletin
APPENDIX I
Key to the species of Crotaphytus
la. Collars on neck absent dorsally and ven-
traUy 2
lb. Collar or collars present on neck dor-
sally and/or ventrally 3
2a. Supraorbital semicircles absent; body pat-
tern with spots but not reticulated — . ivislizenii
2b. Supraorbital semicircles present; occasion-
ally with a vertical black bar on shoulder;
body pattern boldly reticulated reticulatus
3a. Two collars present on dorsum of neck,
first collar often incomplete dorsally and
with spotting in the dorsum between the
two lateral arms of collar collaris
3b. Collars present or absent; when present
reduced, usually incomplete dorsally and
without dorsal spots in the first collar;
body pattern with distinct light cross
bands, often with light elongate spots
between the cross bands iasuZaris
APPENDIX II
Key to subspecies of Crotaphytus collaris and Crotaphy-
tus insularis
la. One or more spots in dorsum of the first
collar; adult males lack a dark pigmented
gular and groin; first collar not united
ventrally; second collar usually extends
onto upper arm (Fig. 3G-H) 2
lb. Spots absent in dorsum of first collar;
adult males have a dark pigmented gular
and groin and first collar unites ventrally
oc nearly so; second collar not extended
onto upper arm 5
2a. Interorbitals form two rows, usually with-
out a fusion of the scales in these two
rows; supralabials usually 11 or more;
gular region of adult males boldly reticu-
lated 3
2b. Interorbitals have one to four scales fused
in the interorbital series; supralabials usu-
ally 10 or less; gular region of adult males
not boldly reticulated C c. collaris
3a. Body dorsum brown with no trace of
green, head cream colored with no trace
of yellow C. c fuscus
3b. Body dorsum green or blue; head yel-
low or light 4
4a. Head yellow or white; if \ellow, not
extending under the chin or caudally past
supraorbital semicircles C. c. haileyi
4b. Head yellow with yellow extending under
the chin and caudally to or just past the
second collar C. c. auriceps
5a. Body dorsum blue, green or gray, with
large light dorsal elongate spots; second
collar length/snout-vent averages from 0.13
to 0.19; number scales from interparietal
to anterior border of first collar 37-43 -.,. C. c.
dickcrsonae
5b. Body dorsum brown or tan, witli trans-
verse bands of yellow, orange, or white;
second collar lengtli/snout-vent length usu-
ally less than 0.14 6
6a. Second collar meets, or nearly meets dor-
sally, usually never more than 10 scales
separate the two dorsal arms of the second
collar; first collar separated by 1-9 scales
(Fig. 3B) C. c. hicinctores
61). Second collar reduced or absent, usually
with distinct white transverse dorsal
bands; first collar separated by 16 to 40
plus scales (Fig. 3A) 7
7a. Second collar very reduced or absent;
fourth toe subdigital lamellae 39-4.5 C. i.
insularis
7b. Second collar less reduced and usually a
portion of it present; fourth toe sub-
digital lamellae 33-37 C. i. vestigium
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LIBRARY
MAR b 1975
HARVARD
LIfI HfStORY AND ECOLOGY OF THE
GREAT BASIN SAGEBRUSH SWIFT,
SCELOPORUS GRAC/OSUS GRAC/OSUS
BAIRD AND GIRARD, 1 852
by
Gary L. Burkholder
and
Wllmer W. Tanner
BIOLOGICAL SERIES— VOLUME XIX, NUMBER 5
DECEMBER 1974/ISSN 0068-1024
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Science Bulletin
LIFE HISTORY AND ECOLOGY OF THE
GREAT BASIN SAGEBRUSH SWIFT,
SCELOPORUS GRACIOSUS GRACIOSUS
BAIRD AND GIRARD, 1852
by
Gary L. Burkholder
and
Wilmer W. Tanner
BIOLOGICAL SERIES— VOLUME XIX, NUMBER 5
DECEMBER 1974/ISSN 0068-1024
LIFE HISTORY AND ECOLOGY OF THE GREAT BASIN
SAGEBRUSH SWIFT, SCELOPORUS GRACIOSUS GRACIOSUS
BAIRD AND GIRARD, 1852
by
Gary L. BiiikhokUi ' ami \\ ilnur \\ . Taniu'i-
IXTHODUCTIOX
Tlie sagebrush swift, Sccloporits iiracinstis
iXraciostis. is tlie most abundant reptile in the
transitional zone of the Great Basin. In this
habitat, being pre\'ed upon bv larger vertebrates
and, in turn, preving upon arthropods, it serves
as an intermediate link in the food ehain. Pre-
vious ecologieal studies ha\'e dealt onl\ with
isolated aspects of the sagebrush swift's natural
history. This studv was undertaken to consoli-
date and to add to our knowledge of this species.
The sagebrush swift is found in western
North America, from northeastern Washington
and west central Montana south to northern Ari-
zona, New Mexico, and soutliern California and
from eastern \\'\()ming to the coast of nortliern
California. Populations are also found in south-
ern California, Baja California, western Texas,
and southeastern New Mexico ( Stebbins, 1966).
One specimen was among those collected b\' thi'
Stansbur\' Expedition to the \'alle\' of the Great
Salt Lake in 1849; this specimen was sent to the
l\S. National Museum, where it was described
h\' Baird and Girard in 1S52.
Stejneger (1893) reported S. g. gmeio.vi/.v to
occupy sagebrush areas at 2,450 m ele\ation in
the White and Invo mountains on the border of
California and Nevada. In 1912 Tavlor obser\ed
that Ne\'ada populations were mainly groimd-
dwellers occupying sagebrush and pin\on-
juniper plant associations. He also notetl
predation of sagebrush swifts b\ Cwtaphi/tiis
wislizenii. Richardson (1915) made a survey of
reptiles of northwestern Nevada and adjacent
territor\' and found considerable morphological
variation between populations. He noted that
sagebrush swifts were abundant on low, sparseh
forested moraines among Arteiiusia tridentata,
Arctostaphi/Ius sp., and Ceanothus sp.
More recently, ^\'oodbur\• and Woodbur\
( 1945 ) observed courtship behavior and gonadal
^Departmenl of Biology, Mt Vernon Nazarene College, Mount V
^Department nf Zoology. Brigham Young University. Prnvn. Utah
cycle. Steiibins (1944 and 1948) and Stebbins
and Robinson (1946) studi(-d territi)rialit\' and
longevitx' in iKirthern California populations of
S. ,g. is,iticilis.
The stomach contents of i^raciosits in Utah
were anahzed in studies by Pack (1921),
Ck'rtsch and WoodburN' (1930). Knowlton and
Janes (1932), Knowlton and Thomas (19.36),
and Knowlton, Maddock, and Wood (1946).
Studies on bod\ temperature liaxc been car-
ried out bv Lieht' (1965). Mueller (1969) re-
corded the mean bodx temperature of ^niciosus
from Yellowstone National Park. Other studies
on thermo-regulation in species of Sceloporns
have been carried out 1)\- Cole (1943), Cowles
and Bogert (1944), Bogert (1949), Larson
(1961). and Brattstrom (1965).
Growth studies on S. f,'. <iraciosus at Yellow-
stone National Park were done bv Mueller and
Moore (1969); however, the results were based
on few indi\iduals. Stebbins (1944) studied the
growth d\namics of California populations of
S. g. gracilis. Tinkle ( 1973) has pro\ ided an
extensive population aiiaKsis of tiie sagebrush
swift in southern Utah.
There ha\e been a nunibi'r of eccjlogical
studies of otiier species of Sreloporiis. A study
by Blair (1960) on S. olivaceus in Texas in-
cludes reproducti\e cycles, population dynamics,
predation, and home range. Mavhew (1962,
1963a, and 1963b) studied the size of home
range, reproductive cycles, and growth of S.
oicutti in California. Fitch (1940) studied the
growth and behavior of S. occidentolis: Wilhoft
and Quay ( 1961 ) reported its testicular histology
and seasonal changes. Other studies on the
western fence lizard have been made by Johnson
(1965-food habits), Davis ( 1967-growth dy-
namics), and Tanner and Hopkin ( 1972)— repro-
duction, growth, and home range). Crenshaw
ernon. Ohio MII5U
84ftnj
Brigham VouNr. University Scienoe Bulletin
(1955) Studied tlie reproductive cycle, habitat,
and growth of S. undulntus; Carpenter (1959)
studied population dynamics. Crisp ( 1964a and
1964b ) examined the female and male reproduc-
tive cycles in S. ct/anogenijs. Zweifel (1949),
Carpenter (I960)," and Goldberg (1970a,
1970b, and 1971 ) studied ovoviviparitv, parturi-
tion and behavior, and histology of the repro-
ductive cycle, respectixclv, in S. jarrovi. Method-
ology and understanding of S. <^. is.rtt(iasus
have been enhanced by these studies.
MATERIALS AND METHODS
Field work was started in the spring of 1970
on a study plot at the eastern edge of the Brig-
ham Young University farm, southeast of
Spanish Fork, Utah. The plot is 1.2 hi-ctares
set with rows of stakes at right angles, each
stake being 12.2 m from other stakes. At the
base of each stake a can trap was buried fliLsh
with the ground level as explained bv Tanner
and Hopkin (1972).
The elevation of the stud\ plot lies between
1,493 and 1,554 m. Tlie shjpe is approximately
40 degrees, facing west and containing a dr\
wash (Fig. 1 and 2A-B). The wash creates
north- and south-facing slopes within the plot.
The substratum on the south-facing slope was
developed from a large talus slope which is
light in color and of a fine gravelly nature.
Small rocky outcroppings and large solitary
boulders occur on both slopes of the wash. The
area soutli of the wash (north-facing slope and
southern end of plot) has a more humus soil
but is still rocky and has a surface covering of
small gravel in some areas.
Two davs of each week were devcjted to
checking traps and two to noosing and return-
ing lizards to the place of capture. Each lizard
Rock Ledge
'y°°fj'-. Ook -Maple
North
Top of Wash NE Side of plot
Bottom of Wash
Top of Wash SW Side of plot
• « o
Fig. 1. Outline i)t stiuK plot with doininaiit \ (■•^(■talioii and topograph) iiulitatcd, liVl' I'arni, Sjianisli i'^irk, Utah.
Biol<)Ck:ai. Series, Vol. 19, No. 5 Liric Histohv of Scelopohus (inAciosus Chacio.sils
J^,
3f'
M
^ .41?
>. ^'
-^rM?^^-^'^
pb-
m
'i:^.'
%"§j!^^
W% *
(!i_
•*&*■'
*^
ii'\
Fig. 2A-B. Study plot hahifat features. A depicts the south-faeini; slope and B the north-facing slope.
RniciiAM Yc.UNf: Unufhsitv Scikxck Bulletin
was marked by toe clipping and its weight, size,
sex, and external appearance were recorded, as
well as body temperature at the time of capture.
Air temperatures at approximately 3 mm and 1
m above the substrate were taken soon after
noting the lizard's temperature. Notes on be-
havior, activity, and feeding were also recorded
while working in the study area. A hydro-
thermograph was used to record weather data
on the study plot, and United States climato-
logical data were obtained from a station located
2.5 km north of the study area. A vegetation
analysis of all plant .species occurring on the
plot was made in 1971, while flowering species
were collected throughout the year.
Lizards were collected throughout Utah
County and parts of Salt Lake and Wasatch
counties. These lizards were autopsied in the
laboratory to analyze reproductive stages,
weight of gonads, stomach contents, and fat
body weights; these data were compared with
external measurements taken from study plot
lizards. Observations of behavior in the labora-
tory were made by keeping lizards in two boxes
partly filled with sand, rocks, and pieces of
wood, creating a suitable habitat. Eggs laid in
captivity were incubated in the laboratory at
approximately 28 C.
Male testicular cycles and meiosis in female
ovaries were analyzed by histological sections.
Stomach contents were analyzed with a stereo-
scopic microscope, and arthropod volumes were
measured by water displacement in a 10-cc
graduated cylinder. All statistical tests of results
were at the 5 percent level of confidence. The
statistical tests used were chi-square, analysis
of variance, and the paired f-test.
Description of the Study Area and
General Observations of the Research
Habitat
Stud\- plot vegetation included annual and
perennial hcrl)s, shrubs, and trees. The domi-
nant plant association consisted of bigtooth
maple, Acer <^mndidentatum; Gambel oak,
Quercits <s,iimhe]u- and sagebrush, Artemisia
tridcntata. Distribution of the oak and maple
is depicted in Fig. 1. Subdominants, abundant
in some areas of the study plot, include snake-
weed, Gutierrezia surothrae: Pctrodoria pumila:
blue bunch wheatgrass, A<:,ropijron spicatiim:
muttongrass, Poa lon>:,ilii;ida: needle grass, Stipa
comato; and junegrass, Koeleria cristafa. Some
perennial and annual herbs were balsam root,
Balsamorrhiza sufiUtattd: and two species of
bromegrass, Bromiis spp. The dominant and
subdominant plant types grew in clusters, with
ample open spaces (Fig. 2A-B). This type of
spatial arrangement appears to be necessar)' for
S. <i. graciosiis to occupy an area.
Collection sites throughout Utah County
where lizards were taken usually had the same
general environmental features as the .study
plot. Most had Gambel oak, bigtooth maple,
and sagebrush as associated dominants. Some
collection sites supported ]uniperus osteospcrma
or sagebrush onh'. Elevation of collection sites
ranged from 1,.'371 to 1,981 m. South-facing
slopes usually provided the best sites for col-
lecting, primarily l^ecause these habitats were
more often chosen by ^raciosiis ( Fig. 2A-B ) .
Summers are hot, with few overcast days
from mid-May to mid-September and with aver-
age daily maximum temperatures ranging from
.30 to 37 C for July. There are approximately
ISO or more frost-free days from April through
September. Howe\er, irregular cold fronts may
periodically reduce activity in March through
the first of May and in late September and
October. Precipitation and temperature on the
study plot are summarized in Fig. .3. The
average seasonal temperatures were similar dur-
ing the years of this study and the precipitation
averaged about 10 inches (25.4 cm) during the
activity season, 1 inch (2.54 cm) per month.
The least precipitation occurred in late June,
July, and the first half of August. The precipi-
tation total was lowest in 1972 and highest in
1970 (March through August).
Emeriiencc
In 1970, 1971, and 1972 the first lizards
emerged when average daily temperatures
reached 10 to 13 C (Fig. 3). These early ani-
mals were juveniles of both sexes and adult
males. The first adult male emerged 19 days
before adult females in 1970 (22 March/10
April), nine days earlier in 1971 (21 March/29
Mareli), and 21 days earlier in 1972 (5 March/
25 March)— an average of 16.3 days between
male and female emergence. Males were caught
soon after emergence ( indicated by the moist
mud on the IxkIv).
Seasonal and Daily Lizard Temperatures
Mean preferred body temperature for study
plot adults taken for the season, June to Sep-
tember, was .32.6 (19.0-38.8) C, juveniles .33.1
(21.0-38.4) C, and hatchlings 31.2 (26.0-35.4) C.
Hatchlings did not tolerate temperatures much
above 36 C. Temperature observations from
the field for hatchlings during August and Sep-
tember rangi'd from 26.0 to 35.5 C. Periods of
ISiouKiicAL Skiues, Vol. 19, No. 5 Line Histokv ok S( klopohus Ciia< losi s Ciiac idsus
Fi
•'S
3. Precipitation (columns) and tcnipfratinc I lines ' l)\ montli tor sliiiK plot tor 1970 through 1972. Shaded
columns represent 1970, \ertic,il lines 1971, anil horizontal lines 1972.
20 to .jO minutes in a paper bag at teinpi'iatiiics
near or ahoxf 38 C provi'd fatal, .simscstiiiy; a
lower critical thermal ma.ximiim for hatcliliiigs
than tor adults.
Lizards emerged from overnight retreats
with bodv temperatures of 16 to 21 C. Basking
range was 21 to 28 C and normal activit\ range
was 28 to 37 C. Adult and juvenile lizards were
able to tolerate temperatures of 38 to 40 C. The
critical thermal maximum in the laborator^â– was
43 C for an adult male and 44 C for an adtilt
female. They were captured in March and
April and kept in captivity at temperatures ot
24 to 27 C.
In Fig. 4, the average temperatures of males
and females are compared to temperatures of
air 3 mm and 1 m above the substrate.
During periods of intense heat in July and
.\ugust, lizards could lie found only in the oak-
maple thickets and under sagebrush, rocks
(crevices and overhangs), and other fonns of
cover. Some remained under cover until the
heat of the day passed, while otliers in the oak-
maple thickets were more active. The contrast
between periods of low and high actixitv l)e-
tween 1000 and 1200 hours in Juh' and August
is represented in Fig. 5. Soiue lizards were at
the basking areas before sun up during June,
July, and August. This activity was seemingly
caused 1)\' early morning ligiit and iiigh tem-
peratures ( 1-5-17 C).
Ddiltj and Seasonal Activili/
\ summary of the daily actixitv for study
plot lizards during the 1971 activity season is
diagrammed in Fig. 5. High temperatures for
40
35
30
25
CD
d
6
^15
10
5
Adults
M
M
Fig. 4. Mean temperatures of adult lizards and of air
(â– 3 mm and 1 m) abo\e substrate. Dashed line re-
presents lizards; solid line, air (3 mm); dotted Une,
,iir dm), 1971.
Brigham Young llNiviiiiSirv Scienck Bullktin
Fig. 5. Number and age ot lizards observed, ]5lolte<l
against time of dav, throughout tlie activity season
of 1971 and March 1972. Solid line represents
adults; dashed line, juveniles; dotted line, hatch-
lings.
air at 1 m ami 3 mm above the .substrate in late
June, Julv, and Augu.st ranged from 34 to 40 C
and from 43.2 to 45.0 C. Temperatures above
50 C for actual .substrate surface were noted
freciuentlv during the afternoons of the warmer
months. Our data indicate that adults and
juveniles have the same daily activity through-
out the year, with the activity of juveniles ex-
tending into September.
Local Wvatlicr Conditions
Periodic cold fronts in March, April, and
early May, if severe enough and over a period
of several days, stopped all activity in the field.
One or two days of wanning were recjuircd be-
fore full activity resumed. Heavy thunder-
storms during June, Julv, and August stopped
;ictivity periodically. Lizards reappeared as soon
as the sky was clear. They basked for 10 to 20
minutes, or until body temperatures rose suf-
ficiently. If a storm came late in the afternoon,
they- did not reappear until the next day. Cold
fronts in September and October interrupted
actiyit^• in much the same manner as earlier
in tlie \'ear.
Slo))c Preference
Tile number of lizards observed per hectare
t)n each slope of the plot is summarized in Table
1. Adults and juveniles preferred the south-
facing slope throughout the activity season.
Hatclilings. after emergence, followed the same
trend. Little acti\'it\ occurred on the north-
and west-facing slopes.
Hibernation
Hibernation of adults began in late August
or earh September and was recognized when
indi\'iduals were no longer seen at their basking
stations. Ju\eniles remained active until mid-
or late September and hatclilings remained ac-
tive until mid-October. One plact- of hibernation
was foimd when an adult male was sighted on
10 March at the opening to a rodent burrow.
He was completely covered with mud, indicat-
ing a recent emergence.
Feeding Behavior
On 28 April 1971 an adult female was
observed climbing on sagebrush and feeding
iunong the foliage. On 13 July 1971 an adult
male ran a few feet from the cover of a small
oak Inisli, grasped a large grasshopper, ate it,
and tlien returned to the oak bush. The lizard
sliook the prey by lateral movements of the
head before swallowing it. On 25 May 1971
an iidtilt male was observed moving up the
trunk of an oak tree and feeding on small in-
sects. Though specimens ate both large and
small prey while in captivity, they rubbed the
sides of their heads on the substrate while
eating the large prev.
Ais,i:,ressive Behavit i r
.\ggression in the field and captivity was
observed in man\ forms in all age classes and
in both sexes. An example of such aggression
between territorial males follows. Chased into
the territory of another by the observer, the
intruder was immediately challenged by the
resident and a melee of biting and rolling en-
Biological Sehies, Vol. 19, No. 5 hut. Hisronv of Scei.opohu.s Ghacio.sis Chaciosu.s
Talile 1. Preference of adults, juveniles, and hatchlings for north-, south-, and west-facing slopes of the study
plot. These values are ratios (number of observations) by month, which are based on the total number of
observations on each slope divided by the area of each slope.
.Adults
S
W
Juveniles
N S
W
Hatchlings
N S \V
March
.\pril
M.iy
|une
liilv
August
September
October
6.2
8.9
.0(
4.3
15.7
1.7
5.5
16.4
25.3
3.7
3.2
8.9
2.9
17.4
38.4
9.6
15.3
46.6
13.3
22.0
26.7
5.0
18.6
19.9
14.0
4. .3
1.5.7
3.3
6.5
22.6
13.9
2.1
.035
1.0
2.7
7.0
17.5 43.2 40.7
12.0 101.4 40.0
4.3 55.5 14.0
sui'd, followed by ;i breakaway and .standoff
of about fi\'e ccntiiiictt'is. Bodie.s were coiii-
prcs.sed latciallv to .sliow the blue sides. Each
stood high on ail linibs with the back slightly
arched and the dewlap lowered. Laboratory
obsi'rvations of territorial males re\ealed that
the resident was alwavs able to vantjuish the
intruder.
The followinti description cin'ers all major
actions <)bser\ed in captive interactions. The
intruder was placed in the bo.\ of the resident,
whereupon he immediately stood high. The
resident moved toward him from the side, mov-
ini;; his head to either side as he approached,
as the intruder moved awa^â– sidewavs. Then
the resident positioned himself parallel to the
intruder, facing the opposite direction, and stood
high on both limbs, compressing his sides to
show the blue and lowering his dewlap. He
then made a ([uiek strike at the intruder, who
rail, followed closelv by the resident. When
tiie\' stopped, the intruder assumed a position of
passi\eness with his head and tail lowered and
with his liodv pressed low to the ground. The
resident repeated the parallel body stance and
then mo\('d at the intruder, who again ran. This
was repeated three times ( Fig. 6A-B ) . Field
(>liser\ations in which a biting melee occurred
often resulted in a temporarv standoff until one
was vanciuished. Juvenile intruders were not
tolerated bv adult males and were iinmechatcK'
chased from the territors'.
Females also reacted aggressivelv toward
males aftt'r they had copulated. Such females,
wiien approached by a male, flattened the body
laterally, lowered the dewlap, pointed the tail
upward and arched the bodv (Fig. 6C-D).
Sometimi's a female would move toward tlK>
male in a challenging postiue and he would
move awa\ . Otlier times the female assumed
tliis pose at tlie mere approach of the male and
hopped on stiff legs toward him.
Baskiiifs, and Panting
All age classes were observed basking on
the substrate, rocks, boulders, and branches of
sagebrush, under bushes, and along the main
branches of oak and maple trees. Basking usu-
all\' l)egan in the morning just after the ravs of
the sun had spread over the study plot. The
lizard's ventral surface was in direct contact
with the substrate. In captivity, basking be-
ha\ior under sun lamps was essentially the
same. On several occasions lizards were ob-
served panting, opening the mouth and going
through the motions of breathing rapidly. In
captivity, panting occurred at room tempera-
tures of 37 to 40 C; field obser\ations were
similar.
Pusliups
Sceloporus iirciciosiis was not included in
the behavioral studies of Purdue and Carpenter
( 1972a and b ) . Because of the specialized na-
ture of their study, we have not attempted to
duplicate it for is^raciasus. Our data are general
and provide onh field obser\ations. Pushups
were observed in all age classes and in both
sexes. Thev appear to be most important in
courtship and territorialitv behavior. Males of-
ten climbed on a rock or some other object and
then engaged in one low pushup and stopped;
this was sometimes followed by a second (high)
and a third (low) one. Other pushup setjuences
are possible, but the above is the most normal
and mav be repeated several times. During the
breeding season, males did pushups as they
mo\ed through their territories. When a female
was encountered, he performed while slowly
approaching her. At this time of the activity
season, or while in the presence of females or
other males, pushups were perfonned very high
on the forelimbs. As he approached to about
7 to 10 em from the female, the sequence was
one low and one or two high pusluiDS.
BiiiciiAM Young UNiMiHsnv Scienck Bulletin
/i'**'"
w
â– :^A
>K—^
B
Fig. 6A-D. Annres.siN'i' Iielun ior iK'tween males (A aiul B) .iiul bctuotn tcmalfs and males (C and D)
BiOLocicAL Skuies, \'ol. 19, No. 5 Lirio IIistohv ok Scei.oi'ohus (..iwr.unvs Chaciosus
r^j^'"" â– ^. %«<>â–
â– <-'â– â– â–
Fij;. 6 A-D (Continued)
10
Brigham Young Univehsity Science Bulletin
Reproductive Behavior
Mating behavior was oliserved eight times in
the field and three times in captivit) . On 4
June 1971 a female moved a distance of approxi-
mately two feet through a bush after being
disturbed. It was then noticed that she carried
a male with her. He had hold of her neck, was
in a dominant position oxer lier body, and was
doing partial pushups while holding on to her.
The posterior part of his body was then brought
under the female, putting the \ents into contact.
The hemipenis was inserted into the cloaca as
the vents came into contact. There was no
movement of the posterior part of the male's
bodv and the hemipenis was held inserted for
approximate! V eight sc-conds, after which the
female broke away, lea\ing the male stationary.
He pursued her persistently for a few feet with
the tail raised and the h(-mipenis still everted.
It took approximately one minute for the liemi-
penis to be retracted. The male exiiibited an
erratic pursuit of the female in contrast to
normal running and courting. As males ap-
proached females in the field, they did pushups
and rapidh- boljbed their lieads. Females were
grasped an\ where from midbody to just behind
the ear, and males remained in a dominant posi-
tion over the female for S-10 seconds Iiefore
attempting to copulate.
Capti\e reproductive behavior was similar,
but with the following \'ariations. When a male
first noticed the female, he approached her with
liis head bobbing. The female was passive, not
mo\ing and remaining low to die ground. Tiie
male grasped the female twice before taking a
firm grip on her shoulder, then assuming a
dominant position o\er lier. .\ few seconds
elapsed without mo\'ement before copulation oc-
curred. The hemipenis contracted rh\thmically
in waves of approximately two per second. After
a short period of separation and reintroduction,
the male no longer approached the female. The
basic posture of the male and female during
copulation is shown in Fig. 7A-C. One male
rubbed the vent on the substrate after the hemi-
piMiis had been retracted. In a few instances the
male did not perform pushups or head bobbing
before he grasped the female. Actual times of
copulation (contact of genital organs) in cap-
ti\itv ranged from 5 to 10 seconds. The tail of
the female in each ease thrashed during copu-
lation.
A complete sequence of egg laying was not
observed; however, laboratory observation re-
vealed that females dig into the substrate 15 to
20 em, lav eggs, and tluMi fill in these lioles as
they return to the surface.
Food Habits
The contents of .39.3 stomachs were examined
in 1971 and 1972. All age groups throughout
the activity season for both years were included.
Data in Table 2 indicate the major families of
arthropods that were preyed upon by S. g.
iiraciosus.
Data indicate a seasonal fluctuation of prey
items taken bv two age groups in 1971 at the
ordinal level. In both \ears lizards were op-
portunistic in their feeding habits, with monthl)'
changes in diet following tlu- changes in abun-
dance of different orders of arthropods. In each
age group, in both years, the priority of num-
bers and consistencv of orders eaten in each
month were essentiallv the same.
The number and size of prey items con-
sumed each month by adult males were notice-
ably different from those consumed by females.
For example, of 1,.3,S3 Hymenoptera, 991 (pre-
dominantly ants) were consumed by females;
and of 284 Coleoptera, 149 were consumed by
males. With regard to total numbers of all
orders eaten by females, 1,494 of 2,157 were
consumed, indicating that females consume a
larger numlx r of items than males. When con-
sidering \olume, the order ot priority for food
items changed. For example, H\menoptera
were the most fre([uent stomaeji item; however,
i]i adults tiiev made up 5.47 cc, while Coleoptera
made ujj 7.26 ee. Total x'ohnne was 14.00 cc
for males and 15.70 ce for females. These \ol-
umes indic;ite that, while females eat more,
thev eat smaller prey; total \-olume consumed is
not signiticauth' different.
OntD^cuctic CVi(/iii;rv in (Uihir diul Color Pattern
.•\ccording to Stebbins (1966), hatchlings
(24-27 mm S\'L) lunc the eharacteristie color
pattern of adults. In the central Utah popula-
tion we find that m;iles at 24.5 to 32.0 nun SVL
l)egin to show faint blue green \'entral belly
patches. Bv the time thev a\i'rage 34 mm SVL
in mid-September, the \'entral patches are edged
with black. The blue is still blended with the
green and is not \ et as dark as in adults. Males
reaching .36 to 39 mm S\L show traces of
throat barring. Females mav show a faint out-
line of \-entrai patches between 32 and 35 mm
S\'L.
Juxenile males eoutiiuu' to dex'elop darker
blue p;itehes \entrall\-, and b\' late April and
.May (at approximate 1\- 40-45 nun SVL)," the\-
have essentialh the same color pattern as adult
males of 55 to 60 mm S\ L. JuNcuile males that
;ue over 47 mm S\'L in nnclNhn are essinfialb
of the same brilliance as adults.
BioL()r.ic:Ai. Sehies, Vol. 19, No. .5 Lii-i; Histohv oi' SoKioronus OitAciosus ()iiac:i()sus
11
TABLE 2. Analysis nl the stomach contents of 183 S. g. gniciosus collected in 1970 and 1971 listed bv order
and family from highest to lowest in nnmber of each.
Taxon
1970
Total
Taxon
Hi/menoplera
2806
Formicidae
86
Unclassified
36
Andrenidae
28
Eulophidae
22
Halictidae
24
Other
3002
Total
Colcojttera
126
Curciilionidae
64
Chr\somelidae
41
Coccinellidae
35
Scarabaeidae
28
Unclassified
20
Elateridae
15
Carabidae
24
Tencbrionida<'
353
Other
Total
IsoplCKl
53
lUiinolerniitidae
40
llcmiptcra
10
Unclassified
1971
Total
Hi/incuoptcra
Formicidae
Andrenidae
Eulophidae
Unclassified
Halictidae
Other
Total
Colcoptcrii
Curculionidae
Unclassifcd
Carabidae
Chrssomelidae
Elateridae
Coccinellidae
Tenebrionidae
Other
Total
Arachnoidea
.\raneida
Unclassified
Lycosidae
Other
Total
liciniptera
Lygaeidae
Unclassified
Other
Total
Hoiiioptcni
Cicadellidae
.â– \phidae
Mcmbracidar
Fiilgoridae
Unclassified
Total
Oithoptcra
Locustidae
Grvllidae
Total
hoptcra
Rhinotennitidae
Lcpidoptcra
Unclassified
Other
Total
Dipteru
Unclassified
hopodn
iru hoplcKi
Other Orders
Grand Total (1970)
103
39
23
16
78
30
15
11
10
5
~7\
47
Lvgaeidac
Other
Total
Aiiiilniitidvd
Arancida
Unclassilicd
l.ycosidae
Acarina
Total
lluimiplcia
.â– \phidac
Cicadellidae
Unclassified
Other
Total
2
l')ipl<i,i
49
Chironomidae
Other
43
Total
34
10
44
39
Lcpidopleid
Unclassified
Ceometridae
Total
Oithoplcra
Locustidae
Other
15
Total
4
MiirocunjphUi
Machilidae
7
Other (Orders
3807
Grand Total (1971)
1981
39
20
16
10
41
2107
131
53
48
45
33
32
27
12
18
399
237
77
57
23
1.57
84
17
28
129
59
28
17
7
TiT
27
18
36
81
56
3
59
51
6
57
15
6
3367
12
Bui(;ii.\M VouNi; Univebsitv Science Bulletin
T
/
Fig. 7A-C. Copulation sequence of lizards in capti\itv irulicating niajm posture of tht- male and female
Biological Seihks, \'ol. 19, No. 5 Like Histouv of Sciii.oroiius C'.raciosus Graciosus
13
iMg. 7 A-C (Continued)
Jmc'iiik' tcMiialfs lost- the Dutlinr of a \t'iitral
pattern as \.\\v\ %to\\ and do not become orange
lateralh' until thev become breeding adults.
When adult females emerge ( March or
.\pril), there is no nuptial color laterally. The
\'enter is white and the normal dorsal and
lateral color pattern is present ( Stebbins, 1966).
In April and May, while the female cycle is
undergoing \itellogenesis and the ovarian eggs
are approaching o\ulation (6.5-7.0 mm diame-
ter), the\' bt'gin to take on the orange colora-
tion laterally. This color may extend from the
hi'ad to the groin region and even onto the tail.
Females reached their most brilliant nuptial
colors wiiile earning o\iducal eggs, between
S and 24 June 1971.
In females tlie \entral patches are usualh
lime yellow and extend to the throat and the
antero\entral part of the tail. This lime \ ellow
suffusion follows the same sequence as the
lateral orange areas, although some females ma\
JKU'e the lateral orange, but with the \ciitral
\ellow greatly reduced. After mid-Jul\- tlie
nuptial colors are lost. Two exceptions to this
wi're a pair of females captured on 27 August
1970 and 20 August 1971 which were \ery
orange, but palpation revealed no oviducal eggs.
Adult males emerging in March luue the
characteristic color and pattern described by
Stebbins (1966). The most brilliant period for
the latero\'entral blue and black is April through
July, though some males may still be brilliantly
colored in August. There is also considerable
blue green in the lateral scales of the shoulder
region and posteriorly onto the base of the tail.
There is no blue green on the tail where re-
generation has occuiTed.
Parasitism and Predation
Stomachs were remo\ed approximately 3 to 5
mm below the pyloric sphincter. While remov-
ing stomachs from 690 specimens, one tapeworm
was found in an adult male and one in an adult
female. These were identified as Oochoristica
scclopori: no nematodes were found. External
parasites were either ticks or mites (Acarina)
and were found on specimens from West Can-
\()n, Utah Coimt\'.
Examination of 690 preserved specimens
from Utah County and parts of ^^'asatch and
Salt Lake counties, re\ealed oiih two cases
where the eardrum of the lizard was discolored
and a heavy growth had replaced the thin trans-
parent structure. Abnomialities were not ob-
14
Brigham YouNr. UNivEHsrrv Science Bulletin
served intemallv on autopsied specimens. Ani-
mals from the study plot were checked ex-
ternally only and no infections other than an
occasional swollen toe from identification clip-
ping or natural loss were observed. Infection or
swelling at the point of tail breakage was not
observed.
Predation was not witnessed in the field.
Utah Countv records indicate the niglit snake
Hypsiglena torqtiata as a predator. Possible
predators in sympatric areas may include pre-
daceous reptiles, hawks, and mammals.
Ontogenetic Changes in the Tail
Ten arbitrary size classes were chosen and
the mean SVL and tail length were taken on 20
animals in each size class except for the smallest
and largest sizes. The mean SVL of each size
class was plotted against the mean tail length
(Fig. 8), contrasting males and females through-
out their growth from hatchlings to adults.
Tail Breakage of Study Plot Lizards
Hatchlings, juveniles, first-year ( after second
hibernation), and older adults were analyzed in
1970 and 1971 for tail breakage. Males and
females were contrasted throughout the activity
season. These data can be summarized as fol-
lows: adults, males 14 of 69 (21 percent),
females IS of 116 (15 percent); juveniles, males,
10 of 97 (10.3 percent), females S of 131 (6.1
percent); hatchlings, males, 5 of 247 (2 per-
cent), females 10 of 231 (4.3 percent). There
was an increase in breakage from hatchlings to
adults (3 percent, S percent, IS percent). One
specimen from the field and one from the stndv
plot had bifurcated tails.
The regenerated portion did not ha\e the
same color or color pattern as an intact tail.
This was strikingh' apparent in males with the
blue green on some lateral scales which extend-
ed onto the proximal half of the tail.
Average Size of Females at Sexual Maturity
Sexual maturity in females was determined
by noting the development of the gonads. The
main criteria used were the presence of yolking
follicles in the ovaries, o\'iducal eggs, or cor-
pora lutea during the reproductive period. Most
reproductive activity took place before 1 July;
therefore, 1 July was used as the date females
were considered mature for that particular year.
All females 53 mm S\ L or larger were consid-
ered mature. Studs' plot females found with
eggs (based on palpation and color and weight
of females) were in this range. Three females
which were 47, 48, and 49 mm S\'L, autopsied
on 28 May 1970 and 7 June 1971, had small
volking ovarian eggs 2 or 3 mm in diameter.
JEgg development in these females was late in
the season compared to other larger females.
If these eggs were exentuallv laid, it would
have been late compared to tlie average time
taken to reach o\ulaton- size and egg laying.
Such individuals made up onlv 1 percent of the
breeding population.
Follicle Growth
As females grew from hatchlings to adults,
the average number of unyolked follicles in-
creased in number and enlarged. Ovaries were
visible as small white tissue masses in the small-
est hatchlings captured (25 mm SVL). The
August hatchlings between 25 and 33 mm SVL
may have none; ]iowe\er, 70 percent had one
to four follicles (Table 4). September and
October hatchlings, just prior to hibernation,
showed an increase in size and number of fol-
licles. Dtuing hibernation there appeared to be
little growtli in the ovaries, although we did
not examine hibernating lizards. In the spring,
follicles continued to increase in size and num-
ber until June, then dceri'ased in Julv and in-
creased again in .\ugust (Table 3).
Adult females underwent seasonal changes
TABLE 3. Number of unyolked folhcle.s in hatchlings and jintiiiks In mouth. 1970-71. .Ml measiiremenl.s are
HATCHLINGS
SVl.
March
Dia.
-April
Foil. No.
SVL
May
Dia. ' Foil. No.
SVL
June
Dia.
Foil. No.
Mean
Range
Number"
JUVENILES
.
Mean
Range
Number"
.34.8
30-37
11
.40
.1-1.1
11.3
7-13
40.0
35-45
11
.70 16.4
.1-1.7 9-23
43.2
10-50
15
.80
.1-2.0
15.8
8-22
"Number of specimens examined.
BiouxacAL Series, Vol. 19, No. 5 Life Histohv oe .Scelopohus Guaciosus Ghacio.sus
15
Mean Tail Lengfh
Fig. 8. Mean -.iiout-vent length of sagebrush suitts
plotted against mean tail length (mm) for 1970-71.
Females are represented by solid line and males by
dotted line.
in the number and .size of unyolked follicles
(Fig. 9). The total number of follicles may
change seasonally, with the lowest number oc-
curring in July after the eggs are laid ( Table
4). Our data indicate that from March to June
a decrease in size and number of unyolked fol-
licles above 1.5 mm occurred. Between 1 Julv
and 15 August only five follicles above 1.5 mm
were seen in 1971 and only one in 1970 (Fig. 9).
This indicates that follicles show a reduction in
size in Juh' and the first half of August. From
mid-August until hibernation there was an in-
crease in mean follicle number and size. In
September fcjllicles ranged between 1.9 and 2.0
mm, appeared to be yellowing, and perhaps were
beginning vitellogenesis.
In Fig. 10 the total number of follicles in
each of five lizard size classes are compared,
indicating that unyolked follicles increased in
number as females became larger and older.
in millimeters.
Meiosis
Serial histological sections revealed that
follicles went through meiosis both in the fall
(August-September) and spring ( March- April).
Eleven of the first 12 females collected from
29 Marcli to 29 April 1971 were undergoing
meiosis. ( )f the last 13 females captured 24
August to 23 Septemljcr. 12 were undergoing
meiosis. The size range (determined from
slides) of follicles undergoing meiosis was .5 to
1.5 mm— axerage size in the fall being .96 and
in the spring JK'ing 1.05.
Yolk Deposition
Females were not yolking follicles in August.
In September follicles were slightly larger and
from criam to light yellowish in color. All
began soon after emergence in the spring. Yolk-
ing eggs continued to increase in size up to the
TABLE 4. Fluctuation in the number of unyolked
follicles in the (uaries of adult females, 1970-71
Number of
Month
Follicles
Lizards
Range
Mean
March
1970
1971
46
â– 1
20-26
23.0
April
197(1
78
4
18-21
19.5
1971
323
15
7-31
21.5
Mail
1970
1.34
6
10-.3()
22.3
1971
.541
27
13-2S
20.0
June
197(1
lOS
5
18-28
21.6
1971
1244
74
4-27
16.8
Jalii
1970
322
18
11-24
17.9
1971
584
37
9-24
15.8
Au..
1970
466
22
12-31
21.2
1971
Svpl.
1970
354
20
12-.35
17.7
112
5
20-25
22.4
1971
219
10
15-33
21.9
HATCHLINGS
8\'L
Julv
Dia. '
Foil. No.
SVL
August
Dia. Foil. No.
SVL
Sept.
Dia.
-Oct.
Foil. No.
Mean
Range
Number"
27.8
25-33
4
.20 2.5
.1-.4 1-4
.33.4
31-41
17
.34
.1-.9
7.8
3-21
JUVENILES
Mean
Range
Number"
43.2
40-53.5
23
.70
.1-1.5
13.7
7-24
.52.1
50-53
.74 1.5.7
.1-16 12-21
16
Buu^iiAM Young Umvehsitv Sc:ience Bulletin
70 71
Sept
Fif;. 9. Mean nuiiilier of iiiiyolkcd lollicles gicatt-T than
1.5 mm in diameter in adnlt fi'males plotted against
30-day intervals throughout the activity seasons of
1970 and 1971. The S represents the size range of
follicles in mm.
time of oMilation. Tlic .siiialli'.st (liaimti-r for
definite \c)lk deposition (l)y eolor ) ranged from
1.5 to 2.0 mm (late Mareh and early .April ),
which is clo.sc to the approximate .size of meiotic
follicles (mean 1.30 mm). Yolking; eggs in-
creased from an axcrage of 1.75 mm in late
Mareh to a peak of 7.09 mm in diameter in the
first half of Jnne, with a range of 6.5 to S.5 mm
jnst prior to oxulation. This was an increase of
approximateh' .07 mm per day. Weight followed
the same trend as si/e. Eggs less than 2.0 mm
in diameter weighed less than 0.1 g. In Jnne
ovidneal eggs averaged .21 g per egg (N-65)
with a range of .14 to .28 g per egg. These were
from clutches ranging from two to se\en eggs.
Yolk deposition occurred in the populations from
29 March to .3 Julw a period of 97 davs.
Ovulation
Eggs passc-d into the ci'olom and from there
into the oviducts. ()\a in the ceolom were flac-
cid and the onh' structiu'e holding the volk was
the vitelline membrane. One female ( BYU
3338, 64 mm S\'L) found on 10 Jun<' 1971 con-
tained some eggs recenth ovulated, two in the
ceolom, and others entering the o\iducts— a total
of seven eggs in the clutch. Judging from the
delicate nature of ovulated eggs, it would seem
that females might reduce their acti\ it\ to pre-
vent breakage; breakage was not observed. A
female captured on 29 \hi\- 1970 had six eggs
( three per o\'ary ) and all were between 6 and 7
mm in diameter. In addition, there were two
eggs (5 mm each) free in the ceolom. Thc\'
were grevish and slighth' crenated in appear-
ance, yet pliable and with discolored volk.
Ovulation in graciosus took place approximateh-
between 2 June and 3 Julv 1971, the 32-day
period between the first and last oviducal eggs.
The majoritx' of females were undergoing ovu-
lation from 2 June to 24 June (22 da\s), at
which time oviducal eggs and the first cijrpora
lutea appeared. In 1970 the development of
eggs was approximateh' 15 davs later than in
1971. The actual time re(|uiied for each stage
( \()lk deposition, etc.) remained fairly con-
stant in terms of acti\(' davs.
Matinii Tiiiic in the Field
Copulations wire obser\cd in the field be-
tween 21 Ma\ and 4 Jimc' 1971 and in captivity
between 2 June and 9 June. At this time males
and femali's wi'rc- seen together in the field in
contrast to the periods before and after the
majoritv of copulations. Observations of field
and captix'e indi\ iduals from 21 May to 9 June
suggest that the majoritv copulate over a period
of approximately 20 days. This time period may
extend earlier into May but probably not later
into June, except in Ncars with nnusualK' late
springs.
20
18
16
14
â– 12
Z
3
Z
Z
<
6
lO
?30 31-40 41-50 51-60 61-70
i'li^. 1(1. l'n\(>lkcd o\arian lollicles plutliil .igainsl
M l-\iiit lensith. Hatclilins's to adults.
BioLocicAL Sf.iues, Vol. 19, No. .5 Life Himohi of .Scflopouus C^hacuosis CIhaciosils
17
E^i; (incl Clutch Size and Weii^hts
Fflllon in^ Oviposit ion
Cliittli size and S\'L for fciiialcs laving
eggs in capti\it\' are listed in Table 5. Twelve
clutches (6i) eggs) were weighed and measured
within one to five davs after oviposition. The
mean size of recently oviposited eggs was 1.3.4
\ 7.9 mm witli a range of 12.0 x 6.5 to 15.5 x
9.0 mm. The mean ^^'eight was .48 g per egg
with a range of .30 to .76 g per egg. Compared
to the weights of oviducal eggs, these recently
laid eggs averaged .20 g more. They were
also larger in size, 13.4 x 7.9 mm as compared
to 12.0 \ 7.1 mm. A few clutches were in moist
sand for a few da\s and undoubtedly increased
in size and wcigiit b\' water absorption.
Size and Wciijit Cd\(inii,cs in
Gravid and \()n!j,iavid Females
Tiie size and weight changes in females be-
fore and after la\ ing and the time that elapsed
between the periods are listed in Table 6.
Compared to the average weight of clutches,
the average wcigiit per egg was .48 g (Table 5).
Tliese data also indicate that females that have
laid eggs with an intervening period of 10 days
or less show no difference in size (SVL) but
weigh 1.5 to 3.0 g less.
Clutch Size, Cdutcltcs per Year, and Comparisons
of Left and Rii^ht Ovaries and Oviducts
Clutch size in <i,raciosus was determined
after the metliod used by Tinkle (1961). The
nunilier of o\arian eggs, oviducal eggs, and
eoipora lutea in ixith ovaries and oviducts are
listed in Table 7. Clutch size was 6.03 eggs per
female. There were slight but not significant
differences in the number of ovarian eggs, ovi-
TABLE ,5. Number and weight of ova 1 to 3 days
after laying (1972) for 12 clutches laid in the
laboratory.
.A\*erage
Number of eggs
Weight of
Wcigiit
in Clutch
SVL
Clutch
per Kgg
8
63
4.80
.600
5
62
2.28
.456
6
62
3.12
..520
6
65
2.05
..341
7
64
3.39
.484
5
62
2.72
.544
5
59
2.31
.462
4
59
2.26
.565
6
61
2.69
.448
.5
58
2.14
.428
5
69
2.46
.410
6
63
2.47
.494
Mean .5.fi6
62.3
2.72
.479
Range 4-8
58-69
2.05-4.80
341 -.600
ducal eggs, and corpora lutea. The mean for
either left or right side was 3.00 eggs each.
The o\-erall mean clutch size for each of the
three methods mentioni-d above was 6.22, 5.85,
and 5.90 t'ggs per femali', respeeti\'ely.
Corpora Lutea
Corpora lutea of I?YU 3.3.38, which had just
ovulated, were used as a standard (jf appear-
ance; thev were red (vascular), and the tissue
proper was thin and translucent in appearance.
lU'ceut corpora luti^a were large (3-4 mm in
diameter) and doughnut shaped, with an in-
dented area on one side near the center. They
regressed in size In one-third (2-3 mm) and
became less \ascular and gre\ish in appearance,
while the eggs remained in thi' oxiducts. Soon
after the eggs were o\ipositioned, the corpora
lutea turned vellowish, beginning a rapid re-
r.\BLE (i. .Size and weight differentials of female lizards before anil after eggs are laid (1970-71
Date
Date
Weight
.SVL
Date not
Weiglit after
Days
Weigli;
,S\L
Cra\ id
Grasid
Cra\ id
Gravid
La\ ill;;
Betw irii
Ditferenee
After
1970
6-26-70
9.32
.58.0
7-11-70
7.56
16
1.76
.59.0
7- 7-70
7.24
.59.0
7-14-70
4.83
7
2.76
.59.0
7-14-70
9.08
61.5
7-31-70
7..54
18
l.,54
63.0
1971
6-18-71
9.11
64.0
7-16-71
7.31
29
1.80
65.0
6-26-71
7.11
.58.0
7- 9-71
5.70
14
1.41
.59.0
6- 4-71
9. .56
64.0
7- 9-71
7.38
35
2.18
64.0
6-1.5-71
8.24
62.0
6-24-71
5.20
10
3.04
60.0
6- 8-71
10,65
66.0
6-21-71
8.03
14
2.62
66.0
6-18-71
8.00
60.0
6-2.5-71
5.17
8
2.89
.57.0
6-21-71
9.06
65.0
7-1.5-71
6.24
r>-
2.82
61.0
7- 7-71
8.14
64.0
7-16-71
5.14
9
2.00
1972°
6-30-72
8.12
62.0
7- 2-72
5.37
3
2.75
61.0
6-30-72
6.85
60.0
7- 4-72
4.96
.5
1.89
58.0
6-30-72
9.80
69.0
7-10-72
7.. 56
11
2.24
69
6-30-72
7.55
64.0
7- 7-72
5..56
6
1 .99
63.0
.\\erage
8..53
62.4
6.24
14
2.24
61.7
°Captiv
18
BlllGHAM VoUNG UNIVERSITY SCIENCE BULLETIN
TABLE 7. Clutcli size in S. f,'. graciosus fin 1970
thiough 1972.
^ olkct! ()\ ai ian
Egg''
()\ iducal
CDrpora
Lutea
Left Right Left Higlit Lfft Right
Total 205 206 134 147 88 83
Number RS 64 48 48 29 29
Range 1-6 1-5 1-5 1-5 2-4 2-4
Mean 3.01 3.21 2.79 3.06 3.03 2.86
Combined
mean 6.22 5.85 5.90
863 Ovarian egg.s, oxicUical eggs, and corpora hitea
143 Lizard.s
2-10 Range
6.03 Mean clnteh size
grcs.sion until tln'\ disappeared in approximate-
!)• 10 to 12 da\s. Corpora lutea were found in
the population from 2 June to 17 July 1970, a
period of 45 day.s, and 15 Jmie to 25 July 1971,
a period of 40 da\'.s.
Allelic FdIUcIcs
Atretie folliele.s were of three t\pcs. First,
yolking follicles were considered to be atretic
if there were corpora lutea or oviducal eggs
present. These follicles appeared to be the
beginning of a second clutch; however, based
on average time for \itellogenesis and oviducal
development, thev were late in the season for
successfid ineul)ation. In adchtion, females were
not found with oNidiical eggs after the third
week in July. Nuptial colors persisted l^cvond
this date in only three individuals, but no eggs
were found. Second, follicles tliat were few in
number and appeared \i)]ked in an o\ar\- but
were turning reddish \ellow were considered
atretic. These were apparently regressing in
size. This condition was also obser\ed before
ovulation in a few females in which some \()lked
o\a changed color and remained smaller in size.
The third condition was that of crenulated o\a
in the ceolom. In all three tspes a total of about
13 percent of tlie females examined had atretie
follicles.
Oviducal Eijt; Pcriixl in (^lidtic
Size and WeiiJU
In 1971 oNiducal eggs were
Jime to 10 JuK, a period of
captive females copulated on 7 ;
laid eggs on 2(S June and 2 Jul
periods of 22, 24, and 27 davs,
24.3 days, wliicli is the approxi
are in the oxiducts, assuming th
curs at or just after copulation.
1971 o\iducal eggs were used
size and weight of eggs. Ovid
Is,
foimd from 2
38 davs. Two
md 9 June and
K . These were
an a\'erage of
mate time eg.gs
at o\uiation oc-
lioth 1970 and
for calculating
ucal eggs were
weighed with the oviduct, which added a slight
l)ias to the weight of the eggs. Thev averaged
12.0 \ 7.1 mm with a range "of 10 x 7.5, 11 -x 6.5
mm to 14 X (S, and 15 x 7 mm. The average
weight per oviducal clutch was .28 g with a
range of .20 to .38 g. This was .07 g greater
than (A'arian eggs. In 1971 oviducts were vascu-
larized from ]() Ma\ to 10 Julv. After o\iposi-
tion the oviducts are no longer conxolutcd and
lose their vascularization.
Ovamignition and Oiidiict Analonaj
Transceolomic migration occurs freciuentlv
in i^raciosiis: Of 48 females ha\'ing o\iducal
eggs, 20 iiad eggs in the right or left oxiducts
that were of a different number than coipora
lutea in the ovaries. Forty-two percent of these
females had undergone transceolomic migration
of ova. Th("refore, o\amigration at the time of
ovulation is essentialh' a 50:50 relationship in
\vhich any ovum has an equal chance of enter-
ing either oviduct. This has a limitation ap-
parently in tliat no more than fixe eggs were
e\er found in one o\'iduct. The trend of migra-
tion follows sexeral patterns. First, there is an
even number of coipora lutea in the o\'aries with
a reduced ninuber in one of the o\iducts.
Second, tlu'y are exactly reversed from ovary
to oviduct— e.g., two corpora lutea in the right
ovar\', but three eggs in the right oviduct.
Third, corpora lutea are of mietiual numljcrs in
the ovaries and there is a highlv ime<|ual num-
ber of eggs in the o\iducts. For example, BYU
32616's right oviduct had three eggs and the
right o\arv had one corpora lutea. The fourth
condition is an e([ual number of corpora hitca
in thi' o\aries with an equal migration of ova
to each o\ iduct.
The o\iducts approaching o\ ulation appear
convoluted and \ ascularized. At their anterior
end the\' open into a slitlike ostium. The ovaries
are attached dorsallv to the l)od\ wall anterior
to the oxidiiet. The n\ iduct is laterally dis-
placed ill relitiou to the ovarv, with the ex-
pandeil ostium lateral to tjie adjaci'Ut oxary
and also at an angle to it.
Oviposition
Egg laxing oeeuis after eggs in tlie oviducts
have receix'ed the membraneous shell. The
period during which females ari' laving eggs,
ON'ipositiou, 'vx as calculated from stud\ plot and
collection data (1970-71). At our study plot the
egg-la\ing period lasted 29 da\s, (3 Julv-31
Juh) in 1970 and 31 days (19 June-20 July) in
1971. Clutch size increased as females grew
in size (Fig. 11).
Biological Series, Vol. 19, No. 5 Life History ok S( ki.oporus Chaciosus Chaciosu.s
19
M 6
O
O
Ul
•.. 5
O
z
z
<
S 2
53-54 57-60
SVL
61-64 65-69
Kis;. 11. Clutch size plotted against .S\'l, in loni size
classes of female S. g. graciosiis.
Incubation Period
Tlu' period from the first captured female
with only corpora lutea to the first hatchling
extended' from 3 July to 24 August 1970, 21
June to 6 .\ugust 1971, and 17 June to 10 August
1972, periods of 43, 48, and 55 davs, rcspec-
ti\'el\'. Tiie incubation periods from field data
were as follows: 1970, 6 Julv to 17 August;
1971, 19 June to 1 August; 1972, 1 July to 10
August (periods of 43, 44, and 41 days, respec-
tiveh', for a combined average of 43 days).
The incubation period for eggs from the time
of o\iposition in 1970, combining the field and
study plot data, was 48 days. In 1971 the period
w as 46 days, and in 1972, 48 da\s. The average
period combined for three years was 47 days.
The data also indicate that the incubation period
of 1970 \\as approximately 15 da\s later than
for 1971, witli 1972 in between. In addition to
this indiri'ct data, nine clutches of eggs were
incubated in the laboratorv at 27 to 28 C in
quart jars partly filled with moist sand. The
overall a\erage for these clutches was 49 days,
a figure similar to the field and stud\- plot data.
The a\'erage period for clutches where the exact
date of laying was known was 46 davs, which
closely approached the overall axerage. From
these data was derived the average incubation
period from the time the eggs were laid: 48
days, with a range of 44 to 52 davs.
Oviducal eggs taken from field females and
those captive females having recently oviposited
each showed a distinct area where the embryo
had probably been growing since the time of
fertilization. Known measurements of captive
females placed the eggs in the oviducts for a
period of approximately 22 to 27 days. This
was particularly true if eggs were ovulated
shortly after copulation ( 1-5 days ) and if they
were fertilized at or about the time they en-
tered the oviduct. Data concerning ovulation
time and copulation period strongly suggest that
fertilization occurred at or shortlv after the eggs
entered the oviducts. If this were the case, it
would add approximatelv 25 da\s to the actual
incubation period.
Sixty-eight eggs were laid and recovered from
the pen where 13 females were kept. Eight eggs
(11.7 percent) were infertile. Seven clutches
did not contain infi'rtile eggs, two had two
each, and four had one each. Infertile eggs ap-
peared softer and amber in color. They did not
attain turgidness, were soon attacked by fungus,
and, unless removed, began to contaminate the
entire clutch. From the remaining 60 eggs, 11
died just before their clutch mates hatched.
The embryos were between 20 and 23 inm
SVL. The yolk sac was intact with a small
amount of yolk still remaining. The 49 eggs
remaining were reduced by fungus contamina-
tion and other causes, leaving only 34 eggs that
hatched.
Avcraiic Size of Males at Sexual Maturity
Sexual maturit\ was detennined by gonadal
developmi'nt, size, and glandular development.
The main criterion used was the presence of
spermatozoa in the testes and epidid\'mis dur-
ing the reproductive period. As in females, 1
Jul\' was used as a seasonal date when males,
on the basis of size, were considered mature.
Some juveniles ma\' reach mature size by the
second week in July and may be mistaken for
adults even though they are not sexually mature.
Based on the abo\e criteria, males 52 mm SVL
were considered sexualh' mature. There were
three exceptions ( 1 percent of total mature
males) in 1970 and 1971 field specimens. These
three were 43, 46, and 50 mm SVL. In 1970
and 1971, none of the breeding population of
plot males were below 52 mm S\'L. This was
based on external appearance of males ( ventral
color, discolored pre-anal area, and swollen tail
base). While a few juvenile males in 1972 ap-
peared as above in late June and early July,
our autops\' data indicate that juveniles are not
functional until after the second liibemation.
20
BiiiciiAM YdUNG Unuehsiiv Science Bulletin
230
320
^.'— ~_^^
210
y^ J\
200
\
HO
'•\
1 180
\
.5 170
V /
»-
V- /'
i 160
\\ //
V
E 150
\\ /;
o
u 1
° HO
V
\\ /*
= 130
\\ /
3
- 120
\ •. /
c
V no
S
\ '■.•'/
100
\\ y'V
90
\ /
80
\ /
20
60
F"ig. 12. Mean sfniiiiifcious tubule diameter (uin) l)\'
montli for 197(1 and 1971. Solid line represents
1970: dotted line, 1971.
Spermatogenic Cycle
Histolog) of the testicular cycle in males
was grouped into eight stages by Mayhew
(1971). Stages of mature males during the 1971
activity season are listed in Table 8. This table
and Fig. 12 to 14 depict mean seminiferous
TABLE 8. Obser\ed spermatogenie stages in sex-
ually mature S. g. graciosns for 1970-71, during
the first and second half of each niOTith of the
actlvitv sea.son."
Marcli April May
Stage 1 2 1 2 12
]unc July Aug. Sept.
12 12 12 12
1
3
2
2
3
7
4
4
1
4
5
1
6
7
8
•otal
13
9
2 ISIS U 3
5 10
.5 8
13 9 G U U 10 14 II 12 26 24 11 3
"Stages of annual spermatogenic cvcle of adnlt males
are after Mayhew (1971).
tubule diameter, gonad weight, and epididymis
epithelial height (thickness) bv month and are
used in constructing tlu- spennatogenic cycle of
1970 and 1971 in the following sections.
In March 1970 male gonads were at stages
3 and 4. Seminiferous tubule diameter averaged
34
33
/" 7 \
32
/' / \
31
/ \
30
â– / 1
29
â– / \
28
â– / H
»
• / \
1"
â– / '\
■S«
/ / \\
X23
// i\
^22
"q.
1U30
/ \\
518
â– \ *
17
/ \\
16
\ •.
15
14
V\
13
Y
12
\
11
V
10
'N
9
M A M J J A S
Fig. 13. .Mean weight (g) ol testis li\ niontli loi fig. 14. Mean epidicK mis i']iith(lial h<i'.^lit ( iim I 1)\
1970 and 1971. Solid line represents 1970; dotted nimilh lor 1970 .iiid 1971. Solid linr r<present's
line, 1971. P)7(); dotted line, 1971.
Biological Series, Vol. 19, No. 5 Life Histohv of Scei.oi'ohus Cuaciosus CiiAtiosus
21
209.6 um (range 185-225 um). Epididymis epi-
thelium was in a pseudostratified condition,
with a height of 18.2 um (range 15-21 um).
Both testes averaged .21 g per lizard in March,
with a range of .13 to .35 g. .\dults in April
ranged from stages 3 to 6 and seminiferous tu-
bules had reached their maximum diameter
(mean 221.0 um, range 196-260 um). Epi-
didymis epithelium was proliferating and had an
average height of 24.7 um (range 10-31 um).
Gonad weight in April averaged .25 g per lizard,
with a range of .14 to .34 g.
Adult males in Mav were at stages 6 and 7.
The average diameter of seminiferous tubules
was 206.4 um (range 164-225 um). The first
males with spemiatozoa in the epididymis were
collected on 29 May 1970. Epididymis epitheli-
um height averaged 34.6 um ( range 32-35 um ) .
Gonad weights reached their peak in Ma\', av-
eraging .30 g per lizard (range .23.37 g).
In June males were still at stages 6 and 7,
with the predominance at stage 7. All had sper-
matozoa in the epididymis and were capable of
inseminating females. Seminiferous tubule di-
ameter was 157.7 um (range 145-203 um). Epi-
didymis height averaged 34.0 imi, with a range
of 26 to 45 mn. Gonads averaged .14 g per
lizard (range .09. 18 g).
In July males were predominantly at stage 8,
with only one at stage 7. Seminiferous tubules
reached their lowest level at 73.8 imi, with a
range of 54 to 90 um. The epididymis contained
spennatozoa until 11 Jul\-. Epididymis height for
Jul)' averaged 22.5 um ( range 17-32 um ) and
began to decline on or before 11 July. Gonad
weights in Juh' reached their lowest level with
a mean of .01 g per lizaid (range less than
.01-.02 g).
In August and September males reached
stages 8 or 1, with the majority at stage 1 (26
out of 31). July and August were secminglv the
end of one reproductive cycle and the begin-
ning of the next. Diameter of seminiferous tu-
bules for August and September averaged 107.2
and 175.6 um, with a range of 71 to 184 and
138 to 195 um, respectivelv. The epididymis
height was at its lowest level \\'ith an .August
average of 15.8 um (range 10-21 um). Septem-
ber epididymis height was uncertain because of
insufficient sample size. Onlv one specimen had
an epidid\niis of 10.0 um. Gonads averaged .04
and .15 g per lizard for August and September.
with ranges of .01 to .17 and .07 to .19 g. re-
specti\el\'.
In 1971 males emerg("d in March, at which
time the gonads were in stages 1 to 3. Semi-
niferous tubule diameter averaged 186.2 um
(range 164-210 um). Epididymis epithelium was
pseudostratified, with an average height of 16.1
um (range 10-20). In March gonad weight was
.21 g per lizard, with a range of .11 to .26 g.
Males in April, May, and June followed es-
sentially the same sequence in development as
in 1970. There were more discrepancies hi July,
however. Individuals were observed to be in
stages 7, 8, and 1. By late August and Septem-
ber males in 1971 were predominantly at stage
1, with only two at stage 8. The seminiferous
tubule diameter began to increase and averaged
110.0 and 165.7 um (ranges of 50-144 and 138-
195 lun ) . Epidid\inides were pseudostratified
and their heights were 18.7 and 9.0 um (ranges
10-27 and 10-18 um). The weights of gonads
were beginning to increase, with averages of
.04 and .15 g per lizard, with ranges of 10 to
.12and .04to.22g.
The spermatogenic cycle in 1970 was ap-
proximately fifteen days later than in 1971. In
March 1970 lizards were at stages 3 and 4,
while 1971 lizards were at stages 1 to 3. It ap-
pears that males in 1970 were more advanced
than those in 1971 even though in both years
they emerged at approximately the same time.
However, by the first half of May 1971, lizards
were more advanced than those in 1970. By
April 1971 epididymis epithelium had prolifer-
ated almost to its maximum for the year, where-
as 1970 epididymides were considerably lower in
epithelial height (Fig. 12). In addition, epi-
didymides in 1970 were not found with sper-
matozoa until 29 May. However, to be compar-
able to 1971 this should have occurred on ap-
proximately 15 Maw In contrast to the date of
15 Ma\ for 1970, 1971 lizards contained sper-
matozoa in the epididymides ten diiys earlier
on 6 Ma\ .
General Trends
Even though adult and juvenile males enter
hibernation with gonads approaching maximum
size, thev continue to increase during hiberna-
tion and develop more rapidly after they emerge.
Peak development of gonads is in the latter half
of April and the first of Mav. Male epididymides
usually contain spennatozoa in late May and
are capable of insemination. Spennatozoa re-
main in the epididymides until mid-Julv. By
the latter part of June, the epididymis epithelium
is sloughing and the lumen is full of cellular
debris. Thus, active spermatozoa are in the
epididymides from 1.5 to 2 months each year, a
period much longer than that normalK' used for
copulation (as based on field obserxations over
three seasons). B\ the end of Mav the gonads
are regressing. Thev continue to regress through
June, and by mid-Julv are at their smallest size
22
Bbigham Vounc Univehsity Science Bum.etin
Teitei > 20a
Seminilerouj Tubule Diameter >190 microns
Spermotoioo in Lumen of Teitei
Epididymis With Motile Sperm
Yolfa Depotition
Ovulation
Co p u la I ion
Oviducti Convoluled and Voscular
T
-r
T
Mo V
June July Aug. Sept.
FiS^. 1.5. Stimniaiy ot vaiioiis rcpiodtit-tivi' stages (it
adult S. <^. firciclo.iits. IJotli 1970 aiul 1971 data arc
comliiiu'd ill older lo simulate tlie a\erag<' Ncariv
trend.
(Fig. 12-14 and Table 8). In Augu.st the gonads
again begin to increase in tlie size of seminif-
erous tubules and weight, continuing until (he
lizards hibernate in late August and rarl\
September.
Male stages are compared with those of fe-
males in Fig. 15. The combined time periods
( 1970-71 ) are used to give the general yearl\
trends of male lizards in relation to females. Our
data for reproductixe cveles in this species are
generally similar to that ri-ported b\' W'oodbuiT
and Woodbury (19451.
Pre-anal Gland
After considerable handling of lizards to ob-
tain field measurements, it was noticed that
males left a moist area on the transparent ruler.
This moist area was directly in front of the \ent
and was discolored in relation to other areas
anterior to it on the alxlomen. At first the moist-
ness was suspeeted to l)e from the normal cloaca)
discharges; however, careful checking deter-
mined it to be a glandular secretion which
darkened the scales in the pre-anal area of adult
males during tlie eopulator\' period. Histological
sections of adult males show that there is indeed
a gland just anterior to the cloaca, under the
epithelial tissue— a duct gland, with cells lining
a lumen. The details are presented in a separate
report (Burkholder and Tanner, 1974).
Color in Males
Adult males are most brilliantly colored from
April until mid- or late July. During this time,
the ventral and v 
