Full text of "Pacific science"

See other formats

Sit

DL. XVII

JANUARY 1963

NO. 1

3ACIFIC SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

HAROLD ST. JOHN

Revision of the Genus Pandanus
Part 14. Malaya and Singapore

GEORGE W. BARLOW
Species Structure of Gillichthys mirabilis

TAKASHI OKUTANI

Molluscan Assemblages of Izu Submarine Banks
WILLIAM A. GOSLINE

Osteology and Systematics of Elongate Perciform Fishes

LINDSAY R. WINKLER and E. YALE DAWSON
Food Habits of California Sea Hares

D. L. INMAN , W. R. GAYMAN, and D. C. COX
Littoral Sedimentary Processes on Kauai

UNIVERSITY OF HAWAII PRESS

BOARD OF EDITORS

O. A. BUSHNELL, Editor-in-chief
Department of Microbiology, University of Hawaii

Robert Sparks, Assistant to the Editors
Office of Publications and Information, University of Hawaii

Thomas S. Austin

Bureau of Commercial Fisheries, Hawaii Area
(U. S. Fish and Wildlife Service)
Honolulu, Hawaii

L. H. Briggs
Department of Chemistry
University of Auckland
Auckland, New Zealand

Ai Kim Kiang
Department of Chemistry
University of Malaya, Singapore

Gordon A. Macdonald
Department of Geology
University of Hawaii

Donald C. Matthews
Department of Zoology
University of Hawaii

Colin S. Ramage

Department of Geology and Geophysics
University of Hawaii

Martin Sherman
Department of Entomology
University of Hawaii

Donald W. Strasburg
Bureau of Commercial Fisheries, Hawaii Area
(U. S. Fish and Wildlife Service)
Honolulu, Hawaii

Albert L. Tester

Department of Zoology and Entomology
University of Hawaii

Miklos F. Udvardy
Department of Zoology
University of British Columbia
Vancouver, Canada

Thomas Nickerson, Managing Editor
Assistant to the University Provost

INFORMATION

Contributions to Pacific biological and physical
science will be welcomed from authors in all parts of
the world. (The fields of anthropology, agriculture,
engineering, and medicine are not included.) Manu-
scripts may be addressed to the Editor-in-Chief,
PACIFIC SCIENCE, University of Hawaii, Honolulu
14, Hawaii, or to individual members of the Board
of Editors. Use of air mail is recommended for all
communications.

Manuscripts will be acknowledged when received
and will be read promptly by members of the Board
of Editors or other competent critics. Authors will be
notified as soon as possible of the decision reached.

FOR AUTHORS

Manuscripts of any length may be submitted, but
it is suggested that authors inquire concerning possi-
bilities of publication of papers of over 30 printed
pages before sending their manuscripts. Authors
should not overlook the need for good brief papers,
presenting results of studies, notes and queries, com-
munications to the editor, or other commentary.

PREPARATION OF MANUSCRIPT

It is requested that authors follow the style of
Pacific Science described herein and exemplified in the
journal. Authors should attempt to conform with the
Style Manual for Biological Journals , Am. Inst. Biol.
Sci. Washington.

( Continued on inside back cover)

PACIFIC SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

VOL. XVII JANUARY 1963 NO. 1

Previous issue published November 21 , 1962

CONTENTS

PAGE

Revision of the Genus Pandanus Stickman, Part 14 . New Species from Malaya

and Singapore. Harold St. John . 3

Species Structure of the Gohiid Fish Gillichthys mirabilis from Coastal Sloughs of

the Eastern Pacific. George W. Barlow 47

Preliminary Notes on Molluscan Assemblages of the Submarine Banks Around the

Izu Islands. Takas hi Okutani 73

Notes on the Osteology and Systematic Position of Hypoptychus dybowskii

Steindachner and Other Elongate Perciform Fishes. William A. Gosline.... 90

Observations and Experiments on the Food Habits of California Sea Hares of the

Genus Aplysia. Lindsay R. Winkler and E. Yale Daivson 102

Littoral Sedimentary Processes on Kauai, a Subtropical High Island.

D. L. Inman, W . R. Gayman, and D. C. Cox 106

note:

Adoption of the Metric System and Celsius Scale 131

Pacific Science is published quarterly by the University of Hawaii Press, in January,
April, July, and October. Subscription price is $4.00 a year; single copy, $1.25. Check
or money order payable to University of Hawaii should be sent to University of Hawaii
Press, Honolulu 14, Hawaii, U. S. A. Printed by Star-Bulletin Printing Company, 420
Ward Avenue, Honolulu 14, Hawaii.

SMITHSONIAN'

JHSTITJJT18N

MAR 2 5 1963

Revision of the Genus Pandanus Stickman, Part 14
New Species from Malaya and Singapore

Harold St. John1

As A RESULT of field work in Malaya and Sing-
apore and of study in the herbarium at Sing-
apore, numerous new species of Pandanus have
been detected. The following novelties are of
this origin.

Pandanus attenuatus sp. nov. (sect. Acro-
stigma)

Fig. 150

DIAGNOSIS HOLOTYPI: Frutex probabaliter
humili, caule in apice 8 mm diametro lutescenti
laevi sed cum dorsis longitudinalibus, foliis 35-
40 cm longis 18-20.5 mm latis chartaceis supra
viridibus infra minime viridibus pallidioribus et
proxima basem purpureo-tinctis 1-costatis et
minime 2-plicatis subplanis in sectione mediali
cum 14-15 nervis parallelis secundariis promi-
nentibus in quoque dimidio nervis tertialis sub-
visibilis infra proxima apicem, lamina ligulata
subito in apice caudato 5.5-6 cm longo 0.7 mm
lato contracto basi amplexicauli inermi valde
nervosa sed ex 2 cm marginibus cum aculeis
0.5-1. 3 mm longis 2-6 mm distantibus subulatis
adscendentibus pallidis, midnervo infra ultra
mediam inermi, in sectione mediali marginibus
cum aculeis 0.7-1 mm longis 1-4 mm distanti-
bus subulatis valde adscendentibus, apice subu-
lato cum marginibus et midnervo infra cum
aculeis 0.5-1 mm longis 0.5-1 mm separatis
subulatis adscendentibus, inflorescentia foeminea
terminali adscendenti cum 1 syncarpio, pedun-
culo 4.5 cm longo 4 mm diametro trigono brac-
teato, syncarpio 3.5 cm longo 2.7 cm diametro
ellipsoideo cum circa 144 drupis eis 13-13.5
mm longis 4-4.5 mm latis crassisque fusiformi-
bus 5-6-angulosis corpore 6-7 mm longo, pileo
7.5-9 mm longo basi 3.5-4 mm alto ovoideo
subanguloso laevi, stylo 4-6 mm longo valde
proxime arcuato osseoso subulato olim in basi
subanguloso, stigmate 4-4.5 mm longo late line-
ari brunneo papilloso in apice extento, endo-

1 B. P. Bishop Museum, Honolulu 17, Hawaii,
U. S. A. Manuscript received June 29, 1961.

carpio in parte lA infera lateribus cartilagineis
pallidis, seminibus 3. 5-4.5 mm longis subdo-
liformatis, mesocarpio apicali hemisphaerico,
mesocarpio basali sparso cum fibris fortibus in
lateribus.

DESCRIPTION OF ALL SPECIMENS EXAMINED:
Shrub, probably low; stem at apex 7—8 mm in
diameter, yellowish, smooth but with longitudi-
nal ridges; leaves 35-40 cm long, 18-20.5 mm
wide, chartaceous, green above, below slightly
paler green and near the base suffused with
purple, 1 -ribbed, slightly 2 -pleated, but nearly
flat, at midsection with 14-15 parallel second-
ary veins in each half, these prominent through-
out, cross tertiary veins barely visible only below
and near the tip, the blade ligulate, abruptly con-
tracted to a 5. 5-7. 5 cm caudate tip, this 0.7 mm
wide, the base amplexicaul, unarmed, very veiny,
beginning 2 cm up the margins with prickles
0.5-1. 3 mm long, 2-6 mm apart, subulate, as-
cending, pale; the midrib below unarmed to
beyonid the middle; at midsection the margins
with prickles 0.7-1 mm long, 1-4 mm apart,
subulate, sharply ascending; the subulate apex
having the margins and midrib below with
prickles 0.5-1 mm long, 0.5-1 mm apart, sub-
ulate, ascending; pistillate inflorescence terminal,
ascending, bearing 1 syncarp; peduncle 3-5 — 4.5
cm long, 4 mm in diameter, 3-sided, bracted;
syncarp 3-3.5 cm long, 2.7 cm in diameter, ellip-
soid, bearing about 120-144 drupes, these 13-
13.5 mm long, 4-4.5 mm thick and wide, fusi-
form, 5-6-angled, the body 6-7 mm long; pileus
7.5-9 mm long, the base 3.5-4 mm high,
ovoid, slightly angled, smooth; style 4-6 mm
long, sharply curved proximally, bony, subulate,
slightly angled only on the base; stigma 4-4.5
mm long, broad linear, brown, papillose, run-
ning to the tip; endocarp centering in lower
but extending to the base, the walls cartilaginous,
pale; seed 3. 5-4. 5 mm long, more or less barrel-
shaped; apical mesocarp hemispheric, with the
center hollow; basal mesocarp sparse, with strong
fibers up the sides.

5 mm

FIG. 150. Pandanus attenuatus St. John, from holotype. a, Habit, X 1; b, drupe, lateral view, X 1; c, drupe, ■
longitudinal median section, X 1; d, drupe, lateral view, X 4; e, drupe, longitudinal median section, X 4;

/, drupe, apical view, X 4; g, style and stigma, oblique view, X 10; h, leaf base, lower side, X 1; i, leaf middle,
lower side, X 1 \ j, leaf apex, lower side, X 1-

2 cm.

Page 230: Revision of Pandanus, 14. Malaya and Singapore — St. John

holotypus: Malaya, Selangor, Gunong lelu
Semangkok, 28 April 1922, I. H. Bur kill 8,876
(SING).

SPECIMENS EXAMINED: Malaya, Selangor,
Gap Sempadang, 1908, Has bin (sing).

DISCUSSION : P. attenuatus is a member of the
section Aero stigma, as is its closest relative, the
Malayan P. unguiculatus RidL, which has the
leaves 21-27 mm wide, at midsection with 17-
19 secondary veins in each half; syncarp 5.5 cm
long; drupes 18-20 mm long; the seed 7-8 mm
long, suprabasal; and the style 2.5-3 mm long.
P. attenuatus has the leaves 18-20.5 mm wide,
at midsection with 14-15 secondary veins in
each half; syncarp 3-3.5 cm long; drupes 1 3—
13.5 mm long; seed 3. 5-4. 5 mm long, basal;
and the style 4- 6 mm long.

The new epithet is the Latin word attenuatus,
drawn out, which is given in allusion to the
attenuate subulate leaf apex.

Pandanus caudatifolius , nom. nov. (sect. Acro-
stigma)

Fig. 151

P. globuliferus RidL, R. Asiat. Soc., Straits Br.,
Jour. 61: 42, 1912; non Thouars (1808).

DIAGNOSIS OF HOLOTYPE: "Stems up to 1 m
tall,” 6-7 mm in diameter, yellowish, smooth;
leaves 33-37 cm long, at the middle 15-17 mm
wide, near the base 10-11 mm wide, firm charta-
ceous, dark green above, pale below, almost flat
except at base, swordlike, tapering in lower lA
to the narrower base, in upper V3 rather sharply
narrowed to the 7 cm subulate apex, this 5 cm
down 1 mm wide, the secondary parallel veins
conspicuous and at midsection 15 in each half,
no visible tertiary cross veins, the base widened
and amplexicaul, unarmed, but beginning at
2-2.5 cm the margins with prickles 0.8-1 mm
long, 2-4 mm apart, subulate, straight or arcuate,
ascending, pale; the midrib below unarmed for
lower 54; at midsection the margins with sub-
ulate tipped serrae 0.2-0.3 mm long, 1.5-3 mm
apart; on the caudate triangular apex the mar-
gins and midrib below with prickles 0. 5-0.9
mm long, 1-3.5 mm apart, stout subulate, arcu-
ate, ascending; peduncle 3 cm long; syncarp
18 mm in diameter, globose, bearing about 40
drupes, these 11-13 mm long, 3.5-4 mm wide,

2-3 mm thick, the body oblanceoloid, com-
pressed, narrowed to the subulate style, 5-6-
angled, the body 8—9 mm long; pileus 6-7 mm
long, its base the widest part of the drupe, the
lower part pyramidal-semiorbicular; style 3-4
mm long, subulate, ascending arcuate; stigma
distal, 2.5-3 mm long, linear, extending to apex,
brown, papillose; endocarp in lower V4, obconic,
the base obtuse, stramineous, dull, cartilaginous,
the walls 0.2 mm wide; apical mesocarp 3.5-4
mm long, a cavern filled with white medullary
membranes and hairs; basal mesocarp sparse, the
margins fibrous, the center fleshy.

HOLOTYPE: Malaya, Selangor, Gunong Se-
mangok, 4,000 ft. alt., April 1894, H. N. Ridley
15,612 (sing).

DISCUSSION: P. caudatifolius is a member of
the section Aero stigma, as is its closest relative,
P. globosus St. John, a Malayan species with the
leaves 11-12 mm wide, little if at all narrowed
towards the base, at midsection the margins with
prickles 1-1.5 mm long, 2-5 mm apart, subulate;
drupes 14-17 mm long; and the style 3-6 mm
long. P. caudatifolius has the leaves 15-17 mm
wide near the middle, ligulate, narrowed below
and near the base 10-1 1 mm wide, at midsection
the margins with subulate-tipped serrae 0.2-0. 3
mm long, 1.5-3 mm apart; drupes 11-13 mm
long; and the style 3-4 mm long.

The epithet is derived from caudatus, having
a tail, and folium, leaf, given in reference to the
appearance of the leaf.

Pandanus globulosus sp. nov. (sect. Acro-
stigma )

Fig. 152

DIAGNOSIS HOLOTYPI: Frutex, caulibus in
apice 6-7 mm diametro laevibus lucidis lute-
scentibus, foliis 38-42 cm longis 11-12 mm lads
firme chartaceis supra obscure viridibus infra
pallide viridibus et ad basem purpureis 1-costatis
licet paene planis, in sectione mediali cum 12-
14 nervis parallelis secundariis in quoque dimi-
dio eis ubique prominentis, nervis tertialis nul-
lis, laminis ligulatis subabrupte contractis in
apice caudato subulato 5. 5-6. 5 cm longo 0.5 mm
lato basi amplexicauli inermi conspicue nervosa
ex 2-3 cm marginibus cum aculeis 0. 5-0.9 mm
longis 2.5-7 mm distantibus subulatis proxime
adscendentibus pallidis, midnervo per 30 cm in-

Fig. 151. Pandanus caudatifolius St. John, from holotype. a, Habit, X 1; b, c, drupes, lateral view, X 1;
d, drupe, longitudinal median section, X 1; e> drupe, lateral view, X 4; f, drupe, longitudinal median section,
X 4; g, drupe, apical view, X 4; h, leaf base, lower side, X 1; i, leaf middle, lower side, X 1; j, leaf apex,
lower side, X 1-

io cm.

Fig. 152. Pandanus globulosus St. John, from holotype. a, Habit, X 1; b, c, drupes, lateral view, X 1 \ d,
drupe, longitudinal median section, X 1; e, drupe, lateral view, X 4; f, drupe, longitudinal median section,
X 4; g, drupe, style, and stigma, apical view, X 4; h, leaf base, lower side, X 1; h leaf middle, lower side,
X 1; j, leaf apex, lower side, X 1-

5 cwl. o 2 cm.

PACIFIC SCIENCE, Vol. XVII, January 1963

ermi, in sectione mediali marginibus cum acuieis
1-1.5 mm longis 2-5 mm distantibus proxime
adscendentibus subulatis, in apice caudato mar-
ginibus et midnervo infra cum acuieis 0.7-1
mm longis 0.7-3. 5 mm distantibus subulatis ad-
scendentibus, inflorescentia foeminea terminal!
erecta cum uno syncarpio, pedunculo 4—6 cm
longo 3-4 mm diametro trigono bracteato, syn-
carpio 2. 5-3. 5 cm longO' 2.3-3 cm diametro
globoso vel elliptici-globoso cum circa 96-120
drop is eis 14-17 mm longis 3-4.5 mm latis cras-
sisque fusiformibus sed apice angustiori 5-6-
anguloso, corpore 9-10 mm longo oblanceoloi-
deo, pileo 7-11 mm longo basi 3-5 mm alta
ovoidea vel oblato-ovoidea cum 5 angulis prorni-
nentibus laevibus, stylo 3-6 mm longO' proxime
arcuato subulate rigido, stigmate 2-3 mm longo
linear! distali brunneo papillose in apice ex -
tento, endocarpio in parte V3 infera obconico
lateribus 0.2 mm crassis cartilagineis stramineis,
mesocarpio apicali suborbiculari cum mem-
branis albis radiatis numerosis, mesocarpio bas-
al! fibroso et carnoso.

diagnosis OF holotype: Shrub; stems at
summit 6-7 mm in diameter, smooth, shining,
yellowish; leaves 38-42 cm long, 11-12 mm
wide, firm chartaceous, dark green above, pale
green below and with a small area near the base
purple, 1 -ribbed, apparently almost flat, at mid-
section with 12-14 parallel secondary veins in
each half, these prominent throughout, no visible
tertiary cross veins, the blade ligulate, rather
abruptly contracted to a 5. 5-6. 5 cm subulate,
caudate apex, this 0.5 mm wide, the base am-
plexicaul and unarmed, prominently veiny, be-
ginning at 2-3 cm up the margins with prickles
0. 5-0.9 mm long, 2.5-7 mm apart, subulate,
closely ascending, pale; midrib unarmed for
about 30 cm; at midsection the margins with
prickles 1-1.5 mm long, 2-5 mm apart, closely
ascending, subulate; on the caudate apex the
margins and midrib below with prickles 0.7-1
mm long, 0.7-3. 5 mm apart, subulate, ascend-
ing; pistillate inflorescence terminal, erect, bear-
ing 1 syncarp; peduncle 4-6 cm long, 3-4 mm
in diameter, 3-sided, bracteate; syncarp 2.5—
3.5 cm long, 2.3-3 cm in diameter, globose
or elliptic-subglobose, bearing about 96-120
drupes, these 14-17 mm long, 3-4.5 mm wide
and thick, fusiform but the tip more slender,

5-6-angled, the body 9-10 mm. long, oblanceo-
loid; pileus 7-11 mm long, the base 3-5 mm
high, ovoid or oblate-ovoid, with 5 prominent
angles, smooth; style 3-6 mm long, curved prox-
imally, subulate, rigid; stigma 2-3 mm long,
linear, distal, brown, papillose; running to the
tip; endocarp in lower obconic, the walls 0.2
mm thick, cartilaginous, stramineous; apical
mesocarp suborbicular, with numerous radial,
white membranes; basal mesocarp sparse, fibrous
and fleshy.

HOLOTYPUS: Malaya, Pahang, Telom ridge,
Nov. 1908, H. N. R [idley\ 13,798 (sing).

discussion: P. globulosus is a member of the
section Aero stigma, as is its closest relative, the
Malayan species P. caudatif olius St. John. Under
that species is given a summary of the contrast-
ing differences.

The epithet is the Latin adjective globulosus ,
like a little sphere, and is chosen with reference
to the shape of the syncarp.

Pandcmus mollifoliaceus sp. nov. (sect. Acro-

stigma)

Fig. 153

DIAGNOSIS HOLOTYPI: Frutex, catile in apice
8 mm diametro luteo laevi, foliis 45-47 cm
longis 23-29 mm latis moll iter chartaceis supra
viridibus infra pallide viridibus et videtur glan-
ds 1-sulcatis 2-plicatis in sectione mediali cum
17-18 nervis secundariis parallelis in quoque
dimidio eis ubique prominentibus, circa apicem
infra nervis tertialis reticulum cum sectionibus
oblongis formantibus, lamina ligulata circa me-
dian! latissima proxima basem 14-15 mm latis,
in 6.5 cm apice caudato subiter diminuentibus
eo 5 cm ex apice 0.7 mm lato, basi amplexicauli
inermi et nervis exevidentibus, marginibus ex 3
cm cum acuieis 1—1.5 mm longis 2-5 mm dis-
tantibus subulatis pallidis in 45° adscendentibus,
midnervo per 35 cm inermi, in sectione mediali
marginibus cum acuieis 0.5-1 mm longis 4-10
mm distantibus subulatis adpresse adscenden-
tibus, in apice contractenti marginibus cum
acuieis 0.8-1.7 mm longis 2. 5-3. 5 mm distan-
tibus divergentibus, midnervo infra cum acuieis
simulantibus 4-7 mm distantibus, in apice cau-
dato marginibus et midnervo infra cum subulato-
serrulis 0.5-0.91 mm longis 2-4 mm distantibus,
inflorescentia foeminea terminal! erecta cum uno

to cm.

l cm. .

Fig. 153. Pandanus mollifoliaceus St. John, from holotype. a, Habit, X 1; b, drupe, lateral view, X 1;
c, drupe, longitudinal median section, X 1; ^ drupe, lateral view, X 4; e, drupe, longitudinal median section,
X 4; f, drupe, style, and stigma, apical view, X 4; g, leaf base, lower side, X 1; ^ leaf middle, lower side*
X 1; i, leaf apex, lower side, X 1-

5 cm.

PACIFIC SCIENCE, VoL XVII, January 1963

capite, pedunculo 3 cm longo 3 mm diametro
trigono bracteoso, syncarpio 2.7 cm longo 2.3
cm diametro subgloboso cum circa 120 drupis
eis 8-10 mm longis (vel in curvatura styli sequi
12-15 mm) 4-5 mm latis 3-4 mm crassis cor-
pore oblongo-ovoideo 5-anguloso lateribus stri-
atis, pileo cum basi 2 mm alta pulviniformi
5-angulato minute striato, stylo 4-5 mm longo
proxime in angulo recto curvato recto vel sinuoso
subulato gracile firmo in dimidia infera angu-
loso, stigmate 3-4 mm longo lineari distali brun-
neo paene in apice extento, endocarpio in tertia
infera 5-6 mm longo late obovoideo lateribus
0.2 mm crassis cum fibris longitudinalibus forti-
bus et membrano gracili pallidi, mesocarpio
apical i cavernoso cum membranis medullosis
albis paucis, mesocarpio basali carnoso et cum
fibris tenuibus.

DIAGNOSIS OF HOLOTYPE: Shrub; stem 8 mm
in diameter at apex, yellowish, smooth; leaves
45-47 cm long, 23-29 mm wide, soft charta-
ceous, green above, pale green and apparently
glaucous below, 1 -ribbed, 2-pleated, at midsec-
tion with 17-18 parallel secondary veins in each
side, these prominent throughout, and on the
lower side near the tip with barely visible cross
tertiary veins making irregular oblong meshes,
the blade ligulate, though broadest at or beyond
the middle, near the base 14-15 mm wide,
abruptly contracted to a 6.5 cm caudate tip, this
5 cm down 0.7 mm wide, the base amplexicaul,
unarmed, the veins obscure, beginning 3 cm up
the margins with prickles 1—1.5 mm long, 2-5
mm apart, subulate, pale ascending at about 45°;
the midrib unarmed for about 35 cm; at mid-
section the margins with prickles 0.5-1 mm
long, 4-10 mm apart, subulate, closely ascend-
ing; on the subapical contracting region the
margins with prickles 0.8-1. 7 mm long, 1.5-
2.5 mm apart, divergent, and those of the midrib
below similar but 4-7 mm apart; on the caudate
tip the margins and midrib below with subulate
serrulations 0. 5-0.9 mm long, 2-4 mm apart;
pistillate inflorescence terminal, erect, bearing
one head; peduncle 3 cm long, 3 mm in diam-
eter, 3 -sided, bracteate; syncarp 2.7 cm long, 2.3
cm in diameter, subglobose, bearing about 120
drupes, these 8-10 mm long (or following
the curve of the style 12-15 mm), 4-5 mm
wide, 3-4 mm thick, the body oblong-obovoid,

5 -angled, the sides striate; pileus with the base
2 mm high, cushion-shaped, 5 -angled, finely
striate; style 4-5 mm long, proximally ascend-
ing at right angles, straight or twisting, slender
subulate, firm, angled in lower half; stigma 3-4
mm long, linear, distal, brown, running almost
to the tip; endocarp centering in lower V3, and
5-6 mm long, broadly obovoid, the walls 0.2 mm
thick, of heavy longitudinal fibers and a thin
connecting membrane, pale; apical mesocarp a
broad cavern with a few white medullary mem-
branes; basal mesocarp fleshy and with weak
fibers.

HOLOTYPUS: Malaya, Negri Sembilan, Gun-
ong Angsi, eastward on ridge, 2,600 ft. alt., 24
Nov. 1923, Mohamed Nur 11,691 (sing).

DISCUSSION: P. mollifoliaceus is a member
of the section Aero stigma, as is its closest rela-
tive, P. caudatifolius St. John, of Malaya, a
species with its leaves 15-17 mm wide, firm
chartaceous, tapering from the middle down-
wards and near the base only 10-11 mm wide,
at midsection the leaf margins with subulate
tipped serrae 0.2-0. 3 mm long, 1.5-3 mm apart;
syncarp 18 mm in diameter, globose, bearing
about 40 drupes; drupes with the body oblance-
oloid, the sides smooth; pileus base pyramidal-
semiorbicular; stigma 2.5-3 mm long; endocarp
in lower 14, obconic; and the apical mesocarp
longer than wide. P. mollifoliaceus has its leaves
23-29 mm wide, soft chartaceous, tapering
downwards and near the base 14-15 mm wide,
at midsection the leaf margins with prickles
0.5-1 mm long, 4-10 mm apart, subulate; syn-
carp 2.7 cm long, 2.3 cm in diameter, subglobose,
bearing about 120 drupes; drupes with the body
oblong-obovoid, the sides striate; pileus base
cushion-shaped; stigma 3-4 mm long; endocarp
in lower V3, broadly obovoid; and the apical
mesocarp wider than long.

The new epithet is formed from the Latin
mollis, soft; foliaceus, leafy, in reference to the
soft texture of the leaves.

Pandanus albibracteatus sp. nov. (sect. Pan-

danus )

Figs. 154, 155, 169, 170^

DIAGNOSIS HOLOTYPI: Arboriformis, caulibus
8 m altis 15 cm diametro viridarium formans,
cortice brunneo muricato, radicibus fulturosis 1

Fig. 154. Pandanus albibracteatus St. John, from holotype. a, Fresh drupe, lateral view, X 1; ^ dried
drupe, lateral view, X 1; ^ drupe, longitudinal median section, X 1; d, drupe, apical view, X 1; e, carpel
apices, stigmas, and proximal sinuses, oblique view, X 4; /, leaf base, lower side, X 1; ft leaf middle, lower
side, X 1; b, leaf apex, lower side, X 1.

PACIFIC SCIENCE, VoL XVII, January 1963

m longis 3—4 cm diametro in lineis muricatis,
foliis 1.72-1.97 m longis 5-3.5 cm latis coriaceis
olivaceo-viridibus supra glaucis infra glauci-
oribus supra midnervum sulcatis 2-plicatis in
sectione mediali cum 41-44 nervis parallelis
secundariis in quoque medio nervis tertialis nul-
lis infero folii ligulato medio exteriori gladi-
formati sensim in apice 40 cm longo caudato
subulato trigono diminuenti eo 10 cm ex apice
0.9 mm lato, basi amplexicauli et inermi sed ex
2-6 cm marginibus cum aculeis 4.5-7 mm longis
10-45 mm separatis subulatis rectis vel arcuatis
apicibus rubro-brunneis plerumque adscendenti-
bus, midnervo infra cum aculeis 3.5 mm longis
20-45 mm separatis conico-subulatis reflexis, in
sectione mediali marginibus cum aculeis 3-5
mm longis 7-20 mm separatis subulatis adpresse
adscendentibus, midnervo infra cum aculeis 3.5-
4 mm longis 22-28 mm separatis crassiter sub-
ulatis adscendentibus, in apice subulato mar-
ginibus et midnervo infra cum subulato-serrulis
0.3-0. 5 mm longis 4-10 mm separatis; inflores-
centia foeminea terminali arcuata cum syncarpio
solitario, pedunculo 27 cm longo 13-20 mm
diametro trigono folioso-bracteato, syncarpio 23
cm longo 17.5 cm diametro ellipsoido-subglo-
boso cum 67 phalangibus, nucleo maturo molli
et carnoso subito contracto in reliquum minp-
tum, phalangibus 5. 5-6.2 cm longis dimidio
supero 3.2-4 cm lato 2. 5-3. 5 cm crasso oblongo-
ellipsoideo vel obovoideo infra aurantiaco-rubro
supra subviridibus, deinde rosaceo 5-6-anguloso
lateribus laevibus sublucidis subconvexis, suturis
lateralibus plerumque nullis rariter 1 (-2), apice
eonvexo, sinibus apicalibus centralibus 0.7-2
mm profundis V-formatis in fondam plerumque
rectis, dimidio infero phalangiis 33-4.2 cm latis
2. 8-3. 9 cm crassis carnoso-incrassatis rosaceo-
aurantiacis cuneato-oblongis humeris 3-6 mm
latis 5-8-angulosis lateribus gradatim curvatis
carne saccharosa innoxia sed non consumpta,
carpellis 4-7 eis centralibus 2A tarn grandibus
quam marginalibus, apicibus depresso-conico eis
marginalibus asymmetricis et aliquis cum piano
vel concavo distalo stigmatis, stigmatibus 2.5-4
mm longis cordatis vel deltoideis vel reniform-
ibus pallide brunneis papillosis sulcatis obliquis
centripetalibus, sinibus proximalibus Vi-Vi ad
fondam extends, endocarpio minime supra-
mediali osseoso mahogani-colorato intra lucido

lateribus 5 mm crassis, seminibus 12-15 mm
longis 5 mm diametro ellipsoideis, mesocarpio
supero in quoque apice cavernam cum fibris
paucis et membranis a lb is medullosis multis
formantibus, mesocarpio basali grande fibroso et
carnoso.

DIAGNOSIS OF holotype: Treelike; stems 8
m tall, 15 cm in diameter, forming a large
clump; bark brown, muricate; prop roots 1 m
long, 3-4 cm in diameter, muricate in lines;
leaves 1.72-1.97 m long, 5-5.5 cm wide, cori-
aceous, olive green, above glaucous and more so
below, broad furrowed above the midrib, and |
with 2 rounded pleats, at midsection with 41-44 j
secondary parallel veins in each half, no ter-
tiary cross veins, lower half ligulate, outer half
sword-shaped, gradually tapering to a 40 cm j
caudate subulate trigonous apex, this 10 cm
down only 0.9 mm wide, the base amplexicaul !
and unarmed, but beginning at 2-6 cm the mar- 1
gins with prickles 4.5-7 mm long, 10-45 mm
apart, subulate, mostly ascending, straight or
arcuate, red-brown-tipped; the midrib below
with prickles 3-5 mm long, 20-45 mm apart,
conic-subulate, reflexed; at midsection the mar-
gins with prickles 3-5 mm long, 7-20 mm apart,
subulate, appressed ascending; the midrib below
with prickles 3.5-4 mm long, 22-28 mm apart,
stout subulate, ascending; on the subulate apex
the margins and midrib below with subulate-
serrulations 03-0.5 mm long, 4-10 mm apart;
pistillate inflorescence terminal, arching, bear-
ing one syncarp; peduncle 27 cm long, 13-20
mm in diameter, 3 -sided, leafy bracted; syncarp
23 cm long, 17.5 cm in diameter, ellipsoid-
subglobose, bearing 67 phalanges, the ripe core
soft and fleshy and quickly shrivelling to a very
small remnant; phalanges 5. 5-6.2 cm long, the
upper half 3.2-4 cm wide, 2. 5-3. 5 cm thick,
oblong-ellipsoid or obovoid, below orange-red,
above greenish changing to pink, 5-6-angled,
the sides smooth, a little shiny, gently curved,
lateral sutures usually none, rarely 1 (—2), the
apex convex; central apical sinuses 0.7-2 mm
deep, V-shaped at the very bottom, mostly
straight; lower half of phalange 33-4.2 cm
wide, 2. 8-3.9 cm thick, fleshy enlarged, pinkish
orange, cuneate-oblong in outline, the shoulders
3-6 mm wide, often eaten by small, black ants,
and reduced to white scars, 5-8-angled, the sides

Page 238: Revision of Pmdanus, 14. Malaya and Singapore — St. John

smooth, gently curved, the flesh sweet, inoffen-
sive, but not eaten; carpels 4-7, the central ones
about 2A as large as the marginal, the apices
very low conic, the marginal ones asymmetric
and some of them having just distal of the
stigma a slight flat platform or even a concavity;
stigmas 2.5-4 mm long, from cordate to deltoid
or reniform, light brown, papillose, creased,
oblique, centripetal; proximal sinus running Vi-
2A way to valley bottom; endocarp slightly
supramedian, bony, mahogany-colored, the inner
surfaces shining, the lateral walls 5 mm thick;
seeds 12-15 mm long, 5 mm in diameter, el-
lipsoid; upper mesocarp forming in the apex
of each carpel a cavern with a few fibers but
filled with aerenchyma of white medullary
membranes; basal mesocarp fibrous and fleshy,
extensive.

DESCRIPTION OF STAMINATE TREE: Tree 7
m tall, 12 cm in diameter; bark brown, muricate;
prop roots none, but tree abused; leaves 1.23 m
long, 3.9-4 cm wide, coriaceous, furrowed above
the midrib, the sides not plicate, at midsection
with 35 secondary parallel veins in each half, no
tertiary cross veins, sword-shaped, tapering into
a 15-20 cm subulate trigonous apex, this 10 cm
down, 3 mm wide; at 3-4.5 cm from the base
the margins with prickles 4-5 mm long, 7-21
mm apart, arcuate subulate, with thickened base,
ascending; the midrib below beginning at 6-7
cm with prickles 3-4.5 mm long, 4-22 mm
apart, similar but reflexed; at midsection the
margins with prickles 1.5-2. 5 mm long, 6-12
mm apart, subulate, closely appressed ascend-
ing; the midrib below with prickles 1.5-2 mm
long, 13-25 mm apart, subulate, arcuate ascend-
ing; the subulate apex with margins and midrib
below with prickles 0. 5-0.7 mm long, 3-7
mm apart, subulate-serrations; staminate inflor-
escence single, terminal, arcuate and pendent, 50
cm long; peduncle fleshy and soon disintegrat-
ing; lower bracts with lower part and the other
bracts all white; lowest floral bract 48 cm long,
2.2 cm wide, the body ligulate except at base
with marginal prickles 2-2.5 mm long, acerose,
ascending, closely appressed, white; the apex
20-25 cm long, subulate, foliaceous; median
bract 19 cm long, 2.6 cm wide, linear-lanceolate,
unarmed except near the tip; spikes about 12,
dense, 2.5-4 cm long, 1.5-2 cm in diameter,

cylindric; staminate flowers very numerous, 1.5-
1.9 cm long; column 4-8 mm long, bearing a
common section of equal length and with about
24 stamens, the free filament tips 1-2 mm long;
anther body 2. 4-3. 6 mm long, linear-lanceolate,
bearing an apical subulate prolongation of the
connective 0.6-0. 7 mm long.

HOLOTYPUS: Malaya, Penang Island, 5 miles
n. of George Town, top of sea beach, June 30,
I960, H. St. John 26,371 (bish).

SPECIMENS EXAMINED: Malaya, Penang Is-
land, Tanjong Tokong, 6 miles n. of George
Town, top of sea beach, staminate, June 30,
I960, H. St. John 26,370 (bish); Penang Island,
Mt. Pleasure, 8 miles n. of George Town, top
of quartz sand sea beach, clump with stems cut
back to 1 m. by 10 cm, old phalanges from the
ground, June 30, I960, H. St. John 26,369
(bish).

DISCUSSION : P. albibracteatus is a member of
the section Pandanus, as is its closest relative,
the Singapore species P. Boryi Gaud., which has
the phalanges with the central apical sinuses
1.5-2. 5 mm deep; marginal carpels with a visor
projecting over the stigma; leaves 3. 5-4.4 cm
wide, sword-shaped, at midsection with 39 sec-
ondary parallel veins in each half, and near the
base the margins with prickles 3-4.5 mm long,
7-25 mm apart. P. albibracteatus has the phal-
anges with the central apical sinuses 0.7-2 mm
deep; stigmas lacking a visor; leaves 5-5.5 cm
wide, the lower half ligulate, the outer half
sword-shaped, secondary parallel veins 41-44 at
midsection in each half, near the base the mar-
gins with prickles 4.5-7 mm long and 10-45
mm apart.

The new epithet is formed from the Latin
words albus, white; bracteatus, with bracts, and
given in allusion to the white floral bracts.

Pandanus ambiglaucus sp. nov. (sect. Pan-
danus )

Figs. 156, 170 b

DIAGNOSIS HOLOTYPI: Arbor juvenalis,
trunco erecto 1 m alto 10 cm diametro, radici-
bus fulturosis ad 1 m longis et 4 cm diametro
brunneis in lineis sparse muriculatis, foliis 2-
2.47 m longis 5.8— 6.3 cm latis coriaceis viridi-
bus et glaucis in lateribus ambis, spinis omnibus
albis, lamina late sulcata et dimidiis subsigmoi-

PACIFIC SCIENCE, Vol. XVII, January 1963

deis in sectione mediali cum 44-46 nervis
parallelis secundariis in quoque dimidio, nervis
tertialis nullis, gladiformatis sensim in apice
trigono subulato caudato 20-30 cm longo di-
minuentibus eo 10 cm ex apice 1.8 mm lato,
basi amplexicauli et inermi sed ex 5-6 cm mar-
ginibus cum aculeis 4-6 mm longis 5-26 mm
separatis valde subulatis arcuatis adscendenti-
bus, midnervo infra ex 7. 5-8. 5 cm cum aculeis
3-4.5 mm longis 20-32 mm separatis arcuatis
subulatis reflexis basibus crassis, in sectione
mediali marginibus cum aculeis majoribus 2.5-
3 mm longis 18-30 mm separatis arcuatis sub-
ulatis adscendenti-adpressis basibus crassis ple-
rumque alternantibus cum aculeis minoribus 1-
2 mm longis, midnervo infra cum aculeis 1.5-2
mm longis 13-24 mm separatis subulatis ad-
scendentibus, in apice subulato marginibus et
midnervo infra cum subulato-serrulis 0.3-1 mm
longis 3-8 mm separatis, inflorescentia foemi-
nea terminali arcuati, pedunculo 40 cm longo

2 cm diametro trigono folioso-bracteato cum
syncarpio solitario eo 21 cm longo 16 cm diam-
etro ellipsoideo (submaturo et viridi) cum 85
phalangibus eis 6-6.6 cm longis 2.8-4 cm latis
2. 8-3. 6 cm crassis obovoideis ad apicem infra
cuneatis, apice convexo 5-6-anguloso lateribus
supra convexis infra concavis (evidente imma-
turis), laevibus lucidis, suturis lateralibus nullis,
sinibus apicalibus centralibus 0.5-2. 5 mm pro-
fundis latis et vadosis rectis vel subcurvatis,
carpelis 7-8 eis centralibus Vi-2A tarn grandibus
quam illis marginalibus eis basalibus mediali-
busque cum apicibus subplanis sed illis centrali-
bus cum apicibus oblato-conicis, stigmatibus 2-

3 mm longis suborbicularibus vel ellipticis ob-
scure brunneis obliquis sulcatis centripetalibus,
ceteris marginalibus truncatis, sinibus proximal-
ibus Vi ad fondam extentis, endocarpio in parte
lA inf era osseoso brunneo, seminibus 12-16
mm longis ellipsoideis, mesocarpio supero in
quaque carpella cavernam cum fibris et mem-
branis pallidis medullosis formanti, mesocarpio
basali fibroso et carnoso.

DIAGNOSIS OF HOLOTYPE: Young tree, trunk
erect 1 m tall, 10 cm in diameter; prop roots
up to 1 m long and 4 cm in diameter, sparsely
muriculate in vertical lines, brown; leaves 2-
2.47 m long, 5.8—63 cm wide, coriaceous, green
and glaucous on both sides, the spines all white,

the blade broad furrowed over the midrib, the
sides slightly sigmoid, at midsection with 44-
46 secondary parallel veins in each half, no
visible tertiary cross veins, sword-shaped, grad-
ually tapering upwards into a 20-30 cm caudate,
subulate, trigonous apex, this 10 cm down 1.8
mm wide, the base amplexicaul and unarmed,
but beginning at 5-6 cm up the margins with
prickles 4-6 mm long, 5-26 mm apart,' stout
subulate, arcuate, ascending; the midrib below
beginning at 7. 5-8. 5 cm with prickles 3-4.5
mm long, 20-32 mm apart, arcuate subulate, ■
heavy based, reflexed; at midsection the mar-
gins with larger prickles 2.5-3 mm long, 18-
30 mm apart, arcuate subulate, heavy based,
ascending appressed, mostly alternating with
smaller ones 1-2 mm long; the midrib below
with prickles 1.5-2 ■'mm long, 13-24 mm apart,
subulate, ascending; on the subulate apex the
margins and midrib below with subulate serru-
lations 0.3-1 mm long, 3-8 mm apart; pistil-
late inflorescence terminal, arching; peduncle
40 cm long, 2 cm in diameter, 3 -sided, leafy
bracted, bearing a single syncarp, this 21 cm
long, 16 cm in diameter, ellipsoid, slightly im-
mature and still green, bearing 85 phalanges,
these 6-6.6 cm long, 2.8-4 cm wide, 2. 8-3. 6
cm thick, obovoid above, cuneate below, the
apex convex, 5— 6-angled, the sides convex-
above, concave below (but this probably due to
immaturity), smooth, shining; lateral sutures
none; central apical sinuses 0.5-2. 5 mm deep,
shallow, the line straight or gently curved; car-
pels 7-8, the central ones Vl -2A as large as the
marginal ones, the basal and median phalanges
with the carpel apices almost flat, but those
near the center with the apices oblate conic;
stigmas 2-3 mm long, suborbicular to elliptic,
dark brown, oblique, creased, centripetal, some
of the marginal ones truncate; proximal sinus
running Vz way to valley bottom; endocarp in
lower J/3, bony, brown; seeds 12-16 mm long,
ellipsoid; upper mesocarp forming in each car-
pel apex a cavern, with fibers and pale medul-
lary membranes; basal mesocarp fibrous and
fleshy.

HOLOTYPUS: Singapore, Woodlands Road, .
14 miles mark, opposite Kranji Nature Reserve,
fresh water swamp, with Dicranopteris linearis,
July 10, I960, H. St. John 26,376 (bish).

Fig. 155. Pandanus albibracteaius St. John, from St. John 26,370. a, Staminal column with anthers, lateral
view, X 10; h, leaf base, lower side, X 1; ^ leaf middle, lower side, X 1; d, leaf apex, lower side, X 1-

FIG. 156. Pandanus ambiglaucus St. John, from holotype. a, Phalange, lateral view, X 1; b, phalange,
longitudinal median section, X 1; c, phalange, apical view, X 1; ^ carpel apex, stigma, and proximal sinus
of a median phalange, oblique view, X 1; e, carpel apex, stigma, and proximal sinus of a phalange near the
base, oblique view, X 1; /, leaf base, lower side, X 1 \ g, leaf middle, lower side, X 1; leaf apex, lower side,

XI.

10 cm/.

O 1 OCYYl.

I 1 1 — . — 1 i. L — l • i I 4

o J

FIG. 157. Pandanus Boryi Gaud., from Singapore, Posher g 36,919. a, Phalange, lateral view, X 1; h,
phalange, longitudinal median section, X 1; o, phalange, apical view, X 1; ^ carpel apex, stigma, and proximal
sinus, oblique view, X 1; ^ leaf base, lower side, X 1 ; leaf middle, lower side, X 1; g, leaf apex, lower
side, X 1-

PACIFIC SCIENCE, Vol. XVII, January 1963

DISCUSSION: P. ambiglaucus is a member of
the section Pandanus, as is P. inclinatus St.
John, also of Singapore, its closest relative, and
a species with the syncarp with 44-61 phal-
anges, these with the apex truncate or sub-
truncate, and the lower half fleshy enlarged;
endocarp median; leaves 1.5— 1.7 m long, 4.4—
4.7 cm wide, above dark olive green, below
green; and the secondary veins at midsection
38-40 in each half. P. ambiglaucus has the
syncarp with 85 phalanges, these with the apex
convex, and the lower half cuneate; endocarp in
the lower lA\ leaves 2-2 .47 m long, 5.8-63 cm
wide, green and glaucous on both sides; and
the secondary veins 44-46 in each half at the
midsection of the leaf.

The new epithet is from the Greek words
ambos, both; glaukos, color of the sea, applied
to the glaucous coating on both sides of the
leaves.

Pandanus Boryi Gaud., Bot. Voy. La Bonite, pi.
22, f. 15, 1841

Fig. 157

Description from Fosberg 36,919, Singapore:
"Small tree,” leaves more than 1.64 m long,

3.5 cm wide, subcoriaceous, green above and
below, channeled above the midrib, 2 -pleated,
in section M-shaped, at midsection with 39
parallel secondary veins in each half, no tertiary
cross veins, sword-shaped, tapering gradually to
the trigonous subulate tip, more than 15 cm
long, and about 10 cm down, only 2.5 mm wide,
the base amplexicaul and unarmed, but begin-
ning at 4-7 cm the margins with prickles 3-

4.5 mm long, 7-25 mm apart, arcuate, subu-
late, ascending, pale; the midrib below be-
ginning at 19 cm with prickles 3 mm long,
25-40 mm apart, similar but reflexed; at mid-
section the margins with prickles 2-2.5 mm
long, 8-15 mm apart, arcuate subulate, ascend-
ing appressed; the midrib below bearing simi-
lar prickles 1.7-2 mm long; on the subulate
apex the margins and midrib below with ser-
rulations 0.3-0. 5 mm long, 3-8 mm apart, red-
tipped. . . fruiting head small cylindric, red
when ripe”; phalange 6.3 cm long, 3.4 cm
wide, 2.8 cm thick, pyriform, compressed, when
dried the apex brown, but yellowish below, the
sides smooth, shining, gently curved, free in

upper V3, the apex low convex, lateral sutures
none (from the direction of the basal fibers and
from the remnant of a shoulder, it is apparent
that when fresh the basal third had fleshy en-
largements); central apical sinuses 1.5-2. 5 mm
deep, broad; carpels 6, the apices very low, de-
pressed pyramidal, the inner ones slightly the
smaller; stigmas 2-2.5 mm wide, reniform, dark
brown, papillose, centripetal and mostly oblique,
some of the marginal ones with the apex trun-
cate by an overhanging visor which terminates
a distal concavity; proximal sinus running Vz
way to valley bottom; endocarp median, 3 cm
long, bony, massive, dark mahogany-colored,
the lateral walls 3 mm thick, the inner surfaces
shining; seeds 12 mm long, 3-4 mm in diam-
eter, ellipsoid; upper mesocarp forming in the
apex of each carpel a cavern with few fibers
but filled with aerenchyma of white medullary
membranes; basal mesocarp fibrous and fleshy.

EXPANDED DESCRIPTION FROM OTHER SPECI-
MENS: Branch tips 4.5-5 cm in diameter, brown,
with crowded leaf scars; leaves 1-1.64 m long,

3. 5- 4.5 cm wide, at midsection with 39—45
parallel secondary veins in each half, near the
base the marginal prickles 3-5 mm long, those
of the midrib below 3-5 mm long; peduncle
terminal, 22 cm long, 15 mm in diameter,
3 -sided, leafy bracted, bearing a single syncarp,
this 16 cm long, 14 cm in diameter, broadly el-
lipsoid, bearing numerous phalanges; phalanges

5.5- 6 cm long, 2-3.4 cm wide, 2-2.8 cm thick,
pyriform to wedge-shaped, compressed; carpels
6-10.

STANDARD specimen: Singapore: edge of
"dry” mangrove swamp, April 17, 1956, F. R.
Fosberg 36,919 (us).

SPECIMENS EXAMINED: Singapore, Sungei

Gunong, Feb. 1893, H. N. Rlidley'} 3,003
(sing); Singapore, Dec. 15, 1904, W. Fox
12,373 (sing).

DISCUSSION: P. Boryi Gaud, is a member of
the section Pandanus. The type specimen is a
single phalange in the Paris museum, collected
by Gaudichaud, and left without data. It was a
misfortune that he abandoned his taxonomic
work half done and devoted his last years to
morphogenesis, producing lengthy papers that
are now forgotten. P. Boryi was published with
the binomial and with a figure showing a

10 ovn.

FIG. 158. Pandanus carnosus St. John, from holotype. a, Fresh lateral phalange, lateral view, X 1; b, dried
subbasal phalange, lateral view, X lj G phalange, longitudinal median section, X lj d, lateral phalange, apical
view, X 1; e, subbasal phalange, apical view, X 1 \ f, apex of carpel in lateral phalange, oblique view, X 4;
g, apex of carpel in subbasal phalange, oblique view, X 4; h, leaf base, lower side, X lj h leaf middle, lower
side, X 1; j, leaf apex, lower side, X 1-

PACIFIC SCIENCE, VoL XVII, January 1963

lateral view of a phalange, nothing more. Since
the genus existed, this was valid publication.
Brongniart reviewed the Pandanus specimens
collected by Gaudichaud and documented them
as far as possible (Ann. Sci. Nat. Bot. VI, 1: 290,
1875). For this species he postulated that it
came from He de la Reunion. However, in the
recent detailed revision of the Mascarene species
(Linn. Soc. Lond. Bot. Jour. 55: 1-32, 1953)
Vaughan and Wiehe found no local species to
match it.

It is of interest, then, to note that the recent
collection from Singapore, Posher g 36,919, is a
perfect match for Gaudichaud’s illustration of
P. Boryi . This collection consists of one phalange
and one leaf minus the tip, so it is scarcely more
adequate than the Gaudichaud collection. Since
it is a perfect match for the Gaudichaud figure,
we offer here a description of this phalange and
leaf^as a standard description to supplement the
minimal publication by Gaudichaud. In the
Singapore herbarium there are also two collec-
tions from Singapore that fall into this same
■species. One has six phalanges and two' leaves
minus the tips; the other had a branch, many
leaf bases, peduncle, core of the syncarp, and
eight phalanges. Together they contribute to our
knowledge of the population. No one of them
furnishes all the details, but of all, the Fosberg
36,919 seems best to take as a standard.

The vessel "La Bonite,” on which Charles
Gaudichaud was the botanist, circumnavigated
the globe in 1836 and 1837, and stopped at
Singapore. Pandanus is still fairly common on
Singapore Island and occurs there in several spe-
cies. One of the tidal estuaries not far from the
harbor is Sungei Pandan, and the general ver-
nacular name of the genus in Malayan is "pan-
dan.” This was the source of our generic name,
Pandanus, latinized and first published by Rum-
phius, later adopted and made valid by Stickman.

Pandanus carnosus sp. nov. (sect. Pandanus)
Fig. 158

DIAGNOSIS HOLOTYPI: Arbor 10 m alta 15 cm
diametro, cortice pallide brunneo sparse muri-
cato, radicibus fulturosis 1-2 m longis 3-3.5 cm
diametro pallide brunneis in lineis muriculatis,
foliis 1.8-2 .4 m longis 5-5.2 cm latis supra
olivaceo-viridibus et subglaucis infra pallide vir-

idibus coriaceis cum midnervo forti et in centra
late sulcatis in sectione mediali cum 45 nervis
parallelis secundariis in quoque dimidio' nervis
tertialis nullis laminis gladiformatis cum apici-
bus 15-30 cm longis subulatis trigonis eo 10 cm
ex apice 0.9- 1.3 mm lato basi amplexicaule et
inermi sed ex 10 cm marginibus cum spinis

5- 6.5 mm longis 11-30 mm separatis valide
subulatis arcuatis adscendentibus pallidis, mid-
nervo infra ex 22 cm cum spinis 6 mm longis
15-35 mm separatis simulantibus sed reflexis,
in sectione mediali marginibus cum aculeis 3.5-
4 mm longis 11-18 mm separatis subulatis valde
adpresse adscendentibus apicibus rubris, mid-
nervo infra cum aculeis 3 mm longis 15-25 mm
separatis simulantibus adscendentibus, in apice
subulate marginibus et midnervo infra cum
subulato-serris 0.5-0.8 mm longis 2-11 mm
separatis, syncarpio solitario terminali 18 cm
longo 1 5 cm diametro subgloboso cum phalangi-
bus numerosis eis 6.7-73 cm longis 2.7-3. 5 cm
latis 2.4-3. 1 cm crassis apice in parte viridi
in parte rubro-aurantiaco 5-6-anguloso, parte

supera libera oblongo-elliptica vel oblongo-
obovoideo apice depresso-convexo lateribus sub-
curvatis rugulosis sed sublucidis, suturis laterali-
bus nullis, sinibus centralibus apicalibus 4.5-

5.5 mm profundis et V-formatis in apicibus
anguste conicis (sed in phalangibus paucis proba-
baliter eis subbasalibus apicibus demissiter pyra-
midalibus sinibus 2-3.5 mm profundis), carpelis

6- 10 apicibus plerumque anguste conicis eis
marginalibus lA latioribus asymmetricalibus et
plus minusve distaliter compressis erectis vel
subdivergentibus, parte lA inf era phalangis
carnoso-incrassata 3-4 mm et deinde 25-35 mm
diametro, stigmatibus 3-3.5 mm longis ellipticis
sulcatis brunneis papillosis, sinibus proximalibus
V5-V2 ad fondam extensis, endocarpio medi-
ali 2.5-3 cm longo mahogani-brunneo osseoso
solido lateribus 3-4 mm crassis intra lucidis,
se.mi.oi bus 13-18 mm longis 4.5 mm diametro
fusiformibus, mesocarpra supero in apice car-
pel! quique cavernam formanti eis lateralibus

2.5 cm longis cum membranis albis medullosis,
mesocarpio basal! fibroso et carnoso.

DESCRIPTION OF ALL SPECIMENS EXAMINED:
Tree 10 m tall, 15 cm in diameter; bark light
brown, sparsely muricate; prop roots 1-2 m
long, 3-3.5 cm in diameter, pale brown, muricu-

Page 246: Revision of Pandanus, 14. Malaya and Singapore — St. John

late in longitudinal lines; leaves 1. 8-2.4 m long,

4.4- 5. 2 cm wide, above olive green and some-
what glaucous, below paler green, coriaceous,
with a strong midrib and above it broadly chan-
neled, at midsection with 45 secondary parallel
veins in each half, no visible tertiary cross veins,
sword-shaped, gradually tapering to the 15-30
cm subulate trigonous apex, this 10 cm down
0.9-1. 3 mm wide, the base amplexicaul and
unarmed, but beginning at 10 cm the margins
with spines 5-6.5 mm long, 11-30 mm apart,
stout subulate, arcuate ascending, pale; the mid-
rib below beginning at 22 cm with spines 6 mm
long, 15-35 mm apart, similar but reflexed; at
midsection the margins with prickles 3.5-4 mm
long, 11-18 mm apart, subulate, closely ap-
pressed ascending, reddish-tipped; the midrib
below with similar ascending prickles 3 mm
long, 15-25 mm apart; on the subulate apex
the margins and midrib below with subulate-
serrae 0. 5-0.8 mm long, 2-11 mm apart; syn-
carp single, terminal, 18 cm long, 15 cm in
diameter, subglobose, bearing numerous pha-
langes, these 6.7-73 cm long, 2.7-3. 5 cm wide,

2.4- 3. 1 cm thick, the apex partly green, the
remainder red-orange, 5-6-angled, the upper 2A
free, oblong-elliptic or oblong-obovoid, the apex
low convex, the sides gently curved, rugulose
but even so somewhat shining, lateral sutures
none; central apical sinuses 4.5-53 mm deep
and V-shaped when with narrowly conic apices
(but on the occasional phalange, probably a
subbasal one, with low pyramidal apices, the
sinuses only 2-33 mm deep); carpels 6-10, the
apices usually narrowly conic, the outer ones of
the same thickness but about lA wider, asym-
metric and more or less flattened distally, erect
or slightly outwardly curved; the lower of
phalange fleshy distended with shoulders dis-
tended to 3-4 mm all around, thus 25-35 mm
in diameter; stigmas 3-33 mm long, elliptic,
creased, brown, papillose; proximal sinus run-
ning Vz-Vz way to valley bottom; endocarp
median, 23-3 cm long, dull mahogany-brown,
bony, massive, the lateral walls 3-4 mm thick,
the inner surfaces shining; seeds 13-18 mm
long, 43 mm in diameter, fusiform; upper meso-
carp forming in each carpel apex a cavern, the
lateral ones 23 cm long, filled with the white
membranes of an aerenchyma; basal mesocarp
fibrous and fleshy.

HOLOTYPUS: Singapore, Kranji village, cut-
over thicket near house, 2 m. alt., July 10, I960,

H. St. John 26,375 (bish).

specimens EXAMINED: Malaya, Pahang, Te-
lok Sisik, Kuantan, near the sea, 4 Dec. 1924,

I. H. Bur kill & Md . Haniff 17,346 (sing).

DISCUSSION: P. carnosus is a member of the

section Pandanus . It is one of the curious Asiatic
species with fleshy, enlarged bases of the pha-
langes. None of the described species are close
relatives, but it is related to an undescribed one
from Okinawa, represented by the collection
St. John 25,950.

The new epithet is the Latin adjective car-
nosus, fleshy, in allusion to the fleshy base of the
phalange.

Pandanus glohosus sp. nov. (sect. Pandanus)
Fig. 159

DIAGNOSIS HOLOTYPI: Arbor ? ramis 3 cm
diametro brunneis rugosis, foliis 81-90 cm
longis 33 cm latis supra viridibus infra pallidi-
oribus et apparente glands late sulcatis coriaceis
in sectione mediali cum 38 nervis parallelis
secundariis in quoque medio nervis tertialis nul-
lis laminis gladiformatis sensim in apice 20 cm
longo subulato trigono diminuentibus eo 10 cm
ex apice 2 mm lato basi amplexicauli inermi sed
ex 5 cm marginibus cum aculeis 33-5 mm
longis 5-12 mm separatis arcuatis subulatis ad-
scendentibus apicibus brunneis, midnervo infra
ex 5 cm cum aculeis 2-3 mm longis 6-18 mm
separatis simulantibus sed reflexis, in sectione
mediali marginibus cum aculeis 2.8-3 mm longis
5-12 mm separatis subulatis valde adpressis
adscendentibus, midnervo infra cum aculeis
0.8-1 mm longis 5-12 mm separatis subulatis
adscendentibus, in apice subulato marginibus et
midnervo infra cum aculeis 03-0.8 mm longis
4-8 mm separatis subulatis adpresse adscenden-
tibus, inflorescentia foeminea terminal! cum
syncarpio solitario, pedunculo 9 cm longo 8 mm
diametro trigono folioso-bracteato, syncarpio 8.8
cm diametro globoso cum 26 phalangibus eis
3. 6-3 .9 cm longis 23-2.8 cm latis 2.2-23 cm
crassis breviter crasse pyriformatis 6-angulosis
lateribus planis vel subcurvatis laevibus lucidis
in sicco pallide brunneis parte 2A supera libera
apice depresse convexo, suturis lateralibus nul-
lis, sinibus apicalibus centralibus 2-33 mm pro-

10 cm.

scm.

Fig. 159. Pandanus globosus St. John, from holotype. a, Syncarp, X Vl\ b, phalange, lateral view, X 1;
c, phalange, longitudinal median section, X 1; d, phalange, apical view, X 1; e, carpel apex, stigma, and prox-
imal sinus, oblique view, X 4; f, leaf base, lower side, X 1; S> leaf middle, lower side, X 1; ^ leaf apex, lower
side, X 1-

I — 1 — — — »

Fig. 160. Pandanus inclinatus St. John, from holotype. a, Habit, X 1/200; b, phalange, lateral view, X 1;
c, phalange, longitudinal median section, i< 1; d, phalange, apical view, X 1; e> f> carpel apices, stigmas, and
proximal sinuses, oblique view, X 4; g, bark with adventitious rootlets, X 1-

o.2 mm.. o 5cm.

PACIFIC SCIENCE, Vol. XVII, January 1963

fundis late V-formatis rectis vel subcurvatis,
carpelis 5-6 apicibus conicis angulatis basi ro-
tundato apice rugoso eis centralibus quando sunt
minoribus, stigmatibus 2.5-3 mm longis cordato-
orbicularibus vel ellipticis brunneis papillosis
prominentibus obliquis centripetalibus, sinibus
proximalibus Vi —2A ad fondam extentis, endo-
carpio minime submediali brunneo osseoso la-
teribus 1.5-2 mm crassis intra obscure brunneis
lucidis, seminibus 10-14 mm longis 3.5-4 mm
diametro ellipsoideis vel obliquiter ellipsoideis,
mesocarpio supero in apice quaque carpellae
cavernam cum fibris paucis et membranis pal-
lidis medullosis formanti, mesocarpio basali fib-
roso et carnoso.

diagnosis OF HOLOTYPE: Apparently tree-
like; branch 3 cm in diameter, brown, roughened
by crowded leaf scars; leaves 81-90 cm long,
3.5 cm wide, green above, below paler green
and apparently glaucous, with a broad V-shaped
furrow above the midrib, coriaceous, at mid-
section with 38 secondary veins in each half, no
visible tertiary cross veins, sword-shaped, taper-
ing gradually to the 20 cm subulate, trigonous
apex, this 10 cm down 2 mm wide, the base am-
plexicaul, unarmed, but beginning 5 cm up the
margins with prickles 3.5-5 mm long, 5-12 mm
apart, arcuate subulate, fully ascending, brown-
tipped; the midrib below beginning at 5 cm
with prickles 2-3 mm long, 6-18 mm apart,
similar but reflexed; at midsection the margins
with prickles 2.8-3 mm long, 5-12 mm apart,
subulate, flat appressed, ascending; the midrib
below with prickles 0.8-1 mm long, 5-12 mm
apart, subulate, ascending; on the subulate tip
the margins and midrib below with subulate,
appressed ascending prickles 0. 5-0.8 mm long,
4-8 mm apart; pistillate inflorescence terminal,
bearing one syncarp; peduncle 9 cm long, 8 mm
in diameter, 3 -sided, leafy bracted; syncarp 8.8
cm in diameter, globose, bearing about 26 pha-
langes, these 3-6— 3-9 cm long, 2. 5-2.8 cm wide,
2.2-2. 5 cm thick, short, thick pyriform, 6-angled,
the sides plane or gently curved, smooth, shin-
ing, when dried light brown, upper 2A free, the
apex low convex; lateral sutures none; central
apical sinuses 2-3.5 mm deep, wide V-shaped,
straight or gently curved; carpels 5-6, the apices
with a rounded base and conic, angled, wrinkled
tip, the central ones, when present, slightly

the smaller; stigmas 2.5-3 mm long, cordate-
orbicular to -elliptic, brown, papillose, prom-
inent, oblique, centripetal; proximal sinus
running l/2-2A way to valley bottom; endocarp'
slightly inframedian, brown, bony, the lateral
walls 1.5-2 mm thick, the inner surfaces pol-
ished, dark brown; seeds 10-14 mm long; 3.5-4
mm in diameter, ellipsoid or obliquely so; upper
mesocarp forming in the apex of each carpel a
cavern with a few fibers and with pale, medul-
lary membranes; basal mesocarp fibrous and
fleshy.

HOLOTYPUS: Malaya, Kedah, w. coast, P.
Dayang Banting, Pfulau] Langkawi, sea level,
on limestone, 27 Nov. 1934, M. R. Henderson
29J33 (SING).

SPECIMENS EXAMINED: Borneo: Karimata
groep, Poelau, Karimata, 23/3/1931, Mondi
133 (BO).

Celebes: Kota Menado, strand, 31 /XII/ 1894,
Ko orders 1 8,462 (bo).

DISCUSSION: P. globosus is a member of the
section Pandanus, as is its closest relative, the
Vietnamese species P. subulatus St. John, a spe-
cies which has the syncarp suborbicular-ellipsoid,
10-10.5 cm in diameter; carpel apices pyramidal
or oblate-pyramidal; stigmas 1.5-2 mm long,
black; endocarp supramedian; the leaf apex at
the point 10 cm down 3.5 mm wide; the mar-
gins near the base with spines 5-6 mm long,
ascending at 45°; P. globosus has the syncarp
globose, 8.8 cm in diameter; carpel apices low
convex; stigmas 2.5-3 mm long, brown; endo-
carp inframedian; leaf apex at the point 10 cm
down 2 mm wide; the margins near the base
with prickles 3.5-5 mm long, fully ascending.

The new epithet is the Latin adjective globo-
sus, spherical, and is given in reference to the
spherical head of fruits.

Pandanus inclinatus sp. nov. (sect. Pandanus).

Figs. 160, 170c

diagnosis HOLOTYPI: Arbor 8 m alta 13 cm
diametro, cortice pall id e brunneo muricato et
cum radicillis adventivis adpresse adscendentibus
ad 1 cm longis, radicibus fulturosis nullis, foliis-
1. 5-1.7 m longis 4.4-4.7 cm lads coriaceis supra
obscure olivaceo-viridibus infra viridibus spinis
omnibus albis cum apicibus brunneis lamina
1-sulcatis et lateribus semirevolutis in sectione

Page 250: Revision of Pandanus , 14. Malaya and Singapore — St. John

mediali cum 38-40 nervis parallelis secundariis
in quoque dimidio, nervis tertialis nullis, lamina
gladiformi in apice caudato trigono subulato
15-20 cm longo diminuenti eo 10 cm ex apice
1 mm lato, basi amplexicauli et inermi sed ex

3- 7 cm marginibus cum aculeis 3.5-6 mm iongis
10-30' mm separatis crassiter subulatis subarcu-
atis adscendentibus, midnervo infra ex 7-8 cm
cum aculeis 3-5 mm Iongis 20-50 mm separatis
crassiter subulatis valde reflexis, in sectione
mediali marginibus cum aculeis 3-3.5 mm
iongis 7-17 mm separatis crassiter subulatis ad-
scendentibus valde adpressis, midnervo infra
cum aculeis 2.5-3 mm Iongis 8-23 mm separatis
subulatis adscendentibus, in apice subulato mar-
ginibus et midnervo infra cum subulato serrulis
0.5-1 mm Iongis 5-10 mm separatis vel in
loculis nullis, inflorescentia foeminea terminali
cum syncarpio uno, pedunculo 30 cm longo 13-
15 mm diametro trigono folioso-bracteato, syn-
carpio 16-23 cm longo 13-20 cm diametro cum
44-61 phalangiis eis 6-6.8 cm Iongis, climb
dio supero 3.2— -4.3 cm lato 23-3.7 cm crasso
oblongo-obovoideo truncate vel subtmncato
aurantiaco-rubro apice subviridi 5-7-anguloso
lateribus subcurvatis vel subplanis laevibus et
lucidis, suturis lateralibus nullis, sinibus centrali-
bus apicalibus 0.7-2 mm profundis rectis vel
subcurvatis, dimidia infera phalangis turbinata
carnosa humeris carnosis 3-6 mm latis carne
subsucrato comes to aurantiaco-rubro, carpelis
6-12 plerumque 7-9 (sed in gemine 18 et pha-
langio 5.5 cm lato 4.4 cm crasso) interioribus
paene minoribus, apicibus planatis sed pauce
oblato-pyramidalibus, stigmatibus 2-3 mm Ion-
gis eilipticis vel late ellipticis obliquis brunneis
sulcatis pluribus rnarginalium truncatis ex piano
oblique et cum concavite parvo distali, sinibus
proximalibus profundis Vi-Vd ad fondam ex-
tent is, endocarpio 3 cm longo mediali osseoso
obscure mahogani-colorato intra lucido lateribus
2.5 mm crassis, seminibus 14-17 mm Iongis

4- 5.5 mm diametro ellipsoideo, mesocarpio
supero in quoque carpelo cavernam cum fibris
panels et aerenchyma cum membranis albis
formanti, mesocarpio basali fibroso et carnoso.

DIAGNOSIS OF holotype: Tree, 8 m tall, 13
cm in diameter; bark light brown, muricate,
and with a few appressed ascending rootlets up
to 1 cm in length; prop roots none; leaves 1.5-

1.7 m long, 4.4-47 cm wide, coriaceous, above
dark olive green, below green, the spines all
white with minute brown tips, blade broad fur-
rowed above the midrib, the sides down curved,
at midsection with 38-40 secondary parallel
veins in each half, no tertiary cross veins, sword-
shaped, tapering upwards into a 15-20 cm sub-
ulate, trigonous, caudate apex, this 10 cm down
1 mm wide, the base amplexicaul and unarmed,
but starting at 3-7 cm up the margins with
prickles 3.5-6 mm long, 10-30 mm apart, stout-
subulate, slightly arcuate, ascending; the midrib
below beginning at 7-8 cm with prickles 3-5
mm long, 20-50 mm apart, stout subulate,
sharply reflexed; at midsection the margins with
prickles 3-3.5 mm long, 7-17 mm apart, stout
subulate, ascending, closely appressed; the mid-
rib below with prickles 2.5-3 mm long, 8-23
mm apart, subulate, ascending; on the subulate
apex the margins and midrib below with sub-
ulate serrulations 0.5-1 mm long, 5-10 mm
apart or even lacking for distances on the mar-
gins; pistillate inflorescence terminal, bearing
one syncarp; peduncle 30 cm long, 13-15 mm
in diameter, 3 -sided, leafy bracted; syncarp lb-
23 cm long, 13-20 cm in diameter, bearing
44-61 phalanges, these 6-6.8 cm long, the upper
half 3.2-4 .3 cm wide, 2.3-37 cm thick, oblong
obovoid, truncate or subtruncate, orange-red,
the color becoming paler upwards to the partly
greenish apex, 5-7-angled, the sides gently
curved or almost plane, smooth and shining;
lateral sutures none; central apical sinuses 07-2
mm deep, the bottom straight or gently curved,
V-shaped, then immediately wide flaring; lower
half of phalange turbinate above the truncate
base, fleshy enlarged, the shoulders 3-6 mm
wide, the flesh slightly sweet, nonirritating,
orange-red; carpels 6-12, mostly 7-9 (but in a
double 18 and this one 5.5 cm wide, 4.4 cm
thick), the central ones only slightly the smaller,
the apices flattened but perceptibly oblate pyra-
midal; stigmas 2-3 mm long, elliptic or broadly
so, oblique, brown, creased, many of the mar-
ginal ones truncate by the visor-like edge of a
distal oblique plane terminating in a small con-
cavity; proximal sinus deep, running Vi-Vi way
to valley bottom; endocarp median, 3 cm long,
bony, dark mahogany-colored, the inner sufaces
shining, the lateral wall 2.5 mm thick; seeds

PACIFIC SCIENCE, Vol. XVII, January 1963

14—17 mm long, 4-5.5 mm in diameter, ellip-
soid; upper mesocarp forming in the apex of
each carpel a cavern with a few fibers and an
aerenchyma of white medullary membranes;
basal mesocarp fibrous and fleshy.

HOLOTYPUS: Singapore, Kranji Nature Re-
serve, brackish swamp with Sonneratia caseola-
ris, Hibiscus tiliaceus, Acrostichum aureum,
July 10, I960, H. St. John 26,378 (bish).

DISCUSSION : P. inclinatus is a member of the
section Pandanus, as is its closest relative, P.
vietnamensis St. John, of Vietnam, a species
with the syncarp 27 cm long, with 106 phal-
anges, these 5.8-6 cm long, the apex low con-
vex; stigma 3.5-5 mm long or wide, black;
leaves 0.99-1.49 m long, 5-6 cm wide, the mar-
gin near the base with spines 4-11 mm long,
the nearby midrib below with spines 6-8 mm
long. P. inclinatus has the syncarp 16-23 cm
long, with 44-6 1 phalanges, these 6-6.8 cm
long, the apex truncate or subtruncate; stigmas
2-3 mm long, brown; leaves 1.5-1. 7 m long,
4.4-4.7 cm wide, the margin near the base with
prickles 3.5-6 mm long, and the nearby midrib
below with prickles 3-5 mm long.

The new epithet is the Latin participle in-
clinatus, inclined, given in reference to the ob-
lique stigmas.

Pandanus incrassatus sp. nov. (sect. Pandanus)
Figs. 161, 171

DIAGNOSIS HOLOTYPI: Arbor 7-8 m alta 8-
10 cm diametro, cortice griseo remote muricato
in ramis vetustioribus et trunco cum radicillis
adventivis paucis adpresse adscendentibus, radi-
cibus fulturosis 5-10 dm longis 10-13 mm
diametro pallide brunneis cum radicillis brevi-
bus spiniformatis in lineis, foliis 1.45-1.7 m
longis 4.2-4.3 cm latis coriaceis supra olivaceo-
viridibus infra viridibus et glaucis spinis albis
deinde apicibus brunneis, laminis 1-sulcatis et
cum plicis duobus rotundatis in sectione mediali
cum 34-35 nervis secundariis in quoque medio
nervis tertialis nullis gladiformatis sensim in
apice 30 cm longo subulato trigono diminuen-
tibus eo 10 cm ex apice 1.1 mm lato, basi arn-
plexicauli et inermi albo sed ex 7—9 cm mar-
ginibus cum spinis 3.5-5 mm longis 15-25 mm
separatis arcuatis crassiter subulatis adscenden-
tibus, midnervo infra ex 9-12 cm cum spinis

3-4 mm longis 20-28 mm separatis valide sub-
ulatis reflexis, in sectione mediali marginibus
cum aculeis 2.5-3 mm longis 10-20 mm sepa-
ratis arcuatis subulatis adpresse adscendentibus,
midnervo infra cum aculeis 1.5-2 mm longis
12-20 mm separatis arcuatis subulatis adscen-
dentibus, in apice subulato marginibus et mid-
nervo infra cum subulato-serrulis 0. 5-0.7 mm
longis 3-5 mm separatis, inflorescentia foeminea
terminali cum syncarpio solitario pendenti,
pedunculo 23 cm longo 1.5 cm diametro trigono
folioso-bracteato, syncarpio 15 cm longo 14.5
cm diametro subgloboso cum 54 phalangibus
eis 4.6-5 .2 cm longis media sup era 2.5-3. 1 cm
longa, 3-3.6 cm lata 2.3-3 cm crassa obovoidea
plano-truncata viridi deinde rubro-aurantiaca 5-
6-angulosa lateribus subcurvatis infra laevibus
et lucidis supra rugulosis et sublucidis, suturis
lateralibus nullis, sinibus apicalibus centralibus
plerumque (et in phalangibus lateralibus api-
calibusque omnibus) 1-2 mm profundis late V-
formatis rectis vel subcurvatis (vel in phalangi-
bus subbasalibus paucis 2-3 mm profundis),
media infera phalangis carnoso-incrassata 2.8- j

aurantiaco-rubra infra pallidiori, humeris 2-5
mm latis undulatis carne saccharosa sed non
consumpta, carpelis 5-8 centralibus l/2~2A tarn j,
grandibus quam marginalibus, phalangibus di- |
midii superi cum apicibus carpeli subtruncatis j
vel oblatiori-pyramidalibus sed eis dimidii inferi
cum apicibus oblato-pyramidalibus, stigmatibus j
2-3 mm longis ellipsoideis obscure brunneis
papillosis sulcatis obliquis prominentibus centri-
petalibus eis marginalibus plerumque cum supra
stigmatem galea distali cartilaginea, sinibus
proximalibus Vi~2A ad fondam extentis, endo-
carpio 2.5 cm longo mediali osseoso obscure
brunneo intra lucido lateribus 2.5 mm crassis,
seminibus 12 mm longis 4 mm diametro ellip-
soideis, mesocarpio supero in apice quique car-
peli cavernam cum membranis albis lucidis
medullosis formanti, mesocarpio basali fibroso
et carnoso.

DIAGNOSIS OF HOLOTYPE: Tree 7-8 m tall,
8-10 cm in diameter; bark gray, remotely muri-
cate, and the older trunk with a few appressed
ascending adventitious rootlets; prop roots 5-10
dm long, 10-13 mm in diameter, light brown,
with remote, short, spinelike rootlets in vertical

Fig. 161. Pandanus incrassatus St. John, from holotype. a, Habit, X 1/100; b, dried phalange, lateral view,
X 1; c, phalange, longitudinal median section, X 1| ^ phalange, apical view, X 1; ^ carpel apex, stigma, and
proximal sinus, oblique view, X 4; /, prop root with spinelike rootlets, X lj S> leaf base, lower side, X 1>
h, leaf middle, lower side, X 1; h leaf apex, lower side, X 1-

5 vnw,.

PACIFIC SCIENCE, Vol. XVII, January 1963

rows; leaves 1.43-1.7 m long, 4.2-4.3 cm wide,
coriaceous, above olive green, below green and
glaucous, the spines white, becoming brown-
tipped, 1 -ribbed and with two rounded pleats,
at midsection with 34-35 secondary parallel
veins in each half, no visible tertiary cross veins,
sword-shaped, long tapering to a 30 cm subu-
late trigonous apex, this 10 cm down 1.1 mm
wide, the base amplexicaul and unarmed, white,
beginning at 7-9 cm the margins with spines

3.5- 5 mm long, 15-25 mm apart, arcuate, heavy
subulate, ascending; the midrib below begin-
ning at 9-12 cm with spines 3-4 mm long, 20—
28 mm apart, stout subulate, reflexed; at mid-
section the margins with prickles 2.5-3 mm
long, 10-20 mm apart, arcuate subulate, ap-
pressed ascending; the midrib below with
prickles 1.5-2 mm long, 12-20 mm apart, arcu-
ate subulate, ascending; near the apex the mar-
gins and midrib below with subulate serrula-
tions 0.5-0. 7 mm long, 3-5 mm apart; pistillate
inflorescence terminal, bearing 1 pendent syn-
carp; peduncle 23 cm long, 1.5 cm in diameter,
3 -sided, leafy-bracted; syncarp 15 cm long, 14.5
cm in diameter, subglobose, bearing 54 phal-
anges, these 4.6-5. 2 cm long, the upper half

2.5- 3. 1 cm long, 3-3.6 cm wide, 2.3-3 cm
thick, obovoid, flat truncate, green, turning red-
orange, the shade deepening downwards, 5-6-
angled, the sides gently curving, below smooth
and shining, above rugulose and less shiny;
lateral sutures none; central apical sinuses
mostly (that is, for the lateral and apical phal-
anges) 1-2 mm deep, wide V-shaped, straight
or gently curving (on a few subbasal ones 2-3
mm deep); lower half of phalange fleshy en-
larged, 2. 8-3. 6 cm wide, 2.4-3. 1 cm thick, wide
cuneate, vermilion above, gradually paling be-
low, the shoulders 2—5 mm wide, scalloped;
pulp sweetish, innocuous, but not eaten; carpels
5-8, the central 1-2 being Vl-2A the size of the
marginal ones; on the phalanges of the upper
half the carpel apices subtruncate or very oblate
pyramidal, on those of the lower half the carpel
apices oblate-pyramidal; stigmas 2-3 mm long,
ellipsoid, dark brown, papillose, creased, oblique,
prominent, centripetal, the marginal ones mostly
with a distal, cartilaginous visor partly project-
ing over the stigma; proximal sinus running
Vl—Vi way to valley bottom; endocarp median

2.5 cm long, bony, dark brown, the inner sur-
faces shining, the lateral walls 2.5 mm thick;
seeds 12 mm long, 4 mm in diameter, ellipsoid;
upper mesocarp forming a cavern in each carpel
apex with white, shining, medullary mem-
branes; basal mesocarp fibrous and fleshy.

holotypus: Singapore, Kranji Nature Re-
serve, mangrove swamp, with Sonneratia caseo-
laris, Hibiscus tiliaceus, Acanthus ilicifolius,
near sea level, 7 July I960, H. St. John 26,374

(BISH).

DISCUSSION: P. incrassatus is a member of
the section Pandanus, as is its closest relative,
the Vietnamese species P. reversispiralis St.
John, a species with the syncarp oval-subglo-
bose; phalanges 4.3-4.6 cm long, orange, the
apex low convex; stigmas all terminal, exposed;
leaves 70-130 cm long, 4.8-6 cm wide, slightly
glaucous above, at midsection the margins with
prickles 3—4 mm long. P. incrassatus has the
syncarp subglobose; phalanges 4.6-5. 2 cm long,
red-orange, the apex truncate; marginal stigmas
covered by a visor; leaves 145-170 cm long,
4.2-4. 3 cm wide, olive green, not glaucous
above, and at midsection the margins with
prickles 2.5-3 mm long.

The new epithet is the Latin adjective incras-
satus, thickened, and is given with allusion to
the thickened base of the phalange.

Pandanus ohtusus sp. nov. (sect. Pandanus )
Fig. 162

DIAGNOSIS HOLOTYPI: Arbor ? foliis 85-88
cm longis 3.4 cm latis coriaceis 1-sulcatis lateri-
bus arcuatis in sectione mediali cum 37-39
nervis parallelis secundariis in quoque medio
eis infra conspicuis, nervis tertialis nullis, lami-
nis gladiformatis sensim in apice 15 cm longo
subulato trigono diminuentibus eo 10 cm ex
apice 2.5 mm lato, basi amplexicauli et inermi
sed ex 5 cm marginibus cum aculeis 2.5-4 mm
longis 8-14 mm separatis arcuatis subulatis pal-
lidis in 45° adscendentibus, midnervo infra ex

8.5 cm cum aculeis 3.5-4 mm longis 12-20 mm
separatis subulatis valde reflexis basibus crassiter
conicis, in sectione mediali marginibus cum
aculeis 3-3.5 mm longis 6-11 mm separatis
arcuatis subulatis adscendentibus apicibus ru-
bris, midnervo infra angusto salienti cum acu-
leis 1.5-2 mm longis 7-10 mm separatis arcu-

5 cm.

Fig. 162. Pandanus obtusus St. John, from holotype. a, Syncarp, X Vl\ b, phalange, lateral view, X 1; i
phalange, longitudinal median section, X 1; d; phalange, apical view, X 1; <?, carpel apex, stigma, and prox-
imal sinus, oblique view, X 4; /, leaf base, lower side, X 1; g, leaf middle, lower side, XI \h> leaf apex, lower
side, X 1.

smm.

PACIFIC SCIENCE, Vol. XVII, January 1963

atis subulatis adscendentibus, in apice subulato
marginibus et midnervo infra cum subulato-
serrulis 0.3-07 mm longis 3-7 mm separatis,
inflorescentia foeminea cum syncarpio unico
terminali, pedunculo 23 cm et plus longo cla-
vato in apice 11 mm diametro subtrigono, syn-
carpio globoso ? 12 cm diametro cum circa 38
phalangibus eis 5-5.5 cm longis 3-3.8 cm latis
27-3.3 cm crassis cuneatis 5-6-angulosis in
sicco pallide brunneis lateribus laevibus lucidis
subcurvatis vel subplanis parte l/$ supera libera
apice convexo vel alte convexo suturis lateral i-
bus nullis, sinibus centralibus apicalibus olim
lineas brunneas, carpelis 8-10 apicibus rotun-
datis interioribus minoribus et l/z tarn grandibus
quam rnarginalibus, stigmatibus 2-2.5 mm
longis cordatis vel reniformibus obliquis centri-
petalibus, sinibus proximalibus brevibus lA~lA
ad fondam extentis, endocarpio mediali 2.5-3
cm longo osseoso extra brunneo intra pallidi
lateribus 2 mm crassis, seminibus 12 mm longis
3.5 mm diametro obliquiter ovoideis, mesocar-
pio supero in apice carpello quoque cavernam
formanti eis lateralibus majoribus cum fibris
longitudinalibus et membranis albis medullosis,
mesocarpio basali fibroso et carnoso.

DESCRIPTION OF ALL SPECIMENS EXAMINED:
Probably treelike; leaves 85-130 cm long, 3 A
cm wide, coriaceous, furrowed above the mid-
rib, the two halves gently arching, at midsec-
tion with 21-39 secondary parallel longitudinal
veins in each half, these conspicuous below, no
visible cross veins, blade swordlike, tapering
upwards and gradually narrowed to a 15 cm
subulate, trigonous apex which 10 cm down
is 2.5 mm wide, the base amplexicaul and
unarmed, but beginning at 5 cm the margins
with prickles 2.5-4 mm long, 8-14 mm apart,
arcuate subulate, pale, ascending at 45°; the
midrib below beginning at 8.5 cm with prickles
3. 5-4. 5 mm long, 12-20 mm apart, the base
heavy conic, the rest subulate, fully reflexed; at
midsection the margins with prickles 3-3.5 mm
long, 6-11 mm apart, arcuate subulate, ascend-
ing, reddish-tipped; the midrib below narrow
and salient, with prickles 1.5-2 mm long, 7-10
mm apart, arcuate subulate, ascending; on the
subulate apex the margins and midrib below
with subulate-tipped serrations 0.3-07 mm
long, 3-7 mm apart; pistillate inflorescence

with a single, terminal syncarp; peduncle more
than 23 cm in length, clavate and at apex 11
mm in diameter, somewhat 3 -sided; syncarp
apparently globose and 12 cm in diameter, and
bearing about 38 phalanges, these 5-5.5 cm
long, 3-3.8 cm wide, 27-3.3 cm thick, cunei-
form, 5-6 angled, when dried pale brown, the
sides smooth and shiny, gently curving or flat-
tish, free in upper Vd, the apex convex or high
convex, lateral sutures none, central apical siT
nuses mere brown lines on the obtuse apex;
carpels 8-10, the apices rounded, not raised,
the inner ones smaller, about Vi the size of the
marginal ones; stigmas 2-2.5 mm long, cordate
to reniform, oblique, centripetal; proximal sinus
a very short crack or hole by the stigma, run- j
ning V4-V3 way to the valley bottom; endocarp j
median, 2.5-3 cm long, bony, pale within, dark j
brown near the margins, the seed cavities shin- j
ing, the lateral walls 2 mm thick; seeds 12 mm j
long, 3.5 mm in diameter, obliquely ovoid;
upper mesocarp forming in the apex of each
carpel a cavern, the lateral ones thrice the larger,
with longitudinal fibers and white medullary I
membranes; basal mesocarp fibrous and fleshy.

HOLOTYPUS: Malaya, Pahang, Beserah, Kuan-
tan, sea level, 18 Aug. 1929, M. R, Henderson
22,746 (sing):

SPECIMENS EXAMINED: Singapore, 15 Feb.
1870, J. S. Gfoodenough}, (SING). This has
leaves only 2 cm wide, and with 21 secondary
veins on a side, but the phalanges seem iden-
tical.

DISCUSSION: P. obtusus is a member of the
section Pandanus. In that section the most simi-
lar species seems to be a Marquesan one, P,
tectorius Soland. var. uapensis F. Br., which has
the phalanges 5.4-6 cm long, 4.5 cm wide, the
apical and lateral sutures very shallow or super-
ficial; apical central sinuses 1 mm deep; stigmas
3 mm long; endocarp supramedian, but the
leaves and syncarp are unknown. P. obtusus has
the phalanges 5-5.5 cm long, 3-3.8 cm wide,
the lateral sutures none, the apical central si-
nuses mere tessellate rulings; stigmas 2-2.5 mm j
long; and the endocarp median.

The new epithet is the Latin word obtusus , j
blunt or obtuse, and is given with reference to
the obtuse phalange and carpel apices.

Fig. 163. Pandanus rubricoloratus St. John, from holotype. a, Syncarp, X V2; dried phalange, lateral
view, XI \ c, phalange, longitudinal median section, X 1; d, phalange, apical view, X 1; carpel apex, stigma,
and proximal sinus, oblique view, X 4; f, leaf base, lower side, X 1; ^ leaf middle, lower side, X 1; ^ leaf
apex, lower side, X 1-

PACIFIC SCIENCE, Vol. XVII, January 1963

Pandanus rubric olor atm sp. nov. (sect. Pan-
danus)

Fig. 163

DIAGNOSIS HOLOTYPI: Arboriformis 3-5 m
alta, foliis 66-73 cm longis 2.8-3 cm latis cori-
aceis supra viridibus infra pallidioribus supra
midnervum sulcatis in sectione mediali cum 27
nervis parallelis secundariis in quoque medio
nervis tertialis nullis laminis gladiformatis in
apice 12-15 cm iongo subulato sensim diminu-
entibus eo 10 cm ex apice 4 mm lato basi am-
plexicauli et inermi sed ex 3—4 cm marginibus
cum aculeis 2.5-5 mm longis 4-18 mm sepa-
rates subulatis pallidis adscendentibus, mid-
nervo infra ex 8-10 cm cum aculeis 2.5-3 mm
longis 16-30 mm separatis simulantibus sed
adpresse reflexis, in sectione mediali marginibus
cum aculeis 2-2.5 mm longis 4-12 mm sepa-
ratis subulatis adscendentibus apicibus brun-
neis, midnervo infra cum aculeis simulantibus
1.5-2 mm longis 6-13 mm separatis; in apice
subulato marginibus et midnervo infra cum
subulato-serrulis 0.6-1 mm longis 2-6 mm sepa-
ratis; inflorescentia foeminea terminal! cum
syncarpio solitario, pedunculo 15 cm et plus
longo 8 mm diametro exclavatis trigono folioso-
bracteato, syncarpio 9 cm longo 8.5 cm diametro
globoso cum circa 34 phalangibus, els 3.6 cm
longis 2 .4-2 .7 cm latis 2-2.4 cm crassis rubris
parte 5/9 supera libera, hac late ellipsoidea 5-
6-angulosa apice depresse convexo lateribus sub-
curvatis laevibus suturis lateralibus nullis, sini-
bus apicalibus centralibus 0.5-2 mm profundis
vadosis, carpelis 6-9 (in gemine 12 et phalangio
3.3 cm lato) apicibus depresse pyramidalibus
angulatis rugosis (in phalangibus basalibus
apicibus depresse convexis) parte 4/9 inf era
carnoso-incrassata, stigmatibus 2-3.5 mm latis
cordatis griseis horizontalibus vel obliquis cen-
tripetalibus, sinibus proximalibus latis Vl ad
fondam extentis, endocarpio mediali 15 mm
longo osseoso obscure brunneo lateribus 1.5
mm crassis intra papillosis et sublucidis, semi-
nibus 8-11 mm longis obliquiter ellipsoideis,
mesocarpio supero in apice quoque carpelo
cavernam cum fibris paucis et membranis brun-
neis medullosis, mesocarpio basali fibroso et
carnoso.

diagnosis of holotype: Treelike, 3-5 m
tall; leaves 66-73 cm long, 2.8-3 cm wide,

coriaceous, green above, paler beneath, fur-
rowed above the strong midrib, the sides arch-
ing, at midsection with 27 parallel secondary
veins in each half, no tertiary cross veins, blade
sword-shaped, tapering upwards and gradually
tapering to the 12-15 cm subulate apex which
10 cm down is 4 mm wide, the base amplexi-
caul and unarmed, but beginning 3-4 cm up
the margins with prickles 2.5-5 mm long, 4-18
mm apart, subulate, ascending, pale; midrib
below beginning at 8-10 cm with prickles 2.5-
3 mm long, 16-30 mm apart, similar but re-
flexed, appressed; at midsection the margins
with prickles 2-2.5 mm long, 4-12 mm apart,
subulate, ascending, brown-tipped; the midrib
below with similar prickles 1.5-2 mm long,
6-13 mm apart; on the subulate tip the mar-
gins and midrib below with subulate serrations
0.6-1 mm long, 2-6 mm apart; pistillate in-
florescence terminal, bearing one syncarp; pe-
duncle more than 15 cm long, 8 mm in diam-
eter, ' not ' clavate, 3-sided, leafy bracted; syn-
carp 9 cm long, 8.5 cm in diameter, globose,
bearing about 34 phalanges, these 3.6 cm long,
2.4-27 cm wide, 2-2.4 cm thick, red, upper
5/9 free, this part broadly ellipsoid, 5-6-angled,
the apex low convex, the sides gently curved,
smooth, lateral sutures none; central apical si-
nuses 0.5-2 mm deep, shallow; carpels 6-9 (in
a double 12, and the phalange 3.3 cm wide),
the apices low pyramidal, angled and wrinkled
(on the basal phalanges the carpel apices low
convex); lower 4/9 of phalange fleshy enlarged
as shown by persistent shoulders; stigmas 2-3.5
mm wide, cordate, gray, horizontal or oblique,
centripetal; proximal sinus wide, running V2
way to valley bottom; endocarp median, 15 mm
long, bony, blackish brown, the lateral walls 1.5
mm thick, the inner surfaces papillose, a little
shiny; seeds 8-11 mm long, oblique ellipsoid;
upper mesocarp forming in the apex of each
carpel a cavern with a few fibers and an aeren-
chyma tissue of brownish medullary mem-
branes; basal mesocarp fibrous and fleshy.

holotypus: Malaya, Pahang, Pul.au Tioman,
Telok Paya, sea level, 19 May 1927, M. R. Hen-
derson 18,444 (sing).

DISCUSSION: P. rubricoloratus is a member
of the section Pandanus , as is its closest relative,
the Vietnamese species P. reversispiralis St.

lent.

FIG. 164. Pandanus brevicornutus St. John, from holotype. a, Syncarp, X 1; ^ drupe, lateral view, X 1;
c, drupe, longitudinal median section, X 1; ^ drupe, lateral view, X 4; e, drupe, longitudinal median section,
X 4; f, drupe apex and style, apical view, X 4; g, style, distal view, X 10; h, style and stigma, proximal view,
X 10; i, style and stigma, lateral view, X 10; j, style and stigma, proximal view, X 10; k, l, style, distal view,
X 10; m, leaf base, lower side, XI \ n> leaf middle, lower side, X 1; o, leaf apex, lower side, X !•

PACIFIC SCIENCE, Vol. XVII, January 1963

John, which has the syncarp 11-14 cm in diam-
eter, oval-subglobose; phalanges 4.3 -4.6 cm
long, orange; central apical sinuses 3.5-5 mm
deep; endocarp lateral walls 2-3 mm thick; and
the leaves 4.8-6 cm wide, at midsection the
margins with prickles 3-4 mm long, and 5-17
mm apart. P. rubricoloratus has the syncarp 8.5
cm in diameter, globose; phalanges 3.6 cm
long, red; central apical sinuses 0.5-2 mm deep;
endocarp lateral walls 1.5 mm thick; and the
leaves 2.8-3 cm wide, at midsection the mar-
gins with prickles 2—2.5 mm long, and 4—12
mm apart.

The epithet is formed from the Latin words
ruber, red; coloratus, colored, in reference to
the red color of the fruit.

Pandanus brevicornutus sp. nov. (sect. Rykia)
Fig. 164

DIAGNOSIS HOLOTYPI: Frutex, ramis assur-
gentibus in apice 17 mm diametro luteis lucidis,
foliis 41-47 cm longis 27-29 mm latis in medio
sed in basi 18-22 mm latis ligulatis chartaceis
proxima basem supra midnervum sulcatis alibi
planis in sectione mediali cum 19-21 nervis
parallelis secundariis in quoque medio nervis
tertialis transversis semievidentis proxima api-
cem in apice 3.5 cm longo 1.5 mm lato trigono
subulato semiabrupte diminuentibus, basi am-
plexicauli inermi sed ex 3-4 cm marginibus
cum aculeis 1.5—3 mm longis 3—7 mm separatis
subulatis adscendentibus obscure mahogani-
coloratis lucidis, midnervo infra ex 9-10 cm
cum aculeo unico 2.5 mm longo arcuato subu-
lato reflexo, in sectione mediali marginibus cum
aculeis 0.5-1. 5 mm longis 6-11 mm separatis
graciliter subulatis proxime adpressi-adscenden-
tibus, midnervo infra cum aculeis 2-2.5 mm
longis paucis remotis arcuatis subulatis reflexis
vel adscendentibus, in apice subulato marginibus
cum serrulis 0.2— 0.5 mm longis 0.5— 1.5 mm
separatis, midnervo infra cum serrulis simulan-
tibus 2-3 mm separatis, infructescentia cum
syncarpio unico terminal! erecto, pedunculo 10
cm longo 6 mm diametro cum folioso-bracteis
multis, syncarpio 5.2 cm longo 4 cm diametro
late ellipsoideo cum circa 960 drupis eis 16—
18.5 mm longis 2-2.5 mm latis 1.5-2 mm
crassis anguste cylindricis compressis 5-angu-
losis corpore 14-15 mm longo, pileo 5-6 mm

longo basi 2 mm alta semiorbiculari, stylo 3-4
mm longo lanceolato osseoso brunneo lucido
proxime curvato integro vel aliquis bifidis, stig-
mate 1-1.5 mm longo elliptico vel late elliptico
sulcato brunneo papilloso paene ad apicem ex-
tento, endocarpio in parte 14 infera cartilagineo
brunneo lateribus 0.1 mm crassis intra pallidis
exlucidis, semine 3-4 mm longo ellipsoideo
truncato, mesocarpio apicali cavernoso angusto
ellipsoideo 9 mm longo, mesocarpio basali in
lateribus fibroso intra carnoso 4-5 mm longo.

DESCRIPTION OF ALL SPECIMENS EXAMINED:
Shrub; branches assurgent, up to 1.3 m long,
sprawling and forming a thicket, 17 mm in
diameter at apex, yellowish, shining; leaves 35-
50 cm long, 24-29 mm wide at the middle, 18-
22 mm wide near the base, ligulate, chartaceous,
yellowish green, not glaucous beneath, near the
base sulcate over the midrib, elsewhere flat, at
midsection with 18-21 secondary parallel nerves
in each half, the tertiary cross veins barely visi-
ble only near the tip, near the tip rather ab-
ruptly tapering into a 3-5-7 cm subulate trigon-
ous apex 1.5 mm wide, the base amplexicaul
and cream-colored, unarmed, but beginning at
3-4 cm the margins with prickles 1.5-3 mm
long, 3-7 mm apart, subulate, ascending, dark
mahogany colored, shining; the midrib below
9-10 cm up with a prickle 2.5 mm long, arcu-
ate subulate, reflexed; at midsection the mar-
gins with prickles 0.5-1. 5 mm long, 6-11 mm
apart, slender subulate, closely appressed ascend-
ing; the midrib below with a few remote
prickles 2-2.5 mm long, arcuate subulate, re-
flexed or ascending; on the subulate apex the
margins with serrulations 0.2-0. 5 mm long,
0.5-1. 5 mm apart; the midrib below with simi-
lar ones 2-3 mm apart; pistillate infructescence
with a single, terminal, erect syncarp (rarely a
smaller, lateral, secondary one, globose and 2
cm in diameter); peduncle 10-15 cm long, 4-6
mm in diameter, with numerous leafy bracts;
syncarp 3. 5-5. 2 cm long, 3-4 cm in diameter,
when young cream colored, broadly ellipsoid,
bearing 600-960 drupes, these 16-18.5 mm
long, 2-2.5 mm wide, 1.5-2 mm thick, nar-
rowly cylindric, compressed, 5 -angled, upper 14
free, the body 14-15 mm long; pileus 5-6 mm
long, the base 2 mm high, semiorbicular,
crowned by a style 3-4 mm long, prolonged

Page 260: Revision of Pandanus, 14. Malaya and Singapore — St. John

into a lanceolate, entire or less commonly bifid
beak, bony, brownish, shining, flattened, proxi-
mally curved; stigma 1-1.5 mm long, elliptic or
broadly so, creased, brown, papillose, running
almost to the apex; endocarp in lower V3, carti-
laginous, brown, walls 0.1 mm thick, the inner
surface pale, dull; seed 3-4 mm long, ellipsoid,

! truncate; apical mesocarp a slender cavity 9 mm
long; basal mesocarp fibrous up the sides, fleshy
within, 4-5 mm long.

HOLOTYPUS: Malaya, Johore, Sungei Kayu,
in swamp, 17 Oct. 1936, Kiah bin Haji Salleh
32,102 (SING).

SPECIMENS EXAMINED: Malaya, Sfungei]
Kayu, Mawai-Jemaluang Road, in swamp, evi-
dently fruits commonly, 11 Oct. 1936, E. ]. H.
Corner 32,460 (SING), in large part, but not
the loose fruit), (sing).

DISCUSSION : P. brevicornutus is a member of
the section Rykia, as is its closest relative, the
Malayan species P. kedahensis St. John, which
has the syncarp 12.5 cm long, 11 cm in diam-
eter; drupes 4. 2-4.4 cm long, 8-13 mm wide,
narrowly oblanceoloid; style 4-5 mm long,
bifurcate, the lobes mostly divergent; endocarp
j with apical narrow pyramidal projection; seeds

15- 16 mm long; leaves 4.5-4.6 cm wide, and
at midsection with 43 secondary parallel veins
in each half. P. brevicornutus has the syncarp
3. 5-5.2 cm long, 3-4 cm in diameter; drupes

16- 18.5 mm long, 2-2.5 mm wide, narrowly
1 cylindric; style 3-4 mm long, lanceolate, less

commonly unevenly bifid; endocarp apex trun-
cate; seeds 3-4 mm long; leaves 24-29 mm
wide, and at midsection with 18-21 secondary
parallel veins in each half.

The new epithet is taken from the Latin
words brevis, short; cornutus, horned, and is
chosen in reference to the short, hornlike style
of this species.

Pandanus distentus sp. nov. (sect. Rykia)

Fig. 165

DIAGNOSIS HOLOTYPI: Planta cum foliis 1.8-
3 m longis 9 cm latis crassiter coriaceis 1-sul-
; catis 2-plicatis in sectione M-formatis in sec-
tione mediali cum 61 nervis parallelis secunda-
riis prominentibus in quoque medio nervis ter-
1 tialis fortibus in dimidia ultima reticulam for-
mantibus lamina ligulata valde contracta in

apice 29 cm longo subulate caudato trigono eo
10 cm ex apice 1 mm lato basi non conservati,
proxima basem marginibus cum spinis 5-7 mm
longis 11-24 mm separatis deltoideo-subulatis
complanatis subarcuato-adscendentibus pallidis
sed apicibus nigris, midnervo infra cum spinis
teretihus 5-6 mm longis 12-30 mm distantibus
subulatis arcuatis reflexis nigris basibus crassis,
in sectione mediali marginibus cum spinis 4-
5.5 mm longis crasse subulatis arcuatis adscen-
dentibus saepe tota pallidis, midnervo infra cum
spinis 4 mm longis 4.5-6 cm separatis crasse
subulatis arcuatis complanatis valde reflexis, in
sectione diminuenti marginibus cum subulato-
serris 1.2-2 mm longis 1-4 mm separatis arcu-
atis, midnervo infra angusto salienti cum aculeis
simulantibus, in apice caudato marginibus et
midnervo infra cum serrulis 0.2-0.4 mm longis
1-4 mm separatis, inflorescentia foeminea "circa
30 cm longa,” pedunculo 10 mm diametro
trigono folioso-bracteato probabaliter cum syn-
carpio solitario, syncarpio circa 15 cm longo et
10-11 cm diametro ellipsoideo, drupis multis
34-36 mm longis (vel stylo sequi 37-41 mm)
8-12 mm latis 7-12 mm crassis anguste oblan-
ceoloideis subcompressis 5-7-angulosis corpore
30-32 mm longo anguste ellipsoideo vel fili-
form! lateribus subcurvatis vel planis, pileo 9-
12 mm longo basi pyramidali-semiorbiculari
laevi, stylo 3-5 mm longo osseoso rigido lucido
compresso proxime curvato basi late oblonga
parte supera acute bifida cum lobis subulatis
sed eis cum lateribus exterioribus extensis
undulatis, stigmate 3-4 mm longo late ovoideo
bifido ad apices extenso brunneo papilloso,
endocarpio submediali osseoso obscure brunneo
lateribus 1 mm crassis intus lucido in apice cum
projectione central! anguste conico, semine 15
mm longo 5 mm diametro ellipsoideo obtuso,
mesocarpio apicali cavernoso cum fibris et
membranis, mesocarpio basali in lateribus fibro-
so intra carnoso.

diagnosis OF HOLOTYPE: Stature not re-
corded; leaves 1.8-3 m long, 9 cm broad, thick
coriaceous, 1 -ribbed and 1 -furrowed, 2 -pleated,
in section M-shaped, at midsection with 61
parallel secondary veins in each half, these rather
prominent except at base, the tertiary cross
veins strong, transverse, forming oblong meshes,
these at least from the middle outwards giving

lo cm,

FIG. 165. Pandanus distentus St. John, from holotype. a, Phalange, lateral view, X 1; ^ phalange, longi-
tudinal median section, X 1; c-k, drupe and style, apical view, X 1; h stigma, proximal view, X 4; n, o,
style, distal view, X 4; p, leaf base, lower side, X 1; leaf apex, lower side, X 1; r, leaf venation, lower
side, near base, X 4. (e, f, h, i, k, Frequent and most typical styles; c, d , g, j, infrequent forms of style.)

o 10 cm.

Fig. 166. Pandanus inundatus St. John, from holotype. a, Infructescence, habit, X 1; ^ drupe, lateral view,
X h e, drupe, longitudinal median section, X 1; ^ drupe, lateral view, X 4; e, drupe, longitudinal median
section, X 4; f, drupe and style, apical view, X 4; g, drupe apex and style, apical view, X 10; h, style and
stigma, proximal view, X 10; i, leaf base, lower side, X 1; j, leaf middle, lower side, X 1; k, leaf apex, lower
side, X 1.

PACIFIC SCIENCE, Vol. XVII, January 1963

both leaf surfaces a reticulate appearance, the
blade ligulate, rather sharply contracted to a 29
cm subulate, caudate, trigonous apex, this at 10
cm down from the tip 1 mm wide, the base not
preserved, near the base the margins with thorns
5-7 mm long, 11-24 mm apart, deltoid, subu-
late, flat, slightly arcuate ascending, pale except
for tiny blackish tips; the midrib below with
heavy based, terete spines 5-6 mm long, 12-30
mm apart, subulate, arcuate, reflexed, black; at
lower third the margins with spines 4-5.5 mm
long, 13—29 mm apart, heavy subulate, arcuate,
ascending, pale throughout or with minute
blackish tips; the midrib with spines 4 mm
long, 4.5-6 cm apart, flat, heavy subulate, arcu-
ate, strongly reflexed; on the contracting area
near the tip the margins with prickles 1.2-2
mm long, 1-4 mm apart, subulate-tipped ser-
rae, arcuate; the midrib below narrow, sharp,
salient, bearing similar prickles; on the caudate
apex the margins and midrib below with ser-
rulations 0.2— 0.4 mm long, 1—4 mm apart; pis-
tillate inflorescence "about 30 cm. long,” pe-
duncle about 10 mm in diameter, 3-sided, leafy
bracted, apparently bearing a single syncarp,
this apparently about 15 cm long and 10 or 11
cm in diameter, ellipsoid (an estimate based
upon a part of the core, two slabs of drupes,
and many separate ones) ; drupes numerous 34-
36 mm long (or following the curve of the
style 37-41 mm), 8-12 mm wide, 7-12 mm
thick, narrowly oblanceoloid, a little com-
pressed, 5-7-angled, the body 30-32 mm long,
narrowly ellipsoid or fusiform, the sides gently
curved or plane; pileus 9-12 mm long, the base
pyramidal-semiorbicular, smooth; style 3-5 mm
long, bony, rigid, shining, flattened, proximally
curved, the base broad oblong, the upper part
sharply bifid with subulate tips but the outer
sides with broad, more or less undulate, ex-
panded margins; stigma 3-4 mm long, broad
ovoid, bifid, running to the apices, brown, papil-
lose; endocarp submedian, bony, dark brown,
the lateral walls 1 mm thick, the inner surface
shining, with a narrow conical projection run-
ning upward into the beak; seed 15 mm long,
5 mm in diameter, ellipsoid, obtuse; apical
mesocarp cavernous, but with fibers and mem-
branes; basal mesocarp fibrous up the margins,
fleshy within.

HOLOTYPUS: Malaya, Pulau Penang, Telok !
Aling, native, common, 23 Oct. 1951, /. Sinclair
39,382 (sing).

DISCUSSION: P. distentus is a member of the
section Rykia, as is its closest relative, the Ma-
layan species P. longicaudatus Holttum & St.
John, a species with the drupes 41-44 mm
long, the lateral ones with styles 4-8 mm long, ,
bifurcate, the forks subulate; endocarp walls
1.5-2 mm thick; leaves 4.7-6 cm wide, the
caudate apex 15-22 cm long, secondary nerves .
48 in each half, and near the base the margins ,
with thorns 3-5 mm long, 6-14 mm apart. P.
distentus has the drupes 37-41 mm long, the
styles sharply bifid, but the lobes with undulate,
expanded outer margins; endocarp walls 1 mm i
thick; leaves 9 cm wide, the caudate apex 29 1
cm long, the secondary nerves 61 in each half,
and near the base the margins with thorns 5-7 i;
mm long, 1 1-24 mm apart.

The new epithet is the Latin participle dis-
tentus, distended, and is given in reference to ;
the lateral extensions of the style forks.

Pandanus inundatus sp. nov. (sect. Rykia )

Fig. 166

diagnosis holotypi: Frutex, caule in apice
8 mm diametro, foliis 37-42 cm longis 30-32
mm latis in medio sed proxima basem 20 mm
latis chartaceis et sulcatis minime 2-plicatis ligu-
latis ad apicem diminuentibus in apice trigono
subulato 4 cm longo 1-1.5 mm lato, in sectione
mediali cum 19-20 nervis parallelis secundariis
conspicuis in quoque dimidio, nervis tertialis :
nullis, basi amplexicauli et inermi sed ex 3-3.5
cm marginibus cum aculeis 2. 5-3. 5 mm longis i
4-8 mm separatis subulatis arcuatis brunneis,
midnervo infra ex 10 cm cum aculeis simulanti- •
bus 3 mm longis reflexis remotis paucis, in sec- I
done mediali marginibus cum aculeis 0.8-1 mm i
longis 7-20 mm separatis subulatis adpresse ad-
scendentibus, midnervo infra cum aculeo unico i
vel pluribus remotis 1.5 mm longis compressis i
arcuatis adscendentibus, in apice subulato mar-
ginibus et midnervo infra cum aculeis 0.3-0. 5
mm longis 0.5- 1.5 mm separatis subulatis ad- f
scendentibus, infructescentia terminali cum cap- |l
ite unico, pedunculo 12 cm longo 4 mm diame-
tro trigono folioso-bracteato, syncarpio 6 cm
longo 4.5 cm diametro ellipsoideo sed minime

Page 264: Revision of Pmdanus, 14. Malaya and Singapore — St. John

3-planato cum circa 832 drupis eis 17—19 mm
longis 2-2.5 mm latis 1.5-2 mm crassis viridi-
bus corpore 14-15 mm longo anguste cylindrico
compresso 5-6-anguloso, pileo 6-8 mm longo
basi 2-2.5 mm alta ellipsoidea laevi, stylo 5-6
mm longo plerumque proximo frequente subre-
flexo, stigmate 2-3.5 mm longo anguste ellip-
tico brunneo papillose ex apice distanto, endo-
carpio in parte lA infera cartilagineo pallide
brunneo lateribus 0.1 mm crasso intra exlucido,
semine 2.7 mm longo ellipsoideo, mesocarpio
apicali cavernoso 8 mm longo, mesocarpio basali
in lateribus fibroso intra carnoso.

DESCRIPTION OF ALL SPECIMENS EXAMINED:
Shrub up to 1.7 m tall, 8-15 mm in diameter,
often decumbent; bark dark mahogany-colored,
shining; internodes 7-11 mm long; prop roots
several and as much as 40 cm long, 7-8 mm in
diameter, brown, with loose spirals of adventi-
tious rootlets 2-5 mm long, divergent, prickle-
like; leaves 37-65 cm long, 25-34 mm wide at
the middle, 20-24 mm wide near the base,
chartaceous, above glossy dark green, below yel-
lowish green, at base channeled above the mid-
rib, slightly 2 -pleated, ligulate, the apex nar-
rowed to a 4-5 cm trigonous subulate tip 1-1.5
mm wide, at midsection with 18-22 secondary
parallel conspicuous veins in each half, but no
clearly visible tertiary veins, the base amplexi-
caul, unarmed, but beginning at 3-3.5 cm the
margins with prickles 2. 5-3. 5 mm long, 4-8 mm
apart, subulate, bent upwards, brownish; the
midrib below beginning at 10 cm with a few
remote similar prickles 3 mm long, reflexed; at
midsection the margins with prickles 0.8-1 mm
long, 7-20 mm apart, subulate, appressed as-
cending; the midrib below with one or several
remote prickles 1.5 mm long, compressed, heavy
based, arcuate ascending; on the subulate tip the
margins and midrib beneath with prickles 0.3-
0.5 mm long, 0.5-1. 5 mm apart, subulate, as-
cending; infructescence terminal, with a single
head, or rarely two heads; peduncle 10-17 cm
long, 4-6 mm in diameter, trigonous, leafy
bracted, sometimes pendent; syncarp 4-6 cm
long, 3-4.5 cm in diameter, ellipsoid but slightly
3-sided, bearing 832-1,040 drupes, these 17-19
mm long, 2-2.5 mm wide, 1.5-2 mm thick,
green, the body 14-15 mm long, narrowly cylin-
dric, compressed, 5-6-angled; pileus 6-8 mm

long, the base 2-2.5 mm high, ellipsoid, smooth;
style 5-6 mm long, mostly proximal in attach-
ment, often somewhat reflexed; stigma 2-3.5
mm long, narrowly elliptic, brown, papillose,
remote from the apex; endocarp in lower
cartilaginous, pale brown, the walls 0.1 mm
thick, the inner surface dull; seed 2.7 mm
long, ellipsoid; apical mesocarp a cavern 8 mm
long; basal mesocarp fibrous up the sides, fleshy
within.

HOLOTYPUS: Malaya, Johore, Sungei Sedili,
30 July 1939, Kiah bin Haji Salleh 63,926
(SING).

SPECIMENS EXAMINED: Malaya, Johore, 5Vl
miles, Kota Tinggi-Mawai Road, in very swampy
places in backwaters of streams, forming more
or less pure stands, 13 April 1935, E. J. H.
Corner 29,231 (BO, sing); Johore Sfungei]
Berassau, Mawai-Jemaluang Road, frequent in
swampy forest, 6 Feb. 1935, E. J. H. Corner
28,744 (sing); ditto, flopping, gregarious in wet
hollows in swampy forest, 7 Feb. 1935, Corner
s. n. (SING); ditto, gregarious in standing water,
low elevation, 28 April 1935, Corner 29,361
(BO, sing); ditto, in standing swamp by the
stream, 5 Jan. 1936, Corner 29,991 (sing).

discussion: P. inundatus is a member of the
section Rykia and in that section is its closest
relative, P. Scortechinii Martelli, a species with
drupes 12-13 mm long; style terminal, 4-5 mm
long; stigma linear; endocarp slightly subme-
dian; seed 6 mm long; leaves 25-27 cm long,
near the base 14 mm wide, at midsection the
secondary nerves 15-16 in each half, tertiary
cross veins visible in the outer half, at midsec-
tion the margins with weak prickles 0.2-0. 3
mm long, flat appressed, close or remote or none.
P. inundatus has the drupes 17-19 mm long;
style of lateral drupes markedly lateral, 5-6 mm
long; stigma narrowly elliptic; endocarp in lower
Vy seed 2.7 mm long; leaves 37-65 cm long,
near the base 20 mm wide, at midsection the
secondary nerves 18-22 in each half, no clearly
visible cross veins; at midsection the margin
with prickles 0.8-1 mm long, 7-20 mm apart,
subulate, appressed ascending.

This was probably the species illustrated by
Martelli (Webbia 4(2): t. 32, f. l-3a, 1914)
as his P. johorensis Martelli. Martelli did not
have the habit of indicating the specimen from

PACIFIC SCIENCE, Vol. XVII, January 1963

FIG. 167. Pandanus kedahensis St. John, from holotype. a, Syncarp, Y Vl\ b, drupe, lateral view, X 1; c,
drupe, longitudinal median section, X 1; d, lateral drupe and style, apical view, X 1; ^ apical drupe and style,
apical view, X 1 \ f, g, stigma, proximal view, X 4; h, leaf base, lower side, X 1; i, leaf middle, lower side,
X 1 ; j, leaf apex, lower side, X i.

Page 266: Revision of Pandanus, 14. Malaya and Singapore — St. John

which he made his illustrations. Evidence is now
accumulating to show that he did not consist-
ently illustrate the types, but rather that he used
the best specimen at hand when making the
drawing. Having seen and illustrated the type
number of P. joborensis, it is evident to the
writer that Martelli’s figures on his plate 32,
published 10 years after he published the spe-
cies, were drawn from another specimen, not
the holotype. This other specimen seems to be
like those here described as the new species
P. inundatus.

The new epithet is the Latin participle inun-
datus, flooded, given in reference to the habitat
of the species.

Pandanus kedahensis sp. nov. (sect. Rykia )
Fig. 167

DIAGNOSIS HOLOTYPI: Planta cum "foliis
grandibus,” pedunculo 22 cm vel plus longo 12
mm diametro trigono folioso-bracteato, bracteis
47-70 cm longis 4. 5-4.6 cm latis coriaceis
1-costatis 2-plicatis in sectione mediali cum 43
nervis parallelis secundariis in quoque latere eis
supra evidentibus et infra prominentibus, nervis
tertialis prominentibus et reticulum cum sec-
tionibus oblongis formantibus lamina ligulata ad
basem spathacea sed ex 14 cm marginibus et
midnervo infra cum aculeis 0.5-1 mm longis
1-4 mm separatis subulatis adscendentibus api-
cibus brunneis, in sectione mediali marginibus
cum aculeis 2-3 mm longis 2-10 mm separatis
arcuatis subulatis adscendentibus adpressis semi-
sombratis in margine undulato, midnervo infra
angusto salienti cum aculeis 1 mm longis 1-4
cm separatis arcuatis subulatis adscendentibus,
in apice caudato marginibus et midnervo infra
cum subulato-serrulis 0.5-0.8 mm longis 1-2
mm separatis, syncarpio 12.5 cm longo 11 cm
diametro late ovoideo cum circa 544 drupis eis
4.2-4.4 cm longis 8-13 mm latis 7-12 mm
crassis anguste oblanceoloideis 5-6-angulatis
parte 2A supera libera corpore 3.8-4 cm longo,
pileo 13-14 mm longo pyramidali-ovoideo 5-6-
angulato laevi plerumque ex apice styli planato
proxime inclinato truncate, stylo 4-5 mm longo
oblongo osseoso brunneo lucido bifurcato lobis
plerumque divergentibus, stigmate 2-3 mm
longo proximo late ovato bifido brunneo papil-
loso paene in apicibus extento, endocarpio in

parte 2A infera obscure brunneo osseoso cum
projectione angusto pyramidali apical i lateribus
2-3 mm crassis intra rugosis lucidisque, semine
15-16 mm longo 2 mm diametro, mesocarpio
apicali cavernoso et cum membranis albis, meso-
carpio basali fibroso et carnoso.

DIAGNOSIS OF HOLOTYPE: Stature not re-
corded; "large leaved”; peduncle more than 22
cm long, 12 mm in diameter, 3 -sided, leafy
bracted, these bracteal leaves 47-70 cm long,
4. 5-4.6 cm wide, coriaceous, 1 -ribbed, 2 -pleated,
at midsection with 43 parallel secondary nerves
in each half, these visible above, prominent be-
low, the tertiary cross veins conspicuous, at right
angles, forming meshes mostly oblong, the blade
ligulate, towards the base spathaceous enlarged;
at 14 cm above the base the margins and midrib
below with prickles 0.5-1 mm long, 1-4 mm
apart, subulate, ascending, brown-tipped; at mid-
section the margins undulate and with partly
countersunk prickles 2-3 mm long, 2-10 mm
apart, arcuate subulate, flat appressed, ascend-
ing, brown-tipped; midrib below narrow, salient,
with prickles 1 mm long, 1-4 cm apart, arcuate
subulate, ascending; the caudate apex having the
margins and midrib below with subulate tipped
serrulations 0. 5-0.8 mm long, 1-2 mm apart;
syncarp 12.5 cm long, 1 1 cm in diameter, broadly
ovoid, bearing about 544 drupes, these 4.2-4.4
cm long, 8-13 mm wide, 7-12 mm thick, nar-
rowly oblanceoloid, 5-6-angled, upper 2A free,
the body 3.8-4 cm long; pileus 13-14 mm long,
pyramidal-ovoid, 5-6-angled, smooth, mostly
truncate by the flattened upper side of the
sharply proximally bent style, this 4-5 mm long,
oblong, bony, brown, shining, bifurcate, the
lobes mostly diverging; stigma 2-3 mm long,
proximal, broad ovate, bifid, brown, papillose,
extending almost to the apices; endocarp in
lower 2A, dark brown, bony, with a narrow
pyramidal apical extension, the lateral walls 2-3
mm thick, in inner surface rugose, shining; seed
15-16 mm long, 2 mm in diameter; apical meso-
carp one large cavern with white membranes;
basal mesocarp fibrous and fleshy.

HOLOTYPUS: Malaya, Kedah, Kedah Peak, in
low forest, not far from summit, 3,500 ft. alt.,
1 April 1925, R. E. Holttum 14,858 (sing).

DISCUSSION: P. kedahensis is a member of
the section Rykia, as is its closest relative, the

Fig. 168. Pandanus serratus St. John, from holotype. a, Syncarp profile, X Vl\ b, syncarp, outline of trans-
verse section, X Vl\ c, drupe, distal view, X 1; drupe, longitudinal median section, X 1> e—k, lateral drupes
and styles, apical view, X 1; h apical drupe, apical view, X 1; m, carpel apex and style, distal view, X 4; n,
carpel apex, proximal view, X 4; o, leaf base, lower side, X 1; P> leaf middle, lower side, X l> lea^ aPex>
lower side, X 1; r, venation near leaf base, lower side, X 4.

Page 268: Revision of Pmdanus, 14. Malaya and Singapore— St. John

Fig. 169. Pandanus albibracteatus St. John, from holotype and paratype. a, Clump of pistillate holotypic
trees at top of beach, with figure of Cheang Kok Choy; b, branch with syncarp; c, foliage of paratype, St. John
26,370 ; d, staminate paratype, St. John 26,370, with figure of Cheang Kok Choy.

Malayan species P. longicaudatus Holttum &
St. John, a species with the peduncle 22 mm
in diameter; syncarp 16=17 cm long, with
about 900 drupes; pileus pyramidal-hemispheric;
stigma 3—3.5 mm long; endocarp walls 1.5-2
mm thick, the inner surface smooth; and the
seed 6 mm in diameter. P, kedabensis has the
peduncle 12 mm in diameter; syncarp 12.5 cm
long, with about 544 drupes; pileus pyramidal-
ovoid; stigma 2=3 mm long; endocarp walls 2=3
mm thick, the inner surface rugose; and the seed
2 mm in diameter.

The new epithet is a Latinization of the name
of the type locality.

Pandanus serratus sp. nov. (sect. Rykia )

Fig. 168

NOM. VERN. : "mengkuang ayer.”

DIAGNOSIS HOLOTYPI: Licet aquatica in aqua
vadosa, fo.li.is 1.5 m longis in media 8.5 cm latis
ad terminos ambos diminuentibus crassiter cori-
aceis supra viridibus infra minirne pallidiori-
bus 1-sulcatis 2-plicatis in sectione depresse
AA-formatis in sectione mediali cum 75 nervis
parallelis secundariis in quaque dimidia in ap ice
longe subulato diminuentibus basi amplexicauli
inermi sed ex 20 cm marginibus cum aculeis
1-2 mm longis 5=10 mm distantibus valde

PACIFIC SCIENCE, VoL XVII, January 1963

midnervo angusto salienti infra cum serris 2-2.5
mm longis 10-15 mm separatis adpresse adscen-
dente subulatis, proxima apicem marginibus
cum serrulis 0.5-1 mm longis 4-8 mm separatis,
midnervo infra cum serrulis minutis remotis
paucis; inflorescentia foeminea erecta cum 1
syncarpio, pedunculo 14 cm longo 1 cm diame-
tro trigono bracteato, syncarpio 20.5 cm longo
et proxima apicem 11 cm diametro, in media

9.5 cm diametro et basi subcylindrica, syncarpio
3-laterato apice obtuso cum circa 4,300 drupis
eis 28-37 mm longis eis medialis basalisque 5-8
mm latis 4-6 mm crassis sed illis superis 9-12
mm latis 7-9 mm crassis oblanceoloideis, cor-
pore 23-32 mm longo 5-6-angulato, pileo 4-8
mm longo basi semiorbiculari-pyramidali laevi,
stylo 3-5 mm longo osseoso brunneo lucido eis
lateralibus et inferioribus 2.5-4 mm latis ob-
longis in basi latioribus gradatim proxime cur-
vatis apice bifurcato lobis 1-3 mm longis ple-
r unique divergentibus eis terminalibus cum stylo
corniformi integro, stigmatibus 3-4 mm longis
lanceolatis in lobis sed infra connatis brunneis
papillosis proximis olim ad apices continuenti-
bus, endocarpio in parte infera osseoso ob-
scure mahogani-colorato lateribus 1 mm crassis
pagina, interiori lucida apice cum extensioni
subulato centrali, semine 1 cm longo anguste
obovoideo, mesocarpio apicali cavernoso 17 mm
longo fibris marginalibus multis centralibus
paucis et cum medulla, mesocarpio basali fibroso
et carnoso.

DIAGNOSIS OF HOLOTYPE: Apparently aqua-
tic, rooting in shallow fresh water; leaves about

1.5 m long, at the middle 8.5 cm wide, tapering
towards both ends, thick coriaceous, green above,
slightly paler below, 1 -ribbed, 2 -pleated, in sec-
tion low M-shaped, at midsection with 75 sec-
ondary parallel veins in each half, tapering
towards the tip into a long subulate apex, but
the actual tip not preserved, the base amplex-
icaul and unarmed, but the margins beginning
at 20 cm up with prickles 1-2 mm long, 5-10
mm apart, stout arcuate subulate, pale, ascend-
ing; the nearby midrib below unarmed; at mid-
section the margins with spines 6.5-7 mm
long, 7-16 mm apart, heavy, arcuate subulate,
ascending, the base much thickened, 4-5 mm
wide, the tips reddish; the midrib below with
appressed ascending subulate tipped serrae 2-2.5

arcuato-subulatis pallidis adscendentibus, mid-
nervo infra inermi, in sectione mediali margini-
bus cum spinis 6.5-7 mm longis 7-16 mm
separatis fortiter arcuato-subulatis adscendenti-
bus basi incrassata 4-5 mm lata apicibus rubris,

Fig. 170. a, Staminate inflorescence and leaf of
Pandanus albibracteatus , paratypic specimen, St. John
26,370; b, Pandanus ambiglaucus St. John, holotype,
branch, foliage, and syncarp; c, Pandanus inclinatus
St. John, holotype, branch, foliage, and syncarp.

Page 270: Revision of Pandanus, 14. Malaya and Singapore — St. John

mm long, 10-15 mm apart, on the narrow 1 syncarp; peduncle 14 cm long, 1 cm in diam-

salient midrib; near the apex the margins with eter, 3-sided, bracteate; syncarp 20.5 cm long,

serrulations 0.5-1 mm long, 4-8 mm apart; the 11 cm in diameter near the tip, 9.5 cm at the

midrib below with a few, tiny, very remote ser- middle, and near the base subcylindric, 3 -sided,

rulations; pistillate inflorescence erect, bearing the apex obtuse, bearing about 4,300 drupes,

Fig. 171. Pandanus incrassatus St. John, holotype, St. John 26,374. a, Habit, foliage, and syncarp, with
figure (right) of Paul Weissich; h, trunk; c, trunk and prop roots; d, branch, foliage, and syncarp.

PACIFIC SCIENCE, Vol. XVII, January 1963

these 28-37 mm long, the middle and lower
ones 5-8 mm wide, 4-6 mm thick; the upper
and apical ones 9—12 mm wide, 7—9 mm thick,
oblanceoloid; the body 23-32 mm long, 5-6-
angled; pileus 4-8 mm long, the base semi-
orbicular-pyramidal, smooth; style 3-5 mm long,
bony, brown, shining, of the lateral and lower
ones 2.5-4 mm wide, oblong, broadening at
base, gently proximally curved and at apex
bifurcate, the lobes 1-3 mm long, mostly diver-
gent, but the terminal drupes with simple, horn-
like style; stigmas 3-4 mm long, lanceolate on
the lobes but united below, brown, papillose,
proximal, running almost to the point of the
style; endocarp centering in lower V3, bony,
dark mahogany-colored, the walls 1 mm thick,
the inner surface shining, the apex with a cen-
tral, subulate prolongation; seed 1 cm long, nar-
rowly obovoid; apical mesocarp a cavern 17 mm
long, with fibers up the margin and a few within,
and with traces of a medullary pith; basal meso-
carp fibrous and fleshy.

HOLOTYPUS: Malaya, Pahang, Sfungei] Bera,

w. Tasek Bera, low elevation, 15 Oct. 1930,

M. R. Henderson 24,137 (sing).

DISCUSSION: P. serr at us is a member of the
section Rykia , as is its closest relative, P. hetero- r
stigma (Martelli) Martelli, a Sumatran species
that has a raceme with 7-10 heads; syncarps
12-15 cm long, 9-19 cm in diameter; drupes 25
mm long, 7-8 mm wide; style spiniform, subu-
late, or narrowly oblong and bifurcate; and the
leaves 2 m or more in length, 10 cm wide. .

P. serratus has the syncarp solitary, 20.5 cm long,

11 cm in diameter; drupes 28-37 mm long,
5-12 mm wide; style normally oblong, bifurcate
but the apical ones stout, hornlike; and the
leaves about 1.5 m long, 8.5 cm wide.

As is the case with many members of the
section Microstigma, this species has the drupes
near the apex of the syncarp markedly longer
and wider than those of the lower parts of the
syncarp.

The new epithet is the Latin adjective ser-
ratus, saw-toothed, given in reference to the leaf
margins.

Species Structure of the Gobiid Fish Gillichthys mirabilis
from Coastal Sloughs of the Eastern Pacific1

George W. Barlow2

A RECENT ISSUE of Systematic Zoology ( I960,
vol. 9, nos. 3, 4) was devoted entirely to a
symposium entitled "The Biogeography of Baja
California and Adjacent Seas.” One recurring
theme was the affinity between forms occurring
on the Pacific Coast in the Californian province
(Hubbs, I960: 134), and those in the northern
part of the Gulf of California. Between these
areas, in the main part of the Gulf, the fauna
was said to differ, being primarily Panamic in
origin (Garth, I960; Hubbs, I960; Walker,
I960).

The outer-coast species found in the upper
Gulf of California are Pleistocene relicts, now
trapped in a cul-de-sac of warm, tropical water
(Hubbs, 1948: 463; I960: 137). According to
Walker (I960: 130), most of the northern
species found there differ from their cognates
on the Pacific Coast. As the symposium re-
vealed, however, there has been no extensive
study comparing populations of one species on
the Pacific Coast with those of the same species
in the Gulf of California.

The immediate objective of this investiga-
tion, although it was initiated before the sym-
posium was conceived, was to establish the
nature of the variation between populations of
Gillichthys mirabilis Cooper. The goal was to
ascertain the degree of differentiation of the
form from the upper Gulf in relation to the
over-all structure of the species. A brief study
(Barlow, 1961^) of G. seta (Ginsburg) has
served as comparative material to gauge the
differences observed in G. mirabilis. G. seta is
a close but distinct species restricted to an ex-

1 A revised portion of the dissertation submitted in
partial fulfillment of the requirements for the degree
of Doctor of Philosophy at the University of Cali-
fornia, Los Angeles, California. Manuscript received
September 21, 1961.

2 Department of Zoology, University of Illinois,
Urbana, Illinois.

treme rocky habitat found only in the upper
Gulf.

G. mirabilis has been cited as one example of
a species from the Pacific Coast which is also
found in the upper reaches of the Gulf of Cali-
fornia (Fig. 1). It occurs in the central region
of the Gulf as well. There is a break in its
distribution around Cabo San Lucas, however,
separating the species into a Gulf group and a
Pacific Coast group (details in Barlow, 1961^).

At the inception of the present investigation
G. mirabilis from the northern part of the Gulf
of California was considered by some ichthyolo-
gists to be a distinct species, G. detrusus Gilbert
and Scofield. Differences of opinion existed
(Barlow, 19 6D), and the issue was clouded by
comparisons with a third species, the then un-
described Gillichthys seta. As will be seen, the
evidence supports the conclusion of Starks and
Morris (1907: 227) that detrusus should be
regarded as a synonym of mirabilis.

In 1930 G. mirabilis from the San Diego
area in southern California was introduced into
the Salton Sea. The fish has become well estab-
lished in that odd saline lake in the Colorado
Desert of California. This has proved to be a
fortunate experiment.

ACKNOWLEDGMENTS

With pleasure I acknowledge the timely as-
sistance of the following people in gathering
material for the study: H. P. Arai, W. J. Bald-
win, Raymond Cannon, L. H. Carpelan, G. L.
Durall, W. I. Follett, R. H. Linsey, J. B. Makern-
son, F. W. Munz, K. S. Norris, P. E, Pickens,
J. H. Prescott, J. C. Quast, E. S. Reese, R. H.
Rosenblatt, Octavio Salazar, George Sims, R. R.
Whitney, and John Wintersteen. Charles Baxter
kindly prepared the chart of the study region,
and Gerta M. Barlow assisted in the preparation
of the manuscript and many of the illustrations.

The preliminary manuscript was read by George
A. Bartholomew, Henry W. Thompson, and
John N. Belkin, all of the University of Cali-
fornia, Los Angeles, and I thank them for their
helpful suggestions. The research and writing
were done under the supervision of Boyd W.
Walker, and I am grateful to him for his guid-
ance and aid.

ECOLOGY AND NATURAL HISTORY

The typical habitat of mirabilis is the inter-
tidal coastal slough with its extensive mud flats.
Less often it is a swampy lagoon, or slough, at
the mouth of a dry river which most of the
time is closed off from the ocean by a sand bar.

Salinity

The water in these habitats is of moderate to
high salinity, hypersaline conditions being com-
mon. The maximum salinity where mirabilis
has been recorded was 82.5 %o at "El Marino,”
north of San Felipe in the upper Gulf of Cali-
fornia ( Lars Carpelan, personal communica-
tion). The fish usually are absent where the
water is fresh or only slightly brackish. Those
kept in fresh-water aquaria were listless and
died within 1-2 weeks, but this may have been
due to the quality of the tap water. The activa-
tion time of the spermatozoa is quickest in
water having a salinity of 35 %o (Weisel, 1948:
47).

Temperature

Aside from illumination, temperature is the
one parameter of the environment to which
it is related that changes more or less regularly
with the latitude of the habitat. Local condi-
tions, of course, may distort this temperature
gradient. Furthermore, the Gulf of California
has a climate different from that of the Pacific
Coast. For these reasons, and in order to point
out the extreme situation found at the Salton
Sea, the meager temperature data available have
been assembled and are presented in Table 1.

Air temperatures have been used instead of
sea surface temperatures. In the backwaters and
ponds where mirabilis occurs, the surface water
temperatures tend to be correlated with air tem-
peratures (Carpelan, 1957: 376; Barlow, 1958:

PACIFIC SCIENCE, Vol. XVII, January 1963

582 ) , even when moderate tidal exchange takes
place (personal observations).

Throughout the year on the Pacific Coast,
air temperatures increase from north to south.
The increase is more pronounced in summer
than in winter. For example, San Diego is
warmer than San Francisco by about 4-5 C dur-
ing the summer, but is only 2-3 C warmer in
the winter.

The protected Alviso ponds of southern San
Francisco Bay, with their increased insolation,
exhibit a special climatological situation (Car-
pelan, 1957: 37 6) that removes them from the
normal gradient of temperature change asso-
ciated with latitude. Air temperatures at the
ponds during 1 year averaged almost 7 degrees
higher than the long-range monthly means for
San Francisco shown in Table 1. The climate
at Alviso ponds evidently is warmer than at
many truly coastal sloughs much further south.

In the Gulf of California region air tempera-
tures increase from north to south during much
of the year. But during the late, hot, summer
season the situation is reversed and the mean
air temperatures decrease, though only slightly,
from north to south. However, the mean sea
surface temperatures for August (Roden and
Groves, 1959: 14) show no latitudinal trend
whatsoever. Chiefly because of the reversal in
the gradient of the air temperatures, the annual
range of means is low in the south but is pro-
gressively greater to the north in the Gulf.

In the tidal sloughs of the Gulf of California
where mirabilis normally is found, the summer
increase from south to north in the range of
water temperatures probably is augmented by
the nature of the tidal change. The tidal range
increases dramatically from south to north up
the Gulf, reaching a maximum of 10 m at the
head of the Gulf (Roden and Groves, 1959:
28). The sills of the sloughs usually are high.
Thus as one proceeds north in the Gulf, the
sloughs daily have a progressively longer pond
phase. This means that the buffering effect of
the tidal exchange will be less, and as a result
the water temperatures in the sloughs will reg-
ister still greater fluctuations.

The winter mean air temperatures at San
Diego on the Pacific Coast, the central and
northern Gulf of California areas, and the Salton

GilUchthys mirabilis — Barlow

TABLE 1

Monthly Mean Air Temperatures

PACIFIC

SALTON

GULF

OF CALIFORNIA3

COAST1

SEA2

West Coast

East Coast

MONTH

San

Francisco

37° 48'N

San
Diego
32° 34'N

33° 18'N

Mulege
26° 53'N

La Paz
24° 10'N

Guaymas
27° 55'N

Topolo-
bampo
25° 36'N

Mazatlan
23° ll'N

January

12.5

12.0

14.0

18.2

17.7

18.6

19.3

February

11.6

13.5

13.0

16.0

19.2

18.9

19.7

19-4

March

12.9

14.4

17.5

17.8

21.3

20.7

20.1

19.7

April «

13.4

15.2

20.5

20.1

23.2

22.8

22.2

21.3

May

14.1

16.9

24.5

22.9

25.5

25.0

23.8

June

14.9

17.8

28.0

27.2

29.1

26.4

July

14.8

19.9

30.5

30.0

29.8

27.5

August

15.1

20.3

31.5

30.4

30.3

30.5

29.7

27.6

September

16.7

19.4

29.5

29.0

30.2

29.7

27.5

October

15.7

17.8

24.5

24.6

27.0

27.3

27.8

26.7

November

14.0

15.9

18.5

19.5

23.2

22.6

24.0

23.5

December

10.4

13.6

13.0

14.9

20.0

18.6

19.7

20.6

Range

7.4

7.8

19.0

16.5

12.1

12.8

11.2

8.3

1 Hubbs, 1948.

2 Carpelan, 1958.

3 Roden, 1958.

Sea are surprisingly similar. The summer tem-
peratures, in marked contrast, diverge greatly.
San Diego and San Francisco remain moderate,
but the mean air temperatures exceed 30 C in
the Gulf and Salton Sea areas.

The climate of the Salton Sea resembles that
of the northern part of the Gulf of California.
Both lie in the same climatological region, but
the Salton Sea is a relatively small and shallow
body of water surrounded by desert (Carpelan,
1958). Consequently, its climate is even more
extreme than that of the upper Gulf.

Dispersal

Pelagic larvae constitute the most likely ave-
nues of genetic exchange between disjunct pop-
ulations of mirabilis. The adults are closely
restricted to their specialized and discontinuous
habitats. At least some of the larvae must be
swept out to sea where they could be distrib-
uted to other bays by the oceanic currents.

There is some evidence that the larvae of
this genus are not adapted to a pelagic life as
prolonged as those of the related genera, Quie-
tula, Clevelandia, and llypnus (but not Eucyclo-

gobius ). This conclusion is based on differences
in postlarval pigmentation and behavior. Pelagic
larvae and nektonic postlarvae characteristically
are translucent with scattered melanophores,
while the completely pigmented condition is
found first in the benthonic stage. Postlarvae
of GilUchthys mirabilis as small as 8-12 mm in
length often are relatively heavily pigmented.
At a comparable length, postlarvae of the other
genera mentioned above are much more trans-
lucent.

Postlarvae of mirabilis in a slough in the
northern Gulf of California, and at the Salton
Sea, were seen to drift or swim into the current
in slowly moving water. The postlarvae de-
scended to the bottom and held station when
the current strength increased. Postlarvae of
Quietula guaymasiae Jenkins and Evermann
were observed to form small groups that swam
vigorously in a strong tidal current in the Gulf,
darting from side to side at the surface in a
manner reminiscent of young atherines; they
seemed well adapted for life in open water.
The larvae of GilUchthys mnabilis evidently are
not dispersed as readily as are those of the re-
lated genera just mentioned.

PACIFIC SCIENCE, Vol. XVII, January 1963

Reproduction

Spawning apparently commences in January
throughout the range of mirabilis, but the
length of the season is uncertain. In southern
California spawning takes place throughout the

period from January to July ( Weisel, 1947:
82). My observations indicate that spawning
also starts in January in the Alviso ponds; fe-
males hatched there in January and February,
however, are mature and full of ripe eggs in
August and September of the same year. It is

Fig. 1. Chart of the study region; latitude N, longitude W.

Gillichthys mirabilis— B arlow

not known whether or not they spawn this late
in the season. In the Salton Sea, the first spawn-
ing occurs in January and the last in May or
June. Postlarvae of mirabilis have been collected
in early February in the northern and central
Gulf of California. There are no data available
on the length of the spawning season in Gulf
populations.

Depending on the size of the female, 4,000-

9.000 eggs are laid at each spawning (Weisel,
1947: 80). Weisel believed that each female
spawns once, rarely twice, in a season. Exami-
nation of ovaries of Salton Sea fish leads me
to believe that each female mirabilis oviposits
2 or 3 times each season, with an interval of
40-50 days between spawnings. MacGinitie
(1939: 500) reported that the related blind
goby, Typhlogobius calif orniensis Steindachner,
spawns twice during one season, with an inter-
val of about 2 months. Females of a more dis-
tantly related goby, the mapo, Bathygobius sopo-
rator (C and V.), can repeat a spawning in 7-16
days (Tavolga, 1954: 432). If the above esti-
mates are correct, each female Gillichthys mira-
bilis has a potential production of about 8, GOO-

27. 000 eggs in one season.

The young of mirabilis reach maturity by the
end of their first year. Rate of growth was in-
vestigated only at the Salton Sea. It is clear
from these data and from the changes in size
composition of collections from other locations
taken at different times of the year, that most
fish attain adulthood and a standard length of
100-140 mm by the end of their first year.
Nothing is known of their life expectancy.

MATERIALS AND METHODS

In the preliminary analysis of mirabilis and
seta, certain counts and measurements were se-
lected as being especially promising (Barlow,
1961a). While investigating these, other useful
characters also came to light.

All counts and measurements, except one,
were carried out in accordance with the methods
of Hubbs and Lagler ( 1949) . The exception,
the height of the anal fin, is defined here as the
distance from the base of the last element of
the anal fin to the tip of the second to last ray,
when the fin is erect.

The study was based almost entirely on speci-
mens deposited in the Fish Collections of the
University of California, Los Angeles. About

4,000 specimens of mirabilis were examined.
The fish from San Francisco Bay ( Alviso ponds )
were obtained from Sim’s Bait Shop, Long
Beach, California, and care was taken to verify
the original source of this material. After fixa-
tion in 10% formalin, the specimens were
washed and placed in 46% isopropyl alcohol.

Statistical Methods

The purpose of each statistical test in this
paper is solely to accept or reject a hypothesis.
The decision to accept or reject a hypothesis was
based on the 5% critical level, to minimize type
II errors (Hoel, 1954: 33). Acceptance of a
hypothesis such as equal means, however, merely
indicates no detectable differences and does not
imply that the populations in question are
proven to be the same. Critical values usually
are given; the subscript notation 0.95 refers to
a one-tail test, and 0.975 to a two-tail test.

Four kinds of tests were employed. The F-test
was used when the hypothesis involved the
testing of two variances. Covariance was applied
to problems when two or more regressions ( fit-
ted by least squares) were tested at the same
time. Decisions to accept or reject differences
between two means were determined by /-tests.
Correlation coefficients were evaluated as out-
lined by Hoel (1954: 124).

Obvious differences were not tested. When
several variances or means were compared, the
two most divergent values were examined first.
If no significant differences were revealed, the
others were not tested.

The statistical parameters of the fin ray counts
are presented graphically. The format is that
of Hubbs and Hubbs (1953). Unless the sep-
aration between means was patent, no statistical
inferences have been drawn from these figures.

RESULTS

Head

The head of fish from the Gulf of California,
as seen from the side, is more pointed than in
Pacific Coast specimens. In this respect the Gulf
populations approach seta. The difference in

PACIFIC SCIENCE, Vol. XVII, January 1963

TABLE 2

Fin Counts for Gillichthys mirabilis, MATERIALS AND LOCALITIES

°N

LOCALITY

COLLECTED

FIN COUNTS
n (SIZE RANGE)

Pacific Coast

37°27'

San Francisco Bay

June 1957

30(39-86)

35°20'

Morro Bay

August 1957

30(86-132)

34°24'

Goleta

September 1957

30(64-125)

34°00'

Venice

June 1957

28(83-138)

33°37'

Newport Bay

October 1955

25(56-154)

33°01'

San Elijo Lagoon

June 1956

30(58-142)

32°47'

Mission Bay

May 1946

30(38-54)

31°43'

Estero Punta Banda

July 1955, 1956

20(80-124)

30°30'

Bahia San Quintin

July 1956

14(91-142)

26°45'

Punta Abreojos

May 1948

16(10-20)

24°47'

Bahia Sta. Maria

April 1955

30(22-38)

Salton Sea

33° 18'

NW Shore

July 19531

30(55-109)

NE Shore

July 19541

30(40-78)

NW Shore

July 1955

30(35-74)

Gulf of California

31°2 r

Bahia Cholla

January 1949

30(20-30)

Bahia Cholla

April 1954

30(48-88)

31 ° 18'

San Felipe

April 1957

31(25-50)

27°57'

Estero Soldado

January 1952

31(15-20)

26° 54'

Mulege

March 1957

30(35-50)

26°22'

Bahia Agiabampo

January 1951

10(66-100)

1 Not included in Figures 6-12.

shape is due largely to the more depressed head
of the Gulf fish, and was one of the main reasons
for naming this form as a new species. In their
description, Gilbert and Scofield (1898: 498)
commented on the depressed head of Gillichthys
detrusus ( = G. mirabilis Cooper ) . This condi-
tion prevails in all the populations investigated
from the Gulf. The mean head depth for fish
from "El Marino” slough, just north of San Fe-
lipe, is 15.6% (13-18%; n = 10) of standard
lenth, which agrees nicely with the value for
Bahia Agiabampo fish (Barlow, 196L*). It
should be noted at this point that the position
of the opercles and the branchiostegals of pre-
served gobies can cause considerable variation
in measurements of head depth and especially
of head width.

The head shape of Salton Sea fish is more like
that of specimens from the Pacific Coast than of
those from the Gulf. The measured head depth,
though, is closer to that of Gulf populations
(Barlow, 196 1^).

The regression of head length on standard
length is slightly, but significantly, greater for
the Pacific Coast populations than for the fishes
from the Gulf when the data are pooled (Fig.
2, Table 3). Along the Pacific Coast the five
populations also differ significantly from one
another ( Table 3 ) . The differences between
Gulf populations are of approximately the same
degree as those between the Pacific Coast groups,
but this was not tested statistically. Fish from
the Salton Sea have the shortest heads, which is
immediately noticeable when inspecting large
adults.

Relative growth differences are seen best by
plotting the logarithms of head and standard
length (Martin, 1949: 8). The regressions for
all populations so presented are nearly parallel
(no figure) . These curves for relative growth of
the head, however, are slightly but clearly curvi-
linear; the slopes steepen with increasing stand-
ard length. The relative degree of difference
between any of the lines is the same for the

Gillichthys mirabilis — BarloW

FIG. 2. Head length versus standard length. Gulf of
California populations: open triangle, San Felipe (El
Marino); open square, Mulege; open circle, Bahia
Cholla, 1954; closed triangle, Guaymas (Estero Boco-
chibampo); closed circle, Agiabampo. Pacific Coast
populations: closed circle, San Francisco Bay (Alviso
ponds); open triangle, Goleta; closed square, Newport
Bay; open circle, San Elijo Lagoon; closed triangle,
Estero Punta Banda; open square, Bahia San Quintin.
Salton Sea population: open triangle, 1953; open
circle, 1954; open square, 1955.

smallest and largest fish measured, i. e., the lines
are parallel. Therefore, differences between
slopes of arithmetic plots presented here are
of no biological significance.

Upper Jaw

The analysis of upper jaw measurements is
complicated by the curvilinear relationship be-
tween jaw length and standard length. (Here-
after, upper jaw is referred to as jaw.) This dif-
ficulty is alleviated somewhat by fitting two re-
gressions through the data; one line for fish
greater than 90 mm, and another for the smaller
specimens (Fig. 3). The change in slope is an
artifact of the arithmetic plot; relative growth is
only slightly curvilinear. The situation is also

complicated by the sexual dimorphism of jaw
length in adult fish. The average length of the
jaw is slightly greater in males, even though the
range is about the same in both sexes. Conse-
quently, the results of the covariance tests (Table
3) should be regarded with skepticism, since
some of the assumptions underlying the test are
not entirely satisfied.

The jaw of fish from the Gulf is about equal
to, or slightly larger than, the jaw of Pacific
Coast fish when specimens of the same length
are compared. On the same basis, the jaw is
much shorter in fish from the Salton Sea.

If jaw lengths are compared as a function of
head length instead of standard length, the jaw
is still relatively longest in Gulf fish and shortest

Fig. 3. Upper jaw length versus standard length.
Gulf of California populations: open triangle, San
Felipe (El Marino); open square, Mulege; open
circle, Bahia Cholla, 1954; closed triangle, Guaymas
(Estero Bocochibampo) ; closed circle, Agiabampo. Pa-
cific Coast populations: closed circle, San Francisco
Bay (Alviso ponds); open triangle, Goleta; closed
square, Newport Bay; open circle, San Elijo Lagoon;
closed triangle, Estero Punta Banda; open square, Ba-
hia San Quintin. Salton Sea population: open triangle,
1953; open circle, 1954; open square, 1955.

PACIFIC SCIENCE, VoL XVII, January 1963

TABLE 3

Results of Co-Variance Tests on Differences in Morphology of Gillichthys mirabilis
(Hypothesis tested is equal means after adjustment for standard length.

Key: F, F ratio; d.f., degrees of freedom; F.05, 5% critical value.)

d.f.

F .95

DECISION

Head length

Gulf/Pacific/Salton Sea

80.3

2,299

3.03

reject

Gulf/Pacific

8.87

1,201

3.89

reject

Pacific (5 populations)

14.8

4,128

2.44

reject

Salton Sea ( 1953-55)

10.1

2,97

3.09

reject

Salton Sea (1954-55)

336

1,66

3.99

Upper jaw length

> 90 mm

Gulf/Pacific/Salton Sea

10.6

2,87

3.10

reject

< 90 mm

Gulf/Pacific/Salton Sea

41.6

2,208

3.04

reject

Salton Sea (1953-55)

7.99

2,84

3.10

reject

Salton Sea (1954-55)

<0.58

1,66

3.99

Anal fin height

Gulf/Pacific/Salton Sea

37.8

2,291

3.03

reject

Pacific ( 6 populations )

19.1

5,129

2.29

reject

Salton Sea (1953-55)

3.13

2,95

3.10

reject

in those from the Salton Sea. The jaw is 81, 74,
and 7 1 % of head length for Gulf, Pacific Coast
and Salton Sea fish, respectively. Thus, the jaw
of the Salton Sea fish is shorter in regard to
head length as well as standard length.

Median Fin Height

The soft dorsal and the anal fins are higher
in fish from the Gulf than in those from Cali-
fornia and the Salton Sea (Barlow, 1961*, table
1 ) . Only the variation in anal fin height is re-
ported in detail.

The anal fin height varies significantly from
population to population along the Pacific Coast
(Fig. 4, Table 3). San Francisco fish, for in-
stance, often fall below the fitted regression;
Newport Bay and Estero Punta Bunda speci-
mens usually are above it. The data are too few
to make an unequivocal statement about trends,
but the fins of northern fish tend to be lower
than those from the south.

Gilbert and Scofield (1898: 498) mentioned
the higher anal fin of detrusus ( = mirabilis
Cooper) when they described it as a new species.

The type locality is the tidal region of the Colo-
rado River delta, but high anal fins are also
typical of the populations to the south in the
Gulf (Fig. 4). The regression of fin length
on standard length for Gulf forms is higher and
steeper than those for the populations from the
Pacific Coast and the Salton Sea. The variation
between Gulf populations is assumed to be
significant. No trend with latitude is apparent.

One sample from the northwestern region of
the Gulf (an isolated salt pond 25 km north of
San Felipe) has remarkably low median fins,
and the dorsal fin profile is less rounded than in
the typical Gulf form (Fig. 5) - The height of
the anal fin averages 8.8% (8. 3-9.5%) of the
standard length (based on 10 specimens, 61-
102 mm in length). Thus the anal fin height
and the dorsal fin shape resemble those of fish
from the Pacific Coast. These data were not in-
cluded in the regression of fin height versus
standard length because the specimens had not
been examined at the time the calculations were
carried out and the figure was prepared. The fish
of this population are thought to be not repre-
sentative of the usual Gulf condition.

Gillichthys mirabilis — Barlow

Fig. 4. Anal fin height versus standard length. Gulf
of California populations: open triangle, San Felipe
(El Marino); open square, Mulege; open circle, Bahia
Cholla, 1954; closed triangle, Guaymas (Estero Boco-
chibampo); closed circle, Agiabampo. Pacific Coast
populations: closed circle, San Francisco Bay (Alviso
ponds ) ; open triangle, Goleta; closed square, Newport
Bay; open circle, San Eli jo Lagoon; closed triangle,
Estero Punta Banda; open square, Bahia San Quintin.
Sal ton Sea population: open triangle, 1953; open
square, 1954; open circle, 1955.

The anal fin height of Salton Sea fish (Fig.
4) is considerably lower than in Gulf groups.
Compared to Pacific Coast populations, the fin
heights are similar but nevertheless lower. The
variation in height from year to year in the
Salton Sea fish is just significant at the 5% level.

The profile of the second dorsal fin is more
rounded in specimens from the Gulf of Califor-
nia than in those from the outer coast or the
Salton Sea. This results from the relatively
greater increase in length of the centermost rays
in the higher fins. Besides the change in shape,
the anterior height of the second dorsal fin
equals or exceeds that of the first dorsal in Gulf
forms (Gilbert and Scofield, 1898: pi. 38). In
fish from the Pacific Coast and the Salton Sea,
the anterior rays are lower and the central rays
are equal to or only slightly higher than the
spiny dorsal.

Distance between Dorsal Fins

The distance between dorsal fins is greatest
in fish from the northern part of the Gulf of
California, ranging from 1.0 to 1.8 times the
eye diameter. The distance between fins of speci-
mens from the central and southern Gulf is
about the same as that of fish from central Cali-
fornia, being 0.7-1. 1 times the eye diameter.
The distance between dorsal fins was one of
the diagnostic characters used by Gilbert and
Scofield (1898: 498) to separate detrusus from
mirabilis.

In Salton Sea fish, the distance between fins
is less than one eye diameter. The posterior
margin of the first dorsal sometimes touches
the anterior margin of the second dorsal. A
greater number of elements in both dorsal fins
as well as an encroachment on the space by two
to three supernumerary spines on the leading
edge of the second dorsal fin cause the decrease
in distance.

Squamation

The scales are slightly larger in specimens of
mirabilis from the Gulf of California than they
are in those from the outer coast and the Salton
Sea. The counts range from 60 to 80 irregular
vertical rows of scales along the body in Gulf
forms, while there are about 80-100 rows in
fish from the Pacific Coast and the Salton Sea.

The ventral squamation is more extensive in
fish from the Gulf than in those from the Pacific
Coast or the Salton Sea. The scales extend an-
teriorly around the base of the pelvic fin in about
30% of the specimens from Bahia Agiabampo
(southern Gulf), in about 10% from Guaymas
(central Gulf), and in about 80% from Bahia
Cholla (northern Gulf). In fish from the Pa-
cific Coast, the scales only occasionally reach
anteriorly beyond the lateral insertion of the
pelvic fin base.

In many respects the scalation is less exten-
sive in Salton Sea fish, although the scales are
about the same size as those on fish from the
Pacific Coast. The ventral, anterior margin of
the scaled area usually just reaches the insertion
of the pelvic fins, but not beyond. The scaleless
area directly under the pelvic fins is greater than
in naturally occurring populations. The naked

area extends posteriorly to just beyond the tip
of the pelvic fins but is not as wide as the pelvic
fins; the size of the area is variable but is always
larger than in other populations. A vertical strip
in the axil, about one-half an eye diameter wide,
is also without scales. Further, the strip of scales
on the nape is shorter than in the other popula-
tions (Barlow, 1961a) . The base of the caudal
fin rays has two to six vertical rows of scales
on it. In this respect only, the Salton Sea fish
occasionally are more fully scaled than those of
the other populations.

PACIFIC SCIENCE, Vol. XVII, January 1963

populations tends to converge in the lower lati-
tudes.

The mean number of dorsal fin rays in the
Salton Sea population is greater by about one
ray than in the other populations.

The large range and variation of the New-
port Bay population are caused chiefly by one
specimen with only eight rays. The fin does
not look aberrant but it must be considered
atypical.

Anal Fin

First Dorsal Fin

The average spine number is about six for
all populations, although some means deviate
appreciably from this value (Fig. 6). The count
never was found to be less than six in Gulf
populations. Five spines, however, are not un-
common in other populations. The only geo-
graphic trend discernible concerns the variances
of the populations; this is reported below.

Second Dorsal Fin

The mean number of rays in the second dor-
sal fin is clearly higher in Pacific Coast than in
Gulf populations (Fig. 6). On the Pacific Coast
the counts are lower in the south, while in the
Gulf the counts are more nearly uniform, or
slightly higher in the south. Thus the average
number of fin rays for Gulf and Pacific Coast

On the Pacific Coast the mean number of
anal fin rays increases from the north, south to
San Diego, then decreases further south. In the
Gulf, where samples are less complete, the
means are about the same in all populations,
or are a little higher in the south. The means
of the Gulf and Pacific Coast populations con-
verge in the lower latitudes (Fig. 7).

The mean number of anal fin rays in the fish
from the Salton Sea is conspicuously higher
than in any of the naturally occurring popula-
tions ( Fig. 7 ) .

Pectoral Fin

The mean number of pectoral fin rays in-
creases progressively from north to south on
the Pacific Coast, but in the Gulf of California
the average number decreases to the south. Con-
sequently, the means of Pacific Coast and Gulf

FIG. 5. Gillichthys mkabilis, female, standard length 100 mm, from "El Marino” slough north of San
Felipe, Baja California. The small black spot on the side of the specimen is a blemish.

Gillichthys mirabilis ■ — -BARLOW

Fig. 6. Fin rays in the first (spiny) and second
(segmented only) dorsal fins, from left to right, pre-
sented as a function of °N latitude. Materials and
place names in Table 2. Large rectangle, one standard
deviation on either side of mean (triangular promi-
nence) ; blackened space, two standard errors on either
side of mean; lower horizontal line, range of counts.
Open rectangle, Pacific Coast populations; vertical lines
in rectangle, Salton Sea population; oblique lines in
rectangle, Gulf of California populations.

populations overlap in the southern latitudes
(Fig. 8).

The average pectoral fin count of the Salton
Sea fish again is much higher than in Pacific
Coast animals. The number of fin rays, how-
ever, is similar to that found in northern Gulf
populations (Fig. 8).

The Venice population, near Los Angeles
on the Pacific Coast, is extremely variable. Some
pectoral fins are deformed and obviously are
aberrant. This habitat is polluted by sewage
and industrial waste products.

Combined Fin Counts

Some of the trends and differences observed
in the study of individual fins disappear when
the fin counts are combined and plotted as a
function of latitude (Fig. 9). At different lati-
tudes the averages of the combined fin counts

are about the same, except in California where
a slight increase from north to south occurs.
Otherwise there is no trend in the mean num-
ber with latitude. When comparisons are made
at the same latitudes, the Pacific Coast popula-
tions and all of the Gulf populations, except
for San Felipe, broadly overlap. Contrariwise,
some divergences are enhanced. The Salton Sea
and San Felipe populations are obvious deviants.
Further discussion of combined fin elements
is reserved for the section on annual variation.

Fin Ray Variance

Over the geographic range of mirabilis there
are dissimilarities in the variances of each of
the fins. The difference between the extreme
variances of the first dorsal fin on the Pacific

Fig. 7. Fin rays in the anal fin (segmented only)
presented as a function of °M latitude. Materials and
place names in Table 2. Large rectangle, one standard
deviation on either side of mean (triangular promi-
nence); blackened space, two standard errors on either
side of mean; lower horizontal line, range of counts.
Open rectangle, Pacific Coast populations; vertical
lines in rectangle, Salton Sea population; oblique lines
in rectangle, Gulf of California populations.

PACIFIC SCIENCE, Vol. XVII, January 1963

Fig. 8. Fin rays in the left pectoral fin presented as
a function of °N latitude. Materials and place names
in Table 2. Large rectangle, one standard deviation on
either side of mean (triangular prominence); black-
ened space, two standard errors on either side of mean;
lower horizontal line, range of counts. Open rectangle,
Pacific Coast populations; vertical lines in rectangle,
Salton Sea population; oblique lines in rectangle, Gulf
of California populations.

Coast (San Francisco Bay and Bahia Santa
Maria), for instance, is highly significant: F =
16.9, F0.975 (29, 29) = 2.1.

The intrinsic variability in the number of
elements in a given fin is associated with the
latitude where the population is situated. In-
creasing variability is associated with higher
latitudes (Fig. 10). All the fins manifest this
increase in variability from south to north,
though not equally well. If one were to rate
the correlations subjectively from good to poor,
they would stand in the order ( 1 ) first and
second dorsal fins, (2) combined counts, (3)
anal fin, and (4) pectoral fin. Only the data
from the first dorsal fin are presented here
(Fig. 10); the other data are available on re-
quest. There also may be a positive correlation

between latitude and variance of the vertebrae
(see below).

The correlation between degrees of latitude
and variance of the first dorsal fin has a coeffi-
cient of 0.848, which differs significantly from
a coefficient of zero. This high degree of cor-
relation doubtless could be increased further
by reducing that portion of the intra-population
variation resulting from sampling techniques,
thereby improving the estimate of the real vari-
ance. As will be seen further on, comparisons
of relatively homogeneous year classes from suc-
ceeding years show that even though the mean
number of elements in the various fins may
change, the variance remains the same. Some

Fig. 9- Combined elements, segmented and unseg-
mented, in the first and second dorsal, anal, and left
and right pectoral fins, presented as a function of °N
latitude. Materials and place names in Table 2. Large
rectangle, one standard deviation on either side of
mean (triangular prominence); blackened space, two
standard errors on either side of mean; lower hori-
zontal line, range of counts. Open rectangle, Pacific
Coast populations; vertical lines in rectangle, Salton
Sea population; oblique lines in rectangle, Gulf of
California populations.

Gillichthys mirahilis — Barlow

Fig. 10. Correlation between population variances
(±) of the first dorsal fin, and °N latitude. Open
circle, Pacific Coast populations; closed triangle, Salton
Sea population; closed circle, Gulf of California
populations.

of the samples represented by a single value
in Figure 10 are heterogeneous, consisting of
more than one year class. Similar dispersion
in each year class, but about different means,
would increase the apparent variation.

Fin Ray Correlations

The segmented rays of the dorsal and anal
fins are positively correlated. When all the
counts are combined, exclusive of the Salton
Sea, the correlation coefficient is 0.603, signifi-
cantly greater than zero. A positive correlation
also is evident within and between year classes.

The number of elements in the pectoral fin
compared to that of the anal, and of the soft
dorsal fin, reveals negative correlations, but only
when means of different populations are com-
pared ( Fig. 11). The correlation coefficient for
the pectoral fin with the second dorsal fin is
-0.737, and with the anal fin it is -0.659; both
values differ significantly from zero. The data
for the sample from San Felipe are shown in
Figure 11 (the highest pectoral fin mean) but
are not included in the computations of the
correlation coefficients; as mentioned before,
the counts are atypical. The inclusion of these
counts does not affect the conclusion that the
pectoral and median fin counts are significantly
correlated. Data from samples from the Salton
Sea also are not included.

Fin Ray Index

Probably the most useful statistic obtained
from the fin counts is a derived index. The
soft rays of the dorsal and anal fins are summed,
then divided by the number of rays in the left
pectoral fin and multiplied by 100. This index
reflects the change in relationship between the
fin ray numbers of pectoral and median fins at
different latitudes (Fig. 12).

On the Pacific Coast, not including San Fran-
cisco and Venice, the index decreases progres-
sively from north to south. In the Gulf of Cali-
fornia the opposite is the case; the index in-
creases to the south. The indices of Gulf and
outer-coast populations consequently converge
at low latitudes, being slightly higher in outer-
coast populations.

Noteworthy is the excellent agreement of
index values for samples from Bahia Cholla and
San Felipe, situated at similar latitudes but on
opposite coasts of the Gulf. This is especially
striking, considering that the San Felipe fish
have unusually high atypical counts.

The most revealing index is that of the Salton
Sea population. It agrees with values obtained
from samples taken in the vicinity of San Diego,
the source of the Salton Sea stock. Further, the
index is considerably higher than in any of the
Gulf populations.

< io
z

19 20 21 22 23

PECTORAL

Fig. 11. Correlation between population means of
number of pectoral fin rays, and number of segmented
rays in the anal (closed circles) and dorsal (open
circles) fins.

PACIFIC SCIENCE, Vol. XVII, January 1963

Fig. 12. Fin ray index: combined number of seg-
mented rays in the second dorsal fin and anal fin,
divided by the number of rays in the left pectoral fin,
multiplied by 100. Open circles, means of Pacific
Coast populations; closed circles, means of Gulf of
California populations; closed triangle, mean of Salton
Sea population; horizontal lines, ranges of values.

Pectoral Fin Asymmetry

Species of fishes are known to differ in de-
gree of asymmetry between the number of ele-
ments in the right and left pectoral fins ( Hubbs
and Hubbs, 1945). The numerical asymmetry
between right and left pectoral fin counts in
mirabilis was investigated to determine if in-
dividual populations, or geographic regions,
could be characterized by this feature (Table 4).
Only the total amount of asymmetry, right plus
left, was found to be useful in this respect.
Values for the Pacific Coast and Salton Sea pop-
ulations are usually higher (mean about 30%)
than those from the Gulf (mean about 20%).
The amount of asymmetry in the two most
southerly populations on the Pacific Coast, how-
ever, is the same as that of the Gulf forms.

The degree of asymmetry is equally divided
between right and left. To facilitate compari-
sons, each population can be characterized ac-
cording to whether there are more fish with
higher counts in the right fin than in the left
fin, or vice versa. If a sample has more fish with
right fins of a higher count than the left, it
is tallied as a "dextral”; and with more lefts
than rights, as a "sinistral.”

Among collections from the Pacific Coast 5
out of 11 are dextral. In the Gulf, 3 out of 6
are dextral, 1 is sinistral, and 2 are equal. The
Salton Sea population vacillates from year to
year between dextral and sinistral. Therefore,
the distribution of rights and lefts within and
between populations is typical of a 50-50
chance situation, such as the toss of a coin.

Vertebrae

The number of vertebrae was compared in
five populations (Table 5). In every sample
the mode is 32, and the average is approximately
the same value. Only the mean of the Salton
Sea specimens differs appreciably from 32; it
is significantly higher (t = 2.02; to. 95 (29) =
1.70).

In contrast to the relative uniformity of the
means, there are patent differences between the
variances; they are greater in the Salton Sea
and Pacific Coast groups than in those from the
Gulf. Each of the variances of the San Francisco
and Salton Sea samples was tested against that
of the northern Gulf collection; the F-ratios
are F = 13.6, F0.975 (18, 39) = 2.4, and
F = 8.6, F 0.97 5 (29, 39) = 2.1, respectively.
The variances of the San Francisco and Salton
Sea groups are not significantly different: F =
1.6, F0.975 (18, 29) = 2.5.

The variances of the vertebrae decrease from
high to low latitudes. If the variances are listed
from north to south, instead of as shown in
Table 5, the following series of values is ob-
tained: 0.577, 0.461, 0.182, 0.157, 0.341, 0.0.
Considering the few populations represented,
the correlation is surprisingly good. The pres-
ence of a similar positive correlation between
latitude and fin ray variance (see above) in-
creases the confidence in the trend indicated
by the variances of the vertebrae.

Gillichthys mirabilis — Barlow
Annual Variation

Material for the study of annual variation
came from the Salton Sea and from Bahia Cholla
(northern Gulf of California). Each sample was
from a year class hatched in the year consid-
ered. This affords a rough estimate of the sta-
bility of the statistical parameters used in the
comparison of separate populations.

The Salton Sea was sampled during the same
month on three successive years. The fish of
one sample are larger than the others ( Table 6 )
and probably were hatched earlier in the year
than those of the other two samples. In addi-
tion, the 1954 collection was from a different
part of thd Sea, but I doubt that this is of im-
portance. Both body measurements and fin
counts were taken.

The two samples from Bahia Cholla came
from year classes that had hatched under strik-

ingly different temperature conditions. The first
were embryos during an unusually cold period
late in the winter of 1949. A moderate fish kill
occurred in the northern Gulf at that time. The
second collection, 1954, consists of fish that had
developed in a relatively normal temperature
situation. Only counts were taken because the
specimens are small.

Slight, but significant, differences occur be-
tween the means of fin counts of different year
classes from the Salton Sea (Table 6). In the
1955 class, all counts are higher than in the
previous two years (except for the first dorsal
fin: 1955 vs. 1953, t = 0.67, *0.95 (58) =
1.67). The mean of the second dorsal fin ex-
ceeds that of the anal fin by about one ray in
each year class.

The ratios of median fin rays to pectoral
rays, as indicated by the fin indices, remain ap-
proximately the same. Thus while the means

TABLE 4

Bilateral Asymmetry of Pectoral Fins
Arranged by Regions, from North to South

PERCENTAGES

AREA

L > R

R > L

R + L

Pacific Coast

San Francisco

Morro Bay

Goleta

Venice1

Newport Bay

San Elijo Lagoon

Mission Bay

Estero Pta. Banda

Bahia San Quintin

Pta. Abreojos

Bahia Sta. Maria

Salton Sea

1953

1954

1955

Gulf of California

Bahia Cholla — 1949

Bahia Cholla — 1954

San Felipe

Estero Soldado

Mulege

Bahia Agiabampo

1 Aberrant; see pectoral fin section.

PACIFIC SCIENCE, Vol. XVII, January 1963

TABLE 5

Vertebral Counts for Gillichthys mirabilis
(Key: x, mean; s, one standard deviation.)

NO.

OF VERTEBRAE

STATISTICAL PARAMETERS

AREA

YEAR

±s

Pacific Coast

San Francisco

1955

32.00

0.577

Pta. Abreojos

1948

31.88

0.341

Gulf of California

Bahia Cholla

1949

32.03

0.182

San Felipe

1955

32.03

0.157

Est. Soldado

1952

32.00

Salton Sea

1955

32.17

0.461

of fin ray counts may change, the relationships
between them are relatively unaffected.

The divergence of means between the year
class samples from Bahia Cholla is greater than
that recorded in the Salton Sea material. The
mean number of rays for each fin, except the
first dorsal, is higher in the "cold” year than in
the "normal” year fish. The differences are sig-
nificant at the 5% critical level. The outstand-
ing deviation is in the mean of the second dorsal
fin which is higher by one full ray.

The relationship between the counts of the
fin rays also is different in the 1949, or "cold”
year class from Bahia Cholla. This results from
the disproportionate increase of the soft dorsal
fin count, which exceeds the anal fin by 1.7 rays.
In most populations, as in the 1954 collection
from Bahia Cholla, the average number of rays
in the second dorsal fin surpasses that in the
anal by only about one ray (0.8 to 1.3).

The fin index also reflects the change in the
relationship between the fin counts. It is higher
by 4% in the 1949 sample, probably due to
the unusual rise in the number of dorsal fin rays.

Although the average number of fin rays and
their interactions may change from year to year,
the variance of each fin remains the same. F-
tests were performed using the maximum and
minimum variance of each fin within the year
classes of each population; all F-ratios were
below the 5% level of significance.

The direction of the annual variation in fin
counts is of particular importance to conclusions

drawn later. In each instance when a year class
deviates to a noticeable degree, the direction of
the changes are the same for all fins with seg-
mented rays. Increases in the dorsal and anal fin
counts, as just shown, are paralleled by an in-
crease in the number of pectoral fin rays; i. e.,
they are positively correlated. Within any one
year class, however, there is no correlation what-
soever between the number of segmented rays
in median fins and in the pectoral fins (Table
7 ) . The pectoral fin counts are the same, on
the average, whether a fish has many or few
median fin rays, when from the same year class.

The bilateral asymmetry of the number of
pectoral fin rays fluctuates between samples of
year classes from the Salton Sea and from Bahia
Cholla. Both the amount and the direction of
the asymmetry vary (Table 4). The total asym-
metry is maximal in the Salton Sea sample for
1955 and in the Bahia Cholla collection taken
in 1949.

The head and jaw length and the anal fin
heights are about the same in the three year
classes collected from the Salton Sea. When the
three groups are tested together, the differences
are significant (Table 2), although the com-
parison of anal fin heights for all three samples
just exceeds the 5% level of significance. If
only the 1954 and 1955 samples are compared,
the differences no longer are significant (Table
3). The size ranges of the 1954 and 1955 sam-
ples nearly coincide, while that from the 1953
year class contains larger fish. The inclusion in

Gillichthys mirabilis— Barlow

the test of the sample with the larger fish ap-
parently causes the significant differences be-
tween the groups. Hence the differences between
the three year classes seem to be caused by
unequal size groups.

Peculiarities of G. mirabilis from the Salton Sea

The mirabilis introduced into the Salton Sea
differ from naturally occurring populations in
several ways. Most of these differences seem
to indicate an interference with the normal de-
velopmental physiology.

Some of the unusual features of the Salton
Sea fish have already been discussed in earlier
sections, and they are enumerated below. All
the counts of fin rays and vertebrae are higher
in these fish than in any others. The relation-
ship between the fin ray numbers as reflected by
the fin index, however, has remained the same
as that of the parental stock. The squamation is

less extensive, and the head and upper jaw are
disproportionately small. The formation of the
interorbital pore is delayed.

The Salton Sea fish differ from naturally oc-
curring forms in still other ways not yet de-
scribed. The otoliths are much coarser and more
opaque than those of specimens from the outer
coast. The outline of the otoliths, as seen from
either flat side, is less angular and is often al-
most round. The edges of the otoliths usually are
only weakly crenulate or smooth, and the char-
acteristic notch is indistinct.

In gobiid fishes, the segmented rays of the
soft dorsal and anal fin typically are preceded
by one slender spine. This holds for mirabilis
in naturally occurring populations, with rare
exceptions. Supernumerary spines, however,
have been found on two specimens of mirabilis
from San Eli jo Lagoon and one from Newport
Bay, and also on one specimen of seta from San

TABLE 6

Annual Variation of Fin Counts of Gillichthys mirabilis
(Key: x, mean; s, one standard deviation; r, range.)

COLLECTED

SALTON SEA

GULF OF CALIFORNIA

NW Shore

18 July

1953

NE Shore

17 July

1954

NW Shore

13 July

1955

Bahia Cholla
27 January
1949

Bahia Cholla
27 April
1954

Size range in mm

55-109

40-78

35-74

20-30

48-82

First dorsal

6.23

6.27

6.07

±s

0.504

0.451

0.521

0.255

0.253

(5-7)

(6-7)

(5-7)

(6-7)

Second dorsal

12.43

12.33

12.57

11.33

10.37

±s

0.679

0.547

0.679

0.480

0.555

(11-14)

(12-14)

(11-14)

(11-12)

(9-11)

Anal

11.27

11.33

11.60

9.60

930

±s

0.583

0.480

0.814

0.564

0.702

(10-12)

(11-12)

(11-14)

(9-11)

(8-11)

Pectoral (left)

21.33

21.47

21.30

20.97

c±S

0.679

0.480

0.628

0.625

0.616

(20-23)

(21-22)

(20-22)

(20-23)

(20-22)

Combined

74.93

74.67

75.70

71.73

69.67

±s

1.68

1.18

1.95

1.53

1.65

(72-79)

(73-77)

(71-80)

(69-76)

(66-72)

Index

112

111

113

PACIFIC SCIENCE, Vol. XVII, January 1963

TABLE 7

Relationship between the Pectoral Fin Rays and the Sum of the Segmented Rays of the
Second Dorsal Fin and Anal Fin in Gillichthys mirabilis

DORSAL PLUS ANAL

SEGMENTED RAYS

San Eli jo Lagoon

mean pectoral rays
(n)

20.0

(9)

20.0

(10)

20.0

(9)

20.0

(2)

Salton Sea

1953

mean pectoral rays
(n)

21.0

(2)

21.1

(ID

21.2

(9)

21.7

(6)

21.0

(2)

1954

mean pectoral rays
(n)

21.5

(17)

21.1

(8)

21.2

(6)

1955

mean pectoral rays
(n)

22.0

(1)

21.5

(ID

21.3

(6)

21.5

(8)

21.0

(2)

22.0

(2)

Bahia Cholla

mean pectoral rays
(n)

21.0

(1)

20.0

(1)

21.0

GO)

21.0

(15)

21.0

(3)

Felipe. But in fish from the Salton Sea, the
segmented rays of each of these fins are com-
monly preceded by two, or occasionally three,
spines. When two spines are present in the soft
dorsal or anal, the anterior-most spine is short.
Usually it is less than one-half the length of
the first segmented ray. Sometimes it is difficult
to find the anterior-most supernumerary spine.
In the soft dorsal only, the first of three spines
is always short (usually a stub) and sometimes
does not reach through the skin, while the sec-
ond usually is moderately well developed and
may be greater than half the length of the first
segmented ray.

Supernumerary spines occur in the second
dorsal fin more often than in the anal fin. In
samples from 3 successive years (in each, n =
30), the percentages of the fins with extra spines
are: second dorsal = 40, 13, and 33; anal =
10, 3, and 3. These data are from direct obser-
vations.

Photographs with X-rays revealed the pres-
ence of additional spines beneath the skin and
of their basal elements. In one collection (n =
30, from 1955, but a different sample than the
one mentioned in the preceding paragraph),

53% of the specimens have two spines in the
soft dorsal and 10% have three (total =z 63%).
In the same sample, 13% of the fish have two
spines in the anal fin, but none have three.

The situation is further complicated in that
the number of spines does not always agree
with the number of basals. Three specimens
with only one spine in the fin have two basal
elements. There is an additional basal anterior
to the basal of the first, and only, spine. In two
specimens, on the other hand, two spines share
a single basal. In these cases, the first spine is
merely a stub. In the specimens with three
spines, three basals are found. Generally, ele-
ments are added anteriorly onto the anal fin,
and especially the soft dorsal fin. This is done
either by adding spines, basal elements, or both.

Axial deformities are conspicuous in fish
from the Salton Sea (Fig. 13). Mostly they are
situated anterior to the origin of the second
dorsal fin. These are predominantly direct dorsal
flexures, but may tend to right or left. Posterior
to the first dorsal fin, the flexures are divided
about equally between dorsal, lateral, and ven-
tral. A few specimens have severe lateral flexures
in the hypural plate (Fig. 13). More than one

Gillichthys mirabilis- -Barlow

flexure per fish is not uncommon. Photographs
with X-rays indicate that the structure of the
vertebrae has broken down at the apex of the
more extreme curvatures. Whether there is a
disintegration and/or fusion of the vertebrae is
not clear from the photographs.

The occurrence of axial deformities increases
during the hottest months of the year (Table
8). During July, the incidence of twisted speci-
mens increases from around the usual 25% to
30'%. The maximum occurs in the period of
October to December. (The very high value
for October is based on a small sample and
therefore is subject to greater errors of random
sampling. There is also a serious problem in
obtaining a truly random sample, because of
the sudden appearance of deformities in the
young fish.)

Axial aberrations usually are manifest for the
first time in subadults during August, the peak
of the hot season. There is no relationship be-
tween size and incidence of deformities in these
young fish. The deformities appear during the
same period of time but not at a given size.

Axial deformities are also apparent in the
sciaenid fishes that have been introduced into
the Salton Sea. Here, too, anterior kyphosis
seems to be prevalent.

Other abnormalities are seen in mirabilis but
are not as common as the axial terata. These
include forward projection of the lower jaw
(Fig. 13), dermal flaps on the upper jaw, and
pronounced elongation of individual rays in
the median fins.

DISCUSSION

Body Form

Differences in size and shape are apparent
in comparisons between various kinds of fishes
from waters of different temperatures. Almost
invariably in the Northern Hemisphere, the
more northern representatives of a species or
of a genus are larger than those to the south
(Hubbs, 1926: 60; Vladykov, 1934: 120). The
changes in body proportions have not been as
thoroughly investigated, but some tentative gen-
eralizations can be made. Northern, slowly grow-
ing races of a species usually have smaller heads,
eyes, maxillas, and fins than do their southern
counterparts, although opposite effects are not
uncommon (Hubbs, 1926: 62; Vladykov, 1934:
118; Martin, 1949: 23; and included refer-
ences ) .

Morphological differences in Gillichthys mira-
bilis, as indicated by lines fitted to part lengths
versus standard length, are apparent between
the aggregates of populations from the Gulf of
California and the Pacific Coast, and between
the populations within these regions. There is
no pattern among the regressions (length of
parts on standard length) which might provide
a due to the nature of the differences. There is
only the slightest tendency for the height of the
anal fin to be greater in more southern popula-
tions of the Pacific Coast region.

The Gulf populations differ, for the most
part, from those on the outer coast in the way
one would predict for faster developing fish.

Fig. 13. Gillichthys mirabilis, female, standard length 101 mm, from the Salton Sea, California, and typical
of extreme terata. Fold below the pectoral fin is from an incision.

PACIFIC SCIENCE, Vol. XVII, January 1963

TABLE 8

Axial Deformities in Gillichthys mirabilis
FROM S ALTON SEA

MONTH

TOTAL

FISH

DEFORMED

FISH

PERCENT

DEFORMED

1955

April

May

June

124

July

August

October

December

1956

January

136

February

The Gulf fish have higher fins and longer jaws,
the head is more depressed, and they tend to be
more completely scaled verier ally and have
slightly larger scales. The head is shorter, how-
ever. Except for the slightly shorter head length,
the form of mirabilis from the Gulf is consistent
with a hypothesis of rapid development. As seen
in the fish from the Salton Sea, shorter head
length is correlated with lower fins, shorter jaws,
and reduced scalation, and presumably this has
resulted from retarded development. If, as sug-
gested, the Gulf fish differ morphologically be-
cause of faster development, then they should
have longer, not shorter, heads. This contradic-
tion probably stems from genetic differences.

No satisfactory explanation can be given for
the distinctly greater distance between dorsal
fins of the specimens of mirabilis from the up-
per Gulf as compared with the other popula-
tions. The form of the fish from the upper Gulf,
however, resembles that of fish from the lower
Gulf, except for the distance between dorsal
fins. Environmentally induced changes usually
manifest themselves in several ways, instead of
being expressed in one character only. The in-
creased space between dorsal fins, therefore,
most likely has a genetic basis.

Countable characters

One of the commonly accepted generaliza-
tions in ichthyology is that the number of ele-
ments in serially repeated characters tends to

be greater at higher latitudes (Vladykov,
1934: 102). Hubbs (1922, 1926) argued that
this relationship depends chiefly on the tem-
perature of the water during early development;
lower temperatures were thought to result in
increasing numbers of elements, and higher
temperatures in lower numbers. Local temper-
ature conditions have been observed to be asso-
ciated with the anticipated changes in counts
(reviewed in Barlow, 1961$). Moreover, any
factor of the environment affecting rate of de- •
velopment, such as salinity or oxygen tension,
seems to have effects similar to those thought
to be produced by the thermal regime (Hubbs,
1926; Taning, 1952; Seymour, 1956).

There are exceptions, however, where the
change in counts goes contrary to- expectations.
In certain instances (Schmidt, 1919: 148;
Hubbs, 1921: 150; 1924; Schultz, 1927: 420)
the counts from one fin only showed an increase
associated with warmer temperatures. The other
fins manifested the normal negative correlation
between number of elements and water tem-
perature.

In spite of laboratory findings that the rela-
tionship between meristic structures and devel-
opmental temperatures is exceedingly complex,
the pattern in natural populations of the same
species is usually straightf orward : higher tem-
peratures during development normally bring -
about lower counts (Barlow, 1961$).

At first glance the changes in countable char-
acters over the geographic range of mirabilis
seem to indicate a gradient resulting from direct
modification by the environment. Presumably,
factors which cause a decrease in the dorsal and
anal fin counts conversely bring about an in-
crease in the number of pectoral fin rays.

Geographically, then, the number of median
fin rays is negatively correlated with the num-
ber of pectoral fin rays. Johnsen (1936: 8-9)
reported the same relationship between fins for
various populations of Gobius flavescens and
Gobius minutus .. These changes could be con-
strued as the result of dissimilar water tempera-
tures or some related factors.

It is more probable, nonetheless, that the
cline in meristic characters results primarily
from different genotypes, not exclusively from
environmental modification. This hypothesis can

Gillichthys mirabilis— BAMLOW

be supported by more than one line of evidence.

The gradient of meristic characters in mira-
bilis follows the temperature gradient of the
late summer period. The number of rays in the
dorsal and anal fins increases from north to
south in the Gulf of California, whereas over
the same latitude on the Pacific Coast the oppo-
site relation holds, and the counts tend to de-
crease from north to south. The same reversal
of trends between the Gulf and outer coast oc-
curs in the pectoral fin.

The fin indices decrease from north to south
on the Pacific Coast, but continue to decrease
in the Gulf from south to north. Thus a con-
tinuous dine in the meristic characters exists
from. San Francisco south, on the Pacific Coast,
and then north up into the Gulf of California.

It might be argued that the late summer tem-
perature gradient in the Gulf is opposite to that
of the outer coast, and that this environmenal
difference directly accounts for the reversal of
the fin-index gradient. This cannot be the case,
because the counts are determined while the
eggs and larvae develop during late winter
through early summer; then the temperature
gradients of the Gulf and Pacific Coast are
parallel, warmer to the south. If any correlation
exists between environmental temperatures and
number of countable elements, it is with the
gradient of extremely warm temperatures found
late in the summer, not those occurring during
early development.

This dine, however, might reflect nothing
more than the progressive physical separation
between populations.

The fin index for seta in the upper Gulf of
California continues the trend noted in mira-
bilis , that is, for lower indices toward the north
in the Gulf. The average value for seta there
is about 82 as compared to about 95 for adja-
cent populations of mirabilis. The temperature
situation that prevails in the habitat of seta
could be regarded as an extension of the gra-
dient seen in the habitats of mirabilis from
south to north in the Gulf. The habitat of seta
is located on the coast in the high intertidal
zone among black lava flows, boulders, and
streams of seepage water (Barlow, 196L*). Cli-
matically this environment is even more rigor-
ous, and surely experiences higher summer

temperatures than do the nearby sloughs where

mirabilis occurs.

Returning to the fin indices of mirabilis,
specimens from the Salton Sea, compared to
their parental stock, have much higher counts
in their dorsal and anal fins, and in their pec-
toral fins as well The differences probably can
be attributed to retardation of developmental
rate. If the interaction between these fins be-
haved as expected from observations on the
geographical dine, the pectoral fin counts should
have decreased while the median fin counts in-
creased. As shown by the fin index, the rela-
tionship between median and paired fins, re-
markably, is unchanged in these fish.

Fluctuations in the mean number of seg-
mented rays from year to year in a given popu-
lation are in the same direction for the median
fins and the pectoral fins. In other words, the
changes in the number of elements in paired
and median fins are positively correlated in their
response to environmental changes. Colder years
induce the formation of more rays in all of
these fins.

In each fish, however, the number of pectoral
fin rays appears to be inherited independently
from the number of median fin rays. In a given
year class, the fish with more numerous rays in
the median fins have, on the average, the same
number of pectoral fin rays as do fish with less
median fin rays. Thus the median fin and the
pectoral fin elements respond to the environ-
ment in the same way, but independently of
each other.

From the foregoing evidence it seems reason-
able to conclude that the change in relationship
between the number of elements in the median
fins and in the pectoral fin, the fin index, over
the geographic range of mirabilis indicates ge-
netic divergence.

Analogous differences between inter- and
i ntra- population al variation have been observed
in other gobies (Johnsen, 1936: 8-9) . In dif-
ferent populations of Gobius flavescens and G.
minutus , the vertebral counts (median fin counts
follow vertebral counts) are negatively correlated
with pectoral fin counts. In contrast, these meris-
tic characters are positively correlated within a
given population; larger fish have more verte-
brae and average more pectoral fin rays as well.

PACIFIC SCIENCE, VoL XVII, January 1963

Johnsen indiscriminately attributed both kinds
of variation to direct modification by the en-
vironment.

A further similarity between the results of
Johnsen s investigation and the present study
lies in the course of divergence of the fin index
(calculated from his data). There is a geo-
graphic cline in the index which is minimal in
the Baltic Sea and increases to the west and
then independently south and north in the North
Sea, but is maximal to the north.

In Gillie bt by s mirabilis , a genetic cline is
also suggested by comparisons of population
means of combined fin ray counts. Variation
between year classes and a comparison of Salton
Sea fish with their parental stock indicate that
slower development is correlated with an in-
crease in the total number of fin elements. If
the developmental rates within the populations
are altered by climatic temperatures, and this
is a reasonable assumption, then the combined
counts should decrease from north to- south. But
the combined counts are about the same for all
populations, with a few exceptions, and the ex-
ceptions are obvious deviants.

One could argue for a cline of combined
counts for the seven populations in California.
But the counts increase, not decrease, steadily
from San Francisco south to San Diego. Such
differences from north to south could be at-
tributed to increasingly saline water to- the south,
or to later spawning seasons to the north. Yet
the northernmost habitat, the Alviso salt ponds,
is highly saline, and spawning commences at
about the same time throughout the range of
mirabilis .

There seems to be homeostasis of the com-
bined counts in spite of changes in latitude,
since over the rest of the range of mirabilis the
means are about the same from population to
population. This suggests genetic compensation,
or relatively complete acclimation of develop-
mental rates. The most obvious deviant, the
population from just north of San Felipe, is
also atypical in body form.

The variance of the countable characters in
each population is even more stable than the
mean of the characters. It does not appear pos-
sible to estimate the respective roles of heredity
and environment in producing this constancy.

Even though the means of the counts were
found to shift within limits from year class to
year class, the variances showed no statistical
differences. The environmental changes respon- j
si hie for the changes in mean counts, most likely
temperature, apparently do not influence the
variances. This would seem to indicate an in-
trinsic homeostasis where variability is con-
cerned.

Photoperiod might determine variability. Day j
length is constant from year to year on the
same day at a given location. Since the change
in photoperiod over the spawning season in-
creases faster at higher latitudes, one can demon- ||
strate a positive, although perhaps false, correla-
tion between degree of change in day length and I
variance. Interestingly, estimates of the variances )
of the different fins in populations of a different
but closely related species, Gillichthys seta ( Bar-
low, 1961*), fall directly in the midst of the
corresponding data (Fig. 10) for mirabilis . \
Somewhat further aside, in two different popula-
tioes of seta the measurements of four body j
parts, not counts, having different means had
the same variances (Barlow, 1961*).

If the variance of each character is largely
genetically determined, then the data indicate
two different types of dines in mirabilis. Based j
on the variances, there would be a simple linear
relationship, north to south, irrespective of
whether the populations are situated on the
Pacific Coast or on the east or the west coasts of
the Gulf of California. The V-like arrangement
deduced from the distribution of the means of
the counts, the fin indices, and measurements of
body parts have already been described and jj
will be summarized below.

The possible relationship between the dif- jj
ferential of the photoperiod and the variance j
cannot be considered as evidence for or against |j
modification or genetic determination. Even ■
though the environment originally may have
induced the variance in question, the phenotype
easily could have become genetically reinforced.

The kind and pattern of the differences found .
between the populations of mirabilis seem too *1
complex to be accounted for on the assumption
that the dissimilarities result chiefly from direct
environmental modification. The most obvious j
part of the explanation is that the divergences i

Gillkhthys mirabilis-— Barlow

reflect the degree of physical separation between
populations. The less the exchange of genetic
material, the greater the opportunity for dis-
similarities to arise and persist.

If we are to understand why the variation
is in a particular direction and not random,
then we must first seek correlations between
environmental factors and the observed geo-
graphic cline. Temperature and illumination are
the only obvious parameters suitable to such an
inquiry. The only evidence available relates to
temperature differences, so illumination will
not be treated. This raises the problem, inherent
in such an approach, of formulating a self-
realizing assumption.

I have already pointed out the crude correla-
tion between the dines of meristic characters
reported on and the gradient of temperatures
found late in the summer. In another article
(Barlow, 1961b) , I have argued that relatively
stable differences in the means of counts be-
tween populations probably reflect physiological
differences, most likely temperature responses,
that are genetically determined. Changes in
counts between populations, as suggested by
Hubbs (1928), might constitute examples of
the well-known Baldwin effect (Baldwin,
1896): phenotypic modification of the counts
(resulting from physiological adjustment) be-
comes genetically augmented and characterizes
the population. A similar argument might hold
for the differences in variances that apparently
are associated with different photoperiods.

Peculiarities of G. mirabilis from the Salton Sea

The Salton Sea is a saline lake situated in the
Salton Sink, a northward extension of the Gulf
of California basin. According to Carpelan
(1958), it has a surface area of about 340
square miles, but a maximum and highly vari-
able depth of only 12 m. The salinity of the
water is about 33 %©, although the Salton Sea
is not of marine origin; it has been formed by
the inflow and evaporation of Colorado River
water. Relative to ocean water, sulfate (2.7
times more concentrated) and calcium (1.9)
ions are especially abundant, whereas potassium
(0.58) and magnesium (0.75) are appreciably
less concentrated. The annual maximum and
minimum surface water temperatures are usu-

ally about 36C and 10C, respectively. Great
daily fluctuation in temperature occurs, espe-
cially in shallow water (Barlow, 1958).

The specimens of mirabilis from the Salton
Sea differ from those on the outer coast in just
the ways one would predict for fish whose
growth had been drastically retarded. All fin ray
counts and the vertebral counts are high (but
the relationship between the fin counts is un-
changed ) , supernumerary spiny rays are formed,
scalation is incomplete, the head and its related
parts are small, the fins are low, and the com-
pletion of the interorbital canal is delayed.
Anomalies, such as a continuous slit for the
anterior and posterior nates, an abbreviated
postorbital canal, and flaps on the jaws, are
holdovers of conditions usually seen only in
postlarvae.

The unusual combination of salts in the water
of the Salton Sea probably interferes with the
development of the fish. The water temperatures
are not to be considered responsible, for they are
moderate during that period of early develop-
ment when the characters are determined. In-
deed, the temperatures are comparable to those
of habitats along the sea coasts. Other water
conditions (oxygen tension, pH, and illumina-
tion) also are similar to those encountered by
naturally occurring populations. The Salton Sea
fish at times are heavily infested with a mono-
genetic trematode, but so are coastal populations.
Besides, many of the characters are determined
before hatching and so could not be modified
by debilitation caused by parasites (Hubbs,
1927).

The possibility exists, of course, that the
mirabilis in the Salton Sea have diverged ge-
netically, and that the structural differences are
the result of such genetic change. The original
stock consisted of merely 500 fish, of which
only a fraction could have reproduced success-
fully. By chance alone, certain alleles must have
been lost, others fixed, even if all 500 fish are
assumed to have reproduced (Wright, 1951).
The population expanded rapidly to a large
number of fish, as did later introductions of
other species. In a large population, genetic
drift would no longer be operative (Wright,
1951). During the 25 years after introduction,
selection has been effected, for the most part,

through the rigorous environment and through
intraspecific competition.

If the gene pool of the Salton Sea fish has
been altered, then the morphological changes
might be due to such genetic differences. This
cannot be known without rearing experiments.
Experiments on the metabolic physiology of the
adults suggest slight genetic divergence has
taken place (Barlow, 1961c).

My opinion is that the morphological pecu-
liarities of the fish from the Salton Sea can be
ascribed primarily to direct modification caused
by the chemical composition of the water in
which they develop; the same could be said of
the San Felipe fish from a saline pool.

Conclusions

The investigation of the species Gillichthys
mirabilis has revealed a complex situation in
which genetic divergence is evidently inter-
twined with phenotypic plasticity.

Over the range of mirabilis, the adaptive
norms seem to shift gradually, but with some
plateaus. A comparison of fin indices and means
of the fin counts indicates the following ar-
rangement : ( 1 ) Populations of mirabilis are
nearly the same genetically on the California
coast, though the Salton Sea and San Francisco
populations may be somewhat distinct. (2) A
genetic cline may exist on the outer coast of
Baja California. (3) The fish from the lower
region of the Gulf of California are similar to,
but slightly different from, those on the outer
coast of southern Baja California. (4) The pop-
ulations found in the northern part of the Gulf
have diverged from those of the southern part
to a greater degree than can be accounted for,
in a consistent manner, by environmentally in-
duced changes.

Investigations of the body form of mirabilis
support the conclusions based on countable
characters. The body form of fish from Califor-
nia is about the same in each of the populations,
although local differences exist. Fish from the
Gulf of California, as a group, differ from those
from California; no adult specimens were avail-
able from the outer coast of southern Baja Cali-
fornia. Within the Gulf populations, fish in the
upper northern region of the Gulf are separable
from those in the southern region.

PACIFIC SCIENCE, Vol. XVII, January 1963

Thus mirabilis is seen as one widely ranging
species whose geographic subdivisions are con-
sistent with known faunal regions. For example,
the fish fauna of San Francisco Bay is isolated
from its southern counterpart, and the degree
of differentiation of the mirabilis population
there reflects this separation. The broad transi-
tion between faunas along the outer coast of
Baja California also seems to be an area of tran-
sition for mirabilis, though more samples are
needed from this region. Likewise, the morpho-
logical separation of the mirabilis of the Gulf !
from those of the outer coast is in keeping with
the known distribution of this species. Finally,
the division of mirabilis from the Gulf into
northern and southern groups is in harmony
with faunistic and oceanographic findings
(Hubbs, I960; Walker, I960; Roden, 1938).

Subspecific names are not proposed for the
geographic variants. Until more collections are
available it would only create confusion to name
the geographic forms as they are known at pres-
ent. Even if they were better known, I doubt
that subspecific names would be of utility here.
None of the evidence suggests that any group, j
or groups, of populations is evolving into a dis- 1
tinct species. The findings indicate only that in |
this widely distributed species appreciable dif-
ferences exist; these are thought to reflect adap-
tive changes.

SUMMARY

1. Gillichthys mirabilis Cooper is found in |
coastal sloughs from Central California south to '
near Cabo San Lucas, and then again in the cen- ;
tral and northern parts of the Gulf of California.

It has been introduced into the Salton Sea.

2. The fish spawn from about January to
June and reach maturity within 1 year. The lar-
vae have a brief pelagic phase, during which
time the species is distributed to new habitats.

3. The head is shortest in specimens of mira-
bilis from the Salton Sea and most depressed
in those from the Gulf of California. The length
of the upper jaw and the height of the median
fins are greatest in samples from the Gulf and
least in those from the Salton Sea. Specimens
from the upper Gulf region have the dorsal fins
separated more than in mirabilis from else-
where.

Gillichthys mirabilis — Barlow

4. The scales are larger and the ventral area
more scaled in specimens from the Gulf than
in those from the outer coast. Scalation is notice-
ably reduced in fish from the Salton Sea.

5. Trends in means of meristic characters,
from north to south along the Pacific Coast,
and then from south to north up into the Gulf
of California, are as follows: spiny dorsal fin —
no change or slight increase; segmented rays of
median fins — decrease; pectoral fin — increase;
combined fin elements — irregular or no change;
fin index (segmented dorsal plus anal rays, di-
vided by pectoral rays, times 100) — decrease;
pectoral fin asymmetry — -decrease; vertebrae —
no change.

6. The variance of the number of elements
in each fin is a function of latitude, greater vari-
ance being associated with higher latitudes.

7. The means of the counts of the median
fin are negatively correlated with the means for
pectoral fin rays when the comparison is be-
tween the various populations. In successive
year classes within a given population, however,
changes in the number of rays in the median
fins are associated with commensurate changes
of the same sign in the counts of the pectoral
fin rays. But within each year class there is no
correlation between the number of pectoral fin
and median fin rays.

8. All meristic characters have higher mean
values in specimens from the Salton Sea, but
the mean of their fin index accords with the
Pacific Coast population from which they were
derived. The Salton Sea fish are aberrant in
many ways, evidently as a result of living in a
marginal environment.

9. The conclusions based on these findings
are condensed on the last page of the foregoing
article.

REFERENCES

Baldwin, J. M. 1896. A new factor in evolu-
tion. Amer. Nat. 30: 441-475, 536-553.

Barlow, G. W. 1958. Daily movements of
desert pupfish, Cyprinodon macularius , in
shore pools of the Salton Sea, California.
Ecology 39: 580-587.

— - — \%la. Gobies of the genus Gillichthys,

with a comment on the sensory canals as a
taxonomic tool. Copeia 1961: 423-437.

1961 A Causes and significance of mor-
phological variation in fishes. Syst. Zool. 10:
105-117.

— 196lc. Intra- and interspecific differ-

ences in rate of oxygen consumption in go-
biid fishes of the genus Gillichthys. Biol. Bull.
121: 209-229.

Carpel AN, L. H. 1957. Hydrobiology of the
Alviso salt ponds. Ecology 38: 375-390.

1958. The Salton Sea. Physical and

chemical characteristics. Limnol. Oceanog. 3:
373-386.

Garth, J. S. I960. The biogeography of Baja
California and adjacent seas, Part II. Marine
biotas. Distribution and affinities of the
brachyuran Crustacea. Syst. Zool. 9: 105-123.

Gilbert, C. H., and N. B. Scofield. 1898.
Notes on a collection of fishes from the Colo-
rado Basin in Arizona. Proc. U. S. Nat. Mus.

(20): 487-499.

Hoel, P. G. 1954. Introduction to Mathematical
Statistics. 2nd ed. Wiley & Sons, New York.
331 pp.

Hubbs, C. L. 1921. Geographic variation of
Notemigonus crysoleucas — an American min-
now. Trans. 111. Acad. Sci. 11: 147-151.

1922. Variations in the number of ver-
tebrae and other meristic characters of fishes
correlated with temperature of water during
development. Amer. Nat. 56: 360-372.

1924. Studies on the fishes of the order

Cyprinodontes, IV. The subspecies of Pseu-
doxiphophorus bimaculatus and Priapichthys
annectens. Misc. Publ. Mus. Zool. Univ. Mich.
(13): 17-23.

1926. The structural consequences of

modifications of the developmental rate in
fishes, considered in reference to certain prob-
lems of evolution. Amer. Nat. 60: 57-81.

PACIFIC SCIENCE, Vol. XVII, January 1963

1927. The related effects of a parasite

on a fish. A retardation of early growth, the
retention of larval characters and an increase
in the number of scales. J. Parasit. 14: 73-84.

1928. An hypothesis on the origin of

graded series of local races in fishes. Anat.
Rec. 41: 49- (Abstract.)

— 1948. Changes in the fish fauna of west-
ern North America correlated with changes
in ocean temperature. Sears Found. J. Mar.
Res. 7: 459-482.

I960. The biogeography of Baja Cali-
fornia and adjacent seas, Part II. Marine bio-
tas. The marine vertebrates of the outer
coast. Syst. Zool. 9: 134-147.

and C. Hu BBS. 1953. An improved

graphical analysis and comparison of series
of samples. Syst. Zool. 2: 49-57.

— and L. C. Hubbs. 1945. Bilateral asym-

metry and bilateral variation in fishes. Papers
Mich. Acad. Sci. Arts and Letters 30 (1944):

229-310.

and K. F. LAGLER. 1949. Fishes of the

Great Lakes Region. Cranbrook Press, Bloom-
field, Mich. 186 pp.

Johnsen, S. 1936. On the variation of fishes
in relation to environment. (Preliminary ac-
count.) Bergens Mus. Aarb. (4): 1-26.

MAcGlNlTlE, G. E. 1939. The natural history
of the blind goby, Typhlogobius calif ornien-
sis Steindachner. Amer. Midi. Nat. 21: 489-
505.

Martin, W. R. 1949. The mechanics of en-
vironmental control of body form in fishes.
Univ. Toronto Stud. Biol. Ser. (58), Publ.
Ontario Fish. Res. Lab. (70) : 1-91.

Roden, G. I. 1958. Oceanographic and meteor-
ological aspects of the Gulf of California.
Pacif. Sci. 13: 21-45.

and G. W. Groves. 1959- Recent

oceanographic investigations in the Gulf of
California. Sears Found. J. Mar. Res. 18: 10-
35.

Schmidt, J. 1919. Racial studies in fishes, II.
Experimental investigations with Lebistes
reticulatus (Peters) Regan. J. Genet. 8: 147—
153.

Schultz, L. P. 1927. Temperature controlled
variation in the golden shiner, Notemigonus
crysoleucas. Papers Mich. Acad. Sci. 7: 417—
432.

Seymour, A. H. 1956. Effects of temperature
upon young chinook salmon. Dissertation
Abstr. 16: 2249.

Starks, E. C., and E. L. Morris. 1907. The
marine fishes of southern California. Univ.
Calif. Publ. Zool. 3: 159-251.

TAning, A. V. 1952. Experimental study of
meristic characters in fishes. Biol. Rev. 27:
169-193.

TA VOLGA, W. N. 1954. Reproductive behavior
in the gobiid fish Bathygobius sop orator. Bull.
Amer. Mus. Nat. Hist. 104: 431-459.

Vladykov, V. D. 1934. Environmental and
taxonomic characters of fishes. Trans. Roy.
Canad. Inst. 20: 99-140.

Walker, B. W. I960. The biogeography of
Baja California and adjacent seas, Part II.
Marine biotas. The distribution and affinities
of the marine fish fauna of the Gulf of Cali-
fornia. Syst. Zool. 9: 123-133.

Weisel, G. F. 1947. Breeding behavior and
early development of the mudsucker, a gobiid
fish of California. Copeia 1947: 77-85.

1948. Relation of salinity to the activity

of the spermatoza of Gillichthys, a marine
teleost. Physiol. Zool. 21: 40-48.

Wright, S. 1951. The genetical structure of
populations. Ann. Eugen. 15: 323-354.

Preliminary Notes on Molluscan Assemblages of the Submarine Banks

Around the Izu Islands

Takashi Okutani1

It is well known that there are several sub-
marine banks along the submerged rise which
extends southwestward from the southern tip
of Izu Peninsula, central Honshu. Small islands
such as Toshima, Niijima, Shikine, and Kozu,
with several other islets, lie on this rise. These,
together with a few other islands situated far-
ther south, are called the Izu Islands. They are
linked by a volcanic system, and there is a con-
siderable number of such banks in the neighbor-
hood. A few papers concerned with hydrograph-
ical, bathymetrical, and faunistic characteristics
of these submarine banks have been prepared by
Suzuki and Sato (1944), Niino (1935, 1952,
1955), and Shirai (1958). On the basis of these
works, together with information furnished by
the present author, Horikoshi (1957) discussed
the topographical peculiarity in relation to the
general molluscan fauna on these banks.

Another group of submarine banks is found
around the O'sumi Islands, south of Kyushu.
Presumably their hydrographical and bathymet-
rical characters are similar to those banks men-
tioned above, but no information about the
molluscan fauna has been available until now.

As a contribution to knowledge about mol-
luscan fauna on the submarine banks and in-
sular shelves around the Izu Islands, this paper
deals with the general account of the molluscan
assemblages of the area and their faunal simi-
larity to another series of banks near the Osumi
Islands in the Kuroshio area. It is based on
biological dredge samples collected chiefly by
research vessels during 1955-59.

The present writer wishes to express his grati-
tude to Dr. Z. Nakai, Tokai Regional Fisheries
Research Laboratory, for the material. Thanks
are extended to Mr. Y. Kurata, Tokyo Fisheries

1 Tokai Regional Fisheries Research Laboratory,
Tokyo, Japan. Manuscript received September 11,
1961.

Experimental Station; Dr. M. Horikoshi, Ocha-
nomizu University; Dr. K. Sakurai and Mr. A.
Teramachi, members of the Japan Malacologi-
cal Society, for their facilities and advice ren-
dered in the course of this study. Thanks are also
due the crews of the research vessels for their
cooperation in collecting the present materiai.

TOPOGRAPHY OF SUBMARINE BANKS AROUND
IZU ISLANDS

The Izu Isiands extend from the mouth of
Sagami Bay to the south. In the northern group
are (Izu-)Oshima, Toshima, Niijima, Shikine,
and Kozu. To the south there are Miyake, Mi-
kura, Hachijo, Aogashima, and Torishima as
the southern extremity (Figs. 1, 2). The banks
are usually isolated from the series of these is-
lands, with depressions deeper than 200 m lying
between them. The tops of such types of banks
are usually flat and about 80-120 m in depth.
They are usually elongate-oval in shape with
the axis in a northeast-southwest direction.

The Hyotanse Bank, one of the representa-
tives of this series of banks, located west of
Kozu Island, has been described by Niino
(1955) as follows: The slopes around the bank
are steep and rocks are exposed there; gravels
and coarse material cover the broad and flat
plain on its top; andesite and basalt, which are
very common in the bedrock, are found mingled
with liparite gravels together with a number of
manganese concretions from the bank; the litho-
logical characters of these rocks are the same
as those of the main islands in the Fuji Vol-
canic Zone. According to gross observation of
the present material, the sediments (gravels
and shells) are heavily coated by calcareous
algae.

The bottom of Zenisu Bank, also studied by
Niino (1935), reveals coarse sand and shells,

PACIFIC SCIENCE, Vol, XVII, January 1963

36°

34°

32°

30°

28°

Fig. 1. Index map for the position of submarine banks and other localities, a, Boso Peninsula; b, Izu Penin- j
sula; c, Kii Peninsula. A, (Izu-) Oshima Is.; B, Hachijo Is.; C, Torishima Is.; D, Goto Islands; E, Amami-
Oshima Is.; F, Kikaijima Is.; G, Okinoshima Is. in the Bungo Straits, a, Omurodashi (bank); P, Kurose
(bank); 7, Shinkurose (bank). (See Figs. 2 and 3 for details of areas I and II, respectively.)

and several steep rocks are exposed above the
sea surface. At Watarinose Bank, which lies
between Kozu Island and Zenisu, the character
of the bottom is supposedly generally the same
as that at Zenisu. The neighboring waters of
those banks are noted as excellent fishing
grounds. Kurose and Shinkurose are the south-
ern banks situated around Hachijo Island. The
bottom characters of these two are known to
be similar to that of Hyotanse. On the other
hand, Omurodashi Bank off Izu-Oshima is said
to be different from the others in having a
sandy mud sediment at its top.

The geographical positions of the submarine
banks of Izu Islands under study are referred
to in Figures 1 and 2.

OCCURRENCE OF SPECIES BY AREA
1. Hyotanse Bank

material: Dredged by the R.V. "Soyo-
maru” on Nov. 20 and 23, 1935, at 7 stations
from depths of 118, 135, 140, 145, 148, 153,
and 170-230 m.

EARLIER WORKS: Niino (1955) reported 11

Molluscan Assemblages — Okutani

species of Pelecypoda and 5 species of Gastro-
poda. Shirai (1958) reported 5 pelecypods and
5 gastropods.

SPECIES IN THE PRESENT MATERIAL: A car
congenitum ( Smith ) ; Mimar carta aizoi Sakurai
(ms), 137 m; Striarca fausta Habe, 137 m;
Samacar pacifica (Nomura and Zimbo), 137 m;
Barbatia tamikoae Sakurai (ms), 140 m; Pseu-
do grammatodon obliquatus Yokoyama, var.;
Nipponolimopsis decussata (A. Ad.), 145 m;
Tucetona shinkurosensis Hatai, Niino and Ko-
taka; Malleus irregularis ( Jousseaume) , var.,
118 m; Chlamys mollita (Rve. ), 118 m; C. lem-
niscata (Rve.), 118 m; C. vesiculosus ( Dkr. ) ;
C. tissotii (Bernard), 153 m; Spondylus anacan-
thus (Mawe); Limatula japonica (A. Ad.);
Lima fujitai Oyama; Septifer gray ana (Dkr.);
Cardita nodulosa (Lamarck), 153 m; Gians sa-
gamiensis Kuroda and Habe; Chama argentata
Kuroda and Habe; Erigidocardium eos (Ku-
roda), 140 m; Meiocardia tetragona (Ad. and
Rve.); Emarginula fragilis Yokoyama, 153 m;
E. teramachii Habe; Microgaza sp. aff. sericata
Kira, 153 m; Talopena lifuana (Fischer), 153 m;
Galeoastraea guttata (A. Ad.), 153 m; Ten-
agodus anguinus (L.), 140 m; Serpulorbis me-
dusae Pilsbry, 140 m; Apollon hirasei Kuroda
and Habe, 145 m; Phanozesta semitort a Kuroda
and Habe, 135 m; Latiaxis pagodus (A. Ad.),
137 m; Bursa ranelloides (Rve.), var., 135 m;
Mitrella sp. cf. lischkei (Smith); Conus gra-
tacapy Pilsbry, 170 m; Conus sp., 153 m.

OTHER SPECIES REPORTED BY NIINO: Euce-
tona hanzawad (Nomura and Zimbo); Limop-
sis tajimae (Yokoyama); Hawaiarca uwaensis
(Yokoyama); Plicatula muricata (Sowerby);
T richomus cuius coralliophaga (Gmelin); Lima
basilanica (Ad. and Rve.), 260 m; Ctenoides
annulata (Lamarck), 260 m; Pecten albicans
(Schroter); Lucinoma spectabilis (Yokoyama);
Perotrochus beyrichii ( Hilgendorf ) , 134, 160,
128 m.

MOLLUSCS REPORTED ONLY BY SHIRAI: Li-
mopsis obliqua A. Ad., 250 m; Septifer excisus
(Wiegmann), 104 m (this may be S. gray ana ) ;
Eragum loochooanum Kira, 250 m (this may be
Gians sagamiensis ) ; Galeoastraea millegranosa
Habe, 260 m (this may be G. guttata) ; Sil-
iquaria cumin gii Morch, 260 m (this may be
Tenagodus anguinus ).

2. Zenisu Bank

MATERIAL: Dredged by the R.V. "Soyo-
maru” on Nov. 20, 1955, at 2 stations from
depths of 85 and 170 m.

EARLIER WORKS: Niino (1935) reported on
the bottom character only.

SPECIES IN THE THE PRESENT MATERIAL:
Acar congenitum (Smith) ; Hawaiarca uwaensis
(Yokoyama), 170 m; Pseudo grammatodon ob-
liquatus (Yokoyama); Pectunculina cernata (A.
Ad.); Limopsis cumingii A. Ad.; E ucetona
shinkurosensis Hatai, Niino and Kotaka, 170
m; Glycymeris rotunda (Dkr.); Polynemamus-
sium intuscostatum (Yokoyama); Chlamys ve-
siculosus Dkr.; C. tissotii (Bernardi); C. lem-
niscata ( Rve. ) ; Plicatula muricata ( Sowerby ) ;
Spondylus anacanthus (Mawe); Pecten albicans
(Schroter); Lima zushiensis (Yokoyama); Vol-
sella sp.; Pycnodonta musashiana (Yokoyama);
Meiocardia tetragona (Ad. and Rve.); Frigido-
cardium eos (Kuroda); Poromya flexuosa (Yo-

34° 30'

34° 20

34° 10

34° 00’

33° 50'

Fig. 2. Detail for I in Fig. 1. Dots indicate the
biological stations for the R.V. "Soyo-maru” in Nov.,
1955.

koyama); V erticordia japonica A. Ad.; Emargi-
nula incisura A. Ad.; Bursa ranelloides ( Rve. ) ,
var.; Distorsio sp.

3. W at arinose Bank

material: Dredged by the R.Y. "Soyo-maru”
on Nov. 20, 1955, at 2 stations from depths of
80-100 m and 140-220 m.

earlier works: None.

SPECIES IN THE PRESENT MATERIAL: A car
congenitum (Smith); Barbatia tamikoae Saha-
ra! (ms); Saniacar pacifica (Nomura and
Zimbo); Tucetona shinkurosensis Hatai, Niino
and Kotaka; Malleus irregularis ( Jousseaume) ,
var.; Chlamys mo Hit a (Rve.); C. lemniscata
(Rve.); Lima fujitai Oyama; Astraea okamotoi
Kuroda and Habe; Galeoastraea guttata (A,
Ad.) ; Casmaria cervica (Sowerby) ?; Tenagodus
anguinus ( L. ) ; Clavus sp. aff. rufovaricosa Ku-
roda (MS).

4 . Toshimatai Bank and Niijima Niskitai Bank

(T oramaguri)

MATERIAL: None is available for the present
study except Gians sagamiensis Kuroda and
Habe and Conus kinoskitai Kuroda from Nii-
jima-Nishitai.

earlier works: Niino (1955) reported 19
pelecypods and 4 gastropods from Toramaguri
and a single gastropod from Toshimatai. Shirai
(1958) reported 9 bivalves and 4 univalves
from Niijima-Nishitai Bank.

SPECIES REPORTED BY NIINO AND SHIRAI:
Toshimatai Bank— Tucetona shinkurosensis Ha-
tai, Niino and Kotaka; Hiatella arctica orientalis
Yokoyama; Perotrochus beyrichii (Hilgendorf),
88 m.

Niijima-Nishitai Bank— -Bar batia plicata
(Dillwyn) (probably A. congenitum ), 120 m;
Tucetona shinkurosensis Hatai, Niino and Ko-
taka, 104 rn; Chlamys pelseneeri Dautzenberg
and Bavay, 120 m; C. vesiculosus Dkr., 78 m;
Lima zushiensis Yokoyama, 73 m; Lima fujitai
Oyama, 104 m; Limatula japonica A. Ad., 104
rn; Cr as satellites oblongatus (Yokoyama), 120
m; Venus toreuma Gould, 78 m; Aloides venusta
(Gould), 120 m; Perotrochus beyrichii (Hil-
gendorf), 104, 120 m; Emarginula fragilis Yo-

PAGIFIC SCIENCE, Vol. XVII, January 1963

koyama, 104 m; E. sp., 104 m; Punctmella fasti -
giata A. Ad., 78 m; Trivirostra oryza (Lamarck),
88 m.

X Ombase Islet ( near Kozu Island)

MATERIAL: A part of the specimens dredged
by the R.V. "Tonan-maru” at depths of 32, 55,
60, and 100 m. These were examined by cour-
tesy of Mr. Kurata.

EARLIER WORKS: Igarashi and others (1956) ■
reported on the bottom character and upon pis-
cifauna from the viewpoint of fishing ground
investigation.

SPECIES IN THE PRESENT MATERIAL: SdMacar
pacifica (Nomura and Zimbo), 60" m; Barbatia
sp.; Tucetona shinkurosensis Hatai, Niino and
Kotaka, 32 m; Glycy metis amamiensis Kuroda,
32 m; Venus toreuma Gould; Galeoastraea gut-
tata (A. Ad.), 55, 100 m.

6. Kurose and Shinkurose

MATERIAL: Collected from Kurose Bank at a
depth of ca. 200 m by Cor allium fishing net of
the R.V. "Tonan-maru.” Examined by courtesy
of Mr. Kurata.

EARLIER WORKS: Niino (1952) surveyed
around Shinkurose Bank and reported 13 gas-
tropods, 20- pelecypods including 3 new forms,
and 1 scaphopod.

SPECIES IN THE PRESENT MATERIAL: KufOSC
Bank- - -A car congenitum (Smith); Area mania
takii Hatai, Niino and Kotaka; Barbatia sp.;
Samacar pacifica (Nomura and Zimbo); Lima
fujitai Oyama; L. quanto ensis Yokoyama; Cras-
satellites oblongata Yokoyama; Perotrochus bey-
richii (Hilgendorf); Emarginula sp.; Tenagodus
anguinus ( L. ) ; Bursa ranelloides (Rve.) var.;
Chicoreus laciniatus (Sowerby); Conus sp.

Shinkurose (Niino)— Area sp., 290 m; Nip-
ponolimopsis nipponica (Yokoyama), 290 m;
Tucetona hanzawai (Nomura and Zimbo), 280
m; Tucetona shinkurosensis Hatai, Niino and
Kotaka, 290 m; Venericardia ryukyuensis No-
mura and Zimbo, 290 m; V asticardium sp., 280
m; Chione chlorotica , 280 m; Cadulus sp., 290
m; Collisella heroldi (Dkr.), 290 m (a littoral
species, may be carried down to the deep by some
means); Margarites cincereus (Couthouy), 290

Molluscan Assemblages — OKUTANI

m; Pseudoliotia micans (A. Ad.), 290 m; Clath-
ofenella reticulata (A. Ad.), 290 m; Mucron-
alia subulata (A. Ad.), 290 m; Tonna luteo-
stoma Kiister, 280 rn; Coralliophaga euginiae
(Bernard), 280 m; Bursa bufonia (Gmelin),
280 m; Conus sp. (identified as C. tone by an
illustration in Niino’s paper).

7. Insular Shelf around Hachijo Island

MATERIAL: None is available here.

EARLIER WORKS: Niino (1952) reported 13
pelecypods and 3 gastropods from depths of
115 and 200 m.

SPECIES REPORTED BY NIINO: Barbatia ha-
chijojimensis Hatai, Niino and Kotaka, 115, 200
m; Area mauia takii Hatai, Niino and Kotaka,
200 m; Tucetona sp., 200 m; Spondylus cruen-
tus Lischke (S. anacanthusl ) , 115 m; Spondylus
sp.; Lima lima L. 115, 200 m; Pycnodonta mu-
sashiana (Yokoyama), 200 m; Pseudochama sp.,
200 m; V asticardium arenicolum ( Rve. ) ; Mere-
trix sp., 115 m; Callista pilsbryi Habe, 200
m; Venus sp., 200 m; Phalium sp., 115 m;
Ocenebra adunca (Sowerby), 115 m; Conus
sp., 115 m.

8. Insular Shelf around Torishima Island

material: Part of specimens collected by
test fishing for Cor allium operated at a depth
of 150-250 m. The material was examined by
courtesy of Mr. Kurata.

EARLIER WORKS: None.

SPECIES IN THE present MATERIAL: Spon-
dylus anacanthus (Mawe); Pycnodonta musa-
shiana (Yokoyama); Plicatula muricata (Sow-
erby); Notolimea sp. cf. tosana Oyama; Chama
argent ata Kuroda and Habe; Tenagodus an-
guinus (L.) ; Talopena lifuana (Pilsbry); Can-
tharus sp.

SIMILAR MOLLUSCAN FAUNA FROM SOME
BANKS SOUTHWEST OF KYUSHU

On the Pacific Ocean side of Japan, groups
of submarine banks other than those mentioned
above are scattered throughout the southwest-
ern waters off Kyushu (Figs. 1, 3). These also
are situated on the submarine rise of the vol-

Fig. 3. Detail for II in Fig. 1. Dots indicate the
biological stations for the R.V. "Soyo-maru” in Feb.,
1959.

canic system on which the Osumi Group lies;
this situation is quite similar to that of the Izu
Banks. Hereafter, this second series of banks is
tentatively called the Osumi Banks. The tops of
these banks are usually at about 100 m, and
they are isolated from the neighboring islands
by depressions of about 200 m in depth. Their
general features are similar to the Izu Banks
in that they have submarine sediments of coarse
sand and gravel which are coated by calcareous
algae. Some rocks exposed at the sea surface are
also found (e.g., the Uji Islets).

Oceanographically, the two series of banks
are decidedly exposed to similar conditions with
respect to the Kuroshio Current, for the usual
main axis of the current passes through the
Osumi Straits as well as across the Izu Sub-
marine Ridge.

The occurrence of molluscan species revealed
by the present survey is as follows:

9. Off Takeshima Island

At a depth of 210 m; dredged by the R.V.
"Soyo-maru” on Mar. 5, 1956; sta. 114: 30° 50'
N, 130° 28' E.

Area mauia takii Hatai, Niino and Kotaka;

Glycymeris amamiensis Kuroda; Chlamys mol-
lita ( Rve. ) ; C. vesiculosus ( Dkr. ) , var.; C.
tissotii (Bernardi); Lima tomlini Prashad; Cte-
noides japonicus (Dkr.); Plicatula muricata
Sowerby; Septifer gray anus (Dkr.); Gians kyu-
shuensis, n. sp.; Chama argentata Kuroda and
Habe; Frigidocardium eos (Kuroda); Vasti-
cardium sp.; Venus toreuma (Gould); Pitar
sp.; Bursa sp.; Galeoastraea millegranosa Habe;
G. tayloriana (Smith); C er at o stoma vespertilis
Kira; Chicoreus laciniatus (Sowerby)?; Polyn-
ices sp.; Conus sp.

FIG. 4. Distribution of Perotrochus heyrichii (Hil-
gendorf). (See section 12 for numerals on dots.)

PACIFIC SCIENCE, Vol. XVII, January 1963

10. Uji Islets

At 3 stations from depths of 126-140 m;
dredged by the R.V. "Soyo-maru” on Feb. 8,
1959.

A car congenitum (Smith), 126 m; Striarca
fausta Habe, 126 m; Samacar pacifica (Nomura
and Zimbo), 126 m; Mimarcaria aizoi Sakurai
(MS), 126 m; Hawaiarca uwaensis (Yoko-
yama), 126 m; Spondylus anacanthus (Mawe),

126 m; Chlamys lemniscata (Rve.), 140 m;
Malleus irregularis ( Jousseaume) , 126 m;
Chama argentata Kuroda and Habe, 126 m;
Atrina penna Habe, 140 m; Penicillus giganteus
(Sowerby), 126 m; Perotrochus salmiana I
(Rolle), 126 m; Serpulorhis medusae Pilsbry,

126 m; Emarginula sp.; Malluvium otohimeae
(Habe), 140 m; Bursa ranelloides (Rve.), var.,
126, 140 m; Semicassis sp., 140 m.

11. Insular Shelf around Goto Islands

Sakurai (1959, and personal communication)
reported the following species which have been
collected by Cor allium fishing nets operated off
the Goto Islands; depths of operation may be
about 100-200 m.

Mimarcaria aizoi Sakurai ( ms ) ; Barbatia
tamikoae Sakurai (MS); A car congenitum
(Smith); Striarca fausta Habe; S. soyoae Habe;
Chlamys lemniscata ( Rve. ) ; Dymia argentata
Habe; Samacar pacifica (Nomura and Zimbo);
Chama argentata Kuroda and Habe; Perotro-
chus hirasei Pilsbry.

TYPICAL SPECIES OR SPECIES-GROUPS FOR
BANKS-ASSOCIATED MOLLUSCA

Because the present data are not based on
quantitative samplings, it is not possible to dis-
cuss the matter from a quantitative point of
view. However, several species-groups may be
indicated as endemic ones (or semiendemic) for
submarine banks or insular shelves, because of
their frequency or abundance in occurrence.

12. Perotrochus beyrichii (Hilgendorf)

Fig. 5

It is well known that this "living fossil” occurs
on the lower shelf around Sagami Bay. The

Molluscan Assemblages — -Okutani

FIG. 5. Perotrochus beyrichii (Hilgendorf) from
Kurose, ca. 200 m, 72.7 mm X 70.0 mm (Mr. Kurata
coll.) .

known localities of the species are as follows
(numbers for records refer to those of localities
shown in Fig. 4) :

1) Off Iioka, Pacific side of Boso Peninsula;
dead shells collected by commercial trawl-
ers ( Sakurai, 1954).

2) Off Hachiman-Saki ("Soyo” sta. 40a; July
4, 1956; 35° 04.7' N, 140° 21.5' E), 250
m in depth; a fragment only.

3) West coast of Boso Peninsula, off Katsu-
yama and Tomiura, about 200-300 m
in depth (Wada, 1954).

4) Sagami Bay, west coast of Miura Penin-
sula. Type locality is off Misaki. Also col-
lected by the H. M. "The Emperor” near
Hayama.

5 ) Okinoyama, a small bank located near the
southwestern tip of Boso Peninsula.

6) Senzu, northwestern coast of Oshima Is-
land; dead shell with hermit crab, col-
lected by Mr. Kurata.

7) Around Omurodashi Bank (Shirai, 1958).

8) Hyotanse, 128, 134, 160 m (Niino, 1955).

9) Niijima-Nishitai, 104, 120 m; and Toshi-
matai, 88 m (Niino, 1955, and Shirai,
1958).

10) Kurose, collected by Mr. Kurata.

As is shown above, this pleurotomariid gas-
tropod is characteristically distributed on the
lower shelf and submarine banks around the
area. Other species of Perotrochus, as ecological
equivalents, are also found in similar habitats in
other districts; Perotrochus hirasei Pilsbry is
known to be distributed around the Pacific coast
of Kii Peninsula and Tosa Bay, and as an in-
habitant of the continental shelf around the
main Japanese islands. According to Mr. Tera-
machi ( personal communication ) , P. hirasei has
been collected by him from depths of 60 fath-
oms (ranging 20-180 fathoms) in such locali-
ties.

1 ) Okezoko Deep located south of Okino-
shima Islet in Bungo Straits. This is col-
lected from thanatocoenose, mainly of
Glycymeris rotunda, with Siphonalia fil-
osa, etc.

2) Off Urado in Tosa Bay, 70-80 fathoms,
sometimes with Erronea hirasei, Chicoreus
dilectus, and some other common shelf
dwellers.

3) Off Tanabe, southwestern coast of Kii
Peninsula.

4) Somewhere in Hyuga-Nada, eastern
waters off Kyushu.

Recently, Kuroda and Habe (1953) have
shown that this species is also distributed in the

Fig. 6. Perotrochus salmiana (Rolle) from Uji
Islet, 126 m, 65.0 mm X 62.7 mm.

PACIFIC SCIENCE, Vol. XVII, January 1963

36°

34°

Fig. 7. Distribution of three species of Japanese Perotrochus. Dots, P. beyrichii; horizontal lines, P. hirasei;
cross lines, P. salmi ana.

western waters off Kyushu and around the Goto
Islands, where some banks-associated assem-
blages are found (Sakurai, 1959, and cf 11).
Perotrochus salmiana (Rolle) (Fig. 6) is also
known from waters off Kii and Tosa provinces.
In the present survey, a new locality for this rare
species was discovered from the Uji Islets
CSoyo” sta. 72a, Feb. 8, 1959, 31° 24.7' N,
129° 37.6' E, 126 m), in company with the
similar banks-associated fauna (cf 10). The
smallness of the specimen in the present ma-
terial may coincide with the fact pointed out
by Parker and Curray (1956: 2436), . .

most of these bank forms were considerably
smaller, although they appeared to be mature
specimens. . . .”

13. Galeoastraea guttata (A. Adams)

Fig. 8

According to Habe (1958), this fascinating
species is distributed around Boso Peninsula
and Sagami Bay. As is shown in the map (Fig.
20 a), this species is characteristically distrib-
uted on the banks around the Izu Islands, i. e.,
Hyotanse, Ombase Islet, and Okinoyama. It is
very rare from the insular shelf, despite the
fact that there is an example from the Izu Penin-

FlG. 8. Galeoastraea guttata (A. Adams) from
Hyotanse, 153 m, 22.2 mm X 30.3 mm.

Molluscan Assemblages — Okutani

Fig. 9. Bursa ranelloides (Reeve) var. from Uji
Islet, 140 m, 48.6 mm X 30.3 mm.

sula coast off Iro-Zaki ("Soyo” sta. 42, Apr. 2,
1958, 34° 34.2' N, 138° 50.7' E, 80 m). No
record from Boso Peninsula is available for the
present study. A possible ecological equivalent,

G. millegranosa Habe, is distributed on the
Osumi Banks as well as in its adjacent waters,
where G. guttata is not found. It is also inter-
esting to note that one of the related species,
Astraea caelata Gmelin, is indicated by Parker
and Curray ( 1956) as a typical banks-associated
mollusca from calcareous banks on the conti-
nental shelf off Texas, U.S.A.

1 4. Bursa ranelloides ( Reeve ) var.

Fig. 9

A rather small form of the species is widely
distributed on the banks mentioned above. It
has been collected from Hyotanse, 135 m;
Zenisu; Kurose, 200 m; off Takeshima, 210 m,
and the Uji Islets, 126 m.

15. Small Arcid Pelecypods

Figs. 10, 11

On the submarine banks under study, some
small species of Arcidae are abundant and are
found together with other members of the
group. The occurrence of species in the present
material is as follows:

IZU BANKS

OSUMI BANKS (KYUSHU)

Hyotanse

Zenisu

Kurose

Goto

Uji

Sta. 114

Acar congenitum (Smith)

—

Area mania takii Hatai, Niino, Kotaka....

—

Mimarcaria aizoi Sakurai (MS)

—

Hawaiarca uwaensis ( Y okoyama )

—

Samacar pacifica (Nomura, Zimbo)

— ?

—

Striarca fansta Habe

—

■ —

16. Small Pectinid Pelecypods
Figs. 12-14

Chlamys vesiculosus (Dkr.) is known to be
very abundant on the continental shelf border-
ing southwestern Japan. Being a sandy bottom
dweller, it is also distributed on the banks of
the northern group at a depth of 32-118 m. A
related species, C. tissotii ( Bernardi ) , often
occurs in the same localities. They are not found

Fig. 10. Area mania takii Hatai, Niino, and Ko-
taka from Kurose ca. 200 m, 28.0 mm X 15.0 mm
(Mr. Kurata coll.).

PACIFIC SCIENCE, Vol XVII, January 1963

on the southern half of Izu Banks but are dis-
tributed on the Osumi Banks. C. lemniscata
(Rve.) is similar in its distribution to the pre-

ceding two species; it occurs rather oftener
than C. mollita (Rve.) does on the banks un-
der study.

IZU BANKS

OSUMI BANKS

Hyotanse

Zenisu

Kurose

Uji

Sta. 114

Chlamys vesiculosus (Dkr.)

—

C. tissotii ( Bernardi )

—

C. lemniscata ( Rve. )

—

C. mollita ( Rve. )

—

— -

17. Tucetona hanzawai (Nomura and Zimbo);
T. shinkurosensis Hatai, Niino, Kotaka
Figs. 15, 16

These small glycymerid species occur on the
banks under study. As the former, T. hanzawai,
originally described from a fossil bed of Ryukyu
( Riu Kiu ) Limestone, was recorded from Hyo-
tanse and Shinkurose (Niino, 1952, 1955).
Hatai, Niino, and Kotaka (in Niino, 1952:
106) stated that "the occurrence of this species

Fig. 11. Samacar pacifica (Nomura and Zimbo)
from off Miyake Island, 170 m, 12.5 mm X 6.5 mm.

FIG. 12. Chlamys tissotii (Bernardi) from Hyo-
tanse, 140 m, 12.6 mm X 13.4 mm.

in the marine fauna of the environs of Hachijo I
Island is interesting, because it may suggest j
the occurrence of a submerged (geological)
formation corresponding in age to the Ryukyu
Limestone.” The latter species has been found
from Shinkurose (the type locality), Hyotanse, J
Zenisu, Watarinose, Niijima-Nishitai, Toshi-
matai, as well as Ombase Islet (Fig. 20 b) .

According to Hatai, Niino, and Kotaka (in
Niino, 1952), T. shinkurosensis (loc. cit. p. 109,
figs. 11, 12) is distinguished from T. hanzawai
(Nomura and Zimbo, 1934, p. 152(44), pi.
5(1), figs. 3a, b: as Glycymeris ) by the follow-
ing points: The shell of T. shinkurosensis is
longer than high, while that of T. hanzawai is
higher than long; the radial ribs of the former
are broad, flat, low, and are 27 in number, while

Molluscan Assemblages— OYJJTANl

there are 25 rounded ribs in the latter species;
the interspaces of shinkurosensis ribs are much
narrower than the ribs, while the interspaces of
the latter species are nearly equal to the ribs;
the hinge teeth of the former are 5 in the an-
terior half and 6 in the posterior half, instead
of 8 and 9 on the anterior and posterior half
of the hinge plate of the latter species.

In the present survey, 40 odd valves from
Izu Banks were examined. The size ranges of
the material were 7.7-17.4 mm in length and

7.6-18.1 mm in height. Of these specimens, no
valve was found which had rib interspaces as
wide as the ribs themselves, despite the fact
that meristic characters vary with individuals,
i. e., height/length, ranges 0.915-1.090; num-
ber of radial ribs, 24-33; anterior hinge teeth,
5-10; posterior teeth, 7-11. On these evi-
dences, all of the specimens are identified as
T. shinkurosensis.

The measurements of two odd valves of topo-

types of T. hanzawai are:

HEIGHT

HEIGHT/ LENGTH

RADIAL RIBS

TEETH

Ant. Post.

No. 1 (left valve)

11.1 mm

1.057

No. 2 (right valve)

12.0 mm

1.190

The interspaces of radial ribs are estimated
to be about half as wide as the ribs. In compari-
son with specimens of two species of similar
size, T. hanzawai is provided with more distin-
guishing features than are pointed out in the
foregoing lines: it has a more prominent umbo,
a deeper shell, and less angular shoulders, thus
a V asicardium- like appearance, partly because
of its shell which is longer than it is high. Nev-
ertheless, the morphological similarity of these
two requires further biometrical study in the
future.

18. Chama argentata Kuroda and Habe
Fig. 17

This is found on many banks and insular
shelves under study. It was collected from the
Niijima-Nishitai Banks, the insular shelf of
Torishima, the Uji Islets, insular shelves of
Takeshima (Sta. 114), and the Goto Islands
(Fig. 20c).

Other species than those enumerated above,
Spondylus anacanthus (Mawe), Plicatula muri-
cata ( Sowerby ) , and Lima fujitai Oyama ( Figs.
18, 19, and 20d), are usually found abundantly
in the area. The occurrence of two or three of
them may be a remarkable faunal characteristic
of the banks.

GEOGRAPHICAL DIFFERENCES AND VERTICAL
LIMITS OF TYPICAL BANKS-ASSOCIATED
ASSEMBLAGES

Since the present material was obtained with
different kinds of gear from a limited number
of stations, a conclusive quantitative analysis
can not be made at present. However, the abun-
dance or probable dominancy of the several

Fig. 13. Chlamys lemniscata (Reeve) from Uji
Islet, 126 m, 22.0 mm X 20.0 mm.

PACIFIC SCIENCE, Vol. XVII, January 1963

species might be estimated. If in the future
these areas are more thoroughly surveyed by
quantitative methods, the faunistic differences
or peculiarities will be more clearly demon-
strated. For the same reason, present data are
presumed to be insufficient for the following
discussion and a supplemental report is to be
expected.

Geographical Differences of the Assemblages

Within the Izu Banks, local differences in the
assemblage are scarcely observed. The typical
species-groups found in the area are roughly
subtropical or warm-temperate forms from the
lower part of the continental shelf bordering
the main Japanese islands. On the other hand,
probably because of the differences in latitude,
a slight difference exists between the faunas of
the northern half and those of the southern
half of the Izu Banks.

Geographically, the Izu Islands and the Osumi
Group are distant from each other. Similar
patches of banks are not found between the Izu
Banks and the Osumi Banks. Therefore, the
continuity or discontinuity of the fauna has
not yet been studied. However, many species
are found in both areas: Area mania takii
Hatai, Niino, and Kotaka; A car congenitum
(Smith); Samacar pacifica (Nomura and
Zimbo); Haavaiarca uwaensis (Yokoyama);
Mimarcaria aizoi Sakurai ( MS ) ; Striarca fausta

Fig. 14. Chlamys mollita (Reeve) from Hyotanse,
118 m, 18.5 mm X 17.0 mm.

Habe; Chlamys vesiculosus ( Dkr. ) ; C. tissotii
( Bernardi ) ; C. lemniscata ( Rve. ) ; C. mollita
(Rve.); Spondylus anacanthus (Mawe); Pli-
catula muricata (Sowerby); Frigido cardium eos
(Kuroda); Chama argentata Kuroda and Habe;
Tenagodns anguinus (L.) ; Bursa ranelloides
(Rve.), var.; and Conus sp.

Of the species that are not common to the
two areas, some forms are apparently ecological
equivalents of banks-associated forms. Several
examples are shown here:

IZU ISLANDS AREA OSUMI STRAITS AREA

Perotrochus beyrichii (Hilgendorf) P. salmiana (Rolle)

Galeoastraea guttata ( A. Ad. ) G, millegranosa Habe or

G. tayloriana (Smith)

Tucetona shinkurosensis Hatai, Niino, and Kotaka

Lima fujitai Oyama L. tomlini Prashad?

Gians sagamiensis Kuroda and Habe G. kyushensis, sp. nov.

Possible Vertical Limits of the Banks-Associated
Fauna

The distribution of the benthic mollusca is
governed much more forcefully by the sub-
stratum of the depths than by other environ-
mental factors. At equal depths in similar lati-
tudinal position, or under the same ocean

climate, the differences in representation are
due to such bottom factors as mud, sand, gravel,
and rock. The banks-associated molluscan as-
semblages shows mainly hard-bottom facies
mingling with some sandy bottom dwellers of
the depths. This is closely associated with the
insular shelf fauna, since the surrounding areas
of almost all the small islands in the region are

Molluscan Assemblages — Okutani

almost entirely rocky. On the other hand, this
is not always similar to the shelf fauna of the
main islands because the shelf bordering the
main islands does not always present a hard
bottom. For instance, a molluscan assemblage
from the soft bottom within Sagami Bay at a
similar depth ("Soyo” sta. T2 6, Nov. 15, 1958,
35° 15.4' N, 139° 27.9' E, 102 m) is made
up of: Glycymeris rotunda (Dkr.) ; Delecto-
pecten macro chrili cola Habe (believed to be
found attached to the carapace of the giant
spider crab, but recently recovered as free living
individuals ) ; Venus faveolatus Sowerby; Onus-
tus exutus (Reeve); Granulifusus niponicus
(Smith). Moreover, collections from a certain
area in Sagami Bay sometimes contain species
similar to those from the banks under study
(cf 12 and 13), while a different assemblage
is found from another hard bottom in the Bay
at the same level. Therefore, the most remark-
able character of a banks-associated assemblage
is the constant occurrence and constant domi-
nancy of the typical species of the area.

The typical banks-associated assemblage ap-
pears at depths ranging from 32 m as the shal-
lowest to 290 m as the deepest, centering around
100-250 m. In waters shallower than this, a
certain upper-shelf assemblage is observed,
which is frequently collected by a lobster-net or
in Gelinidium collecting. Examples of hard-
bottom dwellers in the shallower zones of the
Oshima Islands are: Turbo cornutus (Solander);
F usinus nicobaricus (Lamarck); Fasciolaria gla-
bra (Dkr.); F. trapezium audouini (Jonus),

Fig. 15. Tucetona shinkurosensis Hatai, Niino, and
Kotaka from Niijima-Nishitai, 14 mm in height.

Fig. 1 6. Tucetona hanzawai (Nomura and Zimbo)
from Kikaijima (fossil).

etc. The sandy bottom facies of the upper shelf
in Okada, Oshima Island, is reported by the
dominance of the following species: Terebra
nebulosa Sowerby; Conus tes sulatus Born;
Chlamys vesiculosus (Dkr.); Callista pilsbryi
Habe; Dosinia iwakawai Oyama and Habe; An-
tigona lamellaris Schumacher; Sunetta concinna
(Dkr.), var.; Paphia amabilis (Philippi).

Judging from these observations, the ecotone
between a shallow insular shelf fauna and a
lower insular shelf fauna, which is almost equi-
valent to banks-associated fauna, is presumably
at a depth of 50 m or so. The deeper limit may
be about 300 m. For example, at a station near
Okinoyama, about 300 m deep, the following
species were collected from a gravel bottom
("Soyo” sta. T20’C, Aug. 11, 1958, 34° 59.0' N,
139° 32.3' E) despite the fact that the similar
banks-associated assemblage is observed on the
upper part of this bank: Chlamys sp. aff. mollita
(Rve.); Delectopecten macrocheilicola Habe;
Keenaea sakuraii Habe; Pandora sp.; Turcicula
crumpii Pilsbry; Trophonopsis echinus (Dali);
Japonacteon archibenthicola Habe. Gravels from
such depths are not covered by calcareous algae.

This archibenthal assemblage almost agrees
with that found from similar depths near Omu-
rodashi reported by Igarashi and Kurata (1956).
As has been stated, the molluscan community
on the islands near the banks under study is
allied to those from neighboring banks; while,
out of biological dredge samples from stations
lower than that, at a depth of 470-480 m, near
Hachijo Island, a different archibenthal mollus-
can assemblage is found ("Soyo” sta. B3, March

PACIFIC SCIENCE, Vol. XVII, January 1963

20, 1958, 33° 07.5' N, 140° 03.2' E, 480 m and
do. Nov. 17, 1958, 33° 10.0' N, 140° 02.7' E,
470 m): Fusinus? sp.; Trophonopsis echinus
(Dali); Nep tunea constricta (Dali), var.; Ben-
thovoluta sp.; Conus sp. nov. The bottom is
found to be of volcanic gravel which is not
coated by algal matter.

SUMMARY AND CONCLUSION

As the result of examination of biological
dredge samples taken from submarine banks
around the Izu Islands, it became clear that pos-
sible characteristic banks-associated molluscan
assemblages are found in this area. Since they
are closely associated with the bottom nature
and the depth of the banks, the assemblages are
composed mostly of the hard-bottom lower shelf
fauna found on the southwestern Pacific coast
of the Japanese main islands.

The present material was collected at random
with various kinds of gear, so that a quantita-
tive analysis of the fauna was not possible. How-
ever, constant occurrence and relative abundance
of certain species were assumed as indicators
of the fauna of the areas. The typical banks-
associated molluscan assemblages from the Izu
Islands area are possibly represented by such
gastropods as Perotrochus beyrichii, Galeoastraea
guttata, and Tenagodus anguinus, together with
several pelecypods such as A car congenitum,
Area mauia takii, Hawaiarca uwaensis, Samacar
pacific a, Mimarcaria aizoi, Tucetona shinku-
rosensis, Chlamys vesiculosus, C. tissotii, C. lem-
niscata, C. mollita, Spondylus anacanthus, Plica-
tula muricata, and Lima fujitai.

Fig. 17. Chama argentata Kuroda and Habe from
insular shelf of Takeshima Is., 11.6 mm X 10-1 mm.

Fig. 18. Spondylus anacanthus (Mawe) from Uji
Islet, 126 m, 40.2 mm X 37.5 mm.

The geographical difference of the assemblage
within the area is not so obvious, but it is clear
that the insular shelf fauna is substantially the
same as that under discussion. It seems that the
species found in the area are those distributed
also in warmer districts. This impression may
be corroborated by facts such as these:

1 ) Similar banks-associated assemblages are
found in southern waters off Kyushu
which are much more subtropic in average '
climate.

2) Tucetona hanzawai and Samacar pacifica,
which are found from the Izu Islands area,
were originally described by Nomura and
Zimbo (1934) from the Ryukyu Lime- j
stone of Kikaijima (or Kikai-ga-shima) ,
an island located far to the southwest of ■
the Osumi Group. Concerning the fossil
fauna found from this Ryukyu Limestone, j
Nomura and Zimbo cited Yabe and Han-
zawa’s opinion that the Ryukyu Lime- j
stone was deposited in waters warmer
than are found there at the present time,
and that fossils found there are very sim-
ilar to the recent fauna found around the j

Molluscan Assemblages — Okutani

Fig. 19. Lima fujitai Oyama from Hyotanse, 118
m, 15.5 mm X 12.0 mm.

Ryukyu Archipelago, which is situated
farther south than Kikaijima Island.

3) S triarc a fausta is another example de-
scribed from a fossil bed (at Moeshima
in Kagoshima Bay, Kyushu) containing
warmer water species.

4) Some elements of tropical origin are
found in the banks fauna, e.g., Chicoreus
superbus, which has been known from the

Bonin (or Ogasawara) Islands, and Area
mauia takii, which is presumably an en-
demic form of a Hawaiian species. These
two species are found at Kurose, which is
the most southern bank in the area.

An assumption that the molluscan assem-
blage of the Izu Banks is an isolated shelf fauna
representing species derived from warmer wa-
ter corresponds to that of Parker and Curray
(1956: 2438) for the Gulf of Mexico. They
concluded from their studies that the banks-
associated molluscan assemblages in waters off
Texas represented a population now isolated
from the main centers of abundance ranging
from southeast Florida to the West Indies and
different from the surrounding level-bottom
communities.

According to Teramachi (personal communi-
cation), there are some distributional gaps of
upper-shelf mollusca even within the warm
Kuroshio area, though they are far less con-
spicuous than those existing at about 36° N on
the Pacific coast of Honshu. One such type of
gap presumably is found around Ashizuri-Saki,
for instance. The distributional difference of
certain species of Fusinus, Siphonalia, Ancilla,
and Fulgoraria off the east and west coast of
the cape (about 34° 42' N, 133° E) may be
found there. Regardless of this fact, the typical

Fig. 20. Distribution of bank-associated mollusca. a, Galeoastraea guttata; b, Tucetona shinkurosensis; c,
Chama argentata; d, Spondylus anacant bus -Lima fujitai-Plicatula muricata (or any two of these).

PACIFIC SCIENCE, Vol. XVII, January 1963

Fig. 21. Gians kyushuensis, sp. nov. (holotype),
16.0 mm X 13.0 mm X 5.5 mm.

banks-associated molluscan assemblages are
found from the discontinuous biotopes, i. e.,
southwestern Kyushu (about 129° 30'' E) and
Izu Islands areas (about 140° E). Latitudinally,
they are distributed from the southern tip of
Izu Peninsula (34° 34' N) southward to Tori-
shima Island (30° 30' N).

Above all, this study shows how marine mol-
lusca on the same level under similar ocean
climate are governed in their distribution by
the substratum. Moreover, a particular mollus-
can assemblage (or simply a combination of
the species) can serve to indicate the sedimen-
tary characteristics of the banks in the Kuroshio
area.

REFERENCES

Habe, T. 1958. On the radulae of Japanese ma-
rine gastropods (4). Venus: Tap. Tour. Make
20(1): 43-60, 2 pis.

Horikoshi, M. 1957. Note on the molluscan
fauna of Sagami Bay and its adjacent waters.
Sci. Rept. Yokohama Nat. Univ., sec. 11,
(6): 37-64, 13 tfs., 1 pi, 2 maps.

Igarashi, M.. and Y. KURATA. 1956. Fish and
mollusks collected by trawl net. Survey Rept.
Tokyo Fish. Exp. Sta. 4 (publ. 94): 32-40.
Mimeo. [In Japanese.]

Kuroda, T . and T. Habe. 1952. New locality
of Perotrochus hirasei . Venus: Jap. Jour.
Malac. 17(1): 5. [In Japanese.}

Niino, H. 1935. On the bottom nature of the
banks of Zenisu, Izu Islands. Chigaku Zasshi
47(562): 32-37, 1 pi, 2 tfs. [In Japanese.]

1952. The bottom characters of the in-
sular shelf around Hachijo Island and the
neighboring banks. Jour. Tokyo Univ. Fisher.
39(1): 101-110, 12 tfs.

— — — 1955. On a manganese nodule and Pero- j
trochus beyrichii dredged from the banks j
near the Izu Islands, Japan. Rec. Oceanog.
Wks. Japan. 2(2): 1-5, 1 pi, 1 tf.

Nomura, S., and N. Zimbo. 1934. Marine mol- j
lusca from the "Ryukyu Limestone” of Kikai-
jima, Ryukyu Group. Sci. Rep. Tohoku Imp. ;
Univ., 2nd ser. (Geol.) 16(2): 109-164.
Parker, R. EL, and J. R. Curray. 1956. Fauna ;
and bathymetry of banks on continental shelf, I
Northwest Gulf of Mexico. Bull Amer.
Assoc. Petrol Geol. 40(10): 2428-2439, 6 j
tfs., 1 pi

Sakurai, K. 1954. [A note.] Yumehamaguri
76: 22. Mimeo. [Journal in Japanese.]

— — — 1959. [A note.} Yumehamaguri 97: 68.

Mimeo. [Journal in Japanese.]

Shirai, S. 1958. On some species of mollusca
collected on the banks of Izu-Shichito. Venus:
Jap. Jour. Malac. 20(1): 87-96, 4 tfs. [In
Japanese.}

Suzuki, K., and A. Sato. 1944. On the bottom
nature in the waters from the southern ex-
tremity of the Izu Peninsula to Kozushima
and Toshima. Jour. Oceanogr. Soc. Japan.
3(4): 193-206. [In Japanese.]

Wada, R. 1954. [A note.] Yumehamaguri 75:
170-171. Mimeo. [Journal in Japanese.}

APPENDIX

Gians kyushuensis Okutani, sp. nov. (Carditi-
dae)

Shell equivalve, dosed, stout, gibbous in

Molluscan Assemblages — Okutani

shape; quadrangular in outline; beak anteriorly
oblique; surface rough, uniformly pink in color,
sometimes with delicate wavy pattern of darker
color on posterior region; radial ribs running
from umbo to distal margin, 20 or 21 in num-
ber, as wide as shallow interspacial grooves,
granulated on anteroventral ones, while sparsely
spinose on rest; longest spinose scale as long as
V5 of shell length; escutcheon narrow; lunule
deeply impressed, cordate in outline.

Interior smooth, with shallow radial grooves;
ashy white with pinkish tint; on right valve,
anterior cardinal teeth vestigial, while posterior
ones elongated; on left valve, anterior cardinal
teeth short, small but prominent, posterior ones
elongated, plate-like in shape; ventral margin
crenulated; mantle scar smooth.

locality: 35° 50' N, 130° 28.5' E, 210 m in
depth, gravels and shell.

TYPES: Left odd valve: 16 mm long, 13 mm
high, 5.5 mm thick (holotype). Left odd valve:
13 mm long, 11.5 mm high, 5 mm thick (para-
type). Right odd valve: 18 mm long, 16 mm
high, 7 mm thick (paratype).

REMARKS: This new species is closely allied
to G. hirasei (Dali),2 but the latter has more
radial ribs which are closely scaly. This new
species is distinguished from G. millegranosa
(Nomura and Zimbo), 3 which has a strongly
inflated shell and granulated radial ribs.

DISTRIBUTION: The type locality and off Kii
Peninsula (collected by Mr. Teramachi; con-
joined valve measures 24 X 20 X 18 mm).

2 Dali, 1918: Proc. U. S. Nat. Mus. 54(2234) : 234.

3 Nomura and Zimbo, 1934: Sci. Rept. Tohoku
Imp. Univ., 2nd ser., 16(2): 154(45), pi. v(i),
figs. 13a, b, 14a, b. Also Okutani, 1958: Venus 20:
220, tfs. 2, 3.

Notes on the Osteology and Systematic Position of Hypoptychus dybowskii
Steindachner and Other Elongate Perciform Fishes1

William A. Gosline2

Longer ago that can gracefully be admitted,
Dr. Paul Kahsbauer of the Vienna Naturhis-
historische Museum was kind enough to send
me a specimen of Hypoptychus dybowskii from
Steindachner’s (1880) original series taken off
"Northern Japan.” Steindachner placed this fish
alongside the Ammodytidae, and there has been
a division of opinion ever since as to whether
it should be included in or excluded from that
family (cf, Regan, 1913; Jordan, 1923; Duncker
and Mohr, 1939; Berg, 1940). In order to in-
vestigate its relationships, the Vienna specimen
has been stained and dissected, and its oste-
ology compared with that of the ammodytids
Bleekeria gilli (Fig. la) and Ammodytes tobi-
anus. The specimen of Bleekeria is Hawaiian
and was retrieved from tuna spewings. Am-
modytes is represented by two series, sent to
me from the U. S. National Museum and the
Museum of Comparative Zoology through the
courtesy of Dr. L. P. Schultz and Dr. G. W.
Mead, respectively.

That Hypoptychus belongs to the superfamily
Ammodytoidae seems certain. The relationships
of the superfamily Ammodytoidae are more
obscure. A second objective of the present in-
vestigation has been to look into this matter,
and a preliminary discussion of certain of the
problems involved here will serve as an intro-
duction to the paper.

In a typical percoid fish, e.g., Epinephelus,
there are 24 vertebrae, and the dorsal fin is
composed of an anterior spinous portion and a
posterior soft portion. In such a fish the ma-
jority of the basal supporting elements, i.e.,
pterygiophores, of the spinous dorsal have a
one-to-one relationship with the vertebrae be-
low them; the soft dorsal rays and their ptery-

1 Contribution No. 1176 of the Hawaii Marine
Laboratory and of the University of Hawaii Depart-
ment of Zoology. Submitted July 25, 1961.

2 Department of Zoology, University of Hawaii.

giophores, in contrast, are more closely spaced so
that there is more than one ray and pterygio-
phore to each vertebra. Time and again, how-
ever, the percoids and their derivatives have
become elongate. This change in shape is fre-
quently accompanied by a whole series of other
alterations. Thus, the cranial crests become low
or disappear, the number of vertebrae increases,
the distinction between dorsal spines and rays
becomes reduced, both types of dorsal (and
anal) rays develop an approximately one-to-one
relationship with the vertebrae, and the caudal
fin becomes rounded and its principal rays re-
duced in number. All of these changes are to be
found among the percoids, e.g., the Cepolidae,
trachinoids, ammodytoids, blennioids, schindler-
ioids, and most gobioids.

Indeed, it seems that all of these modifica-
tions occur together in the majority of elongate
percoid derivatives and that those forms, such
as the ophidioids, where there is more than one
dorsal and anal ray per vertebra are the excep-
tion rather than the rule. On the other hand,
the author is aware of no prepercoid teleost
with a one-to-one relationship between soft dor-
sal and anal rays and the vertebrae. If what
has just been said is correct, it follows that any
fish with such a relationship is a percoid deriva-
tive, but that the unit correspondence between
soft rays and vertebrae is of little use in dis-
tinguishing the various lineages of percoid der-
ivation.

Here, the schindlerioids and gobioids will be
dismissed from further consideration, as each of
these groups has peculiarities by which it may
easily be defined. However, Hawaiian specimens
of the trachinoids Parapercis schauinslandi (Par-
apercidae, Fig. 1 c) and Crystallodytes cookei
(Trichonotidae, Fig. lb), and of the blennioid
Tripterygion atriceps ( Tripterygiidae, Fig. Id)
have been stained and dissected. These speci-
mens will be used both for purposes of com-

Elongate Perciform Fishes— Gosline

Fig. 1. Sketches of a, Bleekeria gilli, from Gosline
and Brock, after Fowler, based on a specimen 3 inches
in total length; b, Crystallodytes cookei, based on a
specimen 2 inches long; c, Parapercis schauinslandi,
from a 3 -inch specimen; and d, T riptery gion atriceps,
from a 1-inch fish.

parison with the three ammodytoids and in
an attempt to obtain some understanding of
the lineages to which each of them belongs.

With regard to these six fishes that have been
investigated in some detail, it may be stated
in advance that the author has not had any
great success in discovering significant cranial
differences between them. It is not so much
that such differences do not exist, as that they
appear to have rather slight systematic value.
Though it may be that the author has simply
overlooked significant differences, it would ap-
pear that the percoids and their immediate
derivatives have a rather standardized skull pat-
tern and that the major morphological differen-
tiation of percoid groups has occurred in other
features.

lacrimal (preorbital) followed by about five sepa-
rately movable, canal-bearing ossicles (cf, Ka-
tayama, 1959: figs. 2-5). Above the fifth,
the infraorbital lateral line canal joins the
supraorbital canal. Of the five ossicles the upper-
most is particularly variable and is sometimes
fused to and sometimes free from the sphenotic.

In Parapercis (Fig. 2a) the infraorbital
canal is complete, passing through a lacrimal
and six separate circumorbital bones. The upper-
most of these is firmly attached to the cranium
in Parapercis. Because six circumorbitals ap-
peared to be a high number, the opposite side
of the same specimen and of a larger specimen
of Parapercis schauinslandi were checked; no
variation was found. The circumorbital struc-
tures of T riptery gion differ from those of Para-
percis in having three instead of six circum-
orbital bones and in the failure of the bone to
close over the sensory canal externally. In Crys-
tallodytes the circumorbital canal is still com-
plete but there are only a lacrimal and two
circumorbitals. The lacrimal and second circum-
orbital are large and laminar, but the anterior
circumorbital is quite small.

In Ammodytes (Fig. 2b) there is a large
lacrimal, followed immediately by a moderate-
sized first infraorbital; then there is a broad
gap followed by two small circumorbitals, the

HEAD SKELETON

CIRCUMORBITAL BONES: In the typical per-
coid the circumorbital series is made up of the

FIG. 2. Lacrimal and circumorbital bones of a, Para-
percis schauinslandi, and b, Ammodytes tobianus.
There are no circumorbital bones bordering the por-
tion of the orbit indicated by the dashed line in
Ammodytes. co, Circumorbital bones; and la, lacrimal.

PACIFIC SCIENCE, Vol. XVII, January 1963

upper of which articulates with the skull. In
Bleekeria a similar break in the circumorbital
ring occurs, but it is shorter than in Ammo-
dytes and the posterior series seems to contain
three or four small elements instead of two.
Hypoptychus has the same two anterior ele-
ments followed by a broad gap; posteriorly,
however, there is only a single ossicle, and it
is fused to the sphenotic.

None of the six fishes have any subocular
shelf from the circumorbitals.

JAWS: In sand-diving fishes like Ammodytes
the mouth is usually not terminal; either it is
withdrawn below an overhanging snout, as in
Crystallodytes, or protected by a prognathous
chin, as in Ammodytes. In Ammodytes the
leading, lower jaw is firmly attached, but the
upper has developed excessive powers of pro-
trusion when the mouth is opened (van Dob-
ben, 1935: 34-36). The great protrusibility of
the upper jaw in Ammodytes is accompanied
by a weakening of the bony elements, and it
is probably in relation to this that Ammodytes
and Bleekeria have edentulous premaxillaries.
So far as jaw structure is concerned, Hypopty-
chus is intermediate between the normal per-
coid type and the specializations found in Am-
modytes and Bleekeria.

The premaxillary of Ammodytes (Fig. 3 a)
consists of a long pedicel movably articulating
at its base with the remaining portion of the
premaxillary. The distal half of the maxillary
tapers gradually to a point (Fig. 3 a). The upper
jaw of Bleekeria is essentially similar to that
of Ammodytes except that a number of small
ossicles are to be found in the ligamentous
tissues connected with the jaw apparatus. Thus,
there is an ossicle above the more lateral of the
two pedicels of the premaxillary, another at
the distal end of the premaxillary, and a whole
series in the ligamentous tissue that runs be-
tween the upper and lower jaws.

The upper jaw of Hypoptychus (Fig. 3 b)
differs from those of Bleekeria and Ammodytes
in the following features: the premaxillary
bears a row of teeth (there are about 14 conical
teeth in a single row on each side, not shown in
Fig. 3 h)\ the premaxillary is fused to its pedicel;
and the tip of the maxillary is expanded distally.

A movable articulation between the pre-

Fig. 3. Head skeletons of ammodytoids: superior
views, with the premaxillary somewhat protruded, of
a, Ammodytes tobianus, and b, Hypoptychus dybow-
skii; lateral view, c, of cranium of Ammodytes tobi-
anus. ba, Basisphenoid; bo, basioccipital; ca, cartilage;
co, circumorbital; eo, exoccipital; ep, epiotic; fr, fron-
tal; ha, anterior hyomandibular socket; hp, posterior
hyomandibular socket; in, intercalar; la, lacrimal; le,
lateral ethmoid; me, mesethmoid; mx, maxillary; na,
nasal; pa, palatine; pc, prootic; pi, pleurosphenoid; po,
posttemporal; pr, parietal; ps, parasphenoid; pt, ptero-
tic; px, premaxillary; so, supraoccipital; sp, sphenotic;
tb, tabular; tf, trigemino-facial foramen; and vo,
vomer. In the superior view of Ammodytes, a, the
epiotic is covered by the posttemporal.

maxillary and its pedicel is also found in Crys-
tallodytes, as well as in blennioids such as
Cirripectus and Istiblennius. In Parapercis and
Tripterygion, which have strong premaxillary
teeth, the pedicel is stout and fused to the
toothed portion.

gill covers: In Ammodytes (Fig. 4b) and
Crystallodytes (Fig. 4c) the subopercles are ex-
panded, presumably to protect the throat region.
Indeed, the lower border of the articular in
Crystallodytes is greatly expanded below as well
(Fig. 4c).

SUSPENSORIUM: The suspensorium of Am-
modytes is specialized in a number of regards
(Fig. 4b). Most notable among these is the
elongated palatine strut. The whole length of
this strut from its forward tip to its articulation
with the quadrate is made up of the palatine

Elongate Perciform Fishes — -Gosline

bone; the ectopterygoid is a minute triangular
splint at the very base. In Bleekeria the struc-
ture of the suspensorium is essentially similar
except that the ectopterygoid is somewhat larger
so that the palatine does not meet the quadrate.
The suspensorium of Hypoptychus (Fig. 4a)
is a quite different structure. The palatine and
ectopterygoid are about equal in size and are
united to one another by a digitate suture. The
metapterygoid is a small splint and the meso-
pterygoid appears to be absent.

Undoubtedly the greatest specialization in
the suspensorium is that found in Crystallo-

FlG. 4. Right gill covers, suspeosoria, and lower
jaws, external view, of a, Hypoptychus dybowskii; b,
Ammodytes tobianus; c, Crystallodytes cookei; and d,
Tripterygion atriceps (with the lower jaw dislocated).
an, Angular; ar, articular; de, dentary; ec, ecto-
pterygoid; hy, hyomandibular; io, interopercle; mt,
metapterygoid; mx, mesopterygoid; op, opercle; pa,
palatine; pp. preopercie; qu, quadrate; sb, subopercle;
and sy, symplectic.

dytes (Fig. 4c). Here the suspensorium is
divided into two well-developed and strong
portions with the ectopterygoid forming a long,
delicate strut between them. In the anterior
portion a large, firm mesopterygoid forms a
shelf under a large part of the eyeball; it is
firmly attached to the strong palatine anteriorly,
but only by membrane to the ectopterygoid.

SKULL: In all of these fishes, there are no
frontal-parietal crests, and the minute supra-
occipital crest does not reach above the skull
surface. Tripterygion , however, has a fringed,
backwardly slanted crest running across the
rear of the skull. This crest lies just behind the
tabular ossicle on each side which bears the
apparently incomplete supratemporal sensory
canal commissure. ( In the related Entomacrodus
the supratemporal commissure is almost com-
pletely enclosed in the skull. Laterally, the com-
missure passes through a tabular ossicle that
is fused to the cranium and thence medially
through the rear of the parietals, leaving a large
opening on the middorsal line.)

In Parapercis and Tripterygion the crania are
somewhat more highly arched over the orbit
than in the others. Probably in association with
this, the wings of the parasphenoid extend
farther up the sides of the postorbital bar than
in the remaining four fishes. In all, however,
the prootic extends over the top of the para-
sphenoid wings to the edge of the orbit. (In
Istiblennius, related to Tripterygion, the para-
sphenoid wings meet the frontals in the usual
blennioid fashion.)

In Ammodytes and Bleekeria the two exoc-
cipital condyles lie adjacent to one another and
form the upper portion of the facet for the
articulation of the convex head of the first
vertebra. In the other forms, including Hypo-
ptychus, the exoccipital condyles lie at either
side of the basioccipital articulation; the two
exoccipital bones do not meet below the fora-
men magnum; and there is no rounded articu-
lar head on the first vertebra.

GILL ARCH SYSTEM: In Ammodytes the bran-
chiostegal ray count is 8-7; in Crystallodytes, 7;
in Bleekeria , 7; in Parapercis and Tripterygion,
6; and in Hypoptychus, 4-4.

In all the fishes under consideration the lower
pharyngeals are separate. Ammodytes and

Bleekeria appear to be the only forms with 3
distinct upper pharnygeals on each side; Hypo-
ptychus has separate upper pharyngeals on arches
2 and 3, but appears to have none on arch 4.
Parapercis and Crystallodytes also have two
pairs of upper pharyngeals, but the posterior
pair seems to represent a combination of
pharyngobranchials 3 and 4. Tripterygion ap-
pears to have only a single set of upper pharyn-
geals.

FINS, FIN SUPPORTS, AND AXIAL SKELETON

ANAL FIN: In the six fishes under considera-
tion there is never more than a single unseg-
mented ray at the front of the anal fin, and even
this is lacking in Crystallodytes and Ammodytes.
All of the remaining anal rays are branched in
Parapercis, some in Hypoptychus, only the last
in Tripterygion, and none in Crystallodytes,
Bleekeria, and Ammodytes.

Unlike the other three fishes, there is in the
three ammodytoids a well-developed caudal
peduncle behind the base of the last anal (and
dorsal) ray; this is supported by about five
vertebrae with bladelike neural and hemal
arches.

DORSAL FIN: Parapercis and T rip ter y gion
are the only fishes under consideration that have
spinous dorsals. Furthermore, in these two the
dorsal fins commence farther forward (over the
3rd vertebra in Parapercis, Fig. 3 a, the rear
of the skull in Tripterygion, Fig. 5b) than in
the others (over the 5th vertebra in Bleekeria,
and still farther back in the remaining forms).
Structurally the spinous dorsal fin differs con-
siderably in Parapercis and Tripterygion. In
Parapercis it appears that the spinous dorsal
has undergone some condensation, possibly as
a result of forward movement of the soft dorsal,
for the pterygiophores of the five spines in-
terdigitate between neural arches 2 and 5 (Fig.
5a) \ one supraneural remains (rather than the
three usually found in the lower percoids). In
Tripterygion there are two spinous dorsals, the
first of 3 spines and the second of 14; it appears
very much as if the anterior 3 have appropriated
the usual percoid supraneurals as their support-
ing bases. In the structure of the pterygiophores
supporting the dorsal spines, Parapercis is con-
siderably more generalized than Tripterygion.

PACIFIC SCIENCE, Vol. XVII, January 1963

Fig. 5. Anterior vertebrae, ribs, dorsal rays and
their supports in a, Parapercis schauinslandi, and b,
Tripterygion atriceps. dr, Dorsal soft ray; ds, dorsal
spine; ep, epipleural rib; na, neural arch; ns, neural
spine; pb, pleural rib; pg, pterygiophore; sh, supra-
neural; so, supraoccipital.

In Parapercis the pterygiophores of the spines
(except that of the first 2) retain their basic
bisegmental structure (Fig. 5a) \ whereas in
Tripterygion each pterygiophore is a fused
monolithic unit (Fig. 5b).

In the soft dorsal, as in the anal, all the rays
are branched in Parapercis, some in Hypopty-
chus, only the last in Tripterygion, and none in
Crystallodytes, Bleekeria, and Ammodytes. In
soft dorsal structure, there are again certain dif-
ferences between Parapercis and Tripterygion
on the one hand, and Crystallodytes and the
ammodytoids on the other. In the first place,
Parapercis and Tripterygion have the last dorsal
(and anal) ray cleft to the base; Crystallodytes
and the ammodytoids do not. Second, the ptery-
giophore of each soft dorsal ray in Parapercis
and Tripterygion interdigitates deeply between
a pair of neural spines (Fig. 5a), and there
is an exact correspondence between vertebrae
and soft dorsal rays. In Crystallodytes and the
ammodytoids the pterygiophores of the soft
dorsal ( and anal ) rays are short, weak structures

Elongate Perciform Fishes— Gosline

that interdigitate little if at all between the
tips of the neural spines, and there is a rough
but inexact correspondence between soft dorsal
(and anal) rays and vertebrae.

CAUDAL FIN: The tails of the ammodytoids
are somewhat forked, those of the other fishes
under investigation more or less rounded. In
all, there is a reduction in the caudal ray num-
ber from the typical percoid count of 17 prin-
cipal rays, 15 branched. In Parapercis there are
15 branched rays, but no outer principal un-
branched rays. In Ammodytes and Bleekeria,
there are 15 principal rays, 13 branched. In
Hypoptychus, there are 13 principal rays; ap-
parently 11 of these were branched, but since
the fin rays of the available specimen are broken
the branched ray count cannot be definitely
established (the same is true of the dorsal,
anal, and pectoral fins). Tripterygion and Crys-
tallodytes have 10 principal rays, 8 branched.

With regard to the caudal skeleton, Paraper-
cis (Fig. 6a) is quite typically percoid (Gos-
line, 1961). There are six separate hypurals,
one uroneural, and three epurals; none of these
elements are fused to the urostyle. In the caudal
skeleton of the other five fishes, considerably
more fusion and/ or reduction has occurred. Hy-
purals 4 and 5 are always fused with the urostyle,
and, in Crystallodytes (Gosline, 1955: fig. 7d)
and Hypoptychus (Fig. 6d ), two or three of the
lower hypurals as well. ( Fig. 6d must be viewed
with some reservation, as the specimen from
which it was drawn may have been aberrant
in having the last two vertebrae fused.) There
are two epurals in Tripterygion (Fig. 6b) and
the three ammodytoids, and only one in Crys-
tallodytes.

PECTORAL FIN: The total number of pec-
toral rays in the fishes investigated is 15 in
Tripterygion and Parapercis, 13 in Ammodytes
and Bleekeria, 12 in Crystallodytes, and 9 in
Hypoptychus. Of these, all are segmented in
Tripterygion and the ammodytoids; however,
there is a small, unsegmented, splintlike upper-

Fig. 6. Caudal skeletons of a, Parapercis schauins-
landi; b, Tripterygion atriceps; c, Bleekeria gilli; and
d, Hypoptychus dybowskii. ce, Centrum; ep, epural;
hr, hemal arch; hs, hemal spine; hy, hypural; na,
neural arch; ns, neural spine; un, uroneural; and ur,
urostyle.

PACIFIC SCIENCE, Vol. XVII, January 1963

most ray in Parapercis and Crystallodytes. Judg-
ing from the material available, it appears that
the uppermost pectoral ray is homologous
whether splintlike or segmented, for it has the
same very peculiar basal structure. Like other
soft rays it consists of two halves. However,
in the uppermost pectoral ray the two halves
are usually not mirror images of one another.
Rather, the scapula articulates with a facet that
lies entirely on the inner "half” of the ray
(except, among the fishes investigated, in Hy-
poptychus and Trip ter ygion ) , An inquiry into
this peculiarity has shown that the scopeliform
genus Sy nodus has a small bony nodule that
lies between two equal halves, but is attached
to the inner. As Starks (1930: 238) noted,

Fig. 7. Right pectoral girdles, external view, of a,
Tripterygion atriceps; b, Bleekeria gilli; c, Hypo-
ptychus dybowskii; and d, Ammodytes tobianus. Only
the uppermost and lowermost pectoral rays are shown
in a, b and d; the top of the pectoral girdle is not
drawn in b; and the lower postcleithmm is not in-
dicated in a. ac, Actinost; cl, cleithrum; co, coracoid;
pc, postcleithrum; pe, pelvic girdle; pm, postcleithrum;
py, pectoral ray; sc, scapula; and scl, supracleithmm.

this nodule probably represents a modified ac-
tinost, which in many higher teleosts became
incorporated into the inner, articular "half” of
the uppermost pectoral ray (Fig. lb, d).

The pectoral girdles of Tripterygion and of
the ammodytoids are shown in Fig. 7. Those
of the ammodytoids are peculiar in having the j
supracleithra and cleithra more or less vertically
aligned.

There are two postcleithra in Parapercis,
Tripterygion, Ammodytes, and Bleekeria, and
apparently not any in Crystallodytes and Hy-
poptychus,

PELVIC FIN: Pelvic fins are lacking in the
three ammodytoids studied here. However, two
splintlike pelvic girdle elements are to be found J!
in Bleekeria (Fig. 7b), and a small pelvic fin
of a spine and three rays, located somewhat
ahead of the pelvic bases, occurs in the related
ammodytid genus Embolichthys (Jordan, 1902).
The three other fishes studied here also have
the pelvics originating ahead of the pectoral
bases. Parapercis has a pelvic spine and five
branched rays, the fourth considerably the long-
est. Crystallodytes has a short pelvic consisting
of a spine and five unbranched but segmented
rays. In Tripterygion there are two well-
developed unbranched, but segmented, rays;
there is no spine. In all three fishes the pelvic
girdle articulates anteriorly with the cleithra.'
In Crystallodytes (Fig. 8 d), the two halves of
the pelvic girdle are rather widely separated an-
teriorly; in Parapercis (Fig. 8b) they are united
for nearly their entire length; in Tripterygion
(Fig. 8c) the two halves are not only united,
but anteriorly they seem to have completely
fused. Futhermore, Tripterygion has the pelvic
girdle firmly wedged between the cleithra.

VETEBRAL COLUMN AND RIBS: Parapercis
has 30 ( 10 -j- 20) vertebrae (including the uro-
style); Tripterygion , 34 (10 + 24); Crystal-
lodytes, 55 (29 + 26); Ammodytes lanceo-
latns, according to Regan (1913: 137), 69
(40 + 29); Bleekeria gilli, according to
Duncker and Mohr (1939: 13), 57 (32 + 25);
and Hypoptycbus , 55 (31 + 24).

In Parapercis, Tripterygion, Ammodytes , and
Bleekeria there are two sets of ribs. The lower,
or pleural, ribs start from the 3rd vertebra; the
upper, or epipieural, ribs start from the 1st,

articulating with the upper surface of the pleu-
ral ribs from the 3rd vertebra on (Fig. 4a, b) .
In Crystallodytes there is only one set of ribs
which starts with the 1st vertebra; this set
would appear to represent the epipleurals. In
Hypoptychus, there is also only one series of
ribs, but it starts from the 3rd vertebra and
would appear to represent the pleural series.

DISCUSSION

On the basis of osteological characters, Par-
apercis, Crystallodytes , Tripterygion, and the
three ammodytoids could be grouped in a num-
ber of ways. One such system would separate
the more elongate forms, i.e., Crystallodytes and
the ammodytoids, from the shorter, stockier
Parapercis and Tripterygion. Such a division
could be expressed osteologically as follows.

Parapercis and Tripterygion. Premaxillary
pedicels stout, firmly fused to the toothed por-
tions. Wings of the parasphenoid expanded,
forming the lower portion of the postorbital
bar. The abdominal portion of the vertebral
column shorter than the caudal portion, consist-
ing of 10 vertebrae. Dorsal with an anterior
spinous portion which commences over or ahead
of the 3rd vertebra. Last dorsal and anal rays
cleft to their bases. Pterygiophores of the dor-
sal and anal rays deeply interdigitating between
successive neural and hemal spines, respectively.

Crystallodytes and the ammodytoids. Pre-
maxillary pedicels long and/or movably articu-
lated with their lateral portions. Wings of the
parasphenoid little developed, not extending
up as a portion of the postorbital bar (Fig. 3c).
The abdominal portion of the vertebral column
longer than the caudal portion, of more than
10 vertebrae. No spinous dorsal, the soft dorsal
commencing over or behind the 5th vertebra.
Last dorsal and anal rays undivided. Pterygio-
phores of the dorsal and anal rays interdigitat-
ing little or not at all between the neural and
hemal spines, respectively.

To group the fishes under consideration in

Fig. 8. Pelvic girdles, from above, of a, Caranx
ignobilis; b, Parapercis schauinslandi; c, Tripterygion
atriceps; and d, Crystallodytes cookei. The pelvic rays
are not indicated in a. cl, Cleithrum; pg, pelvic girdle;
and py, pelvic ray.

PACIFIC SCIENCE, Vol. XVII, January 1963

the above fashion is to separate those nearer
the generalized percoid type from those that
are more specialized. Such a division may
merely represent levels of structural organiza-
tion rather than relationships.

Parapercis is much the least differentiated
from the typical percoids of any of the six
fishes dealt with in this paper. All of the re-
maining five fishes ( Tripterygion , Crystallo-
dytes, Ammodytes , Bleekeria, and Hypoptychus )
have in common the following specializations
over and beyond those found in Parapercis :
circumorbital series of bones incomplete or
complete and consisting of a lacrimal and only
2 or 3 circumorbitals; no pungent dorsal spines;
caudal with 13 or fewer principal rays; uro-
style fused to the upper hypurals; pelvics re-
duced (i.e., without branched rays) or absent.

Most of the minor specializations that Para-
percis does exhibit seem to be associated with
fin structure. Thus the pelvics are advanced in
position and have the inner (actually the 4th)
soft rays the longest, but there appears to be
no great specialization of the pelvic girdle (Fig.
8b). The caudal fin lacks the usual principal
unbranched rays, though the caudal skeleton
is typically percoid (Fig. 6a). The spinous por-
tions of the dorsal and anal have been reduced.
In the anal there is a single small unsegmented
splint at the front of the fin. The spinous dorsal
seems to have been pushed forward and con-
centrated as well as reduced in size, for there
is only one supraneural and the pterygiophores
for all five spines extend between neural arches
2 and 5. The soft dorsal and anal retain a typical
percoid condition. However, their pterygio-
phores interdigitate deeply between successive
neural and hemal spines and bear a one-to-one
relationship with the vertebrae. Now in typical
percoid families, though not apparently in the
Cepolidae, the soft dorsal and anal rays usually
do not interdigitate deeply between the neural
and hemal spines, and there are usually about
two pterygiophores per vertebra. This is true
of even fairly elongate forms like the goatfish,
Mulloidichthys samoensis. To change this ar-
rangement to a one-to-one relationship between
pterygiophores and vertebrae requires either an
increase in the spacing between soft dorsal and
anal rays, or an increase in the number of verte-

brae, or both. The cigar-shaped labrid Cheilio
inermis seems to be an instance where a one-
to-one ratio has been brought about by in-
creased spacing between rays, for this fish re-
tains 24 vertebrae. In most elongate fishes, how-
ever, an increase in vertebrae has also taken
place.

On the basis of Parapercis alone it is impos-
sible to evaluate Regan’s (1913) percoid "Di-
vision Trachiniformes” (equals superfamily
Trachinoidae) . Suffice it here to remark that
there does appear to be a somewhat extensive |
group of usually rather deep-water bottom fishes
having essentially the fin characters described
above for Parapercis. Whether these fishes are
really related is impossible to say at this point.
The relationship of Crystallodytes to this group
is also doubtful. Certainly Crystallodytes repre-
sents a much higher degree of differentiation
from the typical percoids than Parapercis. (For
certain characteristics of Crytallodytes , see be-
low.) Some knowledge of less specialized (or
at least of other) forms of the Crystallodytes
lineage should provide far better indications of
its relationships than are available from a study
of this form alone. (Throughout this paper,
Crystallodytes has been considered a trichonotid,
but that this is a correct family allocation is
dubious, cf, Schultz, I960: 273.)

Tripterygion is generally agreed to be a mem-
ber of the perciform suborder Blennioidei. How-
ever, this suborder, since it was defined and later
restricted by Regan (1912, 1929), has under-
gone considerable disintegration and rearrange-
ment (Starks, 1923; Smith, 1952; Hubbs, 1952;
Gosline, 1955; and Makushok, 1958). Even
after certain nonblennioid groups have been re-
moved, Hubbs and Makushok feel that most or
all of the remaining families may be divided
into a northern (cold water) group and a
southern (warm water) group which may have
had independent origins among the percoid
families.

Tripterygion is, in many respects, one of the
more generalized, i.e., percoid, members of the
whole warm water group. This group differs
radically from the remaining fishes under con-
sideration here in its mode of life. The southern
blennies characteristically use their thickened
pelvic rays to prop the anterior end of the body

Elongate Perciform Fishes — Gosline

away from the hard substratum on which they
feed. The more elongate forms at least charac-
teristically rest with the tail bent, and, when
disturbed, retreat into holes in the rock and
coral by means of sinuous movements of the
body. A number of the characteristic external
features of the southern blennies are probably
associated with this mode of life. For example,
the pelvic rays though reduced in number are
stout, and are attached to a short pelvic girdle
that is firmly wedged between the wings of the
cleithra. The dorsal fin extends far forward, in
the extreme case of Xiphasia to above the eye.
There is often a pair of tentacles or a transverse
fringe of them, e.g., Cirripectes, on the nape;
in Tripterygion there is a low transverse fringe
across the nape in exactly the same position as
in Cirripectes , but it is made up of bony flaps
extending upward from the skull.

A few other characters of Tripterygion atri-
ceps may be mentioned because of their bearing
on blennioid classification. The lateral line canal
of the lacrimal and three circumorbitals is not
covered by bone externally (Hubbs, 1952: 48,
50). A basisphenoid is present (Makushok,
1958: 58). The lowermost actinost in the
pectoral girdle is not greatly longer than deep
(Makushok, 1958: 58; compare Fig. la of
the present paper with Makushok’s fig. 25).
Finally, in the caudal skeleton of Tripterygion
(Fig. 6b) the three lower hypurals have fused
to one another. Hypurals 4 and 5 (above the
axis ) have also fused to one another and to the
urostyle and uroneurals. Hypural 6 is a small
separate ossicle (the "minimum hypural" of
Makushok, 1958), and there are two broad
epurals. In the northern blennies, by contrast,
the upper or epaxial hypurals are usually, though
not always, separate from the urostyle (Maku-
shok, 1958: 38, and fig. 22). Also, the northern
blennioids usually have three epurals, rather
than the two of Tripterygion. (In 1955: fig.
7f, I provided a sketch of the caudal skeleton
of Istiblennius gibbifrons. I have not been able
to relocate the specimen from which the draw-
ing was made, but judging from specimens of
Entomacrodus marmoratus and Istiblennius ze-
bra, which have caudal skeletons very like
that of Tripterygion, the figure is incorrect in
showing a fusion of the lower hypurals and the

epurals with the rest of the urostylar vertebra.)

The investigation of Tripterygion reported
on here would support, in its small way, Hubbs’
(1952) and Makushok’s (1958) thesis that
the "northern” and the "southern” blennies are
diphyletic. But whether they are diphyletic in
the sense that the two groups have wholly dif-
ferent origins or in the sense that they have
diverged in two different directions from the
same or from two closely related stocks would
seem to remain an open question. That Trip-
terygion has little relationship to the congrog-
adids (cf, Smith, 1952) also seems clear.

Among the more elongate fishes investigated,
i.e., Crystallodytes and the ammodytoids, Crys-
tallodytes and Arnmodytes at least are sand div-
ers. The pointed heads, long bodies, low vertical
fins, small or absent pelvics, modified scales, and
peculiarities of the lateral line of the body are
probably associated with this mode of life. The
habits of the other two ammodytoids, Bleekeria
and Hypoptychus, are unknown.

That there is any phylogenetic relationship
between Crystallodytes and the ammodytoids
seems extremely doubtful. Crystallodytes differs
immediately from the ammodytoids in the fol-
lowing characters. The lower jaw is included
and the upper appears to be nonprotrusile. The
circumorbital series of bones is complete. The
cleithra are very oblique with the lower ends
far forward of the upper. The dorsal and anal
extend far back, and there is no well-demarcated
caudal peduncle. Both the principal and procur-
rent fin rays are reduced in number. The lateral
line runs low on the body posteriorly, just
above the anal base. Finally there is a series of
specializations related to the eye and suspen-
sorium of Crystallodytes. In the first place, the
eye itself is very characteristic, for Crystallo-
dytes is a pop-eyed fish (Fig. 1 b) with the
small pupil protruding notably beyond the rest
of the eyeball. This eyeball is supported below
by a large and firm subocular shelf composed of
the greatly expanded mesopterygoid. This bone,
together with the palatine, forms a nearly sepa-
rate portion of the suspensorium only weakly
attached to the rest of this structure by the long,
weak ectopterygoid (Fig. 4c). Neither the sus-
pensorium nor the peculiar eyeball shows any
relationship whatever to those structures in

100

Parapercis, which is typically percoid in these
respects. Nor does Crystallodytes show any re-
lationship to any of the other fishes investigated
in these structures. Indeed, the only fishes that
would seem to have a suspensorium anything
like that of Crystallodytes are the congrogadids
(cf, Regan, 1912: fig. 2b, and Smith, 1952:
pi. 6B).

The three ammodytoid fishes may be defined
as follows: elongate fishes with premaxillati.es
highly protrusile. Circumorbital bones incom-
plete, the lacrimal and first circumorbital sepa-
rated from the rest of the series. Fins without
spines or unsegmented rays except for the pro-
current rays of the caudal (pelvic rays of Em-
bolichthys?) . Caudal forked or emarginate, pre-
ceded by a well-marked caudal peduncle which
is supported by five or more vertebrae with
bladelike neural and hemal spines. P'elvics ab-
sent (of a spine and three rays in Embolicbtbys,
Jordan, 1902). Vertebrae 55 to 69, the abdom-
inal vertebrae more numerous than the caudal,
in approximately a one-to-one relationship with
the dorsal and anal rays above and below
them. Cleithra and supracleithra almost verti-
cally aligned.

The analysis of Hypoptycbus indicates that
it is widely separated from Ammodytes and
Bleekeria, Though the three genera hold a num-
ber of features in common it would seem that
Hypoptycbus has evolved in quite a different
direction from the other two. Thus, while Hy-
poptycbus remains more percoid in jaw struc-
ture and skull-vertebral column articulation, it
has become more specialized (degenerate?) in
almost every other feature: the bones are thin;
the branchiostegal rays and fin rays are reduced
in number; the scales have been completely
lost; etc. Hypoptycbus well warrants the sepa-
rate family Hypoptychidae apparently first as-
signed to it by Jordan (1923: 230).

The families Hypoptychidae and Ammody-
tidae may be contrasted as follows.

Hypoptychidae. Scales entirely lacking. Jaws
subequal, the premaxillary with teeth and with
its long pedicel firmly attached. Branchiostegal
rays 4. Articular facets of the exoccipitals widely
separate. Dorsal and anal fins equal in length,
of about 20 rays. Caudal with 13 principal rays;
pectorals with 9.

PACIFIC SCIENCE, Vol. XVII, January 1963

Ammodytidae. At least some scales present.
Lower jaw protruding, sharptipped. Premaxil-
lary without teeth and with its pedicel movably
attached to its lateral portion. Branchiostegal
rays 7 or 8. Articular facets of exoccipitals con-
tiguous. Dorsal extending far forward of the
anal. Caudal with 15 principal rays; pectorals
with 13.

The problem of ammodytoid origin remains
obscure. The majority of features point to a
percoid origin of some sort, but none of the
percoid families known to the author would
seem to provide a suitable ancestor.

The ammodytoids in turn would appear to
have led to nothing with the exception of one
highly speculative possibility. If the terminal
vertebra of Hypoptycbus is not merely the re-
sult of fusion in an aberrant specimen, then a
progressive evolution along many of the lines
already apparent in that fish might end in a
neotenic form very like Schindleria (cf, Gos-
line, 1959).

Whatever the ancestors and derivatives of
the ammodytoids may be, they remain, so far
as known, sufficiently isolated and characterized
to warrant fully the superfamily status among
the Percoidei that has generally (cf, Regan,
1913) been assigned to them.

REFERENCES

Berg, L. S. 1940. Classification of fishes, both
recent and fossil Trav. Inst. Zool. Acad. Sci.
U.R.S.S., Leningrad, 5: 87-517, 190 figs.

VAN Dobben, W. H. 1935. liber den Kiefer-
mechanismus der Knochenfische. Arch. Neer-
landaises de Zool. 2: 1—71, 50 figs.

L> unck.ee, G , and E. Mohr. 1939’. Revision der
Ammodytidae. Mitt. Zool. Mus. Berlin 24:
8-31, 4 figs.

Goslime, W. A. 1955. The osteology and re-
lationships of certain gobioid fishes, with
particular reference to the genera Kraemeria
and Microdesmus. Pacific Sci. 9(2): 158-
170, 7 figs.

— — — - 1959. Four new species, a new genus,
and a new suborder of Hawaiian fishes. Pa-
cific Sci. 13(1): 67-77, 6 figs.

Elongate Perciform Fishes— Gosline

101

- — 1961. The perciform caudal skeleton.

Copeia, 1961: 265-270, 3 figs.

Hubbs, Clark. 1952. A contribution to the
classification of the blennioid fishes of the
family Clinidae, with a partial revision of
the Eastern Pacific forms. Stanford Ichth.
Bull. 4: 41-165, 64 figs.

Jordan, D. S. 1902. Supplementary note on
Bleekefia mitsukurii, and on certain Japa-
nese fishes. Proc. U.S. Nat. Mus. 25: 693—
696, pi. 30, 1 fig.

— — 1923. A classification of fishes includ-
ing families and genera as far as known.
Stanford Univ. Publ, Univ. Sen, Biol. Sci.
3: 79-243.

KATAYAMA, M 1959. Studies on the serranid
fishes of Japan ( 1 ) . Bull. Faculty Educ, Ya-
maguchi Univ. 8: 103-180, 39 figs.

Makushok, V. M. 1958. The morphology and
classification of the northern blennioid fishes
( Stichaeoidae, Blennioidei, Pisces ) . Trudy
Zool. Inst. Akad. Nauk S.S.S.R. 25: 3-129,
83 figs. (In Russian; English translation
mimeographed. )

REGAN, C. T. 1912. The classification of the
blennioid fishes. Ann. Mag. Nat. Hist., ser.
8, 10: 265-280, 4 figs.

— _ — __ 1913. The classification of the percoid
fishes. Ann. Mag. Nat. Hist., ser. 8, 12:
111-145.

— — - — - 1929. Fishes. In: Encyclopaedia Britan-
nica, 14th ed., pp. 305-329.

Schultz, L, P, et al. I960. Fishes of the Mar-
shall and Marianas Islands. Vol. 2. U. S. Nat.
Mus. Bull. 202: 1-438, pis. 75-123, 132 figs.

Smith, J. L. B, 1952. The fishes of the family
Haliophidae. Ann. Mag. Nat. Hist., ser. 12,
5: 85-101, pi. 6, 2 figs.

Starks, E. C. 1923. The osteology and relation-
ships of the uranoscopoid fishes. Stanford
Univ. Publ., Univ. Ser., Biol. Sci. 3: 259-290,
5 pis.

— — 1930. The primary shoulder girdle of
the bony fishes. Stanford Univ. Publ., Univ.
Ser., Biol. Sci. 6: 149-239, 38 figs.

Steindachner, F. 1880. Ichthyologische Beit-
rage (IX) . Sitzungsberichte k. Akad. Wiss.,
Wien, math.-naturwiss. Classe 82: 238-266,
pis. 1-6.

Observations and Experiments on the Food Habits of
California Sea Hares of the Genus Aplysia

Lindsay R. Winkler1 and E. Yale Dawson2

In A PREVIOUS paper the senior author (Wink-
ler, 1959*2) presented the results of fecal pellet
analyses to determine the influence of diet on
the color of Aplysia calif ornica Cooper. Only
the dominant plants were mentioned in con-
nection with that immediate problem. The pres-
ent paper contains more detailed information
from that study and from subsequent observa-
tions and reviews of references to Aplysia food
plants.

The food of A. calif ornica has received only
limited consideration in connection with other
studies. This is also true, however, of Aplysia
species in general. Even in the large monograph
of Mazzarelli ( 1893) the only indication of the
nature of the food is in a discussion of the
method of feeding, where we find the phrase

. . algae (ordinarily Ulva lactuca ) which
serves as nutriment.” In the extensive study of
European Aplysia punctata by Eales (1921),
about half a page is concerned with food. She
notes an apparent change in food habits with
age. The young, she says, live on red seaweed
like Deles seria [probably D. sanguined] and
then slowly migrate through the Laminaria
zone, where they feed on Laminaria, into shal-
low water, where the fully grown adults live on
Fucus and Ulva. However, since she also states
that they are found in the greatest numbers in
the Zoster a beds, it may be that like Aplysia
calif ornica they also feed on Zoster a.

As to the feeding habits of A. calif ornica,
MacGinitie (1935) stated that the animal eats
Zostera in Elkhorn Slough. In a later work
(MacGinitie, 1949) he stated that they eat "sea-
weeds.” Winkler (1959^) referred to three red

1 Department of Pharmacology, School of Medicine,
Loma Linda University, Loma Linda, California. Pres-
ent address: College of the Desert Palm, Desert,
California.

Manuscript received November 6, 1961.

2 Beaudette Foundation for Biological Research,
Santa Ynez, California. Present address: Hancock
Foundation, University of Southern California.

seaweeds used as food by Aplysia calif ornica and
to their effect on the color pattern. Another
paper (Winkler, 1959&) mentions the effect of
diet, both qualitatively and quantitatively, on
the purple secretion. Winkler (1961) mentions
diet in connection with two special habitats
which are reported here in more detail. Marcus
(I96I) stated that A. calif ornica eats seaweeds,
"especially Zostera and C odium.”

MATERIALS AND METHODS

Young specimens of Aplysia calif ornica 4-5
inches long were captured on the central shore
area of Lunada Bay, Palos Verdes, California,
and each was placed in an individual pint fruit
jar until fecal pellets were passed. The pellets
were then removed to small screw-capped vials
and the animal was released at sufficient dis-
tance to preclude resampling. Twenty-five speci-
mens from this area were sampled in this man-
ner. Pellets were also collected from larger
specimens wandering in the deeper waters of
the bay and from 1 mile north of the north
side of Palos Verdes Point. The pellets were
kept under refrigeration until examined and
identified by the junior author. Since the effi-
ciency of sea hare digestion appears to be in
indirect proportion to the rate and quantity of
food ingestion, identification of the algae of-
fered no great problem since these sea hares
were well fed.

Observations on specimens taken from Do-
heny Beach, California, and from a small lagoon
at Lunada Bay, possessing a different flora than
the former, are also reported. The food plants
determined from crop contents were made dur-
ing other studies involving the sacrifice of the
animals.

RESULTS

Results from 25 young specimens taken from
Lunada Bay in 1955 are presented in Table 1.
In this area the following seaweeds were noted

102

Food Habits of Aplysia — Winkler and Dawson

103

TABLE 1

Diet of Young Aplysia calif omica at Lunada Bay

KINDS OF PLANTS*

NO.

SPECIMENS

IN WHICH

FOUND

NO.

SPECIMENS

IN WHICH

DOMINANT

Acrosorium uncinatum

Centroceras clavulatum

Ceramium eatonianum

Corallina vancouveriensis

Gelidium coulteri

G. crinale (?)

Gelidium sp.

Gigartina canaliculata

Hypnea valentiae

Lophosiphonia sp.

Phyllospadix sp.

Plocamium pacifcum

Pterocladia sp.

Pterosiphonia sp.

S per moth amnion snyderae

Sphacelaria sp.

Ulva sp.

* Although marked differences in the relative abundances of
the various algae in the habitat would seem to negate the
validity of this table in indicating selectivity of food by the
sea hares, there is some interesting supporting evidence. Al-
though Gigartina canaliculata was the dominant intertidal
rock-cover alga throughout the area, and was eaten, it appeared
far less dominant in the pellets than the small, relatively in-
conspicuous epiphytic alga Ceramium. Similarly, although
Ulva and Plocamium were so sparse in the areas as generally
to go unnoticed, these, especially the latter, appeared in the
pellets.

to be common, though never found in the fecal
pellets: Laurencia splendens, Chondria calif orn-
ica, Gigartina leptorhynchos, Prionitis lanceo-
lata, Laurencia subopposita, Cystoseira osmun-
dacea, Gymnogongrus leptophyllus, Bossea or-
bigniana, Herposiphonia sp., Rhodymenia sp.,
and Colpomenia sinuosa.

All specimens taken from deeper water in
Lunada Bay and off Palos Verdes Point were
similar in the algal content of their pellets. In
every case the dominant plant was Plocamium
pacificum, but with some admixture. These ad-
mixtures included Phyllospadix sp., Lophosiph-
onia sp., Gelidium (several species), and Cer-
amium eatonianum .

In May 1961 the larger red-striped A. cali-
fornica from Lunada Bay were found feeding
exclusively on the red alga Laurencia pacifica,
which turned their blood and digestive glands
a deep red.

Observations of the crop contents of several
hundred Aplysia specimens were made during
the course of work in 1959 at the Lunada Bay
lagoon. Occasional spot checks were made in
which the contents were determined as to
species, but no single alga was recognized as
dominant. The samples contained: Chondria
calif omica, Lithothrix aspergillum, Ceramium
eatonianum, Centroceras clavulatum, Plocamium
pacifcum, Corallina gracillus, and Ecto carpus
sp., intermixed with fragments of coralline al-
gae. The flora of this small lagoon contained
more coralline algae and much less Gigartina
canaliculata than the afore-mentioned 1955 col-
lection site located Vy mile to the south on
Lunada Bay.

The collecting area at Doheny Beach was
located at the extreme north end of the state
park, where a small lagoon of uniformly shal-
low water is protected on the north by a strati-
fied rocky peninsula submerged at higher tides,
and on the south by a wave-formed reef of
small boulders and gravel. The flora was com-
paratively limited, consisting predominately of
various jointed coralline algae with intertwined
Hypnea valentiae. Algal growth improved as
one moved to the seaweed area of the lagoon. In
this habitat 86 specimens were collected, but
because of apparently grossly similar crop con-
tents only 8 specimens were analyzed for pre-
sentation in Table 2. Collection dates were
March 24, 1959, for numbers D-02 to D-08,
and April 29, 1959, for the remainder. It will
be noted that the latter collection shows a
greater specialization of diet than the former
and may reflect the concentration of sea hares
farther out in the lagoon than at the former
collection date. It may, on the other hand, sug-
gest that the sea hare tends to become more
selective in its food as it becomes older. This
has generally been noted from other areas.

The algal contents of crops of Aplysia vac-
caria have also been examined. Animals col-
lected in their usual haunts in the Egregia beds
have contained only large amounts of Egregia
in their crops. On the other hand, when the
animals are inshore in breeding aggregations
the crops are usually empty or contain sand or
other material of little significance. On one oc-
casion only, several such animals were found

104

PACIFIC SCIENCE, VoL XVII, January 1963

to have ingested a quantity of Pelvetia fastigi-
ata, a food plant that was consistently refused
by aquarium-confined specimens (Winkler,
1955 ) . The diet of the young of this species has
not been determined.

DISCUSSION AND CONCLUSIONS

In northern California Aplysia californica is
known principally from protected bays and es-
tuaries such as Elkhorn Slough, Bodega Bay,
and Tomales Bay. In these areas, beds of Ulva
and Enteromorpha often occur abundantly, and
J. J. Gonor has reported to us in a personal
communication that A. californica lives in these
beds and apparently uses both plants for food.
Marcus (1961), reporting upon specimens
from Bodega Bay and Tomales Bay, indicated
that A. californica feeds on sea plants, especially
Zoster a and C odium. MacGinitie (1935) stated
that sea hares in Elkhorn Slough fed on Zoster a,
but in subsequent personal correspondence has
indicated that they also fed on Enteromorpha,
which is sometimes the most prevalent plant in
the slough. However, he also observed that at
certain seasons the alga was absent from the
area while Zoster a remained. He concluded,
then, that Zoster a was their principal staple
food.

In southern California the eel grass, Zoster a,
does not occur intertidally on the open coast
and is uncommon in bays, and although the

similar surf grass, Phyllospadix, grows in pro-
fusion at low water levels it is rarely ingested
by sea hares. Likewise, neither Ulva nor Entero-
morpha is abundant in open coast habitats of
southern California, and, although captive sea
hares showed preference to Ulva or Enteromor-
pha over any other food offered them in the
laboratory, in nature they resort to feeding on
red algae, which are almost their exclusive food.

Young southern California sea hares seem
to eat a wide variety of seaweeds depending
upon the flora in their immediate surroundings,
although some species are apparently habitually
refused. As they grow older and wander farther J
they seem to become more selective in their
food habits and develop a behavior pattern that
is difficult to break under changed feeding con- ;
ditions in the laboratory.

It has been generally observed that Aplysia
californica in northern California is of larger ;
average size than in southern California, and
the senior author considers that this larger size,
as well as the difference in color (Winkler
1959^), results from dietary differences. Ex-
perimental evidence for the variation in size
similar to that for color variation is needed to j
confirm the homogeneity of the two forms. I
Two distinctive dietary factors offer themselves
for study: (1) the comparative nutritive value
of a high Ulva-Enteromorpha component in the
diet during spring and summer; (2) the effect
of almost exclusive feeding on the marine grass,

TABLE 2

Presence and Relative Abundance of Several Algae in Crops
OF Aplysia californica FROM DOHENY Beach, CALIFORNIA

COLLECTION

NO.

Hypnea

valentiae

Jania

tenella

Corallina

gracilis

Corallina

pinnatifolia

Corallina

vancouveriensis

Sphacelaria

sp.

Lauren cia
sp.

Lithothrix

aspergillum

D-02

XXX

D-04

D-06

XXX

D-08

iiii

D-20

xxxx

D-21

xxxx

D-27

xxxx

D-28

XXX

1 x

Food Habits of Aplysia — Winkler and Dawson

105

Zostera marina, during winter. Further studies
are also needed on the diet of adult Aplysia
calif ornica which, according to Limbaugh (per-
sonal communication ) are most numerous at
depths of 30-40 ft and have been observed
as deep as 100 ft, depending upon concentra-
tions of plants. He mentions them eating the
"thinner red and green seaweeds.”

From evidence now available it would appear
that Aplysia calif ornica and A. vaccaria are not
competitive for food, and that the former uses
red and green algae as well as marine phanero-
grams for food, while the diet of the latter is
essentially restricted to the brown alga Egregia,
at least in the tidal zone. The nutritive value of
Egregia seems to favor growth to large size, for
Limbaugh has reported observations of speci-
mens of A. vaccaria in the subtidal zone measur-
ing up to 30 inches and an estimated 35 lb,
making them the giants of the gastropoda.

SUMMARY

Young Aplysia calif ornica on the open coast
of southern California subsist on a diet mainly
of fleshy red algae but show a preference for
Ulva or Enteromorpha under artificial feeding.
In the northern end of its range, where it ap-
pears to become largely estuarine in habitat, all
reports indicate that it feeds almost exclusively
on green algae and eel grass, Zostera marina. It
is suggested that the dietary differences may in-
fluence its size as well as its color, and that an
adquate winter food supply is an important
factor.

A. vaccaria appears to feed exclusively on the
brown alga, Egregia.

REFERENCES

Eales, Nellie B. 1921. Aplysia. L. M. B. C.
Memoirs, Proc. Trans. Liverpool Biol. Soc.
35: 183-266.

MacGinitie, G. E. 1935. Ecological aspect of
a California marine estuary. Amer. Midi. Nat.
16: 737.

1949. Natural history of marine ani-
mals. McGraw-Hill, New York. 473 pp.

Marcus, Ernst. 1961. Opisthobranch mollusks
from California. Veliger 3 ( Suppl. ) : 9.

Mazzarelli, G. 1893* Monographia delle Ap-
lysiidae del Golfo di Napoli. Mem. Soc. Ital.
Sci. Modena, ser 3, 9: 1-205.

Winkler, Lindsay R. 1955. A new species of
Aplysia on the southern California coast. So.
Calif. Acad. Sci. Bull. 54: 5.

1959^. A mechanism of color variation

operating in the West Coast sea hare, Aplysia
calif ornica Cooper. Pacific Sci. 13(1): 63-
66.

1959 b. Intraspecific variation in the

purple secretion of the California sea hare,
Aplysia calif ornica Cooper. Pacific Sci. 13
(4): 357-361.

1961. Preliminary tests of the toxin ex-
tracted from California sea hares of the genus
Aplysia. Pacific Sci. 15(2): 211-214.

Littoral Sedimentary Processes on Kauai,
A Subtropical High Island1

D. L. Inman,2 W. R. Gayman,2 and D. C. Cox3

ABSTRACT: Beach and shallow water sand samples from the island of Kauai,
Hawaiian Islands, were studied to ascertain the effects of climate on the supply
of sediment, and of wave action on the dispersal and transport of sand along the
shores of this circular island. The littoral sediments of the island are made up of
two components: biogenous material, such as shell, coral, and foraminiferal sands,
formed near the shoreline; and terrigenous material, consisting mostly of volcanic
mineral and rock fragments brought to the beach by rivers.

The northeast trade winds play a dominant part in the climatic and oceanographic
processes affecting the island. On the windward side of Kauai, the annual rainfall
ranges from 30 inches near the coast to over 460 inches at higher elevations; on
the leeward side, annual rainfall is less than 20 inches. This large variation in
rainfall results in a climatic range from tropic-humid to semi-arid. Rivers draining
the wet, windward portion of the island carry relatively little sand, and the shoreline
on this side is characterized by intermittent fringing reefs and beaches of calcareous
sand. A detailed study of several reefs on the windward side of the island shows
that each pair of fringing reefs, divided by a relatively deep inlet, constitute sep-
arate cells for the circulation of water and distribution of sediments.

The highest concentrations of volcanic sand occur on the lee side of the island,
where streams draining the semi-arid region enter the coast. The concentration of
volcanic material in the beach sands decreases logarithmically with distance along
the coast from the rivers.

KAUAI is the next to westernmost of the wind-
ward islands of the Hawaiian Archipelago ( Fig.
1). It is roughly circular in shape, with a di-
ameter of about 25 nautical miles (Fig. 2).
The island is volcanic and has a central plateau
whose eastern rim exceeds 5,000 ft in elevation.
This mile-high rim intercepts the trade winds,
causing moderate to extreme precipitation on
the windward slope and producing a rain-
shadow desert along portions of the leeward
coast.

1 Published as Contribution No. 24 of the Hawaii
Institute of Geophysics and, with the approval of the
Director, as Paper Number 88 in the Journal Series
of the Experiment Station, Hawaiian Sugar Planters’
Association. Manuscript received November 22, 1961.

2 Scripps Institution of Oceanography of the Uni-
versity of California, La Jolla, California.

3 Hawaii Institute of Geophysics, University of Ha-
waii; formerly Experiment Station, Hawaiian Sugar
Planters’ Association.

The contrast between windward and leeward
sides of the island in terms of climate and
source of sediment suggested that this island
was an ideal locality to study the relation be-
tween nearshore sedimentation and the source
and type of sediment. It was found that the
windward streams, although high in runoff, con-
tributed very little sand-size material to the
nearshore areas. On this side, the coastal en-
vironment is characterized by fringing coral
reefs, with protected beaches composed of cal-
careous sand derived from shells, reef detritus,
and foraminifera. Kapaa Reef had been investi-
gated by Kohn and Helfrich (1957) and, since
it seemed typical of the windward reefs, it was
selected for more detailed study.

Consistently high concentrations of terrige-
nous material were found on only a few of the
leeward beaches. These deposits were particu-
larly conspicuous near the mouth of the Wai-

106

Littoral Sedimentary Processes — Inman, Gayman and Cox

107

Fig. 1. Index chart and exposure to waves and winds. Heavy arrows indicate directions of approach of the
five major wave types which affect the island.

mea River, which drains the major canyon on
the island and is in an area of relatively light
rainfall. The most typical of the lee beaches in-
vestigated lay along the shores of the Mana
Coastal Plain.

During the study, it was observed that beach
and dune rock is especially well developed along
portions of the Kauai coast. This was the sub-
ject of a special investigation by Emery and
Cox (1956). Some additional observations are
discussed under the section on sediments.

PHYSIOGRAPHY

Kauai is one of a chain of volcanic islands in
the Hawaiian Archipelago. It formed as a shield
volcano built up from the deep floor of the Pa-

cific Ocean by thin flows of basaltic lava. The
original shield collapsed to form a broad cal-
dera, which subsequently filled with thicker lava
flows and talus deposits. The caldera is now al-
most obscured by faulting, extensive erosion,
and stream cutting. Ninety % of the flows are
composed of normal olivine basalt, while the
remaining rock is a basalt either very rich (pic-
rite-basalt) or very poor in olivine. A compre-
hensive and up-to-date review of the geology of
Kauai is given by Macdonald, Davis, and Cox
(I960).

The ocean floor around Kauai is characterized
by a number of sloping terraces which are ter-
minated on their seaward side by a marked in-
crease in slope. Although the depths of the
breaks in slopes at the outer edges of the ter-

108

PACIFIC SCIENCE, Vol. XVII, January 1963

races range on various sides of the island from
30 ft to 500 ft, their grouping suggests that
terracing occurred at about four separate levels.
The shoalest and widest terrace commonly ex-
tends to depths of 40 to 90 ft; others extend
from about 150 ft to between 240 to 350 ft;
and for the deepest, to about 450 ft. The latter
depth presumably marks the outer edge of a
wave-cut terrace formed during a still stand
near the lowest Wisconsin Sea Level. Generally,
the 450-ft contour ranges from 1 to 2 miles
from shore, although it extends to 4 miles off
the west coast.

The most gently sloping part of the insular
shelf, that part shoaler than 90 ft, is frequently

characterized by many apparently closed depres-
sions, generally ranging from 10 to 30 ft deep.
These are most common on the north coast,
although a very large depression occurs on the
west coast off Makaha Point ( Fig. 2 ) . The con-
figuration of the submarine ridge surrounding
this depression suggests that it may be a relic
coastal sand dune, formed at a lower stand of
the sea, or possibly a relic barrier beach or reef.4

4 Observations made while diving on this reef dur-
ing September 1962 subsequent to submission of this
manuscript show the reef to consist of a framework of
coral. It is believed that it represents a barrier reef
formed during a previous lower stand of sea, and the
name "Kaheko” is proposed for the reef because of
its presumed great age.

Fig. 2. Topography, bathymetry, and sample locations. Numbers along coastline refer to sample designations
listed in text and in Table 2. Intermittent jagged line along shore shows location of major reefs.

Littoral Sedimentary Processes— Inman, Gayman and Cox

109

* SYMBOL a NUMBER INDICATE GAGING STATIONS
SLANTED NUMBERS ARE VALUE OF ANNUAL
RAINFALL IN INCHES

LETTERS DESIGNATE DRAINAGE AREAS

FIG. 3= Rainfall and drainage basins on Kauai. Dotted lines indicate outline of drainage areas. Numbers and
letters, designating stream gaging stations and drainage areas, respectively, are referred to in Table 1. Note area
of lowest rainfall along western coast. Isohytel lines from the Meteorology Department, Pineapple Research
Institute, and the Experiment Station, Hawaiian Sugar Planters’ Association, 1948.

There are at least five well-developed sea valleys
which cut the shallower portions of the insular
shelf. All of these, except the one off Hanalei
Bay, appear to terminate above the 450-ft con-
tour.

CLIMATE

The annual precipitation varies from an av-
erage of about 50 inches, on the windward
(northeastern) shore, to over 450 inches, at the
summit of Mt. Waialeale, and decreases to less
than 20 inches in the rain-shadow produced on
the extreme lee side of the island. Vegetation
varies from tropical rain forest on the windward
side, to arid on the southwestern slopes. Figure
3 shows the drainage pattern, the outlines of

the major drainage basins, and the contour lines
of mean annual rainfall, or isohyets, for the
island, as well as the location of several stream-
flow gaging stations.

Langbein and Schumm (1958) have shown
that the sediment yield from a drainage basin
varies as a function of climate, and that the
maximum yield occurs for an effective precipi-
tation of about 12 inches. The effective precipi-
tation5 is obtained from the actual precipitation
by correcting for evapotranspiration losses to a
standard temperature of 50 F. Sediment yield is
less for precipitation either greater or less than

5 Langbein and Schumm (1958) define "effective
precipitation” as the amount of precipitation that
would be required at a mean temperature of 50 F to
produce the actual annual runoff from the basin.

110

this optimum value of about 12 inches (Fig. 4).
Although sediment yield is influenced by vege-
tation, temperature, rainfall characteristics, and
topography, a rough approximation for sedi-
ment yield can be obtained from the effective
precipitation.

Data indicative of the differences between
the climates of the various drainage basins on
Kauai and of the supplies of sediments in those
basins are given in Table 1. These data include:

( 1 ) a rough calculation of the mean annual
rainfall for the various drainage areas; (2) ef-
fective precipitation calculated from rainfall;
( 3 ) runoff data for the upper portion of some
of the drainage basins, obtained from gaging
stations; and (4) the percentage of carbonate
in the beach sand sample taken nearest the
stream mouths.

In general, the table confirms that the great-
est precipitation and the most runoff occur on
the windward side of the island where the least
amount of terrigenous sediment is found in the
beach sand. The Langbein and Schumm curve
(Fig. 4) shows that the potential sediment yield
is appreciably higher when the effective pre-
cipitation does not exceed about 35 inches, with
a maximum yield corresponding to a value of
about 12 inches. The equivalent values in terms
of annual rainfall on Kauai6 give the band of
high sediment yield as occurring where the mean
annual rainfall does not exceed 46 inches, and

0 After correcting for evapotranspiration, following
Langbein et al. (1949: fig. 2).

FIG. 4. Relation between effective precipitation and
annual sediment yield (taken from Langbein and
Schumm, 1958: fig. 3).

PACIFIC SCIENCE, Vol. XVII, January 1963

a maximum sediment yield corresponding to a
mean annual rainfall of 22 inches. Figure 3
shows that a relatively high percentage of the
drainage basins on the lee side of the island fall
within the range of high sediment yield,
whereas the rainfall in the windward drainage
basins is appreciably greater and thus produces
a somewhat lower yield of sediment. The mean
annual rainfall for each drainage basin and the
corresponding value for the mean effective pre-
cipitation are listed in Table 1, columns 4 and
5, respectively. The Waimea River drainage
basin (Fig. 3, B), which is the principal basin
on the lee side of the island, has mean values
of annual rainfall and effective precipitation of
70 inches and 53 inches, respectively, which are
much nearer the value of maximum sediment
yield than are those for the principal windward
drainage basins (Fig. 3, E, F, G) . In fact, much
of the Waimea drainage basin falls within the
rainfall range of potentially high sediment
yield. Also, the sediment yield in the Waimea
basin is accentuated by the occurrence of ex-
treme topographic relief in the low rainfall area.
This combination of factors, tending to increase
sediment yield, does not occur extensively in
the windward basins.

The high carbonate content of the beach sand
samples on the windward coast tends to con-
firm the difference in sediment yield between
the lee and windward drainage basins (Table 1,
last column). In addition, cursory inspection of
river banks and river mouths indicated that not
only is the yield greater on the leeward side
but also that there is a significant difference in
the size of the sediment: the leeward streams
carry large amounts of sand-size material,
whereas the windward streams carry silt and
clay-size material. A predominance of silt and
clay-size sediment indicates chemical weather-
ing, a process which is enhanced in the wind-
ward basins by greater vegetation density and
slower rate of removal of disintegration prod-
ucts.

EXPOSURE TO WAVES

Although no complete statistical wave data
for spectrums and directions of waves are avail-
able for the island of Kauai, a general descrip-
tion of the prevailing wave conditions can be

Littoral Sedimentary Processes — Inman, Gayman and Cox

111

TABLE 1

Rainfall, Runoff, and Sediment Composition for Drainage Basins on Kauai

BEACH sand

SAMPLE

NEAREST

DRAINAGE BASIN

river basin above gaging station

RIVER

Desig-

nation*

Area

(sq

statute

miles)

Mean

Annual

Temp.f

(°F)

Mean

Annual

Rainfall

(inches)

Effective

Precipi-

tation!

(inches)

Desig-

nation*

River

Drainage

Area

(sq

statute

miles)

Runofff

(inches)

Maximum
Inst. Flow
(cubic ft/
sec)

No.

% Car-
bonate

Waimea

45.0

372 '

1.3

Hanapepe

20.5

52.1

487

4.2

144

128

100

S. Wail u a

22.4

81.0

265

80.0

N. Wailua

14.6

99.8

2045

80.0

Anahola

5.5

53.5

1010

90.0

134

101

Hanalei

19.2

196.0

1130

62.0

Waioli

1.6

277.0

930

76.0

169

130

Lumahai

7.1

230.0

1090

62.0

Wainiha

14.7

91.9

211

55.0

Hanakapiai

2.6

102.0

1600

85.0

* Refers to Figure 3.

t Adjusted for mean elevation of basin, assuming 3 F. decrease per 1000 ft altitude. Station temperature from Macdonald,
Davis, and Cox (I960: 114).

| Precipitation at a mean temperature of 50 F. required to produce the actual runoff. Runoff obtained from mean annual
rainfall (extrapolation from Langbein et al., 1949: fig. 2); effective precipitation obtained from annual runoff (Langbein and
Schumm, 1958: fig. 1).

§ Annual river discharge divided by the drainage area. River discharge from Territorial Planning Board (1939).

obtained by generalizing from the wave sum-
mary given by the Corps of Engineers (1955:
appendix B) and from specific storm descrip-
tions. such as that of Arthur ( 1948) . Review of
these data suggests that the wave and wind re-
gime can conveniently be classed in the follow-
ing manner (Fig, 1):

1. Northeast Trades: Swell generated by the
prevailing northeasterly winds. These waves are
present all year, but are largest from April
through November. Waves typically have pe-
riods of 5-8 sec, are 4-12 ft in height, and
approach from the east-northeast.

2. Kona Storms: Waves generated by intense
winds, associated with local fronts or Hawaiian
lows of extra-tropical origin. The wind may
vary in direction from south through west to
north, and usually blows hardest from the south-

west. These storms are not frequent, but occur
most commonly from November through
March. Waves accompanying these storms typi-
cally have periods of 8-10 sec, are 10—15 ft
high, and are most intense from the southwest.

3. Winter Swell from Aleutian and Mid-lati-
tude Lows : Waves from this source are inter-
mittent, and can occur throughout the year, but
are most intense from October through May.
Typical waves have periods of 10-17 sec, are
8-14 ft high, and may approach from the north-
west, north, and northeast.

4. Southern Swell: Long, low waves generated
by storms in the southern hemisphere, and most
prevalent from June through September. Typical
waves have periods of 14-22 sec, are 1-4 ft high,
and approach from the southeast through the
southwest.

112

PACIFIC SCIENCE, Vol. XVII, January 1963

5. Hurricane: Waves from tropical storms,
usually approaching from the southeast through
southwest. They are most likely to occur in Au-
gust and September, although occurrence may
be between June and December.

Of the five classes of waves, those generated
by the northeast trades are by far the most per-
sistent source of wave energy for the east and
northeast shores of the island. Undoubtedly these
waves account for the greater abundance of cal-
careous reefs on the windward side of the island.
Waves from Aleutian lows are probably the
dominant agent responsible for recent erosion
along the Napali Cliffs on the northwest coast
of the island. These waves produce a southwest-
erly drift of littoral sediment from the Napali
Cliffs towards Nohili Point and have been a
major influence in the formation of the straight
coast line near Barking Sands (between sample
stations 1 to 7, Fig. 11). The meteorological
conditions giving rise to unusually high waves
of this type in January 1947 are described by
Arthur ( 1948). Waves of this type, with break-
ers estimated to be 20 ft high and periods of
14-16 sec, were observed on the north coast of
Kauai on 30 November 1935.

Southern swell is the most common type of
wave on the south and lee sides of the island.
Although these waves are low, they are almost
continuous during the summer months and pro-
duce a westerly transport of sand from the Wai-
mea River toward the Mana Coastal Plain. Hur-
ricanes, although infrequent, are important, and
the local residents attribute some recent erosion
to the combined effects of the hurricane of 1957
and the tsunami of the same year. Kauai is ex-
posed to tsunami waves accompanying seismic
activity in the vicinity of the Aleutian and Ku-
rile Trenches in the northern hemisphere, and,
to a lesser degree, from tsunami waves generated
off the coast of Chile in the southern hemis-
phere. These catastrophic waves cause considera-
ble coastal erosion and, at times, transport large
blocks of reef material inland (Shepard, Mac-
donald, and Cox, 1950).

SEDIMENTS

Eighty-three sand samples and five beach and
dune rock samples were collected from the near-
shore zone along the coasts of the island. Exami-

nation of the sediment showed that it could
easily be divided on the basis of origin into: ( 1 )
biogenous material derived from the skeletons
of molluscs, coral, calcareous algae, and fora-
minifera; and (2) terrigenous material derived
from erosion of the island’s volcanic formations.
These two sediment types were characterized by
important differences in color, density, and solu-
bility in dilute hydrochloric acid. In general, the
biogenous material appeared to constitute the
predominant sediment in the nearshore region
of the island. On the windward coast, the per-
centage of biogenous material was so high that
the terrigenous fraction was rarely visible to
the unaided eye.

Most samples were composed of medium and
coarse grain sand containing 80% to 95% cal-
cium carbonate of biogenous origin. Analysis by
X-ray diffraction showed that, mineralogically,
the biogenous material was composed of three
components: aragonite, magnesium-rich calcite, !
and magnesium-poor calcite. Terrigenous sedi-
ments were common only on the lee coasts of
the island, especially in the vicinity of and down
current from river mouths. The terrigenous ma-
terial consisted mainly of olivine grains and
sand-size, partially-weathered feldspathic rock
fragments, plus a small percentage of opaque
iron oxides.

A brief description of the size distributions of
the sands follows, together with their biological,
chemical, and mineralogical analyses. Detailed
data on each sample are listed in Table 2.

Size Distribution

The size distributions of all the sediments ;
were determined by the Emery settling tube
(Poole et al., 1951), and some, especially the
coarsest, were also analyzed by sieving. The me-
dian diameter of the particle size distribution
is expressed in microns, and measures of the
standard deviation (sorting) and skewness are
given in the graphic phi notation of Inman i
(1952). Sieving was necessary in order to obtain
the parameters for the coarser samples. Since
sieve and settling tube analyses are based on
different properties of the sediment, the para-
meters obtained by the two methods are only
roughly comparable. For example, the partly
hollow tests of foraminifera are retained in a

TABLE 2

Location, Type, Size Distribution, and Composition of Samples

Littoral Sedimentary Processes — Inman, Gayman and Cox

113

OLIVINE to
FELDSPAR
PEAK HEIGHT
RATIO §

ITS

CARBONATE FRACTION §

Mg-Poor

Calcite

trace

ITN

trace

Mg-Rich

Calcite

c<N

h~-

62.8

% .

Aragonite

'sf

r-.

28.7

cCN

72.8

76.0

86.8

85.4

85.0

89.6

87.3

88.1

86.3

86.1

SIZE distribution!

q q

—.08

(—.026)

—.02

(+.043)

CN 00

O r-1

i'X

l/"N „

CN i-i

o o

<N l-l

o o

nt cn

rT> XT

I x

ssf 1—1
CN CN

I x

r- o

CN CN

1 1

NO CN
CN i-h

i x

1-H 00
CCN O

i x

(00‘0)

£Z‘—

.31

(.310)

ir\
ON 00
m m

.48

(.465)

00 o

l rs l/"N

'T no

beach rock

.61

(.48)

.45

(.42)

.56

(.72)

.38

(.46)

q ws

.56

(.64)

H \T\
ITS ITS

.48

(.49)

Median

Microns

253

(227)

309

(297)

451

(435)

4 73
(451)

651

(637) |

707

(646)

779

(747)

245

(243)

243

(230)

281

(330)

257

(262)

no i r\

CN CN

297

(337)

BEACH

SLOPE,

TANGENT

.067

.085

cCN

.101

.096

TYPE

SAMPLE*

LOCATION

Barking

Sands

Barking

Sands

Barking

Sands

Barking

Sands

Barking

Sands

Barking

Sands

Nohili

Point

Mana

Kokole

Kekaha

SAMPLE

NUMBER

rs-

TABLE 2 (continued)

Location, Type, Size Distribution, and Composition of Samples

114

PACIFIC SCIENCE, Vol. XVII, January 1963

olivine to

FELDSPAR
PEAK HEIGHT
RATIO §

i r\

\r\

CARBONATE FRACTIONS

Mg-Poor

Calcite

trace

Mg-Rich

Calcite

ICV

% .

Aragonite

r^-

[

CARBONATE*

52.9

34.9

32.0

7.7

14.7

4.4

6.9

rCV

00.5

0.7

84.6

86.0

8”06

SIZE distribution!

(TV o

\r\ \r\
o o

f +

ON \T\
cn

VO o
m

i-i <N
CM iH

-• — s

V\ 00

o o

+ +

—.13

(0.00)

—.25

(+•14)

i— i srv
CN O

O VO
r-l O

X s

VO CN
r-H O

(TV

—.02

.46

(•42)

.69

(■40)

.49

(.30)

beach rock

00 XT
VO XT

(TV r~-

xp xr

.53

(.57)

.69

(.67)

.62

(.59)

.81

(.65)

.50

(.50)

s — s

(TV ITV

VO ITV

(TV

.54

irv

(TV

Median

Microns

429

(412)

574

(555)

398

(401)

382

(409)

240

(200)

277

(276)

335

(274)

304

(344)

390

(319)

293

(304)

470

(335)

(TV

824

532

BEACH

SLOPE,

TANGENT

t"-

r^-

TYPE

SAMPLE*

LOCATION

Kekaha

Oomano

Waimea

Makaweli

Landing

Koki

Kuunakaiole

Park

SAMPLE

NUMBER

\T\

vrv

F''-

TABLE 2 (continued)

Location, Type, Size Distribution, and Composition of Samples

Littoral Sedimentary Processes — Inman, GAYMAN and Cox

115

olivine to

FELDSPAR
PEAK HEIGHT
RATIO§

ITS

CARBONATE FRACTION!

Mg-Poor

Calcite

20.2

\r\

100

12.2

YOZ

16.5

18.7

16.8

Mg-Rich

Calcite

69.7

i"-

t/"N

50.1

46.7

53.2

% .

Aragonite

25.8

trace

ITN

33.3

36.8

33.3

30.0

carbonate!

4.2

88.6

91.5

86.7

91.2

78.2

90.0

91.1

93.3

90.0

[ 92.4

size distribution!

'S.

—.17

—.33

—.09

—.22

—.20

r +

—.06

—.21

CCN

—.19

—.15

.46

.32

.58

beach rock

.34

.30

.28

dune rock

beach rock

.90

(•69)

(.35)

.63

.40

.46

.39

.57

Median

Microns

259

289

277

285

295

222

518

(543)

(1240)

162

139

r-'

342

361

500

420

BEACH

SLOPE,

TANGENT

.072

r-'

.096

.085

\r\

r-H

TYPE

SAMPLE*

LOCATION

Hanapepe

Bay

Wahiwa Bay

Nahunakueu

Lawai Bay

Poipu

Makahuena

Kapunakea

Kipu Kai

Nawiliwili

Hanamaulu

Military

Reservation

Wailua

Kapaa

SAMPLE

NUMBER

W"N

TABLE 2 (continued)

Location, Type, Size Distribution, and Composition of Samples

116

PACIFIC SCIENCE, Vol. XVII, January 1963

OLIVINE TO
FELDSPAR
PEAK HEIGHT
RATIO§

CARBONATE FRACTION §

I %

1 Mg-Poor
Calcite

18.6

16.7

23.1

16.1

18.5

23.2

VOZ

12.4

trace

Mg-Rich

Calcite

52.1

57.5

54.9

56.5

53.8

I/O

53.8

54.5

rCN

60.4

62.3

1 1

Aragonite

29.3

25.8

26.6

23.1

26.7

L‘LZ

22.2

25.6

29.8

27.2

26.7

CN
i r\

■pq

«!

93.3

89.7

93.8

92.7

87.9

90.7

95.0

93.3

90.3

88.2

88.5

89.7

75.9

85.4

62.0

SIZE DISTRIBUTIONt

is.

or—

—.06

NO GN

o o

r +

CO NO
m ©

1-1 NT

m oo

<N i-H

+ +

(iv— )

—.30

—.07

—.09

—.29

-St

W"N

.60

» rs

\T\

cn m

W~\ rCi

.45

(.37)

.62

(.44)

.60

(.38)

/—N

v/N

.27

.46

.60

.42

.36

.34

.97

.39

Median

Microns

i r\

490

\r\

l/N

555

(651)

(106)

$8 L

i/N

i r\

(599)

(829)

785

(801)

cCN

669

(807)

(959)

289

335

603

254

242

149

190

BEACH

SLOPE,

TANGENT

ins-

.037

TYPE

SAMPLE*

LOCATION

Kapaa

Kealia

Anahola Bay

Moloaa Bay

Kilauea Bay

Kalihiwai

Bay

Kalihikai

Anini

Hanalei Bay

SAMPLE

NUMBER

co>

ITN

VAN

ICN

r-.

TABLE 2 (continued)

Location, Type, Size Distribution, and Composition of Samples

Littoral Sedimentary Processes— Inman, Gayman and Cox

117

OLIVINE TO
FELDSPAR

PEAK HEIGHT
RATIO§

3.9

2.7

CARBONITE FRACTION §

Mg-Poor

Calcite

trace

Mg-Rich

Calcite

r-'

\rs

r"~

.% .

Aragonite

l/A

carbonate!

74.2

76.2

62.5

10.7

no
nf
i rs

11.7

92.5

90.7

64.4

53.0

85.3

23.8

\r\

size distribution!

(—.09)

—.18

(0.00)

i/~\

NT
i rs

.71

vr\

» rs
us

•43

.39

00 us

rCN rCN

| .66

_ ^ _

Median

Microns

341

330

364

611

r~-

369

L 717

871

819

(707)

369

(309)

344

295

620

242

(3732)

BEACH

SLOPE,

TANGENT

.20

.043

TYPE

SAMPLE*

B, BC

q_i

LOCATION

Hanalei Bay

Lumahai Bay

Wainiha Bay

Haena Point

Haena

Ke’e

Hanakapiai 1

Hanakapiai

Hanakapiai |

SAMPLE

NUMBER

rCN

r~-

i r\
i"-

r"-

r-.

84c

r-~

iil

t d C
<u « <u

ui rt c

„„ a 3

2^-3
a, ° jo
K o

c . S
cf

•s §1

^ g
« - 3

..Pi Oh

tiQ c

a .5

& - e

graphic phi measures (Inman, 1952).

$ Gasometric analysis (Bien, 1952).
§ X-ray diffraction analysis.

118

PACIFIC SCIENCE, Vol. XVII, January 1963

Fig. 5. Size distribution curves for sand samples 9 and 15, showing comparison
between settling tube and sieve analysis. Grain size distribution curves are also
presented for the carbonate and noncarbonate fractions, which make up 53%
and 47%, respectively, of sample 15. The size distribution curves for sample 9
(89% calcium carbonate) show a marked bimodal distribution, which is not
typical of most highly calcareous samples of Kauai. In this case, the bimodal
distribution reflects the differences between the coarse carbonate grains arriving
from the north and the finer, but more abundant, carbonate grains from the east.

coarser sieve fraction than their settling velocity
would indicate. Comparisons between sieve and
settling tube analyses are shown in Figure 5.

The most common beach sediment had a me-
dian diameter of about 350 microns, while the
median size of all samples ranged from 140 to
1400 microns. Although the size distributions
of the samples from the windward and leeward
coasts as a group were not markedly different,
smaller coastal segments did show significant
longshore trends of increasing or decreasing
grain size. There is a tendency towards bimodal-
ity in sediment size distribution, especially
where mixing of sediment from different trans-
port directions occurs southeast of Nohili Point
(Sample 9, Fig. 5). Also, the beach face sedi-
ments from Kauai, which had an average phi
deviation measure of about 0.5, are less well
sorted on the average than those from typically
continental beaches. Well-sorted sand of similar
size from continental beaches would have a phi
deviation measure of about 0.3 (Inman, 1949,
1953). The more frequent interruption of lit-
toral drift by points and headlands, and the con-

tinuous addition of a heterogeneous assortment
of material from reefs and from offshore, prob-
ably account for this difference in sorting.

The slope of the beach face increases with
increasing sand size and with decreasing wave
height and intensity. Beaches like those at Ka
paa, which are protected by broad fringing reefs
from the direct attack of large breakers, were
consistently steeper than more exposed beaches.
Also, beaches inside protected bays (as Station
30, Hanapepe; Table 2 ) were steeper than those
exposed to- the open sea (as Stations 1-7 at
Nohili, and Station 86 at Flanakapiai ) . The re-
lation between beach slope and sand size for
various type beaches is given in Figure 6.

Biological Analysis

The only completely identifiable biogenous
components present in significant quantities in
the sands were the benthic foraminifera Hetero-
stegina sub orbicularis and Amphistegina mada-
gascariensis . Other positively recognizable com-
ponents, including other foraminifera, echinoid
spines, sponge spicules, gastropod shells and

Littoral Sedimentary Processes — Inman, Gayman and Cox

119

MEDIAN DIAMETER, MICRONS

62 125 250 500 1000

8 ~

x 6
o
<

4 -

2 -

— i 1 1 r

• PROTECTED BY FRINGING REEF

X " BAY

• EXPOSED BAY

• • •

• •

XX • • • «

X • • • • •

• • •

• • #

3 2 | 0
MEDIAN DIAMETER, PHI UNITS

2000

JRM

Fig. 6. Relation between beach slope and sand size.
Exposed beaches generally have lower slopes than pro-
tected beaches.

opercula, pelecypod valves, and bryozoan frag-
ments, made up less than 5% of the samples.
Most of the remaining nonterrigenous compo-
nent of the sand consisted of rounded calcareous
grains, sometimes etched, pitted, or stained.

A rough count indicated that Amphistegina
and Heterostegina accounted for up to 20% of
the grains in some of the samples. Since these
foraminifera occurred most commonly in the
size range of 500 to 1500 microns, they were
more abundant in the coarser sediments. Am-
phistegina were most common in the finer half
of this range.

Several indentifiable skeletal or shell frag-
ments, collected from the reefs and beaches of
Kauai, or in one case from Oahu, were examined
by X-ray diffraction. The results and a review
of the literature suggest that, in this area, the
aragonite is secreted principally by corals and

molluscs; the magnesium-rich calcite by forami-
nifera, coralline algae, and echinoids; and the
magnesium-poor calcite by molluscs and by Am-
phistegina madagascariensis.

Chemical and Mineralogical Analysis

All of the sand samples were examined quali-
tatively, and representative samples were also
examined quantitatively to determine the chemi-
cal, mineralogical, and biological composition.
The carbonate content of all of the samples was
determined to an accuracy of about 1%, using a
gasometric technique described by Bien ( 1952 ) .
Mineralogical determinations were made with a
petrographic microscope and by X-ray diffrac-
tion, using a North American Phillips Diffrac-
tometer.

TOTAL CARBONATE: The total carbonate con-
tent varied from 0.5%, in the river bottom sam-
ples, to 95% in some beach samples. Fifty-two
of the samples had carbonate contents greater
than 80%, while only 17 contained less than
50% CaC03. Although high carbonate values
occurred on all coasts, samples from the east
coast were composed almost entirely of bioge-
nous material. The sands showing the highest
carbonate values on the north, south, and Mana
coasts were taken from those beaches which
were most distant from large streams (Table 2).
On the east (windward) coast and on the east-
ern part of the north coast, the presence of large
streams appeared to have little effect on the car-
bonate content of the beach sediments. Low car-
bonate values were typical of the beaches west
of the Waimea River mouth on the south coast
between sample stations 15 and 25 and, to a
lesser degree, in Hanalei Bay on the north coast.

CALCITE: The X-ray patterns of the sand sam-
ples showed that the calcite frequently consisted
of two components, magnesium-rich and mag-
nesium-poor calcite (Fig. 7). A quantitative de-
termination of the amount of magnesium car-
bonate in each of the calcite components (i.e.,
the mineral species of the component) was made
by comparing the peak location of each com-
ponent with the curve published by Goldsmith
et al. (1955: fig. 1). In the Kauai samples, the
magnesium-poor peak represents a calcite con-
taining from 2% to 3% MgC03; the magne-
sium-rich peak represents a calcite containing

120

about 13% MgCO.s. Determination of the
amount of both of the calcite components, and
of the aragonite component in the carbonate
fraction of each sample, was made by comparing
the peak ratios of these components prepared
from known mixtures of the components (Gay-
man, in preparation; Lowenstam, 1954). Al-
though recognized corrections have not been
made, it is thought that the relative quantitative
results are significant.

In all of the samples examined, with the ex-
ception of one dune rock sample (40), the
magnesium-rich calcite was much more abun-
dant than the magnesium-poor calcite. This rock
was obtained from a lithified relic dune which
now forms a headland (sample station 40) of
Kipu Kai Valley, south of Lihue. With this one
exception, magnesium-poor calcite never consti-
tuted more than 25% of the carbonate content
of any sample.

Magnesium-rich calcite was always present in
the unconsolidated samples, and it was fre-

Fig. 7. X-ray diffraction analysis of sample 60,
showing the magnesium-rich and magnesium-poor
peaks and the two aragonite peaks. The sample was
taken in a water depth of 27 ft outside of Kapaa Reef
(Fig. 8).

PACIFIC SCIENCE, Vol. XVII, January 1963

quently the only calcite component found. Mag-
nesium-poor calcite was absent from the north
and Napali coasts, and was abundant only in the
dune rock and on the east coast in the vicinity
of Kapaa Reef. The distribution of magnesium-
poor calcite in the Kapaa area and the anomalous
abundance of this component in the dune rock
(sample 40) are discussed subsequently.

According to Chave ( 1954^) , water tempera-
ture and phylogenetic level are principle factors
in determining the magnesium content of bi-
ogenous calcite. High magnesium content is fa-
vored by warmer water and lower phylogenetic
level. Temperature has less effect on the mag-
nesium content of the higher phylogenetic levels. ,
Blackman and Todd (1959) show magnesium
content in foraminifera to vary with family.
Heterostegina, one of the two common species
of foraminifera in the samples, and two uniden-
tified species of coralline algae taken from Ka-
paa Reef, were X-rayed and found to contain j
only magnesium-rich calcite. Conversely, Am-
phistegina , the other common foraminifera, and
a limpet were found to be composed mostly or ,i
entirely of magnesium-poor calcite.

Aragonite: Aragonite is generally consid-
ered to be indicative of warm water deposition.

It is deposited by three quantitatively significant j
groups (Lowenstam, 1954): calcareous algae, j
scleractinian corals, and molluscs. Samples from
Kapaa Reef of two different species of coral [
( Acropora and a scleractinian coral) and one I1
specimen of gastropod (Conus) were X-rayed
and found to be pure aragonite. One limpet, j
mentioned previously, was found to be predom-
inantly magnesium-poor calcite and only 28%
aragonite. Although the aragonite in the car- ,
bonate fraction ranged from 19% to 45%, no
systematic variation with depth or direction was |
apparent.

TERRIGENOUS: A cursory inspection of the
differences in the terrigenous mineralogy was '
made by microscopic and X-ray diffraction tech-
niques. A limited amount of X-ray data on the
terrigenous components of the samples is pre-
sented in Table 2 in the form of the olivine to
feldspar peak height ratios. These data should I
be regarded as only semiquantitative because i
of the variation in peak height introduced by
preferred orientation of the feldspar crystals.

Littoral Sedimentary Processes — Inman, Gayman and Cox

121

Presumably, a difference in the degree of chemi-
cal weathering accounts for the differences in
mineralogy between the terrigenous sediments
on the windward and leeward coasts. Plagioclase-
rich sands occur near the Waimea River mouth,
while on the north coast the same size grades
of sand are plagioclase-poor and olivine-rich;
however, there appears to be little difference
in the average parent rock in the two source
areas. It seems likely that a more rapid decom-
position of feldspar, relative to olivine, takes
place because the feldspar occurs in smaller min-
eral grains. The plagioclase crystals, found only
in sand-size rock fragments, are several orders
of magnitude smaller than the olivine grains
found in the beach sands. The plagioclase crys-
tals are abundant in medium- and fine-grain
beach sands only in the Waimea area. Appar-
ently the easily decomposable volcanic rock frag-
ments in these size ranges are brought to the
shore in mass only where chemical processes are
at a minimum. On Kauai a maximum in sedi-
ment yield and a minimum in chemical weather-
ing occur only in the arid Waimea drainage
basin.

Beach and Dune Rock

The distribution of beach rock on Kauai is
described by Emery and Cox (1956) in a com-
prehensive study of beach rock in the Hawaiian
Islands. They examined 72 miles of sandy
beaches on Kauai and reported beach rock along
21 miles, or about 13% of the entire coastline.
The total occurrence of beach rock may be even
greater, as the presence of beach rock is fre-
quently obscured by a covering of loose clastic
sand. During the present study, a large outcrop
of beach rock that was surveyed and sampled
(No. 18) at Oomana Point in 1955, was com-
pletely covered with fresh sand when the site
was revisited in 1959. Also, in the vicinity of
samples 38 and 39, it was found that a p4-mile
section of beach rock was covered with a uni-
form layer of fresh sand, about 4 inches thick.
Here the slope and general attitude of the beach
rock was the same as that of the fresh beach face.
Water seeping through the beach face was tasted
and found to be fresh, indicating that its origin
was the fresh water ponded behind the beach
ridge.

In fact, there is some correlation on the is-
land of Kauai between beach rock outcrops and
areas of present or past ponding of fresh water
behind calcareous beach ridges. Although it is
recognized (Emery and Cox, 1956) that there
may be several methods of formation of beach
rock, it appears that an important cementation
process on Kauai is associated with the evapora-
tion of fresh calcium carbonate-rich waters seep-
ing through the beach face. This occurs most
generally when fresh water is ponded in calcar-
eous sediments by sand dunes or beach ridges.
Presumably, the fresh or brackish water flowing
through the porous calcareous sediments be-
comes saturated with CaCOa. Evaporation of
this water on or near the beach face causes ce-
mentation, and rock is formed if the beach is
sufficiently stable. This belief is strengthened
by the presence of cemented dune sands, which
are well above sea level and appear to have
been formed in the same manner as the beach
rock, with which these cemented sands are com-
monly associated.

Dune rock (eolianite) was found on the south-
west coast of the island, northeast of Nohili
Point. Near Nohili the dune rock was poorly
cemented and appeared to be contemporaneous
with the formation of the present dunes at Bark-
ing Sands. In the vicinity of Kipu Kai (sample
40 ) several rocky points, such as Alligator Head,
are composed of well-cemented dune rock, which
is now exposed to wave attack. The Kipu Kai
dune rock is older than the present shoreline.

Thin section studies of beach and dune rock
samples (6, 18, 35, 40, 41) confirm the litho-
logic and biologic similarity of the rock samples
and the recent beach sand. The general structure,
texture, and composition of the beach rocks sug-
gest that they were formed either very recently,
or under the same environmental conditions as
those which are now dominant on Kauai
beaches. Examination of thin sections and X-ray
analysis suggest that the cement is calcitic, in
agreement with the findings of Emery and Cox
(1956).

The only dune rock sample, 40, differed from
the beach and beach rock samples in that the car-
bonate fraction is composed entirely of magne-
sium-poor calcite and the cement was consider-
ably more abundant. It does not seem likely that
the single carbonate mineral species now found

122

PACIFIC SCIENCE, Vol. XVII, January 1963

Fig. 8. Bathymetry and sample locations along Ka-
paa Reef. Dotted areas depict breaker zone on outer
edge of reef platform. The circulation pattern of wave
generated currents over the reef is shown in the inset.
(Bathymetry compiled from Board of Harbor Com-

in the rock results from a monomineralic species ;
in the biogenous material of the original deposit.
The presence of the single carbonate mineral spe-
cies may be most easily accounted for by assum-
ing (1) the inversion of an original aragonite
component to calcite (see Jamieson, 1953);
and (2) alteration of magnesium-rich to mag-
nesium-poor calcite by the diagenetic loss of
magnesium ions from an original magnesium-
rich calcite component (see Chave, 1954£). The
time required for such processes to be carried to
completion, and the present physiographic posi-
tion of this fossil dune, both suggest that it is
considerably older than the neighboring beach

rock.

;

KAPAA — A WINDWARD REEF AND BEACH

Kapaa Reef lies along the windward east coast
of Kauai, and has an overall length of about 2Vi
miles along the north-south trending coastline
in the vicinity of the town of Kapaa. It is a
fringing reef and is divided into northern and
southern portions by an inlet through the reef
where two small streams enter the ocean near
the center of the town. The Kealia River enters
the ocean about 1 mile north of Kapaa, and the
Wailua River enters the coastal embayment to
the south of the reef. Although both rivers have
appreciable runoff, they contribute relatively lit-
tle sand-sized material to the littoral zone.

North Kapaa Reef is approximately 4,000 ft
long, and varies in width from about 400 ft at
its northern end to 1,500 ft at the southern end,
where it is terminated by the inlet (Fig. 8).
The reef platform has an average depth of about
2V2 ft below mean sea level, and the tide has a
diurnal range of 1.8 ft. The inshore edge of the
reef is bordered by a beach of medium- to |
coarse-sized calcareous sand. The beach has an
average height of berm above mean sea level of
about 6V2 ft, and has a total thickness of sand
above the reef platform of about 8 ft. A rem-
nant toe of beach rock, which outcrops inter-
mittently on the reef flat 10-40 ft offshore from
the present beach (Fig. 9), indicates a slight re-
cession of the beach. The seaward portion of the

missioners Survey of 1919; photomosaic by R. M.
Towell Inc., 1950; and miscellaneous soundings in
1957 (?) by H. V. Frasier, Hawaiian Dredging Co.,
and in 1959 by Sam Keala, Jr., of Lihue Plantation).

Littoral Sedimentary Processes- — Inman, Gayman and Cox

123

reef is shallow, and causes waves to break in all
weather. The seaward slope of the reef rises from
a depth of about 15 ft, and is made irregular by
numerous channels and solution and abrasion
cavities (Figs, 9, 10). Beyond the base of the
reef, the sea floor has a gentle slope out to the
60- ft deep terrace. This terrace, which is com-
mon to the submarine topography of Kauai, oc-
curs about 1 mile offshore in the vicinity of
Kapaa. A sea-valley extends from near the inlet
between North and South Kapaa Reefs, across
the shelf and into deeper water (Figs. 2, 8).

The surface of the reef platform is very ir-
regular and consists of a series of low intermit-
tent channels and ridges (described by Munk
and Sargent, 1954) which trend east- west and
resemble in miniature the channels on the reef
at Bikini Atoll. The channels are most pro-
nounced on the northern and central portions
of the reef and, although discontinuous, traverse
the reef from the breaker zone to near the beach,
where they enter the north-south trending de-
pression which commonly exists at the. toe of
the beach.

Littoral Processes on the Reef

The circulation pattern of water over North
Kapaa Reef is from north to south; over that
portion of South Kapaa Reef near the inlet, it

is from south to north ( inset, Fig. 8 ) . The cur-
rents flowing from the north and south reefs
converge on the inlet and result in a seaward
flow of several knots through the inlet. The
anomalous depths of 27 and 38 ft, found rela-
tively near shore in the inlet channel, are prob-
ably maintained free from sand by scour as-
sociated with the seaward-flowing current.

Measurements on North Kapaa Reef by Helf-
rich and Kohn (1957) show that both current
velocity and the southerly component of velocity
increase to the south. They obtained average
values of current of about 1 ft per second on the
central portions of the reef and a maximum flow
of about IV2 ft per second near the Moikeha
Jetty. The most intense southerly flow of water
occurs roughly one-third of the way between the
beach and the seaward edge of the reef; currents
at the toe of the beach were somewhat weaker.

The currents appear to result primarily from
the mass transport of water associated with
waves breaking over the reef. Winds may also
be a factor in driving the currents, but their
effect is minor, as evidenced by the prevailing
northerly flowing current over portions of South
Kapaa Reef. Also, on several occasions, southerly
flowing currents were observed flowing over the
north reef in opposition to brisk winds blowing
from the south.

a-a! section across north kapaa reef

SAMPLE 55 56 59 60

DISTANCE IN FEET

* I

Fig. 9. Section across North Kapaa Reef along line shown on Figure 8. Vertical
exaggeration is 10 x. Survey by Sam Keala, Jr., and Hiro Murashige of Lihue Planta-
tion; outer reef and sea floor offshore from the reef based on soundings by authors.

124

PACIFIC SCIENCE, VoL XVII, January 1963

Fig. 10. Generalized sketch of surge channel on Kapaa Reef. Note sediment fill and ripples in bottom of
channel. Sand in channels is moved shoreward by the small waves, which reform from the mass of water
.surging over the reef as larger waves break on outer reef. Prevailing currents, which flow from north to south
(to the right of the viewer), are relatively ineffective in transporting sand on the reef flat.

Investigations indicate that the transportation
■of sand and the circulation of water over Kapaa
Reef do not follow the same paths or cycles.
While the water circulation is principally from
north to south over the north reef, the east-west
trending channels, which are common on the
reef flat, act as traps for the sand, and prevent
any appreciable southerly transport across the
reef flat by water currents. The floors of the
channels are covered with sand and reef rubble,
in some places to a thickness of several feet. The
sand is rippled by small waves moving across
the reef flat from east to west, resulting in a net
transport of sand toward the beach (Fig. 10).
These low waves, which move sand across the
reef flat, are formed from the mass of water
which surges over the reef. The characteristic
pattern is for waves of several feet or more in
height to break on the outer reef, their momen-
tum carrying a considerable amount of water

over the reef in the form of a surge of white
water. Over the reef flat, the surge first forms
into a single wave crest, which may, in turn,
disperse into a train of several smaller waves as
the wave front moves toward the beach. The
reef waves thus formed have heights in the
neighborhood of Vl to 1 ft, and wavelengths In
the order of 20 ft. Water level and shoals on the
reef control the height of the waves, for waves
higher than the approximate depth of water tend
to break and re-form as lower waves. Observa-
tions made while swimming over the reef indi-
cate that the small waves are solitary in nature
and that their particle or orbital velocity is
greater in the onshore than in the offshore direc-
tion. This differential between onshore and off-
shore particle velocity, that accompanies the
small waves, results in a step by step movement
of sand toward the beach.

The east-west trending channels of the reef

Littoral Sedimentary Processes— Inman, Gayman and Cox

125

flat appear to play a unique role in the cycle of
sand transportation over the reef. They function
effectively as traps for sand that might other-
wise be transported laterally along the reef by
water currents and, at the same time, act as open
conduits for the onshore transportation of sand
by wave action.

Once on the beach, the sand probably under-
goes a slow net southerly drift under the com-
bined action of currents and waves impinging
on the beach face. After migrating to the inlet,
the sand is carried out to deeper water by the
relatively strong currents that flow seaward
through the inlet.

Thus, the transportation cycle of sand on the
reef appears to- include: (a) migration of sand
along the channels in the reef flat, which is in-
duced by wave action; (b) southerly transport
of sand along the beach face and at the toe of the
beach by waves and currents; and (c) seaward
transport of sand through the inlet and into
deeper water by the currents which converge
from the north and south reef and flow seaward.

The sand on the beaches apparently is derived
from organisms living on the reef, and from ma-
terial transported in suspension by waves break-
ing over the reef. Diving observations indicated
that the numerous solution cavities and the gen-
eral rough outer edges of the reef cause con-
siderable turbulence in the breaking and near-
breaking waves. This turbulence increases their
capacity for carrying material in suspension. Un-
doubtedly some reef material also undergoes a
net offshore migration by processes of turbulent
diffusion. Whether this loss is greater or less
than the supply of material to the reef from off-
shore is not known.

The position and growth of the reef, in recent
geologic time, appears to be governed by the
combined effects of land runoff and the water
circulation and sand transportation over the reef
system. Inlets tend to occur in the fringing reefs
where streams enter and bring fresh water and
mud, which are detrimental to the reef building
corals and algae. The reef builders flourish where
wave action is most vigorous. Therefore, the reef
platform becomes more extensive in the areas
not influenced by land runoff. The formation of
a wider reef platform causes more and more
water and sand to cross the reef and flow into
the inlets. The scouring action on the inlets by

water and sand derived from the reef maintains
the inlets even when rivers are not flowing. Thus
it would appear that once the position of a po-
tential inlet is determined by land runoff and
coastal configuration, the pattern of water cir-
culation and sand transportation on the growing
reef will tend to perpetuate the location of the
inlet.

Kapaa Sediments

Samples collected from the beach, reef flat,
and sea floor outside the reef all consisted of very
coarse, coarse, and medium-grain sands, contain-
ing about 9'0% calcium carbonate of biogenous
origin. The following trends in the sediment
were observed from north to south along the
north Kapaa reef flat: (1) the sand becomes
finer, (2) the sands become better sorted, (3)
the magnesium content of the cal cite decreases,
and (4) the foraminifera content in the sand
decreases. In general, the reef flat and offshore
samples were coarser than the beach samples.
These trends, although not necessarily definitive
in themselves, are in agreement with the modes
and cycles of sediment transport as discussed pre-
viously. The samples from the Kapaa area con-
tained a higher percentage of magnesium-poor
cal cite than any of the other unconsolidated sed-
iment samples collected from Kauai. In general,
the content of magnesium-poor calcite increases
southward on Kapaa beach and seaward across
the reef. The distribution of magnesium-poor
calcite is consistent with the supposition that the
supply of beach sand is predominantly from
transportation over the reef opposite the beach,
rather than from transportation along the shore.

The only completely identifiable biogenous
components present in significant quantities in
sands were the foraminifera Heterostegina sub-
orbicularis and Amphistegina madagascariensis,
A rough estimate indicated that these species
made up 3% to 20% of the grains in the sam-
ples. All of the tests were considerably worn,
making it quite difficult to separate the two
species under a microscope. A more accurate de-
termination of their relative abundance, based
on their skeletal composition, was obtained by
X-ray diffraction. In sample 59, about 55% of
the foraminiferal carbonate consisted of magne-
sium-rich calcite, indicating deposition by Het-

126

PACIFIC SCIENCE, VoL XVII, January 1963

Fig. 11. Top: Direction of littoral transport and sample locations along the Mana Coastal Plain. Note
beach rock outcrops, and position of multiple beach ridges and recent and ancient dunes.

Bottom: Variation in carbonate content and median grain size with distance along the Mana shoreline.
Median diameters determined by Emery settling tube.

erostegina. The remaining 45% was magnesium-
poor calcite and presumably is derived from
Amphistegina. Very little is known of the living
depth range of either species. The highly worn
appearance of all of the tests suggest that on
Kauai they do not live on the reef but have been
transported some distance.7

MANA COAST — LEEWARD BEACHES

The Mana Coastal Plain (Fig. 11) consists of
calcareous beach, dune, and lagoon sediments,
mainly of marine biogenous origin, but includ-
ing some terrigenous sands near the Waimea

7 Ruth Todd, U. S. Geological Survey, suggests that
both species may thrive in water down to at least 200
ft (personal communication) .

River. Well borings show that these marine sed-
iments are about 125 ft thick near the base of
the Puu Ka Pele slope, and may be 400-500 ft
thick at the seaward edge of the Mana Coastal
Plain. They lie on a gently sloping bench, ap-
parently cut by waves into the original dome
surface during a lower stand of sea level (Fig.

The region west of the Waimea River, con-
sisting of the Mana Coastal Plain and the Puu
Ka Pele slope, is practically a rainshadow desert.
This region receives an average annual rainfall
of 18 inches (Table 1), the lowest rainfall of
any major drainage area on the island. The lee
coast has few fringing reefs, and those that do
occur are more poorly developed than the reefs
on the exposed windward coast.

Littoral Sedimentary Processes — INMAN, GAYMAN and Cox

127

Before man changed the drainage in this area,
most of the water that drained off of the Puu Ka
Pele slope or that was discharged by springs
from the ground-water body beneath the slope
was ponded by the coastal sand dunes, forming a
fresh to brackish marsh. Now even though the
marsh has been drained, it seems unlikely that
significant amounts of sediment are being sup-
plied to the shore by streams other than the War
mea River. The presence of large calcareous
dunes and the dominant marine nature of the
sediments of the Mana Coastal Plain indicate
the importance of the transport of biogenous
material from the sea to the land. Prograding
of the shoreline of the Mana Coastal Plain, dur-
ing the recent geologic past, is indicated by the
occurrence of multiple beach ridges west of Oo-
mano Point, and at Mana by the lithified dunes
situated two miles inland and parallel to the
present coastal dunes at Nohili Point.

The carbonate content of the samples was
high everywhere except near the Waimea River,
where the sediments were principally terrige-
nous. The carbonate content increased towards
the center of the Mana Coastal Plain, attaining a
maximum value of about 90 % at sample stations
8 and 9 (Fig. 11, top). The median diameter of
the samples ranged from about 250 to 800 mi-
crons, with those near Nohili and Oomano Point
being significantly coarser, probably as a result
of increased wave action on the points.

There was a pronounced bimodality in the
size distribution, both by sieving and settling
tube, of sediments from stations 9 and 10. This
seems to reflect the mixing of sediments from
the Napali-Nohili region with those from the
south coast. The major mode occurs at 180 mi-
crons, with a minor near 700 microns (Fig. 5).
Microscopic examination shows the major mode
to consist of unidentifiable fine, angular, car-
bonate grains; the principal constituent of the
coarse mode was foraminifera, mostly Hetero-
stegina suborbiculcwis. In comparison, sample
15, which consists of approximately equal por-
tions of terrigenous and biogenous material, is
better sorted and shows less tendency towards bi-
modality. Sample 15, which was collected nearly
3 miles west of Waimea River, is from a region
of active littoral transport, where the rate of
sorting approaches the rate of mixing of unlike
materials. Carbonate content and size distribu-
tion of the sediments indicate that the terrigen-
ous material is transported along the beach west-
ward from the mouth of the Waimea River and
southward from the Napali Coast, and that as it
is transported it becomes progressively more di-
luted by biogenous material. The logarithm of
the carbonate content is observed to increase
linearly with distance from the Waimea River
and from the Napali Coast (Fig. 11, bottom).

The increase in carbonate content with dis-
tance from the terrigenous source is indicative

Fig. 12. Section through Mana Coastal Plain (along line shown in Fig. 11). Well numbers and well data
taken from Macdonald, Davis, and Cox (I960). Composition of the submarine portion of the profile is
little known and is based principally on the supposition that the section above the line of original slope of
the dome is undifferentiated sediments, probably deposited by littoral currents at a lower sea level.

128

PACIFIC SCIENCE, Vol. XVII, January 1963

of continuous dilution by the addition of bioge-
nous material. An expression giving the dilution
rate can be derived under the following assump-
tions : ( 1 ) sand transported along the shore con-
sists of a mixture of biogenous and terrigenous
material, where the fraction of each is B and 1-B
respectively; (2) this sand is transported along
the shore at a rate Q which is constant with dis-
tance x along the coast; ( 3 ) biogenous material
is supplied to the beach, from offshore, at a
constant rate F{ per unit of coast length, where
it is completely mixed with the sand transported
along the shore; and (4) there is a net deposi-
tion of this mixed sediment at a rate F0 per unit
of coast length, such that Fj = F0.

Conditions 1 through 4 lead to- the following
general expression for the concentration of bi-
ogenous material in the beach sand as a function
of the longshore transport rate Q, the rate of sup-
ply of biogenous material F1? and distance along
the coast x:

In B-rr-^-x + constant of integration

where In B is the natural logarithm of the
carbonate fraction (see Appendix I). This is a
straight line with slope Ft/Q when B is plotted
on a logarithmic and x on a linear scale (Fig.
11, bottom). If the terrigenous material is sup-
plied to the coast in large quantities and at a
single point, as at the mouth of the Waimea
River, B will be almost zero near the river and
will increase exponentially with distance from
the river — as it is observed to- do.

The relation derived here is useful in that it
not only predicts the form of the concentration
change, but also indicates that, if either the long-
shore transport of sediment Q, or the deposition
rate F is known, the other can be computed. The
equation for the curve of concentration change
with distance from the Waimea River is:

In B= 1.4 x — 4.6

where In B is the natural logarithm of the car-
bonate fraction, x is in nautical miles, and
Fj/Qn 1.4 has the units of nautical miles"1.

Neither F; nor Q are known, but very rough
order-of-magnitude estimations can be made for
both. The rate of accumulation of calcareous sed-
iment on the Mana Coastal Plain, divided by the
length of coast line, gives a measure of Fls while

the sediment yield from the Waimea River
drainage basin provides an estimate for the lit-
toral transport rate Q near the mouth of the
Waimea River.

The volume of sediment in the Mana Coastal
Plain down to a depth of 60 ft below sea level is
approximately 4 x 1010 ft3. If it is assumed that
60% of this is of biogenous origin, and that it
has accumulated along a coastal length of 15
nautical miles during a time interval of 8,500
years8, then the rate of supply of biogenous ma-
terial to the coast, Fi? is about 7000 cubic yd per
mile per year. This gives a value for the littoral
transport rate in the Waimea region of:

Q = Fi/ 1.4-= 5,000 yd3 per year.

The Waimea drainage basin, with an average
effective precipitation of 53 inches per year
(Table 1), would be expected to yield about
420 tons of sediment per square statute mile per
year (Langbeinand Schumm, 1958: fig. 3). As-
suming half of this to be sand-size or larger ma-
terial, and using the conversion from weight to
volume of 60 lbs per ft3 (given by Langbein
and Schumm, 1958), one obtains a total yield of
sand-size material from the Waimea basin of
25,000 yd3 per year. Since the Waimea River
does not have a prograding delta, the yield from
the river must nearly equal the littoral transport
rate Q.

Beach surveys of 1926 and 1950 by the Corps
of Engineers (1955: par. 52) indicated a net
loss of sand from Waimea Beach of 5,000 yd3
per year. The littoral transport rate is probably
greater than the loss, and the Corps of Engineers
estimate it to be about 20,000 yd3 per year. Con-
sidering the nature of the assumptions leading
to the calculations of littoral transport rate in
the preceding paragraphs, values of 5,000 and
25,000 yd3 per year are remarkably, if not for-
tuitously, similar. One must recognize that, in
so far as the actual transport rates at Waimea are
concerned, these calculations may eventually
prove to be merely a mental exercise. If this be
the case, it Is hoped that they have at least served
to Illustrate a valid principle.

8 The rate of 60 ft sea level rise in 8,500 years was
selected because this seems to be the most accurate
and significant part on the sea level rise curves, pre-
sented by Shepard (I960: fig. 4), McFaden (1961:
fig. 9), and Jalgersma and Pannekoek (I960': fig. 3).

Littoral Sedimentary Processes- — Inman, Gayman and Cox

129

ACKNOWLEDGMENTS

This paper represents, in part, results of re-
search carried out by the University of California
under contract with the Office of Naval Re-
search. The original field work was performed in
November and December 1955. Supplementary
field study in January 1959 was supported by the
LI hue Plantation Company, Kauai. Valuable ad-
vice and field assistance during the supplemen-
tary study was given by J. T. Orrick and S. L.
Keala, Jr., of the Lihue Plantation. Since this
study, North Kapaa Reef has been modified by
dredging. Unfortunately the dredging was not
performed entirely in accordance with plans
based on the study of the littoral processes over
the reef as outlined here. A north-south channel
was dredged, which may intercept the sand sup-
ply to the beaches and cause the beach to erode.9

Valuable suggestions and guidance during the
course of the study were contributed by R. S.
Arthur, E. D. Goldberg, F. P. Shepard, E. W.
Fager, and M. N. Bramlette of the University
| of Califoria. Suggestions leading to the section
- on exposure to waves and wind were made by
P. L. Horrer of Marine Advisers, La Jolla. Idem
| tification of foraminifera was made by Ruth
Todd of the U. S. Geological Survey, Washing-
ton. In addition, the writers wish to express their
appreciation to H. S. Ladd and J. I. Tracey, Jr.,
I also of the U. S. Geological Survey, for their
; careful reading of the manuscript, and to Edith
Haselwood of the Hawaiian Sugar Planters’ As-
| sociation for her editorial criticism.

APPENDIX I

DERIVATION OE SEDIMENT TRANSPORT
EQUATION

x = x, p x=x2

BEACH ; y> j

LONGSHORE ........ +

CURRENT V |

- A,, »

* Ax *

OCEAN rr

B; while the sediment Q, transported longshore
consists of two components, a terrigenous frac-
tion T, and a biogenous fraction B = 1 — T If
complete mixing takes place within a coastal
segment, having boundaries xi and X2, a distance
Ax apart, then T is a function of the longshore
distance x and can be written T ( x ) . The above
assumptions permit the budget for the fraction
of terrigenous material to be written:

Q • T(Xl) = Q • T(x2) + F0T(xi +Lr) Ax

^ Q T(xi) T(x2) , , Ax,

0= x + F0T(xi. d — - )

Ax 2

but in the limit as Ax

-> o

~ dT
°=Q

F„T(x)

and integration gives

lnT= — ^-x + k

where In is the base of natural logarithms and k
■ is a constant of integration. The relation may be
written in terms of B = 1 — T as:

A small segment of shoreline (see figure) is
! assumed to be in a state of equilibrium, such
i that the rate of longshore transport of material
into the segment (the longshore influx) is con-
stant and equal to the outflux, Qi = Q0- Also, the
onshore influx and outflux are constant and
equal, F| l- F0. The sediment influx from offi
; shore, F:> consists of 100% biogenous material,

9 Comparison of surveys of 1959 and 1962 indicate
| a net erosion of beach immediately westward of the
dredging area (approximately stations 51-57, Fig. 8)
! of about 2,500 cubic yd per year.

I n B= -q- x + k

REFERENCES

Arthur, R. S. 1948. Forecasting Hawaiian swell
from January 2 to 5, 1947. Bull. Amer. Me-
teor. Soc. 29:395-400.

BlEN, G. S. 1952. Chemical analysis methods.
Scripps Inst. Oceanogr. Ref. 52-58.

130

Blackmon, P. D., and R. Todd. 1959. Mineral-
ogy of some foraminifera as related to their
classification and ecology. J. Paleo. 33 ( 1 ) : 1-
15.

Bramlette, M. N. 1926. Some marine bottom
samples from Pago Pago Harbor, Samoa. Car-
negie Inst. Publ. 344. 35 pp.

Chave, K. E. 1954^. Aspects of the biogeo-
chemistry of magnesium 1, calcareous marine
organisms. J. Geol. 62:266-283.

1954 A Aspects of the biogeochemistry

of magnesium 2, calcareous sediment and
rocks. J. Geol. 62:587-599.

Corps of Engineers, San Francisco District.
1955. Beach erosion control report on coop-
erative study of Waimea Beach in Hanapepe
Bay, Island of Kauai, T. H. (Partially pub-
lished as House Doc. 432, 84th Cong. 2nd
Sess.)

Emery, K. O., and D. C. Cox. 1956. Beach rock
in the Hawaiian Islands. Pacific Sci. 10 (4) :
382-402.

Gayman, W. R. In preparation. X-ray diffrac-
tion calibration curves for ratios of aragonite
and magnesium-rich and magnesium-poor cal-
cite.

Goldsmith, J. R., D. L. Graf, and O. I. Joen-
SUU. 1955. The occurrence of magnesium cal-
cites in nature. Geochim. Cosmochim. Acta.
7:212-230.

Hinds, N. E. A. 1930. Geology of Kauai and
Niihau. Bishop Museum Bull. 71.

Inman, D. L. 1949. Sorting of sediments in the
light of fluid mechanics. J. Sed. Petrol. 19:5 1—
70.

1952. Measures for describing the size

distribution of sediments. J. Sed. Petrol. 22:
125-145.

1953. Areal and seasonal variations in

beach and nearshore sediments at La Jolla,
California. Beach Erosion Board, Corps of En-
gineers, Tech. Memo. 39. 134 pp.

Jamieson, J. C. 1953. Phase equilibrium in the
system calcite: aragonite. J. Chem. Physics
21:1385-1390.

PACIFIC SCIENCE, Vol. XVII, January 1963

Kohn, A. J., and P. Helfrich. 1957. Primary
organic productivity of a Hawaiian coral reef.
Limnology and Oceanography 2:241-251.

Jelgersma, S., and A. J. Pannekoek. I960.
Post-glacial rise of sea-level in the Nether-
lands. Geol. Mijnbouw 39:201-207.

Langbein, B. et al. 1949. Annual runoff in the
United States. U. S. Geol. Surv. Circ. 52. 14

pp.

and S. A. Schumm. 1958. Yield of sed-
iment in relation to mean annual precipita-
tion. Trans. Amer. Geoph. Union 39: 1076-

1084.

Lowenstam, H. A. 1954. Factors affecting the
aragonite-calcite ratios in carbonate secreting
marine organisms. J. Geol. 62:284-322.

Macdonald, G. A., D. A. Davis, and D. C.
Cox. I960. Geology and ground water re-
sources of the Island of Kauai, Hawaii. Haw.
Div. Hydrog. Bull. 13. 212 pp.

McF ARLAN, E., Jr. 1961. Radiocarbon dating of
Late Quaternary deposits, South Louisiana.
Geol. Soc. Amer. Bull. 72:129-158.

Munk, W. H. and M. C. Sargent. 1954. Ad-
justment of Bikini Atoll to ocean waves. U. S.
Geol. Surv. Prof. Paper 260-C: 275-280.

Poole, D. M., W. S. Butcher, and R. L.
Fisher. 1951. The use and accuracy of the
Emery settling tube for sand analysis. Beach
Erosion Board, Corps of Engineers Tech.
Memo. 23. 11 pp.

Shepard, F. P. I960. Rise of sea level along
northwest Gulf of Mexico. Recent Sediments,
Northwest Gulf of Mexico, 1951-1958. Amer.
Assoc. Petr. Geol. 1960:338-381.

, G. A. Macdonald, and D. C. Cox.

1950. The tsunami of April 1, 1946. Bull.
Scripps Inst. Oceanogr. 5:391-528.

Territorial Planning Board, Territory
of Hawaii. 1939. Surface water resources of
the Territory of Hawaii, 1901-1938; Sum-
mary of Records. Honolulu Star-Bulletin Lim-
ited. 411 pp.

NOTE

Adoption of the Metric System and Celsius Scale

The Intergovernmental Oceanographic Com-
mission,

Noting that international adoption of the
Metric System and Celsius Scale should be one
of the major keys of promoting marine sciences,

Considering that the use of the Metric Sys-
tem was recommended by the Working Group
on Data Exchange held in Washington from 7
to 10 August, 1962, (Ref: NS/IOC/2-5(c));

Noting further that data are to be submitted

to the World Data Centres in the Metric System
and Celsius Scale.

Bearing in mind that for purposes of inter-
national exchanges of meteorological data, the
World Meteorological Organization (WMO)
has already adopted the said system at its 3rd
Congress,

Recommends that member countries of IOC
take the necessary steps to encourage, as far as
possible, the use of the Metric System and Cel-
sius Scale in their oceanographic publications.

131

ILLUSTRATIVE MATTER

Manuscript Form. Manuscripts should be typed on
one side of standard-size, white bond paper and
double-spaced throughout. Pages should be consecu-
tively numbered in upper right-hand corner. Sheets
should not be fastened together in any way, and
should be mailed flat. Inserts should be either typed
on separate sheets or pasted on proper page, and point
of insertion should be clearly indicated.

Original copy and one carbon copy of manuscript
should be submitted. The author should retain a car-
bon copy. Although due care will be taken, the editors
cannot be responsible for loss of manuscripts.

Introduction and Summary. It is desirable to state the
purpose and scope of the paper in an introductory
paragraph and to give a summary of results at the end
of the paper.

Dictionary Style. It is recommended that authors fol-
low capitalization, spelling, compounding, abbrevia-
tions, etc., given in Webster’s New International Dic-
tionary (unabridged), second edition; or, if desired,
the Oxford Dictionary. Abbreviations of titles of pub-
lications should, if possible, follow those given in
World List of Scientific Periodicals.

Footnotes. Footnotes should be used sparingly and
never for citing references (see later). When used,
footnotes should be consecutively numbered by supe-
rior figures throughout the body of the paper. Foot-
notes should be typed in the body of the manuscript
on a line immediately below the citation, and sepa-
rated from the text by lines running across the page.

Citations of Printed Sources. All references cited
should be listed alphabetically by author at the end
of the paper, typed double-spaced. References to books
and to papers in periodicals should conform to the
following models :

Batzo, Roderick L., and J. K. Ripkin. 1849. A
Treatise on Pacific Gastropods. Rice and Shipley,
Boston, vii + 326 pp., 8 figs., 1 map.

Crawford, David L. 1920 a. New or interesting
Psyllidae of the Pacific Coast (Homop.). Proc.
Hawaii. Ent. Soc. 4(1) : 12-14.

1920&. The sandalwoods of Hawaii. Proc.

Hawaii. Ent. Soc. 4(2) : 374-375, 13 pis.

In the text, sources should be referred to by author,
date, and page, as follows: "It was noted (Rock,
1916: 18) that . . ” or "Rock (1916: 21-24)
says . . .”

Quotations. Quoted matter of fewer than five printed
lines (about 200 characters) should be given in the
text in the usual form, using double quote marks.
Longer quotations should be set flush with left mar-
gin. The author is responsible for the accuracy of
quoted material.

Numbers. Decimals, measurements, money, percent-
ages, time; enumerations in which any figure is 10 or
over; and isolated enumerations of 10 and over should
be given in Arabic figures, rather than spelled out,
except when the number begins a sentence.

Only the minimum number of illustrations required
to supplement the text will be accepted by the editors.
Reproduction costs of illustrations in excess of the
number allowed by the editors will be paid by the
author.

Artwork for illustrations should be SV2 x 11 inches
or smaller, and it should accompany manuscript, on
separate sheets. Often more valuable than a photo-
graph is a good line drawing.

Figures and Graphs. Copy for figures and graphs
should always be drawn large enough to allow for at
least one-third reduction by the engraver. Copy should
consist of carefully prepared line drawings in one
color only, drawn in India ink on plain white draw-
ing paper or tracing cloth. Co-ordinate paper with
lines in light blue (a color which will not show in a
photograph) may be used; but co-ordinates which
should appear in the finished graph must be drawn
in India ink. If original figures may not be conven-
iently submitted with manuscript, duplicate rough
sketches or photographic prints may be furnished to
aid the editors in their decisions.

It is strongly urged that an indication of scale be
incorporated as a part of all drawings in which mag-
nification and size are critical considerations.

Photographs. Photographs should be chosen for clarity
in portraying essential information. They should be
printed for contrast, on glossy paper, and should be
sent unmounted. They should be identified with serial
number written in soft pencil on the back to corre-
spond with list of captions.

Illustrations will be returned to the author.

Tables. Tabular matter should be kept to a minimum.
Each table, prepared to conform with Pacific Science
style, should be typed on a separate page, and its posi-
tion indicated on the manuscript.

Mathematical Formulas. Complicated formulas cannot
be set by the printers. Authors should submit them
as illustrations.

Captions. Readily identifiable captions for figures,
graphs, photographs, and other illustrative matter
should be supplied on a separate page.

PROOF

Proof should be corrected immediately and returned
at once to Robert Sparks, assistant to the editors.
Authors are reminded that the editors will allow only
a minimum number of corrections on galley proof.
Additions to the printed text and changes in style and
content are not permitted.

All queries on proof should be answered. If cor-
rected proof is not received within four weeks after
being sent to the author, author’s changes cannot be
accepted.

REPRINTS

Reprints or separates should be ordered on the
form provided and returned with author's proof. All
correspondence concerning separates must be directed
to the printer, Star-Bulletin Printing Company, 420
Ward Avenue, Honolulu 14, Hawaii.

APRIL 1963

NO. 2

boo- /

P//7

VOL. XVII

PACIFIC SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

TADAO YOSHIDA, TAKE O SAWADA, and
MASAHIRO HIGAKI
Sargassum at Tsuyazaki, North Kyushu

ALBERT TESTER

Role of Olfaction in Shark Predation

EDMUND S. HOBSON
Feeding Behavior in Three Species of Sharks

ALAN G. LEWIS

Life History of Lepeophtheirus dissimulates

MOHAMMED NIZAMUDDIN
Studies on Udotea indica

RITA D. SCHAFER
Effects of Pollution on Mytilus edulis

C. E. PEMBERTON
Insect Pests of Sugar Cane

UNIVERSITY OF HAWAII PRESS

BOARD OF EDITORS

O. A. BUSHNELL, Editor-in-Chief
Department of Microbiology, University of Hawaii

Robert Sparks, Assistant to the Editors
Office of Publications and Information, University of Hawaii

Thomas S. Austin

Bureau of Commercial Fisheries, Hawaii Area
(U. S. Fish and Wildlife Service)
Honolulu, Hawaii

L. H. Briggs
Department of Chemistry
University of Auckland
Auckland, New Zealand

Ai Kim Kiang
Department of Chemistry
University of Malaya, Singapore

Gordon A. Macdonald
Department of Geology
University of Hawaii

Donald C. Matthews
Department of Zoology
University of Hawaii

Colin S. Ramage

Department of Geology and Geophysics
University of Hawaii

Martin Sherman
Department of Entomology
University of Hawaii

Donald W. Strasburg
Bureau of Commercial Fisheries, Hawaii Area
(U. S. Fish and Wildlife Service)
Honolulu, Hawaii

Albert L. Tester

Department of Zoology and Entomology
University of Hawaii

Miklos F. Udvardy
Department of Zoology
University of British Columbia
Vancouver, Canada

Thomas Nickerson, Managing Editor
Assistant to the University Provost

INFORMATION FOR AUTHORS

PREPARATION OF MANUSCRIPT

( Continued on inside back cover)

PACIFIC SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

VOL. XVII APRIL 1963 NO. 2

Previous issue published March 6, 1963

I _

CONTENTS

PAGE

Sargassum Vegetation Growing in the Sea around Tsuyazaki, North Kyushu, Japan.

Tadao Yoshida, Takeo Sawada, and Masahiro Higaki 135

The Role of Olfaction in Shark Predation. Albert Tester 145

Feeding Behavior in Three Species of Sharks. Edmund S. Hobson 171

Life History of the Caligid Copepod Lepeophtheirus dissimulatus Wilson, 1905

( Crustacea : Caligoida). Alan G. Lewis 195

Studies on the Green Alga, Udotea indica A. & E. S. Gepp, 1911.

Mohammed Nizamuddin 243

Effects of Pollution on the Amino Acid Content of Mytilus edulis.

Rita D. Schafer 246

Important Pacific Insect Pests of Sugar Cane. C. E. Pemberton 251

Pacific Science is published quarterly by the University of Hawaii Press, in January,

(April, July, and October. Subscription price is $4.00 a year; single copy, $1.23. Check
or money order payable to University of Hawaii should be sent to University of Hawaii
Press, Honolulu 14, Hawaii, U. S. A. Printed by Star-Bulletin Printing Company, 420
Ward Avenue, Honolulu 14, Hawaii.

mmm

mu-3

JIM 3

Sargassum Vegetation Growing in the Sea around Tsuyazaki,
North Kyushu, Japan1

Tadao Yoshida,2 Takeo Sawada, and Masahiro Higaki3

Ecological studies on the subtidal marine
vegetation, including the Sargassum community,
have not progressed far because it is harder to
make field surveys in subtidal vegetation than
in terrestrial or intertidal vegetations. Some
depth records for species of marine algae have
been obtained using dredges and other instru-
ments. From these data, we know that there is
considerable vegetation developed at subtidal
depths (cf Ueda and Okada, 1938, 1940).

Walker (1947) carried out extensive studies
of the Laminaria beds around Scotland. His at-
tention has been centered chiefly on productiv-
ity. Underwater observation of seaweeds has
been done from time to time in the course of
the development of diving apparatus. Waern
( 1952) published the results of his underwater
survey of seaweeds. More recently Gilmartin
(I960) carried out a survey of the deeper water
algae of Eniwetok Atoll with the aid of a
self-contained underwater breathing apparatus
(SCUBA). Above all, a work published by Gislen
(1930) is the most remarkable. He proposed a
system of classification of life-forms for benthic
plants and animals, and described many "as-
sociations.” When these works are examined
from a phytosociological point of view it is
noticed that they are prescribed by the state of
progress in terrestrial plant ecology at the par-
ticular time of writing. For example, Gislen
only recognized associations in the concept of
Swedish school and based on data from but a
few quadrats.

Artificial stone beds have been laid down on
shallow bottoms at many places around Japan
for the propagation of useful algae, such as

1 Contribution from the Department of Fisheries,:
Faculty of Agriculture, Kyushu University. Manuscript
received November 23, 1961.

2Tohoku Regional Fisheries Research Laboratory,
Shiogama, Japan.

3 Department of Fisheries, Faculty of Agriculture,
Kyushu University, Fukuoka, Japan.

Gelidium and Undaria. In order to evaluate the
usefulness of these artificial stone beds many
surveys have been carried out on the communi-
ties of algae developed upon them. However,
the data secured from them are too rough to
provide satisfactory knowledge of the marine
vegetation and succession in its populations
(Katada, 1958).

Many concepts and methods for the analysis
of vegetational structure have been proposed
recently in the course of the development of
phytosociology. In contrast to the method of
classifying the plant communities used by
Rraun-Blanquet and others, Goodall ( 1953) in-
troduced a new statistical method for classifica-
tion using positive interspecific correlation be-
tween the species as a criterion. On the other
hand, Whittaker (1956) and others developed
the concept of a vegetational continuum. These
are applicable to the study of marine algal vege-
tations.

A Sargassum vegetation covers large areas of
the upper subtidal region of the warmer waters
around Japan. This vegetation affords a good
habitat or spawning bed to many kinds of fish.
Plants of Sargassum and its allied genera, float-
ing at the sea surface after being detached from
their substrata, are known as "floating seaweeds”
or "nagaremo.” The floating seaweeds and many
kinds of fish exist in a close relationship insofar
as the fish spawning and growth of the larvae
of the fish are concerned. The authors have been
engaged in the survey of the floating seaweeds
for several years, and feel acutely the necessity
for knowledge concerning the Sargassum com-
munity from which the floating seaweeds of Ja-
pan are derived. The authors carried out a
survey on the Sargassum community growing
on the rocky shallow sea around Tsuyazaki,
north Kyushu, Japan, and intend to analyse here
the relationship of species populations to the
gradient of environmental factors.

135

136

PACIFIC SCIENCE, Vol. XVII, April 1963

FIG. 1. Map of Kyushu, showing the position of
Tsuyazaki.

DESCRIPTION OF AREA SURVEYED

The peninsula of Tsuyazaki (Fig. 1) lies at
130° 29' E, 33° 47' N, about 20 km north-
northeast of Fukuoka City, and faces on the
Genkai-nada. The peninsula protrudes from
general coast line of sandy beach, and consists
of hills more than 100 m high. Although these
hills are for the most part composed of palae-
ozoic sedimentary rock, there is a granite zone
on the west side facing the sea, and a basalt
layer lying above the 100 m contour line. Along
the middle of the shore of the peninsula, there
is a sandy beach called "koi-no-ura.” The rocky
area, where the Sargassum community develops,
occupies the northern and southern parts of the
peninsula. An outline of the geological structure
of the area concerned is shown in Figure 2.

Although the nature of the bottom is rocky
where the shore is exposed to strong wave ac-
tion from the intertidal zone to a depth of about
10 m, or of boulders in more sheltered places,
such hard bottoms give way to sand at the rela-
tively shallow depth of 15 m or less. Sandy

bottoms are met in deeper places all around the
peninsula.

On the coast of North Kyushu, tides are semi-
diurnal, and the diurnal inequality of the tides
is not remarkable. The spring tidal range is
about 1.9 m, and 1.4 m at neap tide. According
to the "Coast of Kyushu Pilot” (1947), in the
Kuraraseto passage, lying to the north-northeast
about 10 km from Tsuyazaki, the tidal current
flows towards the north from 4 hr after low
tide to 4 hr after high tide, and towards the
south from 4 hr after high tide to 4 hr after low
tide. The tidal current flowing north reaches a
speed of 2.8 knots. Near the shore of the area
surveyed, the tidal current runs even faster
around Tsuzumi islet, lying to the north of the
peninsula, and may reach a maximum speed of
more than 1 knot. The current moves more
slowly near Sone-no-hana and Kyodomari.

Stronger waves and swells are generated with
north or west winds, influenced by the topogra-
phy of the neighbouring area. Hence, the north
and west sides of the peninsula are more ex-

FlG. 2. Geological map of the peninsula of Tsuya-
zaki (after Takehara, 1937).

Sargassum at Tsuyazaki— -Yoshida, Saw ADA, and Higaki

137

°c

Fig. 3. Seasonal variation of surface temperature
(solid line) and specific gravity (dotted line) of the
sea water for I960, measured at the top of the break-
water for the port of Tsuyazaki.

posed to wave action. The eastern sides of Sone-
no-hana and Kyodomari are more sheltered
from wave action, although there is no method
of determining the strength of wave action,
judging from the results of direct observation
of waves and from the characteristics of the in-
tertidal algal zonation.

The tidal current is faster where the shore is
more exposed to wave action, and vice versa in
the area surveyed.

Surface temperatures rise to about 28 C in
August and descend to about 10 C in January
(Fig. 3). Transparency of the sea water is less
than 15 m in the Secchi disc reading almost
throughout the year. After a heavy rain, low
transparency is sometimes recorded around
Sone-no-hana.

METHODS

Field surveys were carried out during 7 days
between July 23 and August 26, 1959, on 13
survey lines In the area (Fig. 4), as follows. A
survey ship was anchored at the offshore end of
each line. A rope was then strung from the ship
to a rock on the shore. Survey spots were se-
lected along the rope every 5 or 10 m depend-
ing on the inclination of the bottom. One or
two observers, a recorder, and a crew got into a
small boat. The boat halted at each survey spot.
The depth of water was measured with a sound-

ing lead and then a quadrat was laid down. The
quadrat used in this study was a 0.5 m square
iron frame. Observations on the plants found in
the quadrat were made by skin diving, using a
face mask only. First, the percentage of vegeta-
tional cover of the area around the quadrat was
recorded. Then, the degree of cover by each
species of the algae present in the quadrat was
measured. The scale for the measurement of
cover degree was as follows :

+ 0-10%

1 11- 20%

2 21- 40%

3 41- 60%

4 61- 80%

5 81-100%

Number and height of the plants were recorded
when this was possible. The species not present
in the quadrat but found in the surrounding
area were also noted.

The number of quadrats placed along each

Fig. 4. Map of Tsuyazaki peninsula, indicating the
13 survey lines.

138

survey line varied from 10 to 20, according to
the inclination of the bottom; 197 sets of
quadrat data were obtained. All data needed for
analysis were recorded on hand-sort edge-
punched cards to facilitate the compilation of
the data. Readings of depth were adjusted to the
low watermark of spring tide, and the quadrat
of 0 m level indicates the quadrats placed above
the low watermark. The depth represented as 1
m concerns the quadrat level laid down between
low watermark and 1 m deep, and so on.

RESULTS

Stratal Structure of Sargassum Community

Twelve species of Sargassum, one species of
Cystophyllum, and one species of Hizikia oc-
curred in the quadrats surveyed (Table 1). The
fruiting period of these species is revealed by
analysing the data obtained from surveys of
the floating seaweeds (Segawa, Sawada, Higaki,
and Yoshida, 1939) . According to these results
and our observations made at the stations in-
vestigated, all of the species growing there ex-
cept Sargassum ringgoldianum had passed their
fruiting period. Most of the individuals of each
perennial species had lost their long branches
bearing the receptacles, and most of the branches
of the next year class had reached a length of
30-40 cm. Therefore, the vegetation was at its
lowest height for the year. However, S . ring-
goldianum was taller than the rest. Annual

TABLE 1

Species of Sargassum and Related Genera
Observed in Quadrats

S. thunbergii (Mertens) O. Kuntze

S, nigrifolium Yendo

S. hemiphyllum C. Agardh

S. confusum Agardh

S. micracanthum (Kiitzing) Yendo

S. ringgoldianum Harvey

S, fulvellum Agardh

S' patens C. Agardh

S. tortile C. Agardh

S. yendoi Okamura et Yamada ?

S. piluliferum C. Agardh
S' serratifolium J. Agardh
Cystophyllum sisymbrioides J. Agardh

Myagropsis my agr aides (Turner) Fensholt)
Hizikia fusiforme (Harvey) Okamura

PACIFIC SCIENCE, Vol. XVII, April 1963

species such as S. horneri were in a very early
stage of growth. At the time of the year when
the survey was done, therefore, all species of
Sargassum were considered to contribute to- the
same layer, and cover degree of the species was
rather easy to determine. Stratal difference may
be developed among the species of Sargassum
during the period from winter to spring, be-
cause of the difference in the growing and fruit-
ing periods of the species concerned (cf Se-
gawa, Sawada, Higaki, and Yoshida, I960).
Species other than Sargassum occurred mainly
at shallower places, and there was almost the
total lack of a frondose lower layer under the
well-developed layer of Sargassum . The crustose
layer was represented by species of unarticu-
lated corallines. In the area where the upper
layer was open because of the absence of Sargas-
sum, several species of frondose algae other
than Sargassum formed a lower layer. Although
the population of Sargassum became sparse in
deeper places, the society of the lower frondose
strata was not developed. This fact may be at-
tributed partly to the survey time, when many
kinds of algae were extremely depauperate in
growth for the year.

Vertical Distribution of Species

The substratum profile and the vertical distri-
bution of some of the major species of Sargas-
sum are presented ( Fig. 5 ) for each station.

At station l(Fig. 5), the rocky bottom is
evenly sloped to a depth of more than 8 m,
then it gives way to- sand. The dominant species
of Sargassum changes with increasing depth:
5'. piluliferum is predominant at depths less
than 3 m, then S. patens predominates over the
other species on rock surfaces about 3 m deep,
and finally S. serratifolium predominates in
deeper places. On the other hand, at station 6
(Fig. 5), the bottom is of boulders for the most
part and there is some solid rock bottom in
places. The slope of the bottom is lower than
at station 1. While three of the species of Sar-
gassum just mentioned are also present here,
their degree of cover and vertical range do not
coincide with what was found at station 1. As
stated above, the vertical ranges of the species
are not the same among the stations surveyed.
An explanation for these differences is desirable.

Sargassum at Tsuyazaki — Yoshida, Saw ADA, and Higaki

139

Fig. 5. Profile of substratum and distribution of species: ri, S. ringgoldianum; pi, S. piluliferum; to,
S. tortile; fu, S. julvellum; pa, S. patens; se, S. serrati folium.

Vegetational Cover

The percentage of vegetational cover was ap-
proximated by eye for several square meters
around the spot where the frame of the quadrat
was laid down. For this measurement, crustose
algae were not included. As shown in a later
section, algae other than Sargassum were found
mostly in the relatively shallow places and had
only a slight effect on the degree of vegetational
cover in the deeper places.

Variation in average vegetational cover with
increasing depth was determined using the
whole quadrat samples. This mean value is in-
dicated with a dotted line in Figure 6. From this
figure it can be seen that the average vegeta-
tional cover is nearly uniform from the low
watermark to 5 m deep, and then decreases
gradually in places beyond the 5 m depth. Here
only depth of water is taken into consideration
as an environmental factor.

In this study, the 13 stations surveyed are
placed in three groups according to the strength
of wave action. For each of the three groups,

then, the relationship between the depth of
water and average vegetational cover is plotted
in Figure 6 by the three solid lines. At the
rather sheltered stations subject to weak wave
action, the maximum value for average vegeta-
tional cover is found at the 1 m depth. As
deeper water is reached the values fall quickly.
For the areas where wave action is moderate,
vegetational cover reaches its maximum 2-3 m
below low watermark and gradually decreases
with increasing depth. An examination of the
values from the stations exposed to the strong-
est wave action shows the values for average
vegetational cover reach 100% above the low
watermark, then fall and again come to a peak
at the depth of 4-6 m. Algae other than Sargas-
sum contribute to the vegetational cover in shal-
lower places exposed to strong wave action.
Therefore, when the degree of cover for Sargas-
sum only is considered, the depth at which the
maximum value for vegetational cover is found
becomes deeper with increasing strength of
wave action.

140

PACIFIC SCIENCE, Vol. XVII, April 1963

Distribution of Species Populations along
Environmental Gradient

The distribution pattern of the populations
of Sargassum may not be understood when only
the depth of water is taken into consideration as
an environmental factor. The effect of the wave
action on it must be considered at the same
time. The survey stations are classed in three
groups according to the degree of wave action,
as shown above. Average degree of cover of
each species is plotted against depth. For five
major species, the curves of average degree of
cover were superimposed to obtain the graph
represented by Figure 7. The curve for each in-
dividual species overlaps the others and the
peaks of the different curves occur at different
depths. For the places moderately exposed to
wave action (Fig. 7, lower graph), the curve
for S. hemiphyllum has its peak at a depth of
1 m below low watermark, that of S. ringgold-
ianum comes second, and next in order are S.
piluliferum, S. patens, and S. serrati folium, from
the shallowest to the deepest. In places subjected
to stronger wave action (Fig. 7, upper graph),
the order of appearance is the same as above,
but the peaks of the curves for degree of cover
occur deeper, except in the case of S. ringgold-
ianum.

The method of gradient analysis developed

Fig. 6. Variation of average vegetational cover in
relation to depth of water.

Average of whole quadrat data

x X Sheltered

O O Moderate wave action

A A Exposed

Fig. 7. Change of cover degree in relation to depth
of water. (Abbreviation of species names as in Fig. 5.)

by Whittaker (1956) is introduced here to
make the distribution pattern of these Sargas-
sum species more clear. The depth of water and
exposure to wave action are considered as the
principal environmental gradients. The popula-
tion nomograms for six major species con-
structed on this basis are shown in Figure 8. S.
serratifolium, S. piluliferum, and 5". hemiphyl-
lum have their peaks at the stations exposed
to stronger wave action, but their peaks oc-
curred at different depths, in the order given,
with S. hemiphyllum at the shallower end of the
series. The species which occurred more abun-
dantly at the places subject to more moderate
wave action are S. ringgoldianum and S. fulvel-
lum. The former grows more abundantly at
places of 2 m depths where the bottom is hard
and wave action is moderate. It has a tendency
to grow in shallower places with both increas-
ing and decreasing strength of wave action.
This bimodal distribution pattern is peculiar
to this species only. Only young individuals of
S. fulvellum, an annual species, were found
when the investigation was carried out, and the
degree of cover of this species was very low.
Nevertheless a population nomogram of dis-
tinctive pattern was obtained. S. patens and S.
tortile grow abundantly on bottoms more shel-
tered from wave action. However, the popula-
tion nomogram for one species differs more or
less from the others in the position of maximum
value, even if the distribution patterns are
similar to each other. Generally speaking, most
species of Sargassum grow in deeper places as
the strength of the wave action increases.

Sargassum at Tsuyazaki — Yoshida, Saw ADA, and HiGAKi

141

Objective Classification of Sargassum Commu-
nity

Goodall ( 1953 ) proposed a method of divid-
ing vegetation samples into statistically homo-
geneous groups. This method involves prepar-
ing a 2 X 2 table whereon the correlation be-
tween every pair of species is analysed by com-
puting a value from the number of quadrats in
which one or both of two particular species are
present or not. An objective classification can
be arrived at when, with respect to all species
growing in the sample quadrats, no interspecific
correlations of significant level occur within
each group of quadrats, so that the quadrats
may be divided into some homogeneous groups.
The significance of interspecific correlation is
decided by x2 test, preparing the 2X2 table,
as shown below:

SPECIES B

present

absent

Species A

present

absent

a + c

b + d

a + b

C+d

n=a+b+c+d

(ad-bc±n/2) 2 n
(a+b) (c+d) (a+c) (b+d)

Values of x2 are calculated between every pair
of the six species of Sargassum contained fre-
quently in quadrats, and are shown in Table 2.
Only two combinations, i.e., the pairs of S. ring-
goldianum-S. piluliferum and S. tortile-S. pa-
tens, have significant correlation at the 5%
level. According to the first procedure of classi-
fication proposed by Goodall (1953: 46-50),
as a result of the elimination of quadrats con-
taining one species of high frequency in those
four species mentioned above, no species was

found which has a significant interspecific cor-
relation with the others in the final two groups
obtained. However, the two groups obtained
from this procedure are not useful because quad-
rats belonging to one group are considerably
intermixed with the others.

Distribution of Algae Other than Sargassum

Although attention was not centered on the
distribution of algae other than Sargassum, ob-
servational data on those algae were recorded
as often as possible. The species observed in
this survey are listed in Figure 9. The frequency
of quadrats containing algae other than Sargas-
sum is calculated for each depth, and the graph
in Figure 10 is the result. This graph illustrates
that the species other than Sargassum occurred
in all quadrats laid above the low watermark,
but that the frequency of their occurrence in
quadrats decreases sharply with increasing
depth. The range of the species growing at dif-
ferent depths is shown in Figure 9. Dictyopteris
undulata was observed in the deepest quadrat.
The marine vegetation was richest in number
of species at the place exposed to the strongest
wave action, and more abundant in number of
individuals near the low watermark. At deeper
places the density of algae is less and the vege-
tational cover may be provided by the species
of Sargassum alone.

DISCUSSION

With respect to the stratal structure of the
marine algal community, Segawa considered that
three layers are recognized, i.e., the upper, lower,
and crustose layers, when a well-developed vege-
tation is being considered (Katada, 1958). The
upper layer is more than 10 cm in height, and
the crustose layer is composed of algae less
than 1 cm high. At the time of the year when

TABLE 2

x2 — Values between Six Species of Sargassum Frequently Encountered

S. hemiphyllum

S. piluliferum

S. ringgoldianum

S. tortile

S. patens

S. serratifolium

6.093

2.652

4.330

2.923

1.661

S. patens

0.517

7.337

6.391

8.405

S. tortile

2.527

0.045

0.000

S. ringgoldianum

2.197

9-776

S. piluliferum

3.772

Depth In meters

142

PACIFIC SCIENCE, Vol. XVII, April 1963

( V

* \ *4\ U.l^

1.2

* \ A ^

1-0

V .7

^.5

- \<4

— .25

S .hemiphyllum

a 1

Wave action

Fig. 8. Population nomograms for six major species of Sargassum.

this survey was made, all the species of Sargas-
sum found in the subtidal region formed but a
single upper layer, and there was no constituent
in this layer other than Sargassum . Katada re-
ported that Undaria pinnatifida may also be in-
cluded in this layer, intermingled with Sargas-
sum.

As already pointed out, in reference to the
population nomograms in Figure 8, the Sar-
gassum community appears to form a contin-
uum. This is confirmed by the application of
Goodall’s objective method of classification.
Hence it is desirable that our attention be cen-
tered on the distribution of each species popu-

Sargassum at Tsuyazaki — Yoshida, Saw ADA, and Higaki

143

lation in relation to the gradient of environ-
mental factors, rather than centering it on classi-
fying the communities.

The junior author has reported that the verti-
cal distribution of a species of Sargassum is
hardly explained when only the depth of water
is taken into consideration, and that, in addi-
tion, presence or absence of a species at a certain
station may partly be determined by the degree
of exposure to wave action (Yoshida, 1961).
At present, our knowledge is insufficient to
elucidate the detailed effect of many environ-
mental factors on the distribution of the species.
For marine plants, increasing the depth of water
has an important effect on growth in connection
with the diminishing intensity and variation in
the spectrum of the light penetrating into the
water. On the other hand, we must take into
account the other environmental factors, such
as wave action or the turbulence of water.
There is also a resultant complex of several
factors. According to Gessner ( 1955: 198-209),
the movement of water may have a close re-
lationship with the assimilation and respiration
rates of marine algae. It has already been re-

(Codium fragile)

Dictyota dichotoma
Dictyopteris undulata
Pachydictyon coriaceum
Padina crassa
P. japonlca

P. arborescens

Zonaria rtiesingiana
Stypopodium zonale
Undaria pinnatifida
Eisenia bicyclis
(Ecklonia kurome)
Pterocladia tenuis
Chondrococcus hornemanni
Jania sp.

Amphiroa ephedra ea
A. erassi ssirna

A. dilatata

Cora Hina pilullfera
Lithophyllum okamural
Laureneia okamurai
L. undulata

(Plocamiua telfairiae)

Fig. 9. Vertical ranges of distribution of algae other
than Sargassum. Species in parenthesis did not occur
in quadrats.

Fig. 10. Frequency in occurence of algae other than
Sargassum (cf Fig. 9) in relation to water depth.

ported that the distribution pattern of inter-
tidal organisms is greatly affected by the turbu-
lence of water (Southward and Orton, 1954).
In this present study the distribution pattern
of the species of Sargassum is clearly shown by
the two-dimensional gradient analysis improved
by Whittaker (1956). Two principal environ-
mental gradients considered here are (a) the
depth of water, and (b) the exposure to wave
action. In the area surveyed the tidal current
is faster where the shore is exposed to the
stronger wave action, as stated above. It is
hardly possible to treat the two factors, namely,
exposure to wave action and speed of tidal
current, independently.

Determining the lower limit of the marine
vegetation was prevented by the diving ability
of the observer. It is observed around Tsuzumi
islet that the species of Sargassum grow at
places deeper than 10 m, where hard bottoms
are found. At most cf the stations, the hard
bottom gives way to sand before what would
seem to be the lower potential limit for Sargas-
sum growth is reached.

SUMMARY

The Sargassum community found in the
upper part of the subtidal region in the sea
around Tsuyazaki, North Kyushu, Japan was
studied phytosociologically in July and August,
1959. It seems that this community can not be
divided successfully into smaller groups by ap-
plying Goodall’s objective method of using posi-
tive interspecific correlation. The gradient analy-
sis method developed by Whittaker (1956)

144

PACIFIC SCIENCE, Vol. XVII, April 1963

was applied here to the major species of Sar-
gassum. Two principal environmental gradients
considered here are depth of water and exposure
to wave action. The stratal structure of this
community is also noted.

ACKNOWLEDGMENTS

The authors wish to express their sincere
appreciation for the guidance of the late Pro-
fessor S. Segawa throughout this study. Their
heartiest gratitude is due Professor T. Hoso-
kawa, of the Faculty of Science, Kyushu Uni-
versity, and to Professor Maxwell S. Doty, of
the University of Hawaii, for their kind advice
and criticism of the manuscript. The authors’
thanks are also due Mr. T. Furukawa in the
Fishery Research Laboratory, Kyushu Univer-
sity, who assisted them in the course of the
held survey.

REFERENCES

Gessner, F. 195 5. Hydrobotanik, Bd. I.
Deutscher Verlag, Berlin. 517 pp., 291 figs.

Gilmartin, M. I960. The ecological distribu-
tion of the deep water algae of Eniwetok
Atoll Ecology 41(1): 210-221.

Gislen, T. 1930- Epibioses of the Gullmar
Fjord II. Skrifts. K. Svensk. Vetensk. Kris-
tenbergs Zoo. Stat. 4: 1-380.

Goodall, D. W. 1953. Objective methods for
the classification of vegetation, I. The use of
positive interspecific correlation. Austral. J.
Bot. 1(1): 39-63.

Katada, M. 1958. Phytosociological considera-
tion on the artificial stone beds for Undaria
pinnatifida. Aquiculture 5(3): 30-39. [In
Japanese.}

Maritime Safety Board. 1947. Coast of Kyu-
shu Pilot. Tokyo. 338 pp.

Segawa, S., T. Sawada, M. Higaki, and T.
Yoshida. 1959. Studies on the floating sea-
weeds, III. Some considerations on the forma-
tion of the floating seaweeds. Sci. Bull. Fac.
Agr., Kyushu Univ. 17(3): 299-305. [In
Japanese.}

I960. Studies

on the floating seaweeds, IV. Growth of some
sargassaceous algae based on the material
secured from floating seaweeds. Sci. Bull.
Fac. Agr., Kyushu Univ. 17(4): 429-435.
[In Japanese.}

Southward, A. J., and J. H. Orton. 1954. The
effect of wave-action on the distribution and
number of the commoner plants and animals
living on the Plymouth breakwater. J. Mar.
Biol. Ass., U. K. 33(1): 1-19.

Takehara, H. 1937. Geology of the Tsuyazaki
area, Fukuoka Prefecture. Mem. Fac. Tech.,
Kyushu Univ. 12(5): 263-276. [In Japa-
nese.}

UEDA, S., and Y. Okada. 1938. Studies on the
vegetation of the marine algae in the seas of
Japan, with special reference to the depth of
the growing zone. Bull. Jap. Soc. Sci. Fish.
7(4): 229-236. [In Japanese.}

1940. Studies on the vegetation

of the marine algae in the seas of Japan, with
special reference to the depth of the growing
zone (II). Bull. Jap. Soc. Sci. Fish. 8(5):
244-246. [In Japanese.}

WAERN, M. 1952. Rocky-shore algae in the
Oregrund Archipelago. Acta Phytogeogr.
Suecica. 30: 1-298.

Walker, F. T. 1947. Sublittoral seaweeds sur-
vey. J. Ecol. 35 (1-2): 166-185.

Whittaker, R. H. 1956. Vegetation of the
Great Smoky Mountains. Ecol. Monogr. 26:
1-80.

Yoshida, T. 1961. A brief study on the Sargas-
sum vegetation around Ushibuka, west Kyu-
shu, Japan. Jap. J. Ecol. 11 (5): 191-194.
[In Japanese.}

The Role of Olfaction in Shark Predation* 1

Albert L„ Tester2

It IS RECOGNIZED that most if not all species
of sharks possess a keen sense of smell which is
used in detecting dead and wounded prey or
other edible material during their well-known
scavenging operations. The early experiments
of Parker ( 1910) , Sheldon ( 1911 ) , and Parker
and Sheldon (1913) established the role of the
paired nasal organs as olfactory receptors. Parker
(1914) demonstrated directional response in
the smooth dogfish (Must elm canis) and pro-
vided a plausible explanation of how this was
accomplished; he postulated that the two sep-
arated nostrils have the ability to detect small
differences in the concentration of odorous ma-
terials enabling the shark to orient in the direc-
tion of equal stimulation and to head "up-
stream” to the source. This tracking ability is
well recognized by skin divers and fishermen
who have involuntarily attracted sharks by re-
taining speared fish or by discarding trash fish
and offal from their boats.

It seems unlikely that any shark species could
maintain itself entirely by scavenging opera-
tions, except perhaps in areas where man pro-
vides forage such as bait, fish offal, or other
forms of edible garbage. Certainly the larger
species are recognized as active predators which
attack uninjured living prey, including man.
Doubtless, vision plays a predominating role in
locating and tracking normal living prey, and
possibly other senses such as hearing are also
involved. In considering olfaction, attention has

Presented at the Tenth Pacific Science Congress of
the Pacific Science Association, held at the University
of Hawaii, Honolulu, Hawaii, August 21 to September
6, 1961, and sponsored by the National Academy of
Sciences, Bernice P. Bishop Museum, and the Uni-
versity of Hawaii.

1 Contribution No. 179 of the Hawaii Marine Lab-
oratory, University of Hawaii, Honolulu 14, Hawaii.

2 Department of Zoology, University of Hawaii,
Honolulu 14, Hawaii. Manuscript received November
23, 1961.

focused largely on feeding activity induced by
the release of attractive substances such as blood
or other body fluids from the wounds of in-
jured prey. The possible part played by olfaction
in the detection and tracking of uninjured liv-
ing prey has been largely overlooked.

In this paper I will present the results of
experiments on the olfactory response of cap-
tive sharks to extracts of natural foods, to hu-
man materials, and to uninjured living fish in
the hope of clarifying the role of olfaction in
shark feeding activity. The results form part of
an investigation of factors affecting the be-
havior of sharks sponsored by the Office of
Naval Research (Contract Nonr 2736(00),
Project nr 104503) over the period 1959-61.
The work was undertaken at the Eniwetok Ma-
rine Biological Laboratory, Eniwetok Atoll, Mar-
shall Islands, and at the Hawaii Marine Labora-
tory, Oahu, Hawaii. I am grateful to the respec-
tive directors, Dr. R. W. Hiatt and Dr. A. H.
Banner, for laboratory facilities. I am indebted
to personnel of the Atomic Energy Commission
and the Pacific Missile Range Facility for logis-
tic and other help. I am particularly indebted to
those graduate students who have assisted in
phases of the project: Edmond S. Hobson, Su-
sumu Kato, Taylor A. Pryor, and Bori L. Olla.

FACILITIES, MATERIALS, AND METHODS

Eniwetok Marine Biological Station

At Eniwetok, small (18-36 inches) blacktip
sharks (Carcharhinus melanopterus ) and small
(20-36 inches) grey sharks ( C . menisorrah)
were readily caught in shallow water by hook
and line and established in captivity. Holding
facilities consisted of two large concrete tanks
housed in a building and illuminated by over-
head fluorescent lighting (Fig. 1). The tanks
were supplied with running sea water pumped

145

146

PACIFIC SCIENCE, VoL XVII, April 1963

Fig. 1. View of shark tanks at Eniwetok Marine Biological Laboratory. (Photograph by Atomic Energy
Commission.)

from the lagoon at a maximum rate of flow of
about 10 gal per min. Lying side by side with
a common middle wall, the tanks were 50 ft
long, 4 ft wide and 3 ft deep with turning
basins 6 ft in diameter at both ends (Fig. 2).
The tanks could be divided into 5 -ft sections
by gates which slid in notches in the walls. Thus
the sharks could be confined in a compartment
consisting of one or several sections. Observa-
tion booths located midway along the tanks on
both sides or blinds erected elsewhere effectively
concealed the observer from the sharks.

Olfaction tests were conducted on both "nor-
mal” and blinded sharks. The sharks were
blinded after anesthesia in a 1/1000 solution
of MS 222- Sandoz (cf Gilbert and Wood, 1957)
by coagulating the proteins of the aqueous hu-
mor with the diode probe of a "Hyfrecator” in-
serted through the cornea. Proof of blindness

was lack of response to a hand waved close to
the surface as they swam by or lack of response
to the beam of a flashlight directed at their eyes.
Within 1 hr after recovery from anesthesia the
sharks circled their compartment, guided by the
tip of the outstretched pectoral fin which
touched the wall. Within a day they were able
to circle the compartment without this tactile
aide. They soon fed avidly on pieces of fish,
squid, or other food which settled to the bottom
before it was eaten. The sharks would detect
the odor while swimming in mid-water and
would spiral down, converging on the food by
swimming in a figure-8 pattern on the bottom.
Our attempts at blinding sharks with contact
occluders (Mishkin, Gunkel, and Rosvold, 1959)
were unsuccessful, perhaps because of faulty
technique in molding plastic "lenses” and fitting
them to the eyes. In general, the response of

Olfaction and Sharks — -Tester

147

the blinded sharks to olfactory substances was
considerably less variable than that of sharks
with normal vision.

Except during starvation experiments, the
sharks were fed sparingly about once or twice a
week; uneaten food was removed to avoid putre-
faction and an unnecessary increase in olfactory
level.

After investigating several different tech-
niques during the early summer of 1959, a stan-
dard method of testing materials which could
be dissolved or suspended in water was adopted.
With natural foods such as fish flesh, usually 5
gm of material was macerated in a Waring
blendor for 5 min with 250 ml of fresh water;
sea water was not used because essential oils
tended to accumulate in the froth. The material
was then filtered and held in a refrigerator until
used. Prior to an experiment a small quantity
(usually 3.0 ml) of the clear solution was di-
luted with sea water to 25 ml in a test tube and
then further diluted to 100 ml with sea water in
a funnel. Substances other than natural foods
were diluted to various concentrations before
being tested.

Prior to testing, a glass funnel and tube lead-
ing from the observation booth to a point just
below the surface in the center of a test area
(e.g., Fig. 2, Tank 1, D) was filled with sea
water to remove air bubbles; the contents were
maintained by closing a pinchcock. The flow of
sea water to the tanks was shut off at the inflow
valve. Usually, five 2 -min control periods were
then run, during each of which the time (sec-
onds) spent by one or more of the sharks in
the test areas (e.g., Tank 1, C and D) was re-
corded by an electric timer activated by a foot
switch. At the same time, observations were re-
corded of the behavior of the sharks and some-
times of the number of passes or turns in the
test areas. The test material was then introduced
silently while the sharks were at the far end of
the compartment, and the activity of the sharks
was again noted and recorded during five (or
more) 2 -min test periods. The nature of the
response was then categorized as attraction, re-
pulsion, etc. on the basis of the graphed data
and the notes.

A similar technique was used in I960 except
that timing was abandoned in favor of counting
the number of turns. Each of the two test areas

was divided by an imaginary line into halves;
turns in the four half-areas were given weights
of 1 to 4, with the weights increasing toward
the half-area of introduction (Fig. 2). The
graphed "count index” of activity seemed to re-
flect our subjective impression of a response

SEAWATER

Fig. 2. Diagram of shark tanks at Eniwetok Marine
Biological Laboratory.

148

PACIFIC SCIENCE, Vol. XVII, April 1963

more realistically than the "time index.” Special
techniques used in studying the response of the
sharks to living fish will be discussed later.

Haivaii Marine Laboratory

At the Hawaii laboratory several grey sharks
of two species {Car char hinus spp. ) , hammer-
head sharks {Sphyrna lewini ) , and tiger sharks
{Galeocerdo cuvier ), all 5 to 7 ft in length,
were readily caught by set line in the adjacent
waters of Kaneohe Bay. They were established
in large seminatural ponds (Fig. 3), screened
by gates and flushed slowly by tidal action. Three
grey sharks and one tiger shark were success-
fully maintained in captivity for 3 years and
were still alive and healthy at the time of writ-
ing (October 1961). Hammerheads were suc-
cessfully maintained in captivity for periods
from 3 months to 1 year. It is suspected that
their death was due either to injury caused by
the other sharks or lack of food. They were un-
able to compete successfully with the fast, ag-
gressive grey sharks; often our efforts at pre-
ferential hand-feeding failed when the food was
taken persistently by the other species. The

sharks were fed sparingly about once or twice a
week on cut or whole fish.

Several experiments were undertaken on a
tiger shark and a grey shark following their re-
spective establishment in Ponds 2 and 3, both
of which were about 100 ft long, 60 ft wide
and 3-4 ft in maximum depth. Observations
were conducted from a 16-ft tower between the
ponds. By means of a pump and hose a continu-
ous flow of water was taken from one pond,
led to the top of the tower, and thence led into
a test area of the other pond. Following a series
of 3 -min control periods, during which quanti-
tative data were collected on the activity of the
shark, notes were made of overt responses and
the path of the shark was diagrammed. The ma-
terial to be tested was then introduced into the
stream of salt water after dilution in a suction
funnel on top of the tower, and the observations
were repeated during a series of 3 -min test
periods.

During the winter of 1959-60 both the grey
and the tiger shark were transferred to Pond 5
(Fig. 4), a much larger enclosure about 360 ft
long and 66 ft wide. Other grey and hammer-
head sharks were added to this pond; eventually

Olfaction and Sharks— Tester

149

they were confined in two-thirds of its length
by a fence. Observations were conducted on the
shark population from the 16-ft tower which
had been moved to a central location along one
side. Usually the sharks would swim back and
forth along the length of the enclosure in a
channel which averaged about 9 ft in depth. Oc-
casionally the tiger shark and the hammerheads
( but rarely the greys ) would traverse the length
of the pond in shallows 1—4 ft in depth along
the side of the pond opposite to the tower. Two
testing techniques were employed which are
henceforth referred to as "point” and "curtain-
funnel” or "curtain-drum” introduction. In both,
activity was recorded during the usual control
and test periods in a test area 50 ft in length
and extending across the width of the pond.
The area, centrally located in front of the tower,
was marked off by cords which stretched across
the pond and were several inches above the
water surface at high tide.

In "point” introduction the material was con-

tained in a 5 -gal funnel on top of the tower and
was introduced at a point either just below the
surface or at a depth by means of a rubber tube
suspended from a boom (Fig. 5). In "curtain-
funnel” introduction the material passed from
the funnel to a perforated hose running trans-
versely across the bottom of the pond at the
center of the test area and extending part way
into the shallows. In a modification, used in the
spring and summer of 1961 and called "curtain-
drum” introduction, a continuous stream of salt
water was pumped into the hose before and
during control conditions; the stream was then,
switched to a 50' gal drum containing about 40-
gal of sea water together with the test material
After introduction, which usually consumed
about three 3 -min test periods, the flow was
again switched to salt water. With both methods
care was taken to prevent the generation of air
bubbles in the curtain for they produced a vari-
able visual response. By using dye it was found
that the curtain was fairly uniform and rela-

FlG. 4. View of Pond
E. S. Hobson.)

5 at Hawaii Marine Laboratory showing the observation tower. (Photograph by

150

PACIFIC SCIENCE, Vol. XVII, April 1963

Fig. 5. Diagram of the arrangement of test apparatus in Pond 5 at Hawaii Marine Laboratory.

tively stable during periods at or near slack
water. When there was a tidal current the cur-
tain lacked uniformity in concentration and was
irregular in shape; it slowly spread to one end
or the other of the test area; sometimes it spread
in one direction at the surface and in the op-
posite direction at or near the bottom. Normally
the sharks would encounter the curtain of ma-
terial during their passage along the deep chan-
nel. They could avoid it by swimming in the
shallows.

Interpretation of a response

Based partly on quantitative data averaged as
an index of activity or plotted in graphic form,
partly on notes of overt responses, and, in the
case of pond experiments, partly on diagrams of
the swimming paths of the sharks in, out, or
through the test area, the result of each experi-
ment was classified as no noticeable response
(O), sensing only (S), weak attraction (A),
strong attraction (AA), weak repulsion (R),
or strong repulsion (RR). Occasionally these
were supplemented by other descriptions such
as "startled reaction,” "alarm reaction,” "agita-
tion,” etc.

The category "no noticeable response” needs
no further explanation. The category "sensing”
was reserved for a response which consisted of
a sudden start or turn on the part of the shark

on first encountering the test material but with
no other noticeable component suggestive of
either attraction or repulsion. Sensing responses
were obtained with a variety of materials in-
cluding weak acids, bases, and salts. The re-
sponse was interpreted merely as an awareness
of any change in the composition of an other-
wise uniform environment. An "attraction” re-
sponse included an initial sensing followed by a
more or less prolonged hunting response, usually
with rapid swimming, circling at or near the
surface, and occasionally with a "gulping” or
flexing of the jaws as when feeding. The shark
would usually make several excited circles in the
test area on encountering the material, and
would then dash down the tank, returning to
the test area for further circling. Almost in-
variably the average index of activity during
test conditions was higher than during control
conditions. A "repulsion” response included an
initial sensing, but this was followed usually
by rapid departure from the test area, a slowing
of swimming speed, and a "cautious” re-ap-
proach to the test area. Often in subsequent
passes the sharks would turn short of the test
area. A strong repulsion was often accompanied
by shaking of the head or flexing of the gill
slits. Almost invariably the average index of
activity during test conditions was lower than
during control conditions.

Olfaction and Sharks — Tester

151

Unfortunately the response to a given ma-
terial varied considerably in repeated testing
and was often difficult to classify. The problems
of bioassay should not be minimized. The re-
sponsiveness of the sharks both at Eniwetok
and Hawaii seemed to vary from day to day
for unknown reasons despite our attempts to
maintain standard conditions of testing and
feeding. Erratic behavior, frequently encoun-
tered during both control and test conditions, in
some cases could be traced to obvious sources
of disturbance, such as noise, but in other cases
could not be explained. Particularly exasperating
was an occasionally exhibited tendency to circle
at one or the other end of the pond or tank for
long periods of time so that tests could not be
conducted. Even though an attempt was made
to conduct tests only after some reasonable uni-
formity in swim pattern persisted throughout
control periods, there was always the question
of whether or not a subtle change in behavior
pattern was related to the material being tested.
In classifying a response, greater reliance was
placed on overt signs such as sudden turns, cir-
cling, gills flexing, and head shaking than on the
quantitative data. Unfortunately the location and
concentration of the material being tested was
not known precisely during tests and could only
be estimated from the use of dyes after an ex-
periment had been completed. Thus even overt
responses could be related to the test material
only by inference.

Our caution in interpretation is reflected in
the large number of responses relegated to
doubtful categories in the results which follow
and the numerous tests which were conducted
on materials of particular interest.

RESPONSE TO EXTRACTS OF NATURAL FOODS

Experiments were conducted on the response
of the tiger shark to extracts of tuna flesh and
decayed shark flesh and on the response of the
blacktip and grey sharks to a wide variety of
potential foods including tuna, eel, grouper,
snapper, parrot fish, jack, giant clam, octopus,
squid, lobster, fresh shark flesh and skin, and
decomposed shark flesh and skin. In general,
the extracts of all food substances tested could
be classed as attractants, although because of

variability in the response of the sharks some-
times several tests of the same substance were
necessary to establish this conclusion. Because
of this variability it was not possible to make
rigorous comparisons between the attractive-
ness of extracts from equivalent quantities of
the various foods. However, it seemed certain
that extracts from moist- or oily-fleshed fish such
as grouper, tuna, and eel generally resulted in
greater activity than those from dry-fleshed fish
such as snappers. In attempting to determine the
response to nonfood substances, frequently
standard extracts of grouper, tuna, or eel were
used either before or after tests of the other
substances to appraise the sharks’ responsive-
ness.

From Table 1, summarizing the results of
169 tests, it will be noted that the response was
classed as a strong attraction in 59, as a weak
or doubtful attraction in 62, as nil or merely a
sensing in 39, and as a weak or a doubtful re-
pulsion in 9. The last, comprising 5% of the
tests, warrants further comment.

The five instances of apparent repulsion in
the 1959 tests at Eniwetok involved extracts of
little tunny ( Euthynnus yaito ) , yellowfin tuna
( Neothunnus macropterus) , and giant clam
(Tridacna) , and occurred early in the summer
when testing techniques were being developed.
Without doubt the response was related to either
incipient pollution of the tanks or decomposi-
tion of the test materials. Excessive quantities
of extracts were being used and excess food was
not being removed from the tanks; one or both
of these factors resulted in the death of several
sharks in one compartment before the condition
was rectified. Our notes state that the tunny ex-
tract, which had been kept for 9 days, smelt
foul.

The four instances of apparent repulsion in
the I960 tests, involving standard extracts of
eel, again took place early in the summer and
involved not pollution of the tanks but decom-
position of the extract even though it was held
at ice-box temperature. The extract was pre-
pared on July 7, I960. On that and the follow-
ing day tests of both greys and blacktips showed
strong attraction (Figure 6A). On July 13, in
seven tests the responses were indicated as weak
or questionable attraction, sensing only, or nil.
On July 16, the material produced erratic re-

152

PACIFIC SCIENCE, VoL XVII, April 1963

TABLE 1

Response of Sharks to Extracts of Natural Food at Hawaii Marine Laboratory (hml)
and Eniwetok Marine Laboratory (embl)

SHARKS, LABORATORY, AND

RESPONSE*

YEAR

MATERIAL

R-R?

O-S

A-A?

Total

1959

Tiger, HML

Fresh tuna extract

Aged shark extract

1959

Blacktips, EMBL

Various extracts

! 5

Aged shark extract

1960

Blacktips and greys, EMBL

Fresh extracts

Aged eel extract

Aged shark extract

Total

169

* RR, strong repulsion; R-R?, weak or doubtful repulsion; O-S, no apparent response or sensing; A-A?, weak or doubtful
attraction; AA, strong attraction.

sponses, some of which were classed as doubtful
repulsion. The notes indicated that the material
smelt foul. Similar results were obtained with
the same material on July 18. The sharks were
tested with freshly prepared standard eel ex-
tract on July 20 and both species showed a
strong attraction response.

In direct contrast to the above results are those
with extracts of decayed shark flesh, which after
a week in the hot sun smelt particularly foul.
Our material consisted of extract of decayed
hammerhead and of decayed tiger shark tested
on the tiger shark at the Hawaii Laboratory (two
tests) and of extracts of decayed blacktip shark
flesh and skin tested on blacktip sharks at Eni-
wetok ( five tests ) . In addition, we tested black-
tips at Eniwetok on an alleged shark repellent,
supplied by a fisherman, which contained ex-
tract of decayed shark flesh as the principle
component ( six tests ) . We also tested fractions
of extract of decomposed shark flesh which were
supplied by Dr. M. A. Steinberg, Bureau of
Commercial Fisheries Technological Laboratory,
Gloucester, Mass. (11 tests ) . No repellent ef-
fects were noted in any of the tests. On the con-
trary, the majority yielded responses which were
classed as either weak or strong attraction. Our
results with the fractions of extract were in
agreement with those reported by Steinberg
(I960) when his material was later tested on
the lemon shark ( Negaprion hrevirostris) , the

reef shark ( Carcharhinus falciformis ) , and the
bull shark ( Carcharhinus leucas ) at the Lerner
Marine Laboratory, Bimini, Bahamas, B. W. I.

Our results with extracts of decomposed shark
flesh seem to be at variance with those of
Springer (1955), who found that the feeding
of the dogshark ( Mustelus canis ) was consist-
ently inhibited by the presence of decayed shark
flesh. Although several hypotheses might be
formulated to account for the difference in re-
sults, no convincing explanation can be made
at the present time, particularly in view of the
apparent repulsion noted with decomposed eel
and other extracts noted in preceding para-
graphs.

BEHAVIOR OF STARVED SHARKS

In considering shark predation, the questions
arise as to how long a shark can exist without
food and whether its olfactory response is modi-
fied by starvation. Some information on these
points was obtained for small sharks at the Eni-
wetok laboratory.

In 1959, following the summer’s work, AEC
personnel at Eniwetok volunteered to keep track
of the fate of four small blacktips under starva-
tion conditions. Three of the sharks died after
about 2 months in captivity. One survived for
3 months but it was not known to what extent
it had maintained itself by feeding on the sharks
which had died.

ACTIVITY INDEX

CONTROLS

TEST

Fig. 6. Activity index during successive 2-min periods, illustrating (A) response of blacktip and grey
sharks to standard eel extract, and (B) response of starved (normal) and fed (blinded) blacktip sharks to
a 1/1000 dilution of standard grouper extract.

154

In I960, four blacktip and four grey sharks
were starved under close supervision. Of the
blacktips, a 20-inch male died after 36 days, a
28-inch female died after 40 days, a 20-inch
female died after 40 days and a 27-inch female
survived for 43 days and was then fed. Of the
greys, a 28-inch female died after 32 days and
a 36-inch male died after 40 days. Two 30-inch
females were starved respectively for 34 and
46 days and were then fed. During starvation,
the sharks became very thin. Those which died
had difficulty in maintaining their equilibrium
for several hours before death; they could not
be revived by forced feeding. These experiments
show that small blacktip and grey sharks can
survive for more than a month without food
but that some will die after 5 or 6 weeks.

Using standard extract, tests were run at in-
tervals to measure the response activity of both
the starved blacktips and greys and, for com-
parison, that of four blinded blacktips which
were fed two or three times a week. Activity
data are given in Table 2.

The activity index for the fed sharks fluc-
tuated from day to day but showed no trend. In
contrast, the index for the starved sharks showed
a more or less steady increase during both con-
trol and test conditions, particularly in the case
of the blacktips. For the latter, the decrease in
index on the last day is due to the moribund
condition of two of the four sharks. As starva-
tion proceeded, the sharks seemed to become
increasingly restless and to respond with in-
creasing vigor to the standard extract. Tests on
the greys were discontinued after August 5,
1961 because of the death of one shark and the
moribund condition of another (starvation be-

PACIFIC SCIENCE, Vol. XVII, April 1963

gan about 2 weeks earlier than with the black-
tips).

Normally the blinded blacktips were more
sensitive to odorous substances than the black-
tips with normal vision. This situation was re-
versed when the latter sharks were starved. This
is illustrated by one experiment (August 20,
I960) when both the blind, fed and the normal,
starved sharks were tested with a 1/1000 dilu-
tion of standard extract (Fig. 6B). Using 0.003
ml (rather than the usual 3.0 ml) the fed sharks
showed a weak attraction response which did
not differ greatly from mere sensing. The starved
blacktips on the other hand gave a strong at-
traction response which included the usual ex-
cited circling and hunting activity.

There is no doubt that hungry sharks are
much more responsive than fed sharks to minute
traces of odorous substances.

RESPONSE TO HUMAN MATERIALS

In this section are reported the results of tests
on the response of normal and blinded blacktip
sharks and normal grey sharks to human urine,
blood and sweat, and to L-serine, a presumed
component of human sweat. Other materials
such as faeces and vomit were not investigated.

Urine

At Eniwetok, in both 1959 (eight tests) and
I960 (three tests) blacktip sharks were pre-
sented with human urine in quantities ranging
from 3 to 80 ml of whole material. The urine
was sensed, as indicated by a swirl or turn on
encountering it, but there was no other con-
sistent response.

TABLE 2

Activity Index of Fed (Blind) and Starved (Normal) Sharks to Standard Extract at
Eniwetok Marine Biological Laboratory, I960

DATE

FED BLACKTIPS

STARVED blacktips

STARVED GREYS

Controls

Test

Controls

Test

Controls

Test

7/20

29.2

41.2

31.0

40.4

33.8

60.4

7/23

21.2

32.0

31.8

44.2

23.8

38.6

7/29

34.8

37.6

57.4

76.8

52.6

74.2

8/5

29.6

54.0

50.6

71.2

86.3

96.8

8/18

27.2

36.2

56.0

97.0

—

8/25

23.4

32.2

36.4

49.2

—

Olfaction and Sharks— -Tester

155

TABLE 3

Response of Sharks at Eniwetok Marine Biological Laboratory to Human Blood

RESPONSE*

YEAR

MATERIAL AND SHARKS

R-R?

o-s

A-A?

Total

1959

Fresh or aged blood

Normal blacktips

Blinded blacktips

Total

1960

Aged blood (4-6 days)

Normal blacktips

Blinded blacktips

Normal greys

Total

1960

Fresh blood (1-2 days)

Normal blacktips

Blinded blacktips

Normal greys

Total

5 1

* RR, strong repulsion; R-R?, weak or doubtful repulsion; O-S, no apparent response or sensing; A-A?, weak or doubtful
attraction; AA, strong attraction.

Blood

Most authors agree that blood in the water
excites sharks. For example, Whitely (1940)
notes that small blacktip sharks on the Great
Barrier Reef would follow persons who had
scratched their legs on coral and would dog their
footsteps through slightly bloodied water. Bige-
low and Schroeder (1948) remark that if per-
sons in the water are bleeding from injuries
the danger from shark attack may be imminent
and the results may prove fatal. Moreover, they
state that the more voracious of the larger sharks
are excited by blood in the water to such a
degree that they will make ferocious attacks
whether the object be fish, whales, or man, dead
or alive. In contrast, based on experience with
releasing turtle and sheep blood while fishing
for sharks, Wright (1948) concluded on ad-
mittedly weak evidence that blood alone, with-
out the presence of some moving object, did
not release the attack pattern. Steinberg ( 1961 )
reports that a captive lemon shark was not at-
tracted by solution of dried beef blood.

The results of experiments with human blood
in 1959 are included in Table 3. The responses
were much more variable and erratic than those
with food extract. A sensing was at times fol-

lowed by an attraction response and at other
times by an apparent flight reaction and a tend-
ency to avoid the test area. The erratic behavior
was unlikely related to a visual stimulus as it oc-
curred in both the blinded sharks and those
with normal vision. Moreover the quantities
used, even when the techniques were being de-
veloped, were not sufficient to produce notice-
able coloration in the water. It was suspected
that the variability in response was related to the
freshness of the blood.

In I960, 32 experiments were conducted with
human blood. The results are summarized in
Table 3 and are given in detail in Table 7.
Quantities ranged from 0.03 to 6.0 ml of a sus-
pension of 5 ml of whole blood in 250 ml of
sea water. With fresh blood tested within 1 or
2 days after collection, 3.0 ml of the suspension
usually produced a moderate or strong attraction
response with the usual behavior components:
excited circling, swirling and hunting. An attrac-
tion response was obtained with 0.3 ml of the
fresh suspension on several occasions and with
as little as 0.03 ml in one test. By the use of
dye it was estimated that the shark first encoun-
tered the material when it had mixed with Va
to Yi of the volume of the test compartment.
If this dilution is assumed, it may be estimated

156

PACIFIC SCIENCE, Vol XVII, April 1963

that the sharks were attracted to human blood
at a concentration of about 0.1 to 0.01 parts
per million of sea water.

A blood suspension held under refrigeration
for 4 days or longer usually underwent hemoly-
sis and acquired a faint to strong putrid odor.
The aged blood produced erratic results with
blacktip sharks as had been suspected in the
1959 tests. At times there was only a sensing
of the material, an avoidance of the area, or
possibly a slight attraction. At other times there
seemed to be a "startled” or 'alarm” reaction
with speeding from the area such as had been
noticed the previous year. This was classed as
repulsion. With grey sharks, on the other hand,
the hemolyzed blood seemed more consistently
attractive.

Our results prove that fresh blood excites
blacktip and grey sharks and promotes a strong
hunting response. They suggest that decomposed
human blood contains a component which is
repellent to blacktips.

Sweat

A large number of experiments were con-
ducted at both the Eniwetok and the Hawaii

laboratories on the response of sharks to human
sweat. They were stimulated by the observation
of Brett and McKinnon (1954) that water in
which human hands had been rinsed retarded
the upstream migration of salmon and induced
an "alarm” response.

Sweat was collected initially by sponging the
body and wringing the sponge in 500 ml of
sea water. Later, at Eniwetok it was collected
directly from the body as it ran down arms,
chest and abdomen in the hot, humid atmos-
phere of the shark house, and at Hawaii it was
collected in the same way by exercising and
subjecting the body to heat lamps. The material
was tested according to the standard procedures
already described. Each sweat test was usually
followed or preceded by standard extract to ap-
praise the sharks’ responsiveness to a known
attractant.

The results of 29 tests conducted on normal
and blinded blacktip sharks at Eniwetok are
included in summary form in Table 4 and are
given in detail in Table 8. The majority of the
tests yielded results which were classed as re-
pulsion. A weak repellent effect (R) was com-
prised of an initial sensing, followed by a slow-
ing of swimming speed, an apparent wariness,

TABLE 4

Response of Sharks to Human Sweat at Eniwetok Marine Biological Laboratory (embl)

and Hawaii Marine Laboratory (hml)

RESPONSE*

YEAR

LABORATORY AND SHARKS

R-R?

o-s

A-A?

Total

1959

EMBL, Normal blacktips

—

Blinded blacktips

Total

I960

EMBL, Normal blacktips

Blinded blacktips

EMBL, Normal greys

Total

1959-60

HML, tiger, Pond 2

HML, tiger, grey,

hammerhead, Pond 5

Total

1960-61

HML, tiger, greys, Pond 5

Total

* RR, strong repulsion; R-R?, weak or doubtful repulsion; O-S, no apparent response or sensing; A-A?, weak or doubtful
attraction; AA, strong attraction.

ACTIVITY INDEX

CONTROLS

0.3ML

SQUID EXTRACT

J I I— J I I I L

TEST

BLINDED
' BLACKTIPS
8-12-59

j t—i i I i

CONTROLS

TEST

Fig. 7. Activity index during successive 2 -mm periods in two experiments, illustrating the difference in
response of blinded blacktip sharks to human sweat and to food extract.

158

PACIFIC SCIENCE, Vol. XVII, April 1963

TABLE 5

Index of Activity in Successive Rings (No. 1, Center) of a Target Area for a Hammerhead
Shark During Successive Tests of "Tilapia Water’’ and Human Sweat in Pond 5,
Hawaii Marine Laboratory, June 16, 1960

RING NUMBER

TEST

AVERAGE

Controls, sea water

4.5

5.5

14.8

20.7

11.4

Tilapia water

27.3

29-3

22.0

11.8

22.6

Sweat

5.4

8.3

13.1

12.3

9.8

Tilapia water

28.4

36.8

29.4

20.6

28.8

Controls, sea water

15.0

19.1

16.9

20.0

17.7

Tilapia water plus sweat

15.1

30.0

21.2

16.9

20.8

and a tendency to avoid the test area. A strong
repellent effect (RR) included in addition a
rapid exit from the area following sensing,
sometimes accompanied by head shaking. When
no noticeable response occurred it was found
usually that this was also the case with the
known attractant. In the one case classed as
doubtful attraction, the first sweat test which
was conducted, it is likely that initial sensing
was interpreted as attraction. The quantitative
data of Table 8, illustrated for two experiments
in Figure 7, give convincing evidence of a de-
pression of shark activity following the intro-
duction of sweat. In each of the 29 tests, the
time spent in the test areas during test condi-
tions was less than during control conditions;
this is in striking contrast to the results with
standard extract where the reverse is almost
invariably encountered.

The apparent aversion to sweat was exhibited
by both normal and blinded blacktips, possibly
to a greater extent in the latter. The sweat of
one donor (ALT) seemed to be effective at a
roughly-calculated concentration of about 1 part
per million. The sweat of a second donor (TAP)
produced no obvious repellent effects in the two
tests which were made.

Because of the possibility that the sharks in
1959 had become conditioned to associate sweat
with punishment, e.g., from handling, the ex-
periments were repeated during I960, taking
precautions against sweat dripping accidentally
into the water and using fresh sharks, some of
which had not been touched by hand. The re-
sults in I960 were similar to those in 1959
(Tables 4, 9). With the blacktips the majority

of the responses were classed as repulsion; in
the one case of apparent attraction, again the
first test of the season, the sharks had just been
introduced and still exhibited erratic behavior.
There was no noticeable difference in response
between the blinded and normal blacktips. With
the grey sharks an aversion to human sweat was
present in the majority of the tests but it seemed
less pronounced than with the blacktips. There
were no obvious differences in response between
the sweat of three donors. The sweat did not
decrease in potency with aging at room tem-
perature for several days; rather, its repellent
properties seemed to increase but this could
not be established with certainty.

In tests conducted during the winter of 1959—
60 on the tiger, hammerhead, and grey sharks
at the Hawaii laboratory, larger quantities of
sweat (10-40 ml) were used because of the
greater volume of the ponds compared with the
Eniwetok tanks. In most cases the results, sum-
marized in Table 4, showed vague repellent ef-
fects following the introduction of sweat. For
the most part, the sharks displayed only a tend-
ency to avoid the test area (R? ), occasionally
with a sharp veering from the presumed loca-
tion of the material and rapid swimming
through or away from the test area (R). Apart
from these latter overt responses which were
displayed on occasion by all three species, the
only strong repulsion concerned the tiger shark
in one test and consisted of obvious "agitation”
and head shaking. Repellent effects were noted
with the sweat of all three donors ( ALT, RJ, and
ESH) but more consistently with that of the
first two than the last.

Olfaction and Sharks — -TESTER

159

A different technique was employed in tests
with the hammerhead which was particularly
responsive to attractants. The introduction hose
running from the funnel on top of the tower
was submerged just below the surface at the
center of a bullseye target area 32 ft in diameter,
the boundaries of which were judged by eye
from reference points on the bottom. During
control conditions, sea water was introduced
from the funnel. During test conditions the fol-
lowing materials were used in varied sequence:

( 1 ) an attractant consisting of water from the
funnel in which fish ( Tilapia ) were swimming,

(2) 50 ml of sweat mixed with sea water in
the funnel, and ( 3 ) a mixture of the attractant
and sweat in sea water. Activity data on one
test are given in Table 5. Based on records of
the time spent and the path followed by the
shark in the target area, activity was calculated
as the distance- swum in each of four rings of
the target per unit of time. It is apparent that,
in general, activity was greatest with the attrac-
tant, intermediate with the mixture of sweat
and attractant, and least (less than controls)
with sweat alone. The sweat depressed but did
not eliminate the response to the attractant.

By tracing the spread of materials in the pond
with the use of dye and calculating the volume
of sea water involved, it was concluded that the
actual sea water concentration of sweat in the
pond experiments at the Hawaii laboratory were
still considerably less than those used in the
tank experiments at Eniwetok. Additional ex-
periments were undertaken during the early
summer of 1961 using much larger quantities
of sweat (100-400 ml per test) and the im-
proved "curtain-drum” method of introduction.
The results are summarized in Table 4. Despite
the larger quantities of sweat which were used
no strong repulsion was noted. In the five tests,
there was weak or doubtful repulsion in three
and sensing only in two. In those tests indicat-
ing repulsion, all three species of sharks, espe-
cially the tiger, showed definite signs of aversion
including veering from the curtain and gill
flexing. The sweat of one donor (sk) seemed
to be more active than that of the other (blo) .

From the above experiments on blacktip,
grey, tiger, and hammerhead sharks one cannot
conclude' that human sweat, per se, is an active
shark repellent. On the other hand, it is certain

that human sweat does contain, at least at times,
a component which is aversive to sharks. Oc-
casionally this induces overt signs of repulsion
such as head shaking, gill flexing, veering, and
rapid retreat; more frequently it induces only
a subtle wariness manifested chiefly by avoid-
ance of the area of introduction. The response
is highly variable. This is unfortunate but almost
inevitable when one considers the uncontrolled
environmental conditions and the many factors
which could contribute to both the variability
of shark behavior and variability of sweat com-
position.

Steinberg (1961) found no evidence of re-
pellent properties in either human sweat or pure
compounds forming constituents of human
sweat in tests with a captive lemon shark at the
Lerner Marine Laboratory. Unfortunately he
gives no information on the concentrations of
material used. Moreover, he reports that the
lemon shark was not responsive to solutions of
dried beef blood nor would it eat chunks of
fresh shark liver which, at other times, had been
particularly attractive to captive sharks. His
negative results are understandable. It has been
our experience that sharks which have not yet
fed in captivity do not respond to either highly
attractive substances such as eel extract or fresh
human blood, nor do they respond to subtle
repellent substances such as human sweat.

In view of the results of this series of tests, it
seems safe to assume that shark attack on hu-
mans is not motivated by the smell of human
sweat.

L-serine

Following the discovery by Brett and Mc-
Kinnon ( 1954) that human hand rinse retarded
the migration of salmon, Idler, Fagerlund, and
Mayoh (1956) undertook an analysis of hand
rinse to determine the repellent component ( s ) .
By employing various fractionation techniques
and testing the fractions on migrating salmon,
they were able to identify the active fractions as
amino acids of which serine was a major com-
ponent. In further tests, the L-isomer of serine
was found to induce the alarm response whereas
D-isomer did not. They stated "L-serine defi-
nitely elicited a typical alarm reaction but the
effects were neither so dramatic nor so long a
duration as the response obtained by hand

160

rinse.” As hand rinse doubtless contained sweat
and as sweat produced a repellent effect on
sharks, it was decided to investigate their re-
sponse to L-serine.

Three tests were conducted at the Hawaii
laboratory on the tiger shark in Pond 2, during
the winter of 1959-60, using 1.0 gm of L-serine
per test. In the first, there was no overt response
except an obvious sensing on encountering the
material. In the second, the shark showed a
sensing of the material, followed by rapid exits
from the area and violent head shaking. In the
third test, there was no noticeable response. In
all three, however, the time spent in the test
area during test conditions was less than during
controls, as had also been the case with sweat.

Three tests were conducted on blacktip sharks
at the Eniwetok laboratory in I960, using 6
ml of a solution containing 1.0 gm of L-serine
(i.e., 0.12 gm per test). No repellent effects
were noted other than a "wariness” in one test.
Again, however, the time spent in the test area
during test conditions was less than during con-
trols.

As definite repellent results had been noted
in one test with the tiger shark, it was decided
to run a third series at the Hawaii laboratory
during the summer of 1961, using much larger
quantities of L-serine despite its high cost. Three
tests were conducted on the tiger and grey
sharks co-inhabiting Pond 5, using the "curtain-
drum” technique.

In all three tests the tiger shark displayed an
aversion to the chemical but only after intro-
duction of the material had been completed. In-
troduction required about 10 min (three to
four 3-min periods). It seemed either that the
response occurred after a threshold concentra-
tion of the material had been reached, or that
there was a latent period between exposure to
the material and response. The overt response
was a violent head shaking either while In or
while leaving the area of concentration. At
times this took place at the surface and caused
considerable splashing. However, in only the
first test (25 gm L-serine) was there frequent
rapid exit from the area on encountering the
material. In the second test (50 gm L-serine)
there was swerving and head shaking after en-
countering the material, but no turning-back
on initial encounter. The response was less pro-

PACIFIC SCIENCE, Vol. XVII, April 1963

nounced than in the first test even though twice
the quantity of material had been used. This
may have been due to a higher tide and thus
a larger volume of water and greater dilution
of the material despite the larger quantity used.
The shark frequently avoided the material by
swimming in the shallows on the far side of the
pond. In the third test (25 gm L-serine plus
10 ml of a 99% nicotine solution), the tiger
shark again displayed agitation and head shak-
ing. The response, however, occurred less fre-
quently than in the other two experiments. The
nicotine had been added in the hope of a syn-
ergistic effect; it had been our impression that
the sweat of smokers was more repellent to
the sharks than that of nonsmokers. Possibly
it tended to inhibit rather than Increase the ef-
fect of L-serine.

The grey sharks, in contrast to the tiger shark,
were not obviously agitated by L-serine; no head
shaking or gill flexing was observed. In the first
test, two grey sharks of the same species veered
sharply on first encountering the material and
returned to the end of the pond. Thereafter all
three grey sharks circled in the end zone for
the duration of the experiment. In the second
test no veering was noted but there was re-
peated circling in the end zone. It was uncertain
whether this could be interpreted as a repellent
effect, for the same habit was noted occasionally
during control periods. In the third test, all three
grey sharks passed through the test area without
signs of awareness, agitation, or repulsion.

Although there is no doubt that the tiger
shark was actively repelled by L-serine, the phys-
iological mechanism producing the response is
unknown. We can offer no satisfactory expla-
nation of the difference in response of the tiger
and the greys to L-serine. It may have involved
species differences in physiological effect or
differences in the concentration of materials
to which they were subjected. The latter is
possible even though the tests were conductd
simultaneously on the tiger and the greys, for
uneven curtains of material were formed by
tidal currents in all three tests and the shallow-
swimming tiger shark may have encountered
different concentrations than the deep-swim-
ming grey sharks. It may be added, however,
that directly opposite results were obtained with
a highly irritating lachrimator which is presently

Olfaction and Sharks — Tester

161

Fig. 8. View of a tiger shark attempting to swallow a spiny puffer. (Photograph by E. S. Hobson.)

being tested as a potential shark repellent. With
this substance, the greys responded violently
with gill flexing, head shaking, and definite
avoidance of the curtain. The tiger shark, on
the other hand, was not noticeably affected until
the concentration of the material had been dou-
bled.

In the foregoing sections it has been shown
that certain species of sharks have an aversion
to and at times are repelled by aged food ex-
tracts, aged human blood, fresh or aged human
sweat, and finally L-serine. Serine, presumably
the D-isomer, is a common amino acid in both
foodstuffs and blood. It may by hypothesized
that at least one of the repelling substances in
all of the above materials is L-serine, which is
presumably present as the L-isomer in human
sweat, but which may be formed in foodstuffs

and blood from the nonrepellent D-isomer dur-
ing decomposition.

It has not been demonstrated that L-serine
has sufficiently active repellent properties to
deter shark attack on prey, including man. How-
ever its repellent properties, its presence in hu-
man sweat, and its possible generation during
the decomposition of foodstuffs and blood war-
rant further investigation.

RESPONSE TO LIVING FISH

Although at times sharks may obtain a large
portion of their food by scavenging dead ma-
terials, they also feed on living prey. When the
prey is wounded, the sharks are doubtless at-
tracted by the odor of body juices as well as
by visual and possibly by other stimuli. It is

162

PACIFIC SCIENCE, Vol. XVII, April 1963

reasonably certain that most species also attack-
healthy, undamaged, living prey, although apart
from attack on man, observations of feeding
activity are singularly lacking in the literature.

In the summer of 1959, a group of biologists
from the Hawaii Marine Laboratory witnessed
the persistent attack of a tiger shark on a spiny
puffer which had inflated itself and was floating
at the surface in Kaneohe Bay. The shark’s at-
tempts to swallow the puffer (Fig. 8) lasted
for about 10 min despite the presence of the
observers who circled in an outboard motor
boat. During the shark’s slow, awkward passes
at the puffer the sound of its jaws clamping
together as it missed the prey could be heard.

Other species of sharks are capable of catch-
ing fast moving prey. For example, Eibl-Eibes-
feldt and Hass (1959) observed both the grey
shark ( Carcharhinus menisorrah ) and the black-
tip (C. melanopterus ) actively feeding on
healthy fish in the Indian Ocean, and even herd-
ing them against the shoreline to facilitate cap-
ture.

Although vision is doubtless the predominat-
ing sense which is used by sharks on converging
on living undamaged prey, it is possible that
olfaction may also be involved. I have found
only one observation in the literature which
supports this possibility, that reported by Shel-
don (1911) and again by Parker and Sheldon
(1913), who found that the dogshark ( Mustelus

canis) was able to locate undamaged living
crabs concealed in a wrapping of eelgrass. The
response of sharks to living, presumably un-
damaged fish was investigated at both the Eni-
wetok and the Hawaii laboratories.

Results

In one series of experiments at Eniwetok in
1959, an empty wire cage (about 6 X 6 X 12
inches) was silently lowered to the bottom of
a test area at the upstream end of a compart-
ment containing four blinded blacktip sharks.
After the usual series of control periods during
which activity was recorded, the cage was re-
moved, a living fish was added, and it was again
lowered into the test area when the sharks were
at the far end of the compartment. Activity was
again observed during a series of test periods.
The water flow was maintained during both
control and test conditions.

The results are included in Table 6. In most
of the experiments attraction responses were
obtained with a 12 -inch grouper ( Epinephalus
fuscoguttatus ) , an 8-inch squirrel fish (Holo-
centridae), and an 8-inch stone fish ( Synancaja
verrucosa) . Although probably excited by con-
finement in the cage, the fish did not move about
much after the cage had been lowered. There
was often a delay of several test periods before
the sharks showed any response. Then, in most
of the experiments, one or more sharks suddenly

TABLE 6

Response of Sharks at Eniwetok Marine Biological Laboratory to Living Fish, 1959

RESPONSE*

SHARKS AND FISH

R-R?

O-S

A-A?

Total

Blinded blacktips

Caged grouper

Caged squirrel fish

Caged stonefish

Total

Blinded blacktips

Grouper water, grouper present

Grouper water, grouper absent

Eel water, eel present

Blacktip water, blacktip absent

Total

* RR, strong repulsion; R-R?, weak or doubtful repulsion; O-S, no apparent response or sensing; A-A?, weak or doubtful
attraction; AA, strong attraction.

Olfaction and Sharks — Tester

163

TABLE 7

Response of Normal Blacktip Sharks (nb), Blinded Blacktip Sharks (bb), and Normal Grey
Sharks (ng) to Human Blood at Eniwetok Laboratory, I960

DATE

(1960)

TIME

TANK

SHARKS

BLOOD

(ML)

DONOR, DATE

ACTIVITY INDEX

RESPONSE *

Controls

Test

7/12

0954

I (F-I)

2BB

3.0

JK 7/8

26.0

9.0

1040

II (A-C)

2NG

3.0

JK 7/8

13.0

20.2

1108

I (A-D)

2NB

3.0

JK 7/8

57.8

39.0

1445

I (F-I)

2BB

6.0

JK 7/8

21.6

6.6

1515

II (A-C)

2NG

6.0

JK 7/8

19-8

24.6

1540

I (A-D)

2NB

6.0

JK 7/8

15.6

13.2

7/18

1529

I (F-I)

2BB

3.0

JK 7/8

21.6

24.6

1552

I (A-D)

2NB

3.0

JK 7/8

37.2

17.8

1645

II (A-C)

2NG

3.0

JK 7/8

11.0

10.0

7/21

1015

I (F-I)

4BB

3.0

ALT 7/20

22.8

35.6

1045

I ( A-D ) |

4NB

3.0

ALT 7/20

27.7

26.4

1135

II (A-C) i

4NG

3.0

ALT 7/20

37.4

49.8

1350

I (F-I)

4BB

3.0

ALT 7/20

30.4

39.2

1415

I (A-D)

4NB

3.0

ALT 7/20

21.8

28.0

1450

II (A-C)

4NG

3.0

ALT 7/20

33.7

44.2

7/22

1015

I (F-I)

4BB

0.3

ALT 7/20

26.4

41.6

1045

I (A-D) |

4NB

0.3

ALT 7/20

31.6

40.0

1135

II (A-C)

4NG

0.3

ALT 7/20

24.6

35.2

1405

I (F-I)

4BB

0.03

ALT 7/20

30.6

35.0

7/26

0910

I (F-I)

4BB

3.0

ALT 7/20

22.8

25.0

1000

I (A-D)

4NB

3.0

ALT 7/20

51.0

46.4

1135

II (A-C) j

4NG

3.0

ALT 7/20

28.9

40.8

2035

I (F-I) j

4BB

3.0

ALT 7/20

34.6

36.8

2150

II (A-C)

4NG

3.0

ALT 7/20

31.0

36.8

7/27

0925

I (F-I)

4BB

0.3

SK 7/26

23.6

27.6

1.0

SK 7/26

—

22.6

3.0

SK 7/26

—

31.8

1045

I (A-D)

4NB

0.3

SK 7/26

45.6

46.2

1.0

SK 7/26

—

56.0

3.0

SK 7/26

—

57.2

1500

II (A-C)

4NG

1.0

SK 7/27

46.8

53.2

3.0

SK 7/27

—

52.2

* RR, strong repulsion; R, weak repulsion; O, no apparent response except sensing; A, weak attraction; AA, strong
attraction.

became excited and engaged in the typical hunt-
ing response. It is assumed that they were stim-
ulated by odors emanating from the fish. In
view of the delayed response, it seems unlikely
that the blinded sharks were attracted by vibra-
tions or sounds that may have been made by the
fish although this possibility could not be ruled
out in these rather crude experiments.

In another series of experiments at Eniwetok
in 1959, an uninjured fish was held in a bucket
of saltwater for 15 to 20 min prior to an ex-
periment. The bucket was slapped or agitated

Fig. 9. Diagram of apparatus used at Eniwetok Ma-
rine Biological Laboratory for investigating response
of sharks to living fish.

164

PACIFIC SCIENCE, Vol. XVII, April 1963

to keep the fish in an excited state. During con-
trol conditions, water from another bucket of
sea water was siphoned into the test area of the
compartment containing the four blinded black-
tip sharks. The siphon was then switched to the
bucket of water which either contained the fish
or from which the fish had been removed. The
usual observations were made during a series
of control and test periods.

The results are included in Table 6. The

blinded blacktip sharks showed an attraction re-
sponse in most of the experiments with the
"grouper water” and in the one experiment with
"eel water” ( Gymnothorax ). It was concluded
that the water in which these fish had been con-
fined contained some substance which was at-
tractive to the sharks. The blinded blacktips did
not show a noticeable attraction response to
"blacktip water.”

During the winter of 1959-60 similar ex-

TABLE 8

Response of Normal Blacktip Sharks (nb) and Blinded Blacktip Sharks (bb) to
Human Sweat at Eniwetok Laboratory, 1959

DATE

(1959)

TIME

TANK

SHARKS

SWEAT

(ML)

DONOR, DATE

ACTIVITY INDEX

RESPONSE*

Controls

Test

8/8

1905

I (A-D)

3NB

0.3

ALT 8/8t

97.7

65.9

1930

I (A-D)

3NB

0.3

ALT 8/8+

—

73.7

2115

I (F-I)

3BB

0.3

ALT 8/8+

67.6

52.7

8/9

1800

I (F-I)

2BB

3.0

ALT 8/8+

29.4

14.1

1945

I (A-D)

2NB i

3.0

ALT 8/8+

54.4

41.7 |

8/10

0845

I (A-D)

2NB

3.0

ALT 8/8+

54.0

52.2

0925

I (F-I)

2BB

3.0

ALT 8/8+

45.0

26.8

8/11

2322

I (F-I)

2BB

1 2-3

TAP 8/11

39.4

25.0

8/12

0035

I (A-D)

2NB

1 2-3

TAP 8/11

70.0 j

58.8

1910

I (F-I)

2BB

3.0

ALT 8/8+

29.5

9.3

2010

I (A-D)

2NB

3.0

ALT 8/8+

73.6

61.9

8/13

1553

I (F-I)

2BB

3.0

ALT 8/8+

33.0

5.9

1710

! I (A-D)

2NB

3.0

ALT 8/8+

68.2

58.5

8/14

1905

I (F-I)

2BB

3.0

ALT 8/14

54.0

| 35.3

! RR

2025

I (A-D)

2NB

3.0

ALT 8/14

67.0

57.7

8/16

1900

I (A-D)

2NB

0.0 6

ALT 8/14

80.8

77.8

1930

I (A-D)

2NB

3.0

ALT 8/14

—

74.2

2010

I (F-I)

2BB

3.0

ALT 8/14

9.2

5.7

8/18

1355

! I (A-D) |

2NB

50.0

ALT 8/8+

34.8

17.5

8/24

2010

| II (G-I)

4BB

10.0

ALT 8/8+

85.7

56.3

8/25

1515

j II (G-I)

IBB

10.0

ALT 8/8+

24.2

1.9

8/31

1930

I (F-I)

2BB

3.0

ALT 8/31

26.9

12.2

9/1

2155

I (F-I)

2BB

3.0

ALT 8/31

37.0

32.0

9/3

1450

j I (F-I)

3BB

3.0

ALT 8/8+

38.6

13.5

1900

I (F-I)

3BB

3.0

ALT 8/8+

53.0

36.0

2145

I (F-I)

3BB

3.0

ALT 8/8+

49.6

35.2

9/4

1930

! I (F-I)

2BB

3.0

ALT 9/4

j 37.0

24.2

2145

I (F-I)

2BB

3.0

ALT 9/4

j 38.0

22.6

9/5

1400

I (F-I)

2BB

3.0

ALT 8/8+

| 21.6

12.2

* RR, strong repulsion; R, weak repulsion; O, no apparent response except sensing; A,
attraction.

f Sweat collected by sponge.

weak attraction; A A, strong

Olfaction and Sharks — Tester

165

TABLE 9

Response of Normal Blacktip Sharks (nb), Blinded Blacktip Sharks (bb), and Normal Sharks
(ng) to Human Sweat at Eniwetok Laboratory, I960

DATE

(1960)

TIME

TANK

SHARKS

SWEAT

(ML)

DONOR, DATE

ACTIVITY INDEX

RESPONSE *

Controls

Test

6/30

1250

I (A-D)

2NG

3.0

ALT 6/29

14.6

13.2

1340

I (A-D)

2NG

3.0

ALT 6/29

12.2

8.0

1350

I (A-D)

2NG

5.0

ALT 6/30

—

7.4

1530

I (A-D)

2NG

3.0

ALT 6/29

13.8

10.0

1555

I (A-D)

2NG

6.0

ALT 6/29

8.7

8.6

7/1

1007

I (A-D)

2NG

3.0

ALT 6/29

17.0

18.0 j

1035

I (A-D)

2NG

6.0

ALT 6/29

17.8

17.6

1105

I (F-J)

2NG

3.0

ALT 7/1

13.2

7.0

1130

I (F-J)

2NG

6.0

ALT 7/1

7.4

8.3

1400

I (A-D)

2NG

6.0

ALT 7/1

13.6

8.8

1422

I (F-I)

2NG

6.0

ALT 7/1

14.0

12.2

1942

I (A-D)

2NG

6.0

ALT 7/1

22.4

17.8

7/2

0903

I (A-D)

2NG

6.0

ALT 7/1

18.4

18.6

1140

I (F-I)

2NB

3.0

ALT 7/2

50.5

92.8

1430

I (F-I)

2NB

6.0

ALT 7/2

68.5

71.1

1445

I (A-D)

2NG

9.0

ALT 7/2

18.4

15.6

7/6

1300

I (A-D)

2NB

5.0

SK-ALT 7/2

52.2

23.2

1400

I (F-I)

2BB

6.0

JK 7/6

31.4

31.0

1430

I (A-D)

2NB

6.0

SK 7/6

47.0

31.4

7/7 |

0905

I (A-D)

2NB {

6.0

JK 7/6

40.8

33.8

7/9

1010

I (A-D)

2NB

3.0

SK 7/6 ;

20.0

7.0

1500

I (A-D)

2NB

3.0

JK 7/6

22.6

5.4

7/11

0839

I (A-D)

2NB

3.0

SK 7/6

33.0

29.0

1549

I (A-D)

2NB

! 3.0

JK 7/6

31.4

10.4

7/19

1055

I (F-I)

2BB

3.0

SK 7/6

24.6

14.2

1120

I (A-D)

2NB

3.0

SK 7/6

31.4

31.6

1145

II (A-C)

2NG

3.0

SK 7/6

7.8

10.0

7/20

1505

! I (F-I)

4BB

i 3'°

JK 7/6

25.4

18.6

1530

| I (A-D)

4NB

3.0

JK 7/6

15.4

15.2

7/23

| 1243

I (F-I)

4BB

3.0

SK 7/11

20.4

24.8

7/24

! 1450

! I (F-I)

| 4BB

3.0

ALT 7/23

23.8

22.6

| 1655

; i (A-D)

! 4NB

3.0

ALT 7/23

28.8

13.2

7/25

i 0940

I (F-I)

4BB

3.0

ALT 7/23

25.2

18.4

| 1020

! I (A-D)

4NB

; 3.0

ALT 7/23

33.8

25.4

i R

8/25

1230

I (F-I)

4BB

3.0

ALT 8/25

28.0

26.2

j 1300

1 I (A-D)

4NB

3.0

ALT 8/25

34.0

32.2

! R

* RR, strong repulsion; R, weak repulsion; O, no apparent response except sensing; A, weak attraction; AA, strong
attraction.

periments were conducted at the Hawaii labora-
tory on the population of sharks in Pond 5,
which at that time consisted of a tiger, a ham-
merhead, and a grey shark. The method of
"point introduction" was used. During tests,
three or four uninjured fish ( Tilapia mosam-

bique ) were confined in the 5 -gal funnel of sea
water on top of the tower. During tests, the
water from the funnel containing the fish was
introduced through the hose. This produced a
strong hunting response in the hammerhead.
The response of the tiger shark and the grey

ACTIVITY INDEX

166

PACIFIC SCIENCE, Vol. XVII, April 1963

STARVED

BLACKTIPS

QUIESCENT A6ITATE0
GROUPER GROUPER
WATER WATER

I I I i ! L I I 1—1 L

0900 0912

TIME - CONTROL AND TEST PERIODS

Fig. 10. Activity index during successive 2-min periods, illustrating response of starved grey sharks to
"'quiescent grouper water” and of starved blackdp sharks to "agitated grouper water.”

shark was positive but less intense. "Tilapia
water” was the attractant used in the experi-
ment involving human sweat which was de-
scribed earlier.

Several, more definitive experiments were
conducted at Eniwetok during the summer of
I960 using an experimental arrangement illus-
trated in Figure 9. Living fish were placed in
a 2 5 -gal plastic container into which sea water
was flowing. The sea water could be siphoned
from the container into the test area of either
Tank 1 or Tank 2, or it could be spilled to the
ground. The compartment of Tank 1 contained
four starved blacktip sharks and that of Tank
2 held four starved grey sharks, all with normal

vision. One observer, manipulating the siphons
and living "prey” fish, was concealed from the
sharks by a blind; a second observer, recording
data on shark behavior, was concealed in the
observation booth.

Only two of several experiments will be de-
scribed in detail. In one (Fig. 10), four group-
ers ( Epinephelus merra ) had been placed in the
container the previous night, with the water
siphoning into the blacktip compartment. In
the morning, following a series of control peri-
ods which started at 0830 (timed on a 24-hr
clock ) , the "quiescent grouper water” was siph-
oned into the grey shark compartment. In the
first test period (at 0842) the grey sharks

ACTIVITY INDEX

Olfaction and Sharks — Tester

167

showed obvious awareness and mild attraction,
with one shark biting the siphon tube, but the
response quickly subsided. Siphoning into the
blacktip compartment was then resumed (at
0900) with no noticeable response from the
sharks; this was anticipated as the water had
been siphoning into this compartment all night.
The groupers were then frightened and excited
by threatening them with a moving stick. There
followed a noticeable hunting reaction by the
blacktips (at 0912), stimulated by the "agi-
tated” grouper water. Similar results were ob-
tained with both starved blacktip and grey
sharks using quiescent and agitated surgeon fish
and mullet in place of the groupers.

In the second experiment to be described in
detail ( Fig. 11), the grey sharks showed a nor-
mal behavior pattern during control periods
which started at 1600. When "quiescent grouper
water” was introduced (at 16 12) they responded,

as above, with a mild hunting reaction; one bit
the tube and others milled near it. In the mean-
time, a small grouper had been removed from
the aquarium and held in a dip net in air for
30 min, at which time it was still alive and ap-
parently undamaged. Wth the water from the
quiescent groupers still flowing into the grey
shark compartment, the "distressed” grouper
was quietly lowered into the container (at 1622)
by a string tied around its body; it was removed
after the third test period (1628). The sharks
displayed a violent hunting reaction with cir-
cling and biting of the tube. The "quiescent
grouper water” continued to siphon into the
compartment for about 1 hr, at which time
(1730) the sharks exhibited normal activity
during control conditions. The small grouper
which had been returned to the dip net and was
still alive after 74 min, was again lowered into
the container (at 1740) for three test periods.

Fig. 11. Activity index during successive 2-min periods, illustrating response of starved grey sharks to
"quiescent grouper water,” to "distressed grouper water,” and to "dead grouper water.”

168

PACIFIC SCIENCE, Vol. XVII, April 1963

Fig. 12. Activity index during successive 2-min periods, illustrating response of starved blacktip sharks
to "quiescent grouper water,” to "distressed grouper water,” and to "dead grouper water.”

The sharks again responded with a violent hunt-
ing response and tube biting. The procedure was
repeated after an additional 4 hrs (at 2200)
during which time the quiescent grouper water
had been flowing into the compartment and the
sharks were responding normally. The small
grouper in the dip net, however, had died. After
the dead fish was lowered into the container
(at 2210), there was a spectacular hunting re-
action by the sharks. Only the string was re-
covered from the container at the end of the
third period: the small dead grouper had been
swallowed by one of the larger "quiescent”
groupers. The water in the container was clear
indicating that the fish had been engulfed whole
without the escape of body juices.

The above experiment was repeated on the
starved blacktip sharks, using the same four
"quiescent” groupers and another "distressed”
grouper. In this case the "distressed” grouper
was not eaten after it had died. The results (Fig.
12) were almost identical to those obtained with

the starved grey sharks. A similar test using
four "quiescent” and one "distressed” mullet
gave similar results (Fig. 13). The sharks even
responded to a small "distressed” blacktip shark
which was held in a dip net for a few minutes
and then, still alive, was lowered into the con-
tainer, in this case in the absence of any "quies-
cent” fish.

Discussion

These experiments show that "quiescent” prey
give off an odor which can be detected by sharks
when it is first introduced into their environ-
ment but to which they soon become habituated.
There is still the question, of course, as to
whether the "quiescent” fish were still under
stress because of the artificial environment of
the plastic container. Regardless of this, the
experiments demonstrate that when the prey
becomes frightened and excited it gives off an
additional or a new odor which again stimulates
the habituated sharks, provoking the typical

Olfaction and Sharks — Tester

169

Fig. 13. Activity index during successive 2-min periods, illustrating response of starved blacktip sharks
to "quiescent mullet water,’’ "excited mullet water,” "distressed mullet water,” and "dead mullet water.”

hunting response. Moreover it seems that shark
activity, and thus presumably the amount of
odorous material released, increases with in-
creased agitation of the prey.

It seems unlikely that the odorous material is
associated with body juices released by direct
injury to the prey on the part of the observer.
The fish used were healthy aquarium specimens
which in some cases, e.g., groupers, were used
over and over again and yet suffered no obvious
ill-effects from being repeatedly "agitated.” It
is unlikely they would be damaged by rubbing
against the sides of the smooth plastic container.
They could, of course, rub against each other
when excited. This may possibly have removed
part of their mucous coating and enabled body
juices to escape through the skin.

That the results were not induced by the arti-
ficial environment of the shark tanks was dem-

onstrated in follow-up experiments with sharks
in the natural environment of Eniwetok lagoon
by Hobson (1963). Water siphoned into the
lagoon from a plastic container in which large,
living, agitated but apparently undamaged
groupers had been placed, attracted both white-
tip ( Triaenodon obesus ) and grey (C. menisor-
rah ) sharks. They detected the "grouper water”
from a distance and followed its path upstream
to the source — a concealed plastic tube.

If the substance which attracts the sharks is
released by some subtle damage to the skin of
the prey it might be similar to that demonstrated
by von Frisch (1941) in the injured skin of
the minnow ( Phoxinus laevis ) . As with von
Frisch’s material, identified as a purine- or
pterin-like substance by Hiittel (1941), it might
produce an alarm reaction among the prey but
still be attractive to the sharks. On the other

170

PACIFIC SCIENCE, VoL XVII, April 1963

hand it is tempting to postulate that the sub-
stance is some metabolite which is released
from gill, vent, or skin by excitement rather
than by injury of the prey.

Whatever may be the source and nature of
the attractant, we have presented evidence that
olfaction is involved in the predation of sharks
on normal, healthy fish. It is suggested that in
the natural environment, fish give off odors to
which the sharks are conditioned. It is further
suggested that when the fish become frightened
or excited, and certainly if they rub against each
other or against a coral head, they give off ad-
ditional or new odors which stimulate the hunt-
ing response in sharks. This hypothesis is con-
sistent not only with our experimental data but
also with our observations of the behavior of
the sharks in their natural environment. For
the most part they display a complete disregard
for the myriad of normal, healthy fish which
surround them. However they are able to track
down and converge on a distressed fish (such
as a live fish suspended from a hook through
the jawbone but otherwise uninjured) with un-
canny speed and accuracy.

REFERENCES

Bigelow, H., and W. C. Schroeder. 1948.
Sharks. In, Fishes of the western north At-
lantic. Sears Found. Mar. Res. Mem. 1, Pt.
1:59-546.

Brett, J. R., and D. McKinnon. 1954. Some
aspects of olfactory perception in migrating
adult coho and spring salmon. J. Fish. Res.
Bd, Canada 11 ( 3 ): 310-318.

Eibl-Eibesfeldt, I., and H. Hass. 1959. Erfah-
rungen mit Haien. Zeit. Tierpsych. 16(6):
733-746.

Gilbert, P. W., and F. G. Wood, Jr. 1957.
Method of anesthetizing large sharks and rays
safely and rapidly. Science 126:212-213.

Hobson, E. S., Jr. 1963. Feeding behavior in
three species of sharks. Pacif. Sci. 17(2).

Huttel, Rudolf. 1941. Die chemische Unter-
suchung des Schreckstoffes aus Elritzenhaut.
Naturwiss. 29:333-334.

Idler, D. R., U. H. M. Fagerlund, and Helen
Mayoh. 1956. Olfactory perception in mi-
grating salmon, 1. L-serine, a salmon repellent
in mammalian skin. J. Gen. Physiol. 39(6):
889-892.

Mishkin, M., R. D. Gunkel, and H. E. Ros-
vold. 1959. Contact occluders: a method for
restricting vision in animals. Science 129:
1220-1221.

Parker, G. H. 1910. Olfactory reactions in
fishes. J. Exper. Zool. 8 : 535—542.

1914. The directive influence of the

sense of smell in the dogfish. Bull. U. S. Bur.
Fish. 33:61-68.

and R. E. Sheldon. 1913. The sense

of smell in fishes. Bull. U. S. Bur. Fish. 32:33—
46.

Sheldon, R. E. 1911. The sense of smell in
selachians. J. Exper. Zool. 10:51-62.

Springer, Stewart. 1955. Laboratory experi-
ments with shark repellents. Proc. Gulf, Car-
ibbean Fisheries Inst. 1954: 159-163.

Steinberg, M. A. I960. Report on progress on
contract NR 104-525, chemical analysis of
shark repelling substances. (Mimeo.)

— 1961. Final report on contract NR 104-

525, chemical analysis of shark repelling sub-
stances. (Mimeo.)

von Frisch, K. 1941. Ueber eine Schreckstoff
der Fischhaut und seine biologische Bedeu-
tung. Zeit. vergl. Physiol. 29:46-145.

Whitely, G. P. 1940. The Fishes of Australia:
Part 1, Sharks. Royal Zoological Society of
New South Wales.

Wright, B. S. 1948. Releasers of attack be-
havior pattern in shark and barracuda. J.
Wildlife Mgt. 12(2): 117-123.

Feeding Behavior in Three Species of Sharks1

Edmund S. Hobson2

This report concerns a study of the feeding
behavior in three species of sharks: Car char -
hinus menisorrah Muller and Henle, the grey
shark (Fig. 1), Car char hinus melanopterus
Quoy and Gaimard, the blacktip shark ( Fig. 2 ) ,
both of the family Carcharhinidae; and Triae-
nodon ohesus Ruppell, the whitetip shark (Fig.
3 ) , of the family Triakidae. The study was con-
ducted in the lagoon at Eniwetok Atoll, Mar-
shall Islands, during the summers of 1939 and
I960. It was a segment of a broad program of
investigation of shark behavior in which labora-
tory and field work were coordinated whenever
possible. The overall program, conducted at
both the Eniwetok Marine Biological Labora-
tory and the Hawaii Marine Laboratory, Coco-
nut Island, Hawaii, was under the direction of
Dr. Albert L. Tester, with financial support
from the Office of Naval Research (Contract
Nonr 2756(00), Project NR 104503).

Observations of sharks in their natural en-
vironment have been the basis for most of the
shark literature which is available today and
yet comparatively little has been offered toward
a realistic understanding of shark behavior.
Most of this material has been written for popu-
lar consumption and is therefore oriented to-
ward the sensational rather than the scientific.

We do find scientifically oriented accounts in
the literature (as for example, Eibl-Eibesfeldt
and Hass, 1959; Limbaugh, 1958; and Wright,
1948) but these observations are limited largely
to incidental encounters.

A critical study of shark behavior, undertaken
with planned experiments in the sharks’ natural

1 Prepared as partial requirement for the Master of
Science Degree, University of Hawaii. Contribution
No. 180, Hawaii Marine Laboratory, University of
Hawaii, Honolulu, Hawaii. Manuscript received No-
vember 23, 1961.

2 Department of Zoology, University of Hawaii.
Presently at the Department of Zoology, University of
California, Los Angeles 24, California.

environment — the aim of this investigation —
has been almost completely neglected.

In this study, a comparison is made of the
feeding behavior of the three species. Behavior
is further related to habitat and to distribution
within the lagoon. Experiments designed to il-
lustrate the roles played by the major sensory
modalities are presented. This last portion of
the investigation, dealing primarily with grey
sharks, includes a consideration of the specific
stimuli involved in releasing feeding behavior.
Consideration is also given to food preferences
and to factors which may possibly inhibit feed-
ing.

GENERAL DESCRIPTION OF STUDY AREAS

The study was centered about two primary
locations: (1) the lee of Engebi Island during
1959, and (2) the lee of Bogen Island, adjacent
to Deep Channel, during I960. The Engebi
Island site was in 1 5 ft of water over a relatively
shallow sand and coral rubble flat extending out
from shore for approximately 400 yd before
dropping suddenly into the deeper regions of
the lagoon. Currents are weak in this area and
the water is generally turbid, with underwater
visibility commonly less than 20 ft. At the
Bogen Island site strong tidal currents are pres-
ent and underwater visibility often exceeds 100
ft. The edge of Deep Channel at this point drops
abruptly from a depth of approximately 10 ft
at the rim to 110 ft at the bottom.

At these locations, observations were made
from the following vantage points:

1. An underwater chamber was fitted to a
vessel moored in the lagoon. This chamber was
a metal cylinder, 14 ft long and 2V2 ft in di-
ameter, open at the top and closed at the bot-
tom, with viewing ports on three sides near the
bottom. An observer in the chamber was situ-
ated approximately 8 ft below the surface.

171

172

PACIFIC SCIENCE, Vol XVII, April 1963

Fig. 1. The grey shark, Carcharhinus menisorrah. (Photo by E. Hobson.)

2. A wire cage was suspended from a raft
anchored on the slope at the edge of Deep
Channel. An observer in this cage was located
immediately below the surface.

3. A canvas screen, anchored on the bottom,
concealed an observer lying prone on the bottom
wearing SCUBA equipment.

4. A 16-ft glass-bottom boat, having the
added advantage of mobility, provided for ob-
servations of activity directly below.

5. In many cases, the observations involved
incidental encounters with sharks during the
course of such routine underwater activity as
installing apparatus, etc.

Experimental procedures and apparatus will
be further described as they pertain to the re-
port.

SPECIES STUDIED

Identification of the sharks is based on Schultz
et al (1953). All three species, Carcharhinus
menisorrah , C. melanopterus , and Triaenodon
obesus, reportedly have a wide Indo-Pacific dis-
tribution. They are reported from the Red Sea
as well as the Maidive Islands by Klausewitz
(1958, 1959). Although positive identification
of the two carcharinids awaits a revision of the
family on a world-wide basis, all three species
appear to be prominent in the shark popula-
tions of most Pacific atolls. Harry (1953) re-
ports them from the Tuamotus, as does Randall
(1955) from the Gilbert group. The author
found them abundant at Palmyra and they were
the only species of sharks consistently seen in
the lagoon at Eniwetok during the present in-
vestigation.

GENERAL OBSERVATIONS OF BEHAVIOR

Often when we were engaged in various
types of activity in the lagoon the resulting
commotion, which commonly involved splash-
ing on the surface or striking metal tools on
hard objects underwater, was immediately fol-
lowed by the rapid approach of an obviously
alerted blacktip or grey shark.

In spite of this initial attraction to many stim-
uli, both species exhibited varying degrees of
caution when encountering unfamiliar situa-
tions. This was particularly apparent in the
blacktip, a species which seemed especially sen-
sitive to potential danger. When work was be-
gun at Engebi, blacktips often circled at the
limit of visibility in the baited area for as long
as 2 or 3 hr before approaching the bait. Then,
when an approach was made, it seemed to be
done reluctantly and was often cut short by
some stimulus, unnoted by the observer, which
startled the shark as it neared the bait and
caused it to swim rapidly away. Usually the
shark repeated the approach, but in these early
tests it was not until several sharks had become
active in the area that a blacktip finally took
the bait. We subsequently found that we were
dealing with the same blacktips day after day
at Engebi and that as the work progressed, their
initial caution steadily declined. This was pre-
sumably due to a growing familiarity with the
situation. Nevertheless, a month of testing
passed before the blacktips took the bait with-
out hesitation. Once released, the attack of the
blacktip was very fast and aggressive.

The grey was a notably bolder species. Al-

Feeding Behavior of Sharks— HOBSON

173

though a black dp often approached rapidly in
response to splashing at the surface, it then gen-
erally reversed direction and retreated just as
rapidly if the investigation of the splashing
brought it into an encounter with a human. In
the same situation, the grey ordinarily reacted
to the encounter with a human by continuing
the approach to approximately 5 yd from the
human, at which point it would veer aside and
circle with an apparent cautious interest. If no
further stimuli were introduced, the grey moved
on.

Black tips and greys showed a marked increase
in excitement when feeding in numbers. This
phenomenon, generally referred to in extreme
cases as the feeding frenzy, has been observed
in many species of sharks. In the blacktips and
greys, the presence of more than one shark ap-
peared to lower the threshold for the release of
feeding behavior.

Whitetips did not show this group effect.
Even when feeding in numbers, members of
this species responded individually and without
a notable increase in excitement. The whitetip
seemed to be relatively unresponsive to many
of the stimulus situations which elicited a sharp
reaction in the grey and blacktip. There was
little overt response seen in this shark when en-
countering a human in the water for the first
time. In this situation we did not see the curi-
osity frequently exhibited by the greys, nor
the start and rapid flight usually exhibited by
the blacktip.

On several occasions whitetips appeared and

took fish from the spears of divers before the
fish could be removed from the water. Even in
these instances, the slow, deliberate actions of
this shark did not give the impression of being
a threat to the diver. This impression may have
been an illusion based on the sluggish behavior
of the animal. On one such occasion a 6-ft
whitetip bit the fish in half and then made a
slow pass at the diver holding the other end of
the spear. This appeared to be a warning pass
at a potential competitor rather than an active-
attack. Such warning passes were noted on sev-
eral occasions directed at grouper or snappers
which approached a bait that was under attack
by a whitetip. Similar behavior was noted in-
greys and blacktips. Tester (personal communi-
cation) observed a small blacktip pursue a
grouper of comparable size from a bait which
had been placed on the bottom in shallow water
near Aniyaani Island. In this case, the blacktip
showed considerably more than a warning pass,
as it actively pursued the grouper among a con-
centration of small coral heads. We did not ob-
serve these sharks exhibiting aggressive behav-
ior toward members of their own species. One
observation made on numerous occasions might
at first glance seem to oppose this view. When
a bait which was too large to be immediately
swallowed was presented to a group of feeding
greys or blacktips, the shark which succeeded
in taking the bait would invariably swim rapidly
away from the area, shaking its head vigorously
from side to side presumably in attempts at
cutting up and swallowing the bait. As it fled,

Fig. 2. The blacktip shark, Carcharhmus melanopterm. (Photo by E. Hobson.)

174

PACIFIC SCIENCE, Vol. XVII, April 1963

Fig. 3- The whitetip shark, Triaenodon obesus. (Photo by E. Hobson.)

the other sharks always followed in close pur-
suit. Although the flight of prey was commonly
noted to release attack in the grey, the aggres-
sive behavior of these other sharks in this case
was believed to have been directed against the
bait rather than the fleeing shark.

There was never any mistaking an alerted
grey or blacktip from one engaged in normal
patrolling activity. Thus we had no difficulty
distinguishing a grey or blacktip entering the
test area in response to a stimulus situation we
had presented from one incidentally passing
through the area. The movements of the alerted
greys and blacktips were markedly accelerated
and the grey in particular seemed tense and
highly responsive to subsequent stimulation.
Movements of the grey immediately before at-
tack were markedly abrupt; its body often ap-
peared stiff, with back slightly arched and head
extending straight out and slightly upward. The
pectoral fins were characteristically pointed no-
ticeably downward. Attack was prefaced in
many cases by such anticipatory movements as
a lateral shaking of the head (noted also by
Eibl-Eibesfeldt and Hass, 1959) and a move-
ment of the jaws as in biting (Hobson et al,
1961).

Although it was not always so easy to make
this distinction with the seemingly unrespon-
sive whitetip, these sharks were undoubtedly
more responsive than they appeared to be. It is
likely that their reactions were simply more
subtle than those of the more excitable greys
and blacktips. Whitetips usually appeared in

an area shortly after divers had undertaken
various types of underwater activity. However,
after their appearance the whitetips would swim
about without any apparent interest in the pro-
ceedings. Nevertheless, the consistency of these
appearances indicated that the sharks sensed ac-
tivity and were interested.

DISTRIBUTION OF SPECIES WITHIN LAGOON

Blacktips were the most commonly observed
shark over the sand and coral rubble flats lying
under approximately 1-40 ft of water at the
perimeter of the lagoon. These flats extend out
from shore for distances ranging from approxi-
mately 50 yd to several miles before the bottom
falls off sharply into the depths. Coral growth
in this area is generally restricted to large iso-
lated heads which, in many cases, reach the
surface of the water at low tide. Although black-
tips exceeding 6 ft in length were seen, speci-
mens of more than 4 ft were not common. Small
blacktips were very common on the seaward
reef flats when the sea covered these flats at high
tide. The seaward flats were largely exposed at
low tide.

The whitetip also frequented the shallow
waters of the lagoon, although unlike the black-
tip, which foraged widely over the flats, the
whitetip centered its activity among the coral
heads and about the coral-rock ledges which
border the seaward passages. Unlike the other
two species, the whitetip was commonly seen
resting motionless on the bottom, often under

Feeding Behavior of Sharks — Hobson

175

ledges and in caves. The whitetip was common
to a length of 6 ft, with individuals of 7 ft seen
on occasion.

We seldom encountered the blacktip when
we moved down the slopes from the shallow
flats into the deeper waters of the lagoon or
seaward passages, but here we found the grey
shark in abundance. A census of the shark popu-

lation of the entire lagoon would probably show
the grey shark to be the most numerous. This
shark was commonly seen up to 7 ft in length.

All three species were generally observed
swimming close to the bottom unless drawn to-
ward the surface to feed.

Figure 5 shows shark sightings by species
during a period of 30 days in the vicinity of

Fig. 4. The test area off Bogen Island, showing the relative position of Deep Channel
(top), raft (at edge of channel), and the barge. The island is out of the picture to the left.
(Photo by R. A. Boolootian.)

176

PACIFIC SCIENCE, VoL XVII, April 1963

Fig. 5. Area in the vicinity of Parry Island, showing the sharks sighted during the
period 26 July to 26 August, I960, and indicating species involved and depth of water
where sighting occurred.

Parry Island and the depth of water in which
these sightings occurred. Two basic rules were
observed in making this count: (1) no more
than one count was made in any one area on
any one day; (2) if there was any question of
whether or not a particular shark had already
been counted on any one day, then this shark
was disregarded. This survey was not intended
to show shark abundance, but rather to illus-
trate the areas and depths in which each of the
three species was normally seen.

The distributional picture which emerges is
consistent with the observations made through-
out the program. For example, during 2 months
of work in 15 ft of water off Engebi, only two
grey sharks, both approximately 2 ft in length,
were seen. On the other hand, the experiments
involved many blacktips and whitetips. In con-
trast, when experiments were conducted along
the edge of the dropoff into Deep Channel,
both whitetips and greys were in abundance,
while blacktips were only occasionally seen.

Feeding Behavior of Sharks — Hobson

177

In December 1959, while fishing outside the
seaward reef slope at Palmyra, we saw only grey
sharks, finding these in considerable abundance.
At the same time, only blacktips were common
on the reef flat a hundred yards away. We saw
comparatively few whitetips at Palmyra, these
in shallow water over the reef. Klausewitz
(1959) found these same species occupying
similar habitats in the Red Sea.

Exceptions to this general distribution pat-
tern were noted. Large grey sharks appeared at
poison stations in water scarcely 6 ft deep,
while whitetips were seen swimming over open
bottom, far from the nearest coral head or rock.
Furthermore, Strasburg (1958) reports the
catching of two blacktips at sea in the Mar-
quesas.

BEHAVIOR AND HABITAT

The sluggish behavior noted in the whitetip
is consistent with a life in and about the caverns
and crevices of the coral reef. This species was
noted as being clumsy and ineffective in at-
tempts at taking baits which were suspended
in midwater. However, this same shark was re-
markably effective in tracking down and cap-
turing prey which had taken shelter deep in one
of the many holes or crevices typical of a coral
reef, thus making available to it prey which
are beyond the reach of both greys and black-
tips. Considerable time was spent placing
wounded fish far back into small holes in the
reef and then watching as a whitetip appeared,
nosed about tentatively for the correct hole, and
then swam in and captured its prey. Large
whitetips were seen disappearing into small
holes from which they presently emerged, al-
ways head first. The experiment described be-
low involved whitetips and greys and illustrates
the division of the food source between these
two species.

Experiment 1

Three small wounded fish, each essentially
identical, were presented simultaneously at three
positions below the raft at the edge of Deep
Channel, where water depth was approximately
35 ft: (1) Suspended mid-way between surface
and bottom; (2) on the bottom in an exposed
position; (3) concealed in a hole beneath a
large rock on the bottom.

Bait 2 was lowered to the bottom at the end
of a weighted line. Baits 1 and 3 were both
secured to a single line which ran from the raft,
down under one side of the rock, through the
hole, out the other side and back up to the raft.
By alternately hauling in one end of the line or
the other, both baits could be simultaneously
hauled aboard the raft or lowered into position.

The experiment was repeated 16 times when
both greys and whitetips were in the vicinity.
Although the two species seemed equally adept
at taking the exposed bait on the bottom, the
suspended bait in all but one instance was taken
by a grey, while the whitetips completely mo-
nopolized the bait concealed in the hole.

When grey sharks encountered humans in
shallow water they often started and fled in
much the same manner described for the black-
tip, rather than exhibiting their usual relatively
bold inquisitive approach. Possibly this apparent
change in behavior was a result of their being
out of their usual habitat. This observation
might offer an insight into the characteristically
timid behavior of the blacktip. Perhaps this ap-
parent timidity is an adaption of these relatively
large animals, which must remain in motion, to
a shallow water habitat. The shark, unable to
take shelter and without room to maneuver,
may find flight the alternative. In the blacktips,
this characteristic timidity is more apparent in
the larger individuals. The suggestion that this
behavior might have some survival value im-
plies the existence of a natural predator. The
only evidence we found of such a predator was
the presence of an 18-inch blacktip in the
stomach of an 80-lb grouper.

DETECTION AND CAPTURE OF FOOD

Sharks are well known to feed avidly on dead
fish, meat, and many other food materials
dumped as garbage or used as bait. It is also
well known that they will consume living fish
impaled on a hook or spear. These, however, are
unnatural situations. Undoubtedly, under nat-
ural conditions these sharks will feed on such
prey as may have been killed or weakened by
disease or injury. The sharks in the Eniwetok
lagoon, as elsewhere, are highly responsive to
stimulus situations which suggest injured
and/or distressed, as well as dead or moribund

178

PACIFIC SCIENCE, Vol XVII, April 1963

prey. Nevertheless, it seems unlikely that this
source of food alone is sufficient to support such
a large shark population. It seems probable,
then, that they act not only as opportunistic
scavengers, but also as predators on healthy
free-moving animals. Eibl-Eibesfeldt and Hass
(1959) report observing both C. menisorrah
and C. melanopterus in the Indian Ocean herd-
ing schools of mullet against the shoreline and
actively feeding on these fishes. Similarly, Stras-
burg (1958) observed pelagic whitetips (Pterol-
amiops longimanus ) herding squid under a
night light. Strasburg also cites other evidence
of pelagic sharks apparently capturing what
would appear to be highly motile elusive prey.

We observed no such activity among the three
species in the lagoon at Eniwetok. These sharks
seemed oblivious to the presence of the numer-
ous reef fishes which were continually present
during the shark’s patrolling activities. This be-
havior might be expected, however, as only un-
der such conditions would the reef fishes allow
sharks to move in amongst them without ex-
hibiting immediate alarm and taking shelter.
This same apparent oblivion to what appears to
be potential prey is also standard behavior seen
in many other reef predators, for example the
groupers, snappers, and moray eels. It seems
likely that this behavior on the part of the
predators is advantageous in allowing them to
catch their prey unawares with a frequency
which, while sufficient to maintain life, does
not destroy the illusion of their non-aggressive-
ness. It is also probable that the threshold for
the release of feeding on healthy prey fluctuates
with the relative availability of more readily
obtainable food items, such as dead or disabled
fish.

The present study is confined to feeding be-
havior with respect to dead, damaged, and dis-
tressed prey. An effort is made to determine
which of several sensory modalities are involved
and which are dominant in the sequence of
events between initial stimulation and the act
of consuming the prey.

RESPONSE TO OLFACTORY STIMULI

A number of experiments were conducted
which elucidated the role of olfaction in detect-

ing dead and living prey. Only three (II, III,
and IV) will be reported in detail.

Experiment II

This experiment was designed to study the
response of these sharks to an uninjured fish
struggling on a line, which might thus produce
visual, mechanical, auditory, and perhaps olfac-
tory cues.

The glass-bottom boat was anchored in 40 ft
of water on the steep slope of Deep Channel
where the current ran in one of two directions,
depending on the tide. Ten trials were con-
ducted, each at a time of strong flood or ebb
current when visibility was good. As a precau-
tion against the sharks becoming conditioned
to feeding at this location, the trials were spaced
over a period of several weeks, with only one
trial on any one day. Each trial involved one
fish, either a grouper ( Serranidae ) , snapper
(Lutjanidae) , or mullet (Mugillidae) , 2 to 3
lb in weight, secured to a line by a piece of
soft, light cord which passed through the mem-
brane behind the maxillary. The fish had been
caught by barbless hook and kept in tanks at
the laboratory until needed; they appeared to
be healthy and uninjured.

Before the fish was lowered to a point 5-10
ft above the bottom, a 5 -min observation period
was conducted to insure that no sharks were in
the area. If sharks were seen the test was de-
layed until at least 5 min after they had disap-
peared.

Observations included species of shark, the
time each took to locate and take the bait, its
general behavior, and particularly the nature
and direction of its approach. It was presumed
that if the sharks approached directly and con-
sistently from downstream the attracting stim-
ulus had been initially olfactory, inasmuch as
only the olfactory stimulus was affected by the
current. If the approach was from random di-
rections, then other sensory cues, such as visual,
were likely to be involved in the initial attrac-
tion.

The results are summarized in Table 1. In 9
of the 10 trials the sharks appeared from down-
stream, swimming rapidly and directly toward
the bait, thus indicating they were responding
to olfactory cues carried by the current. The

Feeding Behavior of Sharks— -HOBSON

179

TABLE 1

Response of Sharks to Fish Which Are Struggling on a Line (Experiment II)

TRIAL

BAIT

TIDAL CURRENT

NO. AND LENGTH OF

SHARKS INVOLVED

TIME TO

APPEAR

DIRECTION

FROM

mullet

flood, moderate

1 4-ft grey

17 min

downstream

mullet

flood, strong

1 4-ft grey

10 min

downstream

grouper

flood, moderate

2 6- ft greys

15 min

downstream

snapper

flood, moderate

2 4-ft greys

20 min

downstream

grouper

flood, moderate

1 6-ft grey

18 min

downstream

mullet

flood, moderate

1 4-ft grey

1 6-ft grey

5 min

downstream

mullet

ebb, moderate

2 4-ft greys

10 min

downstream

grouper

flood, moderate

1 4-ft grey

1 3 -ft whitetip

16 min

downstream

grouper

ebb, moderate

2 6-ft greys

1 6-ft whitetip

14 min

downstream

grouper

ebb, moderate

2 4-ft greys

25 min

upstream

one test which might at first glance appear to
deviate from this pattern deserves quotation
from the field notebook:

Introduction was made at 1035 . . . bait positioned
10 ft off the bottom, being carried 20-30 yd astern by
the current where it struggled vigorously until 1050
when noticeably tired. By 1055 bait no longer strug-
gled, but simply maintained position, looking quite
natural. At 1100, two 4-ft grey sharks appeared from
upstream, cruising slowly along the bottom toward
the test area and giving no indication of having sensed
bait. When still about 20 yd upstream of the boat (and
40 ft down, on the bottom) they both became notice-
ably alerted. At this point apparently they had become
aware of the boat on the surface, as they veered up-
ward and swam at an accelerated rate directly to within
5 yd of the boat, turned aside at this distance and
circled twice. Then, seeming to lose interest, they re-
turned slowly to the bottom and continued at their
leisurely pace downstream, passing within 10 yd of
the bait without apparent notice (as stated, the bait
was not struggling, but merely maintaining position
in the current). The sharks continued downstream ap-
proximately 20 yd below the bait, at which point they
both obviously and simultaneously became alerted,
turned around, and with increasing speed raced back
upstream straight toward the bait, with one of them
taking it.

The first response in the above observation
was apparently one of vision to the boat on the
surface. The second response, in which the
sharks returned upstream to the bait, was ob-
viously one of olfaction.

In this experiment, the sharks appeared to be
following an olfactory cue in a direct line to
an uninjured fish. However, there was no as-

surance that other stimuli were not also in-
volved in the detection and approach to the
bait. The question arises whether these sharks
can follow an olfactory cue directly to its source
in the absence of other cues.

The classical experiments of Parker (1914),
in which he observed the approach of dogfish
to bait when both nostrils were free and when
one was occluded, has been offered as proof that
directional response to olfactory cues is possible
by virtue of the ability of each nostril to detect
minute differences in the concentration of odor-
ous material. In standing water in the ponds at
Coconut Island we have observed the hammer-
head ( Sphyrna lew ini) describing the typical
figure 8 pattern, described by Parker, in which
the shark is apparently locating the source of
stimulation by continually turning toward the
nostril exposed to the greater concentration of
the material. Tester ( 1963 ) also describes how
blinded blacktips spiral down from above in
converging on bait on the floor of the tanks at
the Eniwetok Marine Biological Laboratory. It
seems likely that sharks are capable of following
an olfactory trail in running water, particularly
when the current is strong and the trail narrow,
thus forming what would essentially be an ol-
factory corridor. Under such conditions they
could be expected to make a direct-line ap-
proach by taking advantage of the normal lat-
eral movements of the head, which are part of
the swimming motion, in keeping themselves
oriented in the stimulus trail.

180

PACIFIC SCIENCE, Vol. XVII, April 1963

Experiment III

The purpose of this experiment was to de-
termine if these sharks are capable of orienting
themselves in a current and following an olfac-
tory corridor in a direct line to its source in the
absence of other stimuli.

A location on the edge of Deep Channel sim-
ilar to that used in Experiment II was chosen.
Essentially clear, colorless fish extracts were used
as the olfactory cues. These were prepared from
several species of grouper but always of the
same concentration (600 gm of fish flesh or
skin macerated in a Waring blendor with 6
liters of fresh water and diluted in a large plas-
tic container with 64 liters of sea water). The
material was introduced from the glass-bottom
boat by siphoning from the container through a
clear plastic 34-inch hose which ran down to the
top of the reef. From here it continued down
the slope through a series of holes to a small
cave at the base of a large rock. The hose was
thus effectively concealed for a distance of ap-
proximately 30 yd from the cave.

Eight tests were conducted under various ve-
locities of tidal current over a 15 -day period

with no more than one trial on any one day.
Each test was preceded by a 30-min control
period during which any unusual behavior in
sharks sighted was noted. The duration of the
introduction varied according to the response of
the sharks. Observations included the species
and general behavior of the sharks and particu-
larly the direction and nature of approach. The
results are given in Table 2.

A consistent response was exhibited by both
grey and blacktip sharks during this experiment.
In the presence of a current they appeared from
downstream, swimming at an accelerated rate
directly to the mouth of the cave where, in most
cases, they briefly stuck their snouts inside. They
then turned in very small circles here for a short
period of time before returning slowly, in a
random manner, downstream. After an initial
approach by any one shark, this same shark
often reappeared and repeated the above de-
scribed pattern several times. However, upon
the third or fourth approach by the same shark,
this shark often reversed direction 5-10 yd short
of the hole and returned downstream without
the characteristic circling.

TABLE 2

Response of Sharks to Extracts of Fish Flesh or Skin (Experiment III)
(All Sharks Came from a Downstream Direction)

TRIAL

EXTRACT

TIDAL CURRENT

NO. AND LENGTH OF

SHARKS RESPONDING

TIME TO

INITIAL

APPEARANCE

TOTAL

INTRODUCTION

TIME

frozen

flesh

ebb, moderate

1 6-ft grey

1 4-ft grey

1 4-ft whitetip

10 min

1 hr

frozen

flesh

flood, moderate

1 4-ft grey

1 5 -ft grey

2 4-ft whitetips

12 min

1 hr

frozen

flesh

ebb, moderate

1 4-ft grey

1 5 -ft grey

25 min

30 min

frozen

flesh

ebb, weak

1 4-ft grey

1 4-ft whitetip

35 min

1 hr

frozen

flesh

ebb, weak to
moderate

1 4-ft grey

1 4-ft blacktip

1 5 -ft whitetip

16 min

45 min

frozen

flesh

flood, moderate
to strong

3 4-ft greys

1 4-ft whitetip

9 min

45 min

fresh

flesh

flood, strong

2 6-ft greys

1 4-ft grey

1 4-ft whitetip

1 5 -ft whitetip

7 min

30 min

fresh

skin

flood, moderate
to strong

3 6-ft greys

1 4-ft grey

6 min

30 min

Feeding Behavior of Sharks — Hobson

181

It might be suggested that the olfactory stim-
ulus had not itself been directional, but that
the sharks, having been alerted by the olfactory
cue, had simply turned upstream, and oriented
to the current. However, while a rheotaxic re-
sponse may have influenced the shark in its ini-
tial decision as to which way to go in the cor-
ridor, this observer does not feel that the re-
sponse observed could have been directed by
such a cue. Among other considerations, In an
approach directed solely by current the shark
would have at least briefly overshot the source
of the olfactory stimulus. This would have re-
sulted in a brief but certainly noticeable period
of uncertainty as the shark turned back to pick
up the olfactory cue once again. Nothing of this
sort was seen. Furthermore, the following of an
olfactory corridor was observed in the absence
of current and will be described shortly.

Occasionally the extract elicited a response
from other fish in the area which could have
provided an approaching shark with supplemen-
mentary cues. However, in only 2 of 27 ob-
served approaches might this source of error
have affected the results.

When the current subsided, the approach of
the sharks immediately became less direct —
illustrating the importance of the current in
maintaining the definition of the corridor. No
new sharks appeared during slack water and
those already present milled about continuously
within 30 yd of the cave. Approaches to the
cave during slack water were made in a random
manner and from all directions. At this time it
was apparent that the material was diffusing
out in all directions from its source and was at
the same time being retained in the immediate
area.

The whitetip did not seem to be as respon-
sive to the introductions as were the blacktip
and the grey. On several occasions whitetips
swam directly to the cave in the manner noted
in the grey and blacktip. However, whitetips
just as often swam past the hole without any
noticeable response to the extract. As apparent
unresponsiveness has been noted as characteris-
tic of the whitetip, it is difficult to draw any
direct comparisons between this species and the
grey and blacktip from the observed behavior.

Whitetips do have the ability to follow an
olfactory corridor. The following quotation from

the field notebook describes an incident which
clearly illustrates this and also the formation of
a corridor in the absence of current:

A large parrot fish (Scaridae, 10 lb.) was speared in
about 20 ft. of water. The fish tore itself from the
spear and took shelter in a large coral head. Within 1
min. a 5 ft. whitetip appeared. It became obvious that
the shark had sensed the presence of the wounded fish
as it poked about the holes of the coral head and then
swam into one of them. The chase which followed
was witnessed from the surface. The two fish could
periodically be seen through one or another of the
many holes which honeycombed the coral head. First
the parrot fish would flash by and then the whitetip
in pursuit. The coral head contained an extensive net-
work of caves and the chase seemed to take advantage
of most of them. Suddenly the parrot fish emerged
from a hole. Apparently it had temporarily eluded the
whitetip because there was no immediate sign of the
shark. The parrot fish swam off rapidly on a straight
course for about 30 yd. where it made a 90° turn and
continued on the new course, in a straight line, until
it was out of sight. The whitetip emerged from the
hole just seconds after the parrot fish, but already its
prey was out of sight. The whitetip circled briefly,
then started out along the same path taken by the par-
rot fish. When it arrived at the point of the 90° turn
it continued on a few yards, but quickly slowed and
turned around. After another brief period of circling
the shark picked up the second leg of the trail and
followed it straight out of sight.

In this instance, the corridor was formed by
olfactory substances given off by the moving
wounded fish. There was no noticeable current
at the time.

The sharks involved in Experiment II had
apparently been initially alerted by an olfactory
stimulus emitted by a fish which, while in dis-
tress, was uninjured.

The ability of these sharks to detect the pres-
ence of an unwounded fish in a state of stress
through an olfactory cue was noted by Tester
(1963) early in the program. This point has
been neglected since the observations of Shel-
don (1911), in which a dogfish was noted to
locate an undamaged crab wrapped in eelgrass.
However, in Sheldon’s experiment the attract-
ing stimulus (or stimuli) may have been a
movement or sound made by the crab instead
of, or in addition to, an olfactory stimulus. Any
conclusions derived from Experiment II are
open to the same criticism. Another source of
criticism might be the assumption that a fish
with a line passing through its maxillary mem-
brane may be considered uninjured. The ex-

182

PACIFIC SCIENCE, VoL XVII, April 1963

TABLE 3

Response of Sharks to Water Containing a Grouper Under Stress (Experiment IV)
(All Sharks Came from a Downstream Direction)

TRIAL

TIDAL CURRENT

NO. AND LENGTH OF

SHARKS RESPONDING

TIME TO

INITIAL

APPEARANCE

TOTAL

INTRODUCTION

TIME

flood, moderate

2 5 -ft greys

15 min

30 min

ebb, strong to slack

3 4-ft whitetips

5 min

30 min

flood, moderate

1 4-ft grey

1 3 -ft whitetip

10 min

1 hr

periment described below was designed to mini-
mize these sources of error.

Experiment IV

This experiment was conducted to determine
if these sharks are capable of detecting and
tracking down, exclusively by olfaction, an un-
injured fish under stress.

The experiment was carried out using the
procedure described for Experiment III, except
that, in place of the extract solution, the large
plastic container was filled with sea water and
contained an uninjured grouper (1-10 lb) which
was presumably under stress. In each case, the
fish was caught by line with a barbless hook.
During the experiment the fish was agitated
intermittently with a pole, using care not to in-
flict any damage, while the water was running
from the container to the bottom as described
for Experiment III.

Three trials were conducted over a period of
10 days, with at least 3 days between each trial.
The results are given in Table 3.

The response was essentially the same as that
to the extracts in Experiment III. Greys and
whitetips were involved in the experiment; no
blacktips were seen.

The captive grouper obviously emitted an ol-
factory stimulus which attracted the sharks up-
stream to the cave. Critics of this experiment
may question whether a recently hooked fish can
be considered uninjured. There is justification
for pointing out the hook wound as well as the
skin rubbed and mucus dislodged during han-
dling. These factors offer a possible source of
error. However, the results were consistent with
findings in the laboratory tanks where it was
possible to maintain considerably more control

over experimental conditions. In the latter tests
(Tester, 1963) the strength of the stimulating
olfactory component varied with the degree of
distress of the fish being used. It appeared that
a maximum level was reached shortly after the
death of the fish.

It has been shown that the sharks were able
to track down a distressed but apparently un-
injured fish by olfaction alone in Experiment
IV. Although they could also have made exclu-
sive use of the olfactory sense in tracking down
the hooked fish in Experiment II, it is highly
unlikely that they did so. We shall see that, in
all probability, other sensory modalities not only
contributed to, but in fact dominated, certain
phases of the approach.

RESPONSE TO COMPRESSION WAVES

Compression waves are regarded by many to
have an important effect on the behavior of
sharks, both as an attractant and as a repellent.
Wright (1948) claims that attack patterns in
sharks are released by sounds. Many investiga-
tors (e.g., Eibl-Eibesfeldt and Hass, 1959) re-
port that the vibrations and sounds made by a
wounded fish will attract sharks. On the other
hand, some sounds have been reported to have
a repelling effect, as for example the underwater
shouts of divers (Hass, 1951)- These reports
are based on incidental encounters with sharks
and are generally complicated somewhat by the
presence of stimulating factors other than com-
pression waves which might themselves have
been influential in eliciting the observed re-
sponse. For example, Wrights conclusion is
based to a considerable degree on observations
of sharks appearing just after an underwater
explosion to feed on the dead and stunned fish.

Feeding Behavior of Sharks — Hobson

183

The presence of the disabled fish is mentioned
almost incidentally, although in such a situation
it is impossible to dismiss the olfactory and
visual cues emitted by these fish. Similarly,
the presence of olfactory and visual cues also
complicate reports of sharks being attracted to
a struggling fish in response to compression
waves. In regard to the repelling effect of the
shouts of divers, we did not see any such re-
sponse in the species studied in Eniwetok. How-
ever, sudden movements and/or a sudden burst
of bubbles from the aqua-lung, such as might
readily accompany a shout, often startled the
blacktip and put it to flight.

It is important to consider the ability of
sharks not only to sense these stimuli but also
to locate their sources. Parker (1912), investi-
gating sound as a directing influence on the
movements of some teleosts, noted that there
was a directing effect only during the duration
of the sound. These results cast some doubt on
the ability of such noncontinuous stimuli as a
single underwater explosion to attract sharks.

Two experiments on the perception of com-
pression waves by sharks were conducted in the
field at Eniwetok.

Experiment V

The aim of this experiment was to investi-
gate the effects of compression waves of a variety
of sonic frequencies on these sharks. The fol-
lowing were presented:

A. Continuous pure tones, covering a fre-
quency range in steps from 100-1,000 cycles
per sec.

B. Sounds of mixed frequencies recorded on
a circular tape, including:

1. wood struck against wood, 1/ sec and 6/ sec;

2. rock struck against rock, 2/sec and 6/sec;

3. rasping effect, continuous;

4. rattling of nuts and bolts in a bottle,
continuous.

C. Shouts by the observer, directly projected
through the apparatus.

The experiment was conducted at the Engebi
location, using a tape recorder equipped with an
underwater speaker lowered 6-8 ft below the
surface. Each sound was presented during 20-
min trials, with each trial immediately follow-
ing a 20 -min control period and conducted

under two sets of conditions: (1) following a
different experiment in which sharks had been
drawn into the experimental area and were still
present, and (2) upon arrival at the site, with
no sharks present. All sounds were audible to a
submerged human at distances greater than
150 ft.

Although we watched closely for signs of any
sort of response, for example curiosity, we
saw nothing in the behavior of the sharks which
suggested that they were able to perceive the
sounds.

Experiment VI

This experiment was designed to investigate
the effects of various subsonic compression
waves on these sharks.

A metal hoop, having a diameter of 36 inches,
was fitted with a rubber diaphragm held in posi-
tion by a series of surgical tubing lacings. This
piece of apparatus was installed midway through
a natural tunnel, approximately 20 ft long, in
the coral ridge bordering Deep Channel beneath
the raft. A line was secured to the center of the
diaphragm on that side facing away from the
channel, and this line was run out of the tunnel
and up to the surface where a small skiff rode
at anchor. The diaphragm could thus be vibrated
by jerking on the line.

Two trials were conducted in which the dia-
phragm was vibrated irregularly for a period of
20 min immediately following another experi-
ment in which a number of grey sharks had
been drawn into the area and were still present.
Throughout both trials there were grey sharks
swimming about calmly in the area. At times
they passed within 5 yd of the tunnel’s entrance.

In observations from the raft we looked for
any type of response in the sharks which might
indicate that they were aware of the vibrating
diaphragm. We saw no such response.

The negative results in these two experiments
do not mean that the sharks could not perceive
these stimuli. If releasers of feeding activity are
emitted by struggling fish in the form of com-
pression waves, they probably are of a specific
nature or involve a characteristic pattern. It
would have been a rare stroke of luck if we had
duplicated a specific releaser with our crude
experiments. The problem of delimiting the

184

sensory capacity for stimuli of this sort is prob-
ably best approached with conditioned response
experiments on captive sharks under controlled
conditions before attempting to establish their
role in feeding behavior under natural con-
ditions.

Although the work of Parker (1903, 1911),
Kritzler and Wood (1961), and others has
demonstrated that at least some sharks are sensi-
tive to compression waves of a relatively wide
range of frequencies, it has yet to be proven
experimentally that such stimuli normally re-
lease feeding behavior. Nevertheless, incidental
observations at Eniwetok indicated they do play
an integral role in feeding activity. These ob-
servations are of a nature similar to those men-
tioned at the beginning of this section and are
subject to the same reservations. One such ob-
servation is quoted from the field notebook:

... A 30 lb. grouper was speared on the slope border-
ing Deep Channel . . . the grouper fled, dragging the
spear, into a small cave. . . . Within seconds, 5 greys
4—7 ft. in length swam excitedly into the area from
downstream. As soon as they were in the area, how-
ever, their excited state diminished. They swam about
in the area for several minutes, appearing to steadily
lose interest, before slowly drifting off downstream
and out of sight. After a few minutes ... 5 greys made
another approach which seemed as highly motivated
as the preceding one. Immediately I looked below to
see a small puff of sediment emerge from the cave —
the grouper was obviously thrashing about inside.
Almost as soon as the sharks were in the area, all be-
came quiet in the cave, whereupon the sharks imme-
diately lost their excited state and settled down to
cruising about as before. Two 4-ft. whitetips joined
them before they all again drifted off downstream.
After a few minutes . . . another small puff of sedi-
ment was visible at the mouth of the cave — the
grouper was struggling again. I quickly looked down-
stream to see the greys on their way in as before. The
pattern previously described was repeated, but this
time there were 4 whitetips (3—5 ft.) in addition to
the 5 greys. Several minutes after the sharks disap-
peared downstream for the third time the grouper
scrambled out of the hole and, dragging the spear,
struggled along the bottom toward the bigger, more
protective caves farther up the slope. The 5 greys
charged into the area from downstream before it had
gone 20 ft. At this point, the grouper stopped, and
ceased all movement. Although the grouper was in a
completely exposed position, the excited state of the
onrushing greys diminished almost as soon as the
grouper stopped moving. Instead of attacking the com-
pletely vulnerable grouper as I expected, the greys
settled down to swimming slowly and randomly about
the area. Often they came within inches of the mo

PACIFIC SCIENCE, Vol. XVII, April 1963

tionless grouper, and on at least one occasion grazed
it. The 4 whitetips rejoined the group, along with 2
blacktips (3 and 4 ft.) making a total of 12 sharks,
all obviously interested, but offering no show of ex-
citement or indication that they regarded the wounded
grouper as prey. Nevertheless, the sharks did not
gradually drift off downstream as they had previously,
but continued to swim about in the area. Thirty min-
utes later, when we had to leave, the situation was
unchanged — the grouper was still sitting motionless in
the same spot, while the 12 sharks swam slowly about.
Most of this random swimming about the area, both
at this point and earlier, had occurred downstream of
the grouper.

It was significant that all approaches and de-
partures, as well as most of the random swim-
ming in the area, occurred downstream of the
grouper. This indicated that olfaction was in-
volved throughout the incident. Although the
excited approach of the greys during this inci-
dent appeared to coincide with the periods when
the grouper was struggling, it is possible that
they had first been conditioned by an olfactory
cue. In this case, the olfactory cue may have had
a threshold lowering effect for the postulated
compression wave stimuli. At least one author
(Wright, 1948) doubts that olfaction alone will
release attack patterns in sharks. In this latter
regard, blinded sharks in the tanks fed avidly on
chunks of fish flesh which were lying on the
bottom, thus offering little other than olfactory
stimuli before the sharks came into contact with
them. The question here is whether or not the
behavior of these blinded sharks can be con-
sidered normal.

While the sharks in the latter phase of the
incident did not attack the wounded grouper,
they did not drift off downstream as they had
done when this same fish was concealed in the
cave. We are probably safe in assuming that the
behavior pattern demonstrated in swimming
slowly about in the area was appetitive in nature.
This suggests that an additional stimulus was
required to release the actual attack. This is to
say that a combination of stimuli, e.g., an olfac-
tory element and erratic motion might have been
necessary to release the attack in this case. The
olfactory cue may have simply released an ex-
ploratory behavior pattern which drew the
sharks to the area. Once there, the release of
the attack may then have required a specific
stimulus not presented by the quiet, natural
looking grouper, the coloration of which blended

Feeding Behavior of Sharks — Hobson

185

in very well with the bottom on which it was
resting. The cessation of movement by this
grouper upon the approach of the sharks may
have been a well-established protective behavior
pattern. We will consider this incident further
in the next section. During a later period of the
study, a grouper of the same species and ap-
proximately the same size was caught at the raft
where the underwater action was witnessed from
the observation cage. The grouper wrapped the
line about a piece of coral and by the time the
line was freed and the struggling fish hauled
toward the surface, a single 5 -ft grey came rush-
ing into the area from downstream. The shark
went directly for the struggling fish, took the
tail section into its mouth, and with a few
vigorous lateral shakes of its head came away
with the after portion of the fish.

There is no doubt that rapid erratic move-
ments are a prime releaser of attack patterns in
these sharks, particularly in the presence of spe-
cific olfactory cues. Many authors have noted an
excited state in sharks resulting from the sensing
of movement (for example Limbaugh, 1958).
As both a mechanical disturbance and a visual
stimulus are usually produced by a moving ob-
ject underwater, it is very often difficult to say
with certainty which is more significant as a
releaser of attack in a given situation.

On one occasion, while fishing for sharks, a
3 -ft grey was hooked in the presence of five
larger greys. As the shark struggled vigorously
on the line, the other five sharks became highly
active and appeared to be chasing the hooked
animal. Initially we supposed that an attempt
was being made to attack the hooked shark, but
upon landing this individual after several min-
utes of activity we noted no evidence of injury.
If the larger sharks had been attempting to feed
on the smaller hooked shark there is no doubt
that they could have done so. It is probable that
the activity of the hooked shark excited the
others which were then simply following the
focal point of this activity. Inasmuch as other
species of fish, similarly hooked and struggling,
were immediately taken by the grey shark there
is a suggestion here that this species of shark is
inhibited in some way from attacking members
of its own species. This possibility is further
supported from observations made while fishing
for snapper ( Lutjanus bohar) at Palmyra. Grey

sharks drawn to the fishing area showed no in-
terest in the many free-swimming snapper, but
would immediately attack a snapper which be-
came hooked and began to struggle on the line.
Contrariwise, although the sharks would swim
rapidly about one of their own species which
was hooked and struggling, no attacks were
observed.

RESPONSE TO VISUAL STIMULI

Contrary to many reports on the subject (for
example Halstead, 1958) vision was found to
play a major role in the feeding activity of the
sharks in the Eniwetok lagoon. We briefly men-
tioned vision above in regard to instances in-
volving movement. The following experiments
were designed to further clarify the role of
vision in cases where there was little or no
movement of the bait.

Experiment VII

The object of this experiment was to deter-
mine the role of vision in these sharks when
they are approaching a motionless bait up an
olfactory corridor.

Two baits of similar appearance were pre-
pared for each trial, one of which was a 4-inch
cube of grouper flesh while the other was a
4-inch cube of wood. After we were confident
that no sharks were visible in the area, the two
baits were presented together in a strong cur-
rent, suspended 3 ft apart, midway between the
surface and the bottom. Observations were made
from the cage, and a record was kept of which
bait was hit first, as well as of the nature and
direction of approach of the sharks.

Experiment III had shown that these sharks
are capable of orienting on an olfactory stimulus
in a current and swimming directly to its source.
In every case during the present experiment the
sharks appeared from downstream, swimming
at an accelerated rate, presumably following an
olfactory trail emitted by the grouper flesh. If
the sharks had continued to orient exclusively
on the olfactory stimulus all the way to the bait,
then the fish would presumably have been taken
in all trials. This, however, did not occur. In 20
trials conducted over a period of 2 days, the fish
was struck first 11 times while the block of
wood was hit first 9 times. As no preference was

186

shown, it seems that at least the final phase of
the approach was visually directed.

When the shark selected the wood, the ob-
ject was either simply bumped or briefly taken
into the mouth and then rejected. No teeth
marks were found on the wood following the
test. In all cases the fish-baits were carried away.
Only grey sharks were involved.

If we accept as fact that these sharks orient
visually during the final phase of their approach
to a motionless prey, we must then question the
nature of the stimulating visual image, bringing
us to the subject of visual acuity.

Most elasmobranchs reportedly possess an all-
rod retina with a high ratio of visual cells to
ganglion cells, which provides for low visual
acuity. While this would presumably result in
an inability to see objects in detail, the sharks
would be able to utilize their visual sense under
conditions of relatively little light (Gilbert,
1961). Kato (1962) found no evidence of cones
in the retina of blacktips or whitetips. Perhaps
even more significant, he found no specialized
area, corresponding to the human fovea, which
is generally assumed to be associated with visual
acuity.

At Engebi, we often noted that sharks at-
tracted by a dead fish suspended on a line often
struck objects in the immediate vicinity of the
bait before taking the bait itself. A rock about
IV2 ft square, which was situated in the middle
of a sandy area below the baits, as well as float-
ing seaweed and a lead weight on the line a
foot above the bait were often tested with a
nudge or by actually being taken momentarily
into the mouth. These observations suggest one
of two alternatives: (1) these sharks lack the
visual acuity necessary to distinguish the visual
appearance, i.e., form, markings, etc. of fish from
inedible objects in the immediate vicinity; or
( 2 ) although these sharks do possess the visual
acuity necessary for such a distinction, it was not
utilized by them in this situation.

Tester and Kato (ms) showed that small
blacktips and greys have the ability to discrimi-
nate between a number of objects of different
shape, e.g., squares, triangles, and rectangles,
and that they make this discrimination from a
distance of at least 6 to 12 ft. These species then
possess at least the degree of visual acuity re-
quired to discriminate between these forms.

PACIFIC SCIENCE, Vol. XVII, April 1963

The following experiment was an attempt to
detect evidence of the utilization of a high de-
gree of form discrimination ability in the feed-
ing behavior of these sharks.

Experiment VIII

This experiment was designed to determine
whether or not the visual cues offered by a fish
through its characteristic morphological features
are significant attractive visual stimuli to these
sharks.

Two fresh dead fish were selected for each
trial, both being of the same species, but one
a little larger than the other. Three species were
used: goatfish (Mullidae), 8 trials; surgeon fish
( Acanthuridae), 3 trials; and squirrel fish (Ho-
locentridae), 10 trials. The larger fish was then
decharacterized by removing the head and all
the fins, reducing it in size to approximately
that of the smaller, still natural-appearing fish.
A long, deep incision was then made along the
ventral side of the natural-appearing fish to in-
sure that olfactory stimuli emitted by both baits
were essentially the same. The baits were both
secured to a single line, one spaced approxi-
mately 18 inches above the other, with their
respective position being reversed on alternate
trials. After assurance that no sharks were vis-
ible, the baits were suspended motionless at a
point midway between surface and bottom in a
strong current. Observations were made from
the cage and a record was kept which bait in the
pair was taken first, along with the species of
shark involved and the nature and direction of
approach.

It was assumed that if the characteristic mor-
phological features which had been removed
from one of the pair were significant as attrac-
tive visual stimuli, then the natural-appearing
fish would be favored in the initial choice made
by the sharks. This in turn would demonstrate
a high level of form discrimination ability in
these sharks. However, no apparent preference
was shown to either of the two bait types in
21 trials. The decharacterized fish were hit first
11 times, while the natural-appearing fish were
hit first 9 times. Once both baits were hit simul-
taneously. The relative positions of the baits on
the line had no apparent effect, with the top
bait being hit first 9 times and the bottom bait

Feeding Behavior of Sharks — Hobson

187

1 1 times. As in Experiment VII, only grey sharks
were involved and these approached in every
case at an accelerated rate from downstream. By
positioning the bait in mid-water we favored
this species. A number of whitetips were seen
about the bottom at various times during the
trials.

The baits introduced during this experiment
were the first food given by us to the sharks in
over a month, with the exception of a few un-
injured living fish. This is an important con-
sideration in this experiment, as any condition-
ing of the shark population to feeding on our
presentations would have resulted in a tendency
to take our offerings indiscriminately. However,
no conditioning was noted at this stage of
experimentation.

The failure of these sharks to show a pref-
erence for the natural-appearing fish over the
one which had been reduced to a simple object
suggests that the visual appearance of a fish
per se is not itself significant to these sharks
when feeding. When the results of this experi-
ment are considered along with the observations
at Engebi already mentioned, in which the
sharks struck at inedible objects in the imme-
diate area of the baits, it appears that the sig-
nificant visual cue in these cases was simply that
of an object within the immediate area of the
source of the olfactory stimulus. This evidence
suggests that these sharks do not utilize a high
degree of visual, acuity in their feeding activity.
It follows, then, that if the visual stimulus is
not moving, it must otherwise contrast with the
background against which it is viewed in order
to provide an effective visual cue.

The following experiment involves the rela-
tive effectiveness of two objects in attracting
the attention of sharks, when these objects differ
from one another in degree of contrast in bright-
ness with their background.

Experiment IX

This experiment was designed to determine
which of two baits, one white and the other
black, would be taken first when both are pre-
sented to these sharks together at the surface.

Paired 3 -inch cubes of fish flesh (parrot fish,
snapper, and grouper) were used as bait in this
experiment, with members of each pair identical

except that one retained its natural white color,
while the other was dyed black with nigrosine
dye. It had been previously determined that
nigrosine dye was not sensed by blinded sharks
in the tanks.

The experiment was conducted at the raft
over 40 ft of water when a current was run-
ning. A wounded grouper was placed in the cage
until a number of sharks had been drawn about
the raft. The fish was then removed from the
water, allowing the source-point of the olfac-
tory stimulus to be carried off downstream and
taking the sharks with it. When the sharks were
about 20 yd downstream from the raft, the two
baits were dropped into the water, spaced ap-
proximately 10 ft apart. Movement of both baits
was negligible as they slowly settled. At times
the response of the sharks immediately followed
the entry of the baits into the water, indicating
that the splash had been sensed; at other times
the response was noted after the baits began to
sink, indicating a visual response. These re-
sponses followed the introduction too closely to
have been those of olfaction. In either case, the
sharks came racing back upstream and in all
cases the baits were taken before they had fallen
10 ft. The bait taken first in each case was
recorded, although the alternate bait was always
taken almost simultaneously. As it was antici-
pated that the amount of incident light present
would influence the results, trials were con-
ducted under different light conditions, with
from 10 to 56 trials held each day for 4 days.

The results under each of the different sets of
conditions prevailing were as follows: ( 1 ) day-
light with clear sky, in 96 trials black was taken
79 times, or 82%; (2) daylight with overcast
and drizzle, in 21 trials black was taken 15
times, or 71%; (3) daylight with overcast and
heavy rain, in 14 trials black was taken 8 times,
or 57%; (4) after sunset, with clear sky but
almost dark, in 41 trials black was taken 32
times, or 78%. The total: for 172 trials under
all conditions, black was taken 124 times or
72%.

As the sharks, all greys, raced back toward the
test area they were viewing the baits against
light surface water. The black bait, then, con-
trasted with its background to a greater degree
than did the white bait. This was as noticeable

188

to the observer sitting in the cage as it obviously
was to the sharks.

Recalling some of the observations made
earlier, we might consider again the speared
grouper which rested motionless on the bottom
while 12 sharks swam slowly about in its im-
mediate area. Although this grouper was in
a completely exposed position, its coloration
blended in well with the bottom on which it
was resting. In this position the grouper was
apparently at least temporarily safe from attack
by the 12 sharks, in spite of the fact that it was
wounded and still emitting the olfactory stim-
ulus which presumably had released the ap-
petitive behavior pattern which these sharks
were at that time demonstrating. At Engebi, the
sharks struck at the rock which was sitting in
the middle of the sandy area, floating seaweed,
and the lead weight — all inanimate objects in
the immediate area of the bait, but all of which
sharply contrasted in brightness with the back-
grounds against which they were viewed by the
sharks. These results are consistent with the con-
clusions drawn by Gilbert (1961) from ana-
tomical studies of the shark eye.

Although there is little doubt that vision is
the predominant directing sense within the
visual field, the effective distance involved here
will be highly variable. Such external factors as
water clarity, incident light, and whether or not
the prey is under cover, no doubt determine the
effectiveness of vision in any given situation.

RESPONSE TO GUSTATORY AND
TACTILE STIMULI

While lack of visual discrimination appar-
ently led to selection of the wood as often as it
did the fish during Experiment VIII, the shark
was quick to learn its error. Initially the wood
was actually taken into the mouth, but after
one or two successive trials a nudge was gen-
erally sufficient to dismiss the inedible object.
An appraisal of the bait by the visual or olfac-
tory sense may have been the basis for the
nudge, but when the bait was taken into the
mouth, other senses, e.g., gustation and/or tac-
tile sense, undoubtedly came into play.

A review of the gustatory sense of sharks is
presented by Tester (ms). In this review, he
points out that while the receptors of the gus-

PACIFIC SCIENCE, Vol. XVII, April 1963

tatory sense (e.g., taste buds or terminal buds)
occur in the skin of the body, fins, and barbels
of many species of fishes, in the elasmobranchs
they appear to be associated with papillae which
are confined to the epithelial lining of the
mouth and pharnyx. He also calls attention to
the claim of Budker (1938) that the "pit or-
gans" located on the body of elasmobranchs have
a gustatory function.

An effort was made to design an experiment
which would illustrate the respective roles
played by both gustation and the tactile sense
in accepting or rejecting food taken into the
mouth. As the block of wood in Experiment
VIII differed from the fresh fish in tactile cues
as well as those of olfaction and gustation, it
was impossible to say which of the two might
have been more influential in the rejection of
the wood.

An attempt was made to synthesize a bait
which possessed the visual and tactile properties
of acceptable bait, but which lacked the olfac-
tory and gustatory properties. Tester et al.
( 1955) concluded that in the flesh of many
fishes there is present a substance or substances
which, when extracted with alcohol or water,
can be perceived by a fish through its sense of
smell or taste and which promotes the urge to
feed. Tester commented (personal communica-
tion) that extraction by alcohol was effective in
removing this substance from a piece of flesh.
He observed that squid prepared this way was
not detected by blinded blacktips and although
taken into the mouth by normal blacktips, they
were subsequently rejected. He attributed this
rejection to the lack of acceptable gustatory
stimulation, in as much as the texture of the
prepared squid seemed to be similar to untreated
squid.

The following experiment was an attempt to
duplicate Tester’s results in the field.

Experiment X

This experiment was designed to determine
whether or not a bait offering the visual and
tactile stimuli of food, but which lacks the ol-
factory or gustatory stimuli, will be acceptable
as food by these sharks when they are actively
feeding.

A number of 3 -inch cubes of grouper flesh

Feeding Behavior of Sharks — Hobson

189

were prepared. Half of these baits were soaked
for 4 days in each of the following concentra-
tions of ethyl alcohol: 35%, 75% and 95%, in
that order. After the alcohol treatment the baits
were soaked in sea water for 3 hr. It was hoped
that the product of this process would be a
bait which felt and looked like food, but did
not taste or smell like food. In actuality, how-
ever, the texture of the product was not identi-
cal to that of normal flesh, being notably dry
and leathery. The experiment was nevertheless
continued, with each of the treated baits being
paired with a normal bait. It was assumed that
the normal bait would be taken. If the treated
bait was also taken this would indicate that the
wood in Experiment VIII had been rejected due
to its tactile properties rather than its lack of
acceptable olfactory or gustatory stimuli. If,
however, the treated bait was not taken, Testers
results would be confirmed and it would appear
that gustation (and perhaps the tactile sense
too) had been involved in the rejection of the
wood.

The test was carried out at the raft, in a mod-
erate current, using essentially the same tech-
nique described for Experiment IX, except that
in this case all 15 trials of the experiment were
conducted during a 30-min period of one after-
noon and the response to both baits was noted.

Only grey sharks were involved in the results.
The treated bait was hit first 6 times, while the
natural bait was hit first 9 times, indicating that
the sharks did not make a distinction between
the visual appearance of the two baits. All the
baits were taken into the mouth, but although
in all trials the natural bait was swallowed im-
mediately, the treated bait was rejected within
seconds. As this rejection took place after the
bait had been taken into the mouth, it appeared
to be based on gustatory stimulation (if we
are correct in assuming that the bait offered ac-
ceptable tactile stimulation). A snapper was
seen taking, and apparently retaining, one of
the treated baits rejected by the grey shark.

Another effort to clarify the roles played by
gustation and the tactile sense in feeding in-
volved the use of sponges. It was thought that
perhaps a sponge which had been soaked in an
extract of fish flesh would be accepted by the
sharks as food. If this were the case, then by
pairing the soaked sponge with a normal sponge

it would have been possible to present a choice
of baits which posed the same problem to the
sharks as did Experiment X. In order for the
experiment to have been a success, however, it
was necessary for the sharks to have accepted
the soaked sponge as food, and this proved to
be only temporarily true. Initially, the soaked
sponge was carried off in the manner seen with
a chunk of fish, while the plain sponge was re-
jected as was the prepared bait in Experiment
X. However, after 3 trials the sharks began to
lose interest in the soaked sponges until they
either dismissed them after an unenthusiastic
nudge, or ignored them completely. Further-
more, as the sharks learned that the soaked
sponges were inedible, the extract no longer
excited them. In this latter regard it is difficult
to say whether the sharks actually learned that
the extract did not indicate food or whether
olfactory fatigue was the major factor.

A definite change in general behavior of the
grey sharks was first noted during Experiment
IX and became very evident during Experiment
X. As the sharks became conditioned to feeding
about the raft they responded instantly to any
object which was presented. In addition, there
was a sharp increase in the number of sharks
responding to our test situations. Where ini-
tially a maximum of 5 or 6 grey sharks were in-
volved in our experiments, the number steadily
increased during this period of increased food
supply until more than 20 were being drawn
about the raft. The initial population had been
relatively stable and through successive observa-
tions involving experiments which utilized min-
imum feeding, it had become possible to recog-
nize a certain group of about 6 individuals that
appeared day after day. These same individuals
were seen after the population numbers in-
creased, but apparently many other sharks which
normally frequented other areas were drawn to
this point of concentrated feeding. This change
involved only grey sharks, which is understand-
able as this species was completely monopoliz-
ing the additional food. The increase in num-
bers of sharks resulted in a markedly increased
element of competition. When we witnessed
the simultaneous rush at the baits by up to 10’
sharks, it was understandable that all baits were
taken into the mouth before any discrimina-
tions were made as the slightest hesitation by

190

a shark in this situation immediately eliminated
it from a chance at a bait. Fortunately the pres-
ent tests, which required concentrated feeding,
were concerned with probing questions which
did not suffer from these effects. This change
in behavior, along with the increase in numbers
of the local sharks, was therefore viewed with
interest rather than alarm.

FOOD PREFERENCES

Sharks have been popularly described in a
general way as creatures with an exceptionally
voracious appetite, feeding on such unlikely ob-
jects as tin cans, bottles, and other trash (Linea-
weaver, I960). Although considerable effort was
brought to bear on the problem of food pref-
erences during this study, the experiment de-
scribed below was the only one, of many experi-
ments conducted, which clearly indicated a pref-
erence for one of two food materials presented.

Experiment XI

This experiment was an effort to determine
whether or not these sharks show any preference
between grouper flesh and the flesh of various
species of mollusks, e. g., Tridacna sp., (10
trials); Spondylus sp ., (5 trials); and Cassus
sp., (3 trials).

Mollusk flesh in 2 -inch cubes was paired with
chunks of grouper flesh of the same size, with
both baits thus offering an essentially identical
visual appearance. Prior to the test, large pieces
of grouper and mollusk flesh of the species to
be tested were placed in the cage on the raft
to attract sharks. When the introductions began,
there were 10 grey sharks, 3 to 6 ft in length,
present in the area. The method of introduction
was as used in Experiments VIII, IX, and X, ex-
cept that all trials were conducted during one
session. Observations were made to detect any
evidence of a preference which might be shown
between the two baits.

The fish baits were hit first 8 times, while the
mollusks were hit first 9 times, indicating that
there was no visual preference. However, al-
though both the fish and mollusks were taken
into the mouth with equal vigor, the fish were
presumably swallowed while the mollusks were
rejected almost immediately. The results were
comparable to those of Experiment X, with the

PACIFIC SCIENCE, Vol. XVII, April 1963

response toward the mollusks similar to that
shown toward the treated bait. On one occasion
a single shark took the Tridacna and then the
grouper. After a short period, with both baits
in its mouth, the shark rejected one, presumably
the Tridacna. These same rejected baits were
subsequently taken by groupers and snappers
which waited below the feeding sharks. Al-
though the mollusks tested were unacceptable
as food by these sharks, it is known that they
will feed readily on squid.

It was possible, therefore, to observe a pref-
erence when one of the bait-choices offered was
unacceptable to the sharks. However, when both
bait-choices presented were acceptable, it be-
came difficult to make this distinction, even
though one might have been significantly more
attractive than the other. Thus, most of our ex-
periments concerning food preferences yielded
inconclusive results. For example, a test might
have been conducted to determine the compara-
tive attractiveness of two baits, A and B, both
motionless and presented on the lines within
10 ft of each other. As indicated in Experiment
VIII, although these sharks might have been
drawn in by an olfactory stimulus produced by
bait A, they would then have been quite likely
to have hit B inasmuch as the final phase of the
approach to the bait would have been visually
directed. Once having taken bait B, this bait
would be retained as long as it was not actually
unacceptable.

It has been noted (Tester, 1963) that some
types of fish flesh appear to be more attractive
to sharks than do others. In the ponds at Coco-
nut Island the notably dry-fleshed snapper, Lu-
tianus gib bus, appeared to be far less desirable
to captive sharks as food than did the much
juicier tuna, Katsuwonus pel amis. Springer
(1958) also noted this preference for tuna. Is
it possible that this apparent preference is ac-
tually due to a higher concentration of some
basic attractant which is common to the flesh
of both fish? It has become increasingly appar-
ent that some substance (or substances) in fish
flesh is perceived by the sense of taste and/or
smell of these sharks which is a powerful ele-
ment in the release of a highly motivated feed-
ing pattern. Furthermore, it was indicated in
Experiment X and also in the work of Tester
et al. (1955) that this substance could be ex-

Feeding Behavior of Sharks — Hobson

191

tracted from the flesh, whereupon the flesh it-
self was left undesirable to the sharks. Tester
et al. (1955) expended considerable effort in
attempts at purification, fractionation, and iden-
tification of the attractant extracted from fish
flesh which released a response in captive tuna.
While the precise identity of the substance
eluded the investigators, many of its chemical
properties were determined and presented by
the authors. Tester et al. (1954) suggest that
the so-called attractant which is present in the
body juices of fish, squid, shrimp, and other
forms may be a substance (or substances) which
is common to all these forms.

FEEDING DETERRENTS

The recent increased interest in the habits
and behavior of sharks has stemmed largely
from a growing awareness of the need to de-
velop a more effective means of protecting hu-
mans from shark attack.

A number of tests of proposed repellents were
conducted, including a test of the repellent now
in general use by the armed services. The re-
sults of some of these tests illustrate many of
the problems involved in producing an effective
repellent.

Experiment XII

This experiment tested the effectiveness of
the standard shark repellent (copper acetate-
nigrosine dye) in protecting both dead and
wounded fish from attack by these sharks.

The experiment was conducted at two loca-
tions: (1) on the bottom at the edge of Deep
Channel, and (2) from the raft. In the tests
on the bottom, involving whitetips, three baits
were tied to packages of repellent and anchored
on the bottom, while seven baits, without re-
pellent, were anchored nearby. In the tests from
the raft, which involved grey sharks, all the
baits were tied to packages of repellent and
lowered halfway to the bottom. No attempt was
made to attract sharks before beginning these
tests.

The whitetips took all of the unprotected
baits but did not take any of those tied to the
packages of repellent. On the other hand, al-
though the first grey to appear during the tests
at the raft occasionally exhibited a slight hesi-

tation when making its approach, it was never
long before several greys were in the area and
the bait and repellent package were quickly
taken together. The repellent packages were
shortly rejected, but there was no indication of
a subdued appetite. One grey was seen swim-
ming away trailing a black cloud of shark re-
pellent from its gills with no show of discom-
fort.

The relative effectiveness of the repellent in
protecting the baits from the whitetips may be
related, at least in part, to the absence of the
group feeding effect in this species. This effect,
which was discussed earlier, was noted to lower
the threshold for the release of feeding patterns
in grey sharks.

Experiment Kill

In this experiment the standard shark repel-
lent was presented as a large cloud, both by it-
self and as a protective screen for various at-
tractants. Trials were conducted at the raft in
which 30 gm of repellent were dissolved in 64
gal of sea water in a plastic container. This solu-
tion was presented alone as well as mixed with
each of the following attractant materials: (1)
two small macerated goatfish, (2) 500 ml fish
extract, (3) 1,000 ml of fish extract, and (4)
2,000 ml of fish extract. The extracts were pre-
pared by using 50 gm of grouper flesh per 500
ml of fresh water.

The presentation was made by simply pour-
ing the contents of the container into the water
after a number of sharks had been drawn in
about the raft and noting the results. In each
case, a cloud approximately 6 ft deep and 12
ft wide formed at the surface next to the raft
and slowly drifted downstream.

When the plain repellent cloud was intro-
duced, the sharks rapidly converged on the cloud
to a distance of approximately 5 yd where they
circled slowly, following the cloud as it drifted
downstream. After a few minutes, however, the
sharks lost interest in the cloud and returned to
the raft. When the repellent cloud containing
the macerated goatfish was introduced, the fish
fragments sank beneath the cloud where they
were quickly taken by the sharks. The sharks,
then excited, swam unhesitatingly up into the
cloud where they circled vigorously. Sixteen

192

greys, 3-6 ft long, were involved in the first two
phases of this experiment.

The response to the repellent clouds contain-
ing the extract solutions was essentially the same
in each case. After rapidly approaching the
cloud, the sharks circled briefly about its edges
and then moved slowly inside without notable
excitement. Within 5 min of introduction, all
sharks had lost interest in the introduction.
Eight greys, 3-6 ft in length, were involved in
these tests.

It seems noteworthy that the extract elicited
an unexcited pattern of exploratory behavior
rather than the highly excited aggressive be-
havior seen following the introduction of mac-
erated goatfish. In any case, the most that can
be said for the repelling qualities of the material
used in this test is that the sharks did not swim
into the cloud without incentive to do so. It
may theoretically have served to conceal an
otherwise attractive visual stimulus which might
have been inside.

Experiment XIV

This experiment tested the effectiveness of
an underwater light suspended on a line in
protecting dead fish.

The apparatus was designed so that the bait
(dead goatfish) was suspended 2 ft below the
light in a position where both could be viewed
from the underwater chamber. Observations
were then made of the response of the sharks
to the bait, both in the presence and in the ab-
sence of light. Tests were conducted during four
different nights using both 110- and 300-watt
bulbs. A steady light was used, as well as a light
which flashed at rates of approximately 15, 35,
and 50 flashes per min. Periods of light, 20 min
long, were alternated with 20-min periods of
darkness during both of which the baits were
presented. Sharks had been drawn to the area
by bait placed in a wire basket which was sus-
pended in the water and then raised prior to
each test. A strong current was flowing toward
Deep Channel during each test.

Initially the steady light was effective in pro-
tecting the bait from the sharks for the entire
20-min period, with the bait being taken soon
after the light was extinguished. During the
early periods of light the sharks milled about

PACIFIC SCIENCE, Vol. XVII, April 1963

at the edge of the field, consistently downstream
of the bait and light. Upon continued contact
with the light the sharks became progressively
bolder until despite the light, flashing or steady,
they showed little hesitation in taking the bait.
This boldness seemed to increase notably with
the number of sharks present. There were
usually between 5 and 10 grey sharks present
during these tests; no other species was seen.
This test proved to be a good illustration of
progressive loss of apprehension through grow-
ing familiarity with an initially strange situa-
tion, as well as the mutually stimulating effect
of some species of sharks on one another in
certain situations.

CONCLUSIONS

1. Three species of sharks are common in
Eniwetok lagoon: the grey shark, Car char hinus
menisorrah; the blacktip shark, C. melanopterus;
and the whitetip shark, Triaenodon obesus.

2. Each species inhabits a rather characteris-
tic habitat within the lagoon: (a) the blacktip
in relatively shallow water ( 1-40 ft approx. )
over sand and coral rubble flats which extend
out from shore for distances ranging from a few
yards to several miles; (b) the grey, along the
outer slope of these flats, in deeper water and
in the passages to the sea; and ( c ) the whitetip
about rock-ledges and coral heads.

3. The feeding behavior of these sharks shows
differences which in many cases appears to be
associated with their characteristic habitats.

4. Blacktip and grey sharks show a marked
increase in excitement when feeding in num-
bers. This phenomenon, often referred to as a
"feeding frenzy” in extreme cases, does not seem
to occur in the whitetip.

5. All three species are highly sensitive to
stimuli emanating from or suggesting injured
and/or distressed, as well as dead or moribund
prey.

6. These sharks can detect, by olfaction, both
injured fish and uninjured fish in a state of

stress.

7. Olfaction is the most effective sense in
detecting prey at a distance, providing the ol-
factory stimulus has had sufficient time and
means for effective dispersal.

Feeding Behavior of Sharks — Hobson

193

8. These sharks are able to follow an olfac-
tory stimulus quickly and directly to its source
without benefit of other orienting stimuli when
the olfactory material has been drawn out as a
narrow trail by a strong current or, in standing
water, when the source of the olfactory stimulus
is moving.

9. Generally, olfactory stimuli are effective

in releasing a highly motivated pattern of ex-
ploratory behavior in grey sharks, which is ap-
petitive in nature. Consummation of the feeding
act then requires a subsequent specific stimulus,
as for example one of vision.

10. The grey shark is often highly excited by,
and attracted to, the source of rapid and/or er-
ratic movement. The effect is intensified in the
presence of certain olfactory stimuli but is not
dependent on their presence.

11. Despite displaying varying degrees of ini-
tial curiosity toward many unusual stimulus situ-
ations arising in their environment, the sharks
exhibit caution when encountering a situation
which is unfamiliar. This caution will steadily
subside, however, with an increasing familiarity
with the situation.

12. The final phase of approach to a motion-
less prey by the grey shark is normally directed
by vision, even though the feeding pattern may
have been initially released by some other sen-
sory modality, for example, olfaction.

13. Although vision is an important sense in
the feeding activity of these sharks, a high de-
gree of acuity and form discrimination is not
demonstrated. Rather, the significant visual cues
seem to involve the detection of movement or
contrasting brightness, or both.

14. These sharks may attack any object which
they sense in a high concentration of an olfac-
tory material.

15. The grey shark rejects, from the mouth,
food which does not permit acceptable gustatory
stimulation.

16. The standard shark repellent (copper
acetate-nigrosine dye) now in use by the armed
forces is ineffective in preventing the grey shark
from swimming into a cloud of this material
when the species is present in numbers and is
motivated by food within and adjacent to the
material.

ACKNOWLEDGMENTS

The author is indebted to Dr. Albert L. Tester
for many helpful suggestions during the course
of the investigation and for his constructive
criticism of the manuscript. The study would
not have been possible without assistance in
the field from Dr. Tester and Mr. Francois Mau-
tin, as well as from Mr. John Kay, Mr. Bryant
Sather, Mr. Susumu Kato, Mr. Taylor Pryor,
and Mr. Albert Smith, graduate students, and
Mr. Henry Yoshihara, undergraduate student,
all of the University of Hawaii. The facilities
at Eniwetok were made available by Dr. Robert
W. Hiatt, Director of the Eniwetok Biological
Laboratory, as well as by the Atomic Energy
Commission, the Pacific Missile Range Facility
of the United States Navy, and by the Holmes
and Narver Construction Company. The author
also wishes to thank Dr. William A. Gosline
and Dr. Ernst S. Reese of the Department of
Zoology and Entomology, University of Hawaii,
and Dr. Donald W. Strasburg of the Honolulu
Biological Laboratory, United States Fish and
Wildlife Service, for critically reviewing the
manuscript.

REFERENCES

Budker, P. 1938. Les cryptes sensorielles et les
denticles cutanes des plagiostomes. Ann. Inst.
Oceanog. 18:207-288.

Eibl-eibesfeldt, I., and H. Hass. 1959. Er-
fahrungen mit Haien. Zeit. Tierpsych. 16
(6): 733-746.

Gilbert, P. S. 1961. The visual apparatus of
sharks and its probable role in predation. Ab-
stracts of Symposium Papers, Tenth Pacific
Science Congress, Honolulu, Hawaii, pp. 176—
177.

Halstead, B. 1958. The relationship of trau-
magenous fishes to man. A.I.B.S. Shark Con-
ference, New Orleans, Apr. 8-11, 1958 (MS).

Harry, R. R. 1953. Ichthyological field data of
Raroia Atoll, Tuamotu Archipelago. Atoll
Res. Bull. 18:1-190.

194

Hass, H. 1951. Diving to adventure. Doubleday
and Co., Garden City. 280 pp.

Hobson, E. S., F. Mautin, and E. S. Reese.
1961. Two shark incidents at Eniwetok Atoll,
Marshall Islands. Pacific Sci. 15 (4): 605-609.

Kato, Susumu. 1962. Histology of the retinas
of the Pacific sharks, Carcharhinus melanop -
terus and Triaenodon obesus. M. S. Thesis,
University of Hawaii, Honolulu, Hawaii.

Klausewitz, W. 1958. Die Atoll-Riffe der
Malediven. Natiir und Volk 88 (11): 380 —
390.

1959. Fische aus dem Rotenmeer. I.

Khorpel Fische. Senck. biol. 40:43-50.

Kritzler, H., and L. Wood. 1961. Provisional
audiogram for the shark Carcharhinus leucas.
Science 133 (3463) : 1480-1482.

Limbaugh, C. 1958. Abstracts from field notes
on sharks. A.I.B.S. Shark Conference, New
Orleans, Apr. 8-11, 1958 (ms).

Lineaweaver, T. I960. Shark! Sports Illus-
trated, Feb.: 54-63.

Parker, G. H. 1903. Hearing and allied senses
in fishes. Bull. U. S. Fish. Comm. 1902:45-64.

1911. Influence of the eyes, ears, and

other allied sense organs on the movements
of the dogfish, Mustelus canis. Bull. U. S. Bur.
Fish. 29:43-57.

1912. Sound as a directing influence in

the movements of fishes. Bull. U. S. Bur. Fish.
30: ( 1910) :97-104.

1914. The directive influence of the

sense of smell in the dogfish. Bull. U. S. Bur.
Fish. 33:61-68.

PACIFIC SCIENCE, Vol. XVII, April 1963

Randall, J. E. 1955. Fishes of the Gilbert Is-
lands. Atoll Res. Bull 47:1-243.

Sheldon, R. E. 1911. The sense of smell in
selachians. J. Exper. Zook 10:51-62.

Schultz, L. P., E. S. Herald, E. A. Lachner,
A. D. Welander, and L. P. Woods. 1953.
Fishes of the Marshall and Marianas Islands.
Bull. U. S. Nat. Mus. 202(1): 1-685.

Springer, S. 1958. Field observations on large
sharks. A.I.B.S. Shark Conference, New Or-
leans, Apr.: 8-11 (ms).

Strasburg, D. W. 1958. Distribution, abun-
dance, and habits of pelagic sharks in the
central Pacific Ocean. Fishery Bull. U. S. Fish
Wildlife Serv. 58 ( 138) : 335-361.

Tester, A. L. MS. The olfactory and gustatory
apparatus in sharks.

1963. The role of olfaction in shark

predation. Pacific Sci. 17(2).

, and S. KATO. ms. Form and color dis-
crimination by blacktip and grey sharks ( Car-
charhinidae).

, H. Yuen, and M. Takata. 1954- The

reaction of tuna to stimuli 1953. U. S. Fish
Wildlife Serv. Spec. Sci. Rpt. Fish. 134:1-33.

, P. B. Van Weel, and J. J. Naughton.

1955. Reaction of tuna to stimuli 1952-53.
Part 1. U. S. Fish Wildlife Serv. Spec. Sci.
Rpt. Fish. 130:1-61.

Wright, B. S. 1948. Release of attack behavior
patterns in shark and barracuda. J. Wildlife
Mgt. 12 (2): 117-123.

Life History of the Caligid Copepod
Lepeophtheirus dissimulatus Wilson, 1905 (Crustacea: Caligoida)1

Alan G. Lewis2

Copepods of the family Caligidae are found as
external parasites on both fresh and salt water
fishes throughout the world. Even with their
wide distribution, relatively little is known
about their life history. The presentation of the
life history of Lepeophtheirus dissimulatus , a
caligid found on salt water fishes, should pro-
vide more information on the life history and
general biology of this group of copepods.

Although there are many references to im-
mature and juvenile or larval stages (reviewed
by Heegaard, 1947 : 2 1-3 1 ) , only a few workers
have had the material to describe the complete
life history of a caligid. Through methods de-
scribed below, the author was able to obtain a
series of Hawaiian specimens representing all
of the stages of development of L. dissimulatus.

Lepeophtheirus dissimulatus is one of the
more common parasitic copepods of Hawaiian
acanthurid (surgeon) fishes. The species is not
limited only to Hawaiian fishes but is known
from a wide variety of localities and a wide
variety of hosts (Lewis, in press). Even with
its wide distribution, in both the Atlantic and
Pacific oceans, larval stages have been described
only once, by Shiino (1959:309), from fishes
of the Revilla Gigedo Islands.

Attempts to rear the copepod from the egg to
the adult in culture were only partially success-
ful. Ovigerous adult females were placed in clean
finger bowls partially filled with cotton-filtered
or unfiltered sea water. Circulation of the water
was accomplished by means of a fine jet of
air from a slender glass tube immersed in the
medium. The water was changed at least once

1 A portion of a thesis submitted to the University
of Hawaii in partial fulfillment of the requirements
for the degree of Doctor of Philosophy. Contribution
number 181, Hawaii Marine Laboratory. Manuscript
received November 23, 1961.

2 Department of Zoology, University of New Hamp-
shire, Durham, New Hampshire.

every 8 hr and usually more frequently. At the
time the eggs, carried by the female, hatched,
a portion of the first nauplii, the first larval
stage, was removed with a wide-mouthed pipette
and placed in small stender dishes. The remain-
ing nauplii were left in the finger bowl and
the adult female was removed and placed in a
separate bowl. Water in the finger bowls was
circulated in the same manner as above and
changed at the same time intervals. Water in
the stender dishes was not circulated but instead
was changed at more frequent intervals, every
3-4 hr. As moulting occurred into the second
naupliar stage, the newly moulted second nau-
plii were transferred to new finger bowls or
stender dishes except for three to four indi-
viduals which were preserved in either 10%
buffered formalin or 95% ethyl alcohol. Shortly
after the second naupliar stage moulted into the
succeeding stage, the copepodite, a piece of
loosely woven nylon cloth was placed in the
dishes to offer a substrate for attachment. (It
was found that the copepodite stage is the stage
which attaches to the host.) In addition, pieces
of host tissue were placed in some of the dishes.
The host tissue was left in the dishes for 1 hr
and then removed and the animals transferred
to a new dish in order to reduce contamination
and bacterial action.

The mortality rate between the first and sec-
ond naupliar stages was high and appeared to
be partially due to the inability of the nauplius
to break out of the old cuticle. The rate from
the second naupliar stage to the copepodite was
relatively low. Out of the 30 attempts that were
made to rear the copepod, each with at least one
female carrying from 20-60 eggs, one copep-
odite was obtained that attached to the nylon
cloth placed in the dish. The remaining copep-
odites died without attaching and the single
copepodite that attached died immediately
thereafter. None of the copepodites attached to
the host tissue placed in the dishes.

195

196

PACIFIC SCIENCE, Vol. XVII, April 1963

endo.

~exa

Fig. 1 . Hypothetical thoracic leg of Lepeophtheirus
dissimulatus, showing the various armament compo-
nents. C, Heavy setae; c, fine setae; endo, endopodite;
exo, exopodite; f, membrane; H, well-developed spine;
H', laterally projecting spine appearing as a continua-
tion of segment; h, poorly developed spine; Hm, mem-
brane margined spine; P, large plumose setae; P', large
naked setae; p, small plumose setae; p’ , small naked
seta; prot, protopodite; Q, seta (or spine) plumose
on one side and membrane-margined on other; rh,
spinule; s, hairlike seta; sp, sternal plate.

Because of the failure of the culture material
and in order to obtain a complete timed series
of developmental stages, attached copepodites
were taken from host material and placed in in-
dividual stender dishes. After these copepodites
moulted into the first chalimus (the following
stage ) , the time was noted until the succeeding
moult, into the second chalimus. To ascertain
the time passed in each stage after the first
chalimus, specimens of the previous stage were
taken from the host and treated as above. Due
to the mortality rate under laboratory condi-
tions, many specimens of each stage had to be
collected so that one would pass through the
succeeding stage. Because of the inability to get
any number of specimens to pass through each
stage and due to the necessity of submitting the
larvae to laboratory conditions, the time indi-
cated for each stage in Figures 2-7 should be
regarded as indicative, not conclusive.

The descriptions of the body, appendages,
and processes of all of the developmental stages
and the adult is followed by a discussion of the
general behavior of the various developmental
stages. The graphical figures included in the

text (Figs. 2-7) give the means and ranges of
various measurements of the larval forms.

Larval specimens other than nauplii were
killed and preserved in 95% ethyl alcohol. Speci-
mens to be dissected were placed in 85% lactic
acid to soften and clear them, stained with
methyl blue dissolved in 85% lactic acid, and
dissected in clear 85% lactic acid.

Drawings of the total animal were made from
specimens placed in 85% lactic acid and cov-
ered with a 9 mm cover slip. The cover slip
was raised slightly above the younger specimens
by thin pieces of glass so that the specimen was
not distended. A camera lucida and a Bausch
and Lomb arc projector were used to make the
drawings of the total animal. Drawings of the
appendages and processes were made, with the
use of a camera lucida, from appendages and
processes either in situ on the whole mount or
dissected off and mounted in Hoyer’s mounting
medium. Measurements were made with an ocu-
lar micrometer.

The terminology used, with some exceptions
which are given in the text, is that used in
Lewis (in press). To facilitate the use of the
thoracic leg tables, a hypothetical thoracic leg is
shown in Figure 1, giving all of the component
parts of the armament of the thoracic legs dis-
cussed in tabular form in this publication. Fur-
ther, a table of the hypothetical thoractic leg
is given below in Table 1.

3.5

3.0

2.5

.§2.0

£ 1.5

1.0

0.5

8 -mean

t t

"8*0 120 160 200 240 280

Time in Hours

Fig. 2. Greatest length of body, excluding setae, in
developmental stages of L. dissimulatus. n, Nauplius;
co, copepodite; ch, chalimus.

Lepeophtheirus dissimulatus — Lewis

197

3.5]

40 80 120 160 200 240 280

Time in Hours

FIG. 3. Greatest length of cephalothorax, including
frontal plates, of developmental stages of L. dissimu-
latus. co, Copepodite; ch, chalimus.

DEVELOPMENT AND DESCRIPTION OF
LIFE HISTORY STAGES

As the eggs are extruded out of the oviducal
opening at the posterior end of the genital seg-
ment they are fertilized by sperm from the
seminal receptacle (Wilson, 1905:526). Two
spermatophores are implanted by the male on
the external posterior ventral surface of the fe-
male genital segment and open into the two
oviducts through the seminal receptacles (Wil-
son, 1905:527). The disk-shaped fertilized eggs
are extruded as a uniseriate chain (Fig. 8 a).
Each egg is wrapped in a cuticular material,
presumably secreted by a cement gland in the
genital segment of the female, that remains at-
tached to the female until after the eggs hatch
(Fig. 8£). The covering appears to serve as a
protective structure for the eggs and developing
embryos and as a means of connecting the eggs
together in a string.

Because of the flattening of the fertilized
eggs, development is difficult to observe. From
13-20 hr (at room temperature, approximately
23 C) after extrusion, a reddish-colored eye
spot becomes visible on the yellowish or whit-
ish embryo. From 30 to 40 hr (at room tem-
perature) after extrusion, movement is visible
within the egg. As the movement of the em-
bryo increases, the egg expands and finally rup-
tures. Hatching occurs regularly, the distal egg

hatching first, the proximal last. A lapse of be-
tween 2 and 10 min occurs between the hatch-
ing of one egg and the hatching of the succeed-
ing one. In almost all cases, the egg chamber
ruptures on the inner side of the egg string
(i.e., the surface nearest the median longitudi-
nal axis of the female). The first nauplius, by
sporadic violent movements of its appendages,
works its way out of the egg case and, after a
short period of rest in which some swelling
occurs, assumes the characteristic jerky, free-
swimming habit of the naupliar stages.

Body

The externally unsegmented body of the first
nauplius (Fig. 8c) is of a general obovoid shape
in both dorsal and lateral aspects. The greatest
length is slightly more than twice the greatest
width, and the anterior end is broadly curved.
The lateral margins of the posterior one-half
of the body curve inward regularly to the ir-
regularly-margined posterior end. The posterior
end of the body has a slight median projection
and two indentations, one on each side, from
which the balancers arise. The balancers (Fig.
16k) are unsegmented, slightly curved, rodlike
structures that project laterally and have their
distal end slightly flattened.

The cuticle of the first nauplius is, in most
specimens, slightly larger than the inner body;

3.5

3.0

2.5

I20

I15

1.0

0.5

* It

f <= 1

80 120 160 200
Time in Hours

240 280

Fig. 4. Greatest width of body, excluding marginal
flanges, of developmental stages of L. dissimulatus .
n, Nauplius; co, copepodite; ch, chalimus.

198 PACIFIC SCIENCE, VoL XVII, April 1963

TABLE 1

Armature of Hypothetical Thoracic Leg Shown in Figure 1

MARGIN

STERNAL

PLATE

PROTOPODITE

1 2

EXOPODITE

1 2 3

ENDOPODITE

2 3

Outer

lrh

2H,lHm,lQ

c,C

c,lh

c,2p',2P

Inner

f,lp,ls

1H' c,lP

c,5P

c,2P

a distinct space is visible between the anterior
end of the body and the cuticle. There is in-
distinct evidence of internal segmentation pos-
teriorly. The general body color in life is light
yellow with numerous red and blue pigment
spots scattered over the surface. The eyes are
visible as two minute, clear, round lenses, one
on each side of an irregular, reddish-pigmented
region in the anterior portion of the body. The
alimentary tract is not visible, neither are the
mouth nor the anus. Large globules of material,
presumably yolk, are evident, clustered in an in-
distinctly delimited, oblong mass extending pos-
teriorly from just behind the ocular region.

The first nauplius possesses three sets of ap-
pendages (described later), the uniramous an-
tennules, biramous second antennae, and bira-
mous mandibles in this order proceeding poste-
riorly. All of these appendages possess long,

Fig. 5. Greatest width as a per cent of total body
length, excluding setae, of developmental stages of L.
dissimulatus. n, Nauplius; co, copepodite; ch, chalimus.

lightly plumose setae that presumably function
as flotation mechanisms and in swimming.

The first naupliar stage lasts for 41/2-13 hr
with an average of 6V2 hr (based on nine hatch-
ings at room temperature, approximately 23 C).
After this time the movement of the animal
decreases slightly and the cuticle splits longi-
tudinally at the dorsal anterior end. The nau-
plius then wriggles violently and breaks through
the split, freeing itself from the cuticle.

The body of the second nauplius (Fig. 8 d, e)
is similar to that of the first, although the length
from the mandibles to the posterior end of the
body is greater. The greatest length, excluding
the appendages, is almost 2Vi times the greatest
width. The body is very slightly flattened dor-
soventrally in cross section, the anterior end is
broadly curved, and the lateral margins are flatly
convex. The posterior end of the body is similar
to that of the first nauplius although the bal-
ancers are slightly larger and arise from two
small, circular, platelike processes in the pos-
terior lateral indentations (Fig. 16/).

The bodies of older second nauplii have a
small space between the anterior end and the
cuticle. There is distinct evidence of internal
segmentation, similar to the segmentation of
the copepodite, visible inside the cuticle of older
specimens but not visible on the external sur-
face. The alimentary tract is indistinct and in-
complete, visible only as a cluster of yolk glob-
ules along the median longitudinal axis just
posterior to the ocular region.

The basic structure of the three sets of ap-
pendages of the second nauplius is similar to
that of the first. The armature, however, is
slightly different. This stage lasts for 91/2-19
hr with an average of 14V2 hr (based on eight
hatchings at room temperature, approximately
23 C). Just prior to moulting into the copep-
odite stage, the movement of the animal be-

Lepeophtheirus dissimulatus — Lewis

199

1.00

075-

50.50

0.25

t I

120

160 200 240

Time in Hours

280

Fig. 6. Greatest length of genital segment of de-
velopmental stages of L. dissimulatus. co, Copepo-
dite; ch, chalimus; +, measurement includes both
genital segment and abdomen (one segment in early
stages).

comes sporadic, stopping at times and then
becoming vigorous. During one of the vigorous
periods of movement a crack appears in the
anterior dorsal portion of the cuticle and the
copepodite wriggles out in the same manner as
in the previous moult.

The change from the second nauplius to the
copepodite is the greatest change that occurs in
a single moult during development. In one
moult the body is divided from an externally
unsegmented structure into a cephalothorax con-
sisting of the head, maxilliped-bearing segment,
and first pedigerous segment. Further, four free
segments are visible posterior to the cephalo-
thorax. The first of these segments bears the
second thoracic legs, the second bears a pair of
posteriorly directed spines that may be the pred-
ecessors of the third thoracic legs, the third free
segment is naked but will later give rise to the
fourth thoracic legs, and the fourth free seg-
ment, to which the caudal rami are attached,
will later differentiate into the genital segment
and abdomen.

Not only does the body break up from a rela-
tively undifferentiated second nauplius to a co-
pepodite with a composition comparable to
that of the adult, but also most of the append-
ages and processes are now present. Besides the

antennules, second antennae, and mandibles that
were present on the first and second nauplius,
the postoral processes, maxillae, maxillipeds,
and first two pairs of thoracic legs have been
added and the caudal rami are now present.

The general shape of the copepodite body
(Fig. 8/) is ovoid in both dorsal and lateral
aspects. The greatest width, that of the cephalo-
thorax, is less than one-half the greatest length
(Fig. 5). The anterior cephalothoracic margin
is sharply convex and the lateral margins are
flatly convex. A pair of V-shaped indentations,
or sinuses, are visible in the posterior lateral
margin of the cephalothorax. These posterior
sinuses are found at the junction of the posterior
surfaces of the median and lateral cephalotho-
racic areas and suggest that the lateral areas are
distinct from the median and are an outgrowth
of the cephalon. This is further indicated by
the longitudinal cephalothoracic grooves which,
in the copepodite, run anteriorly for a very short
distance from the apex of the posterior sinuses.
The median cephalothoracic area does not ex-
tend past the posterior lateral area and its pos-
terior margin is irregular. The eyes are distinct
and similar to those of the naupliar stages.

The free second pedigerous segment is wider
than long, its greatest width being 1.3 times the
length. Both the anterior and posterior margins

I.OOi

0.75

JO 0-50

•H

-O

0.25-

40 80 120 160 200 240 280

Time in Hours

Fig. 7. Greatest width of genital segment of de-
velopmental stages of L. dissimulatus. co, Copepodite;
ch, chalimus.

200

PACIFIC SCIENCE, VoL XVII, April 1963

are distinct, the anterior being irregularly con-
vex and the posterior irregularly concave; the
lateral margins are broadly convex. The free
third pedigerous segment is approximately two-
thirds the width of the second and is slightly

wider than long. The lateral margins of the seg-
ment are broadly convex in younger specimens,
straight in older; the width of the posterior end
of the segment is slightly greater than that of
the anterior in older specimens but equal to or

2mm ' 02m

/ *

Fig. 8. Egg string of L. dissimulatus and body of developmental stages from nauplius to first chalimus.
a. Posterior end of ruptured egg string, showing first nauplius just emerging; b, empty egg string after hatch-
ing; c, dorsal view of first nauplius; d, dorsal view of early second nauplius; e, dorsal view of late second
nauplius; f, dorsal view of copepodite; g, dorsal view of first chalimus. Stippled circles, yolk material.

Lepeophtheirus dissimulatus — Lewis

201

slightly less than the width of the anterior end
in younger specimens. The third pedigerous seg-
ment bears a single naked, spinelike process
projecting from each of the posterior lateral
corners (Fig. 12^). The relationship of this
process to the future third thoracic leg is un-
known although Heegaard (1947:63, fig. 8)
figures a pair of lobate appendages in the same
region as the spines and indicates that they be-
come the third thoracic legs of Caligus curtus.
The free fourth pedigerous segment is short, its
greatest width being almost 3 times its length.
The anterior margin is concave, the posterior
convex. There is no indication of the fourth
thoracic legs.

The free fourth segment, the future genital
segment and abdomen, is slightly shorter than
the combined lengths of the third and fourth
pedigerous segments, and the width is slightly
greater than the length. The lateral margins are
flatly convex, and the lateral posterior margins
are concave, forming the attachment surface,
and possibly an articulation surface, for the
caudal rami. The posterior medial margin of the
segment is bilobed, and the anal region is evi-
dent as a minute, V-shaped depression between
the lobes.

The copepodite possesses a spikelike rostrum
on the anterior end of the cephalothorax (Fig.
13 a). The frontal organ is now visible, although
rather indistinctly, and is represented by two
large ovoid bodies at the anterior end of the
cephalothorax that are connected to the rostrum
by a slender, tubelike process. The alimentary
tract is visible as two convex lines extending
from just behind the eyes posteriorly for ap-
proximately one-third the length of the cepha-
lothorax. The tract is visible posterior to the
two convex lines only as a slender mass of yolk
globules along the median longitudinal axis and
extending posteriorly to the free third pediger-
ous segment. There does not appear to be an
oral opening.

The length of time spent in the copepodite
stage is not known due to the inability of get-
ting a copepodite to attach to a substrate and
moult into the first chalimus in the laboratory.
In order to determine the method of attachment,
observations were made on specimens of copep-
odites attached to host material collected in the
field. The copepodite attaches to the host by

means of the second antennae, secretes a frontal
filament, and then after a short period of time
moults into the first chalimus. This process is
described in detail in the portion of the paper
devoted to the general behavior of the develop-
mental stages. The only visible difference be-
tween the free-swimming and attached copep-
odites is the slightly larger frontal organ in
the attached forms. All of the appendages and
processes and the general body structure are
identical. Heegaard (1947:53-65), however,
describes two copepodite stages for Caligus
curtus, the first copepodite moulting into the
second shortly after the host is found. In addi-
tion he describes a "pupal stage” that is passed
within the loosened but unshed second copep-
odite cuticle. Neither the second copepodite
stage nor the pupal stage were found in Lepe-
ophtheirus dissimulatus.

The change from the copepodite to the adult
is gradual for the most part. There are excep-
tions to this in some of the appendages such as
the first and second thoracic legs but, in general,
the transition occurs in a step-by-step manner
through the six chalimus stages which are found
between the copepodite and the adult.

The first chalimus (Fig. 8g), attached to the
host by the frontal filament, is dorsoventrally
flattened. The body is ovoid in a dorsal view
and the greatest length, including the frontal
region but not the frontal filament, is approxi-
mately 1.2 times the greatest width. The cepha-
lothorax consists of the cephalon, maxilliped-
bearing segment, and first pedigerous segment as
in the copepodite. The second, third, and fourth
pedigerous segments, in addition to the single
segment that will later form the genital segment
and abdomen, are free. The anterior margin of
the cephalothorax is broadly curved except for
an irregular median swelling in the region of
the frontal filament. The lateral cephalothoracic
margins flare slightly anteriorly, just posterior
to the antennule base, and then curve flatly pos-
teriorly. The posterior sinuses are distinct but
are formed of the anterior lateral margin of the
second pedigerous segment and the inner mar-
gin of the posterior lateral cephalothoracic area,
not the outer margin of the median cephalotho-
racic area and the inner margin of the posterior
lateral cephalothoracic area as in the copepodite.
The median margin of the thoracic area is flatly

202

convex and extends between the apices of the
posterior sinuses. The eyes are similar to those
of the preceding stages and are situated in the
anterior one-third of the cephalothorax. Cepha-
lothoracic grooves are indistinctly present or
absent although some irregular dorsal grooves
are usually present in the anterior part of the
body.

The free second pedigerous segment is much
wider than in the copepodite, being more than
3 times as wide as long. The lateral margins are
smoothly convex, the posterior margin flatly
convex. The free third pedigerous segment is
approximately 2 Vi times wider than long, the
posterior end is narrower than the anterior, the
lateral margins are flatly convex, and the pos-
terior margin is flat. The third thoracic legs are
now present, the spinelike processes of the co-
pepodite being absent. The fourth pedigerous
segment is shorter than the third, the width is
slightly more than twice the length, and the
posterior region projects laterally slightly at the
junction of the now present fourth thoracic legs.
The posterior margin of the segment is indis-
tinct and irregular.

The combined genital segment and abdomen
(Fig. 12 b), that form the fourth free segment,
are similar in both shape and measurements to
that of the copepodite.

The frontal region (Fig. 13£) is slightly ir-
regular; the rostrum of the copepodite is absent.
The frontal organ consists of two lobate proc-
esses at the base of the frontal filament, an ad-
ditional two lobate processes attached to the
posterior surface of the first two, and an irreg-
ular indistinct area posterior to the second set
of processes. The frontal region is not distinctly
separated from the cephalothorax although an
indistinct groove is present on the ventral sur-
face, extending from the lateral margin, in the
region of the antennules, posteriorly and then
across the ventral surface.

The first chalimus stage lasts for approxi-
mately 40 hr (based on two specimens kept at
room temperature, approximately 23 C) at
which time the specimens moult into the more
elongate second chalimus.

The general shape of the second chalimus
(Fig. 9 a, b) is similar to that of the first chal-
imus. The cephalothorax now includes the sec-
ond pedigerous segment in addition to the first.

PACIFIC SCIENCE, Vol. XVII, April 1963

Late in this stage, however, the division be-
tween the cephalothorax and the third pediger-
ous segment becomes indistinct (Fig. 9b). The
greatest length of the cephalothorax, including
the frontal region, is approximately IV2 times
the greatest width, considerably longer than the
cephalothorax of the first chalimus. The increase
in length is presumably due to the inclusion of
the second pedigerous segment in the cephalo-
thorax. The anterior margin of the cephalo-
thorax is broadly rounded except for the median
protrusion of the frontal filament; laterally, the
margin turns posteriorly sharply, then flares
slightly to the lateral cephalothoracic margins.
The lateral margins are wavy in some specimens
and smooth in others although the entire mar-
gin is generally convex. The posterior cephalo-
thoracic margin is four-lobed in the early second
chalimus, with two large lobes in the lateral
region of each side. In older specimens of this
stage, in which the division between the cepha-
lothorax and the third pedigerous segment is
indistinct, the inner two lobes are not visible
and the median cephalothoracic area, the third
pedigerous segment, projects well past the lat-
eral areas. The posterior sinuses are distinct as
two small, V-shaped depressions between the
two lobes in the younger specimens and between
the inner margin of the lateral areas and the
outer margin of the median cephalothoracic area
in the older specimens. Thus far then, the pos-
terior sinuses appear to be due to the unequal
growth of the lateral and medial regions of
the cephalothorax. The dorsal cephalothoracic
grooves are indistinct, consisting of a single,
very short groove extending anteriorly from the
apex of each of the two posterior sinuses.

The third pedigerous segment, free in the
early second chalimus, tapers evenly from the
junction with the second pedigerous segment
to the junction with the free fourth pedigerous
segment. The greatest length of the segment is
approximately four-fifths the greatest width and
about one-fifth that of the cephalothorax. The
free fourth pedigerous segment flares outward
from the junction of the third to the attachment
of the fourth legs, at the posterior end of the
segment. The greatest width of the segment is
approximately 1.6 times the greatest length,
slightly more than one-half that of the third
pedigerous segment. The division between the

Lepeophth eirus dissimulatus — Lewis

203

third and fourth pedigerous segments is distinct,
between the fourth and the genital-abdominal
segment also distinct although sometimes faint
in younger specimens.

The combined genital segment and abdomen
(Fig. 12c) is slightly shorter than the third

pedigerous segment, and its greatest width is
slightly more than its length. The lateral mar-
gins are flatly convex; the lateral posterior mar-
gins are flat or slightly concave and form the
place of attachment for the caudal rami. The
anal laminae are visible as two knoblike projec-

Fig. 9- Body of second, third, and fourth chalimus developmental stages of L. dissimulatus. a , Dorsal view
of early second chalimus; b, dorsal view of late second chalimus; c, dorsal view of third chalimus; d, dorsal
view of fourth chalimus. Stippled circles, yolk material.

204

tions and form the middle of the posterior sur-
face of the segment.

The frontal organ of the second chalimus
(Fig. 1 3c) is similar to that of the first chalimus
although more distinct. The lobes at the proxi-
mal end of the frontal filament are contiguous
along the median longitudinal axis of the body,
forming a single ovoid structure; the lobes pos-
terior to the first set are slightly smaller than
in the preceding stage. The alimentary tract
now appears complete although the anterior end
is still filled with yolk granules.

The 10 specimens maintained successfully in
the laboratory, at room temperature (approxi-
mately 23 C), remained in the second chalimus
for about 42 hr.

The third chalimus (Fig. 9c) is somewhat
wider than the second. The greatest length, in-
cluding the frontal region but not the frontal
filament, is approximately 1.3 times the greatest
width. The cephalothorax now includes the first
three pedigerous segments, the condition occur-
ring in the adult. The fourth pedigerous seg-
ment is free and forms the connection between
the cephalothorax and the now indistinctly dif-
ferentiated genital segment and abdomen. The
frontal region (Fig. 13 d) is still indistinctly
separated from the cephalothorax. The anterior
margin of the frontal region is flatly tapered
from the projection of the frontal filament lat-
erally. The lateral portions of the frontal mar-
gin are evenly rounded and join the lateral
margins of the cephalothorax at a slight inden-
tation marking the origin of the groove in-
completely dividing the frontal region from the
cephalothorax. The lateral margins of the cepha-
lothorax are smoothly convex, the posterior lat-
eral corners rounded. The posterior sinuses are
distinct and U-shaped; the lateral margins of
the median thoracic area are continuous with
the inner margins of the sinuses. The median
thoracic area extends posteriorly past the poste-
rior extensions of the lateral areas. The reddish-
pigmented regions of the eyes encircle all but
the outer portion of the lenses, a large increase
in the amount of pigmented material over the
preceding stages. The cephalothoracic grooves
are distinct, the major ones forming an H. The
longitudinal grooves of the H extend anteriorly
and curve medially from their origin just lateral
to the apices of the posterior sinuses. The two

PACIFIC SCIENCE, Vol. XVII, April 1963

grooves are connected by an anteriorly curving
cross groove in their anterior region. Other in-
distinct grooves are present in the anterior re-
gion of the cephalothorax.

The free fourth pedigerous segment is short
and partially covered by the posterior portion
of the median cephalothoracic area. The greatest
length of the segment is approximately one-half
the greatest width. The lateral margins are con-
vex; the posterior margin is distinct, completely
separating the segment from the genital seg-
ment.

The genital segment (Fig. 12^) is short, the
length slightly more than one-half the greatest
width, approximately equal to the length of the
fourth pedigerous segment. The lateral margins
are convex anteriorly and indented posteriorly.
The posterior margin is light but distinct, and
the abdomen is now considered distinct from
the genital segment. A pair of one-segmented
dactyliform processes, the fifth legs, are visible
extending posteriorly from the posterior lateral
surface of the genital segment. The distal end
of the fifth leg is rounded and bears two plu-
mose setules.

The abdomen (Fig. 12 d) is short, the great-
est length slightly less than that of the genital
segment, the greatest width approximately 2Vi
times the length. The lateral margins are con-
vex and the posterior margins curve medially to
the bilobed anal region, composed of the anal
laminae.

The frontal organ (Fig. 13 d) is slightly
smaller than that of the second chalimus and
consists of four parts. The first part is formed
of two oval bodies at the base of the frontal
filament, one on either side of the median longi-
tudinal axis, that are contiguous along their in-
ner margins. A somewhat larger circular body
is present, attached to the posterior surface of
the two anteriormost bodies. An elongate, bi-
furcate body is attached to the posterior surface
of the circular body; both distally rounded parts
of the bifurcation extend posteriorly and slightly
laterally into a larger semirectangular fourth
part. The change in the shape of the organ from
the preceding chalimus may be due to a shrink-
ing of the structure, the anterior two parts being
formed from the single ovoid structure present
on the second chalimus; the semirectangular
fourth part can not be traced to any definite

Lepeophtheirus dissimulaius- — Lewis

205

area in the frontal organ of the preceding stage.
Yolk material is still visible in the alimentary
tract and is concentrated in an enlarged area
just posterior to the ocular region.

Eight specimens of the third chalimus lasted
for approximately 33 hr at room temperature
(approximately 23 C) at which time they
moulted into the more elongate fourth chalimus.

The elongate condition of the fourth chalimus
indicates a cyclic condition in the growth of
the first four chalimus stages ( Fig. 5 ) . The av-
erage greatest length of the first chalimus (11
specimens), excluding setae, is 1.54 times the
greatest width, of the second chalimus (45
specimens) is 2.04 times the width, of the third
chalimus (25 specimens) is 1.54 times the
width, and of the fourth chalimus (74 speci-
mens) is 1.95 times the width. This cyclic
tendency, from stubby to elongate to stubby to
elongate, in the relationship between the great-
est length and greatest width in the first four
chalimus stages can not be explained by the
growth of any single structure or body part.
There appears to be a cyclic growth tendency
in several body parts. For example, the cephalo-
thorax length as a per cent of the total length is
79.1, 74.3, 79.5, and 73.6 for the first four chali-
mus stages respectively; the maximum width of
the genital-abdominal segment as a per cent
of the maximum length of the segment is 186.0,
153.6, 180.0, and 153.8 (genital segment only
for this last measurement) for the first four
stages respectively. Certainly, the inclusion of
the second pedigerous segment into the cepha-
lothorax affected the length of the cephalothorax
as a per cent of the total length measurements
in the second chalimus but the fact that the
third pedigerous segment was included in the
cephalothorax in the third chalimus did not ap-
pear to affect the trend in that stage.

The cephalothorax of the fourth chalimus
(Fig. 9 d) is ovoid in outline as in the preceding
chalimus stages. The greatest length of the ceph-
alothorax, including the frontal region but not
the frontal filament, is slightly more than 1.3
times the greatest width. The frontal region
and frontal plates are now distinct and the divi-
sion of the region from the cephalothorax is
complete. The division between the frontal re-
gion and the cephalothorax forms an irregular
groove curving anteriorly from its origin on the

anterior lateral margin. The anterior margin of
the frontal region is broadly curved, the an-
terior lateral surfaces forming laterally project-
ing lobes, the posterior lateral margin connect-
ing these lobes to the lateral margins of the
cephalothorax. The lateral cephalothoracic mar-
gins are generally convex, although wavy; the
posterior lateral corners are rounded. The lateral
posterior margins, outside the posterior sinuses*
are bilobed, consisting of the broadly rounded
posterior lateral extensions of the lateral regions
and the narrow, lobate lateral surface of the
median thoracic area. The posterior sinuses are
distinct and slender, generally V-shaped al-
though sometimes U-shaped. The median tho-
racic area extends posteriorly well past the pos-
terior extensions of the lateral regions and is
irregularly tapered to the slightly rounded pos-
terior margin. The lateral margins of the median
thoracic area are continuous with the inner
margins of the posterior sinuses. The major
cephalothoracic grooves, as in the preceding
stage, form an H. The anterior legs of the H,
however, curve laterally from the junction with
the cross groove anteriorly to their termination
lateral and just posterior to the eyes. The lateral
strengthening regions of the adult are not yet
visible.

The free fourth pedigerous segment is -short,
its greatest length approximately two-thirds its
width. The anterior end of the segment is over-
lapped by the posterior extension of the median
thoracic area of the cephalothorax. The lateral
margins taper both anteriorly and posteriorly
from the widest point, in the middle of the
segment, at the junction of the fourth thoracic
legs. The posterior margin is distinct, forming
a flatly convex groove.

The genital segment (Fig. 12^) is slightly
shorter than the fourth pedigerous segment; its
greatest length, excluding the slightly projecting
fifth legs, is approximately two-thirds its width.
The widest point is in the posterior medial re-
gion of the segment. The anterior lateral mar-
gins curve convexly, the posterior lateral mar-
gins concavely. The fifth legs project as dactyli-
form processes from the concave posterior lat-
eral margins and extend slightly past the poste-
rior end of the segment. The legs are tipped by
two small plumose setae. The posterior margia
of the segment is distinct although irregular.

206

PACIFIC SCIENCE, Vol. XVII, April 1963

0.5mm. <3

Fig. 10. Body of fifth and sixth chalimus developmental stages of L. dissimulatus. a, Dorsal view of
attached female fifth chalimus; b, dorsal view of free-moving male fifth chalimus; c, dorsal view of free-moving
female sixth chalimus; d , dorsal view of free-moving male sixth chalimus.

Lepeophtheirus dissimulatus — Lewis

207

The abdomen (Fig. 12 e) is slightly more
than one-half the length of the genital segment;
its greatest width is approximately twice the
greatest length. The lateral margins are flatly
convex or straight; the lateral posterior margins
are slightly concave and taper to the median,
bilobed anal region.

The bifurcate third part of the frontal organ
(Fig. 13*0 now extends to the posterior end of
the smaller semirectangular fourth part. Both
parts of the bifurcation are club-shaped and do
not extend laterally as in the third chalimus.
The alimentary tract is the same as that of the
preceding stage although there is not as much
yolk material.

Three specimens of the fourth chalimus lasted
for about 36 hr under laboratory conditions
(temperature approximately 23 C) before they
moulted into the fifth chalimus.

The fifth chalimus (Fig. 1 0a, b ) is found both
attached and free on the host although more fre-
quently attached. With the exception of a few
structures such as the genital segment and some
of the appendages, the somewhat flabby condi-
tion of the body and appendages characteristic of
earlier chalimus larvae is absent and the stage is
quite similar to the adult. The greatest length of
the cephalothorax, including the frontal region,
is approximately 1.3 times its greatest width, ex-
cluding the now present marginal flanges. The
frontal plates form most of the frontal region
and a narrow, membranous flange projects from
its broadly curved anterior margin. The lateral
frontal region surfaces project slightly, the pos-
terior lateral margins connecting the lobate pro-
jecting portions to the lateral margins of the
cephalothorax. The division between the cepha-
lothorax and the frontal region is a distinct
three-lobed groove, the median lobe of which
is approximately twice the length of the lateral
lobes. The lateral margins of the cephalothorax
are broadly curved, with distinct marginal
flanges extending from the junction of the
frontal region posteriorly around the posterior
extensions of the lateral regions to the begin-
ning of the posterior sinuses. The lateral pos-
terior margins, outside the posterior sinuses,
are bilobed and consist of the rounded posterior
extensions of the lateral regions and the sharply
rounded lateral surface of the median thoracic
area. The posterior sinuses are distinct and U-

shaped. The median thoracic area extends pos-
teriorly well past the posterior extensions of the
lateral regions; its posterior margin is broadly
rounded but possesses a small, median protru-
sion that forms the junction of the free fourth
pedigerous segment in the now discernible male
but which is absent in the female. The lateral
margins of the protruding median thoracic area
are continuous with the inner margins of the
posterior sinuses. The major cephalothoracic
grooves are similar to those of the preceding
chalimus but appear more definite and not quite
as irregular. Two heavily sclerotized regions,
the lateral strengthening regions, are visible as
posterior laterally curved, rodlike structures in
the cephalothorax, originating just anterior and
slightly medial to the junction of the longitudi-
nal legs of the cephalothoracic grooves and ex-
tending to the region of the lateral margins.

The free fourth pedigerous segment is short;
its greatest length is approximately one-half its
greatest width. The posterior portion of the
median thoracic area of the cephalothorax over-
laps the anterior end of the segment slightly in
the female but not in the male. The lateral mar-
gins are convex, the middle of the segment
being the broadest region and slightly overlap-
ping the proximal end of the fourth thoracic
legs. The posterior margin is distinct although
slightly irregular.

Because of the presence of the sixth thoracic
legs on the male of this stage, the sexes can
now be differentiated, as the female does not
possess this appendage pair. As will be discussed
later, the second antennae can not be used in
differentiating the sexes of any of the chalimus
stages of L. dissimulatus. The presence or ab-
sence of the sixth legs is the best method so far
found by which the sexes can be told apart in
both the fifth and sixth chalimus stages.

The genital segment of the female (Fig. 12 g)
is as wide as long; the lateral margins are flatly
or broadly convex. The anterior end possesses
two lateral, rounded protuberances that are dis-
tinct in a dorsal view but indistinct in a ventral
view. The posterior lateral margins are indented,
the lobate fifth legs arising from the ventral
surface and projecting posteriorly. The fifth leg
does not project past the posterior end of the
genital segment as in the preceding stage. Four
plumose setae, an additional two setae over the

208

PACIFIC SCIENCE, Vol. XVII, April 1963

last stage, are spaced along the outer margin of
the fifth leg. The posterior margin of the female
genital segment is broadly rounded in some
specimens and irregular in others.

The male genital segment (Fig. 12/) is sim-
ilar to that of the female although slightly
longer; the lateral margins are, however, not
indented in the dorsal view. Both the fifth and
sixth legs arise from the posterior ventral lateral
surface, adjacent to each other. Further, both
legs are lobate, the sixth extending slightly past
the division between the genital segment and
abdomen. The fifth leg possesses one plumose
setule on the outer base of the leg and two
plumose setae on the distal end; the sixth leg is
tipped by two plumose setules.

The abdomen (Fig. 12/, g ) is bell-shaped
and short; its greatest length is one-half that of
the genital segment in the males, slightly longer
in the females, and the greatest width is ap-
proximately 1 V4 times the length. The poste-
rior region of the single abdominal segment
is broader than the anterior, the lateral margins
are convex, and the lateral posterior margins
are concavely tapered to the bilobed anal region.

The frontal organ (Fig. 13/) is similar to that
of the preceding stage although the component
parts are less distinct. The attachment filament,
if present, arises from a circular indentation on
the anterior ventral surface just anterior to the
frontal organ. In unattached forms this circular
indentation forms a scar which is the only evi-
dence of the attachment filament. The yolk ma-
terial that was present in the fourth chalimus
now appears to be completely absent.

Only two specimens survived from the moult
into the fifth chalimus to the moult into the
sixth. Both of these specimens spent approxi-
mately 24 hr in the fifth chalimus stage under
laboratory conditions (temperature approxi-
mately 23 C) .

The sixth chalimus stage (Fig. 10c, d) is
the last larval stage. Specimens in this stage of
development were found both attached and free
although, in contrast to the fifth chalimus, more
frequently free. The greatest length of the ceph-
alothorax, including the frontal region, is ap-
proximately 1.2 times the greatest width, ex-
cluding the marginal flanges. The frontal region
is similar to that of the fifth chalimus. The lat-
eral margins of the cephalothorax are broadly

curved and are fringed with a distinct marginal
flange. The posterior lateral corners of the ceph-
alothorax are broadly rounded; the lateral pos-
terior margins, outside the posterior sinuses,
are bilobed, as in the preceding two stages, and
consist of the rounded posterior extensions of
the lateral regions and the sharply rounded lat-
eral posterior surface of the median thoracic
area. The posterior sinuses are distinct and U-
shaped. The median thoracic area extends pos-
teriorly slightly past the posterior extensions
of the lateral areas. The posterior margin of
the extension is broadly rounded in the female
but has a flat median protrusion in the male.
The lateral margins of the extended median
thoracic area are continuous with the inner mar-
gins of the posterior sinuses. The major cephalo-
thoracic grooves are similar to those of the pre-
ceding chalimus stage. Minor cephalothoracic
grooves are visible on some specimens and,
when present, extend posteriorly from the divi-
sion between the frontal region and the cepha-
lothorax. The lateral strengthening regions are
proportionately larger than in the fifth chalimus
but are similar in general shape and position.

The free fourth pedigerous segment is short,
its length approximately one-half the width. The
anterior end is covered by the posterior end of
the median thoracic area of the cephalothorax
in the female but not in the male. The middle
of the segment protrudes laterally, the fourth
thoracic legs being attached to the distal end of
the protrusion; the segment tapers from the pro-
trusion to the narrower anterior and posterior
ends. The division between the fourth pedi-
gerous segment and the genital segment is dis-
tinct although slightly irregular.

The genital segment of the female (Fig. I2i)
is slightly longer than wide, its greatest width
being approximately four-fifths of the length.
The lateral margins are broadly convex, the ante-
rior lateral surface forming an irregular, node-
like formation at the junction with the fourth
pedigerous segment. The fifth legs arise from the
lateral posterior ventral surface but do not ex-
tend to the posterior end of the segment; these
structures are lobate in outline and possess three
plumose setae along their outer margin and one
on the distal surface. The seta on the distal
surface possesses a minute swelling on the prox-
imal outer margin (Fig. 12/) which is absent in

Lepeophtheirus dissimulatus^-LEWlS

209

both the fifth chalimus and the adult but which
was present on all of the female sixth chalimus

specimens examined.

The genital segment of the male sixth chali-
mus (Fig. 12^) is longer than that of the fe-
male; the greatest width is approximately three-
fourths the greatest length. The lateral margins
are flatly convex, angled slightly from a swelling
on the anterior lateral corner, comparable to the
nodelike formation on the female, to the widest
point in the region of the fifth legs. The poste-
rior region of the segment curves medially from
the region of the fifth legs to the abdomen. Both
the fifth and sixth legs are lobate, the fifth ap-
proximately one-half the length of the sixth.
The fifth legs project from the ventral lateral
surface of the posterior one-half of the segment
and possess a single plumose seta on the ante-
rior end of the outer margin and three plumose
setae on the distal surface, an increase of one seta
from the fifth chalimus. The sixth legs project
from the ventral surface just inside and poste-
rior to the fifth legs and extend posteriorly to
the posterior end of the segment. A single plu-
mose seta, absent in the preceding chalimus, is
present on the distal one-half of the outer mar-
gin of the sixth legs and two plumose setae are
present on the distal surface.

The one-segmented abdomen of both sexes
(Fig. 12^?, i) is short, approximately one-third
the length of the male genital segment; the
length is exceeded somewhat by the width. The
anterior end is slightly narrower than the pos-
terior, the outer margin angled slightly from
the anterior end to the middle of the segment.
The posterior surface tapers from the middle of
the segment to the bilobed anal region at the
posterior extremity.

The frontal organ (Fig. 13g) appears to have
degenerated from the previous stage and now
consists of two oval bodies contiguous on the
median longitudinal axis of the body just pos-
terior to the division between the frontal region
and the cephalothorax. These two bodies are
connected to the frontal filament in attached
forms or to the circular depression in unattached
forms by a slender, rod-shaped structure. The
digestive tract is the same as that of the pre-
ceding stage.

Only one specimen survived from the moult
into the sixth chalimus to the moult into the

adult. The time spent by this single specimen
in the sixth chalimus was somewhere between
24 and 30 hr at a temperature of approximately
23 C.

After moulting from the sixth chalimus to
the adult, the last moult in the life history, all
of the appendages and processes remain un-
changed. As seen in Figures 2-7, pertaining to
the growth of L. dissimulatus, the maximum
size of the male is almost completely attained
by the sixth chalimus stage while that of the
female not until the animal is well into the
adult stage. Inasmuch as copulation takes place
shortly after the terminal moult, the male would
be more likely to be near its maximum size in
the last larval stage. During the reproductive
portion of the females life, eggs are stored in
the genital segment before they are extruded,
causing this segment to swell considerably. As
the female does not start to ovulate until some
time after fertilization, the significant increase
in size after the sixth chalimus is due, to a con-
siderable extent, to the enlarged genital segment
as is indicated in Figures 6 and 7. The presence
of eggs in the genital segment also pushes the
fifth legs laterally and, in some instances, ob-
scures them in a dorsal view of the animal.

The cephalothorax of the adult female and
male (Fig. 11a, b) is ovoid; the frontal region is
distinct, separated from the cephalothorax by a
trilobed groove. The lateral margins of the ceph-
alothorax are convex and possess a distinct mem-
branous flange. The posterior sinuses are distinct
and of a shallow V-shape. The median thoracic
region extends posteriorly slightly past the pos-
terior extension of the lateral regions, forming
the junction of the cephalothorax and the fourth
pedigerous segment. The major cephalothoracic
grooves are similar to those of the sixth chali-
mus; two minor grooves also extend posteriorly,
for a short distance, from the junctions of the
lateral and median lobes of the trilobed groove
separating the frontal region from the cephalo-
thorax. The rodlike strengthening regions are
more heavily sclerotized than those of the sixth
chalimus but are similar in shape and are in the
same position.

The fourth pedigerous segment of the fe-
male and male is short; the lateral regions are
drawn out at the fourth leg as in the preceding
stage. The terminal portions of the lateral ex-

210

PACIFIC SCIENCE, Vol XVII, April 1963

tensions are cup-shaped and form socket-like
attachments for the fourth legs. The division be-
tween the fourth pedigerous segment and the
genital segment is distinct and curved convexly.

The shape of the female genital segment (Fig.
12/) is variable, dependent upon the number
of eggs present in the segment. The segment is
flaccid and ovoid in females that have recently
extruded egg strings, and is almost circular in
females whose genital segment is packed with
eggs. Two small knoblike protrusions are pres-
ent on the posterior margin lateral to the junc-
tion of the abdomen; these knobs are charac-
teristic for the female of this species but are
not present until the adult stage. The general
shape of the fifth legs is similar to that of the

sixth chalimus although the proximal seta of
the three plumose setae present on the lateral
surface of the leg of the preceding stage now
arises from the surface of the genital segment,
just lateral to the base of the fifth leg. The
movement of this seta may be due to the swell-
ing of the genital segment.

The genital segment of the male (Fig. 12k)
is ovoid, the width slightly more than three-
fourths the length. The shape of this segment
and of both the fifth and sixth legs is very simi-
lar to that of the sixth chalimus male. The ar-
mature of the legs of both stages is identical.

The abdomen of the adult female and male is
slightly longer than that of the sixth chalimus
(0.12 mm average for 15 sixth chalimus speci-

Lepeophtheirus dissimulatus — Lewis

211

Q05mm QQSmm^ Q05mm^

e f g

0 3 mm

Fig. 12. Genital segment, abdomen, and caudal rami of developmental stages of L. dissimulatus . a, Cope-
podite, ventral view showing third and fourth pedigerous segments, genital-abdominal segment and caudal rami;
b, first chalimus, ventral view of genital-abdominal segment and caudal rami; c, second chalimus, ventral view
of genital-abdominal segment and caudal rami; d, third chalimus, ventral view of genital and abdominal seg-
ments, fifth leg and caudal rami; e, fourth chalimus, ventral view of genital and abdominal segments, fifth
legs and caudal rami; f, male fifth chalimus, ventral view of genital and abdominal segments, fifth and sixth
legs and caudal rami; g, female fifth chalimus, ventral view of genital and abdominal segments, fifth legs and
caudal rami; h, male sixth chalimus, ventral view showing genital and abdominal segments, beginning of
internal spermatophore, fifth and sixth legs and caudal ramus; i, female sixth chalimus, ventral view showing
genital and abdominal segments, fifth legs and caudal ramus; j, enlarged distal seta of female fifth leg show-
ing node on outer proximal surface; k, adult male, ventral view of genital and abdominal segments, fifth and
sixth legs and caudal ramus; l, adult female, ventral view of genital and abdominal segments, fifth legs and
caudal ramus.

212

PACIFIC SCIENCE, VoL XVII, April 1963

mens, 0.14 mm average for 30 adult specimens).
The general shape, however, is the same.

The frontal organ in the adult (Fig. 13^) is
greatly reduced, being visible only as a small
opaque area on the median longitudinal axis of
the body just posterior to the groove dividing
the frontal region from the cephalothorax. The
attachment filament is completely absent, the
only indication of its previous presence being
the circular scar mentioned with reference to
unattached specimens of the sixth chalimus.

Antennule

Throughout development, from the nauplius
to the adult, the antennule is two-segmented.
The changes that occur in this appendage are
changes in the general shape and relative size
of the segments in addition to changes in the
armature.

The antennule of the first nauplius (Fig. \Aa)
is uniramous and is attached to the anterior
ventral lateral surface of the body. The lateral

Fig. 13. Frontal region of developmental stages of L. dissimulatus. Ventral view of anterior surface: a,
Copepodite; b, first chalimus; c, second chalimus; d, third chalimus; e, fourth chalimus; f, fifth chalimus; g, sixth
chalimus; h, adult. A-l, Antennule; A-2, base of second antenna; FO, frontal organ; FP, frontal plate; R,
Rostrum.

Lepeopbtheirus dissimulatus — Lewis

213

Fig. 14. Antennule of developmental stages of L. dissimulatus (ventral view), a, First nauplius; b, second
nauplius; c, copepodite; d, first chalimus; e, second chalimus; f, third chalimus; g, fourth chalimus; h, fifth
chalimus; i, sixth chalimus; j, adult.

margins of both segments are slightly irregular,
more so in some specimens than in others. The
first segment is slightly shorter than the second
and is tapered rapidly in the proximal one-third
to the narrow proximal end, the surface of at-
tachment with the body. The second segment
is tapered from the proximal to the slightly nar-
rower, rounded distal end. The distal end of the
second segment is encircled by a membrane pro-

jecting slightly past the segment. A broadly
angled, heavily sclerotized process projects from
the inner portion of the distal end of the sec-
ond segment and two lightly plumose setae
project from the inner portion of the distal end,
adjacent to the process; these setae are slightly
longer than the greatest length of the appendage.

The antennule of the second nauplius (Fig.
lAb) is attached to the anterior ventral lateral

214

surface of the body. The first segment is now
slightly longer than the second and its proximal
end is almost pedunculate. The lateral margins
of the segment are irregular although basically
parallel; the distal end is slightly convex. The
second segment possesses two long, lightly plu-
mose setae, as in the first nauplius, but also a
short, lightly plumose seta from the inner distal
margin in addition to one short, spinelike proc-
ess on the outer distal surface. The membrane
present on the preceding stage is completely ab-
sent. The copepodite antennule is visible in
late second naupliar specimens, incompletely
filling the appendage; the character of the co-
pepodite appendage is, however, not definable.

The antennule of the copepodite is attached
to the ventral lateral surface just lateral to the
rostrum. The first segment is almost 144 times
the length of the second; its greatest width is
approximately one-half the greatest length. The
proximal anterior lateral margin (the margin
that faces anteriorly in situ ) of the first segment
is strongly convex; the remaining lateral mar-
gin and the posterior lateral margin are some-
what irregular. The first segment possesses two
naked setae from the middle of the anterior
margin and one naked seta from the anterior
distal "margin. The division between the first and
second segments is distinct although slightly ir-
regular. The greatest length of the second seg-
ment is slightly less than the greatest width,
the lateral margins are almost parallel, and the
distal margin is irregular. The second segment
gives rise to 11 naked setae from its distal sur-
face.

The division between the two segments of
the first chalimus antennule (Fig. 14^) is in-
distinct; the appendage is attached to the an-
terior ventral lateral surface just anterior to the
indistinct, incomplete ventral groove dividing
the frontal region from the cephalothorax. The
first segment is slightly more than IV2 times
the length of the second, and the proximal re-
gion is broader than the distal; the rounded
proximal end is almost included in the posterior
margin, and the anterior and posterior margins
are irregularly convex. A single naked seta is
present on the proximal one-half of the ante-
rior surface. The second segment is approxi-
mately twice as long as wide, the posterior and
interior margins are generally flatly convex, and

PACIFIC SCIENCE, VoL XVIIv April 1963

the distal end is rounded. A single naked seta is
present on the middle of the posterior margin,
as in the adult; eight naked setae are also pres-
ent on the distal end.

The first chalimus antennule shows some de-
generation from the copepodite appendage. This
condition is evident in most of the appendages
in the change that occurs from the copepodite
to the first chalimus.

The division between the two segments of
the second chalimus antennule (Fig. I4g?) is dis-
tinct. The first segment is approximately twice
the length of the second, comparatively longer
than in the preceding stage. The proximal end
of the first segment is broad; the segment is
tapered irregularly to the narrower distal end.
The anterior lateral margin is smoothly convex,
longer than the irregularly concave posterior
margin; the distal margin is flat. About 10 naked
setules are present along the anterior lateral
margin of the first segment. The second segment
is twice as wide as long, the lateral margins are
slightly irregular, and the distal margin irregu-
larly rounded. A single naked seta is present on
the middle of the posterior margin of the sec-
ond segment in addition to nine naked setae on
the distal surface.

The third chalimus antennule (Fig. 14/ ) is
attached to a swelling on the posterior lateral
portion of the tear-shaped frontal region. The
greatest length of the first segment is about
twice that of the second; its greatest width is
two-thirds the length. The proximal end of the
segment is angled; the anterior margin is thus
more than twice the length of the posterior; the
distal end is irregular and has two distinct swell-
ings. Six lightly plumose setae arise from the
anterior margin of the first segment and three
from the distal margin. The second segment is
slightly constricted at the junction of the first
segment and curved convexly distally, causing
the segment to appear club-shaped. The greatest
width of the second segment is slightly more
than one-half the length. As in the preceding
two chalimus stages, a single naked seta is pres-
ent on the middle of the posterior margin and
nine naked setae arise from the distal surface.

The fourth chalimus antennule (Fig. 1 4g)
is also attached to a swelling on the posterior
lateral corner of the frontal plates which are
now oriented more in a horizontal manner than

Lepeophtbeirus dissimulatus — Lewis

in the third chalimus. The general shape of the
two antennular segments is similar to that of
the preceding stage although the two distinct
swellings present on the distal end of the first
segment are now absent. The anterior surface
of the first segment bears 10 setae that are dis-
tinctly plumose; the anterior distal surface bears
two plumose setae. The second segment bears
a single naked seta from the middle of the pos-
terior margin, four from the posterior distal
surface, and six from the anterior distal.

The antennule of the fifth chalimus (Fig.
14/?) is attached in the same region as that of the
fourth although the swelling at the junction of
the appendage and the frontal region is not as
distinct. The overlap of the antennule base onto
the anterior ventral surface of the cephalothorax
(see Fig. 13 e-h) in the fifth and sixth chalimus
and the adult may be due, at least in part, to
the incorporation of the swollen portion of the
frontal region into the frontal region of later
stages and the resultant displacement of the pos-
terior portion of the antennule base onto the
cephalothorax. The attachment of the antennule
thus appears to be basically on the frontal re-
gion and secondarily on the cephalothorax. The
first segment of the fifth chalimus antennule is,
with regard to the second segment, slightly
longer than in the preceding stage, being more
than twice the length of the second segment.
The greatest width of the segment is slightly
more than IV2 times the length; the anterior
margin is flatly convex medially and broadly
rounded distally and proximally. The posterior
margin is irregular, with a slight angular pro-
trusion from the distal region. The anterior
surface of the first segment bears 16 plumose
setae, the distal surface three. The length of the
second segment is approximately 21/2 times the
width, the lateral margins are flatly convex, and
the distal margin is rounded. The armature of
this segment is the same as in the preceding
stage.

The antennule of the sixth chalimus (Fig.
1 4i) is attached to the ventral lateral surface of
the frontal region and is adjacent to or slightly
overlaps the anterior ventral surface of the
cephalothorax. The second segment is slightly
longer than that of the preceding stage. The
first segment is broad proximally, its greatest
width being approximately five-eighths of the

215

greatest length, and tapers to the narrow distal
end. The anterior margin is irregular except for
the broadly convex proximal region, the poste-
rior margin is shorter than the anterior, and the
posterior surface bears a small distal protrusion.
Nineteen plumose setae arise from the anterior
surface of the first segment and the rounded dis-
tal surface bears five more. The second segment
is club-shaped; the greatest width is slightly
more than one-third the length. The lateral mar-
gins are generally flatly convex, the proximal
end slightly narrower than the rounded distal
end. The armature of the second segment con-
sists of one naked seta from the middle of the
posterior margin and 11 naked setae from the
distal surface.

The antennule of the adult (Fig. 14;) is sim-
ilar to that of the sixth chalimus except for the
armature. The first segment has a small bifurcate
protrusion on the posterior distal corner, the
spines of which are crenate in the female and
dentate in the male. The first segment bears 20
plumose setae on its anterior and distal surfaces;
the second segment bears one naked seta on the
middle of the posterior surface and 12 naked
setae on the distal surface.

Second Antenna

In contrast to the antennule, the second an-
tenna changes drastically during the life cycle
of L. dissimulatus, from a biramous swimming
appendage in the nauplius to a uniramous pre-
hensile appendage in the adult.

The second antenna of the first nauplius (Fig.
15^) is biramous and attached to the ventral
lateral surface just posterior to the base of the
antennule. The protopodite is one-segmented,
its greatest width approximately three-fourths
of the greatest length. The lateral margins, ex-
cept at the proximal end, are almost parallel;
the distal margin is broadly curved. The five-
segmented exopodite arises from the dorsal lat-
eral surface of the protopodite, not from the
distal end; the division between the exopodite
and the protopodite is indistinct and incom-
plete. The first exopodite segment is slightly
longer than the second, its proximal region tap-
ered evenly to the narrow distal end. The second
segment is tapered evenly from the proximal
to the narrow distal end; a single lightly plu-
mose seta is present on the inner distal lateral

216

PACIFIC SCIENCE, Vol. XVII, April 1963

FIG. 15. Second antenna of developmental stages of dissimulatus. a, First nauplius, ventral view; h, second
nauplius, ventral view; c, copepodite, posterior surface; d, setule of first segment of copepodite antenna (en-
larged over original enlargement); e, first chalimus, posterior surface; f, second chalimus, posterior surface;
g, third chalimus, posterior surface; h, fourth chalimus, posterior surface; i, fifth chalimus, posterior surface;
j, male sixth chalimus, posterior surface; k, female sixth chalimus, posterior surface; l, male adult, posterior
surface; m, female adult, posterior surface.

surface. The third segment is slightly more than
one-third the length of the second; it is slightly
wider distally than proximally and possesses a
spinelike projection on the anterior proximal
surface that extends distally slightly past the
end of the segment. Further, the third segment

gives rise to a single lightly plumose seta from
the inner lateral surface. The fourth segment is
similar to the third although slightly smaller.
The fifth segment is minute, its distal surface
slightly irregular; the anterior lateral margin is
longer than the posterior; a single lightly plu-

Lepeophthewus dissimulatus— Lewis

217

mose terminal seta is borne by this segment.
The endopodite is two-segmented and attached
to the ventral lateral surface of the protopodite.
The division between the endopodite and pro-
topodite is also indistinct and incomplete. The
first segment is about 3 times the length of the
second; its proximal end is narrow and the seg-
ment is curved slightly to the broader distal end.
The lateral and distal margins of the second
segment are continuous and form a broadly
rounded margin. A strong spinelike projection
is present on the middle of the distal end of the
second segment and a slender spinelike projec-
tion is present on the posterior distal end. The
second segment also bears two lightly plumose
setae from its anterior distal surface.

The second antenna of the second nauplius
(Fig. 15 b) is similar to that of the first nauplius
in general outline and make-up. The spinelike
projections on the third and fourth segments of
the first nauplius exopodite are absent in the
second although the remaining armature is the
same. The make-up and armature of the two
endopodite segments is the same as that of the
preceding stage with one exception: the two
spinelike processes on the distal surface of the
second segment appear to be more strongly de-
veloped.

The second antenna of the copepodite (Fig.
15 c, d ) is uniramous and three-segmented. The
appendage is attached to the anterior ventral
surface just medial to the posterior edge of the
antennule base. Wilson (1905:542) indicates
that the caligid second antenna is still biramous
in the copepodite stage and Heegaard (1947:
56, fig. 14) describes and figures the reduced
exopodite on the second antenna of Caligus
curtus. Unless the small setule (Fig. 15 d) or
one of the projections on the proximal segment
of the second antenna of L. dissimulatus is the
remnant of the now absent ramus of the bira-
mous second naupliar appendage, no evidence of
a biramous condition is present. The first seg-
ment of the copepodite appendage is short and
irregular, folded in several places, and possess-
ing two backward-projecting, broadly-angled,
spinelike processes, the proximal of which is
poorly developed and the distal well developed.
The division between the first and second seg-
ments is distinct although irregular. The second
segment is broad proximally but tapered to a

narrow distal end; the lateral margins are regu-
lar, the distal surface concave. The second seg-
ment possesses a small lappet-like process pro-
jecting from the middle of the posterior medial
surface. The third segment is short, less than
one-eighth the length of the second; the proxi-
mal end is slender, the distal is broad, and both
lateral margins are convex. The distal end of the
third segment is flat except for a slight depres-
sion in the middle. A large spinelike terminal
process curves anteriorly from the posterior one-
half of the surface; a slender, seta-like accessory
process projects from the anterior surface.

The first chalimus second antenna (Fig. 15e)
appears to have degenerated somewhat; its shape
is rather indistinct and the entire appendage
appears flabby. The second antenna of this stage
is attached just posterior and medial to the an-
tennule base. The first segment is broad proxi-
mally, somewhat narrower distally, and the gen-
eral outline is irregular. The distal end of the
first segment forms an articulation surface for
the second segment. The second segment is well
developed; its greatest length is slightly more
than the combined lengths of the first and third
segments. This segment is broader proximally
than distally, and the lateral margins are irregu-
lar, the outer longer than the inner; the distal
surface faces inwards and its margin is generally
flat. The third segment is short and truncate,
tipped by a sharply curved, heavily sclerotized,
spinelike process; the accessory process is pres-
ent as a small seta-like structure on the distal
one-half of the inner surface of the segment.

The second chalimus second antenna (Fig.
15/) is attached just posterior and medial to the
base of the antennule. The first segment is broad
and irregular in outline; its distal surface forms
an articulation surface for the second segment.
An indistinct, posteriorly directed, distally
pointed process is present on the inner surface
of the segment. The second segment is well de-
veloped, twice the length of the combined first
and third segments. This segment is broad proxi-
mally and tapers to a narrow distal end. The
outer lateral margins are smoothly convex, the
inner irregularly concave; the distal surface is
angled towards the inner proximal end, and the
distal margin is slightly irregular. The outer
margin of the third segment is convex and con-
tinuous with the outer margin of the short, in-

218

wardly curved terminal process. The inner mar-
gin of the third segment is broken by the lobe-
shaped proximal region. The division between
the terminal process and the segment is indis-
tinct. In addition to the terminal process, a
small seta-like accessory process is present on
the lobate inner proximal region.

The second antenna of the third chalimus
(Fig. 15g) appears to have degenerated even
more when compared with the same appendage
on the copepodite and the previous two chal-
imus stages. This condition is a continuation of
the trend initiated in the first chalimus, a grad-
ual loss of rigidity and distinct form before the
development of the adult appendage. To ex-
emplify this loss of form, the segments of the
third chalimus appendage are distinct but plastic
in nature, the integument is flabby, and the
ridges and grooves present on the second an-
tenna of one side of a specimen are often absent
on the other; further, the sclerotization of the
terminal process appears reduced. The greatest
length and width of the first and second seg-
ments of the second antenna of the third chal-
imus are approximately equal although the shape
is different. The distal end of the first segment
is generally concave, of the second segment al-
most flat. The third segment is slightly shorter
than either of the preceding two and is tapered
from the broad proximal to pointed distal end.
The inner margin of the segment possesses a
distinct protrusion on the distal one-half that
is tipped by a short, pointed process. The more
heavily sclerotized terminal process of the suc-
ceeding stage is visible inside the distal portion
of the third segment.

The second antenna of the fourth chalimus
(Fig. 15 h) is similar in general make-up to that
of the preceding stage except that the first seg-
ment is not as long. Further, it has a flaplike
structure projecting distally from the proximal
end; one specimen also possessed a triangular,
distally directed flap that appeared to correspond
to the posteriorly directed, pointed process on
the first segment of the adult female. The length
of the second segment is greater than that of
the first, the division between the second and
third segments is indistinct, and the lateral mar-
gins of both segments are irregular. The distal
inner margin of the third segment bears a small,
lobate process that will become the accessory

PACIFIC SCIENCE, Vol. XVII, April 1963

process of the adult and is the protrusion that
was present on the same region of the previous
stage. The distal end is bluntly pointed, more
heavily sclerotized than the preceding stage but
not as heavily as the terminal process of the fifth
chalimus, visible inside the fourth.

The second antenna of the female and male
fifth chalimus (Fig. 15 i) are similar. The gen-
eral appearance of the adult female appendage
is now beginning to manifest itself in the fe-
males of this stage. The first segment is irregu-
lar, the distal surface concave and forming the
articulation surface for the second segment. A
spinelike, posteriorly directed process is present
on the posterior surface of the segment. The
second segment is short and strongly developed;
the inner margin is convex, approximately 3
times the length of the concave outer margin
due to the angled distal and proximal surfaces.
The distal surface of the second segment is ir-
regular; the inner portion is concave, the outer
convex although both portions are heavily scle-
rotized and form the articulation surface for
the third segment. The third segment and ter-
minal process in both sexes is approximately
IVi times the length of the second segment;
the proximal surface is lobate and articulates in
the concavity of the distal end of the second
segment. The terminal process of the third seg-
ment is curved inwards strongly at the distal
end; the division between the process and the
segment is indistinct. Further, two seta-like ac-
cessory processes are now present, one on the
posterior proximal region, the second on the
inner distal region.

The second antenna of the sixth chalimus
(Fig. 15;, k) is similar in both sexes. The first
segment is slightly longer in the male but, for
both male and female, is short and the entire
distal surface is concave, forming an articulation
surface for the second segment. The first seg-
ment is irregular in outline, the outer surface
appearing platelike and tapered gradually to the
distal surface, the inner surface somewhat ir-
regular, with a posteriorly directed, spinelike
projection that is more distinct than in the pre-
vious stage. The second segment is strongly de-
veloped, its greatest width slightly more than
four-fifths of the greatest length; the outer mar-
gin is flatly convex, longer than the irregularly
concave inner margin. The width of the distal

Lep e op hth eiruS dissimulatus — -LEWIS

219

end of the second segment is slightly less than
three-fourths of the width of the proximal end;
the distal surface is concave with its outer por-
tion heavily sclerotized and serving as the articu-
lation surface for the outer proximal end of the
third segment. The third segment and terminal
process are similar to those of the preceding
stage although the process is slightly longer and
the distal portion more sharply curved. The
adult female and male appendage is visible
within the second antenna of late sixth chalimus
specimens. Both parts of the bifurcate terminal
process of the adult male can be seen (Fig. 15;)
as well as the single female terminal process
(Fig. 15k).

It was earlier stated that the second antennae
can not be used to differentiate the two sexes
in the late chalimus of L. dissimulatus. As was
just described, the female and the male second
antenna are similar in the sixth chalimus, the
last larval stage. Not until the moult from the
sixth chalimus to the adult is there any signifi-
cant difference between the appendages of the
two sexes in L. dissimulatus. For this reason, the
presence or absence of the sixth legs, found only
in the male, was used as the major differentiat-
ing characteristic. Additionally, in the sixth chal-
imus, the presence of internal spermatophores
in the male was used to verify the differentiation
of the two sexes.

The second antenna of the adult (Fig. 15/, m)
is attached to the ventral surface of the cephalo-
thorax just anterior and medial to the postan-
tennal process and posterior and lateral to the
base of the antennule. The first segment of the
female is short and forms a broad articulation
surface for the second segment. The segment
possesses a posteriorly directed, spinelike pro-
jection from the posterior proximal surface. The
second segment is strongly developed, its great-
est width being equal to its greatest length. The
third segment is slender and heavily sclerotized,
bearing a sharply curved, spinelike terminal
process and two naked, seta-like accessory proc-
esses, one from the inner surface of the proxi-
mal region and the second from the distal lateral
surface. The division between the third seg-
ment and terminal process is indistinct and in-
complete.

The first segment of the adult male second
antenna (Fig. 15/) is longer than that of the

female; the segment is broader proximally than
distally and is attached to the cephalothorax
along the entire length of the proximal surface.
The distal end of the first segment is small and
two-pronged, forming an articulation surface for
the second segment. Additionally, the major
portion of the outer lateral surface forms an
adhesion surface of heavily sclerotized, over-
lapping, platelike structures. The second seg-
ment is strongly developed, its greatest width
slightly more than one -ha If of the greatest
length. The outer margin of the segment is
strongly convex proximally; the inner margin
is irregular due to the presence of two sets of
adhesion surfaces similar to those of the first
segment. The inner distal margin of the second
segment also possesses a finger-like protrusion
that appears to be segmented but presumably
forms an adhesion surface, and the segmented
appearance is due to the overlapping plates. The
protrusion is, in most specimens, curved around
the second segment and appears as a regular
adhesion surface but it is not attached to the
segment except at its proximal end. The third
segment is short and bears a bifurcate terminal
process in addition to a single, naked, seta-like
accessory process from the inner distal end.

Mandible

Comparing the over-all development of the
three naupliar appendages, it is evident that the
mandible has the most drastic change. Almost
all of the change occurs in the moult from the
second nauplius to the copepodite when the
appendage changes from a biramous appendage
to a uniramous, four-parted, rodlike process. The
four-parted condition could not be traced to
either the exopodite or the endopodite of the
naupliar appendage. All four parts appear to
be almost completely fused; there are no muscles
penetrating the appendage and there is no trace
of the second ramus of the biramous naupliar
appendage. These three conditions make the
analysis of the derivation of the mandible of
the copepodite of L. dissimulatus extremely dif-
ficult, and even with the analysis made by Hee-
gaard (1947:59, 196-202), no hypothesis can
be offered until further work has been done.

The mandible of the first nauplius (Fig. 1 6a)
is biramous and attached to the ventral lateral
surface just posterior to the second antenna.

220

PACIFIC SCIENCE, Vol. XVII, April 1963

*0.03 m'rn.

Q03mm 003mm, Q03mm

t U v

QQ5mm

0.03 mm,

GQ5mm

Fig. 16. Mandible, postantennal and postoral processes of developmental stages, in addition to balancers of
naupliar stages of L. dissimulatus. a-j, Mandible: a, First nauplius, ventral view; b, second nauplius, ventral
view; c, copepodite; d, first chalimus; e, second chalimus; f, third chalimus; g, fourth chalimus; h, fifth chali-
mus; i, j, sixth chalimus and adult, k, l, Nauplius balancers: k, First nauplius, dorsal view; l, second nauplius,
lateral view (also posterior end of body of nauplius). m-q, Postantennal process: m, Third chalimus; n, fourth
chalimus; o, fifth chalimus; p, sixth chalimus; q, adult, r—y, Postoral process: r, Copepodite; s, first chalimus;
t, second chalimus; u, third chalimus; v, fourth chalimus; w, fifth chalimus; x, sixth chalimus; y, adult.

Lepeophthekus dissimulates-— Lewis

221

The protopodite is one-segmented, its greatest
width about three-fourths of the greatest length;
the lateral margins are slightly irregular al-
though generally convex, and the distal margin
is broadly curved. The exopodite is four-seg-
mented and attached to the dorsal lateral surface
of the protopodite although the division be-
tween the first segment and the protopodite is
incomplete. The first segment of the exopodite
is more than 2Vi times the combined lengths
of the remaining three segments, its lateral mar-
gins are broadly convex, and the distal surface
has a concavity in which the proximal end of
the second segment is attached. A single lightly
plumose seta is present on the distal portion of
the inner lateral surface of the first segment.
The second segment is short, its greatest width
about equal to the greatest length; the distal
surface is slightly concave and receives the
proximal end of the third segment. A single
seta is present on the distal portion of the inner
lateral surface of the second segment. The third
segment is slightly longer than the second and
flared from the narrow proximal to broader dis-
tal end; the distal surface also has an irregular
concavity for the proximal end of the fourth
segment. A single seta is present on the distal
portion of the inner lateral surface of the third
segment. The fourth or terminal segment is
minute and rodlike, with a concave distal sur-
face that receives the base of the slightly plu-
mose terminal seta. The endopodite is two-
segmented and attached to the ventral lateral
surface of the protopodite although the division
between the first segment and the protopodite
is incomplete. The first segment is approxi-
mately 1 Vs times the length of the second, the
lateral margins are slightly irregular, and the
distal end is flared slightly and bearing a small,
pointed protrusion on the anterior distal corner.
The second segment has slightly irregular lateral
margins, the distal end is slightly broader than
the proximal, and the distal surface is flat and
bears two lightly plumose setae.

The mandible of the second nauplius (Fig.
1 6b) is similar to that of the first nauplius. The
first segment of the exopodite is, however,
longer than that of the preceding stage, being
more than 3 Vi times the combined lengths of
the remaining three segments. The small pointed
protrusion on the anterior distal corner of the

first endopodite segment of the first nauplius is
larger in the second nauplius and projects an-
teriorly although there is some variation in size
in different specimens.

The copepodite mandible (Fig. 1 6c) is uni-
ramous and four-parted as in the adult. The ap-
pendage is attached to the ventral surface of the
cephalothorax just lateral to the division be-
tween the well-developed membranous hyper-
stome and hypostome and projects through the
division into the mouth cone. All four parts are
heavily sclerotized and indistinctly divided, the
division often appearing only as a light area.
The first part is approximately twice the length
of the second, tapered slightly from the proximal
to the narrower distal end. The second part is
strongly tapered from the first to slender third
part. The third part is elongate, the length
slightly less than the combined lengths of the
first two parts; the lateral margins are almost
parallel. The fourth part is short, the length less
than one-half that of the third part. The fourth
part is also angled medially; there is no curva-
ture as the fourth part of later stages has. The
fourth part is tapered to a sharp point distally;
the inner margin is lightly serrated but does not
have the denticulations visible in the adult.

The condition exhibited by the copepodite
mandible is changed but little in the chaiimus
stages. The only part that changes to any extent
is the fourth part, the other three changing only
slightly in their relative lengths. A slight curva-
ture of the fourth part is visible in the first
chaiimus (Fig. \6d)y this curvature becoming
greater throughout the remaining chaiimus
stages (Fig. iGe-i) so that the fourth part is
evenly curved in the sixth chaiimus and adult
(Fig. 1 6i, ;). Further modification of the fourth
part is from the sharply pointed condition of
the copepodite to the bluntly rounded tip of
the adult and the gradual appearance of the
denticulations present on the inner surface of
the adult.

Mouth Cone

The mouth cone is first exhibited by the co-
pepodite, although a slight swelling is visible
on the second nauplius that may be the begin-
ning of this structure. The cone itself is formed
of two membranous structures, a hyperstome
and hypostome, that are usually closely applied

222

to each other or overlap slightly. A slight open-
ing is visible in the proximal portion through
which the mandibles project into the cone. Rela-
tively little change occurs in the mouth cone
from the copepodite to the adult although be-
cause of the size and nature of the cone, a com-
plete analysis was not made. In the adult, how-
ever, both the hyperstome and hypostome pos-
sess several heavily sclerotized strengthening re-
gions on their lateral surfaces. The distal end of
the cone is membranous, very flexible, and finely
setuliferous.

Postantennal Process

The postantennal process is discussed after the
mandible because of the possibility that it is a
true appendage (Heegaard, 1947:59, 203-206).
The terminology used, with regard to the au-
thor’s present beliefs, is the same as that used
in Lewis (in press). The fact that, in L. dis-
simulatus, this process does not appear until the
third chalimus, long after all of the other oral
appendages are present, is an indication that it
is not a true appendage, although Heegaard
(1947:205) presents a hypothesis for its late
appearance. The arguments both for and against
the term "appendage” being applied are, how-
ever, strong enough so that this author prefers
not to commit himself until further studies can
be made.

The postantennal process of the third chal-
imus (Fig. 1 6m) is a blunt-tipped, spinelike
process projecting posteriorly from an indistinct
plate on the ventral surface of the cephalothorax
posterior and lateral to the base of the second
antenna. The proximal end of the process bears
two minute nodules, each with one setule.

The postantennal process of the fourth chal-
imus (Fig. iGn) is more strongly developed
and sharply pointed than that of the preceding
stage. The indistinct plate still bears the process,
from the base of which arise two minute nodules
as in the preceding stage, each with a single
setule.

The postantennal process of the female and
male fifth chalimus (Fig. l6o ) is more strongly
developed than in the preceding stage; the proc-
ess has become distinctly curved medially and,
in addition to the two single setule-bearing nod-
ules on the proximal end, now has a third nod-
ule just medial to the distal region of the proc-

PACIFIC SCIENCE, Vol. XVII, April 1963

ess. The plate from which the process of the
preceding two stages arose is now almost com-
pletely invisible.

The postantennal process of the female and
male sixth chalimus (Fig. 1 6p) is curved me-
dially as in the preceding stage. The proximal
region is, however, more developed, the inner
margin irregular, not evenly curved as in pre-
vious stages. Two nodules are present on the
proximal end but, in contrast to the last three
stages, each now bears two fine setules. Only
one setule was visible on the third nodule, al-
though the extremely small diameter and short
length of the setules made observation difficult
and the number may be incorrect. The plate
from which the process arises is more distinct
than in the preceding stage.

The adult female and male postantennal proc-
ess (Fig. 1 6q) is similar to that of the sixth
chalimus, although the distal end is more sharply
curved. The proximal nodule of the two nodules
on the base of the process now gives rise to
three setules, the distal nodule to four. The
third nodule of the chalimus stages did not ap-
pear to be present.

Postoral Process

The postoral process is of the same general
make-up as the postantennal process, although
it is first present on the copepodite instead of
the third chalimus. The process has been called
the first maxilla and the second maxilla. Its
structure, in addition to the location of the
origin of the subesophageal ganglionic nerve
that innervates it (adjacent to nerves that go
to the lateral musculature and the postantennal
process) and the inability to determine its an-
lage, has caused the author to use the term "post-
oral process.”

The postoral process of the copepodite (Fig.
I6r) consists of a posteriorly projecting, well-
developed, distally pointed process and two
setule-like projections just medial to the proxi-
mal end of the process. Both the process and
the projections arise just lateral and slightly pos-
terior to the base of the mouth cone. The inner
margin of the process is convex, the outer con-
cave with a slight proximal swelling. A minute,
heavily sclerotized, lobelike structure is present
just lateral to the process and appears to be con-

hepeophthewus dissimulatus— Lewis

223

nected to the same poorly defined plate from
which it arises.

The postoral process of the first chalimus
(Fig. 1 6s) is distinctly different from that of
the copepodite. The process consists of a two-
parted, spinelike process and a setule-bearing
nodule, both attached to a well-defined plate just
lateral and posterior to the base of the mouth
cone and contiguous with the plate present at
the base of the cone. The two-parted, spinelike
process projects posteriorly; both parts are
rounded distally, the second part being attached
to the distal end of the first. The greatest length
of the second part is approximately two-thirds
that of the first. The setule-bearing nodule is
just anterior to the base of the spinelike process,
at the anterior end of the plate. The nodule is
small and gives rise to two slender setules from
its distal surface.

A general transition is apparent from the first
chalimus to the adult. The two-parted condition
of the postoral process is still evident in the
second chalimus (Fig. 16/), but in the third
(Fig. 1 6u) the two parts become fused, the
distal portion becomes more sharply pointed,
and the base spreads out to its greatest width, in
the adult (Fig. 1 6y), the width in the adult
being just slightly less than the length. The plate
from which the process and the setule-bearing
nodule arise becomes indistinct in the second
chalimus, third chalimus, and fourth chalimus,
then becomes more distinct in the fifth chalimus
but appears as two parts. These two parts of the
plate are evident in the sixth chalimus and in
the adult. The anterior plate has a posteriorly
directed lobe (within, not projecting from, the
cephalothorax ) just medial to which the setule-
bearing node projects. The posterior plate forms
the base of the spinelike process and is continu-
ous with it. The setule-bearing node enlarges
somewhat throughout development and adds
another setule to its armature, having two slen-
der setules in the second chalimus, three in the
third and fourth chalimus, two (?) in the fifth,
and three in the sixth chalimus and in the adult.
All the setules are borne on the distal surface
of the nodule.

Maxilla

The term "maxilla” is applied to the pair of
oral appendages immediately behind the mouth

cone and postoral processes. This appendage
pair has gone under a variety of names because
of the terminology applied to the postantennal
and postoral processes. Inasmuch as the term
"process” is here applied to the postantennal
and postoral structures, the first pair of recog-
nizable appendages behind the mouth are des-
ignated as the maxillae.

The maxilla is first present in the copepodite
(Fig. 17*) and is found attached to the ventral
surface of the cephalothorax just posterior and
lateral to the postoral process. The first segment
of the two-segmented structure is strongly de-
veloped; the proximal surface possesses a medial
projection that articulates with the cuticle of
the body at the point of attachment of the ap-
pendage. The distal margin of the first segment
is flat; the surface is depressed and receives the
proximal surface of the second segment. The
second segment is elongate, being slightly longer
than the first segment and much thinner. The
lateral margins of the second segment are irreg-
ular, the outer proximal margin curving inward
to the irregular proximal surface, the inner mar-
gin with several flatly convex swellings. The
distal region of the second segment is slightly
swollen, the distal margin broadly rounded.
Two processes are present on the distal region
and extend out from the segment. The outer
process is spinelike and is less, than one-fourth
the length of the segment; this process possesses
a series of minute, tine-like projections along
the inner margin giving it a brushlike appear-
ance. The inner process arises from the posterior
lateral surface of the distal region, not the dis-
tal surface as does the outer process; it is lobate
and flimsy in nature, its length approximately
equal to that of the outer process.

The segments of the maxilla of the first chal-
imus (Fig. 17 h) are similar in general make-up
to those of the copepodite. The proximal articu-
lation surface of the copepodite appendage is,
however, absent in the first chalimus and, fur-
ther, the first segment is somewhat thicker. The
distal surface of the first segment is concave
and forms the articulation surface for the ball-
shaped proximal end of the second segment.
The inner of the two terminal processes on the
second segment is slightly longer than the outer
and is tapered to a sharp point. The outer of
the two processes is simple and lobate, lacking,

224

PACIFIC SCIENCE, Vol. XVII, April 1963

Fig. 17. Maxillae and maxiliipeds of developmental stages of L. dissimulatus. a—h, Maxillae: a, Copepodite;
b, first chalimus; c, second chalimus; d, third chalimus; e, fourth chalimus; f, fifth chalimus; g, sixth chalimus;
h, adult, i-p, Maxiliipeds: i, Copepodite; j, first chalimus; k, second chalimus; l, third chalimus; m, fourth
chalimus; n, fifth chalimus; o, sixth chalimus; p, adult. For minor differences between male and female adult
maxilliped, see description of appendage.

Lepeophthewus dissimulatus — Lewis

225

the series of tine-like projections of the copep-
odite process. The arrangement of the terminal
processes thus appears to be reversing itself
and, as will be described for later stages, does
reverse so that the inner process is complex and
the outer simple.

The second chalimus maxilla (Fig. 17c) ap-
pears to have degenerated slightly from that of
the preceding stage. The shape varies somewhat
in different specimens and, in general, appears
to be of a rather plastic nature. The first and
second segments are of approximately equal
length, the first appearing more strongly de-
veloped than the second. The outer proximal
corner of the first segment is heavily sclerotized
and appears to function as an articulation sur-
face. The distal end of the first segment is
slightly narrower than the proximal, and the
lateral margins are somewhat irregular; the dis-
tal region is heavily sclerotized on the inner
portion although no distinct articulation sur-
faces are visible. The second segment is nar-
rower proximally than distally, the segment ap-
pearing almost club-shaped. The proximal mar-
gin of the segment is almost included in the
inner lateral margin, both lateral margins are
somewhat wavy, and the distal margin is
rounded. The inner terminal process is relatively
longer than in the preceding stage, now being
approximately twice the length of the outer
process; further, the process is tapered to a sharp
point and possesses several minute setules on the
distal end of the outer margin. The inner termi-
nal process is simple and lobate, almost identical
to that of the first chalimus.

The maxilla of the third chalimus (Fig. lid)
differs somewhat from that of the second chal-
imus. The first segment is approximately two-
thirds the length of the second, its proximal end
projecting as a curved, lobate process that serves
as an articulation surface. The outer lateral mar-
gin is broadly convex, the inner flatly convex;
the greatest width is slightly less than one-half
of the greatest length and the general shape of
both of the segments appears to be more distinct
than in the preceding stage. The distal margin
of the first segment is irregular and heavily
sclerotized, the irregularities serving as articu-
lation surfaces for the proximal end of the sec-
ond segment. The second segment is slender,
and the inner and outer margins are generally

flatly convex. The proximal margin of the sec-
ond segment is now completely included in the
inner margin. The proximal surface possesses
two small knoblike projections with a concave
surface between them; both of the projections
and the concave surface articulate with recipro-
cal processes on the irregular distal end of the
first segment. The middle of the inner margin
of the second segment now has a short groove
extending from the surface inward and proxi-
mally; this groove corresponds to the position of
one of the spines present on the adult append-
age. The distal end of the segment is tapered to
a medial point and possesses the two terminal
processes present on preceding maxilla-bearing
stages. The inner of the two processes now
curves sharply outward, over the distal end of
the outer process, and is minutely frilled along
its outer distal margin. The outer process is
lobate, approximately two-thirds the length of
the inner process; the distal margin bears a
fine membrane.

The maxilla of the fourth chalimus (Fig. 17 e)
still bears the somewhat degenerate appearance
of the third chalimus appendage. Except for the
distal end, the first segment is similar to that
of the preceding stage. The outer distal margin
of the segment is concave; the distal end of the
inner lateral margin is also concave, forming a
knoblike projection at the junction of the two
which is heavily sclerotized. The proximal sur-
face of the second segment, included in the
inner lateral surface, is bilobed. The bilobed
condition of the surface appears to be due to a
split or division occurring in the small concav-
ity separating the two heavily sclerotized knobs
of the previous stage. The separation between
the two lobes forms an articulation surface for
the knoblike projection of the inner distal sur-
face of the first segment and the lobes them-
selves articulate in the concavities on the distal
and inner distal lateral surfaces of the first seg-
ment. The middle of the inner lateral margin of
the segment is indented in the region of the
groove of the preceding stage and bears a single
spinule. The second segment is tapered slightly
from the indentation to the distal end, which
is tipped by the two terminal processes. The in-
ner terminal process is similar to that of the
preceding stage, the outer process is slightly
longer, and the fine membrane now extends

226

around the distal end and distal lateral margins.

Both segments of the maxilla of the fifth
chalimus female and male (Fig. 17/) are more
slender and appear more elongate than those of
the preceding stage. The first segment is slightly
shorter than the second and its proximal end
projects as an irregular, lobate articulation proc-
ess. The outer margin of the segment is flatly
convex; the inner is slightly convex proximally,
flat medially and concave distally. The outer dis-
tal margin is tapered abruptly to a knoblike pro-
jection of the inner distal surface that appears
to be the enlarged projection of the distal end of
the first segment of the fourth chalimus. The
second segment is slender, the proximal end
two-parted and included in the inner lateral
margin. The junction of the two parts of the
proximal end forms a concave surface that ar-
ticulates on the knoblike process on the distal
end of the first segment. The outer lateral mar-
gin of the second segment is generally flatly
convex; the inner lateral margin is slightly ir-
regular, with a slight indentation in the middle
that bears a single spinule and a fine membrane
connected to the spinule and extending proxi-
mally for a short distance. The distal end of the
second segment is almost flat and bears two
terminal processes. The inner terminal process
is relatively longer than in the preceding stage,
being 17/2 times the length of the outer process;
the outer margin is fringed with a fine, lightly
frilled membrane. The outer process is lobate
and covered with minute, hairlike projections.

The maxilla of the female and male sixth
chalimus (Fig. 17g) is similar to that of the
fifth chalimus in general shape. The segments,
and especially the proximal projection of the
first segment, are heavier; the knoblike projec-
tion of the inner distal surface is on the inner
corner so that it forms the junction of the inner
lateral and inner distal surfaces. The proximal
end of the second segment, included in the in-
ner lateral surface, again appears two-lobed, the
proximal lobe extending proximally and form-
ing the point of attachment for the muscle that
abducts the second segment. The distalmost of
the two lobes forms an indentation in the mar-
gin that extends for a short distance although
there is a continuation, as a distinct line of di-
vision between the two lobes, running almost
parallel to the lateral margins of the segment.

PACIFIC SCIENCE, Vol. XVII, April 1963

The middle of the lateral surface of the second
segment is tapered slightly but abruptly; the
spinules and membrane present in this region
in the adult and in the preceding stage were not
present on any of the specimens examined. The
two terminal processes on the flat distal end of
the second segment are both slightly larger than
in the preceding stage; the outer is now pointed
distally and both have a fine, membranous mar-
gin along their inner and outer surfaces.

The maxilla of the adult female and male
(Fig. 17 h) is more definitely shaped than any
of the preceding stages and the first segment
more strongly developed than in the preceding
two chalimus stages. The first segment is short
and stocky, the inner proximal end protruding
as an irregular articulation surface. The distal
lateral margins of the first segment are tapered
irregularly to the somewhat narrow distal sur-
face, the median portion of which projects as
a narrow, lobate articulation surface that is heav-
ily sclerotized. The second segment is elongate,
the lateral margins irregularly convex. The
proximal surface, included in the inner lateral
surface, is irregular, with a slight concavity that
forms the place of articulation for the medial
projection of the distal surface of the first seg-
ment. The middle of the inner margin is notched
in two places, the proximal notch bears a minute
spine not present in any preceding stage, and
the distal notch bears a larger spine, about twice
the length of the proximal spine. A fine mem-
brane is also present and connects the two
spines. The two terminal processes arising from
the step-shaped distal end of the second seg-
ment are more elongate than those of the pre-
ceding stage. The innermost of the two proc-
esses is about \l/2 times the length of the outer
and possesses a membrane along the outer mar-
gin. The outer process is fringed by a frilled
membrane along the outer margin.

Maxilliped

The maxillipeds have been termed the first
thoracic appendages in copepods as well as in
other crustaceans and the maxillae as the last
cephalic appendages (Borradaile et aL, 1958:
347-348 ) . This terminology is accepted by most
authors, including those who apply the term
"first maxilla” to the postoral process, but is
not accepted by those workers who call the post-

Lepeophtheirus dissimulatm^—LEW is

antennal process the first maxilla and the post-
oral process the second maxilla. In this latter
case the first pair of identifiable appendages
behind the mouth thus become the first maxil-
lipeds and the last pair of appendages before
the thoracic legs become the second maxillipeds.
With the use of the term "maxilla” for the first
pair of identifiable appendages behind the
mouth, the term "maxilliped” is here applied to
the pair of appendages anterior to the first tho-
racic legs.

The maxillipeds are first present in the co-
pepodite ( Fig. 17 i) as uniramous, two-seg-
mented appendages attached to the ventral sur-
face of the cephalothorax posterior and slightly
medial to the base of the maxillae. The first
segment is slightly less than twice the length of
the second; its proximal end is broad and has a
poorly developed articulation surface that pro-
jects from the middle of the surface. The lateral
margins of the first segment are slightly curved,
the anterior margin flatly concave, the posterior
flatly convex; the distal anterior margin is tap-
ered sharply, inward to the narrow distal end.
Both the distal and inner distal lateral surfaces
have several rather heavily sclerotized projec-
tions and depressions, each fitting depressions
or receiving projections from the proximal sur-
face of the second segment. The second segment
is slender, the inner margin shorter than the
outer although both are slightly irregular. The
distal region of the second segment is tapered
to the short distal surface which bears a strongly
developed, spinelike terminal process that curves
medially evenly. A slender, short, spinelike struc-
ture is present as an accessory process and arises
from the inner distal surface of the segment.
The division between the segment and the ter-
minal process is distinct and complete, that be-
tween the segment and accessory process indis-
tinct and incomplete; the terminal process bears
a membranous fringe along the inner margin.

The first chalimus maxilliped (Fig. 17;) is
more strongly developed than in the copepodite.
The first segment is approximately the same
length, with regard to the second segment, al-
though appearing heavier, its width being
slightly greater than one-third of its length. The
inner proximal surface of the first segment pro-
jects as a tapered articulation process, and the
lateral margins of the segment are flatly convex,

227

the outer margin longer than the inner; the dis-
tal surface is angled inward and downward and
possesses two distinct, heavily sclerotized articu-
lation surfaces for the second segment. The sec-
ond segment, including the terminal process, is
tapered from the broad proximal to pointed
distal end. The proximal region of the segment
has two lobate projections, one from the ante-
rior surface and the second from the posterior,
which articulate with the indentations of the
distal surface of the first segment. A small seta-
like accessory process arises from the inner sur-
face of the segment, at the distinct junction of
the segment and the terminal process. The ter-
minal process is heavily sclerotized, curves in-
ward, and lacks the marginal membrane of the
previous stage.

The changes that occur in the maxilliped
from the first chalimus to the adult (Fig. 17 k-p)
are relatively minor. The general shape remains
the same and the armature of the Second seg-
ment does not change. The articulation surface
that projects from the proximal region changes
slightly, becoming somewhat longer in the sec-
ond chalimus ( Fig. Ilk) and then gradually
curving in the third chalimus (Fig. 17/) and
fourth chalimus (Fig. 11 m) until it becomes
almost recurved in this and the fifth chalimus
stages. The articulation projection then becomes
lobate in the sixth chalimus (not visible in Fig.
17o) and the adult (Fig. Up). The inner sur-
face becomes grooved in the fifth chalimus and
remains that way up to and including the adult.
The division between the second segment and
terminal process becomes rather indistinct in
later stages of development. The adult male
maxilliped has a small shelflike structure that is
present on the middle of the inner margin of
the first segment. This structure appears in the
last moult, from the sixth chalimus to the adult,
and is the only apparent difference between the
maxilliped of the female and the male.

Sternal Furca

The sternal furca is a single process between
and slightly posterior to the maxilliped bases
and is another structure about which little is
known. Most authors have simply reported its
presence or absence and utilized this as a tax-
onomic characteristic. The position of the struc-
ture, posterior to the maxilliped bases, does not

228

PACIFIC SCIENCE, VoL XVII, April 1963

allow any specific interpretation as to its origin.
If the maxillipeds are the first pair of thoracic
appendages, the sternal furca may be the rem-
nant of the sternal plate which exists between
the first three pairs of thoracic legs of L. dis-
simulate and many other caligoids. No defini-
tive evidence for this idea can be offered, how-
ever, other than the relative position to the

maxillipeds and the presence of sternal plates
on some of the thoracic legs. Futhermore, the
sternal furca in L. dissimulatus does not appear
until the fourth chalimus, long after the maxil-
lipeds, the thoracic legs, and the sternal plates
of the first two thoracic legs. The hypothesis
has, therefore, at least one pitfall unless, and no
information is given to support this, a shift can

Fig. 18. Sternal furcae and first thoracic legs of developmental stages of L. dissimulatus. a-e, Sternal fur-
cae: a, Third chalimus; h, fourth chalimus; c, fifth chalimus; d, sixth chalimus; e, adult, f-m, First thoracic
legs: f, Copepodite; g, first chalimus; h, second chalimus; i, third chalimus; j, fourth chalimus; k, fifth chalimus;
l, sixth chalimus; m, adult.

Lepeophtheims dissimulatus — Lewis

229

occur in the time that a structure appears in
development. If this were possible, then the
status of the postantennal process must also be
reviewed. The presence of a short but distinct
muscle that is attached to the sternal furca and
the mobility of the process exhibited by adult
specimens make it difficult to accept Heegaard’s
suggestion (1947:77-78) that the sternal furca
is a cuticular spine.

The sternal furca of the fourth chalimus is
visible developing underneath the third chal-
imus cuticle (Fig. 18*). The process is indis-
tinctly visible between and slightly posterior to
the bases of the maxillipeds but does not pro-
ject from the ventral surface. The proximal end
is broadly rounded, the bifurcation extends less
than one-half of the total length of the process,
and the tines are sharply pointed.

After the moult into the fourth chalimus, the
sternal furca (Fig. 18 b) is distinct, projecting
from the ventral surface of the cephalothorax.
The greatest width of the process is approxi-
mately two-thirds of the greatest length. The
bifurcation extends slightly more than two-
thirds of the length of the process. The tines are
angled outward slightly and are bluntly rounded
distally.

The sternal furca of the fifth chalimus (Fig.
18c) is well developed. The bifurcation of the
process extends about one-half the length of the
process. The tines appear distinct, separated
from the base of the process by a groove; they
are also angled outwards slightly and are bluntly
pointed.

The tines of the sternal furca of the sixth
chalimus do not appear distinct from the base as
in the preceding stage. The bifurcation extends
slightly more than one-half the length of the
process; the tines are angled outward slightly
and are bluntly pointed.

The sternal furca of the adult male and fe-
male (Fig. 18e) is more heavily developed than
in the preceding stages. The tines have a flat
inner surface in contrast to the round inner sur-
face of the previous stages but are still angled
outwards slightly and have a blunt tip.

Thoracic Leg 1

The armature and character of the mem-
branes, spines, and setae comprising the arma-
ture, are given in Table 2.

The first thoracic leg is present in the copep-
odite. The appendage (Fig. 18/) is biramous;
the protopodite, exopodite, and endopodite are
each one-segmented. The protopodite is wider
distally than proximally, its greatest width
slightly more than the greatest length; a single
lightly plumose seta is present just lateral to the
base of the exopodite and an indistinct, seta-like
process is present on the proximal inner margin.
The exopodite is palm-shaped, its lateral and
distal margins continuous. The outer margin of
the exopodite segment bears four spines, the
proximal three of which are simple, the fourth
with a membrane along the outer margin. The
endopodite is also palm-shaped, the lateral and
distal margins again continuous. The outer distal
surface bears a small, triangular, spinelike pro-
jection in addition to the setae shown in Table 2.

The distinct shape of the copepodite first tho-
racic leg is lost in the moult from the copepodite
into the first chalimus. The first thoracic leg of
the first chalimus (Fig. 18g) appears flabby,
the armature seems to have degenerated, and
the long plumose setae borne by the exopodite
and endopodite of the copepodite first leg, and
presumably used in swimming, are lost and re-
placed by short, lightly plumose setae. The pro-
topodite is one-segmented, its width and length
about equal; the distal end is somewhat nar-
rower than the proximal although this is vari-
able in different specimens. A single lightly
plumose seta is present on the distal lateral sur-
face just lateral to the base of the exopodite,
and the seta-like process of the preceding stage
is absent. The exopodite is one-segmented and
dactyliform, the proximal end is wider distally
than proximally, and the greatest length is ap-
proximately 2l/z times the greatest width. The
lateral margins of the segment are slightly ir-
regular, the distal margin broadly curved. A
lightly plumose seta is present on the middle of
the outer lateral surface and six lightly plumose
setae are present on the distal end. The spines
present on the exopodite segment of the copep-
odite appendage are completely absent in the
first chalimus. The endopodite is one-segmented
and irregularly lobate, its length slightly less
than two-thirds that of the exopodite. The en-
dopodite bears two lightly plumose setae on
its distal surface.

Two distinct trends can be noticed in the

230

PACIFIC SCIENCE, Vol. XVII, April 1963

TABLE 2

Armature of First Thoracic Legs of Developmental Stages*

STAGE

MARGIN

PROTOPODITE

EXOPODITE

1 2

ENDOPODITE

Copepodite

outer

3h,lHm

inner

IP'

3P,1Q

Chalimus I

outer

lp,3P

inner

Chalimus II

outer

1P,3P

inner

Chalimus III

outer

lrh

lrh?

inner

3P,1P

Chalimus IV

outer

lrh

lrh?

inner

3P,1P

lrh?

Chalimus V

outer

lrh

lrh?

inner

3P,1P

lrh?

Chalimus VI

outer

lrh

lrh?

inner

3P,1P

lrh?

Adult

outer

lrh

3H,1P

inner

* For explanation of symbols see Fig. 1.

gradual change of the first thoracic leg from the
first chalimus stage to the adult. There is a
gradual reduction in the size of the endopodite,
with regard to the exopodite. Both the exopo-
dite and the endopodite are of approximately
equal size in the copepodite but even in the first
chalimus the endopodite is distinctly smaller
than the exopodite, a change that is carried
through later development until in the adult the
endopodite is a mere rudiment, vastly smaller
than the exopodite. The second trend is the
gradual elongation of the exopodite segment
from the palm-shaped copepodite segment to
the elongate second chalimus segment (Fig.
18/?), then the division of this segment into two
segments in the third chalimus (Fig. 18z) and
the later elongation of each of the two seg-
ments, especially the proximal, to the adult con-
dition (Fig. 18 m).

The armature of the first thoracic leg changes
in character more than in quantity from the first
chalimus to the adult. The second chalimus
exopodite bears seven lightly plumose setae on
the distal surface and one short, lightly plumose
seta from the middle of the outer lateral surface.
The exopodite of the third chalimus, a two-
segmented structure, bears three spines and one
lightly plumose seta on the distal surface and
three lightly plumose setae on the inner lateral
surface of the second or distalmost segment in

addition to a single, spinelike process on the
outer distal lateral corner of the first segment.
The total number of processes on both of the
segments is the same as the number on the sin-
gle exopodite segment of the second chalimus
although the character is quite different. The
armature of the exopodite remains the same as
that of the third chalimus throughout develop-
ment except for the addition of a row of setules
along the inner surface of the first segment.
The two lightly plumose setules on the distal
surface of the endopodite segment of the first
chalimus are found through most of the devel-
opment (see Table 2) although distinctly re-
duced in size until in the adult they appear as
minute, indistinct projections from the distal
surface of the rudimentary endopodite. One
other change occurs in the armature of the first
thoracic leg, the addition of a small, lightly plu-
mose setule on the proximal inner margin of
the protopodite segment of the third chalimus.

Some change occurs in the shape of the first
thoracic leg segments but these are of a rather
minor nature and are shown in Figure 18 f~m.

Thoracic Leg II

The armature and character of the mem-
branes, spines, and setae comprising the arma-
ture are given in Table 3.

Lep eophthewus dissimulatus- — Lewis

231

The second thoracic leg is first present in the
copepodite stage. As with the first thoracic leg,
the appendage appears to degenerate in the first
few chalimus stages and then gradually takes on
the appearance of the adult appendage in the
late chalimus.

The second thoracic leg of the copepodite
(Fig. 19 'a), as in all of the remaining stages, is
biramous. The protopodite is one-segmented,
the width of the proximal and distal ends being
about equal and the greatest length about three-
fourths of the width. The lateral margins of the
segment are broadly convex, and the distal mar-
gin is irregular. A single lightly plumose seta
is present on the protopodite just lateral to the
exopodite base. The exopodite is one-segmented
and ovoid, and the distal margin is distinct from
the outer lateral margin but is continuous with
the inner. The proximal end of the exopodite is
minutely bilobed and heavily sclerotized, form-
ing an articulation surface that is contiguous
with irregularities on the lateral distal surface
of the protopodite. The outer surface of the ex-
opodite bears three spines, the proximal two
of which are simple, the distal long and fringed
by a membrane along its outer margin. The

endopodite is one-segmented, the proximal end
narrow and the distal end broad; the distal two-
thirds of the segment is bent inwards. The lat-
eral and distal margins of the segment are both
distinct. Both the exopodite and endopodite
bear several plumose setae as indicated in Ta-
ble 3.

The second thoracic leg of the first chalimus
(Fig. 19 b) is much more simple than the copep-
odite appendage. The protopodite is one-seg-
mented, and its greatest length is approximately
two-thirds of its greatest width. Both of the lat-
eral margins are continuous with the distal mar-
gin although the entire outline of the segment
is somewhat irregular. The single seta present
on the copepodite protopodite is absent in this
stage. The exopodite and endopodite are of ap-
proximately equal length and both are dactyli-
form. The exopodite is approximately one-half
as wide as long although slightly wider distally
than proximally; the lateral margins are wavy,
the distal broadly rounded. A single naked seta
is present on the middle of the outer lateral sur-
face and five lightly plumose setae on the distal
surface. The endopodite is irregularly rounded
distally; the lateral and distal margins are con-

TABLE 3

Armature of Second Thoracic Legs of Developmental Stages*

STERNAL

PROTOPODITE

EXOPODITE

ENDOPODITE

STAGE

MARGIN

PLATE

Copepodite

outer

2h,lHm

inner

3P,1Q

Chalimus I

outer

lp',3P

inner

Chalimus II

outer

2P,2P

inner

1P,3P

Chalimus III

outer

filH

4H,3P

c,2P

inner

fils

c,2P

Chalimus IV

outer

filH

3H,1Q,2P

2P,2P

inner

fils

C,1P

c,3P

Chalimus V

outer

filH

3H,1Q,2P

c,C

c,lP,3P

inner

fils

C,1P

c,3P

c,3P,2P

Chalimus VI

outer

filH

2H,1Q,3P

c,C

c,3P

inner

fils

C,1P

c,lP

c,2P

Adult

outer

filp

filH

2H,1Q,2P

c,C

c,3P

inner

Is, IP

fils

C,1P

c,3P

c,2P

c,3P

* For explanation of symbols see Fig. 1.

Fig. 19- Second thoracic legs of developmental stages of L. dissimulatus . a, Copepodite; b, first chalimus;
c, second chalimus; d, third chalimus; e, fourth chalimus; f, fifth chalimus; g, sixth chalimus; h, adult.

tinuous. The greatest width of the endopodite
segment is approximately one-half of the great-
est length; the distal surface bears four lightly
plumose setae.

Some enlargement has occurred in the sec-
ond thoracic leg of the second chalimus (Fig.
19c), especially in the protopodite. The protop-
odite is still one-segmented, its greatest width
being approximately three-fourths of its greatest
length. The proximal margin of the segment

is indistinct, and the lateral margins are almost
parallel. The protopodite is approximately twice
the length of the first chalimus protopodite al-
though its width is about the same. The one-
segmented, dactyliform exopodite is attached to
the distal surface of the protopodite. The great-
est width of the segment is approximately one-
half of the greatest length; the lateral margins
are irregular, the distal rounded. Two lightly
plumose setae are present on the distal one-half

Lep eophth eirus dissimulatus^-LEWis

233

of the outer surface and six are borne on the
distal surface. The one-segmented, dactyliform
endopodite is attached to the inner distal sur-
face of the protopodite. The greatest length of
the segment is about two-thirds of the length
of the exopodite, its width about one-half of its
length. The endopodite is tapered slightly to-
wards the rounded distal end which bears five
lightly plumose setae; an additional lightly plu-
mose seta is present on the middle of the inner
margin.

As with the first thoracic leg, the second
shows a rather distinct transition in the moult
from the second chalimus to the third. The ap-
pendage in the third chalimus (Fig. 19 d) has
changed from the rather generalized condition
in the second chalimus to a condition which,
with some exceptions, resembles that of the
adult.

The protopodite of the second thoracic leg of
the third chalimus is two-segmented. The first
segment is narrow and forms a strip between
the sternal plate and the second segment. A
single plumose seta arises from the inner surface
of the first segment. The second segment is
broad and flattened, its greatest width is almost
three-fourths of the greatest length, the outer
margin is irregular, and the inner margin is
convex. The exopodite, which is attached to the
outer distal surface, is two-segmented. The
length of the first segment is slightly greater
than that of the second, and the greatest width
is approximately two-thirds of the length; the
lateral margins are irregularly convex. A long
spine is present on the outer distal surface and
is directed distally; the length of the spine is
slightly less than two-thirds of the segment
length. The second segment is slightly longer
than wide, and the proximal end is narrower
than the distal; both of the lateral margins and
the distal margin are distinct, not continuous,
and are irregular; the outer lateral surface bears
four spines on the distal one-half of the segment.
The proximalmost of the four spines is short
and directed distally and laterally; the second
spine is directed in the same manner as the
first but is considerably longer; the third is
directed laterally, curves distally, and is about
the same length as the second; the fourth or
distalmost spine is directed laterally and is more
strongly developed than the preceding three.

The endopodite, attached to the inner distal sur-
face of the protopodite, is also two-segmented.
The first segment is approximately two-thirds as
wide as long and is curved inwards, the outer
lateral margin being much longer than the inner
and convexly curved. The second segment is
slightly less than IVl times the length of the
first and is longer than wide; both of the lateral
margins are flatly convex and distinct from the
distal margin, which is irregularly rounded. Both
the exopodite and the endopodite bear setae, as
indicated in Table 3.

The second thoracic leg of the fourth chal-
imus (Fig. 19e) is similar to that of the third
chalimus. There has been some expansion of
the protopodite segments and the first exopodite
segment. The setae appear to be more plumose
and, in general, the armature is more like that
of the adult.

The second thoracic leg of the fifth chalimus
(Fig. 19/) is similar to that of the preced-
ing stage although changes have occurred in
the shape and the armature. The protopodite is
two-segmented, the first segment short, being
approximately one-third the length of the sec-
ond segment; the width of the first segment is
approximately three-fourths of the length. The
outer margin of the first protopodite segment
is longer than the inner, the proximal margin
is angled, and the distal is straight. Further, the
outer proximal surface of the segment has a
small knoblike projection that articulates in a
C-shaped depression in the end of the sternal
plate (not shown in the figure), and the inner
margin bears a single plumose seta. The second
segment is swollen distally, the greatest width
of the distal region being approximately four-
fifths of the greatest length; the outer lateral
margin of the segment is irregular, the inner
broadly convex. The exopodite is still two-seg-
mented, the first segment slightly longer than
the second. The outer margin of the first seg-
ment is convex, and the inner margin is slightly
irregular. The outer distal corner of the first seg-
ment bears a long, distally projecting spine that
extends to the distal region of the second seg-
ment. The second segment is palm-shaped, being
wider distally than proximally. The greatest
width of the segment is approximately two-
thirds of the greatest length; the outer lateral
margin is irregular and bears three strongly-

234

PACIFIC SCIENCE, Vol. XVII, April 1963

developed spines, the proximal two extending
distally and laterally, the distalmost extending
laterally and curving distally. The simple fourth
spine of the third chalimus bore a membrane
on the inner margin and a row of setules on the
outer in the fourth chalimus; this condition is
reversed in the fifth chalimus. The endopodite
is two-segmented; the first segment is short, its
width slightly greater than the length which is
approximately one-half that of the second seg-
ment. Both of the lateral margins of the first
segment are convex, the outer twice the length
of the inner. The second segment is irregularly
palm -shaped, and the lateral margins are irregu-
lar although distinct from the distal margin;
the greatest width of the segment is approxi-
mately two-thirds of the greatest length.

The second thoracic leg of the sixth chalimus
(Fig. 19g) is basically the same as that of the
adult. The protopodite is two-segmented; the
first segment is short, less than one-third of the
length of the second. The second segment is
broader distally than proximally, its greatest
width approximately two-thirds of the length.
The lateral margins of the second segment are
slightly wavy, the inner generally convex. The
exopodite is now three-segmented, the division
occurring in the second segment of the previous
stage, just distal to the proximalmost spine. The
first exopodite segment is slightly longer than
the combined lengths of the succeeding two
segments, and its lateral margins are flatly con-
vex. The single spine is slightly denticulated
and strongly developed and extends past the
distal end of the second segment. The second
segment is short, flared from the narrow proxi-
mal to broad, concave distal surface. A single
simple, although well-developed, spine is pres-
ent on the outer distal surface. The third seg-
ment is short, its length approximately equal to
that of the second segment. The distal one-half
of the segment is broadly rounded and a small,
steplike indentation is present in the proximal
one-half of the outer margin. A well-developed,
simple, distally projecting spine is present in
the indentation and a second well-developed,
simple spine projects laterally and curves dis-
tally from the outer lateral surface at the junc-
tion of the curved and indented margins. The
endopodite is also three-segmented. The first
segment is approximately equal in length to the

second; the outer margin is broadly rounded and
approximately 4 times the length of the short,
irregular inner margin. The second segment is
flared from the narrow proximal to broad distal
end, as in the second segment of the exopodite.
The outer margin of the second segment is flatly
convex, the inner slightly concave. The third
segment is short, being approximately three-
fourths of the length of the second segment.
The distal and lateral surfaces are irregularly
rounded, the margins continuous.

With the exception of some minor changes
in the shape and armature of the segments, the
description of the second thoracic leg of the
sixth chalimus can be applied to the adult ap-
pendage (Fig. 19^).

Thoracic Leg III

The armature and character of the mem-
branes, spines, and setae comprising the arma-
ture are given in Table 4.

The biramous third thoracic leg is first pres-
ent as a recognizable appendage in the first chal-
imus (Fig. 20 a). As mentioned earlier, how-
ever, two spinelike processes are present on the
third pedigerous segment of the copepodite and
may represent the third legs in this stage of de-
velopment. The protopodite of the third tho-
racic leg of the first chalimus is one-segmented,
its greatest width and length about equal. The
somewhat irregular outer lateral margin is more
than twice the length of the inner and two naked
setules are present on the outer lateral surface
just lateral to the exopodite base. The exopodite
is one-segmented, broader than long; its lateral
margins are generally convex and the distal
margin is flat although variable in shape in dif-
ferent specimens. Four spinule-like processes
project from the distal region of the exopodite.
The endopodite is also one-segmented and
broadly rounded, both of the lateral and the dis-
tal margins being continuous; the greatest width
of the segment is slightly more than the length.

The protopodite of the third thoracic leg of
the second chalimus (Fig. 20 b) is much larger
than that of the preceding stage, most of the
increase in size being due to the increase in the
lateral surface. The irregular proximal end of
the one-segmented protopodite is broad, ap-
proximately twice the width of the distal end.
The outer lateral margin is almost 4 times the

Lepeophtheirus dissimulatus — Lewis

235

length of the inner. The exopodite and the en-
dopodite differ from those of the preceding stage
in shape and armature. The one-segmented ex-
opodite is broader than long, the lateral margins
are irregular although slightly convex, and the
distal margin is irregularly rounded. Both the
outer and inner lateral margins have several
irregularly spaced setules and the distal surface
bears two spinelike processes. The one-seg-
mented endopodite is about one-half the length
of the exopodite although both the length and
width of the segment are approximately equal;
the lateral margins are almost parallel, the distal
margin slightly rounded, and the inner distal
surface gives rise to two spinelike processes.

The third thoracic leg of the third chalimus
(Fig. 20c) is distinctly different from that of
the second chalimus. It appears that the third
chalimus marks a distinct change in the first
three pairs of thoracic legs. The protopodite is
now broader although the added breadth ap-
pears to have been gained as a result of a length-
ening of the distal margin or an inclusion of
the inner lateral margin with the distal; both
the exopodite and the endopodite are now two-
segmented and the armature has changed.

The protopodite of the third thoracic leg of
the third chalimus is broad and flattened. The
greatest width, across the proximal end, is

slightly less than twice the length; the inner
lateral margin appears to be incorporated into
the distal margin although distinct evidence of
this is lacking. A single plumose seta is present
just lateral to the junction of the protopodite
and sternal plate. The exopodite, attached to
the lateral region of the distal protopodite sur-
face, is two-segmented, the first segment slightly
shorter than the second; the outer margin of
the first segment is convex, the distal margin
sloping to the proximal margin in older speci-
mens but not distinctly so in younger specimens.
The second segment is as wide as long, the lat-
eral margins are irregular, the distal margin is
rounded, and the surface bears three lightly plu-
mose setae. The endopodite is two-segmented
although the division between the first segment
and the protopodite is indistinct in older speci-
mens. The outer margin of the first segment is
longer than the inner, and the proximal and
distal margins are irregular. The second segment
is approximately equal to the greatest length of
the first segment and is almost circular in out-
line.

The protopodite of the third thoracic leg of
the fourth chalimus (Fig. 20 d) is broader than
that of the third and the length has also in-
creased so that the greatest proximal width is
only about three-fourths of the greatest length.

TABLE 4

Armature of Third Thoracic Legs of Developmental Stages*

STERNAL

EXOPODITE

ENDOPODITE

STAGE

MARGIN

PLATE

PROTOPODITE

Chalimus I

outer

2p'

inner

Chalimus II

outer

c,2h

inner

Chalimus III

outer

inner

lP,f

Chalimus IV

outer

lp,2P

inner

lP,f

Chalimus V

outer

f,lP

c,4p'

c,2P

inner

lP,f

1 lp,lH'

Chalimus VI

outer

f,lP

c,lp'

c,3p',lP

c,lp,3P

inner

lP,f,2s

1H'

Adult

outer

f,lP

C,lp'

c,3pMP

inner

lP,f,2s

1H'

C,1P

c,3P

c,5P

For explanation of symbols see Fig. 1.

236

PACIFIC SCIENCE, Vol. XVII, April 1963

Q03mm.

0.03 mm

d 0.05 mm 9

doosmm?

SlrTmCJ?

Fig. 20. Third and fourth thoracic legs of developmental stages of L. dissimulatus. a—g, Third thoracic
legs: a, First chalimus; b, second chalimus; c, third chalimus; d, fourth chalimus; e, fifth chalimus; f, sixth
chalimus; g, adult, h-n, Fourth thoracic legs: h, First chalimus; i, second chalimus; j, third chalimus; k, fourth
chalimus; l, fifth chalimus; m, sixth chalimus; n, adult.

Lep e op ht heir us dissimulatus — Lew is

237

The protopodite tapers distally, the lateral mar-
gins are slightly irregular, and the distal margin
is continuous with the outer lateral margin. The
exopodite is two-segmented, the second segment
2 Vl times the length of the first. The base of
the spine-bearing first segment of the fifth chal-
imus is visible inside the first segment of the
fourth chalimus; the lateral margins of the
fourth chalimus segment are convex. The spine-
bearing segment mentioned above and present
in the fifth and sixth chalimus in addition to the
adult replaces the first segment of the third and
fourth chalimus; the second segment of these
two stages remains as the second segment, which
later divides to form the second and third seg-
ments. The lateral and distal margins of the sec-
ond exopodite segment of the fourth chalimus
are continuous, the distal end is broadly rounded,
and the surface gives rise to three lightly plu-
mose setae. The endopodite is two-segmented
although the division between the first segment
and protopodite is indistinct and incomplete.
The second segment is as wide as long and has
a broadly rounded outline with the lateral and
distal margins being continuous. The distal sur-
face of the segment bears four lightly plumose
setae.

The third thoracic leg of the fifth chalimus
stage (Fig. 20*?) possesses characteristics very
similar to those of the adult. The protopodite
is one-segmented and greatly expanded from the
previous stage. The sternal plate, connecting the
protopodites of the two legs, is broad and forms
a laminate projection from the posterior ventral
surface of the cephalothorax. The exopodite is
still two-segmented, the first segment possessing
a strongly-developed, spinelike projection from
the inner distal surface. The proximal end of the
first segment is distinct from the protopodite
ventrally although it is still fused dorsally to the
protopodite and the distal end to the second
segment. The first segment, other than the proc-
ess, is short, the width being greater than the
length. The second segment is broadly rounded,
the lateral margins continuous with the distal.
The proximal end of the segment is narrow and
the segment appears palm-shaped; the greatest
width is slightly more than two-thirds of the
length. The endopodite is two-segmented; the
first segment is twice as wide as long and ap-
pears as a narrow band with a swollen outer

lateral region. The second segment is attached
to the inner distal surface of the first and is
broadly rounded, the lateral and distal margins
being continuous. The greatest length of the
second segment is slightly greater than that of
the first; the width and length are, however, ap-
proximately equal.

With minor exceptions, the third thoracic
leg of the sixth chalimus (Fig. 20/) and the
adult (Fig. 20g) are the same. The protopodite
of the sixth chalimus is broadly flattened and
laminate; the lateral and distal margins are con-
tinuous. The exopodite is now three-segmented,
the first segment forming the base for a strongly
developed, spinelike process that projects from
the distal inner margin as in the preceding stage.
The lateral margins of the first segment are ir-
regular, the outer being generally convex and
the inner almost straight. The second segment
is short and flared from the narrow proximal to
broad, concave distal end. The third segment is
also short, being approximately equal in length
to the second, and the width is slightly greater
than the length; the lateral margins of the seg-
ment are rounded and continuous with the
rounded distal margin. The endopodite is two-
segmented, the first segment broad, its width
approximately ll/2 times its length. The outer
two-thirds of the distal margin of the first seg-
ment is convex, the inner one-third concave;
the outer lateral margin is continuous with the
distal margin, and the inner margin is irregular.
The second segment is slightly shorter than the
first segment and is attached to the concave in-
ner distal surface of the first. Both of the lateral
margins are continuous with the distal margin
in the second segment.

The protopodite of the adult third thoracic
leg (Fig. 20g) is broader than that of the sixth
chalimus; the first endopodite segment is slightly
longer, although not wider, and the armature
of the appendage has changed slightly in the
moult from the sixth chalimus to the adult.
Other than these differences, the appendages of
the two stages are the same.

Thoracic Leg IV

The armature and character of the mem-
branes, spines, and setae comprising the arma-
ture are given in Table 5.

The uniramous fourth thoracic leg is first

238

present in the first chalimus (Fig. 20 h) . The
appendage in this stage is a one-segmented lobe.
The proximal end is broader than the rounded
distal end, the lateral margins are wavy, and the
distal surface bears three spinule-like processes.

The fourth thoracic leg of the second chal-
imus (Fig. 20/ ) is lobate and somewhat longer
than the appendage of the preceding stage. An
indistinct division is present in the middle of
the appendage, dividing it into two segments.
Whether it is right to call the proximal segment
of the appendage the protopodite and the re-
maining segments the exopodite as this author
has previously done with regard to adult cali-
gids (Lewis, in press) was not determined. For
the present, however, because of the lack of
definite information on this problem, the terms
"protopodite” and "exopodite” will not be used
but the segments simply referred to by number,
with the proximalmost segment being the first.
The proximal end of the first segment of the
fourth thoracic leg of the second chalimus is
broad and angled so that the appendage projects
posteriorly and laterally from its junction with
the fourth pedigerous segment. The second seg-
ment bears three short, terminal, spinelike proc-
esses.

The fourth thoracic leg of the third chalimus
(Fig. 20)) has lost its general lobate shape and
is now three-segmented. The first segment is
short, its greatest length approximately three-
fourths of the width; the segment is tapered
from the broad proximal to somewhat narrower
distal end. The second segment is about twice
the length of the first and almost twice as long

TABLE 5

Armature of Fourth Thoracic Legs of
Developmental Stages*

STAGE

MARGIN

SEGMENTS

2 3

Chalimus I

outer

Chalimus II

outer

Chalimus III

outer

lh,2H

Chalimus IV

outer

lh,2H

Chalimus V

outer

lh,lh,2H

Chalimus VI

outer

Adult

outer

* For explanation of symbols see Fig. 1 .

PACIFIC SCIENCE, VoL XVII, April 1963

as broad; the lateral margins are irregular al-
though basically parallel, and the distal and
proximal margins are both irregular. The third
segment is greater than one-half the length of
the second, the lateral margins are flatly convex,
and the distal margin is rounded and continuous
with the inner lateral margin. Three spinelike
processes are present on the distal surface of
the third segment.

The fourth thoracic leg of the fourth chalimus
(Fig. 20k) is somewhat similar to the third
chalimus appendage. The length and width of
the entire appendage is slightly greater than that
of the preceding stage, the first segment is com-
paratively shorter and wider, the second seg-
ment appears swollen, and the third segment is
not as distinctly separated from the second.

The fourth thoracic leg of the fifth chalimus
(Fig. 20/) is three-segmented. The first seg-
ment is ovoid, the lateral and distal margins
broadly rounded and the outer distal lateral sur-
face bearing a single plumose setule. The divi-
sion between the second and third segments is
indistinct in contrast to the distinct division
present in the preceding stage. The combined
lengths of the second and third segments is ap-
proximately equal to that of the first segment.
One poorly developed spine is present on the
outer distal lateral surface and two well-devel-
oped and one poorly developed spines are borne
on the distal surface of the third segment.

The fourth thoracic leg of the sixth chalimus
(Fig. 20 m) is four-segmented. The first seg-
ment is slightly longer than the combined
lengths of the three distal segments, the proxi-
mal and distal ends of the segment are slightly
narrower than the middle of the segment, and
the lateral margins are wavy. A single plumose
setule is present on the outer distal lateral sur-
face of the segment, as in the preceding stage.
The second segment is short, its greatest length
slightly less than that of the third segment; the
lateral margins are flatly convex, the outer much
longer than the inner. A short, simple spine is
present on the outer distal corner of the second
segment. The third segment is tapered from the
proximal to slightly broader distal end; the lat-
eral margins are flat or flatly convex, the outer
longer than the inner. A single simple spine is
present on the outer distal corner of the seg-
ment. The fourth segment is slightly broader

Lepeophtheirus dissimulatus — Lewis

239

TABLE 6

Mean Length of Attachment Filament

CHALIMUS

COPEPODITE

Measurement (in mm)

0.136

0.111

0.146

0.156

0.155

0.163

0.213

Number of specimens

distally than proximally; the lateral margins are
flatly convex, the distal surface almost flat and
bearing three simple although well-developed
spines. The innermost of the three terminal
spines is longest, the outermost shortest.

The make-up of the fourth thoracic leg of
the sixth chalimus is similar to that of the adult
(Fig. 20#) although the armature is more sim-
ple. The segments of the adult, particularly the
distal three, are more elongate; the terminal
spines on each segment have at least one den-
ticulated or frilled margin and are encircled by
a frilled process at their base.

Caudal Rami

The caudal rami are first present in the co-
pepodite stage (Fig. 12 a). These structures, in
the copepodite, are slightly more than one-half
of the greatest length of the combined genital
segment and abdomen and their greatest width
is approximately equal to the length. Both of
the lateral and the distal margins are irregular.
Two short, plumose setae are present on the
proximal one-half of the outer lateral surface
and two long, plumose setae arise from the distal
surface.

The caudal rami of the first chalimus (Fig.
12b) arise from the posterior ventral surface of
the genital-abdominal segment. The greatest
length of the rami is approximately two-thirds
of the width; the lateral and distal margins are
continuous. Six plumose setae are present on
the rami, two on the distal lateral surface, three
on the distal surface, and one on the inner distal
corner.

The caudal rami of the remaining chalimus
stages and the adult (Fig. 12c-/) are similar
in shape and make-up to the rami of the first
chalimus. Minor differences such as the setula-
tion of the inner margin in the second chalimus
(Fig. 12c) and succeeding stages and a varia-
tion in the length of the setae are the only sig-

nificant changes that occur after the first chal-
imus.

REMARKS ON BEHAVIOR OF DEVELOPMENTAL
STAGES AND THEIR RELATION TO HOST

Both of the planktonic first and second nau-
pliar stages exhibit a strongly positive photo-
tropism in the laboratory. Movement in both of
these stages is sporadic and jerky, the three pairs
of appendages present on each of these stages
moving in unison in a series of strokes and then
remaining motionless for a short period of
time. The animal thus makes a short but swift
movement and then rests, sinking gradually.
Whether it is the response to light or a more
intrinsic factor that controls the amount of time
spent in swimming and in resting was not de-
termined. The function of the balancers present
on the posterior end of the nauplii has been
suggested to be the balancing of the nauplius
when it is at rest (Wilson, 1905:538). Since
these structures were not seen to move to any
extent in L. dissimulatus, their function cannot
be definitely ascertained although their structure
and position do suggest that they play a role in
the positioning of the nauplius during both
movement and rest.

As the age of the second nauplius increases,
the positive reaction to light decreases, a condi-
tion that extends into the planktonic phase of
the copepodite. A light placed on one side of a
finger bowl caused some copepodites to swim
towards that side. The tendency, however, was
not as great as in the naupliar stages and ap-
peared to diminish as the age of the copepodites
increased. The movement of the copepodite was
quite rapid although jerky and irregular; the
periods of time spent in swimming were longer,
and the time spent in resting shorter than in
either of the two naupliar stages. During the
resting phase the copepodite appeared to sink

240

faster than did the nauplius, which can be ex-
plained by the apparent increase in body volume
and the decrease in relative number and length
of setae. Copepodites were kept up to 72 hours
in culture. After about 10 hr they began to prod
the bottom and sides of the dish with the an-
terior end of their body. Inasmuch as the co-
pepodite is the attachment stage, the prodding
may have some relation to the attachment of
the animal to its host.

As was mentioned earlier, attachment was
observed in only one copepodite. Attachment
was accomplished by hooking the strong, claw-
like terminal processes of each second antenna
into the substrate. The formation of the attach-
ment filament was not seen in the single copep-
odite that attached in the laboratory. To de-
termine the means by which the filament was
formed, several copepodites were collected that
had attached to the host and formed a filament.
These animals were separated from the host by
severing the filament close to the body of the
copepodite. A piece of loosely woven nylon
cloth was then placed in a finger bowl and the
detached copepodites placed in with it. Those
specimens that reattached did so by holding
themselves to the cloth substrate by means of
their maxillipeds and second antennae. They
then placed their body at an angle to the sub-
strate, with the anterior end in close proximity
to it, and, by jabbing the substrate with the
middle of the frontal region, they appeared to
secure either the remaining attachment filament
or a new filament formed by a secretion that
hardened almost immediately on the substrate
as an irregular lobe, not the disk found in co-
pepodite and chalimus specimens normally at-
tached to the host. After the filament was at-
tached, the terminal processes of the maxillae
were used to stroke the filament between the
frontal region and the substrate while the co-
pepodite backed off from the point of attach-
ment. The backing off of the animal appeared
to cause the materal secreted by the frontal
organ, either before or after the animal started
backing off, to be drawn out into a filament and
the stroking by the maxillae to make the fila-
ment more even.

After the attachment, the activity of the co-
pepodite decreased. This observed lower activity
rate was noted for the first four chalimus stages

PACIFIC SCIENCE, Vol. XVII, April 1963

although activity did increase to some extent
during the actual shedding of the old cuticle in
moulting.

Heegaard (1947:90-94) indicates that the
various attached stages of Caligus curtus secrete
the attachment filament anew each time a moult
occurs. During the period of moulting observed
in both copepodite and chalimus stages in L.
dissimulates , the original filament remained at-
tached to the animal, the cuticle rupturing on
the anterior dorsal surface and the animal, by
vigorous movements of its body, freeing itself
from the old cuticle and passing it posteriorly
over the posterior end of the body. The shed
cuticle did not remain attached to the host or
to the frontal filament. Heegaard (1947: fig.
32) figures a portion of the fin ray of a cod
with several attachment filaments of Caligus cur-
tus hanging from it and (1947:92) suggests
that these are left by the various chalimus stages.
The only attachment filaments that were ob-
served in host tissue and could be definitely tied
in with a chalimus were those from specimens
of the fifth and sixth chalimus stages of L. dis-
simulatus that had just broken free and assumed
the free-moving adult type of existence.

Gurney (1934:184, 186, 188) indicates that
the attachment filament is increased in length
slightly at each moult as indicated by the small
annuli at the proximal end of the attachment
filament of later chalimus larvae. This could not
be verified or disproved for L. dissimulatus, as
the attachment filament is irregular at the proxi-
mal end at all stages of development, from the
copepodite to specimens of the sixth chalimus
that are still attached. The mean length of the
filament during each stage is given below in
Table 6 and suggests that a small amount of new
material is added although the lack of specimens
in some of the stages does not provide substan-
tial proof of this.

Shiino (1959:305) reports chalimus larvae
and young stages of L. dissimulatus from the
gills of Bodianus diplotaenia captured in the
Revilla Gigedos Islands and includes several fig-
ures of these stages. This report is of interest
because of the place of attachment of the chal-
imus larvae. With only two exceptions, all of
the chalimus larvae of Hawaiian specimens of
L. dissimulatus were found attached to the up-
per surface of the buccal cavity of the host. The

Lepeopbtbeirus dissimulatus — -Lewis

241

two exceptions were chalimus larvae found at-
tached to the gill membrane of Acant kurus tri-
ostegus sandvicensis . The number of larvae pres-
ent in the buccal cavity of infected fishes ranged
from 1 to 203. In the specimen of Acanthums
olivdcem possessing 203 attached copepodites
and chalimus larvae, the roof of the mouth ap-
peared to be actually shingled with copepods.

After breaking free from the attachment fila-
ment, the fifth or sixth chalimus moves out of
the buccal cavity either into the gill cavity or
onto the external surface. The final moult, from
the sixth chalimus to the adult, is accomplished
outside of the buccal cavity. Only one observa-
tion of the final moult was made and this was
after the adult had broken through a split in
the anterior dorsal surface of the sixth chalimus
cuticle. At the beginning of the observation the
animal had pulled the second antennae free
from the old cuticle and was not actively mov-
ing, holding on to the host by means of the
maxillipeds and keeping the second antennae
free. After a short period of time the second an-
tennae were used to grasp the surface and the
maxillipeds and remaining appendages were
pulled free from the cuticle. By means of violent
wriggling of the body and appendages, the cu-
ticle was shed over the posterior end of the
body. After a second short period of time, dur-
ing which the newly emerged adult remained
attached by. the second antennae, all of the tho-
racic appendages and the maxillae were moved
randomly and the animal then assumed the adult
role of skittering over the surface of the host.

Fertilization occurs just after the terminal
moult. The male at this stage of development
is almost completely grown; the female is still
small and will increase in size in the adult stage.
The actual placement of the spermatophores,
held within the genital segment of the male, was
not observed. Several mating pairs were col-
lected and observed, however. The male was
found to clasp the fourth pedigerous segment
or the anterior end of the genital segment of
the female with the second antennae. The func-
tion of the maxillipeds in mating was not de-
termined, although in all observed pairs the
maxillipeds of the male were free. A. Scott
(1901:28) and Wilson (1905:528) indicate
that the caligid spermatophore is viscid and,
as noted for L. dissimulatus , that it forms an

oval or tear-shaped body in the posterior region
of the genital segment. The male presumably
bends the posterior region of its body, the free
segments, underneath itself and, upon contact
with the posterior ventral surface of the female,
discharges the two spermatophores which ad-
here to the genital segment of the female. No
remating was observed and, as suggested by
Wilson (1905:527), the single mating just after
both sexes moult into the adult probably suffices
for the entire egg production of the female.

The young adult female was found in both
the gill cavities and on the external surface of
the body. Large females carrying egg strings
were found primarily in the gill cavity. The co-
pepod appears to have more protection in the
gill cavity, especially against being swept off
of the host by water currents or brushed off
when the host comes in contact with the sub-
strate. The position of the female on the host
thus may be dependant upon its size and upon
the presence or absence of egg strings which, it
appears, add a considerable burden.

The duration of the adult life is not known.
Females kept in the laboratory produced up to
three sets of egg strings, each string being ex-
truded approximately 21 hr after the previous
string had hatched. Thirty to 40 hr is the av-
erage interval between extrusion of the eggs
and hatching. The period of time between fer-
tilization and the production of the first pair
of egg strings is not known, neither is the time
from moulting into the adult until the female
is fertilized. Further, the conditions provided in
the laboratory were far from those in the natural
environment and it is felt that the number of
egg string sets produced is probably more than
the three obtained in the laboratory.

SUMMARY

1. The external anatomy of the 10 stages in
the life history of Lepeophtheirus dissimulatus
is described and figured.

2. The general behavior of the various stages
is discussed. The first two, the nauphar stages,
are planktonic. The third or copepodite stage is
planktonic early in its existence but later at-
taches to the fish host by means of the prehen-
sile second antennae. After attachment, the co-
pepodite secretes a frontal filament that is em-

242

PACIFIC SCIENCE, Vol. XVII, April 1963

bedded in the tissue of the host and enables the
copepodite and following attached larval stages
to remain attached to the host without using
their appendages. At least four of the six chal-
imus stages that follow the copepodite remain
attached to the host by the frontal filament; the
fifth and sixth chalimus stages may either be
found attached or free on the host. The adult is
found moving freely over the surface of the
host.

ACKNOWLEDGMENTS

Part of this study was aided by a grant from
the Sigma Xi-RESA Research Fund, 1961. The
author is grateful to Samuel Kaolulo and to
Lester Zukeran for host materal. He also wishes
to express his appreciation to the members of
his thesis committee for their counseling and
guidance during the preparation of his thesis,
which included the life history of L. dissimu-
latus.

REFERENCES

Borradaile, L. A., F. A. Potts, L. E. S. East-
ham, and J. T. Saunders. 1958. The in-
vertebrata. 3rd ed., revised by G. A. Kerkut.
Cambridge University Press, London, xvii -f-
795 pp.,. 523 figs.

Gurney, R. 1933. British fresh-water Copep-
oda. Ray Society, London, xxix -}- 384 pp.,
2061 figs.

1934. The development of certain para-
sitic Copepoda of the families Caligidae and
Clavellidae. Proc. Zool. Soc. London: 177—
217.

Heegaard, P. 1947. Contribution to the phy-
logeny of the arthropods. Spolia Zoologica
Musei Hauniensis 8:1-227.

Lewis, Alan G. (In press.) Caligoid copepods
(Crustacea) parasitic on fishes of the family
Acanthuridae in the Hawaiian Islands. Proc.
U. S. Nat. Mus.

Russell, F. S. 1925. A new species of Caligus
from Egypt, Caligus pageti , sp. n. Ann. Mag.
Nat. Hist., ser. 9, 15:611-618.

Scott, Andrew. 1901. Lepeophtheirus and
Lernaea . Trans. Liverpool Biol. Soc. 15:1-54.

Wilson, C. B. 1905. North American parasitic
copepods belonging to the family Caligidae,
Part I. The Caliginae. Proc. U. S. Nat. Mus.
28 ( 1404) : 479-672.

Studies on the Green Alga, U do tea indie a A. & E. S. Gepp, 1911

Mohammed Nizamuddin1

ABSTRACT: In Udotea indica reproductive organs are terminal and club-shaped.
Numerous biflagellate zooids are produced in the reproductive organs. Udotea is
named as the type of a new family, Udoteaceae.

Udotea indica A. & E. S. Gepp (1911) was
first collected by J. A. Murray (in 1880-83)
from Karachi near the mouth of the Indus
River. The type specimen is in the British Mu-
seum and the type locality is Karachi. Previous
taxonomic studies have been made, notably by
Boergesen (1930, 1934) and Taylor ( 1950).
This species has also been reported from Bikini
and other areas in the Marshall Islands by Taylor
(1950). It grows in association with Chaeto-
morpha sp. on silt-covered rocks along the edges
of littoral rocky pools, where it generally occurs
in patches, but elsewhere it may be found scat-
tered on rocky platforms.

The genus has been included in the family
Codiaceae by previous authors. The present
study excludes Udotea from the Codiaceae and
places it in a family of its own. The basis for
this conclusion is that in Udotea indica the ter-
minal portions of the filaments are transformed
into reproductive organs, in contrast to the lat-
eral position of the reproductive organs in the
genera of the family Codiaceae.

The discovery, made while examining living
material, that the reproductive bodies are motile
made a thorough investigation of this Udotea
necessary. The present account gives the pre-
liminary results of this study and, while not
completely clarifying many features, shows that
U. indica is not a member of the Codiaceae.
The structure and reproduction of this species
are discussed below.

The thalli are up to 4 cm long, as much broad,
and are slightly calcified. The root-mass forms
a small tuft. The terete stipe is up to 1.2 cm

1 Botany Department, University of Karachi, Ka-
rachi, Pakistan. Manuscript received December 19,
1961.

long and 1 mm thick. The fronds are green,
somewhat rounded, flabellate, orbicular, and
sometimes broadly proliferated above; the base
is cuneate, distinctly zonate. The blade margins
are entire, lobed, or lacerated.

Filaments 31-48 [jl in diameter, radiating
from the stipe to the margin, slightly parallel or
interdigitated, pluriseriate to triseriate, dichoto-
mously branched, supra-dichotomal constriction
uneven. The filaments possess numerous uni-
lateral or bilateral, short or pedicellate or trun-
cate appendages that are simple or lobed. The
appendages are so fitted together as to form a
primitive cortex. The filaments of stipes are ir-
regularly dichotomously branched, having lateral
appendages (Fig. la-e) . Reproductive organs
formed terminally on the filament (Fig. Id, e),
each with a thick constriction in the middle, up
to 16.2 mm long and 2.4 mm broad.

The present effort is the first study made of
reproduction in Udotea . The previous report on
the presence of reproductive organs was con-
sidered erroneous by Ernst (1904) and Fritsch
(1956), who supposed the structures reported
as reproductive to be epiphytes.

The filaments are compact and interdigitated;
but when fruiting occurs the terminal parts swell
and separate, and cytoplasm passes into the swol-
len parts, forming reproductive organs. The
lower part of each serves as a stalk of the repro-
ductive organ (Fig. le). The terminal portion
of the fertile frond becomes loose, and as the
filaments separate the reproductive organs show
( Fig. 2 ) ; but in vegetative fronds the terminal
edge of a blade is compact.

Numerous zooids are produced and they are
liberated successively through an apical pore.
On liberation of the zooids the middle constric-

243

244

PACIFIC SCIENCE, Vol. XVII, April 1963

Fig. 1. Udotea indica . a, Lower part of a filament; b , middle part of a filament; c, upper part of a filament;
d, upper part of a filament, showing reproductive organ; e, upper part of a filament, showing mature reproductive
organ.

Udotea indica— Nizamudmn

245

tion .relaxes, increasing in diameter, and thus
making way for the contents from the basal part
to stream into the terminal part of the repro-
ductive organ. This process is continued until
all the zooids are extruded. These zooids seem to
be morphologically similar. Fusion between
them has not been observed. Their study is still
in progress.

In C odium and Hdimeda special lateral game-
tan gia are formed, whereas in Udotea indica
terminal reproductive organs are formed. Such
terminal club-shaped sporangium formation has
been observed (Howe, 1907) on blades of Av-
rainvillea . This transformation of the terminal
parts of the filaments into reproductive organs
clearly distinguishes these genera from other
members of the family Codiaceae.

The filaments of Udotea indica resemble those
of C odium and Halimeda in their siphonaceous
and dichotomous character, but differ in possess-
ing uni- or bi-lateral appendages. The filaments
in Avrdnvillea are moniliform in appearance,
due to the numerous constrictions, but whereas

Fig. 2. Fertile plant.

appendages are found in Udotea there are none
in Avrainvillea .

Udotea certainly represents a distinctive fam-
ily, which I here name the Udoteaceae, but the
systematic position of this family must await
more complete knowledge of its members’ life
histories.

Udoteaceae fam. nov .

Frond fiabeilate, orbicular, broadly prolifer-
ated above. The base cuneate, distinctly zonate,
margin entire, lobed, or lacerated. Filaments ra-
diating from stipe to the margin, slightly parallel
or interdigitated, pluriseriate to triseriate, di-
chotomously branched, supra-dichotomal con-
striction uneven, uni- or bi-lateral short pedi-
cellate or truncate appendages simple or lobed.
Appendages so fitted together as to form a
primitive cortex. Terminally formed reproduc-
tive organ with a thick constriction.

ACKNOWLEDGMENT

The helpful advice, encouragement, and criti-
cism of Dr. Maxwell S. Doty, Professor of Bot-
any, University of Hawaii, Honolulu, Hawaii,
is gratefully acknowledged.

REFERENCES

Boergesen, F. 1930- Some Indian green and
brown algae especially from the shores of the
Presidency of Bombay. J. Ind Bot. Soc. 9:161.

— - 1934. Some marine algae from the

northern part of the Arabian Sea. Kgl. Dan.
Vidensk. Selsk. Biol. Meddel. 9(6) :20.
Ernst, A. 1904. Beitrage zur Kenntnis der Co-
diaceen. Beih. Bot. Centralbl. 16: 199 et seq.
FRITSCH, F. E. 1956. The structure and repro-
duction of the algae. Cambridge Univ. Press.

2:1-791.

Gepp, A. & E, S. 1911. The Codiaceae of Siboga
Expedition. Siboga-Exped. Monogr. 62:1-
150.

Howe, M. A. 1907. Further notes on Halimeda
and Avrainvillea. Bull. Torrey Bot. Club.
34:491-516.

Taylor, W. R. 1950. Plants of Bikini and other
northern Marshall Islands. Univ. Michigan
Sci. Ser. 18:1-227.

Effects of Pollution on the Amino Acid Content
of Mytilus edulis

Rita D. Schafer1

ABSTRACT: In an attempt to determine the influence of polluted water on the
amino acid content of Mytilus edulis , muscle tissue of specimens from a clean area
was compared with that of specimens from polluted areas and with that of sped-
mens transferred from clean to polluted water. Analyses were made by means of
two-dimensional paper chromatography. The amino acids present under one type
of environmental condition but not under another were cystine, cysteine, methionine,
taurine/asparagine, and proline.

It has long been known that marine waters
containing pollutants affect in various ways the
fauna inhabiting 'such waters. A number of
studies have shown that pollution factors aid the
settlement and growth of some species while
inhibiting others. Wilhelmi (1916) points out
that some species are so favored by such an area
that they occur in numbers sufficient to permit
them to be considered as pollutant indicators.
Reish (1955, 1956, 1957^ 1957 b) calls atten-
tion to the presence of Capitella capitata in all
areas of pollution in southern California; and in
a later study ( I960) , in which he classifies areas
as to degree of pollution, he points out that
each type of area is characterized by the presence
of dominant species of polychaetes. Hartman
(1950) states that near a disposal outfall in
Santa Monica Bay, California, only two major
groups of animals are present, the annelids and
the crustaceans. Filice (1959) in discussing the
distribution of bottom fauna in polluted estua-
rine waters, states that the occurrence of some
species as well as their concentrations will vary
with the nature and degree of pollution. Blevgad
(1932), investigating effects of pollution on
bottom fauna, found "dead” areas at the point
of sewage outfalls. He points out, however, that
at a distance slightly over 100 m away, con-
ditions are the same as at a greater distance.

1 Allan Hancock Foundation, University of Southern
California, and Immaculate Heart College, Los Angeles,
California. Manuscript received December 22, 1961.

McNulty (1957), in a study of the effects of
pollution in Biscayne Bay adjacent to the Miami
shoreline, indicates that pollution in some in-
stances has had a beneficial effect on productivity
of macropfganisms. Hartman (I960), summa-
rizing the results of extensive selected sampling
of 11 southern California coastal areas, states
that productivity and variation of organisms
differ according to types of sediments, depth of
bottom, availability of food, and effects of waste
discharges. She also noted that biomass values are
lowest in shallow depths of Santa Monica Bay,
an area into which the largest of the Los Angeles
disposal plants empties. The biomass values of
this region were comparable to those from oily
samples taken in a more northern area. In addi-
tion to these variations in ability to survive and,
in some cases, to thrive in waters containing
pollutants, it has been noted that the biochem-
ical composition of some forms living in such
an environment differs from that of the same
species from clean waters (Schafer, 1961).
Forms previously used for investigation were
the abalone, Hdiotis cracherodii , and the crab,
Pachygrapsm cmssipes. In these the free amino
adds present In specimens from polluted areas
were compared with those from nonpolluted
areas.

It is the purpose of the present study not
only to evaluate the differences in amino adds
in another organism from clean and polluted
areas, but to investigate as well the effect upon

246

Pollution on Mytilus edulis — Schafer

the free amino acids of transfer from a clean
area to a polluted one.

ACKNOWLEDGMENTS

The author is indebted to the administration
of the Allan Hancock Foundation for the use
of laboratory facilities. This study was supported
by a research grant from the United States Pub-
lic Health Service of the National Institute of
Health, no. RG-491L

MATERIALS AND METHODS

In the present study the wharf mussel, Mytilus
edulis , was selected for investigation. Its wide
occurrence made it easily available from a vari-
ety of areas of varying environmental condi-
tions. Because of its sessile nature and presence
on floats, it was possible to use specimens which
had been constantly subjected to the conditions
prevailing in the area from which it was taken.
No attempt was made to differentiate sexes or
size of specimens, as preliminary tests showed
no amino acid differences in these respects in
this species.

Comparisons of free amino acids were made
on muscle tissue of animals from ( 1 ) different
polluted areas, (2) an area considered free of
pollution, and (3) from animals transferred
from a nonpolluted area to areas of pollution.
In addition, muscle tissue hydrolysates of speci-
mens from the clean area were compared with
hydrolysates of individuals from the polluted
areas.

The sites chosen for study were a small yacht
marina in the Los Angeles Harbor, designated
here and in previous studies of water pollution
as LA. 7; a dock in slip I of the main channel,
identified as LA. 39; and a third area of pollu-
tion in Cerritos Channel of Long Beach Harbor,
LB. 23 (Fig. 1).

Specimens from a nonpolluted area were
taken at Tomales Bay, north of San Francisco.

To establish the normal free amino acid pat-
tern of M. edulis , specimens were taken from
the Tomales Bay area, returned to the laboratory
alive where muscle tissue was excised, quick
frozen with dry ice, lyophilized, and extracted
with 70'% cold ethanol. Hydrolysates were pre-
pared by hydrolysis of excised muscle with 6 N

247

Fig. 1. Areas of pollution from which collections
were made.

HC1 and desalted with 70% cold ethanol. Best
results were obtained by serial studies using dif-
ferent amounts of a given extract. By limiting
the amount of extract on one paper to a minimal
quantity necessary for resolution, those amino
acids ordinarily present in quantities so large
that adjacent spots tended to merge upon de-
velopment remained distinct and compact. By
placing large amounts of extract on a second
paper those amino acids present in trace quanti-
ties could be detected upon development, while
those present in large amounts, already deter-
mined in the first run, merged and became in-
distinguishable.

To determine whether or not differences in
free amino acid metabolism existed in M. edulis
under conditions of pollution, specimens were
collected from polluted areas mentioned above,
and treated in an identical manner.

Assuming that a difference might exist be-
tween these two groups with respect to free
amino acids, as it had in other forms, an attempt
was made to learn whether a change would occur
following transfer from the nonpolluted to the
polluted areas. Specimens from Tomales Bay
were tagged for identification, placed in plastic
containers through which a free exchange of
water was possible, and placed in the polluted

248

PACIFIC SCIENCE, VoL XVII, April 1963

TABLE 1

Free and Protein-bound Amino Acids Present in Specimens of Mytilus edulis
From Three Polluted Areas, One Nonpolluted Area, and Transferred Specimens

ALANINE |

GLYCINE

GLUTAMIC ACID

SERINE

ASPARTIC ACID

THREONINE

LYSINE

HISTIDINE

ARGININE

valine/phenylalanine

LEUCINES

CYSTINE

CYSTEINE

PROLINE

TYROSINE

METHIONINE

asparagine/taurine

UNKNOWNS

Free Amino Acids

Polluted areas

L. A. 7

L. A. 39

L. B. 23

Free Amino Acids

Nonpolluted area

Tomales Bay

Free Amino Acids

Transferred

To L. A. 39

To L. B. 23

Hydrolysates

L. A. 7

L. A. 39

’x~

L. B. 23

Tomales Bay

areas. After 10 days these specimens were re-
moved and analyzed for free amino acid content
in the manner described above. The terminal
survival point for transferred animals occurred
between 10 and 15 days.

RESULTS

Chromatograms made of the free amino acid
extracts of specimens from clean water showed
the presence of alanine, glycine, glutamic acid,
serine, aspartic acid, threonine, lysine, histidine,
arginine, valine, leucines, methionine, proline,
plus three unidentified spots.

Native specimens from L.A. 39, L.A. 7, and
L.B. 23, all polluted areas, contained alanine,
glycine, glutamic acid, serine, aspartic acid, thre-

onine, lysine, histidine, arginine, valine, leucines,
proline, cystine, and methionine, plus two un-
identified spots in the case of L.A. 7 and three
each on the chromatograms of L.A. 39 and L.B.
23. (See Table 1.)

The specimens taken from Tomales Bay and
transferred to the polluted waters of L.A. 39
and L.B. 23 contained well-defined spots of
alanine, glycine, glutamic acid, serine, aspartic
acid, threonine, lysine, histidine, arginine, valine,
leucines, and taurine/asparagine.

The hydrolysate of specimens from Tomales
Bay contained alanine, glycine, glutamic acid,
serine, aspartic acid, threonine, lysine, histidine,
arginine, valine, methionine, leucines, cystine,
and proline.

Pollution on Mytilus edulis — SCHAFER

249

Hydrolysates of specimens from LA. 7, L A.
39, and L.B. 23 contained those amino adds
reported above for the Tomales Bay specimens,
plus tyrosine and cysteine. The spot indicative
of valine assumed a shape which suggests that
phenylalanine may also be present. In the case
of L.A. 7 three unidentified amino adds were
present also; in the case of L.B. 23, two.

DISCUSSION

An analysis of the free amino acids present
in specimens from the three types of situation
indicated above shows that 1 1 of these occurred
in all forms. Those specimens taken from pol-
luted water contained proline, cystine, and
methionine as well. Those from nonpolluted
water contained the proline and methionine but
lacked the cystine. The chromatograms of those
specimens which had been transferred from
clean to polluted waters lacked all three of these
free amino acids, proline, methionine, and cys-
tine. Taurine/asparagine, not found in any of
the others, either from dean or polluted water,
is dearly discernible on the chromatograms of
the transferred specimens.

The nature of the pollutants involved in con-
tributing to these differences has not been spe-
cifically determined. The data compiled by the
Los Angeles Bureau of Sanitation (1957), cover-
ing the periods August 1956 to February 1957,
show oxygen depletion and increased coliform
bacterial counts for the areas covered in this
study. Reish (I960) summarizing characteristics
of the areas classifies L.A. 7 and L.B. 23 as
"healthy bottom” areas and lists a dissolved oxy-
gen at 6.0 ppm for the median at the surface.
He classifies L.A. 39 as "very polluted bottom”
with a dissolved oxygen of 1.6 ppm. In describ-
ing the nature of the substrate he indicated the
presence of black sulfide mud in each instance.
A visually obvious pollutant at all three stations
is an oily substance. The concentration of this
at L.A. 39' is such that all submerged structures
appear to have a tarlike coating. Water circu-
lation at this point seems to be limited to that
produced by tidal fluctuation.

Whatever the composition of the pollutants,
they seem to contain factors which have in-
fluenced the metabolism of the sulfur-containing
amino acids, cystine and methionine. It is known

that methionine can be converted to cystine, and
that cystine and methionine, via cysteine, in
mammals can act as a precursor of taurine. Cys-
tine is present as a free amino acid only in those
specimens from polluted water. It is protein-
bound in M. edulis from both clean and polluted
water. Methionine occurs in free form in all but
the transferred specimens. It is also found as
part of the proteins from M. edulis of both dean
and polluted water. Cysteine appears only in the
protein-bound form in those animals analyzed
from polluted water.

If the spot that appeared in the position of
taurine/asparagine on the chromatogram, is as-
sumed to be, at least in part, taurine, then this
amino add appears in the free form in only
those specimens transferred.

If the specimens taken from dean water can
be considered as those with a normal metabolic
pattern, then It can be safely postulated that
the metabolism of methionine has not been al-
tered in any except the transferred specimens,
where it does not appear in free form. The
presence of cystine as a free amino add in indi-
viduals from polluted water suggests that some
environmental factor has freed a portion of it
from proteins or that an amount in excess of
that utilized in protein formation has been
synthesized. The possible appearance of free
taurine in the transferred specimens and the
total absence of free methionine suggests that
some factor in the change of environment may
have brought about the conversion of the latter
to taurine.

The disappearance of proline In those speci-
mens of M. edulis transferred from a dean to
a polluted environment is unexplainable at
present.

SUMMARY

In an attempt to determine the effects of
water pollutants upon the wharf mussel, Mytilus
edulis, the following analyses were made.

1. Specimens were collected from polluted
waters. From these the free amino acid content
of muscle tissue was analyzed by means of two-
dimensional paper chromatography. These were
found to contain 14 identifiable free amino
adds plus some (two in one instance, three in

250

PACIFIC SCIENCE, Vol. XVII, April 1963

another) unidentifiable ninhydrin-sensitive
spots.

2. Specimens from polluted waters were ana-
lyzed, after hydrolysis, for amino acid content of
muscle tissue. Sixteen amino acids were identi-
fied. Unidentifiable spots also occurred in speci-
mens from two of the three locations.

3. Specimens from nonpolluted water were
treated in the same way. Thirteen free amino
acids were identified. Cystine, present in free
form from polluted water was lacking. Three
unidentifiable spots were also present. Fourteen
amino acids were identified from the protein
hydrolysate. Cysteine and tyrosine, found in hy-
drolysates of polluted water specimens, were
lacking here.

4. Specimens were transferred from clean
water to polluted environments, left for 10 days,
and then analyzed in the usual manner. Chro-
matograms showed them to have 11 free amino
acids in common with the others. Proline and
methionine contained in the others were absent
here. Cystine, present in those native to the
polluted areas, was lacking. Taurine/ asparagine,
not found in any of the others, appeared here.

REFERENCES

Blevgad, H. 1932. Investigations of the bottom
fauna at outfalls of drains in the Sound. Co-
penhagen Rept. Danish Biol. Sta. 37:1-20.

Filice, Francis P. 1959. The effects of wastes
on the distribution of bottom invertebrates
in the San Francisco Bay Estuary. Wasmann
J. Biol. 17(1): 1-17.

Hartman, Olga. 1956. Results on Investiga-
tions of Pollution and Its Effects on Benthonic
Populations in Santa Monica Bay, California.
Allan Hancock Foundation, Univ. So. Calif.
Press. Pp. 1-23.

1959. The benthonic fauna of southern

California in shallow depths and possible ef-
fects of wastes on the marine biota. Waste
Disposal in the Marine Environment. Per-

gamon Press. Pp. 57-81.

McNulty, J. K. 1957. Pollution studies in
Biscayne Bay during 1956. Progress Rept.
Fed. Sec. Agency, PHS, NIH, Feb.: 1-17.

Reish, D. J. 1955. The relation of polychaetous
annelids to harbor pollution. Public Health
Rept. 70:1168-1174.

1956. An ecological study of Lower San

Gabriel River, California, with special ref-
erence to pollution. Calif. Fish and Game
42(1) : 51-61.

1957^. The effect of pollution on marine

life. Industrial Wastes 2:114-118.

195 7 A The relationship of the poly-
chaetous annelid Capitella capitata (Fabricius)
to waste discharges of biological origin. Biol,
of Water Pollution. U. S. Public Health Serv-
ice, Cincinnati. Pp. 195-200.

I960. The use of marine invertebrates

as indicators of water quality. Waste Disposal
in the Marine Environment. Pp. 92-103.

Schafer, R. D. 1961. Effects of pollution on
the free amino acid content of two marine
invertebrates. Pacif. Sci. 15(1) : 49—55.

Wilhelmi, J. 1916. Ubersicht liber die bio-
logische Beurteilung des Wassers. Ges. Naturf.
Freunde Berlin, Sitzber.:297 -306.

Important Pacific Insect Pests of Sugar Cane1

C. E. Pemberton2

SUGAR cane is commercially grown in Hawaii,
Fiji, Queensland, Java, the Philippines, and For-
mosa. From extensive published records, some
257 insect species are known to attack cane
within these several areas. The following sum-

mary will indicate only those species which have
been reported as important, either occasionally
or continuously, together with the nature of
attack and the countries affected.

LBPIDOPTERA

Eucosmidae

Pyralidae

Agrotidae

STALK BORERS

Eucosma schistaceana Sn.
Chilotraea infuscatella Sn.
Proceras venosatus Wlk.
Scirpopkaga nivella F.
Emmalocera umbricostella Rag.
Sesamia inferens Wlk.

Sesamia uniformis Dudg.
Pbragmatiphila tmncata Wlk.

Formosa, Java, Philippines, Guam
Formosa, Java, Philippines
Formosa, Java, Philippines
Formosa, Java, Philippines
Formosa

Formosa, Java, Philippines
Philippines

Queensland, New Guinea

COLEOPTERA

Curculionidae Rkahdoscelus obscurus Boisd.

Trochorrbopalus strangulatus Gyll.
Prionidae Dorystbenes hydropicus Pascoe

Hawaii, Fiji, Queensland, New Guinea
Philippines, Fiji, New Guinea
Formosa

ISOPTERA

Heterotermes philippinens Light
Mastotermes darwiniensis Frogg
Termes meredionalis Frogg
Macrotermes gilvus Hag,
Capritermes nitobei Shir.
Odontotermes formosanus Shir.
Reticulitermes speratus Kolbe

Philippines

Queensland

Philippines, Java

Formosa

COLEOPTERA

Elateridae

Tenebrionidae

DESTRUCTIVE TO SEED PIECES

Lacon mus cuius Cand.
Mel&notus tamsuyensis Bates
Sephilus formosanus Schwarz
Lacon variabilis Cand.
Eutocbia lateralis Boh.

Formosa

Queensland

Java, Hawaii, Philippines

HEMIPTERA

Lygaeidae

SAP SUCKERS ON LEAF OR STALK

Pbenacantha australica Kirk.
Phenacantha saccharicida Karsch
Pbenacantha marcida Horv.
Ischnodemus saccharivoms Okajima

Queensland

Java

Formosa

1 A paper presented at the Tenth Pacific Science Congress, in Honolulu, August 1962. Manuscript received
February 14, 1962.

2 Hawaiian Sugar Planters’ Association Experiment Station, Honolulu, Hawaii.

251

252

PACIFIC SCIENCE, VoL XVII, April 1963

Coccidae

Aphididae

Aleurodidae

Derbidae

Delphacidac

ORTHOPTERA

Gryllotalpidae

COLEOPTERA

Curculionidae

Melolonthidae

Rutelidae

Dynastidae

ORTHOPTERA

Acrididae

LEPIDOPTERA

Pyralidae

Agrotidae

Aulacaspis tegalensis Zhnt.
Aulacaspis madiunensis Zhnt.
Saccharicoccus sacchari Ckll.
Oregma lanigera Zhnt.
Rhopalosiphum maidis Fitch
Aphis sacchari Zhnt.
Neomaskellia hergii Sign.

Proutista moesta Westw.
Perkinsiella saccharicida Kirk.
Perkinsiella vitiensis Kirk.
Perkinsiella vastatrix Bredd.

Formosa, Java, Philippines
Queensland, Java

Queensland, Java, Formosa, Philippines, Hawaii, Fiji
Java, Formosa, Philippines, New Guinea
Hawaii, Philippines, Java, Queensland, Formosa, Fiji
Hawaii, Philippines, Java, Queensland, Formosa
Formosa, Philippines, Java, Queensland, Fiji,

New Guinea

Philippines, Formosa, Java, Guam
Queensland, Formosa, Java, Hawaii
Fiji, Samoa

Formosa, Java, Philippines

DESTRUCTIVE TO ROOTS AND UNDERGROUND PARTS

Gryllotalpa formosana Shiraki

Formosa

Tanymecus circumdatus Wied.

Formosa

Episomoides alhinus Mats.
Ancylonycha horishana Niijima and

Formosa

Kinoshita

Formosa

Ancylonycha vidua Sharp

Philippines, Java

Ancylonycha helleri Brenske

Java

Ancylonycha leucophthalma Wied.

Java

Lepidiota stigma Fabr.

Java

Lepidiota trichosterna Lea

Queensland

Lepidiota frenchi Blackb.

Queensland

Leucopholis rorida Fabr.

Java

Leucopholis irrorata Chevr.

Philippines

Dermolepida alhohirtum Waterh.

Queensland

Apogonia destructor H. Bos

Java

Rhopaea vestita Arrow

Fiji

Pseudoholophylla furfuracea Burm.

Queensland

Anomala limhifera Ohaus

Formosa

Anomala humeralis Burm.

Philippines

Anomala annoguttata Burm.

Philippines

Anomala orient alls Waterh.

Hawaii

Euchlora pulchripes Lansb.

Java

Euchlora viridis Fabr.

Java

Anoplognathus hoisduvali Boisd.

Queensland

Alissonotum impressicole Arrow

Formosa

Alissonotum crassum Arrow

Formosa

Alissonotum pauper Burm.

Formosa, Philippines

LEAF DEFOLIATORS

Oxya chinensis Thunb.

Locusta migratoria migratorioides
Rch. & Fairm.

Locusta migratoria manilensis Meyen
Locusta cinerascens F.

Locusta migratoria solitaria F.
Hieroglyphus annulicornis Shiraki
Gastrimargus musicus F.

Hawaii

Philippines, Queensland
Philippines, Formosa
Philippines

Formosa, Queensland, Fiji, Guam

Formosa

Queensland

Hedylepta accepta ButL
Pseudaletia unipuncta Haw.
Spodoptera exempta Wlk.
Cirphis loreyi Dup.

Hawaii

Philippines, Hawaii, Fiji, Queensland, Formosa, Java
Philippines, Queensland, Hawaii
Philippines, Java, Fiji, Queensland, Formosa

Introduction and Summary. It is desirable to state the
purpose and scope of the paper in an introductory
paragraph and to give a summary of results at the end
of the paper.

Batzo, Roderick L., and J. K. Ripkin. 1849. A
Treatise on Pacific Gastropods. Rice and Shipley,
Boston, vii + 326 pp., 8 figs., 1 map.

Crawford, David L. 1920*. New or interesting
Psyllidae of the Pacific Coast (Homop.). Proc.
Hawaii. Ent. Soc. 4(1) : 12-14.

1920&. The sandalwoods of Hawaii. Proc.

Hawaii. Ent. Soc. 4(2) : 374-375, 13 pis.

In the text, sources should be referred to by author,
date, and page, as follows: "It was noted (Rock,
1916: 18) that . . .” or "Rock (1916: 21-24)
says . .

ILLUSTRATIVE MATTER

Artwork for illustrations should be 8V2 x 11 inches
or smaller, and it should accompany manuscript, on
separate sheets. Often more valuable than a photo-
graph is a good line drawing.

It is strongly urged that an indication of scale be
incorporated as a part of all drawings in which mag-
nification and size are critical considerations.

Illustrations will be returned to the author.

Tables. Tabular matter should be kept to a minimum.
Each table, prepared to conform with Pacific Science
style, should be typed on a separate page, and its posi-
tion indicated on the manuscript.

Mathematical Formulas. Complicated formulas cannot
be set by the printers. Authors should submit them
as illustrations.

Captions. Readily identifiable captions for figures,
graphs, photographs, and other illustrative matter
should be supplied on a separate page.

PROOF

All queries on proof should be answered. If cor-
rected proof is not received within four weeks after
being sent to the author, author’s changes cannot be
accepted.

REPRINTS

Reprints or separates should be ordered on the
form provided and returned with author’s proof. All
correspondence concerning separates must be directed
to the printer, Star-Bulletin Printing Company, 420
Ward Avenue, Honolulu 14, Hawaii.

- '■

5o S. f

YOL XVII p ^!j JULY 1963 NO. 3

PACIFIC SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

K. O. EMERY

Aerial Study of Wave Patterns
MATHILDE SCHWABL

Solenogaster Mollusks from Southern California

SHERWIN CARLQUIST and MARTIN L. GRANT
Studies in Fitchia

JACK T. TOMLINSON

New Burrowing Barnacle from Hawaii

EDGAR J. MARTIN

Toxicity of California Anemones

KELSHAW BONHAM and EDWARD E. HELD
Sea Cucumbers at Rongelap Atoll

DARL E. BOWERS

Field Identification of Californian Beach Hoppers

SIDNEY C. HSIAO, WALTER K. FUJII, and
HELEN H. FINE

Device for Making Successive Photomicrographic

Records

HAROLD ST. JOHN

Revision of the Genus Pandanus

Part 15. Species Discovered in Thailand

S. J. PARAMONOV

Lord Howe Island, Part III

UNIVERSITY OF HAWAII PRESS

BOARD OF EDITORS

O. A. BUSHNELL, Editor-in-Chief
Department of Microbiology, University of Hawaii

Robert Sparks, Assistant to the Editors
Office of Publications and Information, University of Hawaii

L. H. Briggs
Department of Chemistry
University of Auckland
Auckland, New Zealand

John J. Naughton
Department of Chemistry
University of Hawaii

Maxwell S. Doty
Department of Botany
University of Hawaii

Martin Sherman
Department of Entomology
University of Hawaii

Ai Kim Kiang
Department of Chemistry
University of Malaya, Singapore

Walter R. Steiger
Department of Physics
University of Hawaii

Gordon A. Macdonald
Department of Geology
University of Hawaii

Donald W. Strasburg
Bureau of Commercial Fisheries, Hawaii Area
(U. S. Fish and Wildlife Service)
Honolulu, Hawaii

Sidney J. Townsley
Department of Zoology
University of Hawaii

Thomas Nickerson, Managing Editor
Assistant to the University Provost

Donald C. Matthews
Department of Zoology
University of Hawaii

INFORMATION FOR AUTHORS

PREPARATION OF MANUSCRIPT

( Continued on inside back cover )

PACIFIC SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

VOL. XVII JULY 1963 NO. 3

Previous issue published April 29, 1963

CONTENTS

PAGE

An Aerial Study of Hawaiian Wave Patterns. K. O. Emery .......................... 255

Solenogaster Mollusks from Southern California . Mathilde Schwahl 26 1

Studies in Fitchia ( Compositae ): Novelties from the Society Islands;

Anatomical Studies. Sherwin Carlquist and Martin L. Grant.................... 282

Lithoglyptes hirsutus ( Cirripedia: Acrothoracica) , A New Burrowing Barnacle

from Hawaii. Jack T. Tomlinson . 299

Toxicity of Dialyzed Extracts of Some California Anemones ( Coelenterata ).

Edgar J. Martin................... . 302

Ecological Observations on the Sea Cucumbers Holothuria atra and H. leucospilota
at Rongelap Atoll , Marshall Islands.

Kelshaw Bonham and Edward E. Held 305

Field Identification of Five Species of Californian Beach Hoppers

( Crustacea : Amphipoda) . Dari E. Bowers 315

A Simple Device for Making Successive Photomicro graphic Records of
Large Groups of Developing Organisms.

Sidney C. Hsiao, Walter K. Fujii, and Helen H. Fine................... *.321

Revision of the Genus Pandanus Stickman, Part 15. Malayan Species Described

by H. N. Ridley. Harold St. John............................ 329

Lord Howe Island, A Riddle of the Pacific, Part III. S. J . Paramonov. ........... 361

notes:

Additional Plants from the Midway Islands.............. 374

On Malayan Shores: A Review 374

stmpwi

lOTTilW

OCT 1 1 1983

■')

An Aerial Study of Hawaiian Wave Patterns

K. O Emery1

Most of us who have had some experience at
sea on small power or sailing boats have noted
that the sea surface is crossed by usually more
than a single train or set of waves. Each train
can be identified by its direction, period, and
height. Sometimes one train is so dominant that
others may not be noticed. The Marshall Is-
landers of the nineteenth century and earlier,
past masters in the art of handling small boats,
used wave trains as a navigational aid with their
famous stick charts. The dominant train of
waves, produced by the trade winds, was desig-
nated by long parallel sticks tied to a rigid
frame. Trains of smaller waves, some produced
by bending around islands, were shown by small
sticks attached at angles to the long ones. A brief
review of these maps and of pertinent literature
was given by Emery, Tracey, and Ladd ( 1954: 5).

A better platform than a boat for mapping
waves is an airplane because of its wider field
of view and greater speed. The greater speed
permits observations to be made at many points
in an area during a period short enough to re-
duce the effect of changes in winds. The first
such wave maps made by aerial observation were
for the sea off southern California in 1957-58
(Emery, 1958). Each of four surveys at 3-month
intervals exhibited three or four trains of waves,
greatly complicating the identification of waves
refracted or reflected by the island or mainland
shores. In order to simplify the analysis it is
necessary to study an area having only a single
wave train. The Hawaiian Islands were thought
to constitute such an area because they lie
within the belt of trade winds, especially during
summer. An opportunity for study there was
provided by the convening of the Tenth Pacific
Science Congress at Honolulu during the latter
half of August, 1961.

1 University of Southern California, Los Angeles 7,
California. Now: Woods Hole Oceanographic Insti-
tution, Woods Hole, Mass.

Manuscript received January 10, 1962.

Through the interest of the Office of Naval
Research, flight facilities were made available by
Fleet Air Wing Two, Naval Air Station, Barbers
Point, Oahu. The aircraft was a Neptune (P2V),
having a bombardier station with excellent visi-
bility. Acting as recorders and photographers
during one flight were Charles G. Johnson (U.S.
Geological Survey) and Charles Roberts (U.S.
Weather Bureau); and during the other flight
Robert S. Dietz (U.S. Navy Electronics Labora-
tory) and Joe S. Creager (University of Wash-
ington). Appreciation is also due Doak C Cox
(University of Hawaii) and Saul Price (U.S.
Weather Bureau, Honolulu) for their critical
reading of the manuscript.

METHODS

Flights were made on 31 August (1000-1700,
local time) and 1 September (0800-1430). They
covered a total distance of 4200 km at an alti-
tude of 500 m. A total of 375 observation points
were occupied at 2 -min intervals. Positions of
the aircraft were based upon fixes at 10-20~min
intervals using loran, radar, and visual methods.

Observation was purely visual, although some
photographs were taken for illustrative purposes.
A specially devised peloras was held with the
aircrafts true compass heading set on the disk
in a fore-and-aft direction. Moveable arms piv-
oted at the center of the disk were set parallel
to the crests of separate wave trains, requiring
constant visual checking and readjustment. At
2 -min intervals the recorder noted the azimuth
indicated by each arm. Comparison of results
obtained at different headings of the aircraft
indicate that the azimuths are correct within
10°. Estimation of the azimuth of the crests of
each wave train usually required viewing in a
particular direction because the waves were most
dearly seen when viewed away from the sun at
a horizontal angle of about 45° to the wave
crests and at a downward angle of 15 to 45°.
Sometimes about 10 sec of viewing was required

255

256

PACIFIC SCIENCE, Vol. XVII, July 1963

to identify the trains of smaller waves or waves
of similar azimuths. This time requirement and
the different optimum directions of viewing
reduces the value of interpretation from photo-
graphs alone; in addition, photographs them-
selves must be interpreted later — a more diffi-
cult process than direct visual interpretation of
the wave azimuths. Direct visual observation
also permitted the determination of the direc-
tion of wave progress, relative to the positions
of foam patches. Timing the interval between
the appearance of a foam patch at the crests of
successive waves provided an estimate of wave
periods.

In addition to wave data, observations on
wind direction and speed at the water surface
were obtained. The wind direction was taken
as the azimuth of the long narrow parallel wind
slicks which appeared where wind speed ex-
ceeded Beaufort 3. Wind speed was estimated
according to the Beaufort scale of water surface
characteristics (Bowditch, 1943:52), as shown
below.

BEAUFORT

NUMBER

DESCRIPTION

OF SEA

SPEED

(km/hr)

mirrorlike

0-2

wavelets

2-6

short waves

6-12

scattered whitecaps

12-19

many whitecaps

19-30

foam streaks

30-40

The author does not apologize for not using
a more modern and sophisticated technique
involving elaborate camera arrangements or
electronic scanning devices, because direct ob-
servation is simpler and sufficient for the prob-
lem.

WINDS

Weather maps for the period, based upon
ship and shore stations as well as upon TIROS
photographs, are illustrated by Figure 1. The
two high pressure areas north and northeast of
Hawaii are characteristic of the season. A low
pressure area about 800 km southeast of Hawaii
discovered by TIROS satellite was mild and its
effect was not detected during the flights. The
isobars show that the wind in the vicinity of
the Hawaiian Islands should have been from

about 070° (east-northeast); this is typical of
the trade-wind period of the year.

A far more detailed picture of the wind pat-
tern was provided by aerial observation (Fig.
2). On the windward side of the islands the
wind speed was Beaufort 4 except adjacent to
the island of Hawaii, whose blocking effect re-
duced the wind to below Beaufort 2. On the
immediate leeward side of the islands the wind
was reduced to Beaufort 1 by shielding effects.
Between the islands the wind was sped by fun-
nelling, so that a speed of Beaufort 5 occurred
between Hawaii and Maui. A tongue of Beau-
fort 4+ extended from between Oahu and Mo-
lokai to a position off the south coast of Oahu.
Verification of its presence was obtained by
anemometer readings aboard R/V "Argo” of
Scripps Institution of Oceanography, which
worked in that area during 2 September. These
tongues of high wind speed expanded and joined
laterally well to the lee of Maui. However, still
farther to the southwest is an area of winds less
than Beaufort 3 having an unknown extent.
Possibly this area is the result of the combined
lees of all the islands; if so, it would be of in-
terest to learn how much farther southwestward
the area extends.

Wind direction was estimated from slicks
which were long (to 500 m), narrow (a few
meters), and occurred in parallel series (5-50
m apart ) parallel to the wind. These wind slicks
are zones of convergence between cells of surface
water caused to rotate around horizontal axes by
the wind (Woodcock, 1944). Oil (perhaps from
phytoplankton) collects in the convergences and
changes the light reflectivity and wave pattern,
as observed by Dietz LaFond (1950). Other
debris, including sugar mill wastes, were locally
present in some slicks. The wind slicks were re-
stricted to areas having wind speeds of Beaufort
3 or more, and are one of many phenomena
which are dependent upon this critical wind
speed (Munk, 1947). As shown by Figure 2 the
windflow lines, deduced from the wind slicks,
spread apart to pass around the islands, particu-
larly around large and high Hawaii. Their con-
centration in the straits causes the high wind
speeds there.

In areas having wind of less than Beaufort 3,
wind slicks were usually absent. In the lee of
Hawaii and Maui their place was taken by slicks

Hawaiian Wave Patterns— -Emery

257

Fig. 1. Weather map for 1600 Honolulu time (0600 G.C.T.) on 31 August 1961. The winds and isobars
are typical of the season for the Hawaiian Islands, except for the minor low to the southeast. Courtesy of
Charles Roberts of Honolulu office of U. S. Weather Bureau.

of another kind, ones which are commonly sev-
eral km long, 20-200 m wide, broadly sinuous,
and 1 or more km apart. Their general trend
happened to be more or less perpendicular to
the wind and parallel to shore. These slicks are
typical of convergences above shallow internal
waves (Ewing, 1950; LaFond, 1959). A third
kind of slick was noted behind a ship — straight,
parallel-sided, about 30 m wide, and oily brown.
It clearly resulted from ship wastes, probably
from pumping of bilges or fuel tanks.

WAVES

Instead of the anticipated single train of
waves, the area was found to contain three trains
of large long-period waves and several trains
of small short-period ones. Many measurements
of the former waves revealed an average of
7 -sec period, whereas the latter were 1 sec or
shorter. Most of the wave energy was confined

to the two trains of 7-sec waves coming from
the east (090°) and southeast (130°). The
train from the east was slightly sharper crested,
perpendicular to the wind slicks ( compare Figs.
2 and 3 ) , and most frequently topped by white-
caps. Interference between these two trains
caused the sea surface to be broken into a
diamond-shaped pattern, each block of which
contained a short section of wave crest.

Along the sides of the islands, wave refraction
caused the waves to bend and to approach paral-
lelism with shore, but the width of the zone
was generally too small to map on the scale of
the survey. At the straits the waves took a circu-
lar front, evidently due to diffraction (Arthur,
1951). The shapes of the wave fronts are similar
to those of the tsunamis studied by Shepard,
Macdonald, and Cox (1950) in the Hawaiian
region, but the latter are controlled more by
refraction than by diffraction owing to their
great wave length. Spreading of the circular

258

PACIFIC SCIENCE, Vol. XVII, July 1963

fronts beyond the straits partially filled the area
leeward of the islands. Locally, complex inter-
ference patterns resulted from crossing of wave
trains from different straits. In general, however,
the leeward sides of the islands had only low
waves of the 7-sec period. Southwest of Oahu
a third train of 7-sec waves was observed mov-
ing northward; it is of unknown origin.

The waves of about 1-sec period were too
short to have been produced by other than local
winds. However, their round crests and large
angle with the wind slicks indicate that these
waves were not being formed during the time
of the survey. Possibly the ones northeast of
the island chain and southeast of Hawaii re-
mained from a local wind which blew just prior
to the survey; these areas were surveyed only
during 31 August. The waves southwest of the
chain were more irregular in pattern and may
have owed their origin to cold air masses flowing
down the slopes of several islands.

EFFECTS ON SHORES

During a study of beachrock in the Hawaiian
Islands by Emery and Cox (1956) the shores
of most of the islands were examined in detail.
Others were viewed from the air during the

wave survey. These shores can be classified as
follows : rocky ( irregular and commonly rugged,
with sea cliffs and only narrow beaches or pocket
beaches), sandy (long, broad sand beaches), and
muddy (mangroves, narrow beaches or none,
locally with lagoons or artificial fish ponds).
Ignoring transitional forms, the shore types of
the islands are illustrated by Figure 4. A re-
lationship of the shore type to the chief wave
pattern, that from a generally easterly direction,
can readily be observed. High storm waves from
any direction (Arthur, 1948) also can control
the shore types, but infrequency of the storms
and inconstancy in direction of their waves
reduces their importance.

Rocky shores characterize the sides of the
islands most exposed to direct wave attack,
except at Kaneohe Bay (east side of Oahu)
where a wide offshore reef protects the shore.
In some areas, such as almost the entire circum-
ference of Hawaii, rocky shores dominate, re-
gardless of exposure direction, probably because
of the recency of vol can ism. Longshore currents
developed by a diagonal approach of waves to
the shore have caused much of the sediment
produced by wave attack of the rocky shores or
of adjoining reefs to be carried beyond the
areas of greatest exposure to waves. Some of

Fig. 2. Results of aerial observation of wind at sea surface on 31 August and 1 September 1961. Wind
flow lines (wide lines) are based upon wind slicks observed at each position (note weather-vane line at each
position). Wind speeds are based upon condition of sea surface expressed in Beaufort scale (indicated by

narrow numbered lines).

Hawaiian Wave Patterns — Emery

259

Fig. 3. Wave pattern based upon aerial observation at positions indicated by circles on 31 August and 1
September 1961. The wide lines show changing directions of crest lines of each of three different trains of
7- sec waves. The narrow lines show the same for several trains of short-period (about 1-sec) waves.

this material has been deposited to form sandy
shores chiefly in the immediate lee of prominent
rocky headlands where the intensity of wave
attack and of longshore currents is sharply re-
duced. Generally this lee is west of headlands,
but on Lanai it is on the east side because
that side is sheltered by nearby Maui. Muddy
shores are not common, but those which do
exist are in areas which have the greatest pro-
tection from waves, well into the lee of the
islands or of headlands.

In recent years there has been erosion of
certain sand beaches, particularly those on the
east side of Kauai, the north side of Oahu, and
the north side of Maui. In part, this erosion has
probably resulted from man’s activities. How-
ever, the close relationship between shore type
and degree of exposure exhibited by Figure 4
suggests another cause. As shown by Wentworth
(1949), there may have been a directional
change of trade winds at Honolulu from a mean
of 050° in 1908 to 085° between 1925 and
1939 to 063° in 1943. If the trend is real and
if it continued into recent years, the average
wind direction would again have been about
050° sometime between 1950 and I960. Al-
though the wind direction at Honolulu may not
correspond exactly with the direction of the
winds at sea because of the deflecting effects of
nearby mountains, the changes in wind direction

at Honolulu may well reflect changes at sea.
Such a change of wind direction at sea would
have produced a corresponding change of wave
approach. Possibly the beaches are being shifted
somewhat as weather vanes in response to this
change in direction of wave approach.

SUMMARY AND CONCLUSIONS

Mapping of surface winds and wave patterns
from the air is a simple procedure. A 2 -day
survey in the Hawaiian Islands showed that the
wind was from the east, but was subject to

Fig. 4. Relationship of type of shore to degree of
exposure to dominant waves. Radiating lines at upper
right-hand corner show common azimuths of wave
crests and directions of wave approach.

260

PACIFIC SCIENCE, Vol. XVII, July 1963

local deflection by the islands. It was probably
typical of the trade-wind period of the year.
May through September, when the winds blow
from the northeasterly quadrant 80-9596 of the
time. From October through April the trade
winds still blow from the same general direction
most (65-80%) of the time (Blumenstock,
1961). The wave pattern was more complex
than anticipated, with three trains of 7-sec waves
and several trains of about 1-sec period. Most of
the wave energy was in two trains coming from
the east and southeast, evidently produced by
the trade winds, and agreeing well with the
general observation of predominant waves from
the east in the Hawaiian region. Shore types of
the islands appear to be closely related to degree
of exposure to these waves. Rocky shores are
generally in the most exposed sectors, sandy
shores are in the immediate lee of prominent
headlands, and muddy shores are in the most
protected sectors. Recent changes of sandy shores
may be due to possible long-term changes in
the trade winds, with attendant changes in the
waves produced by them.

REFERENCES

Arthur, R. S. 1948. Forecasting Hawaiian swell
from January 2 to 5, 1947. Bull. Amer. Me-
teorological Soc. 29:395-400.

1951. The effect of islands on surface

waves. Bull. Scripps Inst. Oceanogr. 6: 1-26.

Blumenstock, D. I. 1961. Climate of Hawaii.
U.S. Dept, of Commerce, Weather Bur. Cli-
matography U.S. 60-51:1-20.

Bowditch, N. 1943. American practical navi-
gator. U.S. Navy Hydrogr. Off. 9:1-387.

Dietz, R. S., and E. C. LaFond. 1950. Natural
slicks on the ocean. J. Mar. Res. 9:69-76.

Emery, K. O. 1958. Wave patterns off southern
California. J. Mar. Res. 17:133-140.

and D. C. Cox. 1956. Beachrock in the

Hawaiian Islands. Pacif. Sci. 10:382-402.

, J. I. Tracey, Jr., and H. S. Ladd. 1954.

Geology of Bikini and nearby atolls. U.S.
Geol. Survey Prof. Pap. 2 60- A: 1-265.

Ewing, G. 1950. Slicks, surface films and in-
ternal waves. J. Mar. Res. 9: 161-187.

LaFond, E. C. 1959. Slicks and temperature
structure in the sea. U.S. Navy Electronics
Lab., San Diego Res. Rept. 937: 1-35.

Munk, W. H. 1947. A critical wind speed for
air-sea boundary processes. J. Mar. Res. 6:
69-76.

Shepard, F. P., G. A. Macdonald, and D. C.
Cox. 1950. The tsunami of April 1, 1956.
Bull. Scripps Inst. Oceanogr. 5:391-528.

Wentworth, G K. 1949. Directional shift of
trade winds at Honolulu. Pacif. Sci. 3:86-88.

Woodcock, A. H. 1944. A theory of surface
water motion deduced from the wind-induced
motion of the Physalia. J. Mar. Res. 5:
196-205.

Solenogaster Mollusks from Southern California

Mathilde Schwabl1

The solenogaster mollusks named below
were collected off southern California by the
research vessel "Velcro IV” of the University
of Southern California, Los Angeles, Califor-
nia, during the course of quantitative surveys
made along the shelf and slope marine bottoms
from Santa Barbara, California, to south of the
Mexican border. Depths of bottoms ranged
from shallow shelf (about 20 m) to deep off-
shore basins (about 1800 m). Most station
numbers referred to in the text have been pub-
lished in the Pacific Expedition series of the
Hancock Foundation; they can be consulted
in volumes 19 and 22. Data for additional sta-
tion numbers are given in the Appendix to
this report. The types of all new species and
duplicates after the first are deposited in the
collections of the Allan Hancock Foundation
at the University of Southern California; a
first set of duplicates is retained by the author.

Solenogasters have been previously recorded
from California, based on collections of the
USS "Albatross,” and described by Heath ( 1911
and 1918). They were taken during dredging
operations and the specimens can be only ap-
proximately located with respect to precise
place, depth, and kinds of sediments.

Figures 1 and 2 plot the distributions of
solenogasters taken from sea bottoms between
the mainland of California and Santa Catalina
Island. The dots locate the positions of inter-
sections; they mark the even-numbered min-
utes of latitude and longitude, and are about
2 nautical miles apart. This area is more com-
pletely detailed in Hartman (1955). A large
impoverished subsill area in the deepest part
of the Channel sustains almost no life. The
surrounding fringe and shelf in shallower
bottoms supports characteristic kinds of soleno-
gasters, as indicated by the symbols used in
the two charts.

1 Berlin Lichterfelde Kommandantenstrasse 18, Ger-
many. Manuscript received November 22, 1961.

The diagnoses of the species require a few
comments. The illustrations of the spines show
only typical kinds, as these structures show
strong individual variations. Nevertheless cer-
tain features, such as comparative size, being
strongly keeled or not, and others, remain con-
stant throughout the species. The small letters,
a. b, Cj cl, used in the figures, refer to the body
regions from which the spicules were taken
and can be ascertained by reference to the en-
tire figure of Crystallophrisson recisum n. sp.

I have come to the conviction that the so-
called subradular organ of the Crystallophris-
sonidae is primarily the papilla bearing the
outlets of a third group of salivary glands,
which up to now has escaped the attention
of other authors. It is not a sensory organ. A
small number of gland follicles is situated ven-
tral !y of the fore-gut and just in front of the
radula. The cells of these follicles resemble
those of the lateral salivary glands, which empty
their secretion through more or less well-defined
ducts. An unpaired, subradular ganglion is sit-
uated among these gland follicles and was found
present in each species examined; therefore,
it has not been specially mentioned.

Limifossor fratula, Crystallophrisson nanu-
lum, and C. scahrum have been described by
Heath (1911, 1918); for these, abbreviated
diagnoses are given.

GENUS Limifossor Heath, 1904
Type L. talpoides Heath, 1904

Limifossor fratula Heath, 1911

Heath, 1911, pp. 44, 72.

COLLECTIONS:

2189 (9) in 228 fm, with Crystallophrisson
hartmani and C. scabrum.

2190 (1) in 186 fm.

2218 (1) in 249 fm, with C. hartmani.

2220 (3) in 180 fm, with C. hartmani and
C. nanulum .

261

262

PACIFIC SCIENCE, Vol. XVII, July 1963

Fig. 1. The San Pedro area between the mainland of southern California and Santa Catalina Island, show-
ing the distribution of species of solenogasters in different locations. Other data are from Hartman, 1955.

2221 (2) in 147 fm, with C. nanulum.

2230 (1) in 300 fm, with Prochaeto derma
calif ornicum and C. hartmani .

2231 (3) in 115 fm, with Crystallophrisson

sp-

2293 (1) in 252 fm, with C. sp.

2297 (1) in 181 fm.

2306 (3) in 215 fm, with C. scahrum.

2329 (1) in 260 fm.

2337 (1) in 170 fm, with C. sp.

2361 (4) in 167 fm, with C. nanulum , C.
hartmani and C. sp.

2362 (2) in 352 fm, with C. hartmani,
2404 (1) in 438 fm.

2412 (1) in 324 fm, with C. hartmani and
C. hancocki.

2418 (2) in 185 fm, with C. scahrum and
C. nitidulum pacifica.

2430 (2) in 80 fm, with C. sp.

2500 (1) in 450 fm.

2612 (1) in 100 fm, with C. nanulum.

2625 (6) in 230 fm, with C. hartmani and
C. nanulum .

2644 (4) in 310 fm, with C. hartmani and
C. rectum .

2723 (2 ) in 325 fm, with C. hartmani and
C. hancocki.

2738 (1) in 342 fm, with C. hartmani and
C. sp.

2749 ( 1 ) in 277 fm, with C. scahrum ,

2792 ( 1 ) in 300 fm, with Crystallophrisson
sp.

2851 (1) in 230 fm, with C. sp.

3704 (1) in 115 fm, with C. nitidulum
pacifica.

3731 (2 ) in 275 fm, with C. incrassatum.
4756 (2) in 202.2 m.

Solenogaster Mollusks — -Schwabl

s :

4776 (1) in 167.1 m, with Crystallophrisson
sp.

4778 (1) in 215 8 m. with C. sp.

4835 (1) in 140 m.

DIAGNOSIS: The body is short, about 10 mm
long; the radula is very large, distichous, and
has 28 transverse rows. This organ with its
muscles is heavier than that in L. talpoides.
Dorsal salivary glands are present. The stomach
and digestive glands are well developed and
distinct from the intestine. Color is slate gray
with a yellowish cast. Spicules from the middle
of the body measure 0.5 mm long. The length
index is 1. 3-4.7.

DISTRIBUTION : Two individuals were re-
ported by Heath (1911) from southern Cali-
fornia in 260-284 fm. The present collection
contains 64 specimens; 14, or 21.9'%, come
from depths of 77 and 150 fm; 31, or 48.8%,

263

are in 150 to 245 fm; 19, or 29.7%, in depths
greater than 245 and 450 fm. The habitats are
on various substrata, but certain kinds were
selected: 52 individuals, or 81.2%, on clay, 1
on silt, 1 on rocks and mud, and 3 on clayey
mud.

The station numbers refer to localities that
are either in the San Pedro channel area (Fig.
2), in Santa Monica Bay to the north, or in
Redondo canyon or the San Pedro Sea valley.

GENUS Plathymenia Schwabl, 1961
Type P. branchiosa Schwabl, 1961

Schwabl, 196l7‘. p. 100, figs. 1-5.

Neomeniid with short and plump body; the
spicules are hollow needles. The body has a
subterminal atrial opening and a terminal slit-
like dorsoventral cloacal opening. Cirri in the
atrial chamber are reduced; the fore-gut is

- 33%0'

35®2$ -

Fig. 2. The San Pedro area between the mainland of southern California and Santa Catalina Island, show-
ing the distribution of species of solenogasters in different locations. Other data are from Hartman, 1955.

264

PACIFIC SCIENCE, Vol. XVII, July 1963

Fig. 3- Plathymenia branchiosa. Sta. 2324.

without a radula but has unicellular subepithe-
lial dorsal diffuse) and very small ventral
salivary glands; a well-developed oesophagus
with villi is surrounded by numerous gland-
follicles. The mid-gut is largely divided into
diverticles, and the gonads also have large
lateral diverticles. The brain is bilobed, has a
dorsoterminal organ of the Entonomenia type.
The shell-gland has subepithelial glands; there
are no copulatory organs.

The genus is known for a single species,
P. branchiosa Schwabl.

Plathymenia branchiosa Schwabl, 1961
Fig. 3

Schwabl, 1961 b, p. 100, figs. 1-5.

COLLECTION: 2324 (1) in 400 fm, with
Crystallophrisson riedli n. sp.

DIAGNOSIS (translated from the German):
The species has the characteristics of the genus.
The body is flattened ventrally; length of the
body is 10 mm, width 2.5 mm. There is no

dorsal keel but a tapering front end and a
broadened or rounded hind end. Color in alco-
hol is muddy gray. The cuticle is heavy and
there are no epidermal papillae. The spicules
are hollow, straight or slightly curved needles,
the radial ones up to 700 /x long, the tangential
ones up to 100 /x. The ventral furrow is dis-
tinctly separated from the cloacal opening, one
single ventral fold. About 30 sensory setae in
epidermal pockets surround the atrial opening.
The muscles are very weak. The parenchyma
of the hind end has a gelatinous basic sub-
stance. The pericardium is very large and has
short distal appendages. The ventricle for half
its length is dorsally connected with the peri-
cardium, the auricle is free, and has two atrio-
ventricular openings. A large cloacal chamber
has about 110 simple radial gill-folds. There
are no abdominal spicules.

DISTRIBUTION: This species is known
through a single specimen from the San Pedro
area, in 400 fm, in mud with glass sponge.

FIG. 4. Prochaetoderma californicum n. sp. Sta.
2802-54.

Solenogaster Mollusks — SCHWABL

265

FIG. 5. Prochaetoderma calif ornicum. Diagram of entire animal, showing arrangement of internal organs.
b, Brain; bp, buccal plate; g, gill; gd, gonoduct; gpd, gonopericardial duct; gl, ganglion; h, heart; l, liver; m,
mandible; oe, oesophagus; p, pericardium; ph, pharynx; r, radula; ra, radula cartilage; rc, rectum; rg, repro-
ductive gland; s, stomach.

GENUS Prochaetoderma Thiele, 1902
Type P. raduliferum (Kowalevsky) 1901

Prochaetoderma calif ornicum n. sp.

Figs. 4-7

COLLECTIONS:

2228 (1) in 293 fm.

2230 (3) in 300 fm, with Limifossor fratula
and Crystallophrisson hartmani.

2321 (34) in 385 fm, with C. hartmani and
C. sp.

2363 (2) in 429 fm, with C. marinellii and

C. sp., juv.

2411 (4) in 400 fm, with C. hartmani, C.

hancocki, and fragments.

2798 (5) in 386 fm, with C. hartmani and
C. red sum.

2802 (1) in 420 fm, with C. hartmani.

2837 (2) in 454 fm, with C. hartmani.
2839 (6) in 446 fm, with C. hartmani and
C. hancocki.

2899 (3) in 335 fm.

3037 (1) in 360 fm.

DIAGNOSIS: These very small animals meas-
ure 2.25 to 3.05 mm in length; greatest diam-
eter is 0.42-0.85 mm; they show a very char-
acteristic habitus ( Fig. 4 ) ; a short head region
is not separated from the following trunk re-
gion ( Fig. 5 ) but is distinguished from it
by the spines, which are small leaflike structures
(Fig. 7), densely arranged and mostly standing
upright on the body wall. The following region,
occupying the greater part of the body and
containing such main organs as stomach or
mid-gut, liver, and gonads, is covered by flat
imbricating spines arranged obliquely towards
the dorsal midline of the body. The tail-like
hind part of the body is provided with spicules
resembling those of the head region but of
larger size. The terminal end, containing part
of the gonoducts, pericardium, and cloacal
chamber with its organs, is protected by a

266

PACIFIC SCIENCE, Vol. XVII, July 1963

brushlike wreath of long and pointed thorn-
like spicules.

The mouth opening is flanked by the paired
triangular buccal plate, and the entire front
end can be withdrawn into the semicircular
groove. The subepidermal muscle layers, with
the exception of those in the tail region, are
very weak, and the diagonal one is almost
totally lacking. Three pairs of distinct retrac-
tors in the front end, and six pairs of gill re-
tractors, pass laterally from the pericardium
and are present with the thick radula muscles.

The nervous system consists of paired brain
lobes, the two halves connected by a strong
commissure and of three pairs of precerebral
ganglia. The buccal connective leaves the brain
separately, whereas the lateral and ventral ones
have a common root. The buccal system con-
sists of three ventral commissures and a single
dorsal one; an unpaired subradular ganglion is
present.

The digestive tract begins with a wide phar-

FlG. 6. Prochaetoderma calif ornicum. A transverse
row of the distichous radula, showing basal plate,
tooth, and lateral plate.

FIG. 7. Prochaetoderma calif ornicum. Differ-
ent kinds of spines from body regions a, b, c, d,
illustrated in Fig. 9.

ynx, its epithelium bearing a distinct cuticle
which, in two large lateral pockets, is differen-
tiated into a pair of mandibles. Small salivary
glands are developed at the junction of the
pharynx and the short ciliated oesophagus. A
glandular subradular organ lies in front of the
large distichous radula; it consists of 9-12
transverse rows of teeth (Fig. 6) and is situ-
ated in a distinct radular sack. The oesophagus
connects with a short wide stomach from which
the long rectum and the short sacculate liver
originate.

Solenogaster Mollusks — Schwabl

The reproductive organs consist of a short
reproductive gland, a long unpaired ( ! ) gono-
pericardial duct, and the typical gonoducts with
their ciliated glandular parts. The pericardium
is wide and short, its appendages unite ven-
trally and surround the supra-rectal ganglion
as well as the distinct gill-sinus, but do not
extend backwards along the cloacal chamber.
Blood cells are present. The animal possesses
two pairs of gills of a rather primitive structure.

DISTRIBUTION: Southern California, chiefly
in the San Pedro area; 62 specimens all from
considerable depth, between 293 and 454 fm.
Eleven samples were taken, 9 from mud ( 5 in
the glass sponge area), and only 2 samples
( with 4 specimens ) came from other than mud
(gravel or sandy mud). Figure 4 shows one
from station 2802.

Fig. 8. Crystallophrisson incrassatum. Sta. 3831-55.

267

GENUS Crystallophrisson Mobius, 1875
Type C. nitidulum (Loven) 1844

Includes Chaetoderma Loven, 1845, preoccu-
pied in Pisces by Swainson, 1839 (see Thiele,
1932).

Crystallophrisson incrassatum n. sp.

Fig. 8

COLLECTIONS:

3731 (2) in 275 fm, with Limifossor fratula.
3733 (2) in 305 fm.

DIAGNOSIS: The animals were very poorly
preserved; the diagnosis therefore is largely
based on external appearances.

Spicules were totally lacking but the species
obviously possessed these structures originally.
The habitus is very characteristic (Fig. 8).
The body is very stout; the length ranges from
12-17 mm, the greatest diameter between 3
and 4.5 mm, and the length index is 4-5. The
prothorax is white, highly swollen, and may
attain a third of the total length of the animal.
Color of the body is gray, covered with mud.
The metathorax is slender and short; the pre-
abdomen is swollen and contains the particu-
larly large liver. The postabdomen forms a
characteristic conical ring around the cloacal
opening; the gills are mostly protruding from
the cloacal chamber. Buccal, dorsal, and lateral
salivary glands are present. A "tongue” is pro-
vided with a sharp chitinous edge on both
sides behind the main tooth, but real teeth
are lacking. There are four pairs of precerebral
ganglia. The muscles consist of one pair of
dorsal and some pairs of lateral retractors at the
front end. Granular cells are also situated on
the numerous septa dividing the liver into
countless lobules.

DISTRIBUTION: This is known only from the
Santa Barbara basin, southern California, in
about 300 fm, in mud. Fig. 8 an individual
in station 3731.

Crystallophrisson recisum n. sp.

Figs. 9, 10

COLLECTION: 2895 (1) in 265 fm.

DIAGNOSIS: This is a small species, measur-
ing 11 mm long, with an average diameter of

268

PACIFIC SCIENCE, Vol. XVII, July 1963

2 mm. Color of the body is light brown. The
rather swollen prothorax (Fig. 9) measures
about a third of the entire length. The rounded
buccal plate is pierced by the mouth opening.
The spicules (Fig. 10), including the small
ones, are distinguished by a distinct keel; the
pointed thornlike spicules of the hind end bear
conspicuous brown incrustations. With the ex-
ception of the gill retractors, the muscles are
fairly weak; the longitudinal ones split up into
five or six bundles on each side before giving
origin to two pairs of dorsal and three pairs
of lateral retractors of the front end. Among
the usual six pairs of gill retractors, the dorsal
posterior ones are markedly thick and partly
attached to the wall of the cloacal chamber
between the two gills. The posterior vertical

FIG. 9- Crystallophrisson reel sum. Entire animal in
side view, showing the body regions, a, h, c, d. from
which are taken the spines named in the text.

FIG. 10. Crystallophrisson recisum. Spines from
body regions, showing characteristic ones from regions
a, b, c, and d. (Compare with Fig. 9.)

diaphragm is very strong. Three free gill-
lamellae on each side arise from the cloacal
wall. The nervous system is typical; the location
of the brain is rather far back above the radula.
The cerebro-buccal connective is split off the
common trunk shortly behind its origin. The
narrow fore-gut possesses a very strong sphinc-
ter some distance from the mouth opening;
buccal glands are to be found only in front
of it. The three kinds of salivary glands are
present but are unusually small; the radula is

Solenogaster Mollusks — Schwabl

269

provided with two small denticles; the cuticle
of the tongue is thin but highly chitinous. The
cuticular ridge on the dorsal wall of the mid-
gut is near the opening of the liver, and a
rectum is present. Granular cells occur on the
dorsal wall of the liver and on the edges of
the septa which divide the liver into numerous
lobules. The pericardium is spacious; the heart
is typical and has two atrioventricular openings.
The gonoducts open on papillae; their epi-
thelium is not continued on the wall of the
cloacal chamber.

DISTRIBUTION : This has been found only
once, in the San Pedro area of southern Cali-
fornia, in 265 fm, on gray green mud.

Crystallophrisson scabrum Heath, 1911

Heath, 1911, pp. 44, 63, pi. 4, figs. 2, 16, pi.
29, figs. 6, 7, 9-11, pi. 30, figs. 1, 3, pi. 37,
%• 19.

COLLECTIONS:

2189 (12) in 228 fm, with Limifossor fratula
and C. hartmani.

2306 (1) in 215 fm, with Limifossor fratula.
2357 (1) in 100 fm.

2418 (13) in 185 fm, with Limifossor fratula
and C. nitidulum pacifca.

2749 ( 1 ) in 277 fm, with Limifossor fratula.
2793 (2) in 251 fm, with C. sp.

DIAGNOSIS: This is a rather small species,
10-12 mm in length, and 2 mm in greatest
diameter. The region of the liver is greenish,
and the hind end has brown incrustations. The
prothorax is short and swollen, and the second
half of the body is considerably thickened. The
mouth opening is rather large and surrounded
by the round buccal plate. Compact groups of
gland cells occur mainly on the dorsal side of
the fore-gut. The pericardium is large and the
heart highly muscular. The cloacal chamber is
small.

DISTRIBUTION: This species was originally
described from Monterey Bay in 715 fm
(Heath, 1911); the present records are from
the San Pedro area, including Redondo can-
yon, in depths of 100-277 fm, in m.ud.

Crystallophrisson hartmani Schwabl, 1961
Figs. 11, 12

Schwabl, 1961a, pp. 258-277, figs. 1-10.
COLLECTIONS:

2150 (3) in 310 fm, with C. rubrum and C.
riedli.

2189 (1) in 228 fm, with C. scabrum and
Limifossor fratula.

2218 (1) in 249 fm, with Limifossor fratula.

2220 (1) in 180 fm, with Limifossor fratula

and C. nanulum.

2223 (1) in 480 fm, with C. sp.

2230 (2) in 300 fm, with Limifossor fratula
and Prochaeto derma calif ornicum.

2299 (1) in 360 fm.

2301 (1) in 335 fm.

2302 (1) in 185 fm, with C. sp. (used for

Fig. 12, left).

2321 ( 1) in 385 fm, with Prochaetoderma
calif ornicum and C. sp.

2361 (1) in 167 fm, with Limifossor fratula,
C. nanulum and C. sp.

FIG. 11. Crystallophrisson hartmani. Characteristic
spicules from body regions a , b, c, d.

270

PACIFIC SCIENCE, Vol. XVII, July 1963

Fig. 12. Crystallophrisson hartmani. Left, Sta. 2302-53- Right, Sta. 2644-54.

2362 (1) in 352 fm, with Limifossor fratula.

2410 ( 1 ) in 410 fm, with C. hancocki.

2411 (2) in 400 fm, with Pro chaetoderma
calif ornicum and C. hancocki.

2412 (1) in 324 fm, with Limifossor fratula
and C. hancocki.

2441 (3) in 340 fm.

2625 (2 ) in 230 fm, with Limifossor fratula
and C. nanulum.

2642 (4) in 422 fm.

2644 (2 ) in 310 fm, with Limifossor fratula
and C. rectum (see Fig. 12, right ) .

2723 (1) in 325 fm, with Limifossor fratula
and C. hancocki.

2738 (2) in 342 fm, with Limifossor fratula
and C. sp.

2795 ( 1 ) in 290 fm, with C. sp.

2798 (3) in 386 fm, with Prochaetoderma
californicum and C. rectum.

2802 (2) in 420 fm.

2837 (2) in 454 fm, with Prochaetoderma
californicum and C. sp.

2839 (4) in 446 fm, with Prochaetoderma
californicum and C. hancocki.

2846 (2) in 612 fm.

3019 (4) in 425 fm, with C. sp.

DIAGNOSIS: Length of body 5-18 mm, width
of body in front 0.8-2 mm, posteriorly 0 .4-0.6
mm. The anterior half of the body, containing
the short liver, is swollen; the posterior half
is slender and tail-like, terminating in a button-
like posterior end (Fig. 12). The head region is
whitish and the rest of the body is yellowish-
brown with dark incrustations on the hind end.
The buccal plate is round, and the terminal
opening of the cloacal chamber is surrounded
by longer and shorter thorns. The spicules
(Fig. 11) of the anterior end are small and
squamiform; those of the rest of the body are
lancet-like or leaflike and have a keel. The
four groups of longitudinal muscles are lim-
ited to the posterior half of the body; one pair
of dorsal and one of lateral retractors occur
anteriorly. There are five pairs of gill retractors.
The nervous system consists of four pairs of
precerebral ganglia. One pair of small ganglia
connects with the buccal ganglia near the junc-

Solenogaster Mollusks — Schwabl

271

tion of fore- and mid-gut. Buccal glands and
lateral and dorsal salivary glands are present.
The radula has one pair of sickle-like teeth, one
pair of small lateral teeth, and an unpaired
piece. The mid-gut bears a cuticular stripe
dorsally. The liver has granular cells which are
present also at the end of the irregular septa.
Gill lamellae number 15-20. The heart is mus-
cular and has two atrioventricular openings.

DISTRIBUTION : This species occurs commonly
in the San Pedro area of southern California,
in depths of 330-1100 m, in sediments of mud
or clay. It is frequently found in glass sponge
associations.

Crystallophrisson nanulum (Heath) 1911
Fig. 13

Chaeto derma nanula Heath, 1911, pp. 66-67,
pi. 4, figs. 1, 12, pi. 27, fig. 3, pi. 28, figs.
7, 10, 11, 12, pi. 37, fig. 18.

COLLECTIONS:

2220 (7) in 180 fm, with C. hartmani and
Limifossor fratula.

2221 (2) in 147 fm, with Limifossor fratula.
2361 (2 ) in 167 fm, with Limifossor fratula,

C. hartmani and C. sp.

Fig. 14. Crystallophrisson hancocki. Characteristic
spicules from body regions c and d.

2612 (1) in 100 fm, with Limifossor fratula
and C. sp.

2625 (1) in 230 fm, with Limifossor fratula
and C. hartmani.

DIAGNOSIS: The body is small, comparatively
stout (Fig. 13), and measures 7-9 mm long
and up to 1.2 mm at its greatest width. The
spines are keeled and appear heavy. The color
of the body is yellowish-brown. The radula
is small and has two small denticles. Salivary
glands are reduced to a small group of cells
near the radula.

DISTRIBUTION: This was originally described
based on a single specimen from southern Cali-
fornia, in 260-284 fm. The present collections
include 13 specimens, all from the San Pedro
area, in depths of 100-230 fm, and always as-
sociated with Limifossor fratula in sediments of
green or sandy mud.

Fig. 13. Crystallophrisson nanulum. Sta. 2220.

272

PACIFIC SCIENCE, Vol. XVII, July 1963

Crystallophrisson hancocki n. sp.

Figs. 14, 15

COLLECTIONS:

2411 (2) in 400 fm, with Prochaetoderma
calif ornicum and C. hartmani.

2412 (1) in 324 fm, with Limifossor fratula

and C. hartmani.

2475 (1) in 405 fm.

2497 (1) in 24 fm.

2628 (1) in 350 fm.

2641 (1) in 373 fm.

2723 (2) in 325 fm, with Limifossor fratula

and C. hartmani.

2839 (2) in 446 fm, with C. hartmani and
Prochaetoderma calif ornicum.

2888 (1) in 390 fm.

2901 (2) in 310 fm.

DIAGNOSIS: This species is of moderate size
and measures between 19 and 25 mm long,
with a fairly large prothorax (Fig. 15). The
color in alcohol is brownish-gray. The spines
are large but weak, and most of them lack a
marked keel (Fig. 14). The body is distinctly
narrowed between the preabdomen and the
slender and somewhat elongated postabdomen;
on the latter the slitlike dorsoterminal sense
organ is clearly visible. The buccal plate is
rounded and pierced by the mouth opening

FIG. 15. Crystallophrisson hancocki. Sta. 2839-

Fig. 16. Crystallophrisson nitidulum var. pacifica.
Characteristic spicules from body regions a, h, c, d,
and other spines, labeled E, F, G, discussed in the text.

near its dorsal edge. A glandular zone on the
postabdomen and moderate incrustations of
the latter are present. The subepidermal mus-
cle layers are fairly strong, even in the region
of the liver; in the prothorax the longitudinal
layer splits up at once into numerous bundles.
One finds several pairs of ventral, dorsal, and
lateral retractors; the gill-retractor muscles are
typical. The nervous system is characterized by
a brain more clearly bilobed than usual, by a
cerebrobuccal nerve split off the one common
trunk shortly behind its origin, and by a supra-
rectal ganglion surrounded by the pericardium.

The digestive tract begins with a narrow
fore-gut; its sphincter muscle is very strong.
Buccal glands are observed only in front of it.
Small ventral salivary glands with outlets on a
very marked papilla; small dorsal ones behind

Solenogaster Mollusks— -SCHWABL

273

FIG. 18. Crystallophrisson nitidulum var. pacifica.
Sta. 2418.

Crystallophrisson nitidulum (Loven) 1844,
stem species

Crystallophrisson nitidulum var. pacifica n.
var.

Figs. 16-18
COLLECTIONS:

2121 (1) in 32 fm, with C. sp.

2168 (1) in 12 fm.

2176 (3) in 28 fm.

2417 (1) in 34 fm, with C. sp.

2418 (2) in 185 fm, with Limifossor fratula
and C. scahrum.

2614 (2) in 155 fm.

2457 (1) in 111 fm.

2743 (1) in 150 fm.

3204 (1) in 115 fm, with Limifossor fratula.

3205 (1) in 88 fm, with C. sp.

the radula and particularly large lateral ones
extend from the radula to the junction of the
fore-gut with the mid-gut. The radula is large
and highly chitinous but without denticles. It
extends far into the lumen of the fore-gut
and has unusually large supports. The mid-gut
and rectum are spacious. The pericardium is
not very wide but extends far proximally. The
ventricle is muscular and has two atrioventri-
cular openings. The dorsal vessel begins with
a bulb. The epithelium of the gonoducts con-
tinues for some distance on the wall of the
cloacal chamber.

DISTRIBUTION : All specimens come from
the San Pedro area, southern California, and
all but one from depths between 310 and 446
fm. Eleven, or 78.5%, are from mud, and 5,
or 35.7%, from glass sponge areas.

Fig. 17. Crystallophrisson nitidulum var. pacifica.
Sta. 2176.

274

PACIFIC SCIENCE, Vol. XVII, July 1963

DIAGNOSIS: Overall length is 20-40 mm;
greatest width 1.0-2. 5 mm, the largest speci-
mens coming from greater depths. This variety
of the well-known C. nitidulum Loven agrees
with the stem species in its outer appearance.
Characteristic are the not very swollen pro-
thorax, the outline of the buccal plate, the short
but clearly visible dorsoterminal sense organ,
and its inner organization showing the devel-
opment and number of muscles of the stem.
The nervous system is like that of its stem
species, and the radula similarly has two den-
ticles. Other similarities include the structure
of the liver, the epithelium of the gonoducts

Fig. 19. Crystallophrisson riedli. Characteristic
spines from body regions a, b, c, and d.

and anus which continues for a great distance
on the cloacal wall, and above all by the pres-
ence of only one atrioventricular opening.

The variety differs from the stem species
by its greyish color and by the short but widely
expanded postabdomen (Figs. 17, 18), which
contains a large blood sinus surrounding the
cloacal chamber, reducing the latter to an un-
usually small cavity. The spines (Fig. 16) cor-
responding to those of type C (see Fig. 9),
which are characteristic of the stem form, oc-
cur only sparsely, whereas spines of types B,
E, and G (the last only on the postabdomen)
are abundant. On smaller specimens the spicula
of type F could be found in great numbers on
the metathorax, pre- and postabdomen; on
larger specimens they were replaced by spines
of type D, showing a distinct keel. Character-
istic of the variety is the large, clearly outlined
atrium of the heart which surrounds the supra-
rectal ganglion.

DISTRIBUTION: The stem species has been
widely recorded from both sides of the north
Atlantic Ocean, the west coast of Sweden, Nor-
way, Iceland, the North Sea, the eastern coast
of North America from New Foundland to
Virginia, and the West Indies; depths have
ranged from 20-2250 m. The new variety is
known through 14 specimens coming from
the San Pedro area of southern California, in
depths of 12-185 fm, in sediments of mud,
sandy mud ( 2 specimens ) , and sandy clay ( 1
specimen ) .

Crystallophrisson riedli n. sp.

Figs. 19, 20

COLLECTIONS:

2150 (2) in 310 fm, with C. hartmani and C

rubrum,

2324 (1) in 400 fm, with Plathymenia bran-

cbiosa.

2352 (1) in 420 fm.

2414 (1) in 177 fm.

2838 (1) in 394 fm.

DIAGNOSIS: Individuals range in length from
15-22 mm and have an average diameter in
the prothorax of 2 mm. Alcohol preserved
specimens are gray with a whitish prothorax.
In fixed state the posterior half of the prothorax

Solenogaster Mollusks— Schwabl

275

Fig. 20. Crystallopbrisson riedli. Sta. 2838.

is frequently more contracted than the anterior
half (Fig. 20). The dorsoterminal organ is
very short but continuous with the opening
of the cloacal chamber. The buccal plate is
rounded and has a dorsal cleft containing the
mouth opening. The abdominal gland zone is
present. Spines (Fig. 19) are of medium size
and have a marked keel.

Dorsal, lateral, and ventral retractor muscles
are present at the front end; the longitudinal
muscle layer in the prothorax is gradually split
up into numerous bundles. The gill retractors
are short; the ventral anterior ones are very
heavy; the second pair of dorsal posterior ones
are lacking or, if present, attach to the cloacal
wall between the gills. Lateral, ventral, and
part of the dorsal anterior gill retractor unite
before entering the gill, which gives the trans-
verse section of the gill a very characteristic
appearance. The gill axis consists of a bulbous

ventral half, and a very slender dorsal half.
The posterior vertical diaphragm is unusually

strong.

The nervous system is typical; the buccal
nerve leaves the common root very soon. The
fore-gut has a strong circular and longitudinal
muscle layer, circular in outline. Numerous
buccal glands occur behind the sphincter mus-
cle. All three kinds of salivary glands are pres-
ent but the ventral ones are very small. The
radula is typical. Granular cells in the liver
are not only concentrated on the dorsal wall
but irregularly distributed. The circulatory or-
gans are typical, with two atrioventricular open-
ings. The gonopericardial duct is fairly long
and curved; the ciliated part of the gonoducts
is comparatively long. The openings of the
gonoducts are mostly on flat papillae, and their
epithelium is not continued on the wall of the
cloacal chamber.

FIG. 21. Crystallopbrisson inf latum. Characteristic
spicules from body regions a, b, c, and d.

27 6

PACIFIC SCIENCE, Vol. XVII, July 1963

Fig. 22. Cry stall op hris son inflatum. Sta. 2791.

DISTRIBUTION: This species is known
through six specimens, all from the San Pedro
area including Redondo canyon, southern Cali-
fornia, in depths of 177-420 fm. Three were
found in mud, one in mud and gravel, and
one in coarse sand.

Crystallophrisson inflatum n. sp.

Figs. 21, 22

COLLECTIONS:

2110 (3) in 427 fm.

2791 (2) in 415 fm.

DIAGNOSIS: This is a large species, with total
length ranging between 25 and 35 mm, and
greatest width 4 mm; the prothorax region is
greatly inflated (Fig. 22). The hind end is
provided with two lateral prolongations of the
body wall which can be withdrawn into the
circlet of thorns protecting the opening of the
cloacal chamber and the adjacent glandular

zone. Spines ( Fig. 21) are only very slightly
or not at all keeled. The color in alcohol is
dark gray with a brown tinge. The prothorax
is whitish-brown, and the buccal plate is darker
and shows three incisions on its dorsal edge
of which the median one contains the mouth
opening. The muscles are fairly thick, consisting
of two pairs each of dorsal and lateral retrac-
tors at the front end. In the anterior half of
the prothorax there is a characteristic isolated
small bundle of longitudinal muscles on both
sides of the mid-dorsal line. Gill-retractor mus-
cles are typical. The nervous system is also
typical, with a brain situated very near the

Fig. 23- Crystallophrisson marinellii. Characteristic
spicules from body regions a, b, c, and d.

Solenogaster Mollusks — ScHWABL

277

anterior end of the body, with three pairs of
precerebral ganglia, and the cerebrobuccal con-
nective split off the common trunk immediately
after leaving the brain. The fore-gut is short
and unusually wide, with small lateral folds oc-
curring only in the radular region. The buccal
glands are in front of the sphincter muscle,
and all three kinds of salivary glands are pres-
ent, the lateral ones being notably large and
surrounding also the junction of mid- and fore-
gut. The radula is characteristic, large, and
lacking real teeth; the tongue has a very stout,
highly chitinous cuticle which on either side
of the median toothlike structure may attain a
thickness of 25 p. The mid-gut is narrow. The
liver has granular cells on the dorsal wall and
on the edge of the numerous septa.

Reproductive organs are typical; the open-
ings of the gonoducts are on papillae, their
epithelium not continued on the wall of the
cloacal chamber. The pericardium is very spa-
cious and extends far backward; the ventricle
is highly muscular, and there are two atrio-
ventricular openings.

DISTRIBUTION: Five specimens, all from the
San Pedro area of southern California, come
from depths of 415-430 fm, in mud.

Crystallophrisson marinellii n. sp.

Figs. 23, 24

COLLECTIONS:

2363 (1) in 429 fm, with Prochaetoderma

calif ornicum and C. sp.

2473 (1) in 16 fm.

2753 (1) in 322 fm.

2884 (1) in 190 fm.

DIAGNOSIS: This is a comparatively large
specimen (Fig. 24), measuring from 20-40
mm long in a state of contraction; the greatest
width is 3 mm in the region of the pinkish-
white prothorax which may attain a length of
5-7 mm. The terminal buccal plate is brown
and rounded in shape. It has a single deep
cleft on its dorsal side which contains the wide
mouth opening. In alcohol the body is gray
with a distinct pink shade. The spines (Fig.
23) are large but weak, and have a slight keel;
abdominal spines are very short. The dorso-
terminal sense organ is covered with small

Fig. 24. Crystallophrisson marinellii. Sta. 2884.

spines. It is continuous with the opening of
the cloacal chamber in front of which is found
a glandular region. Except for the six pairs of
very strong gill retractors, the muscles are
weak. The longitudinal ones in the prothorax
region are split up gradually into numerous
bundles. Two pairs of dorsal and several pairs
of lateral and ventral retractor muscles are
present.

The brain is located near the radular region
and is not attached to the dorsal wall of the
fore-gut but is surrounded by numerous groups
of piriform gland cells belonging to the glands
of the buccal plate. The common trunk of the
three large connectives follows its course for
some distance before the buccal connective is
free. Four pairs of precerebral ganglia are pres-
ent. The suprarectal ganglion may be sur-
rounded by the pericardium and even by the
atrium. Behind the mouth opening the fore-
gut narrows conspicuously and does not widen

278

PACIFIC SCIENCE, Vol. XVII, July 1963

much along its course. The radula is powerful
but without real teeth or denticles. All three
kinds of salivary glands are present. Buccal
glands occur only in front of the sphincter
muscle. The mid-gut is very wide. The peri-
cardium is notably large, the ventricle highly
muscular and there are two atrioventricular
openings. The dorsal vessel originates as a con-
spicuous ampulla situated in the pericardium.

Reproductive organs are without peculiarity.
The ciliated portion of the gonoduct is rather
long; the gonoduct openings are on papillae,
their epithelium not continued on the wall of
the cloacal chamber.

distribution: Four specimens, all from the
San Pedro area of southern California, were
taken in depths of 16-429 fm, in mud or
sandy mud.

Fig. 25. Crystallophrisson rectum. Characteristic
spicules from body regions c and d.

FIG. 26. Crystallophrisson rectum. Sta. 2798.

Crystallophrisson rectum n. sp.

Figs. 25, 26

COLLECTIONS:

2644 (1) in 310 fm, with Limifossor fratula
and C. hartmani.

2798 (1) in 386 fm, with Prochaetoderma
calif ornicum and C. hartmani.

DIAGNOSIS: These animals are moderately
large (Fig. 26) being 10-18 mm long, and
having an average diameter of 1—1.5 mm. The
prothorax may reach 2.5 mm in diameter. The
color is yellowish-brown. The spines (Fig. 25)
are weak and have little structure. The buccal
plate is semicircular and has a deep and wide
dorsal cleft which contains the mouth opening.
The cuticle is very thin, the abdominal gland
zone present. The subepidermal muscle layers
are weak; there are three pairs of dorsal, three
pairs of lateral, and several pairs of ventral

Solenogaster Mollusks- — SCHWAB L

279

retractor muscles. Only five pairs of gill re-
tractors are present; the ventral posterior ones
are lacking.

The nervous system shows a very compact
mass of three pairs of precerebral ganglia and
a markedly bilobed brain, both attached to
the dorsal wall of the fore-gut; the buccal nerve
is shortly freed. The suprarectal ganglion is
surrounded by the pericardium. The fore-gut is
narrow, triangular in outline where located

>6ocy<.

Fig. 27. Crystallophrisson rubrum. Characteristic
spicules from body regions b, c, and d.

Fig. 28. Crystallophrisson rubrum. Sta. 2150.

between the mouth opening and the brain,
then semicircular in the brain region, and some-
what widened and quadrangular in the region
of the radula. Numerous buccal glands are vis-
ible behind the sphincter and all three kinds
of salivary glands are present. The radula is
typical and lacks denticles. The mid-gut is wide
and has a cuticular region near its opening to
the liver. The rectum is slightly winding.

The pericardium is spacious, the ventricle
slender and not very muscular, with two atrio-
ventricular openings. The dorsal vessel begins
with a bulb. Gills are conspicuously small and
have only a few gill lamellae. The epithelium
of the gonoducts is not continued on the wall
of the cloacal chamber.

DISTRIBUTION: Two specimens come from
the San Pedro area, southern California, in
310-386 fm, one from clay sediments, the other
from mud with glass sponge.

280

PACIFIC SCIENCE, Vol. XVII, July 1963

Cry st alio phr is son rubrum n. sp.

Figs. 27, 28

COLLECTION :

2150 (1) in 310 fm, with C. hartmani and

C. riedli.

DIAGNOSIS: This species is readily distin-
guished from others by its outer purple color
and its plump body ( Fig. 28 ) . The single speci-
men measured 17 mm long and 2.5 mm at its
greatest width. The front end was withdrawn
and the exact outline of the buccal plate is
thus not known, but it is certainly pierced by
the mouth opening. The spines (Fig. 27) are
large and stout and have a distinct keel; the
abdominal needle-like spines attain a length
of 800 n. A glandular zone in front of the
opening of the cloacal chamber is present and
"giant cells" are frequently encountered in the
epidermis of the entire body. The animal is
highly muscular; there are two pairs of dorsal,
one pair of lateral, and one pair of ventral
retractors of the front region, all of them very
strong. These retractors split off from the longi-
tudinal muscle layer which is located dorsally
in the prothorax, dividing only into a few
rather large bundles. The six pairs of gill-re-
tractor muscles are typical; the anterior dorsal
one has a double origin. The diaphragms are
very strong, and an anterior vertical one can
be distinguished.

The nervous system shows three pairs of
small compact precerebral ganglia, a small
brain, separated from the dorsal wall of the
fore-gut by a large sinus, and a cerebrobuccal
connective split off the common trunk at some
distance from the latter’s origin. The fore-gut
is provided with strong longitudinal muscle
fibres and with a stout sphincter, with buccal
glands anterior to the last and with all three
kinds of salivary glands, of which the dorsal
ones open into an unusually large pouch of
the fore-gut. The strong radula consists of the
typical parts in addition to a pair of small
pointed, curved denticles.

The reproductive organs are typical; the
gonoducts are simple without sacculations;
their dorsal ciliated parts are long and slender
tubes. The epithelium of their outlets contin-
ues on the wall of the cloacal chamber. The

pericardium is very large and together with
the posterior part of the atrium surrounds the
suprarectal ganglion. The muscular ventricle,
connected with the atrium by two atrioventricu-
lar openings, is attached at its front end to
the dorsal and ventral pericardial wall. The
dorsal vessel begins as a large bulb.

distribution: This species is known
through a single specimen from Redondo cafi-
yon, in the San Pedro area, southern California,
in 310 fm, in mud.

ACKNOWLEDGMENTS

The author is indebted to all those who made
these collections possible. The operations of the
"Velero IV" were supported by funds provided
partly by the Allan Hancock Foundation, the
California State Water Pollution Control Board,
and the National Science Foundation. The sort-
ing of solenogasters was done by staff members
of the Hancock Foundation; the anatomical
preparations were prepared in the laboratory of
the Department of Zoology, University of Vi-
enna. The author is also indebted to the Aus-
trian Department of Education and the Depart-
ment of Zoology of the Free University of West
Berlin.

APPENDIX

Additional Station Data

Physical data are published for most of the
station numbers listed above. Those not sep-
arately listed below are to be found in Hart-
man (1955) and Hartman and Barnard (1958,
I960). The following are newly listed; all
localities are in southern California.

2457-53. Oct. 17, 1953. 4.0 miles SE of Point
Fermin lighthouse, 33° 39' 19" N. lat., 118°
14' 20" W. long. In 21.5 fm. A snapper
sampler took a very small sample of sand.
2884-54. July 27, 1954. 5.6 miles SW of end
of Newport pier. 33° 31' 58" N., 118° 00'
00" W. In 190 fm. The orange-peel-grab
took 1.93 cu ft of gray green mud.

Solenogaster Mollusks — SCHWABL

281

2888-54. July 27, 1954. 14.7 miles EWE of East
end, Santa Catalina Island. 33° 21' 58" N.,
118° 02' 00" W. In 390 fm. OPG took 2.67
cu ft of gray green mud.

2895-54. July 28, 1954. 9.9 miles SSW of end
of Newport pier. 33° 27' 57" N., 118° 01'
58" W. In 265 fm. OPG took 2.58 cu ft of
gray green mud.

2899-54. July 28, 1954. 8.25 miles SW of end
of Newport pier. 33° 30' 00" N., 118° 02'
02" W. In 266 fm. OPG took 2.64 cu ft of
gray green mud.

2901-54. Aug. 11, 1954. 12.8 miles SW of East
end, Santa Catalina Island. 33° 26' 00" N.,
118° 06' 06" W. In 312 fm. OPG took 3.02
cu ft gray green mud.

3037-55. May 8, 1955. 3.75 miles from North
light, Santa Barbara Island. 33° 31' 17" N,
118° 58' 00" W. In 360 fm. OPG took 2.58
cu ft of gray green sandy mud.

3204- 55. July 7, 1955. Santa Monica Bay. 33°
58' 10" N., 118° 39' 10" W. In 115 fm.
OPG took 2.08 cu ft of green silty mud.

3205- 55. July 7, 1955. Santa Monica Bay. 33°
58' 00" N, 118° 37' 00" W. In 88 fm. OPG
took 2.08 cu ft of green silty mud.

4756-56. Dec. 8, 1956. 4.1 miles from Scripps
pier, La Jolla. 32° 51' 55" N., 117° 20' 30"
W. In 102 fm. OPG took 2.39 cu ft of green
mud and some broken shells; temperature of
sediment 9.8 C.

4776-56. Dec. 10, 1956. 2.2 miles from Dana
Point. 33° 26' 00" N., 117° 43' 30" W. In
87 fm. OPG took 3.46 cu ft of green mud;
temperature of sediment 10.2 C.

4778-56. Dec. 10, 1956. 5.2 miles from Dana
Point. 33° 30' 20" N., 117° 47' 45" W. In
100 fm. OPG took 2.96 cu ft of green mud.
Temperature of sediment 9.7 C.

4835-57. Feb. 6, 1957. 11 miles from Port
Hueneme Light. 34° 02' 15" N., 119° 01'
45" W. In 77 fm. OPG took 1.38 cu ft of
olive green sandy silt.

REFERENCES

Hartman, Olga. 1955. Quantitative survey
of the benthos of San Pedro basin, southern
California, Part 1. Preliminary results. Allan
Hancock Pacific Exped. 19 ( 1 ) : 1-185, 9 pis.

— and J. Laurens Barnard. 1958, I960.

The benthic fauna of the deep basins off
southern California. Allan Hancock Pacific
Exped. 22 (1, 2): 1-297, 19 pis., map.

Heath, Harold. 1911. Reports on the scien-
tific results of the expedition to the tropical
Pacific, in charge of Alexander Agassiz, by
the US Fish Commission Steamer "Alba-
tross,” from August, 1899, to June, 1900.
The Solenogasters. Mem. Mus. Comp. Zool.
Harvard 45: 1-182, 40 pis.

1918. Solenogasters from the eastern

coast of North America. Mem. Mus. Comp.
Zool. Harvard 45:183-263, 14 pis.

SCHWABL, Mathilde. l%la. Crystallophrisson
( = Chaetoderma ) hartmani, nov. spec., eine
neue Aplacophore aus dem Ostpazifik. Zool.
Anz, Bd. 166, Heft 7-8:258-277, 10 figs.

— — — 1961 A Plathyucenia branchiosa, nov.
gen., nov. spec., ein neuer Vertreterder Neo-
meniidae aus dem Ostpazifik. Zool. Anz., Bd.
167, Heft 3-4:100-115.

Thiele, J. 1932. Die Solenogastres des arkti-
schen Gebietes. Fauna Arctica, Hena, Bd. 6,
Lief. 4:379-382.

Studies in Fitchia (Compositae) : Novelties from the
Society Islands; Anatomical Studies

Sherwin Carlquist and Martin L. Grant1

The genus Fitchia has attracted interest be-
cause of its arborescent habit, its distribution
in southeastern Polynesia, the endemism of its
species, and its phylogenetic relationships. In
a monograph of this genus, six species occur-
ring on live islands were recognized ( Carlquist,
1957). Following the appearance of that mono-
graph, Grant notified Carlquist of his collec-
tion, while on a Bishop Museum Fellowship
in 1931, of specimens of Fitchia from two ad-
ditional islands (Bora Bora and Tahaa) in the
Society Group. The genus had not been pre-
viously reported from these islands, and manu-
script names, as a new species and subspecies,
respectively, had been applied to the taxa. On
Grant’s invitation, Carlquist made the anatomi-
cal studies presented here which, it is felt, puts
the recognition of the two new entities on a
much firmer basis than gross morphology alone
could have provided. In addition, descriptions
are given of one additional specimen each of
two previously known species of Fitchia, one
from Tahiti, and one from Rapa. The occur-
rence of the two novelties is of more than or-
dinary interest because of the nature of this
genus, and permits more extended remarks on
the nature of speciation in Fitchia. Flowering
material of F. rapensis, not available for the
monograph, has been studied here. This paper,
therefore, may be considered an addendum to
the monograph. The taxonomic descriptions be-
low have been prepared by Grant. The anatomi-
cal account which follows is the work of Carl-
quist, and has been prepared to conform with
the types of data offered in the monograph.

1 Claremont Graduate School, Claremont, California;
and Iowa State Teachers College, Cedar Falls, Iowa.
Manuscript received December 28, 1961.

TAXONOMIC DESCRIPTIONS

1. Fitchia cor data M. L. Grant et S. Carlquist,
sp. nov.

Fig. 31

Frutex arboreus 2 m altus glaber, caule basi-
liter 4 cm crasso. Folia simplicia petiolata petio-
lis 25-65 mm longis, longitudine media (N =
42) 47 mm. Laminae late obovatae vel subor-
biculares luteovirides 40-72 mm longae 36-65
mm latae, magnitudine media foliarum grandi-
orum circa 58 X 54 mm, basi cordata vel ro-
tundata, apice obtuso acutove vel parum acu-
minate, margine crenulato vel subintegro, venu-
lis lateralibus 5-10 (numero medio 8.5) pro
unico latere. Pedunculi terminales 20-40 mm
longi, longitudine media ( N = 7 ) 33 mm. Capi-
tula solitaria circa 40-florata. Involucra 25 mm
longa 25 mm lata, bracteis 3-4 seriatis viride-
lutei coriaceis marginibus- scariosis, exterioribus
late triangulatis 3 mm longis 9 mm latis, medi-
oribus semiorbicularibus, interioribus lanceo- i
latis 13-17 mm longis 5-7 mm latis. Bracteae
receptaculares 12-17 mm longae 2-3 (-5) mm
latae. Corollae 15-20 mm longae, tubis circa 10
mm longis, sinibus subaequalis. Antherae saccis
5 mm longis, apicibus staminum cuneato-lance-
olatis 1-1.5 mm longis. Ovaria ad anthesin 7-8
mm longa, stylis 20-22 mm longis. Achaenia
matura 13-16 mm longa, aristis 8-10 mm longis
plerumque triangulatis in sectione transversa
subdeciduis.

Tree-like shrub, 2 m tall, 4 cm in diameter \
at the base, glabrous. Bark 1 mm thick, brown,
almost smooth, with fine longitudinal ridges
0.5 mm deep. One-year twigs on flowering
shoots greenish, 2.5-3 mm in diameter; two-

282

Studies in Fitchia — CARLQUIST and GRANT

283

FIGS. 1—6. Secondary xylem. Figs. 1-4, Fitchia cuneata ssp. tahaaensis. Figs. 5-6, F. cordata. Figs. 1, 2:
Tran sections; note tyloses in vessels; Fig. 2, a sclerified tylosis in which the lumen is nearly occluded. Figs. 3, 4:
Tangential sections; Fig. 3 is a portion showing long, thick-walled fibers; Fig. 4 shows storied apotracheal
parenchyma cells. Fig. 5: Transection; note band of apotracheal parenchyma above center of photograph. Fig.
6: Tangential section. Fig. 2, X 300; others, X 117.

year twigs pale brown, 4-5 mm in diameter;
internodes 4-12 mm long, on vegetative shoots
to 20 mm long. Leaves simple. Petioles 25-65
mm long, the average being 47 mm ( N = 42 ) ;
the stipular sheaths 0.4-0.6 mm high. Blades

broadly ovate to suborbicular, yellowish-green,
with a narrow (0.3 mm) stramineous margin,
and with a dark callous at the apex; 40-72 mm
long, 36-65 mm wide, the average ( N = 8 )
size of the largest blade on the flowering shoots

284

PACIFIC SCIENCE, Vol. XVII, July 1963

being 62 X 55 mm, the second largest 58 X 54
mm, and the smallest of the fully-developed
leaves 64 X 43 mm; the base cordate to a
depth of 1-2 (-3) mm or rounded; the apex
obtuse to acute, with the margins near the tip
slightly rounded to straight, or, if barely acu-
minate, departing at most 1 mm frorn^a straight
line; the margin crenulate, with 2-3 indenta-
tions (up to 0.5 mm deep) per cm, to suben-
tire, the base entire for 20 mm adjacent to the
petiole; with 5-10 (average 8.5) pairs of major
lateral veins, usually curving somewhat towards
the tip of the leaf. Peduncles terminal, 20-40
mm long, the average (N = 7) 33 mm, often
arcuate and reflexed. Head solitary, with about
40 flowers. Involucres 25 mm long and 25 mm
wide; the bracts in 3-4 series, greenish-yellow,
coriaceous, scarious-margined, the outermost
broadly deltoid (3 mm long, 9 mm wide), the
middle ones semicircular, and the inner ones
lanceolate (13-17 mm long, 5-7 mm wide).
Receptacular bracts 12-17 mm long, 2-3 (-5)
mm wide. Corollas 15-20 mm long, the tube
about 10 mm long, the shallower sinuses nearly
equalling the ventral cleft, and the lobes with
terminal hairs 0. 1-0.6 mm long. Anther sacs
5 mm long, the stamen tips cuneate-lanceolate,
1-1.5 mm long. Ovary at anthesis 7-8 mm long;
the styles 20-22 mm long. Ripe achenes 13-16
mm long, with awns 8-10 mm long; awns gen-
erally triangular in transection and somewhat
deciduous.

DISTRIBUTION: Society Islands, endemic to
Bora Bora.

SPECIMEN EXAMINED: Bora Bora, in dwarf
rain-forest on the summit of Mt. Tarapaia, alti-
tude 645 m, Jan. 3, 1931, M. L. Grant 4968
(bish, type; RSA, ISTC, isotypes).

This species was found in several places on
Mt. Tarapaia, none less than 25 m in altitude
below the summit, but only four inflorescences
were discovered, two in bud, one at anthesis,
and one in fruit. Associated woody plants were
Alstonia, Metrosideros, Freycinetia, and Glochi-
dion. Native name: anei.

Mt. Tarapaia appears on U. S. Hydrographic
maps as Mt. Pahia, elevation 2165 ft (— 660
m), the name and elevation being taken from
French maps. It has been recorded as "Tarao-
pai’a.” The only higher point on Bora Bora is

Otemanu (Mt. Temanu of the maps), eleva-
tion 725 m, a peak which apparently has never
been climbed by anyone other than Polynesians.

2. Fitchia cuneata J. W. Moore, Bishop Mus.

Bull. 102:48. 1933 ssp. tahaaensis M. L.

Grant et S. Carlquist, subsp. nov.

Fig. 32

Frutex arboreus 2.5 m altus glaber, caule
basaliter 2.5 cm crasso. Folia simplicia petiolata
petiolis 10-37 mm longis, longitudine media
(N=30) 20 mm. Laminae plerumque ellipti-
cae vel parum ovatae obovataeque vel rhom-
boidales subatrovirides 40-110 mm longae 25-
58 mm latae, magnitudine media foliarum
grandiorum circa acuminato, margine crenu-
lato serratove vel subintegro, venulis lateribus
5-10 ( numero medio 7.5 ) pro unico latere.
Pedunculi terminales 25-50 mm longi, longi-
tudine media (N= 10) 33 mm. Capitula soli-
taria circa 75-florata. Involucra 20-40 mm longa,
15-25 mm lata, bracteis 3-4 seriatis, viride-
lutei coriaceis marginibus . chartaceis, exteriori-
bus breviter ovatis reniformibusque 2-5 mm
longis 6-15 mm latis, medioribus semiorbiculari-
bus 8-12 mm longis 9-13 mm latis, interioribus
cuneatis 12-20 mm longis, 4-8 mm latis. Brac-
teae receptaculares lanceolatae 12-17 mm longae
2-4 mm latae. Corollae ad anthesin 15-22 mm
longae, tubis 4-10 mm longis, dentibus 5-7 mm
longis. Antherae saccis, 4.5-5 mm longis, apici-
bus staminum cuneato-lanceolatis 1.5 mm
longis. Ovaria ad anthesin 8-10 mm longa, 3-4
mm lata, stylis 24-30 mm longis. Achaenia
matura 10-20 mm longa, 4-6 mm lata, aristis
10-13 mm longis plerumque teretibus sub-
persistentibus.

Treelike shrub, 2.5 m tall, 2.5 cm in diameter
at the base, glabrous. Bark 1 mm thick, brown,
roughened with irregular scaly ridges. One-year-
old twigs on flowering shoots greenish, 2 mm
in diameter; two-year twigs pale brown, 3 mm
in diameter; internodes 4-8 mm long, on vege-
tative shoots to 12 mm long. Leaves simple.
Petioles 10-37 mm long, the average (N = 30)
20 mm; the stipular sheaths 0. 5-0.8 mm high.
Blades mostly elliptical, varying to slightly
ovate, obovate, or rhomboid, dull green, with
a narrow stramineous margin, and with a dark

Studies in Fitchia — Carlquist and Grant

285

callous at the apex; 40-110 mm long, 25-58
mm wide, the average size (N=10) of the
largest leaf on flowering shoots 84 X 45 mm,
the second largest 75 X 42 mm, and the small-
est of the fully developed leaves 59 X 30 mm;
the base cuneate (usually) to barely rounded;
the apex acute, occasionally slightly acuminate;
the margin crenulate to toothed or subentire,
with 2-4 indentations (up to 1 mm deep) per
cm; with 5-10 (average 7.5) pairs of major
lateral veins, mostly curving slightly toward the
tip of the leaf. Peduncles terminal, 25-50 mm
long, the average (N=10) 39 mm long, al-
ways reflexed and usually arcuate at anthesis.
Heads solitary, with about 75 flowers. Involucres
15-25 mm long, 20-40 mm wide; the bracts in
3-4 series, greenish-yellow, coriaceous, charta-
ceous-margined, the outermost short ovate or
reniform (2-5 mm long, 6-15 mm wide), the
middle ones semi-circular (8-12 mm long, 9-13
mm wide), and the inner cuneate (12-20 mm
long, 4-8 mm wide ) . Receptacular bracts lanceo-
late, 12-17 mm long, 2-4 mm wide. Corollas
at anthesis 15-22 mm long, the tube 7-10 mm
long, and the teeth 5-7 mm long, the shorter
teeth about half the length of the limb, or
eventually equalling it; lobes with hairs about
0.75-1 mm long. Anther sacs 4.5-5 mm long,
the stamen tips cuneate-lanceolate, 1-1.5 mm
long. Ovary at anthesis 8-10 mm long, 3-4 mm
wide; styles 22-30 mm long, the stigmatic
branches 0.8 mm long, barely separating at an-
thesis. Mature achenes 10-20 mm long, 4-6
mm wide; the awns 10-13 mm long, generally
rounded in transection, subpersistent.

DISTRIBUTION: Society Islands, endemic to
Tahaa.

SPECIMEN EXAMINED: Tahaa, district of
Ruutia, slopes of Mt. Ohiri, in rain-forest of
Crossostylis, Alstonia, and Morinda, altitude
465 m, Jan. 25, 1931. M. L. Grant 5161 (bish,
type; RSA, ISTC, isotypes).

This subspecies was observed in three other
nearby localities, all within an altitudinal range
of 15 m, and about 70 m below the top of
Ohiri, the highest point on the island. Other
associated woody plants were Xylosma, Meryta,
Metrosideros, Wikstroemia, F agar a, Astronia,
and Hernandia.

3. Fitchia nutans Hook f., London Jour. Bot.
4:640, t.23-24. 1845.

Since so few specimens of this species are
available in the United States, and the several
Kew and Paris sheets are without detailed habi-
tat data, the citation of an additional collection
may be of interest: Tahiti iti, district of Teahu-
poo, on Mt. Ronui, in Metrosideros- Weinman-
nia forest, altitude 890 m, July 2, 1930, M. L.
Grant 3925 (BISH, istc). The following de-
scription is from this collection alone.

Treeshaped glabrous shrub, 2.5 m tall, with
a basal diameter of 5 cm; bark 2 mm thick,
brown, with longitudinal ridges about 1 mm
deep; the wood very sweet-smelling. One-year
twigs 2-3 mm in diameter; two-year twigs 3.5-
4.5 mm thick; internodes 7-10 mm long. Peti-
oles 30-60 mm long; stipular sheaths 2-3 mm
high. Blades ovate, yellowish-green, 60-130 mm
long, 45-80 mm wide, the average of 12 leaves
being 96 X 61 mm; truncate or cordate at
the base, occasionally barely rounded, often
somewhat oblique; the apex slightly acuminate;
the margin irregularly and shallowly crenulate
to entire; with 8-10 major lateral veins. Pedun-
cles two in each of the four inflorescences pres-
ent, 60-65 mm long, reflexed. Heads shattered
with age, the involucral bracts and corollas hav-
ing dropped. Receptacular bracts 13-14 mm
long, 3-5 mm wide. Ripe achenes 9-10 mm
long, 2-2.5 mm wide, with persistent awns 7-8
mm long.

This collection fits the general description
of the species (Carlquist, 1957: 63), except
as follows, with Carlquist’s measurements and
notes in parentheses: plant smaller (4. 5-7. 5 m
tall), blades often crenulate (entire), maximum
leaf size greater (115 X 70 mm), base often
truncate to cordate (acute to obtuse), heads
paired (solitary), receptacular bracts shorter
(20-22 mm long), achenes much shorter (lb-
17 mm long at anthesis) and narrower (3-4
mm). These differences, however, do not seem
significant enough to warrant separation, and
the material matches Hooker’s plate closely.

Although the type of the species was sup-
posedly from "Elizabeth Island,’’ all the evi-
dence (Carlquist, 1957: 63) suggests that it is
confined to Tahiti. The present specimen comes

FIGS. 7-12. Figs. 7, 8, 11, Fitchia cordata. Figs. 9, 10, 12, F. cuneata ssp. tahaaensis. Fig. 7: Leaf transec-
tion, adaxial face above. Secretory canal is above vein at right. Fig. 8: Transection of awn from an achene,
adaxial face at right. Note lignification of hypodermal layers, presence of a single vascular bundle. Fig. 9:
Leaf transection, adaxial face above. In addition to secretory canals adjacent to vein, left, secretory cavities
may be seen, center and right. Fig. 10: Transection of awn from an achene, adaxial face upper right. Tri-
chomes in section, above and below. Fig. 11: Transection of pith. Secretory canal with adjacent fibrosclereids,
above; inner margins of vascular cylinder, below; note thin-walled cells at top of figure. Fig. 12: Longitudinal
section of pith. All cells have secondary walls, some such are thicker walled. Figs. 7, 9, X 145. Figs. 8, 10,
X 255. Figs. 11, 12, X 135.

Studies in Fitchia — CARLQUIST and GRANT

from the smaller mountain (Tahiti-iti) of the
two which make up the island.

Hoffmann (1890: 353) reported F. nutans
from Tabuai ( = Tubuai ) , as well as from
Tahiti and Elizabeth (now Henderson) I., but
as Hemsley (1885: 20) had earlier shown,
this supposed locality is based on the type sheet
(Cuming 1424) which Hooker had earlier at-
tributed to Elizabeth Island.

Erdtman (1952: 124) cites a specimen of
F. nutans as "Fiji, Moseley, anno 1875.” Carl-
quist (1957:63) suggested that this is in error,

287

because no other collections have been made of
Fitchia in the Fiji group. Also, Hemsley ( 1885:
20 ) reports that there is at Kew a specimen
collected by Moseley in Tahiti at an altitude of
4,000 ft, and this is probably the specimen
cited by Erdtman.

Papy (1955: 325) mentions F. nutans as
growing on "Tahiti, Moorea, Raiatea, Rapa.”
The record from Rapa is probably based on the
report of Riley (1926: 55), which was made

before F. rapensis had been described as a
separate species (Brown, 1935: 366) , although

FIGS. 13-16. Fig. 13: Fitchia cordata. Transection of involucral bract taken about midway along length of
bract; adaxial face at left. Fig. 14: F. cordata. Transection receptacular bract, adaxial face at left. In Figs. 13,

14, tracheary elements shown in bold outline, fibers and sclereids stippled. Fig. 15: "Typical” leaf of F.
cordata, showing main veins. Fig. 16: "Typical” leaf of F. cuneata ssp. tahaaensis. Figs. 13, 14, X 105. Figs.

15, 16, X 2/3.

288

PACIFIC SCIENCE, Vol. XVII, July 1963

Papy was aware of this. Neither Moore nor
Grant found F. nutans on Raiatea, and Papy
was aware of the description of F. cuneata from
that island. Moorea is certainly a likely locality
for F. nutans, but for none of these localities
does Papy cite specimens.

Thus, the geographic distribution offered by
Carlquist (1957:2) must be broadened to in-
clude the occurrence of F. nutans on Tahiti-iti,
F. cordata on Bora Bora, and F. cuneata ssp.
tahaaensis on Tahaa.

4. Fitchia rapensis F. Brown, Bishop Mus. Bull.
130:366. 1935.

Since the ten previously recorded collections
of this species are all without flowers (Carl-
quist, 1957 : 62 ) , the description of a flowering
specimen should be of interest: Rapa, moist
zone, altitude 520 m, Jan. 3, 1922, W. B. Jones
372 ( Whitney Expedition) ( BKL ) .

"Tree,” glabrous. Petioles 40-75 mm long.
Blades ovate, the larger ones averaging 90 mm
long and 84 mm wide, rounded or scarcely
cuneate at the base, the apex acute; entire; with
about 12 pairs of major lateral veins. The sin-
gle inflorescence is terminal, with two heads,
the peduncles 50 and 60 mm long, reflexed.
Heads 45 mm long and 45 mm wide. Bracts in
about 4 series, those of the second layer semi-
circular, 10 mm long and 22 mm wide, and
those of the inner layer 20 mm long and 12
mm wide. Receptacular bracts cuneate, 16-17
mm long, 3-4 mm wide. Mature corollas 24-26
mm long, the ventral cleft reaching to 6-8 mm
of the base, the segments eventually separating
to within 10 mm of the base; the tips of the
lobes often densely covered with sclerified cells,
which are up to 0.75 mm long. Anther sacs
5.5-6 mm long, caudate, the tail 0.3-0.4 mm
long; the stamen tips cuneate, 1.5 mm long.
Mature style ( broken off) at least 32 mm
long. Achenes, at and just after anthesis, 10-12
mm long, 4 mm wide, the awns 12-15 mm
long, strongly bent and twisted at the base.

ANATOMICAL DESCRIPTIONS

Anatomical studies were based on the her-
barium specimens cited above. Liquid-preserved
heads of F. cuneata ssp. tahaaensis and wood
samples of both F. cuneata ssp. tahaaensis and
F. cordata , for which the herbarium specimens
serve as vouchers, were available. The methods
of preparation of these materials are the same
as those employed for such materials in the
two papers (Carlquist, 1957, 1958) dealing
with the anatomy of this genus.

SECONDARY XYLEM: With respect to quali-
tative features, the species of Fitchia other than
F. speciosa are much alike in wood anatomy
(Carlquist, 1958). Fitchia cuneata ssp. tahaaen-
sis and F, cordata share the qualitative features
of those species. The bands of apo tracheal
parenchyma are clearly shown by the two new
taxa ( Figs. 4, 5 ) . Thick-walled fibers are readily
apparent In F. cuneata ssp. tahaaensis (Figs. 1,
3 ) ; libriform fibers are less thick-walled in
F cordata (Figs. 5, 6). Sclerosed tyloses have
been reported in F. nutans and F. speciosa
(Carlquist, 1957, 1958); they are abundant in
F. cuneata ssp. tahaaensis (Figs. 1, 2), although
none were observed in F. cordata .

Quantitative features of the two new taxa
are as follows: F. cordata : vessels per group:
1.37; average vessel diameter: 61.5 /x; diameter
of widest vessel: 88" "p; average length vessel
elements: 325 ft; average length libriform fibers:
502 /a; average length apotracheal parenchyma
cells: 356 g; average height multiseriate rays:
1.17 mm. Fitchia cuneata ssp. tahaaensis: ves-
sels per group: 1.66; average vessel diameter:
62.8 /a; diameter of widest vessel: 92 /a; average
length vessel elements: 326 /a; average length
libriform fibers: 507 /a; average length apo-
tracheal parenchyma cells: 380 /a; average
height multiseriate rays: 1.51 mm.

Quantitative as well as qualitative data point
up a close similarity between F. cuneata ssp.
tahaaensis and the typical F. cuneata. Quanti-
tative and qualitative data for F cordata re-

FIGS. 17-19. Fitchia cuneata ssp. tahaaensis. Fig. 17: Transection of involucral bract taken about midway
along length; adaxial face below. Fig. 18: Transection of receptacular bract; ad axial face and adjacent achene at
left. Fig. 19: Transection of style. At center of style, stigmatoid tissue. Exterior to each bundle is a secretory
canal, except for bundle at upper left, which is flanked by a pair of canals. At right, anthers in transection.
Note secretory canal in each of the two anthers. Fig. 17, X 100; Fig- 18, X 80; Fig. 19, X 120.

290

PACIFIC SCIENCE, Vol. XVII, July 1963

semble the figures given (Carlquist, 1958: 6-7)
for F. cuneata and F. nutans.

PITH: Pith anatomy proved important in dis-
tinguishing species of Fitchia (Carlquist,
1957). Likewise, the two new taxa possess
characteristics useful for taxonomic purposes.
Pith of F. cordata (Fig. 11) consists of both
thick-walled, lignified, and thin-walled, non-
lignified, cells. Secretory canals are present; each
is surrounded by an eccentric zone of fibro-
sclereids. Although secretory canals surrounded
by small lignified thick-walled cells occur in F.
nutans and F. tahitensis, the presence of these
combined with occurrence of thin-walled non-
lignified cells in pith is characteristic only of
F. cordata. Pith of F. cuneata ssp. tahaaensis
(Fig. 12) lacks secretory canals and consists
of thin-walled and thick-walled lignified cells.
The only pith pattern which matches this is
that described in the monograph for typical F.
cuneata.

LEAF: Leaf dimensions are mentioned in the
taxonomic description above. However, aver-
ages of dimensions of leaves from a collection
were used in the monograph, and these may be
compared with those of the new taxa. Mature
leaves of the collection of F. cordata (Fig. 15)
show an average lamina width of 52.5 mm, an
average lamina length of 53 mm, and a petiole
length of 49.7 mm. The only Fitchia which re-
sembles F. cordata both in form and dimensions
is the extreme population of F. rapensis from
the summit of Rapa. Leaves of F. cordata have
longer petioles than that collection, and there
is certainly no close relationship between F.
cordata and F. rapensis. Leaves of F. cuneata
ssp. tahaaensis (Fig. 16) which could be termed
mature show the following average dimensions:
lamina width, 48.0 mm; lamina width, 87.5
mm; petiole length, 26 mm. When compared
with the chart in the monograph (Carlquist,
1957: 50) the lamina dimensions fall in the
vicinity of F . nutans and some collections of
F. rapensis. The leaves are wider than those of
F. tahitensis but longer than those of the typical
F . cuneata. Thus, leaf shape distinguishes both
F . cordata and F. cuneata ssp. tahaaensis.

Histological features of leaves of the new
taxa also distinguish them. Fitchia cordata pos-
sesses fibers in bundle sheaths of many veins,

but they are lacking in sheaths of finer veinlets.
No secretory cavities are present in the meso-
phyll, although secretory canals occur adaxially
or abaxially to the veins, or both (Fig. 7).
Mesophyll is about 10 layers thick. These char-
acteristics are not matched by any other Fitchia
collection. Presence of a few fibers would ally
F. cordata to F. nutans or F. tahitensis, but
these two species possess secretory cavities. Ab-
sence of secretory cavities does characterize F.
speciosa and the typical F. cuneata, but these
taxa lack sclerenchyma in bundle sheaths.

Leaves of F. cuneata ssp. tahaaensis (Fig. 9)
lack sclerenchyma along all but the largest
veins. In addition to the secretory canals above
and below veins (Fig. 9, left), secretory cavi-
ties are present in the mesophyll (Fig. 9, right) .
The mesophyll is about 10 cells thick. This
description agrees closely with that of typical
F. cuneata except for the presence of secretory
cavities. Although such a difference might con-
ceivably arise from a difference in maturity of
a plant, it seems more likely a valid difference,
for specimens of both subspecies were of flow-
ering age.

INVOLUCRE: The heads of F. cordata are
small, and the involucre of a living specimen
would probably measure about 2 cm in diam-
eter— the smallest involucre size in the genus
except for F. tahitensis and F. cuneata. Involu-
cral bracts of F. cordata (Fig. 13) have a thick-
ness comparable to some bracts of F. nutans, but
differ in their lack of the abundant scleren-
chyma which characterizes bracts of F. nutans.
The mature state of the bracts of F. cordata
illustrated is certain, because they came from
a head in fruit which was in the process of
shattering. Occasional lignified thick -walled
cells may be seen along the inner face of the
bract and around the larger veins. In presence
of sclerenchyma and other histological charac-
teristics, the involucral bracts of F. cordata are
midway between those of F. nutans and F. cu-
neata. The receptacular bracts of F. cordata
(Fig. 14) show similar characteristics. As might
be expected, lignified thick-walled cells are more
frequent than in the involucral bracts.

Fitchia cuneata ssp. tahaaensis also has very
narrow involucres, like those of the typical F.
cuneata or F. tahitensis. Histologically, involu-

Studies in Fitchia— CARLQUIST and Grant

291

FIGS. 20-24. Figs. 20, 23, Fitchia cordata . Figs. 21, 22, 24, F. cuneata ssp. tahaaensis. Figs. 20, 21: Tips of
corolla lobes, showing exterior surface. Sclerenchymatous cells are shown in black. Figs. 22, 23 : Stamen tips,
showing inner surface. Anther sacs shown in outline, below. Fig. 24: Base of anther and adjacent filament.
Note caudate anther sac, right. All, X 65.

292

cral bracts of F. cuneata ssp. tahaaensis (Fig.
17) resemble closely the description given for
typical F. cuneata in the monograph. This re-
semblance is especially evident in the lack of
sclerenchyma. Only an occasional lignified cell
(usually near a vein) can be found. Collenchy-
matic thickening may occur near outer or inner
faces of the bract. In the section shown, more
than a single series of bundles may be seen;
some of these bundles have distorted orienta-
tion of xylem and phloem, a condition men-
tioned in the monograph for F. speciosa. This
condition is probably occasional among larger
bracts of several species. The thickness of the
involucral bract, as well as its lack of scleren-
chyma, identifies the two subspecies of F. cu-
neata closely with each other. The receptacular
bract (Fig. 18) of a head at anthesis certainly
lacks sclereids, although a greater degree of
lignification might be apparent in bracts of a
head in fruit.

ACHENES: Awn shape and anatomy proved
useful characteristics in distinguishing species
of Fitchia. In comparing illustrations of these
( Carlquist, 1957: plate 8) with these of the
new taxa (Figs. 8, 10) this also proves to be
true. In F. cordata (Fig. 8), the awns tend to
be rounded in transectional shape, like those of
F. nutans. However, they show a tendency to
be wider than those of F. nutans. They have a
single vascular bundle and, unlike at least some
awns of F. nutans, they lack a secretory canal.
Awns of F. cuneata ssp. tahaaensis (Fig. 10) are
like those figured for the typical F. cuneata in
all respects. Presence of three longitudinal zones
of trichomes tends to make them triangular as
seen in transectional view. Where such tri-
chomes are absent, awns may have a more
rounded shape.

COROLLA: In the monograph, considerable
attention was focused on the importance of
venation patterns of Fitchia corollas, both for
their phylogenetic importance within Compos i-
tae and for their usefulness in distinguishing
species. Corolla venation patterns are not illus-
trated here for the new taxa because they con-
tribute relatively few novel features and can
be referred to patterns illustrated in the mono-
graph. Although few flowers of F . cordata could
be studied in this respect (on account of the

PACIFIC SCIENCE, Vol. XVII, July 1963

ravages of the living plant by insects ) , the vena-
tion pattern of this species is substantially like
that figured for F. cuneata or F. tahitensis. There
is some variation in the levels at which adja-
cent lateral veins in each lobe join beneath
sinuses, but differences in these levels are not
nearly so extreme as the condition figured for
F. nutans (Carlquist, 1957: 10). In their rela-
tively short length (ca. 18 mm) the corollas
of F. cordata approximate those of F. tahitensis
also.

Corollas of F. cuneata ssp. tahaaensis have
similar size and venation features. Typically,
the patterns are the same as mentioned for F.
cordata. However, occasionally a pair of adjacent
laterals do not join beneath a sinus, but con-
tinue into the tube and achene without joining.
This condition does occur occasionally in the
other species, such as F. speciosa. A feature of
interest in the corolla of F. cuneata ssp. taha-
aensis is the presence of one or two supernu-
merary veins in each lobe. In addition to the
median and lateral veins, these fine supernu-
merary veins may be present for a short distance
near the base of the lobe. Such veins were not
seen in the typical F. cuneata, for which, how-
ever, mature corollas were not available. Super-
numerary veins are occasional in corollas of F.
speciosa, and characteristically abundant in those
of F. mangarevensis.

The occurrence of larger numbers of veins
in corollas of F. mangarevensis is particularly
interesting, since that species is close to F.
rapensis. Corollas, now available, of F. rapensis
confirm this relationship in the similarly elabo-
rate venation pattern (Fig. 26). This similarity
is shown in the presence of five veins per lobe,
rather than three. Such a condition is basic in
the construction of the F. rapensis corolla, al-
though additional veins or vein fragments may
be present. The outermost veins of each lobe
fuse at the tips of many corolla lobes. The co-
rolla venation of F. mangarevensis (Carlquist,
1957:14) is somewhat more complicated than
this, because although the five-vein condition
may be observed in some lobes in that species,
additional veins are more frequent. The pres-
ence of complex corolla venation in both F.
rapensis and F. mangarevensis is interesting in
that it raises the question of how complex

Studies in Fitchia— CARLQUIST and GRANT

293

venation may have been present ancestrally in
Fitchia . Is the more complex condition primitive
or a specialization? The question cannot be
answered in terms of the data available, and
cytological information would be very desirable
as a line of supporting evidence.

Distribution of thin-walled and thick-walled
trichomes and occurrence of sclereids on corolla-
lobe tips have proved useful features in the
systematics of Fitchia. In the two new taxa these
features are also of interest. Fitchia cordata ( Fig.
21) possesses lobe tips not identical to those of
any species previously figured. The frequency
of sclereids at the lobe tip is notable. Long
multiseriate trichomes, all the cells of which are
sclereids, are present. No trichomes composed of
nonlignified cells were observed. This condition
is most closely matched in the genus only in
the new subspecies of F. cuneata (Fig. 20). In
F. cuneata ssp. tahaaensis sclereids are rare or
absent at the lobe tips. The trichomes are long,
multiseriate (rarely unisef. fate ) , and composed
wholly of sclereids. This condition is not unlike
the condition in typical F. cuneata, except that
the trichomes in the Tahaa plants are few and
long.

The corolla-lobe tips of F. rapensis (Fig.
25) are, as might be expected, rather similar
to those of F. mangarevensis (Carlquist, 1957:
27 ) . They are different in that in F. rapensis the
sclerified trichomes are slightly longer, in gen-
eral, and extend farther down the lobe. Oc-
casional thin-walled hairs may be seen on the
terminal portion of lobes of F. rapensis, but
the extremely dense coating of thick-walled lig-
nified trichomes clearly marks this pattern as
closest to that of F. mangarevensis.

STAMENS: Stamen tips likewise offer taxo-
nomic criteria. The long stamen tips of F. cor-
data (Fig. 23) exceed those of F . nutans in size,
and are thus the longest of the Society Islands
species of Fitchia. The stamen tips of F. cuneata
ssp. tahaaensis (Fig. 22) are longer than those
of the typical F. cuneata, but do not differ mark-
edly from those of F. tahitensis or F. nutans.

The stamen tips of F. rapensis (Fig. 27) are
1.5 mm long, thus matching those of F. man-
garevensis. This is particularly interesting be-
cause the two species are so much alike in this
respect and because they have longer stamen

tips than the other species of the genus, with the
exception of F. speciosa.

A feature of some interest not previously
observed is the occurrence of somewhat caudate
anther sacs in F. cuneata ssp. tahaaensis (Fig.
24) and the other taxa of Fitchia. In view of
Cronquist’s (1955) dictum that anthers of Com-
positae are primitively tailless, this would seem
curious, because Fitchia possesses so many ana-
tomical and morphological characteristics which
appear primitive for the family. Cronquist’s
dictum, for which no appreciable evidence is
adduced, seems highly questionable.

In transection the anthers of F. cuneata ssp.
tahaaensis (Fig. 19) exhibit a feature of interest.
One or two secretory canals may be observed in
the connective of each anther. This condition
occurs sparingly in F. speciosa, but was not
observed in other species of Fitchia. The oc-
currence of anther secretory canals has been
noted in other genera which may be related to
Fitchia, such as P etrohium and Oparanthus
(Carlquist, 1957).

STYLE: Like the anthers, the style of F. cuneata
ssp. tahaaensis (Fig. 19) seems to exhibit more
numerous secretory canals than those of other
species of Fitchia. One canal (or occasionally a
pair) is present exterior to each of the four
style bundles. Liquid-preserved material is re-
quired for accurate demonstration of this phe-
nomenon. In the only other species for which
such material was available, F. speciosa , canals
were much less abundant in styles (Carlquist,
1957 ) . However, as indicated in the monograph,
many other Heliantheae do show abundance of
stylar secretory canals.

pollen GRAINS: The drawings of pollen
grains of Fitchia (Figs. 28-30) show some dif-
ferences in representation compared with those
of the monograph. Although the outer sculp-
tured layer (ectosexine) is composed of fine
rods and spaces, as illustrated earlier, a lacunose
inner sculptured layer (endosexine) has now
been observed. The endosexine consists of rods
interspersed in large spaces. In addition, an
inner and outer layer of nexine (below in each
figure) may be distinguished. Size and spine
shape are of especial interest, however, in com-
parison of the species. The markedly blunt
spines of F. cuneata ssp. tahaaensis (Fig. 29)

294

PACIFIC SCIENCE, Vol. XVII, July 1963

Figs. 25-27. Fitchia rapensis. Fig. 25: Tip of corolla lobe, showing exterior surface; sclerenchymatous cells
in black. Fig. 26: Corolla, showing venation. Fig. 27: Stamen tip, showing inner surface. Figs. 25, 27, X 65.
Fig. 26, X 8.

Studies in Fitcbia—C ARLQUIST and GRANT

match those of the typical F. cuneata ( Carlquist,
1957:32). In F. cordata (Fig. 28) the spines
ate even more markedly blunt. Pollen grains
of F. cordata (range, 39-51 u; average, 42 fx)
are the smallest in the genus. The pollen grains
of F. cuneata spp. tahaaensis (range, 45-51 fi;
average, 48 /*) closely match those of ssp. cu-
neata in size. The slightly smaller size of grains
in the latter may be caused by the fact that
flowers of F. cuneata ssp. cuneata from which
grains were taken were not quite mature. Pollen
grains of F. rapensis (Fig. 30) average 60 u
in diameter, and have a range from 51 to 66 /x.
They are thus slightly larger than those of F.

295

mangarevensis, although the difference is not
significant. The spines of pollen grains of F.
rapensis have an inverted-funnelform shape, like
those of F. mangarevensis, but appear to be
more sharply pointed. In addition, lacunae in
spine tips were observed for the first time in
the genus in F. rapensis. This feature, however,
is not unexpected in Fitchia, since other Helian-
theae have lacunae in spine tips (Carlquist,
1957:33).

SPECIES CHARACTERISTICS: In the mono-
graph three species groups were recognized:
( 1 ) F. speciosa; (2) F. nutans, F. tahitensis,
and F. cuneata; and (3) F. mangarevensis and

Figs. 28-30. Optical sections of equator of pollen grains, showing a third of each grain. Germ pores indi-
cated at left and right in each. Fig. 28, F. cordata. Fig. 29, F. cuneata ssp. tahaaensis. Fig. 30, F. rapensis.

296

PACIFIC SCIENCE, Vol. XVII, July 1963

Figs. 31-32. Type specimens of Fitchia. Fig. 31, F.cordata. Fig. 32, F. cuneata ssp. tahaaensis.

Studies in Fitchia — Carlquist and Grant

F. rapensis. The new taxa clearly belong to the
second group. The present study also emphasizes
the distinctness of the third group and the close
relationship between the two species it contains.

Fitchia cuneata ssp. tahaaensis agrees with
typical F. cuneata in most respects. Those by
which it differs include: presence of sclerified
tyloses in wood (not yet observed in wood of
typical F. cuneata ) ; leaves with wider and longer
laminae; secretory cavities present in leaf meso-
phyll; involucral bracts thicker; sclerified tri-
chomes on corolla-lobe tips longer and fewer;
anther tips longer. These differences are rela-
tively minor. Close identity of the two sub-
species is revealed by such critical features as
general leaf shape, pith structure, bract anatomy,
corolla vascularization, awn shape and anatomy,
and pollen size and ornamentation. The two
taxa are therefore best regarded as subspecifically
different. This situation is precisely what one
would expect on account of the closeness of
Tahaa to Raiatea, sister-islands (within a com-
mon fringing reef).

The nearest island to the sister-islands of
Raiatea is Bora Bora. Qne might expect, there-
fore, a similarity between F. cuneata and F.
cordata. These two do appear similar in their
small heads, relative lack of sclerenchyma in
involucral bracts, venation and size of corollas,
lack of secretory cavities in leaves (excepting
ssp. tahaaensis ) and blunt shape of spines on
pollen grains. They are dissimilar in that F.
cordata possesses secretory canals and thin- walled
nonlignified cells in the pith, has cordate leaves,
and some sclerenchyma along veins of leaves.
These characteristics are reminiscent of F. nu-
tans. From F. nutans , however, F. cordata differs
by such respects as those in which it resembles
F. cuneata. Features of F. cordata which do not
match those of any species in the F. nutans-F.
cuneata-F. tahitensis group include the presence
of nonlignified cells in pith, the small leaf size
and cordate shape combined with relatively long
petiole length, the presence of both long mul-
tiseriate sclerified trichomes and sclerified epi-
dermal cells on corolla-lobe tips, long stamen
tips, small pollen size, and blunt spines on
pollen grains. These characteristics, combined
with the distinction furnished by its geographi-
cal isolation on Bora Bora, mark F. cordata as
worthy of recognition as a new species.

29 7

PHYLOGENETIC CONSIDERATIONS: The new
taxa described here, although they contribute
no anomalous features to the genus, do enlarge
our picture of it and permit more detailed ob-
servations in this regard. The monograph of the
genus provided some notions on the relation-
ships of Fitchia. These ideas do not seem to
require alteration. The new taxa do enhance the
diversity of the Society Islands Fitchias. Indeed,
Papy ( 1955 ) notes that the Society Islands form
a center of diversity for Fitchia, and that the
variability of Fitchia on Tahiti is suggestive of
this. The latter phenomenon remains a problem
worthy of study when more material from Tahiti
becomes available.

Postulation of phylogenetic trends within
each species group seems inadvisable, both
because of the limited quantity of material
known and the fact that the species seem to
be variations on a basic theme rather than stages
along a phylogenetic pathway. However, some
interpretations of relationship among the species
groups may be offered here.

The larger size — in all parts of the plant —
of F. speciosa, as compared with the remaining
species, seems specialized. The larger pollen size
suggests the possibility of a greater chromosome
number, and the morphology of the awn base
seems clearly a specialized feature. The fact that
F. speciosa has such an isolated geographical
position, the westernmost species of Fitchia, in
contrast with the likelihood of an American
ancestry of this helianthoid (e.g., Brown, 1935;
Papy, 1955), reinforces this supposition. The
extremely large seeds of F. speciosa seem less
adapted to long-distance dispersal than the
smaller seeds of the Society Islands species. The
most likely interpretation, seemingly, is that
seed size, and the peculiar adnation of achene
summit to awn base in F. speciosa are charac-
teristics acquired subsequent to arrival of its
ancestors in Rarotonga. Fitchia speciosa may be
regarded as a highly distinct derivative of the
Society Islands stock. Although corollas of F.
speciosa are the largest in the genus, their vena-
tion is like that of the Society Islands species,
not that of F. rapensis and F. man gar even sis,
which have complex corolla venation.

If pollen-grain size is an indication of chro-
mosome number, then one would expect the
Society Islands species to have the lowest num-

298

PACIFIC SCIENCE, Vol XVII, July 1963

ber, and progressively higher chromosome num-
bers would be indicated for the F. rapensis-F.
mangarevensis group and F. speciosa. If this
were true, and it were a criterion of phylogeny
within the genus, the complex floral venation
of F. rap en sis and F. mangarevensis would be
a specialization. Information on comparative
cytology within the genus, in any case, is highly
desirable.

REFERENCES

Brown, F. B. H. 1933. Flora of southeastern
Polynesia, III. Dicotyledons. Bishop Mus.
Bull. 130:1-386.

Carlquist, S. 1957. The genus Fitchia (Com-
positae). Univ. Calif. Publ. Bot. 29:1-144.

1958. Wood anatomy of Heliantheae

(Compositae). Trop. Woods 108:1-30.

Cronquist, A. 1955. Phylogeny and taxonomy
of the Compositae. Amer. Midi. Nat. 53:478-
511.

Erdtman, G. 1952. Pollen morphology and
plant taxonomy. Chronica Botanica, Waltham,
Mass.

Hemsley, W. B. 1885. Report on present state
of knowledge of various insular floras. Bot.,
Challenger 1 ( 1 ) : 1-75.

Hoffmann, O. 1890. Compositae. In: Engler
and Prantl, Die natiirlichen Pflanzenfamilien
4(5) : 87-391.

Papy, H. R. 1955. Tahiti et les lies voisines. La
vegetation des lies de la Societe et de Makatea.
2e partie. Trav. Lab. Forest. Toulouse 5(2/1:
3): 162-386.

Riley, L. A. M. 1926. Notes on the flora of
Rapa. Kew Bull. 1926:51-56.

Lithoglyptes hirsutus (Cirripedia: Acrothoracica),
A New Burrowing Barnacle from Hawaii

Jack T. Tomlinson1

Two samples of coral from Kaneohe Bay,
Oahu, Hawaii, have each yielded a number
of specimens of a new species of acrothoracican
burrowing barnacle of the family Lithoglypti-
dae. Samples of Psammocora v err mm Vaughan
collected by Stephen A. Wainwright,2 and of
Porites compressa Dana collected by Charles
Stasek,3 were referred to me by William A,
Newman.3 These barnacles are the first repre-
sentatives of the order Acrothoracica known
from Hawaii.

Family LITHOGLYPTIDAE Aurivillius 1892

Lithoglyptidae emend. Tomlinson and New-
man I960.

Mouth cirri well developed, on a 2-jointed
pedicle; 4-5 pairs of terminal cirri, but if only
4 pairs, caudal appendage present; no gut teeth
or gizzard in digestive tract; with adhesive disc
on mantle; lateral bar absent; burrows in coral
or mollusc hard parts.

Lithoglyptes Aurivillius 1892 {emend.)

Four pairs of terminal cirri on a 2-jointed
pedicle with oblique sutures at first joints; cau-
dal appendage present; mouth cirrus with 2
rami of 3 to 6 articles (5 to 6 in original
description ) .

Lithoglyptes hirsutus n. sp.

DIAGNOSIS: Female (Figs. 1-7): Mantle
aperture strongly arched on both sides of an
acute projecting tip which is not a movable

1 Biology Department, San Francisco State College,
San Francisco 27, California. Manuscript received
December 19, 1961.

2 Zoology Department, University of California,
Berkeley 4, California.

3 Scripps Institution of Oceanography, La Jolla,
California.

spine or hook; aperture length exceeds Vi of
mantle width, aperture armed with numerous
teeth and long flexible hairs, especially on the
outer edge of the lip area; anterior and pos-
terior rami of mouth cirri with 5 and 3 articles,
respectively; caudal appendage with 2 seg-
ments; head with acute projection opposite
mouth area; burrow pointed oval in surface
view. Holotype 1.2 X 0-67 mm; about 30 dried
specimens in Psammocora verrilli from a depth
of 3-6 ft on Sand Bar Reef and in Porites com-
pressa from NE side Checker Reef, Kaneohe
Bay, Oahu, Hawaii. The species is named for
the presence of numerous hairs on the mantle
aperture.

TYPE MATERIAL: Holotype USNM 107544.
Paratype material: San Francisco State College,
San Francisco, California; California Academy
of Sciences, San Francisco, California; Plymouth
Laboratory, England; Seto Marine Biological
Laboratory, Japan; Portobello Marine Station,
New Zealand.

DESCRIPTION: Female: The mantles of 16
adults averaged 1.95 mm in height x 1.18
mm in width (maximum 2.1 mm and 1.6 mm),
and are flattened laterally. The mantle is studded
with numerous small T-shaped teeth, short
spinules, and three- or four-pointed star-shaped
teeth, all scattered about on the surface, but
particularly near the aperture. The mantle at-
tachment disc adheres strongly to the burrow,
but is readily removed if the coral is decalcified.

The average length of the slitlike aperture
of 12 specimens is 0.97 mm. Near the anterior
end of the aperture two acute projections ex-
tend to a point, but are not movable or hooked.
The outer edges of the thickened apertural
area are heavily set with teeth and long hairs.
The inner edges have fewer teeth and hairs,
but have a row of comblike projections which
extend into the aperture and tend to occlude
it. This structure has been called the "comb-
collar” by previous workers.

299

FIGS. 1—7. 1, Lithoglyptes hirsutus, n. sp., female, side view of adult removed from burrow; abbreviations:
AC, anal cirrus; AK, attachment knob; CC, comb collar; H, head; L1M, lip of mantle aperture, inner margin;
LOM, lip, outer margin; MC, mouth cirrus; AIP, mouth parts; TC, terminal cirri; TO, tooth.

2-4, Mouth parts of Lithoglyptes hirsutus, n. sp., female; 2. mandible with palp; 3, first maxilla; 4, second

maxilla.

5-7, Lithoglyptes hirsutus, n. sp., female; 5, middle portion of fourth (last) terminal cirrus, side view;
6, burrow in coral, surface view; 7, mouth cirrus, slightly foreshortened in comparison with some specimens.

Lithoglytes hirsutus — Tomlinson

301

The mouth parts (Figs. 2-4) are typical for
the genus, being composed of a pair of man-
dibles with palps and two pairs of maxillae.
Each mandible has four strong teeth, the inner
one bearing short bristles. The mandibular palp
exceeds the mandible in length; with long, soft
bristles on the edges near the tapering point.
Each first maxilla is armed with two strong
teeth, numerous bristles, short teeth along the
cutting edge, and has the usual apodeme. The
second pair of maxillae are large and soft, and
have numerous flexible bristles distributed along
their edges. These close-set appendages serve
as posterior limits to the mouth field.

The mouth cirri ( Fig. 7 ) have a two-jointed
pedicle upon which the two bristle-bearing
rami articulate. They arise below the mouth
parts but terminate above them, frequently
curving outward at the tips. The anterior ramus
has five segments and the posterior ramus has
three segments. The segments of the rami are
equipped with numerous bristles, which make
the determination of the number of segments
very difficult.

There are four pairs of biramous, multiseg-
mented terminal cirri, on two-segmented pedi-
cles. The slanted articulation between these
pedicular segments is characteristic of the genus.
The rami are armed with single long setae on
the outer edges of every second to sixth artic-
ulation. The setae on the inner edge of the
terminal cirri alternate between pairs of long
setae at the articulation and pairs of shorter
setae toward the middle of the segment

(Fig. 5).

A pair of two-jointed uniramous caudal ap-
pendages occurs at the posterior end of the
body. The total length of the caudal appendage
is about Vi the length of the pedicle of the
posterior terminal cirrus.

The burrow of the female appears in the sur-
face view as a pointed oval (Fig. 6). It is
deeper than it is long. The walls are nearly
perpendicular to the surface, and rounded at
the bottom. The depth of the burrow appears

to be slightly more than the greatest diameter
of the animal in dried specimens. It is assumed
that the living animal closes the burrow open-
ing with the hardened, hairy apertural area
of the mantle.

MALE: Dwarf males resembling cyprid lar-
vae were seen, but being dried they were much
distorted, and whether or not they were truly
mature stages could not be determined. Of the
11 males studied, none had the very reduced
baglike shape of many acrothoracican males,
nor the attenuated stalk of the male of Lit ho -
glyptes spinatus. They were of a size and shape
one would expect of a cyprid larva which had
lost its bivalved carapace. The presence of a
penis could not be ascertained.

L. hirsutus is the first member of the genus
to be found in the central Pacific. Of the other
species in the genus L. indicus Aurivillius, L.
ampulla Aur., and L. hicornis Aur. are from
the Java Sea and Indian Ocean, and L. spinatus
Tomlinson and Newman is from Jamaica.

L. hirsutus can be distinguished from L. bi-
cornis, L. ampulla , and L. spinatus by the ab-
sence of the apertural hooks and/ or spines
borne by the latter three species. This new
species may be distinguished from L. indicus,
which also lacks the hooks and spines, by the
presence of the "hairs” on the mantle aperture
and by its small size (2 X 1 mm rather than
6X^ mm).

REFERENCES

Aurivillius, C. W. S. 1892. Neue Cirripeden
aus dem Atlantischen, Indischen und Stillen
Ocean. Ofversigt af Kongl. Vet.-Akad. Forh.
Stockholm 49 (3): 133-134.

Tomlinson, Jack T., and William A. New-
man. I960. Litho glyptes spinatus, a burrow-
ing barnacle from Jamaica. USNM Proc.
112 (3443): 517-526.

Toxicity of Dialyzed Extracts of Some California
Anemones (Coelenterata)1

Edgar J. Martin2

ABSTRACT: Anemones of the California coast, collected from November I960
to March 1961, were compared with respect to their toxicity. Aqueous extracts
of the anemones were dialyzed and assayed by intraperitoneal injection to mice,
and the survival times of the injected mice were compared.

The extracts of Anthopleura elegantissima and A. xantho gram-mica were by far
more toxic than those of Metridium senile, Corynactis calif ornica, Tealia crassicornis,
T. lofoiensis, and T. coriacea.

The differences in potencies of the extracts may indicate species variations in
( 1 ) toxicity, (2) efficiency of the extraction method, or ( 3 ) both.

The AQUEOUS extracts of many coelenterates
are poisonous (Dodge, 1957). The poisons are
mixtures of biologically active compounds ( Cre-
scitelli and Geissman, 1962), some of them
amines (Welsh and Prock, 1958), others heat-
labile colloids (Martin, I960; Mathias et al.,
I960). The poison occurs in both the nema-
tocysts and the nematocyst-free structures of
the anemones (Lane, I960; Martin, to be pub-
lished ) . Both the chemical composition and the
biologically active substances of extracts vary
from one species of anemone to another (Ma-
thias et al., I960; Bergmann et al., 1956). The
present study was designed to compare the
lethality in mice of dialyzed extracts of some
anemones of the California coast.

MATERIAL AND METHODS

SPECIES AND SITES OF COLLECTION: Ane-
mones were collected at various sites on the
California coast from November I960 to March
1961: Anthopleura elegantissima from Bodega
Bay, Tomales Bay, Monterey, and Corona del
Mar; A. xanthogrammica from Tomales Bay and

1 Supported in part by National Institutes of Health
Grant RG 7626. Manuscript received February 19,
1962.

2 Laboratory of Comparative Biology, Kaiser Foun-
dation Research Institute, Richmond, California; and
Pacific Marine Station, University of the Pacific, Dil-
lon Beach, California.

Monterey; Metridium senile from Monterey;
Corynactis calif ornica, Tealia lofotensis and T.
crassicornis from Bodega Bay, all intertidal; and
T. coriacea off Marineland from a 30-ft depth.
In addition, the intertidal Nudibranch Diaulula
sandiegensis and the Mollusca Crassostrea gigas
and Mytilus calif ornianus were collected at To-
males Bay. Anthopleura elegantissima is com-
mon all along the California coast. Its extracts
were used as the base of comparison with those
of the other species available.

The anemones were placed in the laboratory
in aerated aquaria, with sea water which was re-
newed twice a week. Water temperature varied
between 13 and 20 C. The anemones were ex-
posed to the light of the laboratory. They re-
ceived no food supplement. Specimens which
did not show normal vitality in the aquaria were
discarded. After 2-4 weeks the anemones were
removed from the aquaria and cleaned from ad-
herent objects. During this manipulation they
contracted firmly. Their body wall was then
punctured to empty pockets of trapped water.
Then the anemones were rinsed with distilled
water for 10-15 sec, blotted with mild pres-
sure, weighed in air, and their volume deter-
mined. The specimens weighed from 0.3 to
34.2 g.

EXTRACTION: The anemones were homoge-
nized with three times their volume of distilled
water at high speed in a Waring blender for

302

Toxicity of Anemone Extracts — Martin

303

two 1-min periods. After being centrifuged for
30 min at 1000 g the supernatant was decanted
and centrifuged again. The resulting superna-
tant extract was turbid but did not contain cells
or debris. It was either decanted or, when it
was found covered with a lipid layer, removed
with syringe and needle; then its pH was deter-
mined. The extract was dialyzed for 4 hr through
commercial cellophane tubing against ten times
its volume of 0.001 M sodium phosphate (pH
6.2) in 0.8% sodium chloride with 1% acti-
vated charcoal (Weinberger, 1936). The dial-
ysis was repeated once with fresh solution. All
operations were performed at room temperature.

Extracts of the other lower metazoa were pre-
pared in the same manner. All extracts were as-
sayed immediately after their preparation. The
potency of extracts of each species was com-
pared with that of an extract of A. elegantissima
prepared on the same day.

ASSAY: The extract was given intraperi-
toneally to DAL-Swiss albino mice weighing
17 to 23 g. The mice were placed in cages in
lots of four to eight (Russell et al., I960), and
observed for 24 hr; their survival time was
recorded. The abdominal cavities of the mice
were inspected post mortem and mice with
intraperitoneal hemorrhage were discarded. This
condition was found in animals that showed
severe pain reaction immediately after the in-
jection and died within 4 min. The discarded
animals were replaced by supplementary ones
to complete the series. Each extract was as-
sayed on a series of at least four mice.

EXPERIMENTS AND RESULTS

Preliminary experiments explored the effect
of varying doses of extracts of A. elegantissima
on the survival time of the injected mice. Only
the area of LD<)<* was considered. A dose effect
curve showed that with decreasing doses the
survival time increased. At the dose of 20 cc
of extract per kilo of mouse, both the range
and the mean of the survival time were within
practical limits; therefore, this dose was se-
lected as a convenient reference.

The pH of all the extracts varied from 6.0
to 6.6.

At a dose of 20 cc of extract of A . elegantis-
sima per kilo of mouse, the mean survival time
of the injected mice varied from 8 to 36 min
among the ten series tested. Between some of
these series the difference in survival time was
significant at the 2% level by the Chi-square
test. The cause of these differences is unknown.
The differences showed no correlation with vari-
ations in the size, sites of origin, or dates of
collection of the specimens, or with the pH
of the extracts. They were not relevant for the
present study.

Table 1 shows that the potency of extracts
may vary from one species of anemone to
another. The highest potency, as estimated by
the survival time of mice after intraperitoneal
injection, was found for the extracts of A. ele-
gantissima and A. xanthogrammica. Extracts of
other anemones were far less potent even when
their dose was tripled. With extracts of T. lofo-

TABLE 1

Comparison of the Survival Time of Mice after Intraperitoneal Injection

of Extracts of Anemones

SPECIES

NO. OF

BATCHES

NO. OF

MICE

DOSES CC OF

extract/kg

OF MOUSE

SURVIVj

Mean

\L TIME IN

INUTES

Range

PROPORTION
OF MICE
SURVIVING
24 HR

Anthopleura elegantissima

15.7

5-47

1/48

A. xanthogrammica

9.2

6-15

0/10

Metridium senile

174.1

80-240

0/6

Corynactis calif or nica

138.5

118-180

0/5

Tealia crassicornis

342.8

252-480

0/6

T. lofotensis

336

3/4

T. lofotensis

460-540

2/4

T. coriacea

264-640

2/4

T. coriacea

480-900

2/4

304

PACIFIC SCIENCE, Vol. XVII, July 1963

tensis and T. coriacea it was not possible to

obtain an LD99.

The extracts of the Nudibranch Diaulula
sandiegensis and the Mollusca Crassoslrea gigas
and Mytilus calif ornianus, each assayed on six
mice at the dose of 60 cc extract per kilo of
mouse, caused transitory depression of activity
of the injected mice, but not death.

The extracts of all anemone species lost their
toxicity when they were heated at 90 C for
20 min.

DISCUSSION

The storing of the anemones alive for 2
weeks or longer before preparing the extracts
was intended to minimize the amount of pos-
sibly toxic products from food ingested by
them. The dialysis of the extracts was aimed at
minimizing their content of biologically active
amines. The nature of the non-dialyzable tox-
ins is not known. They may be proteins similar
to those demonstrated by various authors in
other coelenterates ( Mathias et ah, I960; Far-
ber et al., 1961).

We found information concerning the chem-
ical components of the anemones here investi-
gated only in the literature on A. elegantissima
(Bergmann and Landowne, 1958). In general
the lipid content of coelenterates Is high, and
the proportions of various lipids and soaps
and their chemical nature vary from one species
to another (Bergmann et al, 1956; Bergmann
and Landowne, 1958). The solubility of pro-
teins is affected by lipids, and if it is assumed
that the poisons here studied are proteins, it can
be speculated that the solubility of these poi-
sons may vary from one species to another.

The data of Table 1 suggest that the toxicity
of anemones may vary from species to species.
This interpretation is supported by the great
difference between species with respect to po-
tency of extracts, which was observed even
when the dose of the weaker extracts was
tripled. But it is also possible that the efficiency
of the extracting method, and hence the poison
content of the extracts, may vary from species
to species. This consideration applies equally
to our study and to other authors’ reports of
species variation in toxic compounds. These
causes for variation may coexist.

Lethal effects on mice were observed with
extracts of the anemones but not with extracts
of the Nudibranch and the two bivalves. This
suggests that toxicity Is not a property of all
littoral lower metazoa.

REFERENCES

Bergmann, W., S. M. Creighton, and W. M
Stokes. 1956. Marine products. XL. Waxes
and triglycerides of sea anemones, j. Org.
Chem. 21:721-728.

— — — and R. A. Landowne. 1958. Marine
products, XLVI. Phospholipids of a sea anem-
one. J. Org. Chem. 23:1241-1245.
Crescitelli, F., and T. A. Geissman. 1962.
Invertebrate pharmacology: Selected topics.
Ann. Rev. Pharmacol. 2:143-192.

Dodge, E. 1957. A study of the contents of
the nematocysts of Metridium senile (L.),
Thesis. University of Washington, Seattle,
Wash. 63 pp.

Farber, L., and P. A. Lerke. Preliminary ob-
servations on the toxic property of the sea
anemone, Rbodactis howesii. In: Abstracts of
Symposium papers, 10th Pacif. Sci. Cong.,
Honolulu, p. 447.

Lane, C. E, 1961. Pbysalia nematocysts and
their toxin. In: H. M. Lenhoff and W. F.
Loomis, eds., The Biology of Hydra and of
Some Other Coelenterates. U. Miami Press,
Coral Gables. Pp. 169-178.

Martin, E. J. I960. Observations on the toxic
sea anemone, Rbodactis howesii. Pacif. Sci.
14:403-407.

Mathias, A. P., D. M. Ross, and M. Schach-
TER. I960. The distribution of 5 -hydroxy ~
tryptamine, tetramethyl ammonium, homa-
rine and other substances in sea anemones.
J. Physiol. 151:296-311.

Russell, F. E., J. A. Emery, and B. G. Bow-
ers. I960. A comparison of mouse and chick
in toxicity determination. Toxicol. AppL
Pharmacol. 2:558-563.

Weinberger, E. 1936. Improvement on dial-
ysis. Brit. Patent No. 452,000.

Welsh, John J., and P. G. Frock. 1958. Qua-
ternary ammonium bases in the coelenter-
ates. Biol. Bull. 115:551-561.

Ecological Observations on the Sea Cucumbers Holothuria atra
and H. leucospilota at Rongelap Atoll, Marshall Islands

Kelshaw Bonham and Edward E. Held1

In a distributional study of the black sea cu-
cumbers, Holothuria atra (Jager) and H. leu-
cospilota ( Brandt ) , at Rongelap Atoll ( Fig. 1 )
the beaches of most of the larger islets were
surveyed during September 1959, March 1961,
and September 1961. Estimates of abundance
and size were made, and observations on tem-
perature tolerance, predation, asexual reproduc-
tion by fission, and some other ecological aspects
were considered.

These two species of sea cucumbers are fre-
quently, although not always, found together in
the intertidal area of the seaward sides of the
islets at the edge of the reef-flat nearest shore.
They occupy pools of water from a few centi-
meters to about one-half meter deep at low tide
(Figs. 2, 3), or are occasionally entirely exposed.
H. atra usually covers itself with a coating of
light coral sand held in place by tube feet, al-
though occasionally it is naked and black. The
integument is thick and firm. This species lacks
the Cuvierian organ and thus may be handled
without the annoyance of adhesive threads. H.
leucospilota is always naked and almost black,
has a softer integument with fine protuberances
giving it a prickly appearance. The intricate,
microscopic calcareous deposits in the integu-
ment differ markedly in the two species: in H.
atra (Fig. 4 A) the "tables” are tall and there
are no "buttons,” while in H. leucospilota (Fig.
4B ) the tables are short and squat, and there are
many buttons.

H. atra also occurs in deeper water in the
lagoon, where it does not coat itself with sand,
and to an undetermined extent on the vast areas
of reef-flat between islets. At high tide the water
at the seaward shores of the islets usually covers
both species to a depth of I-IV2 m, and they

1 University of Washington, Laboratory of Radia-
tion Biology, Fisheries Center, Seattle 5, Washington.
Manuscript received December 20, 1961.

are then frequently subjected to wave action
from the surf whose initial force has been
broken by the outer reef. A conspicuous feature
of this habitat is the churning agitation and
aeration of the water. H. atra appears to main-
tain its position by inhabiting depressions and
holding onto the relatively smooth reef-flat by
means of its tube feet. H. leucospilota usually
anchors the posterior portion of its body under-
neath a stable rock or in a hole in the reef floor.
In the absence of other cover it may find its way
under a cluster of H. atra (Fig. 3). At appro-
priate times it extends its highly mobile and
extensible anterior end outward for feeding.

Concentrations of these two species almost
always occur where large slabs of beachrock are
situated on the lower edge of the sandy beach
slope bordering the inner edge of the reef-flat,
and where, after the recession of the tide, the
sea water may be observed to issue from the
sand onto the reef-flat below the slabs of rock
until the next tide. Below a sandy beach without
slabs of rock no such prolonged runoff is evi-
dent, and the sea cucumbers usually are few or
lacking. It is in the pools fed by the relatively
cool water from underneath the rock slabs that
H. atra and H. leucospilota thrive. Concentra-
tions of sea cucumbers were occasionally seen
in the absence of slab-rock, and rarely slab-rock
was found without sea cucumbers. They occupy
an ecological niche almost devoid of other
macroscopic organisms; occasionally small snails
occur, but frequently the sea cucumbers are
alone. There is no obvious food in the sand they
ingest.

CENSUSING OF SEA CUCUMBERS

The surveying procedure consisted of observ-
ing, photographing, and taking notes on the
abundance of Holothuria while walking around

305

306

PACIFIC SCIENCE, Vol. XVII, July 1963

the periphery of 16 islets completely, and 5
others partially, thus covering 67% of the sea-
ward shores of all islets in the atoll. The islets
are numbered clockwise on the map of Rongelap
Atoll shown in Figure 1 starting at the north-
west extremity. Only about 2 % of the inter-islet
reef area and shallower lagoon bottom regions
supposedly suitable for H. atra was censused.

Distances were measured by chain on the
seaward shores of four islets, Aniiru, Kabelle,
Namoen, and Rongelap, while at other islets
distances were either determined by pacing, or,
in most cases, estimated. On the east rim of the
atoll high-altitude vertical, and elsewhere low-
altitude oblique, aerial photos were employed to
aid in orientation. Photographs taken on the
reef-flat of undisturbed sea cucumbers were
sometimes used in the estimations of abundance.

The places of concealment under overhanging
ledges were inspected, but rocks were seldom
lifted to check for specimens.

Table 1 shows the distribution of H. atra and
H. leucospilota at the localities surveyed. Section
numbers representing 100-m units of seaward
shoreline appear under the appropriate fre-
quency group. The central frequency groups in-
crease by a factor of 10 while smaller ranges of
numbers delimit the extremes. Along with the
range in the column headings are the means in
parentheses— geometric except for the first,
where the zero limit precludes use of the geo-
metric mean. Geometric and arithmetic means
were tested for agreement with totals of the
last two columns showing sums by localities
taken directly from a previous tabulation, which
also supplied the data for the body of Table 1.

Sea Cucumbers at Rongelap Atoll— Bonham and Helb

307

TABLE 1

Distribution of H. atra and H. leucospilota on the Seaward Shores of Islets
by 100-m Section Numbers

(The body of the table shows section numbers in which the abundance of sea cucumbers fell within the range
indicated in the column heading. Sections for each islet were numbered clockwise starting at the counterclock-
wise extremity (left-hand end when facing seaward) of the seaward shore.)

Islet

Dat

100-m

section number

Remarks

Total individu-
als observed
or estimated

Name

it)

Q«

Nun

0-2

(U*

nbers 0

3-10

5.62)

f H. at

11-100

(33.2)

-a

101-

1000

(318)

1001-

3000

(1730)

Numbe

0-2

(U*

rs of :

3-10

(5.62)

H. leuc

11-100

(33.2)

ospili

10J,-

1000

(318]

Dta

1001-

3000

(1730!

R. atra

U- imissLz

SPilota

Naen

15,

1-5

W. Yugui

Pigani-

yaroyaoro

1-4

E. shore, many

210

2000

Aerik

1-3

1-2

12 atra on W. shore 12

E. Yugui

1-4

2-4

8 atra on E, 16 on

W. 24

shore

Lomuilal

1-5

Geien

1-4

13 Inner

(10,000 atra in lagoon.

I. 13 6

i I. 15)

10000

13 Outer

240

18-

Reef area

(700 meters north of Aniiru I. had 83,000 atra, 2600 leuco.and 4300 Stichopus

83000

2600

chloronotus)

Aniiru

1-2

200

60 j

Kabelle

10-

3,8,9

0,1,4,

(10)

2-10

0, 1

200 atra & 100

3000

150

leuco. in N.E.

channel

Namoen

4 leuco. S. side

2 300

15 |

Mellu

2-5

1, 15

6-9,13,

10-12 ,

1-23

3 atra, 3 leuco., E

3500

14,16-

19,20,

channel

18,21

22,23

Gog an

1-8

3-6

1-2,

8 atra, E. channel.

330

7-8

Slab rock at 1-3 & !

Kieshi-

1-6

600

echi

Eniaetok

2-3,

1.12,

4,9

11, 14

1-9,

10,14

5 atra & 1 leuco.

3600

5-8

13,15-

11-13

N. side

15-23

Rongelap

1-2,

3-5,

6, 7,15,

28-30,

1-24,

25,41

46-48,

some atra, lagoon

3400

990

11,

8-14,

36, 42,

22,27,

39-41,

26-40,

44,45

50-52,

side

20/

16-21,

45,55,

37,43,

44,46 ,

42-43,

49,56

54,55,

31-35,

57,61,

47-52,

53,67-

63,66

57-62,

37,53,

66,71

54,58-

71,83,

72,74

64,65,

67-69,

60,62-

84,88-

73,75,

83-92

65,70,

77-79,

72-79,

80-82,

81,82

85-87

Arbar

1, 2,'

3-5,

1-26

27-29

6-12,

13-26

27-29

Tufa

200 atra , lagoon ,

mid-islet only, seen

Burok

1-12

1 atra, lagoon sidel

| 60

Pokoreppu

‘ " 1

Total sections

160

(Total 110421

6327)

‘Arithmetic mean. Means in other columns are geometric.

308

Better agreement resulted from the use of geo-
metric than from arithmetic means.

Analysis of the table shows that the islets had
an estimated 1.7 X 104 H. atra and 3.7 X 108
H. leucospilota. In addition, an extensive area
(4 X 104m2) of shallow (4-6 m) lagoon bot-
tom off Island 13 and Island 15 (Fig. 1) was
observed from a skiff to be populated with an
estimated 104 30-cm black (not sand-coated)
H . atra lying on the light sandy portions among
the darker predominating, staghorn corals. The
estimate for the 700 m of reef -flat north of
Aniiru L, as far as a transverse ridge across the
reef, was based on three traverses from 3 to 6
m wide: first, walking north along the middle
of the reef; then back, one-third of the way in
from the seaward edge of the reef; and finally,
pacing west across the reef on a line 200 m
north of Aniiru I., for a distance of 500 m
lagoonward from the seaward reef edge. Extra-
polations of the above observations to the entire

Fig. 2. Inner edge of reef-flat, Rongelap I., section
29, showing Holothuria in pools. Sept. 5, 1959, 1025
hr.

PACIFIC SCIENCE, Vol. XVII, July 1963

atoll suggest the following numbers of H. atra
and H. leucospilota , respectively: for islet shores,
2.5 X 104 and 5.5 X 103; for inter-islet reefs,
4 X 106 and 105; and for shallow lagoon bot-
tom areas, 5 X 105 H. atra, or a total for the
entire atoll of 4.5 X 10° atra and 1.6 X 105
leucospilota.

OTHER SPECIES OF SEA CUCUMBERS

Species observed other than H, atra and H.
leucospilota are mentioned in order of decreas-
ing abundance. Stichopus chloronotus , the green-
ish-black prickly sea cucumber on the inter-islet
reef-flat and on the sandy, shallow lagoon shore
near the north channel of Kabelle I. was most
common. Also on the shallow, sandy bottom,
lagoonward from Kabelle I. we observed in mid-
September 1961, during a period of extreme
tides, a heavy concentration (about 1/m2) of
10 to 20 -cm specimens of a light-colored Ac-
tinopyga sp. with small brown spots. Dead
specimens were common on the lagoon shore of
Kabelle I. The large brown Actinopyga mauriti-
ana occupied the outer edge of the reef-flat.
Holothuria gyrifer, a small tan-colored sea cu-
cumber with white spots, was common under
rocks on the seaward reef and especially in the
channel north of Kabelle I. Occasional speci-
mens of Ophiodesoma spectahilis were encoun-
tered under rocks.

SAND PASSING THROUGH HOLOTHURIA

The holothurians have been called the earth-
worms of the sea. The amount of sand passing
through the gut of Holothuria is of interest
because of the possibility of profound alteration
of the ecology in those areas where sea cucum-
bers are abundant. Crozier (1918) has estimated
that in 1.7 sq miles of bottom in Harrington
Sound, Bermuda, 500-1000 tons of sand pass
through Stichopus moehii annually (considering
their average length to be 27 cm and the dry
weight of the contained sand to be 46 g). He
concluded that the primary effect of the feeding
habits upon the environment is the moving of
sand from one place to another.

Trefz (1958:14) timed the passage of sand
through captive H. atra at about 12 hr for an
average-sized specimen of 30 cm. Assuming an

Sea Cucumbers at Rongelap Atoll — Bonham and Held

309

Fig. 3. H. leucospilota (black) extending from
underneath H. atra in a pool on the eastern shore of
Rongelap I., Aug. 23, 1958, 1700 hr.

average gut content of 100 g dry weight, this
would be equivalent to 1 gram in 7.2 min.
Larger specimens required more time for pass-
age of sand through the gut than did smaller
specimens, according to the following relation-
ship derived from her data for specimens of
25-38 cm (omitting two outlying observations):

t=5.3 + 0.233 L

where t represents time in hours for passage
through the gut, and L is length of sea cucumber
in cm.

Yamanouti (1939:614) weighed the gut con-
tents of 65 small (5-25 cm) H. atra and re-
corded an average of 17.1 g dry weight (range,
0.4-40.5 g). Time required for passage through
the gut of captive specimens averaged 4.8 hr.
Thus, 1 gram would be ingested every 16.8 min
on the average.

Direct observation during low, daytime tides
of undisturbed, feeding individuals of H. atra

30 cm long in their natural habitat at Rongelap
Atoll showed egestion of sand at the rate of 1
g (dry basis) in from 5 to 10 min, agreeing
with deductions from the data of Trefz on cap-
tive specimens. Assuming continuous feeding, at
Rongelap Atoll the estimated 5 X 10r> H. atra
and H. leucospilota probably ingest and egest
about 2.4 X 108 kg of sand yearly.

It has been pointed out by Crozier (1918)
for Stichopus moebii, by Yamanouti (1939) for
H. atra and other species, and by Trefz ( 1958)
for H. atra, that there is virtually no grinding
action or reduction in size of sand particles upon
passing through the gut. Trefz has demonstrated
that even such minute and delicate calcareous
structures as the integumental "anchors and
plates” of the sea cucumber, Ophiodesoma spec-
tabilis, do not undergo perceptible dissolution
in the gut of H. atra, but that they appear the
same after egestion as before ingestion, thereby
negating the idea of chemical dissolution in the
gut.

In some of the scoured depressions on the
seaward reef where very little sand is present,
H. atra serves an additional function by retain-
ing within and around itself much of the sand
required in its feeding that might otherwise be
washed away.

SIZE OF SPECIMENS

The length of undisturbed H. atra in the open
ranged from about 2 to 60 cm, with estimated
weights of 10 to 2000 g. We have not observed
smaller ones. Frequent estimates of length dur-
ing the censusing led to the opinion that the
geometric mean length of H. atra on the islet
shores is about 17 cm and of the more slender
H. leucospilota about 25 cm. In contrast to the
elongate shape of the larger specimens, the
smaller specimens of H. atra were short and
thick; in fact, the smallest (2 cm) were as thick
as they were long. At the northeastern reef of
Kabelle I. on September 14, 1961, 407 speci-
mens of H. atra in a single pool were individ-
ually measured or estimated for length and
diameter in inches, while still undisturbed,
and were then placed in a container of water
for volume measurement in cm3 by displace-
ment. The temperature of the pool ranged
from about 35 to 39 C during the 5 hr re-

310

PACIFIC SCIENCE, VoL XVII, July 1963

TABLE 2

Length-Frequencies of 407 H. atra Measured
in a Pool on the Northeastern Shore of
Kabelle Island, Sept. 14, 1961

LENGTH

INCHES

NUMBER

LENGTH

INCHES

NUMBER

159

quired for measuring. Both the high tempera-
ture and the slight, unavoidable disturbance of
adjacent individuals when picking up specimens
for volume measurement resulted in a small
amount of contraction. After handling, the speci-
mens were deposited in another part of the pool.
Table 2 gives the length-frequencies. These
measurements show a single major peak at about
5 inches ( 12.7 cm), and fail to affirm the pres-
ence of other expected frequency groups.

Figure 5 is a histogram showing the volume-
frequency. As with length-frequency there is
a single peak ( at about 60 cm3 ) .

From lines fitted by inspection to the scatter
diagram of Figure 6 the relationship of volume
(V) to length (L) in the size range 5-15 cm
was estimated to be:

V= 5.4 L0-94

The rather abrupt increase in slope beyond
15 cm is emphasized by the visually fitted
line:

V = 0.019L30

A few additional measurements of specimens
longer than 25 cm suggest that this slope of
3.0 continues throughout the upper size-range.

TEMPERATURE TOLERANCE

Consideration of their habitat, exposed to the
sun in shallow pools at low tide, would suggest
a high temperature tolerance for H. atra. On

partly cloudy days after at least 2 hr of con-
tinuous sunshine, the temperatures of 69 pools
containing H. atra ranged from 31.1 to 39.4 C,
with mean and standard deviation of 36.19 and
2.14 C. The temperatures, determined with clin-
ical centigrade thermometers, of eight black sea
cucumbers were consistently higher than simul-
taneous temperatures in the sand-coated sea
cucumbers alongside them (by an average of
0.25 C, standard deviation 0.16 C). After more
prolonged exposure to sunlight this differential
would be expected to increase. Shaded thermom-
eters on the bottoms of the pools gave readings
lower than within the sand-coated sea cucum-
bers, while thermometers in direct sunlight on
the pool bottoms approximated those within
the black sea cucumbers. Extreme differences
between pool- and specimen-temperatures at any
one place and time barely exceeded 1 C Some
degree of reflective insulation from the radiation
of the sun is imparted to H, atra by the coating
of light-colored sand with which it habitually
covers its body, enabling it to retain a slightly
lower body temperature than it does when not
sand-coated, or than does the naked, black H,
leucospilota when out in the open.

The warmest pool in which a H. leucospilota
was found was 38.4 C, but the animal was not
feeding. The usual habitat of H. leucospilota is
cooler than that of H. atra because of its habit
of seeking the shade, and protection of rocks.
Water coming in over the reef at high tide has
a temperature of about 29 C. When this cool
water saturates the beach it remains cool if
overlain by strata of slab rock, but is warmed
by the sun if the sand is exposed. H. atra oc-
curred in pools of higher temperature, 39.4 C.
The upper limit was not determined. Crozier
(1915: 281) gives the maximum temperature
of the habitat of H. surinamensis as 31.8 C and
states that muscle coagulates at 42 C. This is
only about 3 C above the highest pool tempera-
tures in which H. atra has been encountered.
H. atra is unusually tolerant of heat, and is al-
most the only macroscopic organism living in
these warm pools at low tide on the warmest
days.

Fig. 4. Photomicrographs showing calcareous deposits in the integument. A, Tables of H. atra. B, Tables
and buttons of H. leucospilota.

Sea Cucumbers at Rongelap Atoll — Bonham and Held

311

312

PACIFIC SCIENCE, Vol. XVII, July 1963

PHOTOTROPISM

H. atra, and to an even greater extent, H.
leucospilota are negatively phototropic since
both protect themselves from strong light, H .
atra by its sand coating, and H. leucospilota
by seeking shelter. They usually react to quick
changes in light intensity, as by withdrawal of
tentacles when covered by a shadow during feed-
ing. H. leucospilota reacts to a shadow by sud-
denly contracting under the protection of the
rock where its posterior end is anchored, thus
frequently requiring a quick grasp and a firm
pull in order to collect this species.

PREDATION

Besides reflective insulation the sand coating
of H. atra affords concealment. This function,
however, would seem to be subordinate to that
of insulation, because of the small amount of
predation to which H. atra is apparently sub-
jected, unless, of course, the converse obtains,
that the small amount of predation may be due
to the concealing effect of the sand coating. A
damaged sea cucumber has been observed only
once. This specimen, presumably pecked by a
bird, occurred in an area where curlews had
been active. In no case has an animal been ob-
served actually preying upon H. atra. Grazier
(1915:246) observed lightly colored parts of

Fig. 5. Histogram showing volume-frequency of
407 H. atra from a pool at Kabelle I., Sept. 14, 1961.

Fig. 6. Relationship of volume to length of 407
H. atra at Kabelle I., Sept. 14, 1961. Lines drawn by
inspection.

H. surinamensis which he suspected of being
regenerated following either autotomy or injury
by enemies such as fishes or crabs, but states
that there is no evidence that Holothuria has
enemies of this kind. Frizzell and Exline ( 1955:
29) ascribe predation on intertidal sea cucum-
bers to sea gulls, and note the general concur-
rence among authors that predation is slight, but
propose that the apparent protective mecha-
nisms of holothurians suggest predation to be
more common than is supposed.

TRANSVERSE FISSION

Fission has been reported in holothurians by
Crazier (1915:291), who found regeneration
of either end of the body in about 10% of the
H. surinamensis encountered. Frizzell and Exline
(1955:15 and 27) state that a few forms in-
cluding Holothuria parvula (Selenka), Cucum-
aria lactea, and C. planci (Brandt) sometimes

Sea Cucumbers at Rongelap Atoll — Bonham and Held

313

reproduce by transverse fission, and that Holo-
thuria difficilis Semper of the tropical Pacific
commonly does so.

In September 1959 in a warm pool of about
37 C on the southwest shore of Burok I. two
short specimens of H. atra were encountered
lying end to end. The integument of the two
adjacent terminal end portions was devoid of
sand because of the absence of tube feet. The
specimens were saved, and later examination
revealed a calcareous oral ring in only one, in-
dicating that the two individuals resulted from
transverse fission. Another specimen in an ad-
jacent pool was photographed in an advanced
stage of constriction; but without disturbance
by even so slight a stimulus as a shadow it vol-
untarily relaxed the constriction and resumed
normal shape within a period of about 5 min.

During the September 1961 census, enough
fissioning ( Fig. 7 ) and fissioned specimens were

Fig. 7. H. atra, about 23 cm long, undergoing
transverse fission in a pool 10-cm deep. Mellu I., sea-
ward side near west end, Sept. 18, 1961, 1520 hr.

encountered to verify that fission, or asexual re-
production, does commonly occur in Holothuria

atra .

The importance of fissioning as a means of
reproduction is emphasized in Figure 6 by the
relative thickness of the small specimens and the
consequent reduced slope (0.9) of their logarith-
mic relationship of volume to length as com-
pared to the usual cubic relationship (slope of
3 ) for the larger specimens.

If it may be assumed that sexual reproduction
would be seasonal in nature, while fissioning
would occur at all seasons, then the lack of ap-
parent age classes or frequency modes in the
volume-frequency histogram of Figure 5 is also
consistent with the concept of reproduction by
fissioning.

SUMMARY

1. The preferred habitat for H. atra and H.
leucospilota on the seaward reef-flat is described
and is linked with the presence of slabs or
shelves of rock at the junction of the sandy
beach with the reef-flat. On hot days runoff
water from such areas is of relatively low tem-
perature (because of the insulating effect of the
slabs of rock).

2. Estimates of numbers of these two sea
cucumbers are based upon notes and photo-
graphs made while traversing the peripheries of
the larger islets and some reef and lagoon areas,
resulting in an estimated total of about 5 X 10°
H. atra and 2 X 10 5 H. leucospilota.

3. The passage of sand through the gut of
H. atra is at the rate of 1 g (dry weight) in
from 5 to 10 min, resulting in an estimate for
the atoll of about 2 X 108 kg of sand ingested
and egested yearly. The primary ecological role
of the sea cucumbers in this feeding action is
to move the sand from one place to another.

4. Length of H. atra observed ranged from 2
to 60 cm. Measurements of 407 specimens in
one pool showed that for specimens up to 15
cm in length, volume increased slowly with
length, while for those from 15 to 25 cm, vol-
ume increased almost as the cube of length. This
is construed as evidence of reproduction by
fissioning.

5. H. atra appeared healthy and feeding in
small pools with temperatures up to 38.9 C.

314

PACIFIC SCIENCE, Vol. XVII, July 1963

The internal temperature is slightly lower in
the sand-coated than in the naked black speci-
mens because of the reflection of light and
heat by the coating of sand characteristic of
this species.

6. Evidence of predation upon H. atra or H.
leucospilota is practically lacking.

7. Transverse fission in H. atra has been ob-
served in various stages of progress, and is
considered to be an important means of repro-
duction.

REFERENCES

Crozier, W. J. 1915. The sensory reactions of
Holothuria surinamensis . Zool. Jb. Phys. 35:
231-297.

1918. The amount of bottom material

ingested by holothurians. J. Exp. Zool. 26:
379-389.

Cuenot, L. 1948. Anatomie, Ethologie et Sys-
tematique des Echinodermes. Classe de Holo-
thuries. In: P. P. Grasse, Traite de Zoologie,
Anatomie, Systematique Biologic. Tome 11,
Echinodermes, Stomatocordes, Procordes,
Paris. Pp. 82-120.

Frizzell, D. L., and Harriet Exline. 1955.
Monograph of fossil holothurian sclerites.
Bull. Univ. Missouri School of Mines & Met-
allurgy, Tech. Ser. 89: 1-204.

Trefz, Shirley M. 1958. The physiology of
digestion of Holothuria atra Jager with spe-
cial reference to its role in the ecology of coral
reefs. Doctoral thesis, University of Hawaii.
Pp. 1-149.

Yamanouti, T. 1939. Ecological and physio-
logical studies on the holothurians in the
coral reef of Palao Islands. Palao Trop. Biol.
Sta. Stud. 4:603-635.

Field Identification of Five Species of Californian
Beach Hoppers (Crustacea: Amphipoda)

Darl E. Bowers1

While studying the correlations between the
distribution of five species of beach hoppers of
the genus Orchestoidea and the physical and
biotic factors of their sandy beach habitat, it
became necessary to identify with certainty the
animals collected from the many beaches sam-
pled up and down the Californian coast. Since
it was desirable tentatively to name the hoppers
captured in the field, I made an effort to find
characters which were easily observed on the
beach and would allow such identification. As
a result of my examination of many thousands
of animals from over a hundred collecting locali-
ties, I evolved a recognition of the five species
mainly on the basis of pigmentation patterns,
elements of which are relatively consistent in
spite of the many variations to be found in the
total pigmentation.

The species under consideration are the two
large hoppers, Orchestoidea calif orniana
(Brandt) 1851, and O. corniculata Stout 1913;
the two somewhat smaller species 0. columbiana
Bousfield 1958, and O. pugettensis (Dana)
1853; and the small species, O. benedicti Shoe-
maker 1930.

Laboratory identification of these has been
made possible by E. L. Bousfield ( 1957, and
particularly 1959), to whom I am indebted for
his help with some of the key characters. In
these papers Bousfield has described a new spe-
cies from southern California, 0. minor, similar
to O. columbiana, but with a few subtle anatom-
ical differences. These will require statistical
analysis in order to show more clearly the affini-
ties of the two species. This newly described
form will not be considered here since I have
been mainly concerned with the beaches of
central and northern California.

The pigmentation patterns may be relied on
for identification of both sexes for almost all

1 Mills College, Oakland, California. Manuscript re-
ceived January 30, 1962.

sizes of animals, with the following cautions:
( 1 ) The largest individuals of calif orniana al-
most invariably have lost the dorsal pigmenta-
tion, but color and form of the second antennae,
as described below, will usually make this no
problem; (2) the largest individuals of cornicu-
lata may have lost the dorsal pigmentation, but
again, color and form of the antennae will
usually suffice; (3) the smallest individuals of
all species (under 4 mm in body length) may
be so lightly pigmented as to make identification
difficult or to require magnification and lighting
available only in the laboratory; (4) animals
living on white or very light-colored sands
usually are only faintly marked, in which case
combinations of other characters may have to
be used.

Some general beach characteristics and hop-
per habits may also be of use in identification
but are not dependable alone. O. calif orniana
is typically found on long exposed beaches
composed of fine sand, with few if any rocks
present, beaches that are wide from the fore-
shore to back shore with dunes into which the
hoppers may retreat during high spring tides
and for breeding, beaches which usually have a
quite flat foreshore, which is correlated both
with exposure to wave action and with fine
sand particles. O. corniculata, on the other hand,
is typically found on shorter, more protected
beaches composed of coarser sand, possibly with
rocks or boulders present, beaches which may
be narrow from foreshore to back shore with
cliffs or piles of boulders preventing retreat dur-
ing high spring tides (breeding animals are
commonly found mixed with the rest of the
population), beaches which may have a fairly
steep foreshore, which is correlated both with
protection from wave action and with coarser
sand particles. These two animals are not often
found together, but occasionally on a calif or-
niana-dommated beach, as at Carmel, California,

315

316

PACIFIC SCIENCE, VoL XVII, July 1963

corniculata hoppers may occur at the ends of the
beach where conditions are not optimal for
calif ornicma. I have never found calif orniana on
corniculata-&omimx£<\. beaches.

For columbiana and pugettensis I have not
gathered enough data to make very meaningful
statements here. They often occur together on
coarse sand beaches, some of which are quite
long, onto which relatively little seaweed is cast
as food for the hoppers. 0. benedicti is com-
monly found with calif orniana, and otherwise
lives on beaches of the finest, silty sand.

It is not always easy to find mature individuals
in the daylight on a beach and much sample
digging is often required to locate a ''bed” of
hoppers. Small individuals are readily found but
are not so readily captured. The distribution of
hoppers on a beach is not of a random nature,
but is dictated by the location of seaweed, the
main food of these omnivores, on the shore, and
by the tidal cycles, which they tend to follow up
and down the beach. Since these animals feed
mostly at night on the freshest drift present and
many remain under this drift at dawn, the most
productive searching is usually done under the
seaweeds brought ashore by a high tide of the

previous night. Large individuals, however, may
move farther up the beach; and in calif orniana,
at least, the mature males and some females in
the breeding season (probably February to
October) with eggs or young in their brood
pouches will be found scattered in fairly high
and dry zones that may not have been reached
by the surf for some days.

The location of the burrows of these large
animals requires careful observation in the early
daylight hours. In digging the burrow, or clean-
ing out an old one, large calif orniana kick sand
out onto the surface in two opposite elongated
rays. If this sand is of a different color because
of a higher moisture content or because it was
obtained from different-colored material below,
it may be quite noticeable. However, during
the day, drying action of the wind and sun may
reduce the color difference between the burrow
sand and the surface sand, and the winds level
the elongated heaps, making them increasingly
difficult to see. The burrows may be open shafts
up to 12 inches in depth, with a plug of sand
in the mouth of the burrow.

O. corniculata make burrow mounds much
more like those of a pocket gopher. The sand is

Fig. 1. Sketch of O. calif orniana to show features mentioned in the text.

Californian Beach Hoppers — Bowers

317

pushed up less vigorously than by calif orniana
and thus simply falls all around the burrow
mouth in a rounded heap. Large individuals are
usually found with smaller animals in mid- to
high-tide zones rather than higher up on drier
parts of the beach, as in calif orniana. The bur-
rows are commonly open just around the hopper
and are usually less than 6 inches deep, but I
have found these hoppers more than 2 ft down.

The three smaller hopper species dig less
than the large hoppers and thus do not make
very obvious surface markings. They are more
often found associated with washed-up debris.

I have found no sexual dimorphism with re-
spect to pigmentation pattern, but the sex of
these animals is easily distinguished by the
structure of the 2nd gnathopods (the 2nd pair
of large ventral appendages) except as noted
below (see Fig. 1). In females the sixth seg-
ment is a fleshy paddle-shaped structure. In
young genetic males the same form is present,
but it becomes modelled through a series of
molts into a larger subchelate "hand” with a
curved dactyl closing against an emminence of
the "palm.” As far as I know, males can be dis-
tinguished from females only when the change
in the gnathopods begins to be apparent. In
calif orniana and corniculata this occurs when the
animals reach about 9.0 and 13-0 mm in body
length, respectively (measured from the anterior
of the head through the chord of the straight-
ened but still somewhat curved body to the
back edge of the 3rd abdominal segment ) . Sex-
ually active females possess oostegites under the
thoracic region. These are thin plates with a
fringe of hairs, which overlap to form a mar-
supium in which eggs and young are carried.
However, some large females lack these hairs,
their presence apparently being dependent on
the breeding cycle rather than on size. The 2nd
antennae of females of all species are shorter
and less robust than those of males, and are less
colorful.

In midday, as the waves of a rising tide
moisten a beach, it is common to see small ani-
mals moving around in the just-wet area, feeding
on fresh bits of seaweed, washed up sand crabs,
or seemingly just wandering. Probably all species
do this, but in my experience this movement
has occurred mostly with columhiana, less fre-

Fig. 2. Left 2nd antenna of O. corniculata male.
Compare this with appendage in Figure 1.

quently with calif orniana. Under such condi-
tions, an observer lying face down on the sand
can use pigmentation patterns to distinguish the
animals without even touching them.

The most obvious character that distinguishes
large male calif orniana from male corniculata is
the form and color of the second antennae. In
calif orniana the flagellum of the antenna is rela-
tively thin, longer than the combined length of
the segments of the peduncle, and gradually
tapers to the tip, which reaches back beyond the
middle of the body when folded from the pe-
duncle base. The color of the peduncle is usually
rosy-red. The peduncle segments are longer but
not as massive as those of corniculata. In cornic-
ulata the flagellum of the antenna is thicker than
in calif orniana, is shorter than the combined
length of the peduncle segments, and tapers
rapidly to the tip, which does not reach the
middle of the body when folded ( see Fig. 2 ) .
The color is usually salmon-pink, but sometimes,
on a darker substrate, may be tinged with blue
or brown. Newly molted individuals of both
species may be white.

The antennae of columhiana are calif orniana-

like, but in large animals are bluish-white rather
than red. The antennae of pugettensis are cor-

318

PACIFIC SCIENCE, Vol. XVII, July 1963

Fig. 3. Dorsal views of thoracic segments 6, 7, and
8, and abdominal segments 1, 2, and 3, showing two
variants of each pigmentation pattern: calif orniana-
like group— O calif orniana; b, O. Columbiana; c,
O. benedicti — corniculata-\ike group: d, O. cornicu-
lata; e, O. pugettensis.

niculata-Yike but are more commonly bluish or
brownish rather than salmon-pink. The anten-
nae of benedicti are proportionally the shortest
of the genus but are shaped like those of cornic-
ulata.

The longest males I have measured (see above
for the reference points) are: O. calif orniana,
26.0 mm; O. corniculata, 25.0 mm; O. Colum-
biana, 19.0 mm; O. pugettensis, 15.5 mm; and
O. benedicti, 10.0 mm. The females are a few
millimeters shorter.

Because of exceptions to the above general
statements about antennae, one finds that with
subadult individuals, females, and indeed even
mature males from some beaches, other obvious

Californian Beach Hoppers — Bowers

319

traits are needed for field identification. Gen-
erally, the overall pigmentation tone of these
hoppers tends to match the color of the sandy
substrate; but even on light sands where pig-
ment is reduced, there are a few key spots that
usually show up. I will be concerned mainly
with the dorsum of the first three abdominal
segments, which appear in a top view as the
last three body segments ahead of the urosome,
and the 7th and 8th thoracic segments.

A study of the sketches in Figure 3 will show
that there are two main groups of pigmentation
patterns. The broad "butterfly” spots of the 1st
and 2nd abdominal segments of the "calif or-
niana- like” group are found with modifications
in O. calif orniana, O. columbiana, and O. bene -
dicti. The T-shaped spots of the 1st, 2nd, and
3rd abdominal segments of the " corniculata-
like” group are seen in O. corniculata and 0.
pugettensis.

The calif orniana-/fi£e group. In calif orniana
and benedicti there is a sagittal line showing in
most variants of this pattern. Note the absence
of pigment from the 3rd abdominal segment on
calif orniana, but the presence of pigment in
benedicti. In columbiana the midline marking
is missing and the "butterfly” design is more
nearly a flattened X, again with no pigment in
abdominal segment 3. The characteristic mark-
ings of abdominal segments 1 and 2 are suf-
ficient to separate calif orniana from columbiana.
Benedicti, besides being small, is more heavily
pigmented with discontinuous spots, which thus
give the impression of a checkerboard design
along the whole body.

The corniculata -like group. The two species
in this grouping, corniculata and pugettensis,
are the most difficult to separate, although they
are easily distinguished from all the others. The
most obvious dorsal pattern differences seem to
be in the last two thoracic segments in which
pugettensis has more delicate markings, but I
have detected no easily recognizable differences
here. A more consistent and reliable set of marks
may be found at the lower margins of the lateral
body wall in the last three thoracic segments of
pugettensis (see Fig. 4). The third of these is
missing in corniculata, which thus shows only
two spots which are usually more diffuse and
less intensely pigmented than in pugettensis.

Fig. 4. Side views to show key characters of thoracic
segments 6, 7, and 8. a, O. corniculata; b, O.
pugettensis.

(These spots do not serve to set off these two
species from the other three, however, for these
others may have such markings as well. ) These
two hoppers are found on the same beach oc-
casionally, and in such cases it may be necessary
to rely on the key characters presented by Bous-
field in the papers cited. However, these pig-
mentation patterns, antennae colors, and other
more subtle clues allow one acquainted with the
animals to make fairly reliable preliminary de-
terminations.

Whatever "hedging” the reader may detect in
this presentation is due to the wide range of
variability to be found in these animals and to
the fact that exceptions keep cropping up as

320

PACIFIC SCIENCE, Vol. XVII, July 1963

study continues. However, it is hoped that this
discussion will make it easier to identify these
animals in the field. I am grateful for financial
support for this study which came in part from

the American Academy of Arts and Sciences,
and in part from the Mills College Faculty Re-
search Fund. My thanks are here expressed to
Dr. Joel Hedgpeth for helpful suggestions.

FIELD KEY TO species OF Orchestoidea

1. Mature animals 2

1. Immature animals and others not distinguished by first part of key 5

2. Second antennae when folded reaching back to or beyond middle of body; flagellum longer

than peduncle 3

2. Second antennae when folded not reaching back to middle of body; flagellum shorter than

peduncle 4

3. Color of 2nd antennae rosy-red calif orniana

3. Color of 2nd antennae bluish-white columbiana

4. Color of 2nd antennae usually salmon-pink corniculata

4. Color of 2nd antennae not so 5

5. Dorsal pigmentation pattern containing "butterfly” designs 6

5. Dorsal pigmentation pattern containing T-shaped designs (Lower limb of T may be faint

or missing) 8

6. Mid-dorsal line absent, "butterfly” spots flattened X’s columbiana

6. Mid-dorsal line present 7

7. No markings on 3rd abdominal segment; sides of body relatively free of pigmentation

marks calif orniana

7. Markings on 3rd abdominal segment; sides of body blotched with checkerboard pattern

...benedicti I

.corniculata
pugettensis Ij

8. Two diffuse spots on sides of body..
8. Three discrete spots on sides of body.

REFERENCES

Bousfield, E. L. 1957. Notes on the Amphipod
genus Orchestoidea on the Pacific Coast of
North America. Bull. So. Calif. Acad. Sci.
56(3): 119-129.

1959. New records of beach hoppers

(Crustacea: Amphipoda) from the Coast of
California. Nat’l Mus. Canada Contr. Zook,
Bull. 172:1-12.

A Simple Device for Making Successive Photomicrographic Records of
Large Groups of Developing Organisms1

Sidney C. Hsiao, Walter K. Fujii,
and Helen H. Fine2

In OUR analysis of the effect of ionizing radia-
tion upon the cleavage of sea urchin zygotes
we found it necessary to take successive photo-
micrographs of a large number of eggs, in
different samples, which had been exposed to
graded doses of ionizing radiation (Hsiao and
Daniel, I960). In order to estimate the rate
of cleavage of the irradiated samples of ferti-
lized eggs it is highly desirable to follow the
cleavage of each egg in every sample and make
photomicrographic records for later analysis.
In other words, we need to take time-lapse
pictures of the developing eggs subjected to
different amounts of radiation so as to calcu-
late the rate of cleavage and correlate it with
dosage. After some preliminary trials we have
put together, using commonly available ma-
terials, a simple device capable of taking pho-
tomicrographs repeatedly from the same field
in a series of samples of irradiated eggs at
specified time intervals. By lining up, accord-
ing to time, prints made from each field, each
egg can be identified and its cleavage followed
from the first to the last frame in the series,
and its rate of cleavage can be calculated. It
occurs to us that investigators who have occa-
sion to record developmental and other recur-
rent phenomena may find this simple device
useful. A brief description of its method of con-
struction and manipulation is reported in this
paper.

PRINCIPLES OF CONSTRUCTION

The simplest way to obtain a sequential rec-
ord of a number of fertilized eggs after their
exposure to irradiation or other treatment would

1 Work supported by U. S. AEC contract AT (04-3)
330. Hawaii Marine Laboratory contribution No. 189.

2 Department of Zoology, University of Hawaii.
Manuscript received February 6, 1962.

be to place the eggs on the stage of the photo-
micrographic instrument and take cinemato-
graphic or time-lapse pictures. This requires
one set of instruments for each sample exposed
to one specific dose. The cost of equipment and
the limitation of laboratory space would rule
out all sample series of reasonable size. How-
ever, if the samples are immobilized and the
photomicrographic camera is brought over an
exactly predetermined point in each sample to
take time-lapse photomicrographs during the
course of cleavage, it would be possible to make
photographic records of a number of samples
with one instrument. This can be done by: ( 1 )
Immobilizing samples of eggs contained in
uniform-sized culture vessels, such as a culture
cell on a 1- X 3 -inch micro slide, in a straight
line on a stage supported by a steady stand
independent of the photomicrographic camera,
so that when the microscope moves no motion
is transferred to the samples. ( 2 ) Using a pho-
tomicrographic camera carriage made to move
back and forth along a straight track placed
parallel to the line of the immobilized sample
cultures and adjusted to bring the microscope
objective across the middle of the cultures. (3)
Employing a series of uniform spacers (such
as 1-inch rectangular bars for use with the
1- X 3-inch micro slide containing samples of
egg cultures), whose width equals the distance
between the centers of the culture vessels of
two neighboring samples, to stop the photo-
micrographic camera carriage on the track each
time the objective of the microscope reaches the
center of a sample on the immovable stage. In
this way a single photomicrographic camera
can be used to photograph the eggs in the exact
center of each sample culture vessel, and a
series of the time-lapse photomicrographs is
obtained for later analysis.

321

322

MATERIAL USED AND METHOD
OF CONSTRUCTION

PACIFIC SCIENCE, Vol. XVII, July 1963

The device built on the above principles con-
sists of two independent parts: (A) a steady
stand for supporting the cultures to be photo-
graphed, and ( B ) a movable carriage on which
a microscope and its camera and substage lamp
are mounted so that the microscope and its
accessories can travel from culture to culture
without disturbing the stand or the organisms
on it. The materials used for the construction of
this device are limited to the few simple items
listed below:

7-ft length of 2 X 2-inch wood
piece of 28 X 8 X 1-inch plywood
12 -ft length of Vi -inch wide V^-inch thick
angle aluminum

20 X 8 X Vi-inch sheet of aluminum
1 dozen No. 8, 1-inch wood screws
8 self-tapping sheet metal screws, size
3 /32-i/4-inch

3 aluminum sheaves, 5i-inch in diameter
11 machine screws with nuts, size 3/32-
V4-inch

6-ft length of 3-sided 1 -inch-square ex-
truded aluminum channel
piece of 5 Vi X 28 X Vi -inch plate glass

All these are easily obtainable from local hard-
ware stores.

A. The stand. This independent part of the
instrument is made of a rectangular wooden
base, two aluminum trapezoidal end supports,
and a rectangular glass toppiece or stage. The
construction of these three parts is summar-
ized below. ( 1 ) From the 2 X 2 -inch lumber,
two 11- and two 31-inch lengths are cut. These
four pieces are joined at right angles by means
of either end-lap joint or pinned tenon-and-
tusk joint into a rectangular base measuring
31 X 11 inches on the outside. (2) From the
Vs- -inch-thick sheet of aluminum two 8X8-
inch pieces are cut and each is shaped into a
near trapezoid (it will be referred to as trape-
zoid in this paper for simplicity), as shown
in Figure l A, by cutting off a triangular piece
whose two sides adjacent to the right angle are
2 and 6 inches long (area ABC in Fig. 1 A).
After rounding the corners, two 5/32-inch holes

Fig. 1. Parts of the stand. A, Aluminum end sup-
port used as left end of stand. 1, 2, 5/32-inch holes;

1 being 1 inch from vertical edge and 2 5 inches
from 1; 3, 4, 3, are three beveled holes 3 inches apart,

3 being 1 inch from vertical edge, and all Yl inch
from the bottom. B, Diagram of angle aluminum end
frame for plate-glass toppiece or stage. Two upper
holes 5/32 inches, two lower ones Ys inch. C, End
portion of plate-glass toppiece or stage showing plate
glass l , mounted between two 28Yl -inch strips of
angle aluminum, 2, which are attached on end piece,

3. The two VXinch holes, H, are for mounting the
toppiece or stage on the trapezoid end support. D,
Sectional view of right end of stand, showing method
of mounting trapezoidal end supports. 2, on wooden ,
base, 3; and toppiece or stage, 1, on the trapezoid, ;
2. S, No. 8 wood screw; H. sheet metal self-tapping i

screw.

Photomicrographic Records — Hsiao, FUJII, and Fine

323

5 inches apart are drilled Vi inch from the top
and Vi inch from the vertical edge of one of the
trapezoids. Three 3/ 16-inch holes spaced 3
inches apart are next drilled Vz inch from the
bottom and 1 inch from the vertical edge. The
holes on the second trapezoidal end support are
similarly drilled but are done from the oppo-
site surface so that this piece of aluminum
forms a mirror image of the first one. ( 3 ) The
rectangular glass toppiece or stage is made
from a 5^2 X 28-inch strip of V4-inch plate
glass and a rectangular frame made of VXinch
wide angle aluminum. Two 554-inch sections
are cut from the aluminum stock and four holes
drilled in each piece. Along one side of the
aluminum angle two 5/32-inch holes are drilled
each Vs inch from the end, along the other side,
two Vs-inch holes, each Vz inch from the end,
are also drilled, as shown in Figure 16. Two
28]/2-inch lengths of aluminum are cut from the
same stock and two Vs-inch holes are drilled on
each piece, each hole being Ys inch from the end.
The 28-inch long plate glass is placed lengthwise
between the inner .surfaces of the long alumi-
num angle strips whose ends are then placed
across the two short (554 inches ) end pieces
in such a way that the outer surfaces of the
four angle strips are in contact and the pre-
drilled pg-inch holes on the long strips match
the 5/32-inch holes on the short ones, as shown
in Figure 1C, leaving the Vs-inch holes of the
short strips exposed at the ends of the frame.
Four 3/32-1/4-inch self-tapping sheet metal
screws are driven through the 5/32-inch holes
in the short end aluminum section tapping into
the Vs-inch holes in the long sections to hold
the aluminum frame and glass together. It will
be noticed that all the 5/3 2 -inch holes are for
the passage of, and the Vs-inch holes for the
tapping by, the self-tapping sheet metal screws.

To assemble the stand the two trapezoidal
end supports are used to join the glass toppiece
or stage to the wooden rectangular base. After
placing the wooden base on a flat bench or
table top, with its long sides running from left
to right in front of the operator, the right
trapezoidal end support is clamped with a C-
clamp on the inside of the 11 -inch-long end
piece of the wooden base so that the vertical
edge of the trapezoid faces the front of the
stand, farthest from the operator and Va inch

from the right distal corner of the rectangular
base, the slanting edge nearest the operator, the
bottom of the trapezoid flush with the bench
top and the smaller openings of the three bev-
eled holes against the inside surface of the
wooden end piece. After drilling three holes
into the wood, using the holes in the alumi-
num trapezoid as guides, three No. 8, 1-inch,
wood screws are driven into the wood until
their heads are flush with the aluminum (see
Fig. ID at S ).

The C-clamp is released. Similarly the left
trapezoidal end support is screwed on the in-
side of the left end piece of the wooden base.
The aluminum framed glass toppiece or stage
is fastened to the trapezoidal end supports by
apposing the two Vs-inch holes on each end
angle aluminum piece (6 in Fig. 1C) to the
two 5/32-inch holes near the top of each trape-
zoid and a round-head self-tapping sheet metal
screw driven from the outside through the
trapezoidal end support, tapping into the angle
aluminum as shown at H in Figure ID. When
properly assembled this stand is very steady
and can support a large number of culture
slides or vessels on the plate glass.

B. The carriage. This part of the instrument
consists of a platform with track, a cart for
carrying the microscope and its accessories,
and a number of spacers of uniform size. ( 1 )
The platform is made from a piece of 1 X 28
X 8-inch plywood and two strips of angle
aluminum. Each piece of aluminum is cut 28
inches long and two 5 /32-inch holes drilled
through the apex of the angle, Vz inch from
each end, and beveled. One piece of the drilled
aluminum is placed along the edge of the ply-
wood board with the apex pointing upward
and a No. 8, 1-inch, wood screw driven through
each hole into the wood. The second piece of
angle aluminum is placed parallel to the first,
but with its apex 5 Vs inches from that of the
first, and fastened on the wood in a similar
way. These two pieces of angle aluminum form
the track on which the microscope-carrying
cart moves. (2 ) The microscope-carrying cart
is constructed from a piece of the V^-inch sheet
aluminum, 5 X 6Va inches in size, with three
pieces of angle aluminum and three Vs -inch
aluminum sheaves attached to its under surface.
Six 5/32-inch holes, as indicated by No. 1-6 in

324

PACIFIC SCIENCE, Vol. XVII, July 1963

Figure 2 A, are drilled through the sheet alumi-
num which forms the floor of the cart. A 44 X
44-inch slit is cut at A and a round 44-inch
hole at B, as shown in the same figure. Slit A
is used to adjust and fix the substage illumi-
nator in place, and B is for the microscope set
screw, the same screw which came with the
microscope for fastening it to the bottom of
the microscope case during shipment. Two 5-
inch sections of angle aluminum are cut, and
two 5/32-inch holes drilled on each limb of
the right angle. Each hole is 44 inch from the
end of the aluminum section. One of these two
sections is mounted on the front undersurface
of the floor of the cart with two 3/32 machine
screws and nuts, and the other section similarly
mounted on the right undersurface at right
angles to the first (Fig. 2 B at 4). A third
piece of angle aluminum, 144-inch long is cut

A. B.

Fig. 2. Diagrams showing microscope carrying cart
floor and top view of stand in use. A, Top surface of
cart floor, showing openings 1 to 6 for attaching angle
aluminum on undersurface; A, slit for adjusting and
fixing illuminator; B, opening for microscope base set
screw. Broken line indicates outline of the base of the
microscope. B, Undersurface of cart floor. 1—3, sheaves;
4, 5 -inch angle aluminum for buffering against spacer.
C, Right portion of plate-glass toppiece of stand show-
ing culture slides, 1-6 in place. Stippled areas repre-
sent masking tape. Other slides and glass (or metal)
are positioning guides.

and a 5/32-inch hole drilled at a distance of
44 inch from each end. It is mounted along
the back border on the undersurface of the cart
floor with metal screws and nuts through open-
ings 3 and 4 (see Figs. 2 A, 2 B). Three alumi-
num sheaves (1, 2, 3 in Fig. 2 B) are mounted
with machine screws and nuts on the vertical
limb of the angle aluminum pieces on the
undersurface of the cart floor so that sheaves
No. 1 and 2 are 444 inches apart and each 44
inch from the end of the 5 -inch section of angle
aluminum. Sheave No. 3 is mounted on the
center of the vertical limbs of the 144 -inch
section of angle aluminum and at a distance
544 inches from a line joining sheaves No. 1
and 2. When the cart is placed on the platform
the sheaves should ride smoothly but snugly
on the track. (3) The spacers are six 5 3/ 16-
inch sections cut from the 1-inch extruded rec-
tangular aluminum channel. One spacer is
mounted on the right-hand end of the plat-
form with No. 8 wood screws, after the proper
holes are drilled, to serve as the starting or
ending point in the side to side movement of
the microscope-carrying cart.

OPERATION OF INSTRUMENT

To employ this instrument for taking photo-
micrographs a microscope and a substage il-
luminator are mounted on the microscope-carry-
ing cart. In our laboratory an old E. Leitz
Wetzler microscope with its mirror removed
is fastened on the car r floor by its set screw
driven through opening B into the horse-shoe
shaped base, and a Kohler illuminator through
slit A. The light is centered and adjusted by
standard procedures (Gray, 1958). The stand
is placed on the laboratory table with its length
parallel with the edge of the table and the
slanting edges of the aluminum end supports
near the operator. The platform is placed inside
the stand so that a 44 -inch clearing is produced
on all four sides between the platform and the
base of the stand. The microscope with its cart
and illuminator is carefully put in place by
sliding the microscope stage immediately under
the plate-glass toppiece of the stand from back
forward across the stand until the sheaves are
over the track. The cart is lowered onto the
track and pushed toward the right until it is

Photomicrographic Records — Hsiao, FUJII, and Fine

325

stopped by the fixed spacer at the end. A cul-
ture slide containing the sample is placed on
the plate glass with its length perpendicular to
the long axis of the glass toppiece and directly
under the low power objective of the micro-
scope. In our experiments a glass or aluminum
ring ^4-inch I. D. and 5/32-inch high is
mounted in the center of a 1- X 3 -inch micro
slide with epoxy cement to serve as a culture
cell. When the culture slide has been centered,
a piece of metal or glass is placed snugly against
its right side and fixed on the plate glass with
masking tape to serve as a slide end-stop. To
facilitate lining up the other culture slides on
the left side of this extreme right one the
1-inch wide space between the 28-inch-long
aluminum frame on the side nearest the oper-
ator and the culture slide is filled with another
1 X 3 -inch slide at right angles to the end-
stop (see Fig. 2 C). Other culture slides are then
easily placed in a straight line on the left of the
original one as shown in Figure 2C by push-
ing them toward the right lower corner against
the slide end-stop. After all the culture slides
are placed in a straight line from right to left,
the microscope-carrying cart is moved toward
the left by drawing it along the track and the
1-inch-wide aluminum spacers are placed be-
tween the cart and the end piece on the plat-
form, the number of spacers used being equal
to the number of culture slides. As both the
spacers and the culture slides are 1-inch wide
the slide on the extreme left will be centered
directly under the microscope. The culture slides
and spacers are numbered correspondingly, both
starting with No. 1 on the left and ending
with the highest number on the right.

A photograph of the assembled device with
the photomicrographic instrument in place is
shown in Figure 3.

To take photomicrographs with this device,
a camera is mounted on the microscope, and
the specimens in the culture cell brought into
view. Usually, before the first exposure is made,
the specimens are centered and arranged on
each slide, using a glass needle, so that a large
number of individuals can be observed in each
microscope field. Once the specimens are prop-
erly arranged, no other manipulation is needed,
and they will stay in place throughout the
course of the experiment. After the specimens

Fig. 3. Photograph of device, with camera, micro-
scope, and illuminator in place.

are brought into sharp focus, a photograph is
taken with the time of exposure and light in-
tensity adjusted by the usual method for photo-
graphic work (Shillaber, 1944). To take a
photograph of the second culture, the first
spacer on the left is removed, the microscope
pushed 1 inch toward the right tightly against
the remaining spacers, the specimens brought
into sharp focus, and a second frame of the film
exposed. This procedure is repeated until the
microscope-carrying cart is against the stopping
spacer on the extreme right of the platform,
and each culture has been photographed.

To start the second series of photomicro-
graphs, the microscope is moved to the left, all
the aluminum spacers replaced on the plat-
form, maintaining their original order, and the
cart pushed tightly against them. This auto-
matically places the microscope objective over
the center of the first culture. After a predeter-
mined time lapse, the process of photomicrog-
raphy described above is repeated, taking a sec-
ond photograph of each one of the cultures,
covering, in each case, exactly the same field
as in the first run. The exact time of day a
photograph is taken is noted down, and later

326

PACIFIC SCIENCE, Vol. XVII, July 1963

1 H

'•••••••Si

: ' :

• , 5JWs**1 v «»**•.*«

• .*?*•*•* *A!‘*.»* • *««*.*.

I*.* ••«•<*# ,*#♦*•■ *f*V *.♦•**«*;?

«**;<*::* :&5 k *;*%:*:«

*• » • i*i*y*«* ♦•**••, * * *****1 *

& * , *.* • * * « IK M *

n*v’Vt

* **v ? *t* V.+t *.» * 5* #;

****»*>'*•*

■> ; a „

Fig. 4. Examples of time-lapse photomicrographs taken with the device. Each vertical column represents one
sample culture in an X-irradiation experiment. Column 1 is the control, or Or; column 2 was subjected to 5r;
and column 3 to 20r. The top horizontal row was taken at 1:25 P.M. or 85 min post-fertilization. The second
horizontal row was taken at 1:30 P.M.; the third row at 2:15 P.M.; and the fourth row at 3:00 P.M. or 180
min post-fertilization.

Photomicrographic Records-— Hsiao, Fujii, and Fine

327

transcribed onto the print. In one of our ex-
periments, for instance, six cultures of eggs,
each culture having been treated with a differ-
ent dose of X-irradiation, were photographed
successively within 3 min. In this way, a series
of time-lapse photomicrographs, each bearing
the time of day when it was taken, was made
for each sample culture from the zygote to a
desired stage, such as morula, and the resulting
photographic prints analyzed and the rate of
cleavage calculated. In Figure 4 selected exam-
ples of sequential photographs are reproduced
to show the stability of the relative position of
the eggs and their structural change during
cleavage.

To identify each egg and estimate its rate
of cleavage, a tracing of the first photograph
of each sample is made on transparent paper.
The drawing of each egg to be followed is
given a number. By superimposing the tracing
on the second or any subsequent time-lapse
photograph of a particular sample culture, the
eggs can be individually identified. The time
it takes for an egg in a given sample culture
subjected to a specific treatment to attain a
certain developmental stage, (for instance, the
attainment of the four-cell stage by a zygote
after acute exposure to 20 roentgens), is deter-
mined by subtracting the time of fertilization
from the time the picture was first taken which
shows the desired stage of the egg. Thus, in
one experiment the eggs were fertilized at
12:00 noon, the two-cell stage of egg No. 38
first appeared in the photograph taken at 1:30
P.M.; the four-cell stage at 2:15 P.M.; and the
eight-cell stage at 3: 15 P.M. Thus we can deduce
that it took 90 min for 1st cleavage, 135 min
for 2nd cleavage, and 195 min for 3rd cleavage.

DISCUSSION

1. Blum and Price (1950) described an ap-
paratus for following photographically the
cleavages of sea urchin eggs. They pointed out
two advantages of the photographic method:
( 1 ) more accurate timing of the cleavages than
fixing followed by counting; (2) the cleavages
of each individual egg can be followed and
recorded. Blum and Price used two microscopes
in the inverted position, one for U-V-ray irra-

diated eggs, and the other for controls. The
present device requires only one microscope.

2. In our instrument, observation is not lim-
ited to one treatment, i. e., one sample of treated
animals compared with its control. In the pres-
ent device, a large number of samples, each re-
ceiving a special treatment, can be compared
with one or more untreated control or controls,
using only one photomicrographic apparatus. A
stand 28-inch long as used in our apparatus,
can accommodate 20 or more samples at one
time. The only limit to the number of samples is
the time required for making one run of pho-
tography through the whole set of samples,
and that required for making photographic
print-analyses later. With this apparatus (as
with Blum and Price’s) the change in cleavage
rate due to treatment such as X- irradiatic® 'Can
be recorded. For example an increase or de-
crease in the rate of cleavage can be observed;
or initial retardation followed by recovery is
made apparent. Similarly, the course of abnor-
mal development can be followed; some of the
abnormalities may disappear, others may per-
sist. Fixing of eggs and later examination can
not provide a sequential series of recorded
events, especially recoveries. Furthermore, the
eggs from a single female can be used in all
of these samples, thus eliminating individual
variations resulting from the use of batches of
eggs from different females.

3. The instrument described here is one ver-
sion of our device. Other modifications both in
the material used and in details of construction
are possible. For instance, we have also applied
two coiled springs symmetrically to the micro-
scope-carrying cart in order to pull the entire
carriage tightly against the aluminum spacers.
Another modification is the use of a light cool-
ing and filtering device. A shallow transparent
tray containing cold water can be placed on
the plate glass between the culture samples
and the light source.

ACKNOWLEDGMENT

We are indebted to Dr. Walter R. Steiger
and Mr. George Yokotaki of the University of
Hawaii physics department for their advice
and machine-shop help.

328

PACIFIC SCIENCE, Vol. XVII, Inly 1963

SUMMARY

A simple device for taking time-lapse pho-
tomicrographs of a group of cultures of eggs
to show the development of each individual
has been described. This device is made of
readily available material. It consists of two
parts: (1) A piece of plate glass supported
by a frame which is steady and independent
of the motion of the photomicrographic appara-
tus. It serves to replace the stage of the micro-
scope and to carry the eggs in cultures in a
motion-free state. (2) A carriage for the pho-
tomicrographic apparatus which can be made
to travel back and forth in a straight line from
left to right, repeatedly bringing the micro-
scope and its camera to a specific field in each
culture for sequential photomicrography.

The device has the advantage of being sim-
ple, easy to construct and operate, and it can
be used to follow photographically an individ-
ual egg during its early development from zy-
gote to morula as well as to record the appear-

ance, disappearance, or persistence of develop-
mental abnormalities. Other recurring phe-
nomena can also be studied photomicrograph-
ically by this instrument.

REFERENCES

Blum, H. F., and J. P. Price. 1950. Delay of
cleavage of the Arbacia egg by ultraviolet
radiation. J. Gen. Phys. 33:285-304.

Hsiao, S. C., and P. C. Daniel. I960. Studies
on the effect of ionizing radiation upon de-
veloping sea urchin. Anat. Rec. 137: 365.

Gray, P. 1958. Handbook of Basic Microtech-
nique. McGraw-Hill Book Co., New York.
Pp. 24-39.

Shillaber, C. P. 1944. Photomicrograph in
Theory and Practice. John Wiley & Sons,
New York.

Revision of the Genus Pandanus Stickman, Part 15
Malayan Species Described by H. N. Ridley

Harold St. John1

As IN MANY OTHER GENERA Henry N. Ridley,
director of the Botanic Garden, Singapore, made
noteworthy contributions to the knowledge of
the Malayan species of Pandanus. His species
were validly published in local journals or in
his books. He described them briefly, usually
giving some details of the stem, leaves, appear-
ance of the syncarp, length of style, and width
of the exposed tip of the drupes. He cited
specimens but did not adopt the type method.
None of his species were illustrated. The writer
has investigated Ridley’s species, sought the
holotypes or has chosen lectotypes in the Singa-
pore herbarium. These specimens were kindly
made available by the present director, Dr. H.
M. Burkill.

Ridley’s species of Pandanus are nearly all
good, and these are to be maintained. He placed
most of his species in either the section Rykia
or in Aero stigma. As will be noted in the treat-
ment that follows, additional sections are to be
found among his species. The large majority
of Ridley’s species are treated in this article;
and the few remaining ones will be illustrated
and described in subsequent parts when their
type specimens have been located and studied.

SECTION Aero stigma

Pandanus aurantiacus RidL, Roy. Asiat. Soc.,
Straits Branch, Jour. 41: 49 (1904) =
1903; FI. Malay Penin. 5: 81, 1925;
Martelli, Webbia 4(2): t. 30, figs. 10-
13, 1914; (sect. Aero stigma)

Fig. 172

DIAGNOSIS OF LECTOTYPE: "Large branching
shrub with stems 3.6 m. tall, 5-7.5 cm. through”;
leaves "glaucous green”; the lower bracteal leaf
1.26 m long, 4.2 cm wide, coriaceous, 1 -ribbed

1 B. P. Bishop Museum, Honolulu 17, Hawaii,
U. S. A. Manuscript received July 19, 1961.

but 2 -pleated and in section M-shaped, the
blade sword-shaped, from the base gradually
tapering to the approximately 20 cm long subu-
late apex which about 10 cm down is 1.5 mm
wide, at midsection each side with 42 parallel
secondary nerves, the base amplexicaul and un-
armed but beginning 7-8 cm up the margins
with serrations 0.6-0.9 mm long, 1-2.5 mm
apart, pale; the midrib below from 21 cm up
with serrations 0. 3-0.5 mm long, 2-10 mm
apart, subulate-tipped; at midsection the mar-
gins and midrib below with serrae 1-1.5 mm
long, 1-3 mm apart; near the apex the margins
and midrib below with rigid serrulations 0.4-0.6
mm long, 2-4 mm apart; pistillate inflorescence
43 cm long, 12-13 mm in diameter, trigonous,
bracted, spicate, bearing in the terminal 20 cm
5 syncarps which are orange and subequal; syn-
carps 6-7 cm long, 5-5.5 cm in diameter, ovoid
to broadly ellipsoid, bearing about 160 drupes,
these 21-22 mm long, 3.5-5 mm wide, 3-4
mm thick, lower 34 oblanceoloid, upper lA
subulate, 4—6-angled, the body sides smooth,
later with exposed fibers; pileus 10-11 mm
long, of a lower half narrowly conic-pyramidal,
the surface smooth, sharply angled, the angles
continuing along the stout subulate style form-
ing the upper half, arcuate proximally; stigma
3 mm long, linear, in a distal cleft running al-
most to the apex; endocarp centering in lower
lA, and 7-8 mm long, the walls pale, 0.3 mm
thick; apical mesocarp a broad cavern 2 mm
long; basal mesocarp fibrous up the sides, fleshy
within.

EXPANDED DESCRIPTION: Shrub to 5 m tall;
stem with the leaf scars inconspicuous; prop
roots 60 cm long; leaves to 2 m long, 4-5 cm
wide, above clear green, below paler and glau-
cous; foliage leaves little broadened at base,
the margins beginning at 4 cm up with prickles
1.6-2 mm long, 2-5 mm apart, stout subulate,
ascending; the midrib below beginning at 9

329

a cm. o zcvei

FIG. 172. Pandanus aurantiacus Ridl., from lectotype. a, Infructescence, X Vl\ b, drupe, lateral view,

X 1; c, drupe, longitudinal median section, X 1; d, drupe, lateral view, X 4; e , drupe, longitudinal median

section, X 4; /, drupe, style, and stigma, apical view, X 4; g, leaf base, lower side, X 1; b, leaf middle,

lower side, X 1; h leaf apex, lower side, X 1-

Page 274: Revision of Pandanus, 15. Malaya — St. John

331

cm with prickles 3-3.5 mm long, 8-20 mm
apart, very stout subulate, strongly reflexed; pis-
tillate scape 15-40 cm long, bearing 3-9 syn-
carps, these 5-7 cm long; drupes very hard until
ripe when they become bright reddish orange
and soft.

LECTOTYPE: Singapore, Bukit Timah [Bukit
Mandai Road], 1894, H. N. R[idley ], (SING).
Specimen examined and herein designated as
lectotype.

SPECIMENS EXAMINED: Singapore, Pasir Pan-
jang, 17 Oct. 1912, I. H. Bur kill & M. R. Hen-
derson 6,802 (bo, SING). Malaya: Johore, Pulau
Kukub, 1904, H. N. Ridley (paratype), (SING).

Ridley, when publishing this species, cited
three collections and mentioned Sarawak, but
did not designate a holotype. The most com-
plete of his original specimens is here chosen
as the lectotype.

Pandanus collinus Ridl., Mater. FI. Malay Penin.
2: 228, 1907; FI. Malay Penin. 5: 79,
1925; Martelli, Webbia 4(1): 10, 1913;
(sect. Acrostigma )

Fig. 173

DIAGNOSIS OF LECTOTYPE: Shrub 1. 2-2.1 m
tall; stem at summit 6 mm in diameter, brown-
ish, smooth; leaves 25-45 cm long, 8-11 mm
wide, firm chartaceous, apparently pale green
on both sides and flat except near the base
where 2 -pleated, 1 -ribbed, above unarmed, at
the middle with 10-11 secondary parallel veins
in each half, below the tertiary cross veins
barely visible, gradually narrowed in the last
10 cm to a subulate tip 15-20 mm long and
0.6-0.8 mm wide, the base amplexicaul, un-
armed, blackish purple, beginning at 2 cm
from the base the margins with prickles 0.7-1
mm long, 3-7 mm apart, subulate, ascending,
yellowish but red-tipped; the midrib below un-
armed to well beyond the middle; at mid-
section the margins with subulate tipped ser-
rulations 0. 2-0.3 mm long, 1.5-3 mm apart,
appressed ascending, red-tipped; near the apex
the margins with serrulations 0. 1-0.2 mm long,
0.3-0. 5 mm apart; the midrib below with ser-
rulations 0. 2-0.3 mm long, 0.5-1. 5 mm apart;
pistillate inflorescence terminal, erect, bearing
1 head; peduncle 8 cm long, 3 mm in diameter,

bracted; syncarp 3 cm long, 2.5 cm in diameter,
subglobose, bearing about 88 drupes, these
14-15 mm long, 4-5 mm wide, 3-4 mm thick,
the body ellipsoid to obovoid, 5-6-angled, 10
mm long; pileus with the base 3.5-4 mm high,
semiorbicular or slightly narrower, rather
smooth, but with longitudinal ridges and lines;
style 4-5 mm long, abruptly subulate, angled,
sharply proximally curved, bony; stigma 3-4
mm long, elliptic-linear, brown, papillose, ex-
tending to the tip; endocarp centering in lower
Vb but extending to the base, the walls 0.1 mm
thick, cartilaginous, stramineous; seed 7.5-8 mm
long, lance-obovoid; upper mesocarp an oblate
hemisphere, filled with white medullary mem-
branes; basal mesocarp fibrous up the sides,
fleshy within, sparse.

LECTOTYPE: Malaya, Kedah, Kedah Peak,
woods, 1893, H. N. R[idley }, (SING). Specimen
examined and here designated as lectotype!

DISCUSSION: P. collinus belongs in the section
Acrostigma. It was described by Ridley from
three collections from Malaya. The best of these
and the one labeled P. collinus! by Ridley is
here chosen as the lectotype. Contrasting dif-
ferences are given for this in our treatment of
P. alticola.

Pandanus glaucophyllus Ridl., Roy. Asiat. Soc.,
Straits Branch, Jour. 41: 50, (1904) =
1903; ( sect. A cro stigma )

Fig. 174

DIAGNOSIS OF holotype: "Stems 3-4 m. tall,
12-25 mm. in diameter,” the internodes brown
to yellowish, smooth, mostly higher than the
nodes; leaves 52-60 "or more” cm long, near
the base about 4 cm wide, near the apex 5. 7-6.5
cm wide, appearing narrowly oblong, rather
flaccid and when dry chartaceous, glaucous, 1-
ribbed but 2 -pleated and in section low M-
shaped, the secondary parallel veins conspicuous
and 34 of them in each half, the apex abruptly
contracted to a 6-7 cm trigonus subulate caudate
prolongation which near its base is 1 mm wide,
the base of the blade amplexicaul and unarmed,
but beginning at 4 cm up the margins with
prickles 0. 5-0.8 mm long, 1-3 mm apart, slen-
der subulate, pale, ascending or diverging and
some of them doubles or triples; the midrib

5 mm

Fig. 173. Pandanus collinus RidL, from lectotype. a, Habit and syncarp, X 1; drupe, lateral view, X 1;
c, drupe, longitudinal median section, X 1;^ drupe, lateral view, X 4; e, drupe, longitudinal median section,
X 4; f, drupe, style, and stigma, apical view, X 4; g, style and stigma, distal view, X 10; k, leaf base, lower
side, X 1; i, leaf middle, lower side, X 1 \ j, leaf apex, lower side, X i.

acm.

Fig. 174. Pandanus glaucophyllus Ridl., from holotype. a, Habit and syncarp, X Vl\ b, drupe, lateral view,
X 1; c, drupe, longitudinal median section, X 1; d, drupe, lateral view, X 4; e, drupe, longitudinal median
section, X 4; f, drupe, style, and stigma, apical view, X 4; g, leaf base, lower side, X 1; h, midrib, lower
side, near base, X 4; i, leaf middle, lower side, X 1; i> leaf margin, near middle, X 4; k, leaf apex, lower
j side, X 1*

334

PACIFIC SCIENCE, Vol. XVII, July 1963

below beginning at 4-9 cm with prickles 0.8-1
mm long, 0.5-6 mm apart, slender subulate,
brownish, ascending, oblique, closely appressed;
at midsection the margins with prickles 0.5-0.7
mm long, 1-3 mm apart, slender subulate, as-
cending flat appressed and occupying recesses
in the margin; the midrib below glabrous or
with a few prickles like those lower down; at
the bend of the apical contraction the margins
with prickles 0.6-0.8 mm long, 0.6-1. 5 mm
apart, slender subulate, somewhat ascending or
divergent; the midrib with prickles 0.6-1 mm
long, 1-2 mm apart, arcuate subulate, ascend-
ing; on the caudate tip the margins and mid-
rib below with prickles 0. 3-0.7 mm long,
0.3-1. 5 mm apart, stout arcuate subulate, as-
cending; pistillate peduncle 8 cm long, 7 mm
in diameter, erect, bearing a single syncarp 5.7
cm long, 4.5 cm in diameter, broadly ellipsoid,
bearing about 320 drupes, these 12-13 mm
long (or 16-18 mm following the curve of the
stigma), 5-6 mm wide, 3. 5-4.5 mm thick, the
body narrowly oblong ellipsoid, 5-6-angled, the
sides smooth; pileus smooth, the base 3-3.5 mm
long, conical or broadly so, bearing the terminal
style 5-6.5 mm long, stout subulate, strongly
curved proximally, having a distal cleft nearly its
whole length in which is the stigma 3-4 mm
long, linear, brown, papillose, running to the tip;
endocarp pale brownish, centering in the lower
(4, its walls 0.1 mm thick; seeds 7-8 mm long,
oblance-oblong; apical mesocarp a cavern 2-2.5
mm long, invaded by trichomes and hairs; basal
mesocarp with fibers up the sides and the center
fleshy.

HOLOTYPE: Malaya, "Perak, Thaiping Hills,
in dense forest about 2500 feet altitude," above
tea border, Dec. 1898, H. N. Ridley (sing).
Holotype examined!

Pandanus parvus Ridl., Roy. Asiat. Soc., Straits
Branch, Jour. 33: 71, Jan. 1900; FI.
Malay Penin. 5: 78, 1925; P. flagellifer
Warb., in Engler’s Pflanzenreich IV, 9:
80, Dec. 21, 1900; Martelli, Webbia
4(1): 14, 1913; 4(2): t. 33, f- 7, 1914;
(sect. Aero stigma)

Figs. 175, 176

DIAGNOSIS OF HOLOTYPE: "Stem 2.92 cm.
tall, 12 mm. in diameter, slender, branched

above,” rooting throughout; leaves 27-37 cm
long, near the tip 32-36 mm broad, near the
base 16-20 mm broad, appearing ligulate, but
actually tapering downwards, thin subcoriaceous,
dark green above, pale and glaucous beneath,
2 -pleated and in section low AA-shaped but only
1 -ribbed, the midrib below narrow and raised,
the secondary parallel veins prominent through-
out, and at midsection 18-21 on a side, the terti-
ary cross veins faintly visible below near the tip,
making small squarish meshes, near the apex ab-
ruptly rounded contracted to a 3-6 cm caudate
deltoid subulate tip 0.8 mm wide, the base am-
plexicaul, unarmed for 4-5 cm, then the margins
with prickles 0.7-1. 3 mm long, 2-7 mm apart,
slender subulate, straight or slightly curved,
brownish, ascending; the midrib below unarmed
to near the apex; at midsection the margins with
similar prickles 0.5-1 mm long, 4-12 mm apart;
on the rounded apical contraction the margins
with prickles 0.5-0.8 mm long, 0.5-2 mm apart,
stouter subulate, slightly ascending; the midrib
below with prickles 0.8- 1.4 mm long, 1-7 mm
apart, arcuate subulate, slightly ascending; on
the caudate tip the margins and midrib below
with stout prickles 0. 3-0.8 mm long, 0.5-1. 5
mm apart, subulate, arcuate, ascending; in the
series of bracteal leaves the first several 2-4
cm long, lance-navicular, entire; pistillate in-
florescence 1 -headed; peduncle 5 cm long, 4-5
mm in diameter, 3 -angled, few bracted; syncarp
erect, 3.5 cm long, 3.3 cm in diameter, ovoid-
globose, bearing about 150 drupes, these 11-12
mm long or following the curve 14-15 mm
long, 3. 5-5. 5 mm wide, 2. 5-3. 5 mm thick, fusi-
form, the body 7-8 mm long, oblanceoloid,
the surface smooth, later fibrous; pileus smooth,
apparently glaucous, 9-10 mm long, the base
oblong-pyramidal and with the apex oblique
and proximally curved, tapering into a hornlike
style which is 5-6 mm long, all (except the
apical ones) sharply proximally curved, stigma
4-5 mm long, lance-linear, dark brown, papil-
lose, extending to the tip; endocarp in lower
V4, stramineous, bony, the walls 0.3 mm thick,
and the lateral walls extending upwards to
bound the apical cavity; seed 5 mm long, ob-
lanceoloid, truncate at tip; apical mesocarp a
cavern filled with white medullary ascending
hairs; basal mesocarp sparse, fibrous and fleshy.

Fig. 175. Pandanus parvus Ridl., from holotype. a, Habit and syncarp, X 1| b, drupe, lateral view, X 1;
c, drupe, longitudinal median section, X 1 \ d, drupe, lateral view, X 4; e, drupe, longitudinal median sec-
tion, X 4; f, drupe and stigma, apical view, X 4; g, leaf base, lower side, X 1; h, leaf middle, lower side,
X 1; i, leaf apex, lower side, X 1-

336

PACIFIC SCIENCE, Vol. XVII, July 1963

1 1

7 cm

O-J

Fig. 176. Pandanus parvus Ridl., from Singapore, cult., 1895, H. N. Ridley, a , Staminate inflorescence,
X 1; b, stamens, lateral view, X 10; Q floral bract, lateral view, X 1-

expanded DESCRIPTION: Pistillate stems
0.8-2.92 m tall, 6-12 mm in diameter; prop
roots up to 50 cm long, 4-5 mm in diameter,
smooth, green to brown, nearly vertical; leaves
27-50 cm long, 21-48 mm wide, the caudate
apex 3-9 cm long; peduncle 5-12 cm long;
syncarp 2.5-4 cm long, 2.5-33 cm in diameter,
globose or ovoid-globose.

DESCRIPTION OF staminate PLANT: Stami-
nate inflorescence 15-18 cm long, bracts white
and somewhat fleshy; peduncle 3-4 mm in di-
ameter, fleshy, soon withering and caducous;
lowest bracts 8-13 cm long, 26-30 mm wide,
sterile, the body 6-9 cm long, navicular, with
prominent parallel veins, the lower half with
margins unarmed; near the apex the margins

Page 280: Revision of Pandanus, 15. Malaya — St. John

337

with prickles 0.5-1 mm long, 0.3-1 mm apart,
subulate, diverging or subascending; the mid-
rib with stouter prickles 0.7-1 mm long, 0.5-1
mm apart, slightly ascending; on the caudate
tip the margins and midrib below with prickles
0.5-0.9 mm long, 0.5-1 mm apart, arcuate
subulate, ascending; median bracts 3-4.5 cm
long, 1.6-2 A cm wide, lanceolate, the margins
and the midrib below towards apex minutely
prickly; spikes 5-8 and 1.5-2. 5 cm long, 0.8- 1.3
cm in diameter, ellipsoid, dense; stamens very
numerous, attached singly and directly to the
axis; filament 0.2 mm long; anther 4.5-5 mm
long, 0.2 mm wide, bearing an apical subulate
prolongation of the connective 0.4-0.7 mm
long.

HOLOTYPE: Singapore, Kranji, 1892, H. N.
R[idley J, (SING). Holotype examined!

SPECIMENS EXAMINED: Singapore, Hinterin-
dien, Dec. 16, 1905, A. Engler 3,862 (b); Singa-
pore, Bidadari, 1898, H. N. R{idley ] 9, 171
( FI ) ; Singapore, cult., first male flowers seen,
1895, H. N. R[idley ], (fi, sing); without col-
lector, ex Hb. (SING), Forest Iurong, low grow-
ing plant, Jan. 10, 1889, Flora of Singapore 92
( B ) , the holotype of P. flagellifer.

Malaya: Johore, Sungei Pelepah Kiri, 26 June
1928, E. J. H. Corner 33,336 (sing).

Pandanus perakensis Ridl., Mat. FI. Malay Penin.
2: 231, 1907; FI. Malay Penin. 5: 81,
1925; (sect. Aero stigma)

Fig. 177

diagnosis of LECTOTYPE: (Probably acau-
lescent); leaves 1.5-1. 8 m long, 2-3.6 cm wide,
subcoriaceous, light green, linear, rather ab-
ruptly narrowed to the 9 cm subulate, trigonous
apex, this 10 cm down 6 mm wide, in section
the blade AA-shaped, at midsection with 39—41
secondary veins in each half; near the base the
margins with spines 2-3 mm long, 5-13 mm
apart, stout subulate, arcuate, pale, broad based,
ascending; the nearby midrib below with spines
2.5-3 mm long, 6-13 mm apart, similar but
even thicker subulate and reflexed; at mid-
section the margins with subulate-tipped serrae
0.5-1. 5 mm long, 4-8 mm apart, the midrib
below with prickles 0.5-1 mm long, 8-15 mm
apart, subulate; near the apex the margins with

serrulations 0. 3-0.5 mm long, 2-3 mm apart;
the midrib below with similar ones but 2-4
mm apart; peduncle 15 cm long, 8-9 mm in
diameter, straight, bracted; syncarps 3-4, spicate,
crowded, 4. 5-6. 5 cm long, 3. 5-4. 5 cm in di-
ameter, elliptic-cylindric (rarely subglobose),
appearing spiny from the numerous curved as-
cending styles ( but a few at the very base
reflexed), bearing about 430 drupes; drupes
20-22 mm long, 5-6 mm wide, 4-5 mm thick,
oblance-fusiform, compressed, apparently green,
upper l/z free, the body 13-14 mm long, ob-
lanceoloid, the apex truncate, pileus at last
shedding, the base semiorbicular, bearing a style
6-7 mm long, stout subulate, ridge angled,
gently arcuate towards apex of syncarp; stigma
4-5 mm long, linear, distal, running to the very
apex; endocarp 6-7 mm long, centering in
lower V3, ellipsoid, truncate, stramineous, car-
tilaginous, the sides 0.15 mm thick; apical meso-
carp 3-3.5 mm long, a fat discoid cavern with
a few white membranes; basal mesocarp fibrous
along the margins, fleshy within.

EXPANDED DESCRIPTION: Pistillate plants
with peduncle 13-15 cm long; syncarps 2-3
and 4.5-7 cm long, 3. 5-5. 5 cm in diameter.

LECTOTYPE: Malaya, "Perak, Maxwell’s Hill”;
June 1893, H. N. R{idley ], (sing). Lectotype
examined!

SPECIMENS EXAMINED: Malaya, Pahang, Ku-
ala Lipis, 1892, A. S. Machado 11,633 (SING);
Kedah, Baling Hill, 21 Nov. 1941, E. J. H.
Corner & J . C. Nauen (SING).

Pandanus spinulosus (Ridl.) comb. nov. (sect.
Acrostigma) \ P. collinus Ridl. var. spinu-
losus Ridl., FI. Malay Penin. 5: 79, 1925.

Fig. 178

DIAGNOSIS OF HOLOTYPE: Stem slender, just
below the leaves 4 mm in diameter; leaves
67-68 cm long, 16 mm wide, subcoriaceous,
green above, below paler and apparently some-
what glaucous, with a central furrow over the
midrib, 2 -pleated, and towards the tip the 2
pleats spinulose above, at midsection with 16-18
parallel secondary veins in each half, these
prominent throughout, below the tertiary cross
veins visible, remote and forming elongate
meshes, the blade ligulate, in outer (4 tapering

Fig. 177. Pandanus perakensis Ridl., from lectotype. a, Infructescence, X 1; b> c, drupe, lateral view, X 1;
d, drupe, longitudinal median section, X 1; e, drupe, lateral view, X 4; f, drupe, longitudinal median sec-
tion, X 4; g, drupe, style, and stigma, apical view, X 4; h, leaf base, lower side, X h leaf middle, lower
side, XI \ i> apex, lower side, X 1-

Fig. 178. Pandanus spinulosus (Ridl.) St. John, from holotype. a, Infructescence, X 1; ^ drupe, lateral
view, X 1; c, drupe, longitudinal median section, X 1; d, drupe, lateral view, X 4; e, drupe, longitudinal
median section, X 4; f, drupe, style, and stigma, apical view, X 4; g, style and stigma, lateral view, X 10;
h, leaf base, lower side, X 1; i, leaf middle, lower side, X 1; j, leaf apex, upper side, X 1; ^ leaf apex, lower
side, X 1-

o loom.

J i _i _J „ 1 , -i.,.. ... L___ _l

Fig. 179- Pandanus unguiculatus Ridl., from holotype. a. Syncarp, X 1; b, drupe, lateral view, c,

drupe, longitudinal median section, X 1; d, drupe, lateral view, X 4; e, drupe, longitudinal median section,
X 4; f, drupe, style, and stigma, apical view, X 4; g, leaf base, lower side, X 1; h, leaf middle, lower side,
X 1 ; i, leaf apex, lower side, X 1 •

Page 284: Revision of Pandanus, 15. Malaya-— St. John

341

gradually to the subulate, deltoid apex about

3 cm long and 1.5 mm wide, the base amplexi-
caul, unarmed, partly dark purple, beginning at

4 cm up the margins with prickles 0.7- 1.5 mm
long, 3-5 mm apart, subulate, ascending at 45°,
the base thickened, pale; the midrib below un-
armed to beyond the middle; at midsection the
margins with prickles 0.7-0.9 mm long, 2-4
mm apart, subulate, ascending; near the tip the
margins and midrib below with subulate ser-
rations 0.4-0. 6 mm long, 0.7-1 mm apart; pis-
tillate inflorescence erect, bearing a single syn-
carp; peduncle 10 cm long, 4-5 mm in di-
ameter, 3-sided, bracteate; syncarp 4.5 cm long,
3.1 cm in diameter, oblong-ellipsoid, bearing
about 120 drupes, these 15-17 mm long, 3-4.5
mm wide and thick, pale green, fusiform but
the upper end contracted and more slender, 5-6-
angled, the body 11-12 mm long; pileus 8-9
mm long, the base 2.5-3 mm high, narrowly
semiorbicular, with 5-6 sharp angles, these
mostly single; style 5-5-5' mm long, sharply
curved proximally, flattened and ridged below,
pale, bony; stigma 3-4 mm long, distal, linear,
running to the apex, brown, papillose; endocarp
centering in lower but extending to the
base, the walls 0.1 mm thick, cartilaginous, pale;
seed 8-9 mm long, extending to the base; apical
mesocarp spherical, filled with white, medullary
membranes; basal mesocarp sparse, fibrous.

HOLOTYPE: Malaya, Kelantan, along the
ridge of Gunong Sitong, 2,500 ft. alt., 6 March
1924, Mo harried Nur & Foxworthy 12,252
(sing).

DISCUSSION: P. spinulosus is a member of
the section Aero stigma, and in that section is
found its closest relative, P. collinus Ridl., which
species has the leaves 25-45 cm long, 8-11 mm
wide, unarmed above, each half at midsection
with 10-11 secondary parallel veins, at mid-
section the margins with subulate-serrations
0.2-0. 3 mm long; syncarp 3 cm long; drupes
14-15 mm long, the pileus base hemispheric,
abruptly contracted to the subulate style which
is 4-5 mm long. P. spinulosus has the leaves
67-68 cm long, 16 mm wide, above towards the
tip the two pleats spinulose, at midsection with
16-18 secondary parallel veins in each half, at
midsection the margins with subulate prickles
0.7-0.9 mm long; syncarp 4.5 cm long; drupes

15-17 mm long, the pileus base narrowly hemi-
spheric, gradually tapering into the thick subu-
late style which is 5-5.5 mm long.

Pandanus unguiculatus Ridl., Mat. FI. Malay

Penin. 2: 229, 1907; FI. Malay Penin.

5: 78, 1925; (sect. Aero stigma)

Fig. 179

DIAGNOSIS OF HOLOTYPE: "Plant small”;
leaves 26-48.5 cm long, near the base 2-2.3
cm wide, near the middle 2. 4-2. 7 cm wide,
ligulate, but perceptibly tapering both ways
from the middle, dark green above, pale green
below, thick chartaceous, only the midrib thick-
ened, but the blade pleated and M-shaped in
section, the secondary parallel veins conspicu-
ous, at midsection 17-19 in each side, the ter-
tiary cross veins visible in upper third below,
oblique, forming short, rather oblong meshes;
near the apex abruptly contracted to a subulate
tip more than 20-25 mm long, 0.9 mm wide,
the base amplexicaul and entire but beginning
at 4-5 cm the margins with prickles 0.7-1 mm
long, 2-6 mm apart, subulate, ascending, pale;
the midrib unarmed for lower 34 s; at the mid-
section the margins with prickles 0.5-0.7 mm
long, 2-5 mm apart, subulate, appressed as-
cending, "red-tipped” but when dried brownish;
near the apex the margins and midrib below
with subulate-tipped serrulations 0.2-0. 3 mm
long, 0.5-2 mm apart; peduncle 5 cm or more
long, 4 mm in diameter, bracted; syncarp sin-
gle, 5.5 cm long, 4 cm in diameter, broad ellips-
oid, bearing about 220 drupes, these 16-18 mm
long (or 18-20 mm, following the curve), 4-5
mm wide, 3.5-4 mm thick, oblance-fusiform,
the body 14-15 mm long, narrowly ellipsoid;
the pileus smooth, pyramidal-ovoid, 5-7 mm
high, tapering into the hornlike style which is
2.5-3 mm long, mostly upwards curved to about
right angles; stigma 2. 5-3. 5 mm long, distal,
linear, papillose, brown; endocarp slightly sub-
median, 9—1 1 mm long, oblanceoloid, the apex
truncate, but with ascending shoulders partly
surrounding the apical cavity, stramineous, carti-
laginous, the lateral walls 0.15 mm thick; seed
7-8 mm long; apical mesocarp forming a
broadly ovoid cavern with a few white medul-
lary membranes; basal mesocarp scant, fibrous
and fleshy.

Fig. 180. Pandanus stelliger Ridl., from lectotype. ar-d, Drupe, lateral view, X 1; e> drupe, longitudi-
nal median section, X 1; f, g, drupe and stigma, apical view, XI \ h, drupe, lateral view, X 4; i, drupe,
longitudinal median section, X 4; j, drupe and stigma, apical view, X 4; /, leaf base, lower side, XI \ m,
leaf middle, lower side, X 1; n, leaf apex, lower side, X 1-

Page 286: Revision of Pandanus , 15. Malaya- — St. John

343

EXPANDED DESCRIPTION FROM ALL SPECI-
MENS EXAMINED: Stems up to 1.8 m tall, 9-13
mm in diameter, smooth, yellowish, erect or
ascending, with aerial roots; leaves arching out-
ward and the tips pendent, glaucous below, the
subulate tip 20-46 mm long; inflorescence erect;
peduncle 5-6 cm long.

HOLOTYPE: Malaya, Selangor, Bukit Kutu,
May 1896, H. N. R{idley'] 7,659 (sing). Holo-
type examined!

SPECIMENS EXAMINED: Malaya, Pahang, Fra-
sers Hill, 19 Aug. 1937, */. H. Corner 33,233
(SING).

SECTION Aster o stigma

Pandanus stelliger Ridl., Roy. Asiat. Soc, Straits
Branch, Jour. 41: 49-50, (1904) =
1903; FI. Malay Penin. 5: 76, 1925; Mar-
telli, Webbia 4(2) : t 26, f. l-2a, 1914;
(sect. Aster o stigma)

Figs. 180, 181

DESCRIPTION OF LECTOTYPEl Shrub 1.5-33
m tall, the slender stems pale brown, 2.5 cm in
diameter, branched, bearing a few mucros;
leaves dark green, 55-60 cm long, near the base

Fig. 181. Pandanus stelliger Ridl., from lectotype, habit and infructescence, X Vi.

344

PACIFIC SCIENCE, Vol. XVII, July 1963

1.8 cm wide, near the middle 2. 8-3. 8 cm wide,
in section with a broad, central channel, and
hence M-shaped, the secondary parallel veins at
midsection 19-20 in each half, prominent, the
tertiary cross veins visible below, transverse or
oblique, forming short, somewhat oblong
meshes, the actual base amplexicaul and un-
armed, but beginning at 2-2.5 cm the margins
with prickles 1.5-2. 5 mm long, 4-9 mm apart,
broad subulate, ascending, pale; the midrib
unarmed except near the tip; at midsection the
margins with prickles 2. 1-2. 5 mm long, 7-23
mm apart, similar; near the 3-8 cm apex the
margins and midrib below with serrulations
0.2-0. 3 mm long, 2-6 mm apart, brown; pe-
duncle terminal, 5 cm long, leafy bracted, bear-
ing 4 syncarps in a dense spike about 10 cm
long; syncarps subequal, 5-6 cm long, 3-5 — 4.5
cm in diameter, ovoid to broadly ellipsoid, bear-
ing 64-88 drupes, these 19-21 mm long (in-
cluding the stigma), 7-10.5 mm wide, 6-8 mm
thick, oblanceoloid with an obtusely convex
apex, the sides 5-6-angled, smooth, when dried
brown, the body 15-17 mm long, cuneiform;
pileus 6-7 mm long, oblate semiorbicular; style
subterminal, 0.5-1 mm long, vertical except on
the basal drupes where sharply reflexed distally;
stigmas on the expanded apical surface, 4—7
mm Jm diameter, 4-7 -pointed, horizontal or
mushroom-shaped, papillose, brown; drupe body
14-17 mm long; endocarp centering in lower
34, bony, light brown, the inner surface shining,
the lateral walls 0.5 mm thick, the apical plug
1—1.3 mm thick; seeds 6-7 mm long, obdeltoid;
apical mesocarp with a cavity 6-7 mm long,
with a few white membranes near the margins;
basal mesocarp sparse, fibrous and fleshy.

EXPANDED DESCRIPTION OF ALL PISTILLATE
PLANTS EXAMINED: Plants 1.5-4 m tall; leaves
55-113 cm long, at midsection with 1 9—2 9 sec-
ondary parallel veins in each half; peduncle 5—7
cm long, bearing 2-5 syncarps, these 4-6 cm
long; drupes 7-13 mm wide, 6-10 mm thick,
bright yellow.

DESCRIPTION OF STAMINATE PLANTS: Leaves
63-70 cm long, near the base 20-21 mm wide,
near the middle 26-29 mm wide, at midsection
the parallel secondary veins 22-24 in each half,
and at midsection the margins with appressed
ascending serrulations 0. 3-0.4 mm long, 7-12

mm apart; staminate inflorescence 47 cm long,
leafy bracted; lowest bract 47 cm long, 2.2 cm
wide, for the lower 10 cm slightly distended,
sheathing and paler, the rest foliaceous; floral
bracts thin, apparently white, the median one
1 5 cm long, 4.6 cm wide, lanceolate, on the outer
34 the margins and midrib below with serrula-
tions 0.1 mm long, 0.2-1 mm apart; the 7
spikes 3-6 cm long, 12-15 mm in diameter,
cylindric, dense; fascicle 5-8 mm long, with
19-22 stamens, the naked common column 3-4
mm long, rhachis 2-2.5 mm long, clavate; free
filament tips 1-2 mm long, divergent; anthers
1.5-2 mm long, linear-oblong, bearing a pale
subulate projection of the connective 0.2-0.3
mm long.

LECTOTYPE: Malaya, "Selangor, on the Tras
Route at the 20th mile (8775),” 2,000 ft. elev.,
July 1897, H. N. Ridley (SING). Lectotype here
designated; specimen examined! Isotype ( FI ) !

SPECIMENS EXAMINED: Malaya: Perak, Larut,
top of mountains, open junfgle], 2,000 to 3,000
ft. alt., April 1883, H. Kunstler 4,166 (cal);
Haram Parak, June 1884, Scortechini ( CAL ) .

Pahang: Gunong Tahan, 7 Sept. 1937, E. J.
H. Corner (SING) ; Sungei Chelia, Chegar Perah,
14 Oct. 1927, M. R. Henderson 19,370 (sing).

Penang: Government Hill, Dec. 1895, C.
Curtis (SING); Waterfall Hill, 500 ft. alt., June
’88, L. Wray Jr. 2,227 (cal) .

SECTION Hombronia

Pandanus irregularis Ridl., FI. Malay Penin. 5:
76-77, 1925; Henderson, M. R., Roy.
Asiat. Soc., Malay Branch, Jour. 17: 82,
1939; (sect. Hombronia)

Figs. 182, 183

DIAGNOSIS OF HOLOTYPE: "Leaves long, very
broad,” 9-3 cm wide, thick and leathery, 1-
ribbed, 2 -pleated, in section depressed, AA-
shaped, the secondary parallel veins visible be-
low, scarcely so above and at midsection 83-92
in each half, the base amplexicaul and unarmed,
but beginning at 12 cm up the margins with
thorns 4.5 mm long, 1-2 cm apart, heavy deltoid
subulate, black-tipped, ascending; the midrib
below with similar reflexed thorns 2-3 cm apart;

to cm.

Fig. 182. Pandanus irregularis Ridl., from holotype. a, Infructescence, X lA ; b, drupe, lateral view, X 1;
c, drupe, longitudinal median section, X 1; d, drupe and stigmas,' apical view, X 1; e> drupe, transverse
median section, X 1; f, leaf base, lower side, XI \ g> leaf middle, lower side, X 1-

346

PACIFIC SCIENCE, VoL XVII, July 1963

Fig. 183- Pandanus irregularis Ridl., from holotype, leaf apex, lower side, X I-

on a leaf section cut higher up (exact place
unknown) the margins with spines 3-3.5 mm
long, 25-36 mm apart, heavy subulate, ascend-
ing; the midrib below with scars of fallen spines
3-6 cm apart; pistillate inflorescence 30 cm
long, with about 7 syncarps; peduncle 15 mm
in diameter, obtusely trigonous, bracted; syn-
carps 6. 5-7.5 cm long, subglobose, bearing
about 36-44 drupes, these 1-3- but mostly 2-
celled, 3-3.5 cm long, 1.8-2.4 cm wide, 1.2-1. 5
cm thick, "dark green,” narrowly obovoid to
oblanceoloid, compressed, the apex entire, ob-
tuse, the sides smooth, convex, lateral sutures
none; stigmas almost all eroded but 1 basal
drupe bears a single stigma 5 mm long, lanceo-
late, sharply bent to the horizontal, the recep-
tive surface distal; endocarp 22 mm long, cen-
tering in upper 36, at midsection filling the
drupe, bony, massive, dark brown but including
pale fibers, the lateral walls 4-5 mm thick; seeds

8-9 mm long, 3 mm in diameter, ellipsoid; api-
cal mesocarp cavernous, with strong, longitudi-
nal fibers and soft pith; basal mesocarp fibrous
and fleshy.

EXPANDED DESCRIPTION OF ALL SPECIMENS
EXAMINED: Leaf more than 1.4 m long, near
the base 9 cm wide, near the middle 12 cm
wide, near the apex 10.5 cm wide, broad ligu-
late; margins near the base with thorns 3. 5-4.5
mm long, 1-2.5 cm apart; midrib below with
thorns 3-3.5 mm long, 2-5 cm apart, subulate,
reflexed; at midsection the margins with spines
4-4.5 mm long, 20-25 mm apart, sharply as-
cending, subulate but the base much thickened;
the midrib below concealed; the apex apparently
damaged in growth and without a caudate apex,
rather contracted to 2 broadly obtuse lobes and
deeply emarginate, and even prickly on the
margins of the cleft, all this, however, appar-
ently an abnormality; peduncle 30 cm long;

Page 290: Revision of Pandcmus, 15. Malaya — St. John

347

spike 14 cm long, bearing 4 syncarps; stigmas
4 mm long, sharply proximally bent, the recep-
tive surface usually proximal.

HOLOTYPE: Malaya, Kelantan, Sungei Ketah,
Batu Bau (Bow), 15 Feb. 1924, Mo h anted Nur
& F oxworthy 12,104 (SING). Holotype exam-
ined!

SPECIMENS EXAMINED: Malaya, Pahang, Gua
Tipus, the common pandan on the limestone, 17
Oct. 1927, M. R. Henderson 19,468 (sing).

DISCUSSION: P. irregularis was placed by Dr.
Ridley in the section Rykia. It has, however, the
drupes 1-3 -celled, and usually 2 -celled, with
the cells in linear arrangement, and the large
stigmas terminal, but distal and laterally di-
rected. There is no question but that it must
now be placed in the section Hombronia.

It is one of the numerous plant species known
to occur only on limestone knobs. These pre-
cipitous pinnacles or narrow ridges rise spec-
tacularly from the rolling uplands of Thailand
and Malaya. Their sides are often precipices of
shining, white limestone, sparsely vegetated, but
their crests have a dark evergreen, virgin forest
cover. To a botanist they are very attractive.
Four species of Pandanus in Malaya are re-
stricted to these calcareous habitats. Besides this
species published by Ridley, two more are now
being described, and one more has yet to be
found in fruit. A good account of this localized
flora is given by Dr. M. R. Henderson (Roy.
Asiat. Soc., Malay Branch, Jour. 17: 13-87, pi.
Ill — XI, 1939).

Malaya, sect. nov.

Phalangibus pluricarpellatis, stylis terminali-
bus adscendentibus vel arcuatis corniformatis in-
tegris vel bifidis, stigmatibus centripetalibus el-
lipsoideis vel lanceolatis; plantis masculis in-
cognitis.

Phalanges of several united carpels; styles ter-
minal, ascending or arching, stout, hornlike with

a subulate apex or compressed and more or less
oblong, with the apex sharply bifurcate; stig-
mas centripetal, ellipsoid or lanceolate; stami-
na te plants unknown.

HOLOTYPUS: Pandanus Klossii Ridl., Fed.
Malay States Mus., Jour. 6: 190-191, 1915.

DISCUSSION : The single known species occurs
on the mountains of Pahang in Malaya. It com-
bines the characteristics of the section Rykia
in its hornlike, simple or bifurcate styles, and
of the section Pandanus, in its many-carpellate,
large phalanges. Since these two sections are
very distinct and are only remotely related, it
is best to make the very bizarre P. Klossii the
basis of a new section.2

Pandanus Klossii Ridl., Fed. Malay States Mus.,
Jour. 6: 190-191, 1915; FI. Malay Penin.
5: 74, 1925; (sect. Malaya)

Figs. 184, 185

DIAGNOSIS OF HOLOTYPE: Shrub 2-7 m tall;
stems usually solitary, 7.5 cm in diameter, gray,
erect or arcuate; leaves 1.3-1. 8 m long, 6.8-72
cm wide, thick coriaceous, above dark green,
below pale green, 1 -ribbed, but 2-pleated and
in section low M-shaped, the longitudinal paral-
lel secondary veins 0.5-0.8 mm apart, 49-52
in each half, evident on both sides as are the
tertiary cross veins that form a network of
meshes, oblong or rhombic, longer than broad,
the blade ligulate, and only near the tip rather
abruptly narrowing to a 5 cm stout subulate tip,
trigonous and 2-3 mm wide, the very base not
collected, near the base the margins with stout
thorns 5-8 mm long, 8-30 mm apart, arcuate
ascending, black; the midrib below with similar
but reflexed thorns; at midsection the margins
with prickles 3 mm long, 15-50 mm apart,
subulate, ascending, flat appressed; the midrib
below with prickles 2 mm long, 35-50 mm
apart, heavy, arcuate, ascending; near the tip

2 The new section can be inserted in the author’s key in this Revision of the Genus Pandanus (p. 2) by
replacing the 11th line from the bottom by:

L. Stigmas elongate, acerose, or stout hornlike, simple or bifurcate,

a. Stigmas stout hornlike, simple or bifurcate Malaya

a. Stigmas elongate, acerose,

FIG. 184. Pandanus Klossii Ridl., from holotype. a, Phalange, lateral view, X 1; phalange, longitudi-
nal median section, X 1', c, phalange, and stigmas, apical view, X 1; d, carpel apices and stigmas, apical
view, X 4; e, style and stigma, oblique view, X 4; f, leaf base, lower side, X 1; ^ leaf middle, lower side,
X 1; I), venation in outer third, lower side, X 4; i, leaf apex, lower side, X 1-

Page 292: Revision of Pandanus, 15. Malaya — St. John

349

the margins and midrib below with serrations
0. 3-0.5 mm long, those of the margin 1-1.5 mm
apart, those of the midrib below 2-5 mm apart;
pistillate peduncle 15-30 cm long, 12-18 mm
in diameter, 3-sided, leafy bracted; syncarp sin-
gle, globose or ellipsoid, about 15 cm in di-
ameter; phalanges numerous and 6-6.5 cm long,

3. 6- 3. 9 cm wide, 2-2.7 cm thick, cuneiform,
4-6-angled, "showing signs of turning red or
orange,’’ the sides smooth, gently convex, free
in upper J4, the apex low convex, lateral sutures
visible only in upper Vy central apical sinuses

1- 4 mm deep, straight or arcuate, narrow; car-
pels 9-10 (in a double 16-17 and the phalanges

5.7- 6 cm wide), the apices semiorbicular ,
faintly angled; styles 4-6 mm long, dark, bony,
either simple and hornlike or flattened and
sharply bifid, mostly centripetal; stigmas as long
as the styles, ellipsoid or lanceolate, brown, pap-
illose, covering the proximal face; proximal
sinus none; endocarp in lower 14 and 2.7 cm
long, bony, massive, dark brown, the lateral
walls 2 mm thick, the inner surfaces shining
and rugulose; seeds 14-16 mm long, 2-3 mm
in diameter, oblong-ellipsoid; upper mesocarp

2- 2.5 cm long, of dense pith and a few fibers;
basal mesocarp fibrous and fleshy.

ADDED DESCRIPTION FROM ALL SPECIMENS
EXAMINED: Phalanges about 96; carpels 5-10;
syncarp 9-15 cm in diameter.

HOLOTYPE: Malaya, Pahang, Gunong Tahan,
July 1911, H. N. Ridley 16,037 (SING). Iso-
type (bo). Ridley stated, "Common all over
the Padang. In the more open exposed spots
the stem is short and erect, about 6 to 8 feet
tall; in the woods the stems are long and weaker,
often falling about at all angles, 20 feet or
more long. No trace of male flowers could be
seen anywhere. The plant is very abundant, al-
most filling up the woods in some places." The
collection, Ridley 16,037, was made by Ridley
previous to his publication of the species in
1915, was labeled P. Klossii by him, and seems
to have been the only collection in the Singa-
pore herbarium up to that time. In his Flora
of the Malay Peninsula (1925) he accepted the
species and cited only his collection from Gu-
nong Tahan, so this one seems acceptable as
his holotype.

SPECIMENS EXAMINED: Malaya, Pahang,

Fig. 185. Pandanus Klossii Ridl., from holotype,
syncarp, X Vl-

Gunong Tahan, 14 Sept. 1937, E. J. H. Corner
(SING) ; Pahang, Gunong Tapis, Kuantan, 4,600
ft. alt., 14 June 1934, C. F. Symington & Kiah
28,843 (sing).

DISCUSSION: P. Klossii Ridl, a very unique
species, is here made the holotype for the new
section Malaya. This combines the characters of
the two sections Pandanus and Rykia. When
Ridley described the species in 1915 he did not
place it in a section, but later in 1925 in his
Flora of the Malay Peninsula he put it in the
section Keura, which is now called Pandanus.
As usual he had a good eye for species, but his
description seems to have been hastily drawn.
He stated ( 1915: 191 ) "the drupes are separate

350

PACIFIC SCIENCE, Vol. XVII, July 1963

till the fruit is nearly ripe, when from 6 to 7
become adnate and remain so as the whole fruit
breaks up.” The holotypic collection consists of
two sheets, one with overripe fruit, the other
with a small, immature syncarp. In both the
phalanges are of several fused carpels. The car-
pels are fused from the base to within 4 mm
of the apex, and the sides lack sutures.The writer
sees no evidence on these specimens to confirm
Ridley’s statement that the drupes are separate
and only tardily connate; on the contrary, all
the structural evidence indicates that the pha-
langes are of several carpels united from the
beginning.

Though Ridley did not explain the source
of his specific epithet, it was obviously given in
honor of Cecil Boden Kloss, an English zoolo-
gist and to a lesser degree a botanist. In 1903
and 1907 he was employed by the Singapore
Botanic Garden. He was later director of the
Raffles Museum.

Multidens, sect. nov.

Frutices, foliis angustis, syncarpio solitario el-
lipsoideo, stylo singulo terminali osseoso prox-
ime curvato basi cylindrico margine (2-) 3-6-
lobato lobis deltoideis, drupis 1-loculatis. Plantis
masculis cum inflorescentia bracteata 1-spicata,
staminibus distinctis in axili affixis, filamentis
4-plo tarn longis quam antheris.

Bushes; leaves narrow; syncarp solitary, el-
lipsoid; style single, terminal, proximally curved,
bony, shining, the base cylindric, the margin
(2-) 3-6-lobed, the lobes deltoid; drupes 1-
celled. Staminate plants with inflorescence brac-
teate, bearing a solitary terminal spike; stamens
separate, attached directly to the axis; filament
4 times as long as the anthers.

HOLOTYPUS: P. pentodon Ridl., FI. Malay
Penin. 5: 75, 1925. Also in the section is P.
tetrodon Ridl. of Singapore.

Pandanus pentodon Ridl., FI. Malay Penin. 5:

75, 1925; (sect. Multidens)

Fig. 186

diagnosis OF lectotype: Stem short,
branched, up to 2 cm in diameter; leaves said
to attain more than 100 cm in length, 11-12
mm wide, coriaceous, dark glossy green, pale
green beneath, channeled above the midrib and
with 2 lateral pleats, in section low M-shaped,
narrowly ligulate, outer halves of leaves missing,
at midsection with 17-22 secondary parallel
veins in each half, the tertiary cross veins barely
visible below near the base, the base amplexi-
caul and unarmed, apparently copper-colored,
beginning at 3-5 cm up the margins with pale
serrulations 0. 5-0.8 mm long, 1-6 mm apart;
the midrib unarmed to at least beyond the mid-
dle; at midsection the margins with slender ser-
rulations 0.3 -0.6 mm long, 4-8 mm apart, the
upper surface unarmed; the apex tapering to a
10 cm trigonous subulate tip which 10 cm
down is 4 mm wide, the margins and midrib
below with serrations 0. 2-0.3 mm long, 0.7-3
mm apart; pistillate peduncle 13 cm long, 8 mm
in diameter, bracted, bearing a single, erect
syncarp; the highest peduncular bract 30 or
more cm long, at base 2.2 cm wide, narrowly
lanceolate and tapering noticeably right from
the base, the margins with serrulations similar
to those of the leaves, but on the upper surface
beginning 10 cm up the 2 lateral pleats with
similar serrulations, the lower ones 5-13 mm
apart, the upper ones 2-5 mm apart; partly
hiding the fruit are 3 series of floral bracts, 3
deep, and subequal, 5 cm long, 2 cm wide, lance-
elliptic, fleshy, cream-colored; syncarp 5 cm
long, 3 cm in diameter, bearing about 400
drupes (when young these are cream-colored),
12-13 mm long, 2-3.5 mm wide and thick,
linear-oblanceoloid, 5-6-angled, the body 10-11
mm long, the summit rounded; pileus 2.5-3

Insertion in Key

On p. 225 (p. 2 of reprint) for the second J, read
J. Styles 1 (-2), often cartilaginous or bony,
z. Styles toothlike to broad subulate, entire or bifid; staminate spikes bearing stamens

fascicled at apex of column Rykia

z. Style with cylindric base, then curving proximally and the margin (2-) 3-6-lobed
with deltoid lobes; staminate spike solitary, bearing separate stamens attached directly
to the axis Multidens

Fig. 186. Pandanus pentodon Ridl., from lectotype. a, Infructescence, X 1; ^ drupe, lateral view, X 1;
c, drupe, longitudinal median section, X 1 \ d, drupe, lateral view, X 4; e, drupe, longitudinal median sec-
tion, X 4; /, drupe and style, apical view, X 1; g, drupe apex and style, apical view, X 10; h, stigma, oblique
view, X 10; i, drupe apex, style, and stigma, lateral view, X 10; j, leaf base, lower side, XI; ^ leaf middle,
lower side, X 1; l, leaf apex, lower side, Xl;», staminate inflorescence, X 1; «, stamens and axis, X 10.

io cm.

| —I — T“ ' ' I

o 5 cm.

FIG. 187. Pandanus tetrodon Ridl., from holotype. a, Syncarp, X 1; b, drupe, lateral view, X 1; drupe,
longitudinal median section, X 1; bl-g, drupe, style, and stigma, apical view, X 1; b, drupe, lateral view, X
4; i, drupe, longitudinal median section, X 4; j, k, drupe, style, and stigma, apical view, X 4; /. stigma,
proximal view, X 4; m. leaf base, lower side, X 1; n> middle, lower side, X 1-

i cm.

Page 296: Revision of Pandanus, 15. Malaya — St. John

353

mm long, the base pyramidal-semiorbicular,
smooth, when dried somewhat yellowish; style
1-1.5 mm long, apical, bony, yellowish, shining,
its base short cylindric, then the rest sharply
curved proximally and flabellate with the mar-
gins (3-) 4-5 (-6) dentate; stigma 0.5-1 mm
long, obdeltoid to flabellate, papillose, brown,
on the distal side of the style; endocarp central,
bony, pale brown, the upper lateral walls 1 mm
thick; seed 5-5.5 mm long; apical mesocarp
an empty cavern; basal mesocarp fibrous up the
sides, fleshy within.

STAMINATE PLANT: Herbage not seen; stam-
ina te inflorescence dense, leafy bracted, the main
part 14 cm long; lowest floral bract about 30
cm long, the upper part green, foliaceous, 10
mm wide, with serrulations 0.2 mm long, 2-5
mm apart, the 10 cm basal part 19-20 mm wide,
folded, firm, apparently pale, with similar or
smaller serrulations; median floral bract 7 cm
long, 3 cm wide, navicular, elliptic, acute, ser-
rulate near the apex on margins and midrib
below, firm, apparently pale; spike single, ter-
minal, 7.5 cm long, 13 mm in diameter, cylin-
dric, dense; stamens distinct, attached directly
to the axis; filaments 4.2-4.4 mm long, anthers
0.8 mm long, narrowly oblong, bearing at tips
a 0.15-0.2 mm subulate prolongation of the
connective.

LECTOTYPE: Malaya, Johore, Ulu Kahang,
abundant along edges of Kahang river, 250 ft.
alt., 1 June 1923, R. E. Holttum 10,861 (sing)
— the pistillate plant here designated as lecto-
type. Mingled on the same sheet is a staminate
plant.

Pandanus tetrodon Ridl., Roy. Asiat. Soc., Jour.
68: 13-14, 1915; P. singaporensis Ka-
nehira, Jour. Jap. Bot. 14: 173-177, figs.
6-8, 10, 1938, a superfluous name; non
Barrotia tetrodon Gaud., Bot. Voy La
Bonite, t. 13, figs. 1-8, 1841, an invalid
name; non P. tetrodon Balf. f., Linn.
Soc. Bot. Jour. 17: 63, 1878, a name
not accepted by its author; non P. tetro-
don Balf. f. ex Kanehira, Bot. Mag. To-
kyo 52: 236-239, fig. 70, 1938, a later
synonym of P. compressus Mar tel li
( 1905 ) ; ( sect. Multidens )

Fig. 187

DIAGNOSIS OF HOLOTYPE: "Large bushy
plant; branches at apex 2.5 cm in diameter”;
leaves well over 97 cm long, 2.6 cm wide at
the middle, 2 cm wide near the base, coriaceous,
1 -ribbed, 2 -pleated, in section low M-shaped,
about at the midsection with 25-26 secondary
parallel veins in each half, these visible at least
below throughout, tertiary cross veins visible in
outer half, transverse, forming oblong meshes,
mostly short, the blade ligulate, the apex not
preserved, the base amplexicaul, unarmed, be-
ginning at 5 cm up the margins with prickles
1.5-2. 5 mm long, 5-10 mm apart, subulate-
tipped serrae, pale, with reddish tips; the mid-
rib below unarmed for at least lower %’s; near
the midsection the margins with similar serrae,
1.3-1. 5 mm long, 6-13 mm apart; leaf apex
"acuminate,” but not preserved; pistillate in-
florescence terminal, more or less concealed; pe-
duncle 19 cm long, 7 mm in diameter, 3-sided,
sinuous, leafy bracted, bearing 1 syncarp, this
8.8 cm long, 5 cm in diameter, cylindric with
obtuse ends, bearing about 176 drupes, these
22-24 mm long, 6—10 mm wide, 5-10 mm
thick, increasing upwards in size and the apical
ones definitely the largest, cuneate-oblanceoloid,
obtuse, 5-6-angled, upper % free, the sides
nearly smooth; style 3-4 mm long, bony, dark
shining brown, at base cylindric, then quadrate
and curved proximally, and all the lateral ones
with an oblique plane to the 2-3-4-dentate
margins; stigma 1—2 mm wide or long, rugose,
of various shapes, filling the cavity proximal
of the style apex, with 2-4 (-5) deltoid lobes;
endocarp centering in lower Vs, bony, dark
brown; seed 6 mm long, obdeltoid; apical meso-
carp 6-7 mm long, a rounded cavity with a few
pale membranes; basal mesocarp fibrous and
fleshy.

EXPANDED DESCRIPTION OF PISTILLATE
PLANTS: Stems at apex 1.5-2. 5 cm in diameter,
brown, striate, covered with ascending adventi-
tious rootlets 1-2 cm long; leaves 78 to more
than 97 cm long, the tip gradually diminishing
to a trigonous subulate apex about 10 cm long,
and at base 4 mm wide, on the subulate apex
the margins and midrib below with serrations
0.3-0. 5 mm long, 2-5 mm apart; syncarp sin-
gle or with a secondary one below and 4.5 cm
long, ovoid.

Fig. 188. Pandanus immersus Ridl., from holotype. a, Syncarp, X 1; h, drupe, lateral view, XI \ c, drupe,
longitudinal median section, X lj d—i, drupe, style, and stigma, apical view, X 1 ’> j> drupe, lateral view, X 4;
k, drupe, longitudinal median section, X 4; l, drupe apex and style, apical view, X 4; m, drupe apex, style,
and stigma, proximal view, X 4; n, leaf middle, lower side, X 1; venation of leaf middle, lower side.
X 4; p, leaf apex, lower side, X 1-

Page 298: Revision of Pandanus, 15. Malaya — St. John

355

HOLOTYPE: Singapore, on the west coast road
near Pasir Panjang, Dec. 11, 1911, H. N. Ridley
15,465 (SING). Holotype examined!

SPECIMENS EXAMINED: Singapore, Botanic
Gardens, Aroid Rockery, cult., 20 April 1936,
E. J. H. Corner 30,993 (kep, sing).

SECTION Rykia

Pandanus immersus Ridl., Asiat. Soc., Straits
Branch, Jour. 41: 48-49, 1903; Mater.
FI. Malay Penin. 2: 224, 1907; FI. Malay
Penin. 5: 76, 1925; Martelli, Webbia
4(1): 18, 1913; (sect. Rykia )

Fig. 188

NOM. VERN.: "mengkuang ayer.”

DIAGNOSIS OF HOLOTYPE: Aquatic, "the stem
rising but little above” the surface of the river;
leaves "many feet long,” 10.2 cm wide, coria-
ceous, green above, pale and glaucous beneath,
the longitudinal veins strong, visible above and
conspicuous below, about 1 mm apart, at mid-
section with 70 secondary parallel veins in each
side, the venation also consisting of tertiary
cross veins 1-3 mm apart, these easily visible
above and conspicuous below, the base not pre-
served but the blade 1 -ribbed, 2 -pleated, in sec-
tion M-shaped, ligulate, narrowing upwards and
in the upper quarter tapering to a subulate del-
toid apex about 30 cm long, and at a point
about 10 cm down from the tip only 1.3 mm
wide; at about the midsection the margins with
thorns 2.5-4 mm long, 9-18 mm apart, subulate
but the base thickened, nearly straight, ascend-
ing, pale except for the very tip which may be
reddish; near the tip the margins with serrae
0.8-0. 9 mm long, 1.5-3 mm apart, stout; the
midrib below nearly unarmed; pistillate inflores-
cence apparently emersed; peduncle more than
8 cm long, 15 mm in diameter, 3 -sided, brac-
teate; syncarp perhaps a third mature, 9.5 cm
long, 6 cm in diameter, ellipsoid, 3 -sided, bear-
ing about 1,440 drupes, these 17-19 mm long,
5-6 mm wide, 3-4 thick, the body 12-13 mm
long, arcuate spatulate, upper free, cuneate
to the base; the pileus 7-8 mm long, the base
subsemiorbicular, minutely papillose; the style
4-6 mm long, hornlike, mostly gently curved
proximally, the apical ones unbranched, of the

median and lower ones about half are entire
with a single horn but the other half are bifur-
cate and the 0.5-2 mm lobes often are wide-
spreading; stigma 3.5-5 mm long; endocarp
(immature) oblanceoloid, nearly as long as the
body; mesocarp fibrous and fleshy.

HOLOTYPE: Malaya, "Selangor, in the Labu
River, forming dense thickets,” May 1891, H.

N. R[idley] , (SING). Holotype examined!

Pandanus johorensis Martelli, Soc. Bot. Ital., Bui.
302, 1904; P. muarensis Ridl., Mater.
FI. Malay Penin. 2: 226-227, 1907, and
FI. Malay Penin. 5: 77, 1925; (sect.
Rykia )

Figs. 189, 190

NOM. VERN.: "rassau.”

DIAGNOSIS OF ISOTYPE: Leaves 1.3-1.45 m
long, 4.2 cm wide, coriaceous, paler beneath,
1 -ribbed, 2 -pleated, at midsection with 43 sec-
ondary parallel veins in each half, no visible
cross veins, ligulate, gradually tapering to a
27 cm subulate deltoid apex, this 10 cm down

O. 7 mm wide, the base amplexicaul, unarmed,
but beginning at 10-11 cm on one side, and
at 18 cm on the other the margins with prickles
1.5-2 mm long, 4-11 mm apart, subulate, as-
cending, dark reddish; the midrib below begin-
ning at 8 cm with prickles 3-3.5 mm long,
12-22 mm apart, stout subulate, arcuate, re-
flexed; at midsection the margins with prickles
1-1.5 mm long, 4-14 mm apart, stout subulate,
arcuate, closely ascending; the midrib below
with prickles 1.5-1. 8 mm long, 5-45 mm apart,
stout subulate, arcuate, reflexed or ascending; on
the subulate apex the margins with serrae 0.5
mm long, 1-3 mm apart; those of the midrib
below similar but 3-7 mm apart; peduncle 29
mm long, 8 mm in diameter, 3 -sided, slightly
sinuous, leafy bracted; syncarp single, 6 cm long,
3 cm in diameter, narrowly ellipsoid (very
young), bearing about 4,096 drupes, these (im-
mature) 11-12 mm long, 1.5-2. 5 mm wide and
thick, 5-6-angled, upper 2A free, the body 4.5-5
mm long, oblong-ellipsoid, papillose, with thick
longitudinal fibers; style 7 mm long, setiform,
hard, shining and yellow; stigma linear, proxi-
mal, brown, as long as the style; endocarp and
seed not yet formed, but inside the ovary wall

Fig. 189. Pandanus johorensis Martelli, from isotype, a, Syncarp, XI \ b, c, drupe, lateral view, X 1; d,
drupe, lateral view, X 4; e, drupe, longitudinal median section, X 4; /, drupe apex and style, apical view,
X 4; g, style and stigma, proximal view, XI 0; h, leaf base, lower side, X 1; i> leaf middle, lower side, X 1;
j, leaf apex, lower side, X 1.

UtOOT

Fig. 190. Pandanus johorensis Martelli, from Corner 28,161. a, Staminate inflorescence, X l/3\ b, spike,
X 1; c, staminate column, scale, and stamens, X 10; d, leaf base, lower side, X 1; ^ leaf middle, lower side,
X 1; f, leaf apex, lower side, X 1 •

UUUS

io cm.

358

PACIFIC SCIENCE, Vol. XVII, July 1963

Fig. 191. Pandanus pevangensis Ridl., from holotype. a, Infructescence, X 1/5; b, drupe, lateral view,
X 1; c, drupe, longitudinal median section, X 1; d, drupe apex and style, X 1; drupe apex and stigma,
proximal view, X 4; /. leaf base, lower side, X 1; g, leaf middle, lower side, X 1; ^ leal apex, lower side,

X I-

Page 302: Revision of Pandanus, 15. Malaya — St. John

359

of tissue and heavy longitudinal fibers there is
a single large cavity, with indefinite traces of
inner tissue.

EXPANDED DESCRIPTION FROM ALL PISTIL-
LATE PLANTS EXAMINED: Shrub, to 6 m tall,
upper branches 3-4 cm in diameter, dark brown,
forming dense thickets, with a few thornlike
axillary rootlets 4-6 mm long, divergent; leaves
1-1.8 m long, 2.9-4. 1 cm wide; syncarp solitary,
pendulous, 12-16 cm long, 6-10 cm in diame-
ter, ellipsoid.

DESCRIPTION OF STAMINATE PLANTS: BraC-
teal leaves up to 80 cm long, 3.5 cm wide, coria-
ceous, apparently pale, ligulate, the upper lA
long tapering to a trigonous subulate apex, this
10 cm down 2 mm wide, the lower wholly
unarmed, at midsection with 5 1 secondary veins
in each half, no visible tertiary veins; near the
apex the margins and midrib below with subu-
late serrulations 0.5-0.7 mm long, 2-6 mm
apart; staminate inflorescence 45 cm long, with
8 spikes, these 15-25 cm long, 12-15 mm in
diameter, densely flowered but somewhat in-
terrupted; flowers sweet and sickly scented; fila-
ment column 4 mm long, stout, clavate, bearing
an apical parasol-like scale 3-5 mm in diameter,
the apex hispidulous with hairs 0. 3-0.4 mm
long; free filament tips 0.5-2 mm long, pendent
from the lower surface, glabrous or sparsely
hispidulous; anthers 10-16, and 0.8-1. 3 mm
long, mostly cylindric, bearing an apical mucro
0.2 mm long, formed of a projection of the
connective (description from Corner 28,161).

ISOTYPE: Malaya, Johor, Muar, Bukit Wiligu,
Sungei Pauh, 1902, Fox 11,326 (SING). Isotype
examined!

SPECIMENS EXAMINED: Malaya, Johore, Ma-
wai, generally flowering of staminate and pistil-
late on riverside, flowers visited by bees and
beetles, May 13, 1934, E. J. H. Corner 28,161
(SING).

DISTRIBUTION: Malaya and Sumatra, form-
ing dense thickets in estuaries, forming a zone
beginning where Nip a stops, in tidal fresh wa-
ter. Plants with similar leaves have been seen
from Borneo.

DISCUSSION : P. johorensis Martelli is a mem-
ber of the section Rykia. It was briefly pub-
lished in 1904 by Martelli as only a few phrases
in a key to certain species. His holotype was:

"Stato di Johor. Penis. Malese, no. 11326 Ridl.
(H. Becc.).” The specimen above cited is an
isotype in the herbarium in Singapore. It bears
the data: "Malay Peninsula, Johor, Bukit Wilgu,
Sungei Pau, 1902, Fox 11,326.” In his Materials
for a Flora of the Malayan Peninsula 2: 226,
1907, Ridley accepted Martelli ’s species, and
cited the single collection, "Muar: Bukit Wilgu,
Sungei Pauh (W. Fox) 11326.” Later, Ridley
in his Flora of the Malay Peninsula 5: 77, 1925,
treated this plant as P. johorensis sensu Ridl.,
not of Martelli, and redescribed it as the new
species P. muarensis Ridl. For this he had the
holotype: "Muar, Sungei Pauh (Fox).” What-
ever misconception Ridley thought there was,
he had no right to rename the species using the
same holotype. Certainly the collection Fox
11326 from Bukit Wilgu, Sungei Pauh, Johor,
is the same as [Ridley] 11,326 with the same
locality data. The number is a Singapore Her-
barium number, as the staff members were re-
quired to use this single series of numbers, not
their own personal collection numbers. The hol-
otypic specimen in the herbarium at Firenze was
not a duplicate distributed from Singapore.
Rather, it is a fragment consisting of about two
score of the drupes placed in a paper pocket on
which in Martelli’s hand is written the data.
These fragments were taken from the syncarp
of the isotypic specimen, then in Calcutta, now
in Firenze. It lacks stem, leaves, inflorescence,
and syncarp. Nevertheless, the actual holotype
is this fragmentary specimen, Fox (or Ridley ),
once in Calcutta, and now in Firenze. However,
there seems no doubt but that P. muarensis Ridl.
is a typonym and must be placed in the syn-
onymy of P. johorensis Martelli. The isotype in
Singapore has a syncarp with immature drupes
that have not matured their endocarp and seed.

Pandanus penangensis Ridl., Asiat. Soc, Straits

Branch, Jour. 41: 50, (1904) = 1903;

FI. Malay Penin. 5: 81, 1925; (sect.

Rykia )

Fig. 191

diagnosis OF HOLOTYPE: "Tree 7 m. tall,
10 cm. in diameter”; leaves 2 m or more long,
near the base 12 cm wide, when young marbled
light and dark green, beginning at 15-16 cm

360

PACIFIC SCIENCE, Vol. XVII, July 1963

the margins with prickles 1.3- 1.5 mm long, 4-7
mm apart, pale, subulate, flat appressed ascend-
ing; the midrib below sharp salient and with
similar ascending prickles, the secondary parallel
veins conspicuous and in lower part 69 in each
half, and except at base the tertiary cross veins
prominent on both sides, transverse or oblique
and forming short, rather oblong meshes, mid-
section not seen; at 45 cm down from the apex
the blade 11.5 cm wide, the apex abruptly nar-
rowed to a subulate tip about 5 ( ? ) cm long, the
margins with subulate serrae 1.5-2. 5 cm long,
3-10 mm apart; the midrib below with serrae
0.8-1 mm long, 7-20 mm apart; drupes 42-47
mm long, 7-9 mm wide, 6-7 mm thick, 5-6-
angled, the sides smooth, gently curved, prob-
ably only the pileus free, the body 37-40 mm
long, drupes and pilei shedding separately;
pileus 9-11 mm long (or if measured along
the curve of the style 13-16 mm long), 10-13
mm wide, 8-10 mm thick, 6-angled, broad
ovoid-pyramidal, the tip proximally curved;
style 5-7 mm long, heavy subulate, curved

proximally almost to a right angle; stigma 4-5
mm long, lanceoloid, papillose, brown; endocarp
32-33 mm long, narrow ellipsoid, bony, ebony-
colored, the inner surface shining, the lateral
wall 1-1.5 mm thick; seed 15-17 mm long,
4-6 mm in diameter, ellipsoid; apical and basal
mesocarp fibrous and fleshy.

EXPANDED DESCRIPTION FROM ALL SPECI-
MENS examined: Leaves to 4 m long, the mid-
section 10 cm wide, the margin with prickles
2.8-3 mm long, 5-14 mm apart, stout, heavy
based, arcuate subulate, appressed ascending,
brown; pistillate inflorescence of 4-5 syncarps,
each with numerous drupes; syncarps 15-20 cm
long, 9.5-10 cm in diameter, ellipsoid.

HOLOTYPE: "Malaya, Penang Hill from base
to the top, in woods.” The specimen is labeled:
Malaya, Pulau Penang, Fort Hill, 1900, H. N.
Ridley (SING). Holotype examined!

SPECIMENS EXAMINED: Malaya, Pulau Pe-
nang, Government Hill, common on the hill
top, but not often in fruit, 29 July 1907, L H.
Burkill 2,666 (sing).

Lord Howe Island, A Riddle of the Pacific, Part III

S. J. Paramonov1

In this final part (for parts I and II see
Pacif. Sci. 12 ( 1 ) : 82-91, 14 (1): 75-85) the
author is dealing mainly with a review of the
insects and with general conclusions.

INSECTA

Our knowledge of the insects of Lord Howe
Island is only preliminary and incomplete. Some
groups, for example butterflies and beetles, are
more or less sufficiently studied, other groups
very poorly.

Descriptions of new endemic species and
records of the insects of the island are dis-
persed in many articles, and a summary of our
knowledge in this regard is lacking. However,
a high endemism of the fauna is evident. Al-
though the degree of endemism is only at the
specific, or at most the generic level, the con-
nection with other faunas is very significant.

Olliff (1889) wrote an interesting review
of Coleoptera of the island. In his time about
80 species of Coleoptera were recorded, but
among them were 46 new species and 5 doubt-
fully new. That is, more than 56% were en-
demic species. Similar reviews for other groups
are lacking.

An interesting large, wingless phasmid with
a body length of 106 mm, Dryococelus (for-
merly Carabidion) australis Montr., is now prob-
ably extinct. Gurney (1947: 391) wrote: "The
present status of Dryococelus is uncertain. A
note in Ward’s Natural Science Bulletin (vol.
8, No. 2, p. 11, 1935) indicated that the species
is extinct, but I have been informed by John
W. H. Rehn, of the Academy of Sciences of
Philadelphia, that he has seen a recently pub-
lished note suggesting that the species is main-
taining itself.”

1 Commonwealth Scientific and Industrial Research
Organisation, Canberra, Australia.

Manuscript received February 27, 1962.

During two visits to the island, in 1954 and
1955, the author failed to find the insect. An
official enquiry was made recently to the Ad-
ministration staff of the island, and the author
received a letter from the Superintendent of
the Island, Mr. H. Ward, on Nov. 3, 1961, in
which he states: "A number of the old inhabi-
tants have been questioned and all have advised
that it is at least 30 years and possibly 40
years since this insect has been seen on the
Island. A member of the staff, aged 33 years,
has never seen or heard of the insect, nor has
any pupil of the local School.”

The only possibility is that the insect may
still exist in one of the biggest banyan trees
on the slope of Mt. Gower, on the lagoon side.
The area is well isolated from the settlement
where the rat concentration was probably the
greatest, and may have survived in crevices of
the tree.

The presence of the insect in the past is
evidence of an ancient connection with Aus-
tralia or some other continent by a land bridge;
any other type of transportation is highly
improbable.

A very interesting example of distribution
is found in an archaic Homopterous family,
the Pelorididae, with very limited possibilities
of movement (nearly wingless), and adapted
to very restricted habitat. There are 15 species
recorded now for the whole world. Thirteen of
them are associated in Chile, New Zealand,
Tasmania, and eastern Australia with the south-
ern beech Nothofagus. The plant also grows
in New Caledonia, and Dr. J. W. Evans (Di-
rector of the Australian Museum) visited this
island especially to search for Pelorididae. None
were found, nor was an environment discov-
ered which would be favourable for their
existence.

In 1959 J. W. Evans, having been informed
that a single nymph was recorded from Lord
Howe Island, visited the summit of Mt. Gower

361

362

PACIFIC SCIENCE, Vol. XVII, July 1963

Fig. 1. A typical cloud "carpet” covering the summit of Mt. Gower while the rest of the island is in sun-
shine. This "carpet” preserves the humidity and coolness over long periods, in strong contrast with other parts
of the island. (Photo by Miss Z. Liepa.)

and during 3 hr (sic) found two new species
of this family.

It is interesting to note that Nothofagus is
absent from Lord Howe Island, but 24 adults
and 7 nymphs of Pelorididae were found in
moss. Probably the Pelorididae are not espe-
cially associated with Nothofagus , but require,
above all, high humidity with cool temperatures.
It is evident that this family has lived on the
island in a very specialised habitat for millions
of years. The possibility of transportation must
be excluded. It means that the general condi-
tions for life are unchanged: the summit of
Mt. Gower is a refuge for species ecologically
close to recent New Zealand fauna and flora
( see Fig. 1 ) , whereas at sea level are preserved
elements from the north, from purely tropical
areas.

It is noteworthy that, of the endemic palms,
two dwarf species grow not at sea level, as do

two others (see Fig. 2), but much higher up,
closer to the summits of the two large moun-
tains.

The submerged land "Howeania” was iso-
lated from Australia, or connected only in the
far north, and the present "Australian” elements
we must regard as casual elements, not as hav-
ing come by a direct land bridge. The Austra-
lian elements play an important role in the
composition of the flora and fauna of the
island, but it is remarkable that the most typical
Australian forms, the Australian "sui generis,”
are absent.

The Diptera of the island have not been
reviewed, the main obstacle being the lack
of knowledge of the fauna of surrounding
areas. The dipterofauna in general is very im-
poverished, because the island lacks fresh water
basins, and the running streams are of tem-
porary character. Therefore the families and

Lord Howe Island, Part 3 — PARAMONOV

363

genera connected in their life history with
water are absent. Sandy ecological niches are
also poorly represented, and in general the eco-
logical uniformity is a factor which limits the
richness in flies.

However, many new species were discovered
from the families Asilidae, Muscidae, Ortali-
dae, Leptidae, Calliphoridae, Tachinidae, etc.
(see references).

The author has the impression, however,
that most of the endemic species are repre-
sented in the surrounding areas by closely re-
lated species. Nothing extremely old or "extra”
was found; however, only the lower part of the
island has been comparatively studied. The
middle zone (about 1,000 ft) and the high
zone (2,000 ft and above) are practically un-
touched. The small fresh-water streams and
pools deserve the special attention of collectors.

The author observed the hatching of enor-
mous numbers of hmttia and Sarcophaga from

the sand on the beach. They emerged early
in the morning, and with wings still undevel-
oped walked onto the small rocks on the shore,
the surface of some of the rocks being abso-
lutely covered with them. Here they waited
until their wings were developed and then
flew away. The sandy area is covered by sea
during high tide, but the puparia apparently
are not harmed. The author observed no hatch-
ing from the sandy areas not reached by the
high tide.

To illustrate the spate of our knowledge of
the Lord Howe Island fauna we may quote
the results of J. D. Bradleys collecting in 1953:
in Wi days of collecting, among 22 specimens
of Microlepidoptera there were 12 species, 8 of
which were new, and 4 others which were Aus-
tralian species not previously recorded for the
island.

The island is also very suitable for experi-
ments in insect biological control. In October

Fig. 2. Two endemic species of palms which dominate the lower part of the island. Before the last war
the export of their seeds was an important industry of the island. (Photo by Miss Z. Liepa. )

364

PACIFIC SCIENCE, Vol. XVII, July 1963

Fig. 3. An "Arcade” on Admiralty Island. The greyish dots in the air are sea birds which nest there. The
basalts form the main mass of the island. (Photo by Miss Z. Liepa.)

1959, specimens of Optus oophilus were liber-
ated there to control the Queensland fruit fly,
Strumeta tryoni, and the parasite was recov-
ered in March 1961. The small size of the
island greatly facilitated the experiments.

It is impossible to go into details about the
insect fauna of the island as the available data
are too fragmentary. However, some points are
very significant, especially those concerning
wingless insects (and birds); they show a close
relationship to the New Zealand fauna and
to the eastward land masses in general. The
Australian element, considered numerically,
may be larger than that of New Zealand or
Polynesia, but the proximity of the Australian
continent, its great extension northwards and
southwards, and the direction of the prevailing
winds in the southwest Pacific no doubt ac-
count for this preponderance. If we take as an
example the plant genera which are confined
to Australia and Lord Howe Island (Notelaea,

Melaleuca , Lyonsia, Lagunaria, and W estringia) ,

we must not overestimate the importance of
this fact, because the most characteristic Aus-
tralian genera are either- entirely absent from
Lord Howe Island or represented by only one
or two species. This means that these species
have had more facilities for transportation,
natural or by man.

Summarizing all the data in our hands, we
can say from its fauna that Lord Howe Island
is not a part of the Australian zoogeographical
region, but is closer to the New Zealand sub-
region.

GEOLOGY

In order to understand the history of the
fauna and flora the geology also must be con-
sidered. The general geology of the island is
very simple; the island consists mainly of two
principal formations: the volcanic rock which

Lord Howe Island, Part 3”— PARAMONOV

363

forms the general mass (see Fig. 3), and the
stratified beds resting on it (see Fig. 4). Two
thirds of the island is composed of volcanic
rocks, comprising three isolated masses.

Edgeworth (1889: 132) summarized his
study of rocks as follows:

1. All the igneous rocks of Lord Howe Island (so
far as represented by the collection examined)
belong to the Basalt group.

2. A vast period of time must have elapsed between
the eruption of the diabasic basalt and that of
comparatively recent olivine basalt.

3. All the basalts, with the exception of the diabasic
types, are probably not earlier than Tertiary,
and some may be Post -Tertiary.

4. The diabasic basalt is probably Pre-Tertiary, or
may be Paleozoic.

Immediately overlaying the volcanic rocks,
and between them and the succeeding coral-
sand rock series, occurs a bed of stiff unctuous
red or yellow clay. It does not appear to be
fossiliferous.

More interesting is the coral-sand rock se-
ries; this deposit is often from 30 to 40 ft
high. The coral-sand rock is the chief fossilifer-
ous deposit of the island, and has yielded the
remains of the interesting reptile Meiolania,
eggs of turtles, bird-bones, and recent species
of both land and marine shells.

A systematic boring of this deposit, as well
as of coral beds in the sea, may give us a de-
tailed picture of the island’s physical history.
Unfortunately, geological study is only at a
preliminary stage, and when it was stated that
no minerals of economic value existed on the
island further investigations were stopped.

The island deserves, however, our special
attention as a very convenient object where
the history of Australia, of the surrounding
areas to the east, and of the Pacific may be
studied much more easily than at other points.

The impoverished fauna and flora and the
simple geological structure may give us all

Fig. 4. A typical stratified deposit at the eastern seashore east of Old Gulch; the coral-sand strata are
prominent. (Photo by Miss Z. Liepa.)

366

PACIFIC SCIENCE, Vol. XVII, July 1963

Fig. 5. Map showing the area of "Howeania.”

the necessary data, because of the small volume
of material for study. Lord Howe Island may
be the focal point for the study of this area
of the Pacific.

An enormous number of facts about the
Australian continent remain to be studied, and
this will require many years of work, but the
general outline of the history of organisms
and environment may be elucidated much more
rapidly if we know the history of Lord Howe
Island. For developing synthetic conclusions
the island represents an extremely useful area.

COMPOSITION OF THE FAUNA AND FLORA

The fauna and flora of the island probably
consists of three different layers of animals
and plants:

1. The remnant from the very large land
mass, now submerged, which we have called
"Howeania.” This remnant is the oldest part
of the fauna and flora.

2. Immigrants from surrounding areas, ar-
riving after the land was submerged, i. e., very
long ago, in the geological sense. These ele-
ments have had time to develop into new
species.

3. Recent immigrants, helped directly or in-
directly by man. We have some data which
indicate that in recent time several species
arrived on the island, but finding conditions
unsuitable for permanent settlement they dis-
appeared.

The history of the fauna can be illustrated
with the help of Figure 5. If we suppose that
the shallow area of the sea, shown by the white
area, was the outline of the submerged land

Lord Howe Island, Part 3-— Paramonov

367

( Howeania ) , then the peculiarities of the fauna
and flora of the island can be explained easily.

The absence of typical Australian elements
was caused by the very wide and deep part of
the sea in between, which acted as an impas-
sable barrier.

The presence of Australian elements which
are not typical for the Australian fauna can
be explained by the very narrow strait in the
north (see dotted line of Tropic). It permit-
ted the penetration of tropical elements, whilst
the elements more adapted to moderate cli-
mate could reach the island from the south,
where there was a broad land connection with
New Zealand.

The very essential difference between the
faunas of Lord Howe and Norfolk Island may
be explained by the very large and deep sea
area extending eastwards from Howeania. Since
it is about halfway between New Caledonia and
New Zealand and was connected to them by
land, Norfolk Island contains elements of both
countries.

The New Zealand elements are not repre-
sented so well on Lord Howe Island because
the ecological conditions are very limited spa-
tially, and they are confined mostly to the moun-
tain summits of the island.

We have reached all these conclusions after
attentive study of the animals and plants, de-
tails of which can not be demonstrated in a
work of this size. The author reached these
conclusions before consultation of the above
map showed the distribution of the deep and
the shallow portions of the sea. The illustrated
structure of the sea bottom fully supports the
author’s considerations, which are not purely
theoretical speculations based on the study of
biology, but reflect also the history of the land
masses in the discussed area. Here we have a
working theory for reconstruction of the his-
tory of the island and its life. The theory also
explains satisfactorily the history of the basic
part of the island’s fauna.

With regard to the immigrants which
reached the island after the submergence of
Howeania, the position is less satisfactory: first,
there is a lack of data about the fauna of the
island; second, data are lacking about the dis-
tribution of organisms in the surrounding areas.

Without these data, reconstruction of the his-
tory of the fauna and flora is impossible.

Regarding comparatively recent immigrants,
we must consider the role of man in the dis-
tribution of plants and animals. Early whalers
must have played a considerable part. If, for
example, we find on the island a plant which
has a strange, widely interrupted distribution,
such as South Africa and the island, and grows
near the sea shores, we have grounds to sus-
pect that the plant has been transported by
ships. As an example, we may cite the case
of the so-called "African water lily” in Aus-
tralia ( Aponogeton distachyum, or Cape Pond
Lily). The author found the lily in 1947 in the
lagoon at Lome, Victoria. This plant is very
abundant there, and in July the entire lagoon
is covered with the white flowers. The lily is
rapidly diminishing in numbers, however, and
we must predict that sooner or later it will
disappear from the lagoon.

During the last century whalers introduced
many animals and plants along the sea shores
which they visited. One of the island’s chief
industries began in this fashion, i. e., the cul-
ture of the high quality onion, some bulbs of
which had been washed ashore, found by an
island woman, and planted.

COMPOSITION OF THE FLORA

Oliver (1917) gave an analysis of the com-
position of the island’s flora. Although the data
are outdated in nomenclature, the conclusions
may be regarded as valid.

Of the 169 genera of vascular plants repre-
sented on Lord Howe Island 4 are endemic.
Of these 4 genera, Colmeiroa and Hedyscepe
are allied to New Zealand forms, Negria to
both New Zealand and New Caledonian genera,
and Howea to Malayan and tropical Australian
genera.

If the five species belonging to these 4 genera
be taken as modified descendants of species
which arrived via a land bridge, then they
would indicate a New Caledonia-New Zealand
migration, with the land connection severed
first at the southern end, thus accounting for
a greater degree of peculiarity for the species
related to New Zealand forms.

368

PACIFIC SCIENCE, Vol. XVII, July 1963

Of the nonendemic genera 93 are widely
distributed, occurring in New Zealand, Aus-
tralia, and Polynesia; 47 others range widely
through tropical countries, but do not reach
New Zealand; 1 occurs in New Zealand only;
5 in Australia only; 3 in Polynesia only; 11 in
New Zealand and Australia only; 2 in New
Zealand and Polynesia only; 1 in Africa only.

It is interesting to state, and Oliver omitted
to do so, that most of the genera are repre-
sented only by a few species: 169 genera are
represented by 209 species, which means that
speciation is not strong on the island. Progres-
sive evolution was almost absent on the island,
and we have only transformation, probably
due to genetic impoverishment in the popu-
lations.

It is possible also that some endemics of the

island are only relics of species more widely
distributed in the past, and now preserved only
on the island.

If we compare the number of the very widely
distributed genera (see above) and the genera
with very restricted areas, we see a great dif-
ference: of 165 nonendemic genera 95 are
distributed in New Zealand, Australia, and
Polynesia; 47 only in Australia and Polynesia;
and only 23 belong to the different countries,
but are isolated.

The presence of a large portion of widely
distributed genera and species might have been
expected in the flora of an isolated island, be-
cause species possessing facilities for wide dis-
persal would naturally form the bulk of immi-
grants after the land connection had been
severed.

Fig. 6. View of the lagoon from the south; in the background are Rabbit Island and Mt. Eliza; on the
seashore are three Araucarias introduced from Norfolk Island, growing well among the native plants. Taken at
low tide. (Photo by Miss Z. Liepa.)

Lord Howe Island, Part 3 — Paramonov

369

Fig. 7. The lagoon viewed from the south during low tide. In the foreground are coral reefs, of interest
to the marine zoologist; behind these is low Rabbit Island, and in the background is the northern part of the
island with tooth-shaped Mt. Eliza. (Photo by Miss Z. Liepa. )

This means that we must be very careful
in making conclusions based only on numerical
data. For the history of the flora and fauna the
easily distributed elements are practically value-
less.

Let us turn to the analysis of the species. Of
209 species of the flora 70 (or 33%) are en-
demic. The percentage of specific endemism is

TABLE 1

TOTAL
NO. OF
SPECIES

ENDEMIC
SPECIES
INCLUDED
IN TOTAL

PERCENTAGE
OF ENDEMIC

FORMS

Whole flora

209

Australia

168

New Zealand

109

Polynesia

116

very high. Oliver gives the following table
which is very instructive:

From Table 1 (from Oliver, 1917) it is evi-
dent that endemic forms of Lord Howe Island
are distinctly more closely related to New Zea-
land and Polynesia than to Australia. As a re-
sult, the line of Gressitt (1956) must be trans-
ferred westwards of Lord Howe Island, separat-
ing it from the Australian continent. The zoo-
logical data also have shown that the fauna of
the island is more closely related to the lands
to the east than to Australia.

A direct connection of Lord Howe Island
with Norfolk Island, however, is doubtful, be-
cause the Araucarias of Norfolk Island have not
reached Lord Howe Island naturally, although
those transplanted by man are growing very
well (see Fig. 6).

370

PACIFIC SCIENCE, Vol. XVII, July 1963

NORTH ROCK

ADMIRALTY ISLETS

LORD HOWE ISLAND

.SUGARLOAF PASSAGE

SUGARLOAf *

-Hi MALABAR

WIRELESS STN
WELL
CEMETERY
WIND SOCK
WILD GOATS
WILD PIGS
BANYAN TREE
JETTY

MUTTON BIRD ISLAND

SAIL ROCK •/*£> i6S

LION POINT

ROCKY POINT

WOLF ROCK

\\ BOAT EDMANOCH POINT O
lHAVEN / *

> CUT GRASS POINT
SUGARLOAF POINT

Mt GOWER iSHfe)
t» 2840' 3;^

LARGEST t
BANYAN TREE

GEORGE ROCK

Q (awash)

TRACK

GOWER ISLAND O

Lord Howe Island, Part 3— PARAMONOV

371

CONCLUSIONS

1. Lord Howe Island is not an "oceanic”
island owing its existence to the activity of
corals, although some part was played by them;
the island’s fauna and flora are not a casual
"mixtum compositum” of elements arrived by
various methods from different directions, dur-
ing different epochs.

2. Lord Howe Island is a small part of a
sunken continent or large island, preserving
a very specific fauna and flora, with very high
endemism (in some groups as high as 70%).

3. Some endemic species developed here
probably as a result of very long isolation from
the closely related population in the surround-
ing areas; other endemic species (some birds)
probably are unchanged species preserved from
the sunken land, being mostly destroyed by
man in the surrounding areas.

4. The sunken land ("Howeania”) was
never connected with the Australian continent
after creation of Eucalyptus and Acacia species,
because on the island they are absent, although

growing very well when introduced. It appears
that the island’s isolation is very old.

5. Lord Howe Island was probably never
directly connected with Norfolk Island. The
Araucarias of Norfolk Island, so typical of that
island, are absent from Lord Howe Island. The
species common to both islands are probably
species with very strong capacity for dissemi-
nation.

6. Having no land connection with Australia
and Norfolk Island, Lord Howe Island received
its fauna and flora mostly from the north and
south. Warmth-loving elements of the flora
and fauna probably arrived from the northeast
( palms, for example ) . All the tropical elements
show a connection with this direction.

7. The elements of subtropical or moderate
areas, liking cool climate and high humidity,
probably arrived from southeastern and south-
ern directions (the so-called New Zealand ele-
ments have their origin in these sectors).

8. Since it is a highly elevated island (up to
2,800 ft) Lord Howe Island has preserved its
character, and its flora and fauna, for a long
period of geological time. The flora and fauna
of its summits (for example the plant Dixonia

Fig. 8.

1. Mt. Eliza

2. Caves in this area

3. North Hills

4. Dawson Peak

5. Poole’s Lookout

6. North Ridge

7. Grassy Hill

8. North Scab

9. Old Settlement Creek

10. Thomson’s Lookout

1 1 . Settlement Creek

12. Hine’s Curio Shop

13. "Ocean View’’ Guest House

14. The Peg

15. "Somerset" Guest House

16. Thomson’s General Store

17. Electric Power Station

18. "Waverley’s” Tea Rooms
19- "Dignam’s” Tea Rooms

20. The Hall

21. "Leanda Lei’’ Hotel

22. Doctor

23. Post Office & Bank Agency

24. Hospital

25. Government House

26. School

27. "Pine Trees’’ Guest House

28. Nichol’s Clear Place

29. Blinky Beach Picnic Hut

30. Lagoon

31. Soldier’s or Big Creek

32. "Smoking Tree"

33. "Goat House”

Asterisks indicate localities known by more than one name. Those currently used on the island are shown
on the map and alternative names are listed below:

Admiralty Islets: Roach Islets
Blinkenthorpe Bay: Ross Bay
Lion Point: Mutton Bird Point
Mt. Malabar: North Peak
North Bay: Callam’s Bay
North Rock: North Islet

Old Gulch: Collin’s Cove
Rabbit Island : Blackburn or Goat Island
Rabbit Island Passage: Boat Passage
South Entrance: Erscott’s Passage
Sugarloaf: Soldier’s Cap
Transit Hill: Lookout Mountain

372

antarctica, and the Pelorididae insects) are
very old elements, preserved in a nearly un-
changed condition for many millions of years.
It is evident that after the ancient land sank,
there was no re-elevation of it which deserves
serious attention (there were only slight oscil-
lations of ocean).

9. At the present stage of our knowledge
of the geology, paleontology, and biology of the
island, it is impossible to draw any conclusions
concerning the theory of drifting continents:
there are no positive and no negative data.

10. Finding of interesting data can be ex-
pected after a study of Ball’s Pyramid. This
colossal rock has never been touched by ex-
ploration, and no specimens either of plants or
of animals have been collected there. The pos-
sibility is not excluded that some forms which
have been destroyed on the island may have
survived on this rock.

11. There are numerous caves on the island,
some very long, which have not been explored;
we have reason to think that interesting re-
mains of recent and past fauna may be pre-
served there.

12. The presence of a coral reef gives an
opportunity to gather additional data about
oscillations of the sea level, and their magni-
tude (see Fig. 7).

13. The preliminary data at our disposal
give evidence that in the past there existed a
large land mass, probably southeastward from
Lord Howe Island, where the flora and fauna
of the island developed. On the island itself
we can find only remnants of this life of the
past.

14. Lord Howe Island is a particularly in-
teresting subject for the study of the Pacific
and surrounding countries. The scientific value
of the island has been underestimated, almost
neglected. Only one expedition, organised by the
Australian Museum in 1887, has worked on
the island. Since then any study that has been
done has been private, and by amateurs.

This article shows that the island deserves
more serious attention: its interest is not local
but international. Only longer expeditions of
geologists and biologists, working together, can
supply us with the necessary data.

PACIFIC SCIENCE, Vol. XVII, July 1963

REFERENCES

Ashton, J. H. 1921. A revision of the Aus-
tralian Cicadidae, I. Proc. Roy. Soc. Vic.
33:87-107.

Bradley, J. D. 1956. Microlepidoptera from
Lord Howe Island. Bull. Brit. Mus. Ent.
4 (4) : 145-164.

Carter, H. F. 1920. Notes on some Australian
Tenebrionidae, etc. Proc. Linn. Soc. N.S.W. 45
(2): 222-249, figs.

1920. Revisional notes on the Family

Cistelidae. Trans. Roy. Soc. S. Aust. 44:198-
217.

1921. Description of . . . New Mosquito

from Lord Howe Island. Proc. Zool. Soc.
Lond. 1920 (4) : 62 3-628.

1926. Revision of Athemistus and Mi-
crotragus (Fam. Cerambycidae) with notes,
and description of other Australian Coleop-
tera. Proc. Linn. Soc. N. S. W. 51 (4) : 439,
506.

Chopard, L. 1951. A revision of the Austra-
lian Grylloidea. Rec. S. Aust. Mus. 9:397-
533.

Cunningham, G. H. 1926. The Gasteromy-
cetes of Australasia, IV. Species of the genus
Geaster. Proc. Linn. Soc. N. S. W. 51 (2) : 87.

Dodd, A. P. 1924. Chalcidoidea and Procto-
trupoidea from Lord Howe, etc. Trans. Roy.

S. Aust. 48:162-186.

(

Evans, J. W. 1941. Concerning the Pelorididae.
Aust. Jour. Sci. 4(3) : 95-97.

1956. Insect distribution and conti-
nental drift. A symposium . . . Geolog. Dept.
Univ. Tasmania, pp. 134-161.

1959. The Pelorididae of Lord Howe

Island. Rec. Aust. Mus., Sydney 25(3): 57-

61, figs.

Gressitt, J. L. 1956. Some distribution patterns
of Pacific Island Faunae. System. Zool. 5(1):
11-33, 47.

Lord Howe Island, Part 3 — Paramonov

373

Gurney, A. B. 1947. Notes on some remark-
able Australian Walkingsticks, etc. Ann. Ent.
Soc. Amer. 40(3) : 373-396.

Hale, H. M. 1926. Studies in Australian Aqua-
tic Hemiptera, No. 7. Rec. S. Aust. Mus.
3 (2): 195-217, figs.

Hincks, W. D. 1947. Dermaptera from New
Hebrides and Lord Howe Island. Ent. Mo.
Magazine, London 83:65-67.

Johnston, H. T., and Hardy, G. H. 1923.
Some Sarcophagid flies from Lord Howe
Island. Rec. Aust. Mus. 14(1): 62—7 1, figs.

Laing, F. 1925. Descriptions of some new
genera and species of Coccidae. Bull. Ent.
Res. 16(1): 51-66, figs.

Lea, A. M. 1909. Australian and Tasmanian
Malacodermidae. Trans. Ent. Soc. Lond., 1909
( 1 ) : 45—2 5 1, pis.

— 1909. Revision of the Australian Cur-

culionidae belonging to the subfamily Cryp-
torhynchidae ( Coleoptera ) . Proc. Linn. Soc.
N.S.W. 33 (4), 1908:723.

1916. Notes on Lord Howe Island

Phasma and on associated Longicorn Beetle.
Trans. Roy. Soc. S. Aust. 40:145-147, pis.

1917. Notes on some miscellaneous

Coleoptera, with descriptions of new species,
Part 3. Trans. Roy. Soc. S. Aust. 41:121-322
(with list of distribution, pp. 212-214).

19 18. On Australian Coleoptera, Part

1. Rec. S. Aust. Mus. 1(1): 96.

Miscellanea. 1917. Notes on Diatomaceous
Earth from Lord Howe Island. Trans. Roy.

Soc. S. Aust. 41: 659.

Ollxff, A. S. 1889. The insect fauna of Lord
Howe Island. In: Lord Howe Island, its Zool-
ogy, Geology and Physical Characters. Mem.
Aust. Mus. 2:77-98, pi. 6. (The first, most
important record about the island.)

Paramonov, S. J. 1957. On some new Ortalids
( Acalyptrata ) . Ann. Mag. Nat. Hist. (12)
10:779-781.

1958. A review of Australian species

of Laphria (Asilidae), with descriptions of
three new species from Lord Howe Island.
Pacif. Sci. 12 (1):92— 105, figs.

I960. A review of Australian Pygo-

phora-species (Muscidae). Ann. Mag. Nat.
Hist. (13)3:505-512.

Ramsay, E. P. 1882-3. Description of a new
species of Coris from Lord Howe Island.
Proc. Linn. Soc. N.S.W. 7(2): 301.

Tonnoir, A. L. 1923. Notes on Australian
Rhypliidae. Ann. Mag. Nat. Hist. (9) 12:
502-505.

Turner, A. J. 1917. On some moths from
Lord Howe and Norfolk Islands in the South
Australian Museum. Trans. Roy. Soc. S. Aust.
41:53-120.

1918. Further notes on some moths

from Lord Howe Island, etc. Trans. Roy. Soc.
S. Aust. 42:276-289.

1922. Some Australian moths from

Lord Howe Island. Proc. Linn. Soc. N.S.W.
47(4): 439-440.

Tilly ard, R. J. 1917. Odonata, Plannipennia,
and Trichoptera from Lord Howe and Nor-
folk Islands. Proc. Linn. Soc. N.S.W. 42(3) :
529-544, figs.

Waterhouse, G. A. 1897. The Rhopalocera
of Lord Howe Island. Proc. Linn. Soc. N.S.W.
22(2) :285-287.

1920. Description of new forms of

butterflies from the South Pacific. Proc. Linn.
Soc. N.S.W. 45(3): 468, 470.

Wheeler, W. M. 1919. The ant genus Lodo-
myma. Psyche, Boston 24:97-106.

1919. The ants of the genus Metapone.

Ann. Ent. Soc. Amer. 12(3) : 173—191, figs.

1927. Ants of the genus Amhlyopone.

Proc. Amer. Acad. Arts and Sci., Boston 62:
1-4, 15-17, figs.

NOTES

Additional Plants from the Midway Islands

A collection of plants from the Midway Is-
lands, made by Dr. Hubert W. Frings, Depart-
ment of Zoology, University of Hawaii, on
April 16, 1962, includes four species not previ-
ously recorded from there. These are:

GRAMINEAE

Polypogon monspeliensis (L.) Desf.,
Eastern Island.

CRUCIFERAE

Coronopus didymus (L.) J. E. Smith,
Eastern Island.

EUPHORBIACEAE

Euphorbia peplus L., Sand Island.

PRIMULACEAE

Anagallis arvensis L., Sand and Eastern
Islands.

All four of these species, although not native
to Hawaii, are now found on the major islands
of the Hawaiian chain and were probably in-
troduced accidentally to Midway from Oahu.
At various times ornamental plants, and the soil
in which to grow them, have been taken to
Midway from Honolulu. It is likely that seeds
of the plants reported here reached Midway in
such soil.

Dr. Frings’ collection included 27 species of
vascular plants, which he described as the more
common species on the atoll. It is of interest
to note that only 5 of these species ( Boerhavia
diffusa L.; Tribulus cistoides L.; Ipomoea indica
(Burm. f.) Merr.; I. pes-caprae (L.) Sw.; Scaevola
sericea Vahl) are native to Midway. This pro-
vides some indication of the changes in the
flora, and concomitant changes in the ecosystem,
which are related to man’s activities on Midway.

This collection has been deposited in the
herbarium of the B. P. Bishop Museum in Ho-
nolulu— Charles H. Larnoureux, Department of
Botany, University of Hawaii, Honolulu, Hawaii.

On Malayan Shores: A Review

S. H. Chuang. 1961. On Malayan Shores. Pp.
xvi + 225; 112 numbered plates and frontispiece
(13 plates and frontispiece in color), 28 text
figures. 5.5 X 7.5 inches (14 X 19 cm). Pub-
lished by Muwu Shosa, P. O. Box 1813, Singa-
pore.

It is for this incredibly rich center of the
great Indo-West Pacific marine flora and fauna
that Dr. Chuang’s small volume is intended as
an introduction. He did not write the book for
the specialist in any one group, or even for the
trained marine biologist, but for the "amateur
naturalist.” With this aim he has avoided as
much of the specialized language of zoology as

possible; his descriptions of plants and animals
are short and point out only one to several
salient characteristics; he uses no keys and gives
no synonyms. On the other hand his illustrations
are numerous and excellent: some 50 species
are illustrated by line drawings in the text fig-
ures and over 500 are represented by clear
reproductions of photographs in the 112 plates.
He has limited himself to the more common
and conspicuous species, as would be expected;
he does not touch upon the fishes which have
been dealt with in previous books.

The book is divided into two parts, the first
is a general review of aspects of marine biology

374

On Malayan Shores: A Review

375

applicable within the scope of the work, the
second is a systematic review. The first section
includes chapters on the chemistry of sea water,
zonation of the shore, coral reefs, plankton, etc.
There is also a chapter valuable for a trained
biologist visiting the peninsula which discusses
Malayan collecting localities ( Chapter VI ) . The
second portion is divided among seven phyla,
although eight or nine additional phyla are dis-
cussed, with species named, in the chapter on
animal life of Part I. The appendix gives faunal
lists for four typical shore habitats. A glossary
gives Malayan names for over 100 species. The
bibliography is short and covers only those
books cited in the text, not the systematic works
from which the nomenclature was derived. The
index is very complete, listing all systematic
names applied to all taxa, English common
names, topics discussed in the first part of the
book, and authors cited; it even includes a
glossary to some of the more technical terms
used, as "siphonoglyph” and "periostracum.”

While this book is addressed only to the

amateur, it obviously will be used by two other
groups: students of biology in Malaya, and ma-
rine biologists visting these waters for the first
time. Having spent a year working on the ad-
jacent Gulf of Thailand, and having briefly vis-
ited Malayan shores, I can attest how extremely
valuable this volume will be to visitors. More-
over, because the fauna from India to Indonesia,
from the Coral Sea to the South China Sea is
basically the same, it will be of use to all visitors
to the Western Pacific and adjacent Indian
Ocean.

Admittedly the book has limitations: it is
not an exhaustive scientific monograph on the
flora and fauna. Yet if it were, it would not be
one small volume, but many large volumes.
Rather than disparage the book for omissions,
we should commend the author for producing a
volume of considerable value to biologists, and
regret that books such as this are not available
for other areas in the tropical Pacific. — A. H.
Banner, Department of Zoology, University of
Hawaii, Honolulu, Hawaii.

Introduction and Summary. It is desirable to state the
purpose and scope of the paper in an introductory
paragraph and to give a summary of results at the end
of the paper.

Batzo, Roderick L., and J. K. Ripkin. 1849. A
Treatise on Pacific Gastropods. Rice and Shipley,
Boston, vii + 326 pp., 8 figs., 1 map.

Crawford, David L. 1920^. New or interesting
Psyllidae of the Pacific Coast (Homop.). Proc.
Hawaii. Ent. Soc. 4(1) : 12-14.

1920&. The sandalwoods of Hawaii. Proc.

Hawaii. Ent. Soc. 4(2): 374-375, 13 pis.

In the text, sources should be referred to by author,
date, and page, as follows: "It was noted (Rock,
1916: 18) that . . .” or "Rock (1916: 21-24)
says . . .”

ILLUSTRATIVE MATTER

Artwork for illustrations should be 8Y2 x 11 inches
or smaller, and it should accompany manuscript, on
separate sheets. Often more valuable than a photo-
graph is a good line drawing.

It is strongly urged that an indication of scale be
incorporated as a part of all drawings in which mag-
nification and size are critical considerations.

Illustrations will be returned to the author.

Tables. Tabular matter should be kept to a minimum.
Each table, prepared to conform with Pacific Science
style, should be typed on a separate page, and its posi-
tion indicated on the manuscript.

Mathematical Formulas. Complicated formulas cannot
be set by the printers. Authors should submit them
as illustrations.

Captions. Readily identifiable captions for figures,
graphs, photographs, and other illustrative matter
should be supplied on a separate page.

PROOF

All queries on proof should be answered. If cor-
rected proof is not received within four weeks after
being sent to the author, author’s changes cannot be
accepted.

REPRINTS

NO. 4

VOL. XVII

OCTOBER 1963

PACIFIC SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

JEN-HU CHANG

Climatology in Hawaiian Sugar-Cane Industry

D. L PLUCKNETT, J. C. MOOMAW, and C. H. LAMOUREUX
Root Development in Aluminous Hawaiian Soils

BENJAMIN C. STONE

New and Critical Species of Pelea

ROBERT L. WILBUR

A Prior Name for Gouldia terminalis

WILLIS E. PEQUEGNAT

Population Dynamics in a Sublittoral Epifauna

THOMAS S. HIDA and ROBERT A. MORRIS
The Marquesan Sardine in Hawaii

DONALD C. MATTHEWS

Folliculinids from Submerged Wood

CHARLES P. HOYT

Rhinoceros Beetles in West Africa

V. S. SOBOLEV

Features of Vo Icanism of the Siberian Platform

MAXWELL S. DOTY

Gibsmithia hawaiiensis gen. n. et sp. n.

HAROLD ST. JOHN

Revision of the Genus Pandanus
Part 16. Thailand and Vietnam

INDEX

UNIVERSITY OF HAWAII PRESS

BOARD OF EDITORS

O. A. BUSHNELL, Editor-in-Chief
Department of Microbiology, University of Hawaii

Robert Sparks, Assistant to the Editors
Office of Publications and Information, University of Hawaii

L. H. Briggs
Department of Chemistry
University of Auckland
Auckland, New Zealand

Maxwell S. Doty
Department of Botany
University of Hawaii

Ai Kim Kiang
Department of Chemistry
University of Malaya, Singapore

Gordon A. Macdonald
Department of Geology
University of Hawaii

John J. Naughton
Department of Chemistry
University of Hawaii

Martin Sherman
Department of Entomology
University of Hawaii

Walter R. Steiger
Department of Physics
University of Hawaii

Donald W. Strasburg
Bureau of Commercial Fisheries, Hawaii Area
(U. S. Fish and Wildlife Service)
Honolulu, Hawaii

Donald G Matthews
Department of Zoology
University of Hawaii

Sidney J. Townsley
Department of Zoology
University of Hawaii

Thomas Nickerson, Managing Editor
Assistant to the University Provost

INFORMATION FOR AUTHORS

PREPARATION OF MANUSCRIPT

( Continued on inside back cover)

PACIFIC SCIENCE

A QUARTERLY DEVOTED TO THE BIOLOGICAL
AND PHYSICAL SCIENCES OF THE PACIFIC REGION

VOL. XVII OCTOBER 1963 NO. 4

Previous issue published September 17, 1963

CONTENTS

PAGE

The Role of Climatology in the Hawaiian Sugar-Cane Industry: An Example of

Applied Agricultural Climatology in the Tropics. Jen-hu Chang 379

Root Development in Aluminous Hatvaiian Soils.

D. L. Plucknett, J. C. Moomaw, and C. H. Lamoureux 398

Studies in the Hawaiian Rutaceae , IV. New and Critical Species of Pelea A. Gray.
Benjamin C. Stone

A Prior Name for the Hawaiian Gouldia terminalis (Ruhiaceae).

Robert L. Wilbur

Population Dynamics in a Sublittoral Epifauna. Willis E. Pequegnat

Preliminary Report on the Marquesan Sardine, Harengula vittata, in Hawaii.
Thomas S. Hida and Robert A. Morris

Hawaiian Records of F olliculinids ( Protozoa ) from Submerged Wood.

Donald C. Matthews

Investigations of Rhinoceros Beetles in West Africa. Charles P. Hoyt

Characteristic Features of the Volcanism of the Siberian Platform.

V. S. Sobolev

Gibsmithia hawaiiensis gen. n. et sp. n. Maxwell S. Doty

Revision of the Genus Pandanus Stickman, Part 1 6. Species Discovered in
Thailand and Vietnam. Harold St. John

Index

407

421

424

431

438

444

452

458

466

493

fliftHSORUff

issTirunon

FEB 2 6 1964

The Role of Climatology in the Hawaiian Sugar-Cane Industry:
An Example of Applied Agricultural Climatology in the Tropics1

Jen-hu Chang

Climatological study in the Hawaiian sugar-
cane industry has a long, noteworthy history.
Meteorological observations on the plantations
were initiated in 1883, preceding the establish-
ment of the first official weather bureau station
in Hawaii by fully 20 years. The climatological
network in the cane-growing areas expanded by
leaps and bounds to 50 stations at the turn of
the century, and to 500 stations in I960 in an
area of only 350 sq miles (Fig. 1).

In the late 1920’s and early 1930’s Das ( 1928,
1931 a, 1931 b9 1932) used the rainfall and tem-
perature records to define the effect of climate
on crop yield and juice quality in a simple, direct
manner. He also advocated the use of day-degree
as a guide to irrigation control (Das, 1936).
Investigations along the same line of climate-
and-plant complex were carried further by Wil-
liams (1933), Borden ( 1940, 1949), Clements
(1940), Swezey (1942), and others into the
early 1940 s.

At the end of World War II meteorologists,
freshly relieved from their wartime duties, were
able to turn their attention to a multitude of
peacetime problems. The Hawaiian sugar in-
dustry promptly seized this opportunity. The
Experiment Station and the Pineapple Research
Institute founded their joint Meteorological De-
partment in 1946 and contracted with a group
of meteorologists at the University of Chicago
to investigate dynamic and regional climatology
of Hawaii. The culminating results of this in-
tensified research were a series of papers in the
Meteorological Monographs, discussing the gen-
eral circulation, weather types, local flow pat-
terns, rainfall statistics, and the like. Such studies
inevitably led to the improvement of forecasting
and furnished valuable information for opera-
tional planning.

At the half-century mark, when weather

1 Contribution No. 121, Experiment Station, Ha-
waiian Sugar Planters’ Association, Honolulu, Hawaii.
Manuscript received June 14, 1962.

modification was a new subject, the industry
embarked upon an intensive study of cloud seed-
ing. Leading scientists from seven research in-
stitutes in the U.S.A. and abroad participated
in the so-called "Project Shower.” It was hoped
that the Hilo coast, with its humid trade winds,
would provide an ideal environment where rain-
fall could be induced by the addition of chemi-
cals to the warm clouds; it was soon realized,
however, that scientific know-how of artificial
rainfall was probably still decades away. The
project nevertheless produced valuable informa-
tion in the field of cloud physics.

Realizing the limitations upon man’s ability
to modify weather on a grand scale, the industry
sought instead to improve the efficiency of water
use through the study of micrometeorology. In
1957 an evapotranspiration project was initiated
at the Experiment Station. Apart from its ob-
vious application in irrigation planning, the de-
termination of potential evapotranspiration pro-
vides a vital link for solving the energy budget
and water balance equation in the soil-plant
system. The solution of these equations renders
the climate-and-yield relationship amenable to
quantitative treatment.

Climatological study in the Hawaiian sugar
industry thus encompasses a wide variety of
topics. More than 50 papers have appeared in a
dozen scientific journals. The industry ranks
among the leaders in the study of agricultural
meteorology. The experience gained during the
past three decades may very well benefit research
workers in other parts of the world, especially
those in the tropics. This paper attempts to
summarize these studies. It is hoped that such a
stock-taking will not only serve as a reminder
of our past accomplishments but also as a guide-
post for planning future research.

TRADE-WIND WEATHER

The subtropical high pressure cell in the Pa-
cific Ocean with its attendant trade wind is the

379

380

PACIFIC SCIENCE, Vol. XVII, October 1963

Fig. 1. Location map.

basic circulation in Hawaii; all other weather
types are perturbations in this basic current. The
eastern portion of a subtropical anticyclone is
characterized by divergence and subsidence and
consequent adiabatic heating and pronounced
dryness aloft. There is, however, a mixing layer
near the surface where the air in contact with
the cold ocean currents becomes cold and humid.
These two layers of air are separated by a zone
of temperature inversion which, along the Cali-
fornia coast, occurs at an altitude of about 1,500
ft. Downstream, toward the west, as water tem-
perature rises and subsidence weakens, the trade-
wind inversion reaches a height of about 6,000
ft in Hawaii.

Figure 2 shows the vertical structure of trade
wind in Hawaii. The air below the inversion is
moist, with a mixing ratio of 12-15 gm/kg at
the ground surface. In the absence of orographic
uplift this moist air is a poor rain-producer be-

cause the trade inversion acts as a lid to oppose
the development of convective clouds; therefore,
in lowland areas, the trade wind is atmospheri-
cally moist in terms of humidity but ecologically
dry in terms of rainfall.

The air below the trade inversion has a lapse
rate close to dry adiabatic. It requires only a
little uplifting to produce rainfall. Baer ( 1956)
has demonstrated theoretically that if the rela-
tive humidity is approximately constant below
the inversion layer, rainfall would increase with
height exponentially up to the base of inversion.
The logarithmic distribution of rainfall with
height is a unique climatic feature in Hawaii.

The prevalence of trade-wind weather in Ha-
waii is dependent upon the location and strength
of the Pacific subtropical anticyclone. In July
and August, when the subtropical ridgeline in
the high troposphere is located to the north of
the islands, the trades prevail during 97% of

Climatology and Sugar Cane — CHANG

381

the time (Yeh et al., 1951^) . In winter, when
the subtropical anticyclone moves southward,
the frequency of trade-wind weather decreases
to 40-50% (Fig. 3).

OTHER WEATHER TYPES

During one-third of the year, when the trades
are not prevalent, weather in Hawaii falls into
two general types: mixed and cyclonic. The
mixed type is characterized by the presence of
an essentially east-west polar front north of the
islands. This type is a very poor rain-producer
as the front is located at a distance. The mixed
type often ushers in cyclonic weather.

Cyclonic weather occurs in Hawaii during
22% of the year. The cyclonic weather may be
further divided into four subtypes: (1) extra-
tropical cyclones, ( 2 ) kona storms, ( 3 ) easterly
waves, and (4) tropical cyclones or hurricanes.
The first two subtypes occur most frequently in
winter while the latter two subtypes are pri-
marily summer disturbances.

Uncomplicated extratropical cyclones occur
only when the zonal westerlies move southward
into the tropics. They are, however, weakened
on their journey southward and produce only

TYPICAL LAPSE RATE CURVE SHOWING THE
STRUCTURE OF TRADE WIND IN HAWAII

I4°F 23 32 41 50 59 68“F

FIG. 2. Structure of the trades in Hawaii.

Fig. 3. Monthly variation of frequencies of three
weather types.

small rains. Some of these cyclones reach only
the northern island of Kauai.

The kona storm is essentially a cold-core low
developed in the tropics. Simpson (1952) has
described two main processes in its formation:
( 1 ) the transformation of mid-latitude cyclones
which have been trapped at low latitudes by
blocking action of a warm high; (2) the cy-
clogenesis in the easterlies triggered by the
building down of a pre-existing cold upper low
usually at the southern extremity of a polar
trough. Once formed, all kona storms seem to
assume the same characteristics. The term kona
indicates that the wind during such a synoptic-
period is southerly. Kona storms occur about
three or four times a year, bringing heavy rain-
fall throughout the islands. In the dry lowland
areas they may account for more than half of
the annual rainfall.

The occurrence of winter cyclones and kona
storms in Hawaii is closely related to the general
circulation. Yeh et al. (1951 b) have shown that
winter rainfall in Hawaii increases as the lati-
tude of the jet stream increases, unless the latter
penetrates to the latitude of the islands them-
selves. They explained that when the jet stream
is weak and far to the north, Hawaii will ex-
perience a period of low zonal index. Low index

382

flow pattern is characterized by large meridional
flow and frequent interaction between tropical
perturbations and troughs in the westerlies.

On a hemispherical scale the mid-tropospheric
westerlies are composed of roughly sinusoidal
waves. In winter an upper air trough is climato-
logically anchored off the east coast of Asia.
The location of the next downstream trough
depends upon the zonal velocity according to
the well-known Rossby formula. The zonal ve-
locity in the western Pacific is such that the
trough is usually located to the west of the is-
lands in January and to the east in February
(Namias and Mordy, 1952). Since the trough
is characterized by convergence ahead and di-
vergence behind, cyclones in Hawaii are more
numerous in January than in February. The low
rainfall in February is especially evident in the

PACIFIC SCIENCE, Vol. XVII, October 1963

northern island of Kauai (Leopold and Stidd,
1949).

The easterly waves are trade-wind perturba-
tions that may form during any part of the year
but with a maximum during summer. In August
the easterly waves occur along the east coast of
Hawaii every 3 or 4 days, bringing with them
decks of high clouds and rainfall; however, they
dissipate readily over the rugged land and sel-
dom reach Oahu and Kauai.

Since 1904 there have been only four tropical
storms with wind of 74 rnph or more passing
through the islands, all in the 1950’s. The for-
mation of a hurricane in the central Pacific is
somewhat of a climatic anomaly and requires a
combination of favorable conditions. In 1957,
when two hurricanes passed through Hawaii,
Frazier (1957) noted that a well-developed

Fig. 4. Median annual rainfall (inches) in Hawaii.

Climatology and Sugar Cane — CHANG

383

trough extending to low latitudes persisted along
the west coast of North America throughout the
summer and fall season. This trough, through
an energy dispersion mechanism, sharpens the
next downstream subtropical trough, thus pro-
moting cyclogenesis in the latter area ( Ramage,
1959). The increase in radiation and tempera-
ture in Hawaii during the last decade, which
will be discussed in a later section, was probably
also an important cause for the increased hurri-
cane activity.

RAINFALL

If the islands of Hawaii did not exist, the
annual rainfall over the ocean would be in the
neighborhood of 30 inches; the actual average
rainfall over the islands is about 70 inches. This
increase of 40 inches of rainfall is the result of
orographic uplifting of trade wind and its per-
turbation-easterly waves. The effect of easterly
waves is strongest near Hilo and diminishes
northward toward the Hamakua coast. The me-
dian annual rainfall at Hilo is 139 inches, con-
siderably higher than any other coastal area in
the state (Fig. 4).

The trade-wind rainfall increases with alti-
tude exponentially up to a certain point and
then decreases again. The belt of maximum
rainfall varies from 3,000 to 4,000 ft, depending
upon the effect of local topography on the flow
patterns. This effect has been discussed in detail
by Leopold (1949). The steep isohyetal gradient
in a trade-wind climate is best illustrated in
Kauai, where Mt. Waialeale, with a mean annual
rainfall of 465 inches, is only 15 miles away
from the semiarid west coast. Most of the sugar
plantations are located in lowlands with a me-
dian rainfall of less than 50 inches. A few cane-
growing areas on Hawaii, however, receive as
much as 200 inches of rainfall in a year.

Except along the Kona coast of Hawaii, where
the upslope sea breeze produces frequent sum-
mer afternoon showers, winter is the wet season
throughout the state. The contrast between sum-
mer and winter rainfall is most accentuated in
the dry lowlands, where the monthly rainfall
in summer is often less than 0.5 inch. Summer
rainfall minima are found only in a very few
places in the tropics and are designated as "As”
climate in the Koppen classification.

In spite of the large areal and seasonal varia-
tions, the rainfall distribution for any particular
month bears a close resemblance to that of the
annual. Thus Stidd and Leopold (1951) were
able to express the monthly rainfall as a function
of annual rainfall in the following manner:

y = a (x — 30) -f b

where y is monthly rainfall; x, annual rainfall; a,
the gradient factor of observed orographic in-
crease of rainfall through increment of average
annual rainfall; b, a geographic constant quan-
tity derived from a rainfall blanket of uniform
thickness over the islands and adjacent ocean.
The constant 30 is inserted because the mini-
mum annual rainfall in the dry lowland is about
30 inches.

This same concept was later expanded to de-
scribe daily rainfall distribution during trade-
wind weather. For the island of Oahu, for ex-
ample, a dry index of zero is assigned to a
theoretical station having a zero mean annual
rainfall, and a wet index of 100 to a station
having 250 inches mean annual rainfall. A daily
forecast chart is constructed by plotting the in-
dices as the abscissa and the daily rainfall
amounts as the ordinate. The forecaster fore-
casts the daily rainfall for the dry and wet index
stations. From the straight line connecting these
two reference points, daily rainfall for any actual
station may be read off as the ordinate corre-
sponding to the abscissa of the station’s mean
annual rainfall.

The trade-wind rainfall is more likely to occur
during the night or in early morning than during
the day. Loveridge (1924) attributed the noc-
turnal rainfall to the radiative cooling at the
top of the clouds. Leopold (1948), however,
added that cooling at night would lower the
condensation level. Nocturnal rainfall, in distinct
contrast to afternoon showers in many tropical
countries, is in many ways beneficial to agricul-
ture.

Trade-wind showers are very light, with drop
size less than 2 mm in diameter (Blanchard,
1953), and with intensities usually much less
than 0.2-0.3 inch per day. Only kona storms
and hurricanes are capable of causing severe
crop damage. On January 24, 1956, a kona storm
deposited 38 inches of rain at Kilauea, Kauai.
Such severe storms are rare, however, averaging

CUMULATIVE PERCENTAGE OF RAINFALL

Fig. 5. Relationship between per cent of days with rain and per cent of total rainfall.

one in every 5 or 6 years. Furthermore, storm
damages are usually restricted to Kauai and the
east coast of Hawaii.

The frequency distribution of daily rainfall in
Hawaii is extremely skewed. Figure 5 shows the
relationship between the percentage of rainy
days and the percentage of rainfall amounts
cumulated from the least to the heaviest. At
Ewa, for example, 10% of the days with the
heaviest rain accounted for nearly 60% of the
total, while 50% of days with least rainfall
amounted to only 6% of the total rain. The
skewness of the daily rainfall frequency curve

decreases slightly with the increase of annual
rainfall.

The skewed rainfall distribution has at least
two important implications. First, a large
amount of the annual rainfall will probably be
lost as runoff. Second, the mean monthly rain-
fall will be considerably higher than the median,
and the former is a poor indicator of the
"normal” condition. For this reason median
rainfall is used extensively in Hawaii.

As rainfall variability is very high in the
tropics (Biel, 1929), even the use of the
median is inadequate for many agricultural pur-

Climatology and Sugar Cane — CHANG

385

poses. A knowledge of the extremes and fre-
quency probabilities is indispensable. Because
the basic cause of rainfall variation in Hawaii
is rather uniform, the rainfall probabilities for
different stations can be related in a simple,
empirical manner (Landsberg, 1951). Figure 6
shows the probabilities (slanting lines) of hav-
ing an annual amount of less than a given quan-
tity (ordinate) as a function of the median
annual amounts (abscissae), based on at least
60 years’ records of 20 plantation stations. It is
evident that Figure 6 could be used as a risk
chart. Monthly rainfall probabilities could also
be presented in the same manner.

TEMPERATURE

The mean annual temperatures in the low-
land plantation areas vary from 72.5 F along
the east coast of Hawaii to 75 F for the drier
stations. The temperature decreases with eleva-

MEDIAN ANNUAL RAINFALL (INCHES) g

Fig. 6. Probabilities of having individual rainfall
amounts in Hawaii as a function of median annual
rainfall.

tion at an average rate of 4 F per 1,000 ft. The
coldest plantation stations, at an elevation of
about 3,000 ft, have a mean annual temperature
of 62-63 F.

All the stations in Hawaii below an altitude
of 5,000 ft have an annual temperature range
of less than 9 F (Jones, 1942) ; thus a station at
an elevation of 2,000 ft with an annual temper-
ature of 65 F would have a minimum monthly
temperature as high as 62 F. The isothermal
climate is favorable for the growth of a peren-
nial crop like sugar cane.

In Hawaii, as in many other tropical coun-
tries, the daily temperature range exceeds the
annual mean daily temperature range and ex-
hibits greater areal differences than does the
mean annual temperature. Figure 7 shows the
distribution of the annual mean daily tempera-
ture range in the cane-growing areas. In general
the dry leeward stations have the greater tem-
perature range. The daily temperature range is
subject to a small seasonal variation, being
slightly higher in the winter. According to the
results of phytotron experiments at the Cali-
fornia Institute of Technology, daily tempera-
ture range exerts a profound influence on fruit
quality (Went, 1957).

The distribution of soil temperature in the
tropics can be deduced fairly accurately from
observations of air temperature (Chang, 1958).
The mean annual soil temperature at any depth
differs only slightly from the mean annual air
temperatures. The annual temperature range at
the soil surface under the cover of a sugar-cane
crop is reduced to half of the air temperature
range. Thus, in most of the lowlands in Hawaii
soil temperature exceeds 72 F throughout the
year. Studies at the Experiment Station have
established that temperatures of 62 F are ex-
tremely limiting for cane growth and nutrient-
and water-uptake. Such low temperatures are
observed only in the mountains above 2,000 ft.
The lowest soil temperature recorded on sugar
cane land was 61 F at Hamakua plantation at
an elevation of 3,000 ft.

RADIATION

Radiation measurements have been taken at
35 plantation stations by photochemical tubes
which use oxalic acid as agent and uranyl sul-

386

PACIFIC SCIENCE, Vol. XVII, October 1963

Fig. 7. Annual mean daily temperature range (°F) in Hawaii.

fate as catalyst. The photochemical method is
accurate enough for general agricultural pur-
poses (Chang, 1961). Apart from the direct
effect on local climate and plant growth, the
radiation data could be used for assessing yield
potential, estimating irrigation water needs,
directing fertilizer practices, etc. (Borden, 1940).

Figure 8 shows the distribution of mean daily
radiation in Hawaiian sugar plantations. Maui
has the highest radiation. Pukalani, in central
Maui, receives 607 langleys/day or over 220,000
langleys/year. This is exceeded only by a very
few desert areas in the world (Budyko, 1958).
The average radiation for the cane-growing
areas in Hawaii is in the neighborhood of 510
langleys/day, well above Houghton’s (1954)
estimate of 415 langleys/day for the land areas
in the latitudinal zone 0°-20° N.

In Hawaii the minimum radiation is recorded

in December, while the maximum is usually in
June or July. The maximum monthly radiation
is about 15% higher, and the minimum some
2 5 % lower, than the annual mean. Radiation in
December is about two-thirds that of June or

July-

SECULAR CLIMATIC CHANGES

Since the cause for climatic variation in Ha-
waii is rather uniform, and since the leeward
stations in particular are sensitive to circulation
regimes (Landsberg, 1951), the climatic change
at Makiki, in Honolulu, can be considered to 1
illustrate that of the islands in general.

The average daily radiation at Makiki in-
creased some 10% from 491 langleys/day in
the 1930’s to 544 langleys/day in the 1950 s
(Fig. 9). The rising trend probably started in
1936, but records before 1932 are not available.

Climatology and Sugar Cane — CHANG

387

Increased radiation results in a rise of tempera-
ture, though with a large time lag in a maritime
climate. The temperature at Makiki (Fig. 10)
did not start to rise until 1948.

Increased radiation could conceivably bring
about a change in general circulation. Went-
worth (1949) noted that the prevailing wind
in Honolulu shifted from northeast to east from
1907 to the late 1930’s and veered back there-
after. Increased frequency of east wind suggests
a southerly location of the subtropical anti-
cyclone, less frequent trade-wind weather, but
not necessarily an increase in cyclonic activities;
therefore, the trade-wind rainfall should de-
crease when the east wind is prevalent. At
Makiki there was a trend to decreased rainfall
from the 1920’s to 1941 and to increased rain-
fall thereafter (Fig. 11). This explanation of

the rainfall trend is tentative, however, and re-
quires further study.

Whatever the causes for climatic change may
be, its impact on agriculture is varied and pro-
found. In general, the climate in Hawaii has
become more favorable for sugar-cane culture
during the last 20 years. This must account for
part of the yield increase during that period.

MODIFICATION OF CLIMATE

The climate of Hawaii is, in general, well
suited to the growth of sugar cane. Water econ-
omy is the one area where the climatic environ-
ment could be altered significantly to bring
about a higher yield. Irrigation in the Hawaiian
sugar industry started in 1852. At the present
time 54% of the plantation area is irrigated. In

Fig. 8. Solar radiation (langleys/day) in Hawaii.

388

PACIFIC SCIENCE, Vol XVII, October 1963

Fig. 9- Solar radiation at Makiki, 1932-60.

general the irrigated areas have less than 60
inches of rainfall in a year.

Intelligent management of irrigation requires
a knowledge of the water need by a particular
crop. Das (1936) advocated the use of day-
degree as a measure of water need and as a
guide to irrigation interval control. Numerous
field experiments were conducted in Hawaii to
determine the relationship between sugar yields
and the number of day-degrees between irriga-
tion. It was found that temperature is a poor
indicator of the solar energy, which determines
to a large extent the water needs of a crop. In
fact, the monthly temperature and radiation at
Makiki for the period 1932-60 have a correla-
tion coefficient of only 0.57. The correlation is
even poorer when the maximum temperature
is used instead of the mean. In other parts of
the world also, the use of day-degree as a
phenological index has been refuted (Schneider,
1952; Wang, I960).

As interest in the day-degree approach waned,
Baver (1954) properly called attention to the
rapid development of the evapotranspiration
and other micrometeorological concepts and
their application in irrigation. He contended
that the meteorological approach has the ad-
vantage of simplicity of operation when com-
pared with methods based upon measurement
of soil moisture change. More important, how-
ever, are the many applications of evapotrans-
piration in determining the regional water
balance and in disentangling the climate-yield
relationship. With these ideas in mind the Ex-

periment Station started an intensive study of
micrometeorology in 1957.

EVAPOTRANSPIRATION AND ENERGY BUDGET

Sugar cane is a tall, ungainly plant with an
aerodynamically rough canopy. Wind profile
measurements indicate that the roughness pa-
rameter of a mature sugar cane of 4 cm height
is 9 cm, as against 2.3 cm for thick grass of 10
cm height and 0.1 cm for short lawn grass
(Sutton, 1953). The high roughness of a sugar-
cane crop renders the aerodynamic approach to
evapotranspiration a difficult task (Deacon et
al., 1958). This, together with the lack of a
suitable instrument for measuring vapor flux,
accounts for the fact that the aerodynamic
method was not assayed in our experimental
work.

The potential evapotranspiration of sugar
cane was measured by drainage lysimeters and
the data were analyzed by the IBM computer
according to the Penman (1948) and Thorn-
thwaite (1948) formulae. Detailed discussion
of the instrumentation and the results has been
reported elsewhere (Chang, 1961). It needs
only to be emphasized that in a tropical mari-

FlG. 10. Five-year moving mean of temperature
at Makiki, recorded at middle year, for the period
1918-61.

Climatology and Sugar Cane — CHANG

389

Fig. 11. Five-year moving mean of rainfall at
Makiki, recorded at middle year, for the period
1920-58.

time climate the Penman estimates are far better
than the Thornthwaite formula.

The various radiation components were meas-
ured at Makiki to evaluate the energy budget
approach to evapotranspiration. Over a mature
cane field the partition of incoming radiation is
approximately as follows: 16% reflected radia-
tion, 17% back radiation, and 67% net radia-
tion (Fig. 12). The relationship between the
net and incoming radiation is almost a constant
throughout the year and over different vegeta-
tive surfaces. It was observed at Wahiawa that
over short grass and pineapple fields, which
have an albedo of about 5%, the net radiation
remains two-thirds of incoming radiation.
Theoretically it is difficult to reconcile the fact
that the net radiation is not affected by the de-
crease of albedo, but from a practical standpoint
this is convenient.

In the tropics, especially under the cover of
tall vegetation, heat flux to and from the soil
is negligible; therefore, the net radiation is
consumed either in heating the air or in evapo-
transpiration. At Makiki 82 % of the annual net
radiation, or 55% of the incoming radiation, is
used in evapotranspiration. There is a small sea-
sonal variation of this percentage value, which
is some 10% higher in summer than in winter.

There is also a regional variation of the

energy partition. For instance, the percentage of
insolation used in evapotranspiration increases
with total radiation. In Hawaii the regional
variations of energy budget are rather small, and
the equation at Makiki could be used to esti-
mate with reasonable accuracy the monthly
potential evapotranspiration for other cane-
growing areas in Hawaii. For estimating short
term, say weekly, potential evapotranspiration
it is advisable to use an evaporimeter which
integrates not only radiation but other meteoro-
logical elements as well.

The simple energy budget presented here is
applicable only in the humid tropics. In middle
and high latitudes the energy budget is subject
to enormous seasonal variations. In an arid cli-
mate the advective heat, which is extremely dif-
ficult to evaluate, may introduce a significant
error.

PAN EVAPORATION AS A MEASURE OF
EVAPOTRANSPIRATION

Until a cheap and readily installable piece of
equipment capable of recording vertical vapor
transfer is available, agricultural meteorologists
will continue to use evaporation pans. In an arid
climate pan evaporation is accentuated by the
oasis effect to such an extent that its usefulness
as a climatic parameter is greatly impaired. In
humid climates, pan evaporation has been found
to be more accurate than the Penman and
Thornthwaite estimates (Suzuki and Fukuda,
1958), and a satisfactory guide to irrigation
control (Krogman and Lutwick, 1961). In this

INCOMING RADIATION (100%)

FIG. 12. Energy budget over a sugar-cane field at
Makiki.

390

PACIFIC SCIENCE, Vol. XVII, October 1963

respect Hawaii is ideally suited to the use of
evaporation pans, thanks to the existence of a
moist mixing layer below the trade inversion.

Lysimeter and pan-evaporation records in Ha-
waii indicate that the potential evapotranspira-
tion of mature sugar cane is approximately the
same as evaporation from a ground-level U.S.
Weather Bureau Class A pan. The ground-level
pan, however, evaporates some 10% less than an
elevated pan at cane top level, and some 15%
more than a buried pan in an irrigated plot.

The one-to-one ratio obtained by using the
ground-level pan is very much in line with
ratios found for many other crops. From the
standpoint of water use, crops may be divided
into two groups: conventional and nonconven-
tional. The potential evapotranspiration of a
nonconventional crop is influenced to a large
extent by its physiology. Rice, with a ratio of
1.2 (Suzuki and Fukuda, 1958), and pineapple,
with a ratio of 0.35, are good examples. The
water need of a conventional crop is deter-
mined primarily by the ^weather conditions, al-
though it increases with the roughness of the
crop, especially in areas of large advective heat
(Tanner and Pelton, I960). Thus the ratio in-
creases from 0.75 for short grass (Penman,
1948) to 0.87 for corn (Fritschen, 1960^), and
to 1.0 for sugar cane.

The fact that potential evapotranspiration of
sugar cane approximates pan evaporation does
not necessarily mean that irrigation based on a
ratio of one-to-one is most economical. A new
experiment has been set up at Waipio to de-
termine sugar-cane growth and yield by using
different pan ratios, i.e., 1.30, 1.15, 1.00, 0.85,
0.70, and 0.55. The yield data over a number of
years should permit determination of the most
economical level of irrigation.

Fig. 13- Monthly water balance at Opaeula.

MONTHLY AND ANNUAL WATER BALANCE

The monthly water balance of a place can
be portrayed by comparing the pan evaporation
and the median rainfall. At Opaeula, for ex-
ample, rainfall exceeds pan evaporation during
the winter months from November to February
(Fig. 13). From March to October, however,
supplemental irrigation is needed to fill the
water deficits, which total 29.6 inches in a year.

By analyzing the water balance of some 30
stations we have been able to construct an an-
nual water-deficit map for the cane-growing
areas in Hawaii (Fig. 14). The map is tentative,
as the stations are not well distributed. Recently
we have added 20 pan stations in order to
strengthen the network. In constructing the map
the radiation records were also used to fill the
gap.

By planimetering the deficit areas in the map,

TABLE 1

Estimated Annual Water Deficit for Growth of Sugar Cane

ISLAND

AREA
( acre )

AVERAGE DEFICIT
( inches )

TOTAL DEFICIT
( acre-inches )

Hawaii

98,601

16.5

1,628,390

Maui

42,424

72.8

3,087,612

Oahu

33,223

41.0

1,361,141

Kauai

47,088

33.4

1,573,502

Total

221,336

34.6

7,650,645

Climatology and Sugar Cane — CHANG

391

the annual water deficits of the four islands are
estimated as in Table 1. It is noted that there is
a large regional variation. The average annual
deficit for plantation areas on Hawaii, which
are almost exclusively unirrigated, is only 16.5
inches. By contrast, the average deficit for Maui
is as high as 72.8 inches.

The total annual water deficit for all the
plantations is estimated at 7.6 million acre-
inches, or 208 billion gallons. The industry uses
about 400 billion gallons of water a year for
irrigation. At first glance, the camp crop in Ha-
waii seems to be irrigated more than adequately.
This is, however, not true. Much of the irriga-
tion water is wasted due to maldistribution. For
instance, a plantation irrigation at a high level
of adequacy often loses as much as 60% of its
water through deep penetration or runoff. Fur-

thermore, the water supply is often such that a
plantation over-irrigates in one season and suf-
fers from drought in another.

The monthly water-balance charts and the
water-deficit maps are a valuable guide to agri-
cultural planning. Such questions as whether or
not to construct a reservoir, what irrigation sys-
tem to adopt, and what crop to grow can be
answered, at least in part, through the use of
climatic data.

The monthly water balance presented above
is by no means precise because the median rain-
fall is only an approximation of the effective
rainfall. The amount of effective rainfall varies
with the rainfall intensity as well as with the
moisture storage capacity of the soil. To credit
rainfall correctly, it is necessary to compute the
daily water balance.

Fig. 14. Annual water deficit (inches) in Hawaii.

392

PACIFIC SCIENCE, Vol. XVII, October 1963

I960 JULY AUG SEPT OCT NOV DEC

FIG. 15. Running daily soil-moisture balance at Waipio. Dashed lines indicate irrigation.

DAILY WATER BALANCE

If the soil storage capacity is known, the daily
water balance can be computed by comparing
the daily rainfall and potential evapotranspira-
tion. Figure 15 shows the running daily soil
moisture balance at Waipio, where the storage
capacity is 2.5 inches. The rainfall in excess of
the amount that a soil can hold is regarded as
surplus. By subtracting the surplus from the
rainfall, the effective rainfall can be determined.
When the daily water balance reaches zero,
drought occurs and irrigation is called for. The
water deficits during the drought days can be
added to determine the total water deficit dur-
ing the growing season of a crop.

This daily water-balance scheme assumes that
the depletion rate of soil moisture is equal to
potential as long as the soil moisture is above
the wilting point. This assumption is adopted
primarily for simplicity of computation. It
should not be taken to mean that we endorse
the well-known Veihmeyer and Hendrickson
argument (1955). Although the relationship
between the depletion rate and the moisture
tension is still debatable, there are indica-
tions that the relationship probably varies with
weather conditions (Denmead, 1961). The con-
stant depletion rate reported by Veihmeyer and
Hendrickson is probably less in error in a humid,
cloudy climate, especially under the cover of

tall vegetation, than in a dry continental climate.

The computed daily water balance at Waipio
has been compared with gypsum-block readings.
They agree in general. When the water balance
reaches zero, the gypsum-block resistance is
usually between 3,000 and 5,000 ohms. The
latter value calls for irrigation. In general, the
irrigation date determined by the pan evapora-
tion is one or two days earlier than that indi-
cated by the block reading.

KOHALA RESULTS

Kohala is a plantation on northern Hawaii
with an annual water deficit varying from 20
to more than 50 inches. In I960 only about
27% of the plantation was irrigated. The ques-
tion then arises as to whether an expanded irri-
gation program will be profitable in the long
run. Apparently this question cannot be an-
swered without an investigation of the climate-
and-yield relationship.

The daily water balances during more than
10 years for one irrigated and five unirrigated
stations in Kohala were computed by using a
storage capacity of 3 inches. The total water
deficits for different crops were summarized
and correlated with the yields (Fig. 16). The
correlations are highly significant for Hawi,
Niulii 9, Halawa 3, and Upolu 6, but are poor
for Puakea 6 and Union 8. In general they are
very good for field crops, especially in view of

Climatology and Sugar Cane — CHANG

393

the fact that there were considerable differences
in crop age, solar radiation, cultural practice,
variety, etc.

The average slope of the regression lines for
the five unirrigated fields is 4.6 tons of cane
per acre for every 10 inches of water. This is an
important figure in determining the economy
of irrigation. It is thought that by extending the
regression lines to the point of zero deficit the
approximate yield potential of the area could be
estimated. The average yield for the five unirri-
gated fields was 65.2 tons of cane per acre. The
estimated potential would be 91.4 tons cane per
acre, or an increase of 40% over the present
yield. This estimate is probably reasonable when
compared with the yield of 109 tons of cane per
acre at Waialua, Oahu, an irrigated plantation
with slightly better climatic conditions.

Admittedly the use of linear extrapolation as
a means of estimating yield potential is a crude
procedure. Yet this seems to be the most rea-
sonable method we can adopt at present. The-
oretically the relationship between water and
yield should be curvilinear, as shown in Figure

17. The exact shape of the curve cannot be
determined until results of the Waipio experi-
ments become available. The difference in yield
potential between the two curves in Figure 17
is caused primarily by radiation, if soil and
other factors are constant. Experiments are also
underway to assess the effect of solar radiation
on yield. It is hoped that in future this con-
certed research program will enable us to draw
with certainty the climate-and-yield relationship
now presented hypothetically in Figure 17.

By analyzing the daily water balance at Ko-
hala, we also derived an empirical relationship
between the actual rainfall and the effective
rainfall for plant use. The computation was car-
ried out by assuming varying soil-moisture
storage capacities of 2, 3, and 4 inches. Figure
18 shows the curvilinear relationships between
the annual rainfall and effective rainfall for all
the five stations combined. Multiple regression
analyses were carried out to the fourth power
and a few selected values are given in Table 2.
It is noted that the effective rainfall increases
with the storage capacity only slightly, espe-

U 60

-I — J 1 1 I 1 I _L l

10 20 30 40 50 60 70 80 90

WATER DEFICIT (INCHES) IN 24 MONTHS

FIG. 16. Relationships between cane yield and water deficit for six fields at Kohala.

394

ANNUAL RAINFALL (INCHES)

Fig. 17. Hypothetical curves showing the relation-
ship between cane yield and climate in Hawaii.

dally when the annual rainfall is low. The Ko-
hala results could be applied to other areas for
estimating effective rainfall, as the rainfall
characteristics are more or less similar through-
out the islands.

JUICE QUALITY

The dry matter production of a sugar-cane
crop can be related in a quantitative manner to
the solar radiation and the amount of effective
water applied, if cultural practices remain the
same. The problem of juice quality is more
complicated, however, and at present can be
dealt with only in a rough empirical manner.

The weather factors that favor high sucrose
content are:

PACIFIC SCIENCE, Vol. XVII, October 1963

1 ) High radiation favors photosynthesis and
formation of sucrose.

2) Water supply should be adequate during
the growing season, but should be followed by
a dry period of about two months before har-
vest. High rainfall during the ripening period
will dilute the juice and cause a part of the
sucrose to break down into other sugars.

3) It is generally known that cool weather
preceding the harvest season improves the su-
crose content of the cane. Less well known,
however, is the beneficial effect of large diurnal
temperature range. Harrington (1923) and
Morinaga (1926) have shown that alternating
high and low temperatures hasten the germina-
tion of seeds and bulbs. If the buds of sugar
cane react as favorably towards a great range of
temperature, then earlier suckering could be ex-
pected, which would mean a greater number of
mature stalks at harvest and hence a higher
sucrose content.

In summary, the ideal weather for high juice
quality would be abundant sunshine, high tem-
perature range, adequate water supply during
the growing season, and dry and cool weather
during the ripening period. A study of Figures
7 and 8 would indicate that the Ewa, HC&S,
Pioneer Mill, and Kekaha plantations have the
best climate for good juice quality.

It is true that we cannot modify the climate
to such an extent as to appreciably improve the
juice quality. Nevertheless, an understanding of

0 10 2030 40 50 60 70 80 90 100
WATER DEFICIT (INCHES) IN 2 YEARS

Fig. 18. Relationships between annual rainfall and
effective rainfall in Kohala, assuming varying soil-
moisture storage capacities of 2, 3, and 4 inches.

Climatology and Sugar Cane- -CHANG

395

TABLE 2

Effective Rainfall as a Function of Annual Rainfall at Kohala
(Assuming varying soil moisture storage capacities of 2, 3, and 4 inches)

Annual rainfall (inches)

Effective rainfall for:

100

2 -inch storage

34 -

3 -inch storage

4~inch storage

the climatic effect on juice quality has at least
two practical applications. First, we can schedule
planting and harvest dates accordingly, in order
to best fit the seasonality of a climate; in this
connection a study of climatic singularity would
be valuable. Second, an understanding of the
role played by climate could help us to interpret
the results from many agronomic experiments.

CONCLUDING REMARKS ON CLIMATOLOGICAL
RESEARCH

This review has demonstrated the broad scope
of climatological research in the Hawaiian sugar
industry. The application of this research, as
Curry (1952) pointed out 10 years ago, could
ultimately give new life to many an economic
activity. Toward that end we have made only a
beginning. Much work remains to be done.
Future research should be planned in three
general areas:

L GENETIC CLIMATOLOGY: Only 20 years
ago climatology was treated primarily as a study
of statistical meteorology, replete with records
but almost devoid of explanations. The post-war
studies of the climate of Hawaii are among the
pioneers in the field of genetic climatology.
These studies bring a fair measure of systematic
order into the otherwise incoherent climatic
facts and explain many seemingly local phenom-
ena in the light of general circulation. A better
understanding of the dynamism and genesis of
climate could conceivably lessen or avert agri-
cultural hazards in many other parts of the
world. It is hoped that Trewartha’s new book
(1961) will stimulate interest and hasten de-
velopment in this field.

2. ENERGY BUDGET AND WATER BALANCE:
Genetic climatology focuses its attention on the

free atmosphere. Local variations, caused by
the interaction of the atmosphere and the ter-
rain, can best be understood by a study of the
energy budget and water balance. Conven-
tional observations, tied to the needs of synoptic
forecasting, are often inadequate for the study
of topoclimate. Urgently needed is an inter -
nationally-standard field instrument for the
measurement of solar radiation. The recent de-
velopment of an instrument which measures
net radiation (Soumi et al, 1954; Fritschen,
1960£), or even net radiation minus soil heat
flow (Portman, 1954), is most encouraging.

Measurements of evapotranspiration by in-
expensive field instruments are also needed on
a wider scale. In humid climates evaporimeters,
such as evaporation pans or atmometers (Halkais
et al, 1955), have been of value. In arid climates,
however, the problem is vexing. There the very
concept of potential evapotranspiration is elu-
sive and unrealistic. For potential evapotranspi-
ration requires, by definition, a homogeneous soil
moisture regime infinite in horizontal extent,
and once the area upwind is adequately watered
the climate is no longer arid. In actuality the
effect of advective heat exists even in a very
large irrigated area. Gal’tsov (1953), for in-
stance, has observed decreasing water require-
ments from the border towards the center of an
irrigated region in Kazakhstan. It is difficult to
evaluate the effect of advective heat. Perhaps an
approach similar to but more refined than Be-
lasco’s study ( 1952 ) on the modification of air
mass is in order.

To solve the water-balance equation the soil
moisture storage capacity needs to be known.
This requires the cooperation of soil scientists.
On a global scale, representative figures of
moisture storage capacity of the major geo-
graphical regions are useful for many purposes.

396

PACIFIC SCIENCE, Vol. XVII, October 1963

In view of the importance of energy budget
and water balance in characterizing the regional
climate, it is a pity that almost all the textbooks
in English on climatology make no reference to
the subjects. The Russian work by Budyko
(1958) is the only general reference. In this
respect, Thornthwaite’s sustained interest in
these areas is commendable, and his recent plea
( 1961 ) should be well heeded.

3. EFFECT OF CLIMATE ON GROWTH AND
YIELD: We have demonstrated that determina-
tion of water balance is a step toward solving
the climate-and-plant relationship. More field
experiments are needed, however, to define the
hypothetical curve as presented in Figure 17.
Eventually we hope that a method will be de-
veloped to estimate dry-matter yield and juice
quality from climatic data.

It has been argued that climatologists are not
responsible for the study of the effects of climate
on vegetation (Trewartha, 1957). To be sure,
plant scientists are better qualified to run the
field experiments. But the development of tech-
niques for evaluating the climatic elements that
affect plant growth falls squarely within the
responsibility of a climatologist. Without these
techniques the meaning of climatic data is often
obscured. Whoever solves the problem of cli-
mate-and-plant relationship will contribute work
of enormous practical value.

REFERENCES

Baer, L. 1956. Orographic rainfall distribution
with application to Hawaii. Trans. Amer.
Geophys. Un. 37:546-547.

Baver, L. D. 1954. The meteorological ap-
proach to irrigation control. Hawaiian Plant.
Rec. 54:291-298.

Belasco, J. E. 1952. Characteristics of air masses
over the British Isles. Geophys. Mem. 11:
1-34.

Biel, E. R. 1929. Die veranderlichkeit der jah-
reeummon des niederschlage auf der Erde.
Geogr. Jber. Ost. 14:46.

Blanchard, D. C. 1953. Raindrop size-distri-
bution in Hawaii rains. J. Met. 10:457-473.
Borden, R. J. 1940. Nitrogen-potash-sunlight
relationships. Hawaiian Plant. Rec. 44:237-
241.

1949. Sunlight and sugar yield. Ha-
waiian Plant. Rec. 53:43-45.

Budyko, M. I. 1958. The Heat Balance of the
Earth’s Surface. U.S. Weather Bureau. 259 pp.
Chang, J. H. 1958. Ground Temperature. Har-
vard University. 300 pp.

1961. Microclimate of sugar cane. Ha-
waiian Plant. Rec. 56:195-225.

Clements, H. F. 1940. Integration of climatic
and physiologic factors with reference to the
production of sugar cane. Hawaiian Plant.

Rec. 44:201-233.

Curry, L. 1952. Climate and economic life: A
new approach. Geogr. Rev. 42:367-383.
Das, U. K. 1928. The influence of weather on
the production of sugar in a typical unirri-
gated plantation of Hawaii. Hawaiian Plant.
Rec. 32:79-107.

1931^. The problem of juice quality.

Hawaiian Plant. Rec. 35:163-201.

1931 b. Weather and the quality of juice

at Ewa. Hawaiian Plant. Rec. 35:135-162.

1932. Weather and crop relationships at

the Honokaa Sugar Company. Hawaiian Plant.
Rec. 36:255-271.

1936. A suggested scheme of irrigation

control using the day-degree system. Hawaiian
Plant. Rec. 40:109-111.

Deacon, E. L., C, B. Priestley, and W. C. Swin-
BANK. 1958. Evaporation and water balance.
Climatology, UNESCO, pp. 9-34.

Denmead, O. T. 1961. Availability of soil
water to plants. Iowa University Ph.D. thesis.
Frazier, H. M. 1957. The weather and circula-
tion of October, 1957. Mon. Weath. Rev.
85:341-349.

FRITSCHEN, L. J. 1960^. Transpiration and evap-
otranspiration as related to meteorological
factors. Iowa State University Ph.D. thesis.

I960 A Construction and calibration of

the thermo-transducer type net radiometer.
Bull. Amer. Met. Soc. 41:180-183.
Gal’tsov, A. P. 1953. O Klimatisheskom
vzaimodeiztvii oroshaemykh i neroshaemykh
ploshchadei. Akad. Nauk SSSR, Ser. Geogr.
3:11-20.

Halkais, N. A., F. J. Veihmeyer, and A. H.
Hendrickson. 1955. Determining water
needs for crops from climatic data. Hilgardia

24:207-233.

Climatology and Sugar Cane — CHANG

397

Harrington, G. T. 1923. Use of alternating
temperatures upon the germination of seeds.
J. Agric. Res. 23:295-332.

Houghton, H. G. 1954. On the annual heat
balance of the northern hemisphere. J. Met.

11:1-9.

Jones, S. B. 1942. Lags and ranges of tempera-
ture in Hawaii. Ann. Assoc. Amer. Geogr. 31:

68-97.

Krogman, K. K., and L. E. Lutwick. 1961.
Consumptive use of water by forage crops in
the upper Kootenay River Valley. Canad. J.
Sci. 41:1-4.

Landsberg, H. 1951. Statistical investigations

into the climatology of rainfall on Oahu. Met.
Monogr. 1:7-23.

Leopold, L. B. 1948. Diurnal weather patterns
on Oahu and Lanai, Hawaii. Pacif. Sci. 2:
81-95.

— 1949. The interaction of trade wind and

sea breeze, Hawaii. J. Met. 6:312-320.
and C. K. Stidd. 1949. A review of con-
cepts in Hawaiian climatology. Pacif. Sci. 3:
215-225.

Loveridge, E. H. 1924. Diurnal variation of pre-
cipitation at Honolulu, Hawaii. Mon. Weath.
Rev. 52:584-585.

Morinaga, T. 1926. Effect of alternating tem-
perature- upon the germination of seeds. Amer.
J. Bot. 13:141-166.

Namias, J., and W. A. Mordy. 1952. The Feb-
ruary minimum in Hawaiian rainfall as a
manifestation of the primary index-cycle of
the general circulation. J. Met. 9: 180-186.
Penman, H. L. 1948. Natural evaporation from
open water, bare soil and grass. Proc. Roy.
Soc Ser. A. 193:120-145.

PORTMAN, D. J. 1954. Direct recording of the
difference between the radiation and conduc-
tion components of the energy balance of the
earth’s surface. Bull. Amer. Met. Soc 35:4.
Ramage, C. S. 1959. Hurricane development. J.
Met. 16:227-237.

Schneider, M. 1952. Sumen, mittel und mitt-
lere extreme der temperatur von phanologis-
chen zeitspannen. Ber. dtsch. Wetter. 7:276-

281.

Simpson, R. H. 1952. Evaluation of the kona
storm, a subtropical cyclone. J. Met. 9:24-35.

Soumi, V. E., M. Franssila, and N. F. Is-
litizer. 1954. An improved net radiation
instrument. J. Met. 12:276-282.

Stidd, C. K., and L. B. Leopold, 1951. The
geographical distribution of average monthly
rainfall, Hawaii. Met. Monogr. 1:24-33.
SUTTON, O. G. 1953. Micrometeorology. Mc-
Graw-Hill Book Co. 240 pp.

Suzuki, S., and H. Fukuda. 1958. A method
of calculating potential evapotranspiration
from pan evaporation data. J. Agric. Met. 13:
81-85.

Swezey, J. A. 1942. Rainfall evaluation as an
aid to irrigation interval control. Hawaiian
Plant. Rec. 46:75-100.

Tanner, C. B., and W. L. Pelton. I960. Po-
tential evapotranspiration estimates by the ap-
proximate energy budget method of Penman.
J. Geophys. Res. 65:3407.

Thornth WAITE, C. W. 1948. An approach
toward a rational classification of climate.
Geogr. Rev. 38:55-94.

1961. The task ahead. Ann. Assoc.

Amer. Geogr. 51:345-356.

Trewartha, G. T. 1957. A reply to what one
geographer wants from climatology. Prof.
Geogr. 9:8-9.

I96I. The Earth’s Problem Climates.

University of Wisconsin Press. 334 pp.
Veihmeyer, F. J., and A. H. Hendrickson.
1955. Does transpiration decrease as the soil
moisture decreases? Trans. Amer. Geophys.
Un. 36:425-448.

Wang, J. Y. I960. A critique of heat unit ap-
proach to plant response. Ecology 41:785-
790.

Went, F. W. 1957. Climate and agriculture. Sci.
Amer. 196:83-94.

Wentworth, C. K. 1949. Directional shift of
trade winds at Honolulu. Pacif. Sci. 3:86-88.
Williams, W. L. S. 1933. Weather charts for
plantation use. Hawaiian Plant. Rec. 37:97-
106.

Yeh, T. C, C. C. Wallen, and J. E. Carson.
1951a. A study of rainfall over Oahu. Met.
Monogr. 1:34-46.

J. E. Carson, and J. J. Marciano.

1951 b. On the relation between the circum-
polar westerly current and rainfall over the
Hawaiian Islands. Met. Monogr. 1:47-55.

Root Development in Aluminous Hawaiian Soils1

D. L. Plucknett,2 J. C Moomaw,3 and C. H. Lamoureux4

ABSTRACT: Roots of Rhodomyrtus tomentosa and Melastoma malabathricum
were excavated in three soil series from the bauxitic area of Kauai. Root systems
of R. tomentosa and M. malabathricum in Kapaa and Halii soils were very shal-
low, with tap roots turning laterally at shallow depth and with long lateral roots
very close to the soil surface. Deeper tap-root penetration of R. tomentosa and
M. malabathricum was observed in the Koolau soil.

Lime and phosphorus treatments were added to bauxitic subsoils of the Kapaa
and Halii series in pots and Leucaena glauca ( L. ) was planted in the pots. Tap roots
of L. glauca were stimulated by phosphorus treatment, but were restricted in
untreated subsoils. Increased root development with phosphorus treatment seemed
to be more related to phosphorus supply than to decreased aluminum effects. No
evidence of root damage due to aluminum was found.

L. glauca roots were sectioned with a freezing microtome and stained, using
hematoxylin without a mordant. Although all staining obtained could not be at-
tributed to aluminum, since other metals can act as mordants for hematoxylin,
intensity of staining was assumed to be related to aluminum concentration in the
tissues. Cell walls, nuclei, and cytoplasm stained in all tissues, and outer walls of
epidermal cells stained very heavily. Staining was more intense in roots from check
and P-treated plants than in roots from lime-treated plants.

Studies of the root development of natural
plant communities (Weaver 1920, Weaver
and Albertson 1943, Weaver and Darland,
1949) and of cultivated plants (Weaver, 1926;
Troughton, 1957; Crider, 1955) under a vari-
ety of conditions and treatments, are well
known to ecologists. Normally, a well-devel-
oped root system is essential to the vigor-

1 Published with the approval of the Director of
the Hawaii Agricultural Experiment Station, Univer-
sity of Hawaii, as Technical Paper 583. Manuscript

received June 8, 1962.

3 Department of Agronomy and Soil Science, Uni-
versity of Hawaii.

3 Department of Agronomy and Soil Science, Uni-
versity of Hawaii. Present address: The Interna-
tional Rice Research Institute, Manila Hotel, Manila,
Philippines.

4 Department of Botany, University of Hawaii.

ous growth and successful competition of the
dominants in most plant associations. Shallow-
rooted plants, however, are known from many
different habitats, especially those with poor
physical properties but also from those with a
low nutritional status with respect to nitrogen,
phosphorus, calcium (Fox, Weaver, and Lipps,
1953) and potassium (Haynes, 1943). Root
damage and restricted root development in such
crop plants as barley and tobacco have been
associated with high levels of soil aluminum,
but little is known of root development in un-
cultivated plants growing in soils with high
concentrations of active aluminum.

In a study of vegetation on gibbsitic Ha-
waiian soils Moomaw and Takahashi (I960) re-
ported shallow root systems, but detailed studies
of root development and distribution in these

398

Root Development in Aluminous Soils— PLUCKNETT, Moomaw, and Lamoureux

399

soils were not made at that time. Two particu-
larly aggressive plant species, Rhodomyrtus
tomentosa (Ait.) Hassk. and Melastoma mala-
bathricum L., are known to grow well on Ha-
waiian bauxitic soils, where Rhodomyrtus tends
to form dense thickets excluding all other
vegetation. These two species were introduced
on Kauai about 50 years ago and Rhodomyrtus
has now been declared a noxious weed. It is the
object of this study to characterize root growth
and distribution of these and other tap-rooted
species on the bauxitic soils of Kauai.

Aluminum has been shown to be toxic to
plants. One of the frequently-reported toxic ef-
fects of aluminum is root injury (Lignon and
Pierre, 1932; McLean et al., 1926; Bortner,
1935) in which roots may be brown in color
with few rootlets and discolored root tips (Gil-
bert and Pember, 1931) or with root tips
blackened and thickened to twice normal size
(Bortner, 1935). Restricted lateral root de-
velopment in rye has been reported using water
cultures (Magistad, 1925). Trenel and Alten
(1934) concluded that aluminum may be a
root poison. Using a divided-root technique corn
plants were exposed to nutrient solutions with
and without aluminum. Injury was restricted to
roots in the high-aluminum solutions. Nagata
(1954) found that over 5 ppm aluminum in
culture solutions hindered barley growth and
that aluminum seemed to accumulate in the
roots. Growth hindrances were decreased by
adding phosphorus or calcium. He concluded
that translocation of phosphorus from the root
to the top in the barley plant was hindered by
aluminum in culture solutions.

The precipitation of phosphate and alumi-
num in the plant as an aluminum phosphate
has been suggested. Burgess and Pember (1923)
proposed that aluminum was fixed as relatively
insoluble aluminum phosphate in plants, espe-
cially in roots. McGeorge ( 1925 ) suggested that
internal precipitation of aluminum by phos-
phorus may be important in plants but listed
no specific location.

Wright ( 1937 ) divided root systems of bar-
ley plants, placing each half in different culture
solutions with and without aluminum. Plant
analysis indicated plant damage resulting from
poorly developed root systems in solutions con-

taining aluminum, and internal precipitation of
phosphorus and aluminum where large amounts
of aluminum and phosphorus were present in
roots. Wright (1943) found a higher percent-
age of phosphorus in aluminum-treated barley
plants than in nontreated; this was particularly
marked in the roots. The water-soluble phos-
phorus in the aluminum-treated plants was low,
while a H2SO4 solution (pH 3.0) extracted
practically all P from untreated plants but much
smaller amounts from plants grown in contact
with Al. The precipitation was listed as occur-
ring primarily in roots, and sharp reductions in
yield were attributed to P deficiency in meri-
stematic regions due to root precipitates. Wright
( 1945 ) , using microchemical tests to determine
inorganically and organically bound P, found
abundant inorganic P in roots grown in contact
with Al and little or none in roots from solu-
tions without Al.

Problems of plant growth on acid soils have
long been ascribed to the "active” Al in the soil
and to problems of phosphate nutrition due to
fixation of phosphates by Al. Longnecker and
Merkle (1952) studied root development of
crimson clover in relation to lime placement
and found most root growth in layers which
had been limed. The beneficial effect of liming
was attributed to decrease in solubility of Al
and Mn and an increase in solubility of P. Rag-
land and Coleman (1959) applied lime at sev-
eral rates to subsoils of the Norfolk catena in
pots and found grain sorghum root growth
into unlimed subsoils was related inversely to
amounts of exchangeable Al. Root growth into
subsoils increased substantially with lime treat-
ment. Root development of sorghum grown in
suspensions of acid clay was restricted severely
unless 80% of the acidity was neutralized.

DESCRIPTION OF EAST KAUAI

The area studied is referred to by McDonald
et al. (I960) as the Lihue Depression. It is a
nearly circular basin with the rim being formed
by the Haupu ridge on the south, the main
mountain mass of central Kauai on the west,
the Makaleha mountains on the north, and
Nonou and Kalepa ridges on the east. The basin
is floored with lavas of the posterosional Koloa

400

volcanic series. Two vents from the Koloa vol-
canic series, Hanahanapuni Crater and Kilohana
Crater, lie within the basin.

The general topography of the basin is of
gently sloping to moderately steep ridges and
plains dissected by perennial streams, notably
the Wailua River and its tributaries.

The average annual rainfall in the Lihue De-
pression ranges from 40 or 50 inches near the
ocean to over 170 inches near the mountains.
Rainfall is usually highest in winter months but
there are no months during which no rain falls.

Mean monthly temperatures from nine sta-
tions below 300 ft elevation on Kauai range
from 69 F in February and March to about 77
F during August through October (McDonald
et al., I960). Although no temperature records
are available for higher elevations, there is a
decrease in temperature with increase in eleva-
tion, of about 3 F for each 1,000 ft.

Prevailing winds are the northeast trade
winds but cyclonic storms occasionally upset this
pattern, especially in winter months.

DESCRIPTION OF SOILS STUDIED

Three soils occurring in the Lihue Depression
were selected for study. These soils are located
in or near the main area of infestation of R.
tomentosa and Al. malabathricum and either
comprise or are associated with the major baux-
itic soils of East Kauai. The principal mineral
form of Al present in these bauxitic soils is
gibbsite, the trihydrate of aluminum oxide
(Sherman, 1958). Detailed soil descriptions have
been made by the Soil Conservation Service
(Womack, I960).

Kapaa Series

The Kapaa series is a deep, well-drained,
Aluminous Ferruginous Latosol developed on
gently sloping to steep uplands on Kauai. These
soils occur mainly in association with the Halii
soils which lie above, and Puhi soils which lie
below. The Halii series is developed from parent
material similar to that of the Kapaa series,
namely the melilite and nepheline basalts of the
Koloa volcanic series. The Kapaa soils occur be-
tween 200 and 1,000 ft elevation with mean
annual rainfall from 60 to 100 inches. They are

PACIFIC SCIENCE, Vol. XVII, October 1963

clayey in texture but feel like silty clay in the
A horizon (0-6 inches) owing to the strong,
very fine granular structure. They are extremely
hard when dry but sticky and plastic when
moist, containing many roots and few pebbles
in the surface layers. With depth, some mottling
occurs on the blocky structures and pebbles are
more numerous. These soils are used mainly for
pasture, nonirrigated sugar cane, and pineapple.

Halii Series

The Halii series is a deep, well-drained Alu-
minous Ferruginous Latosol on gently sloping
to moderately steep uplands on Kauai. This
series is associated with the Koolau series at
higher elevations and with Kapaa series below.
It occurs in belts at about 300 to 1,000 ft eleva-
tion with mean annual rainfall from 80 to 120
inches. The A horizon (0-9 inches) of this
series is a gravelly clay that feels like silty clay
and is grayish-brown in contrast to the yellow-
ish-brown of the Kapaa series. It is somewhat
less hard when dry and more plastic when wet
than the Kapaa. It is used principally for non-
irrigated sugar cane and small acreages of pine-
apple.

Koolau Series

The Koolau series is a deep, poorly-drained
Hydrol Humic Latosol developed on gently slop-
ing to moderately steep uplands. It is associated
with the Halii soils, and occurs between 400
and 4,000 ft elevation with mean annual rain-
fall of 120 and 200 inches. Most of the Koolau
series is covered by rain forest, but some is used
for sugar cane and pasture.

The Koolau series is a grayish-brown clay that
feels like a silty clay loam, with a structure that
appears to be massive to very weak medium
granular. It is only slightly sticky and plastic
when moist, with matted roots and many worm
holes and casts making it quite porous. Lower
horizons have distinct strong brown mottles.

EXPERIMENTAL PROCEDURE

Roots of Melastoma malabathricum and Rho-
domyrtus tomentosa were excavated in the
Halii, Koolau, and Kapaa soil series and de-
scriptions and measurements were made of

Root Development in Aluminous Soils — Plucknett, Moomaw, and Lamoureux

401

depth of penetration of tap roots, location and
length of lateral roots, relation of roots to
soil profile, and possible evidence of causes of
thicket formation. A few plants of Norfolk
Island pine ( Araucaria excelsa) (Lamb.) R. Bn.
were also excavated in the Wailua Game Refuge.

Root description and measurement were also
made of R. tomentosa seedlings used in a pot
experiment designed to measure plant and soil
Al. Notes taken included color, thickening, black
tips, number of lateral roots, length of tap roots,
and number of active buds.

Because tap roots were observed turning
laterally in Halii and Kapaa soils in the field, a
pot experiment was established using the soil
layer in which these roots turned as "subsoils.’’
These soils were collected from horizons where
tap roots were observed to turn laterally and
were sacked carefully to prevent dehydration.
They were screened through wire mesh con-
taining approximately 4 meshes to the inch.
Weighed samples of the screened soil were used
to form the "bottom” 5 -inch soil layer in 11-
inch plastic pots, and were treated with six lime
and phosphate treatments. After the treatments
were mixed thoroughly in the subsoils, a 5 -inch
layer of untreated Kapaa surface soil was added.
Leucaena glauca 5 seeds were planted in the sur-
face soil and after germination plants were
thinned to two per pot.

At harvest the soils were carefully removed
from the pots and washed from the roots. Meas-
urements of tap-root penetration and lateral
root development were made. Roots were ex-
amined for blunted and blackened tips, and root
tips from each treatment were preserved for
staining studies. Yields of tops and roots were
recorded and plant Al concentrations were de-
termined. The pH, extractable Al, exchangeable
calcium, and cation exchange capacity were de-
termined for each soil.

Root tips of L, glauca plants from treated pots
were sectioned on a freezing microtome and
stained, using hematoxylin without a mordant
(Johansen, 1940). Slides of the root sections
were made and photomicrographs were taken.

6 In a personal communication Dr. F. R. Fosberg
has indicated the correct name for this plant should
be Leucaena leucocepbala.

Fig. 1. Rhodomyrtus tomentosa excavated in the
Kapaa soil series, Wailua Game Refuge. The tap root
turned laterally at 4-inch depth and lateral roots pene-
trated diagonally before ascending toward the surface.

RESULTS

Six R. tomentosa plants were excavated in
the Kapaa soil series. All plants were extremely
shallow-rooted, with tap roots turning laterally
at depths of from 3 to 10 inches below the sur-
face (Fig. 1 ) . Lateral root development irnrhese
plants was especially pronounced and one large
10-ft shrub had a lateral root 24 ft long. Lateral
roots displayed a tendency to grow downward
and outward for 2 to 4 inches and then to as-
cend toward the surface. Lateral roots were
frequently found just at, or slightly under, the
soil surface.

Two thickets of R. tomentosa were excavated
in the Kapaa soil series. These thickets con-
tained shrubs up to 10 ft in height with trunks
1.5 to 2.25 inches in diameter. Roots were ob-
served with diameters up to 2 inches. Tap roots
turned at a depth of 10 inches and no roots of
the thickets were found below this depth. There
was no evidence of root fusion.

M. malabathricum plants excavated in the
Kapaa soil were found to have root development
similar to that in R. tomentosa, with tap roots
turning laterally at shallow depth and long lat-
eral roots. One plant (Fig. 2) had a small,
twisted and deformed tap root with 2 main
lateral roots 5 ft long. Fibrous roots were almost
lacking in both R. tomentosa and M. mala-
bathricum.

Norfolk Island pine trees (A. excelsa) planted
about 10 years ago in the Kapaa soils were

402

excavated to determine root development in
species planted in these soils. Tap roots of these
trees penetrated to what appeared to be the
bottom of the planting hole before turning up-
ward toward the surface.

Two individual plants and a thicket growth
of R. foment os a were excavated in the Koolau
soil near Hanahanapuni Crater. Tap roots of
the two individual plants penetrated 9 inches
downward before turning diagonally for 1 or
2 inches. Small plants of M. malabathricum
nearby, like the R. tomentosa, had tap roots
which penetrated from 8 to 10 inches.

The main tap root of the R. tomentosa thicket
penetrated 24 inches downward in the Koolau
soil without turning, even though water from
the soil filled the hole at a depth of 1 ft, placing
it well below the water table at the time of
sampling. One lateral root of this thicket was
observed from which numerous stems had
arisen. This was the only observation of this
type in the plants excavated. It is possible that
this "root” could have been a stem buried by
road construction since the thicket was located
dose to a forest preserve road. The Koolau soil
in this area has a grey surface 8 to 10 inches in
thickness with a reddish-brown, yellow-mottled
layer below.

A 5 -ft M. malabathricum plant about 20 ft
from the R. tomentosa thicket was also ex-
cavated and the tap root was traced to a depth
of 18 inches where water rapidly filled the
trench.

A series of R. tomentosa plants in the Halii

Fig. 2. Melastoma malabathricum excavated in the
Kapaa soil series, Wailua Game Refuge. Note the
shallow lateral roots and the small tap root.

PACIFIC SCIENCE, Vol. XVII, October 1963

Fig. 3. Plants of Rhodomyrtus tomentosa from the
Halii soil series on Kilohana Crater, Kauai. The lateral
root of the larger plant was 11 ft long.

soil series were excavated on the northern slopes
of Kilohana Crater. These plants had extremely
shallow root systems with tap roots turning
laterally at about a 4-inch depth and with lateral
roots almost at the soil surface (Fig. 3). R. to-
mentosa shrubs in this area were easily pulled
up without digging, and tracing of lateral roots
was accomplished by pulling. Excavation of such
roots was difficult because of long overlapping
lateral roots of surrounding plants. One 40-inch
plant had a lateral root which arose from the
tap root about 2 inches below the soil surface
and which was traced at depths of 1 inch or less
for 11 ft.

No evidence of root injury was found in
R. tomentosa plants grown in pots except for
an unusual blunting of root tips in the Halii
soil. This blunting is best described as a curving
and thickening of the root tip which then re-
sembled a miniature chicken’s head. Some black-
ened tips were also found in the Halii soil, but
since the concretionary iron-rich surface of the
Halii soil was used in these pots it is doubtful
that these root abnormalities were due to A1
injury.

The only evidence for thicket formation of
R. tomentosa in this pot study was the presence
of a number of buds on the tap root just below
the soil surface. Young shoots were observed
arising from these buds and often 6 to 10 young
shoots were growing simultaneously on one
plant. The number of shoots arising from buds
increased with time. Bud numbers of single
plants ranged from 5 to 30.

Root Development in Aluminous Soils— Plucknett, Moomaw, and LAMOUREUX

403

Root: Development of Leucaena glauca

•Root development of Leucaena glauca in
treated subsoils of the Halii and Kapaa soil
series was investigated separately in pots. In
both the Kapaa and Halii subsoils root develop-
ment was usually restricted to the untreated top-
soil if subsoils were untreated. The greatest root
development in subsoils was produced by P
treatments. Treatments in which lime was added
in addition to P stimulated root growth more
than lime alone. The addition of 1,000 lb of ele-
mental P without lime produced the most root
penetration into the subsoils and also the
highest plant yields.

Comparative root systems produced by treat-
ments in the Kapaa subsoil are illustrated in
Figure 4. Tap roots of L. glauca in the P treat-
ments were straight and penetrated to the bot-
tom of the pot. Tap roots of check plants did not
develop in the untreated subsoils. The effect of
treatment on tap-root penetration and root de-
velopment in treated Kapaa subsoil ranked as
follows: P > lime plus P > lime > check.

Figure 5 shows comparative root systems in
Halii subsoils. Roots of check plants in the
Halii subsoils penetrated slightly into the sub-
soil, but total root development in the check was
much less than total development in subsoils
treated with P and with lime plus P.

Two types of tap-root development with
treatment were observed: A long straight tap
root was characteristic of P treatment, while a

Fig. 4. Plants of Leucaena glauca grown in treated
subsoils of the Kapaa series. Treatments from left to
right are: 500 lb P, 1,000 lb P. 5 tons lime, 5 tons
lime plus 1,000 lb P, and check.

Fig. 5. Plants of Leucaena glauca grown in treated
subsoils of the Halii series. Treatments from left to
right are: check, 5 tons lime plus 1,000 lb P, 5 tons
lime, and 1,000 lb P.

branching tap root was characteristic of the
lime plus P treatment.

An interesting result of treatments added to
Halii subsoils was the number of nodules pro-
duced on roots of L. glauca. Most nodulation
occurred with lime plus P, but lime alone also
stimulated nodule formation. Only one nodule
was found with P treatment, and no nodules
were found in the check.

Staining of Leucaena glauca Root Tips

Hematoxylin staining of cell walls, nuclei, and
cytoplasm was evident (Fig. 6d) . Outer walls of
epidermal cells were especially heavily stained
( Fig. 6a, b) , which may indicate a precipitation
of aluminum in this region. In P- treated plants
two darkly stained areas at the periphery of the
stele were observed, which may also represent
areas of aluminum precipitation. One of these
areas is shown in Figure 6c. The cells of the
stele appeared to stain more intensely than those
of the cortex (Fig. 6a, b), but the stelar cells
are less highly vacuolated and thus contain
more stainable material per unit volume than
the cortical cells.

Staining was more intense in roots from the
check (Fig. 6a) than in roots from lime-treated
soils (Fig. 6b). Phosphate-treated roots stained
more heavily than Jime-treated roots. Roots
treated with a combination of 5 tons lime
plus 1,000 lb P stained more deeply than those
treated with lime alone, but in most cases

404

PACIFIC SCIENCE, Vol. XVII, October 1963

stained less deeply than those treated with P
alone. Staining of nuclei was especially pro-
nounced in the 1,000 lb P treatment.

The results obtained in this study differ from
those reported by McLean and Gilbert (1927)
for corn and cabbage, and by Wright and
Donahue (1953) for barley, in that cells from
stelar regions stained readily in all our prepara-
tions. Although McLean and Gilbert noted stain-
ing of nuclei and cytoplasm, this staining was
restricted to the epidermis and outer cortex, and
none of the stelar cells were stained. Wright and
Donahue found that staining occurred from the
epidermis to the outer wall of the endodermis,
but that there was very little staining in the
stelar region.

DISCUSSION

The importance of the effect of high soil A1
on root growth cannot be minimized, but shal-
low root development in the Halii and Kapaa
soil series was not interpreted as resulting from
"A1 toxicity.” High soil A1 can cause conditions
in the soil which may limit root development,
however. From previous work, it is known that
A1 can interfere with phosphate nutrition of
the plant both by precipitation of phosphorus in
the soil and possibly by precipitation within
the plant. In addition, A1 is thought to con-
tribute to the acidity of the soil. If high alu-
mina content of these subsoils is important in
limiting root development of plants in bauxitic
soils, any decrease in extractable A1 and plant A1

Fig. 6. Photomicrographs of Leucaena glauca roots sectioned and stained with hematoxylin. A, Root sec-
tion from the untreated (check) subsoil of the Kapaa soil series. B, Section from Kapaa subsoil treated with
2.5 tons lime. C, Root from Kapaa subsoil treated with 500 lb P. Note heavily stained area in xylem region.
D, Cross-section of root from Kapaa subsoil treated with 5 tons lime plus 1,000 lb P. Note staining of nuclei,
cell walls, and cytoplasm.

Root Development in Aluminous Soils — Plucknett, Moomaw, and Lamoureux

405

should result in stimulated root growth. An ex-
amination of results of liming treatments shows,
however, that although pH was increased and
extractable A1 and plant A1 concentrations were
decreased by liming, no marked stimulation of
root development or plant growth of L. glauca
occurred with liming. The increased root de-
velopment of L. glauca in P- treated Halii and
Kapaa subsoils in pots is interpreted more as a
response to P than a decrease in A1 effects. In
addition, of course, a mass-action effect may be
operating in which active aluminum in the soil
is being supplied with enough phosphorus to
permit complete precipitation as aluminum
phosphate with enough remaining to supply the
plant with adequate P.

Deep tap-root development in the Koolau
soils was unexpected because of the extremely
poorly-drained condition of this soil. Root
growth in the wet Koolau soil in pots also ap-
peared normal even though figures for extract-
able A1 were high.

The lack of root damage in plants used in this
study was probably related to the evolutionary
background of the plants, which seem to thrive
in areas of low fertility and high rainfall. Roots
of plants sensitive to Al, like rye or barley,
would probably be severely injured in these
soils. But Leucaena glauca, like many other
tropical plants, does not respond markedly to
lime.

Thicket formation in R. tomentosa is prob-
ably caused by the large number of adventi-
tious shoots which arise from buds on the tap
root just below the soil surface.

Heavy staining of roots with hematoxylin
cannot be definitely attributed to Al alone, since
iron and other metals may also act as a mordant
for hematoxylin. However, since in the present
study chemical analysis showed Al to be present
in large amounts, much of the staining obtained
is attributed to Al. The particularly intense
staining of the outer walls of the epidermis, and
of two areas in the outer part of the stele, are
interpreted as an indication that Al precipita-
tion may have occurred in these areas.

Our results appear to differ significantly from
those of McLean and Gilbert (1927) and of
Wright and Donahue (1953). The reasons for
these differences are not immediately apparent.

In all three studies root sections were stained
in hematoxylin without the addition of a mor-
dant, and the staining which occurred was inter-
preted as an indication that aluminum was
already present in the tissues.

REFERENCES

Bortner, C. E. 1935. Toxicity of manganese to
Turkish tobacco in acid Kentucky soils. Soil
Sci. 39:15-33.

Burgess, P. S., and F. R. Pember. 1923. Active
Al as a factor detrimental to crop production
in many acid soils. R. I. Agri. Expt. Sta. Bull.
194. 40 pp.

Crider, F. J. 1955. Root growth stoppage re-
sulting from defoliation of grass. USDA
Tech. Bull. 1102. 23 pp.

Fox, R. L., E. J. Weaver, and R. C. Lipps.
1953. Influence of certain soil profile charac-
teristics upon the distribution of roots of
grasses. Agron. J. 45:583-589.

Gilbert, B. E., and F. R. Pember. 1931. Fur-
ther evidence concerning the toxic action of
aluminum in connection with plant growth.

Soil Sci. 31:4, 267-273.

Haynes, J. L. 1943. Effects of pasture practices
on root distribution. J. Amer. Soc. Agron.
35:10-18.

Johansen, D. A. 1940. Plant Microtechnique.
McGraw-Hill, New York.

Lignon, W. S., and W. H. Pierre. 1932. Sol-
uble Al studies, II. Soil Sci. 34:307-322.

Longnecker, D., and F. G. Merkle. 1952.
Influence of placement of lime compounds on
root development and soil characteristics. Soil
Sci. 73:71-75.

Magistad, O. C. 1925. The Al content of the
soil solution and its relation to soil reaction
and plant growth. Soil Sci. 20:181-225.

McDonald, G. A., D. A. Davis, and D. C.
Cox. I960. Geology and Groundwater re-
sources of the Island of Kauai, Hawaii. Ha-
waii Div. Hydrogr. Bull. 13.

406

PACIFIC SCIENCE, Vol. XVII, October 1963

McGeorge, W. T. 1925. The influence of Al,
Mn, and Fe salts upon the growth of sugar
cane, and their relation to the infertility of
acid island soils. Hawaiian Sugar Planters’
Assoc. Expt. Sta. Bull. Agri. Chem. Sec. 49.

McLean, F. T., and B. E. Gilbert. 1927. The
relative aluminum tolerance of crop plants.
Soil Sci. 24(3): 163-174.

Moo MAW, J. Q, and M. Takahashi. I960.
Vegetation on gibbsitic soils in Hawaii. J.
Arnold Arb. 61:4, 391-411.

Nagata, T. 1954. Influence of Al-ion concen-
tration on the absorption of nutrients by
plants. I, II. J. Sci. Soil Manure Japan 24:
255-262. (C A. 48, 7710).

Plucknett, Donald L. 1961. Root growth in
bauxitic soils. Hawaii Farm Sci. 10(1) :8-9.

Ragland, J. L., and N. T. Coleman. 1959-
The effect of soil solution aluminum and cal-
cium on root growth. Soil Sci. Proc. 23:35 5—
357.

Sherman, G. D. 1958. Gibbsite-rich soils of the
Hawaiian Islands. Hawaii Agri. Expt. Sta.
Bull. 166.

TRENEL, M., and F. ALTEN. 1934. Die physi-
ologische Bedeutung der mineralischen Bo-
denaziditat. Angew. Chem. 46:313-320.

Troughton, A. 1957. The underground or-
gans of herbage grasses. Commonwealth Bur.
Pastures and Field Crops Bull. 44. 163 pp.

Weaver, J. E. 1920. Root development in
grassland formations. A correlation of root
systems of native vegetation and crop plants.
Carnegie Inst., Wash. Publ. 292. 151 pp.

1926. Root Development of Field

Crops. McGraw-Hill Book Co. Ltd., N. Y.
291 pp.

and F. W. Albertson. 1943. Resur-
vey of grasses, forbs, and underground plant
parts at the end of the great drought. Ecol.
Monogr. 13:63-117.

and R. W. Darland. 1949. Soil-root

relationships of certain native grasses in vari-
ous soil types. Ecol. Monogr. 19:303-338.

Womack, Durwajrd W. I960. Tentative De-
scription Legend of the Soil Survey of the
Island of Kauai. May, I960. Soil Conservation
Serv., State of Hawaii. (Mimeo.)

Wright, K. E. 1937. Effects of phosphate and
lime in reducing aluminum toxicity of acid
soils. Plant Physiol. 12 ( 1 ): 173-181.

— - — — 1943. Internal precipitation of P in
relation to Al toxicity. Plant Physiol. 18:708-
712.

— 1945. Aluminum toxicity: Microchem-

ical tests for inorganically and organically
bound phosphorus. Plant Physiol. 20:310-
312.

— and B. A. Donahue. 1953. Aluminum

toxicity studies with radioactive phosphorus.
Plant Physiol. 28:674-680.

Studies in the Hawaiian Rutaceae, IV
New and Critical Species of Pelea A. Gray1

Benjamin C. Stone2

The following notes, including descriptions
of four proposed new species, are the partial
outcome of monographic studies of Hawaiian
genera. Pelea is a genus confined to the Hawai-
ian and Marquesan archipelagoes, with all but
two of its species endemic to the Hawaiian
Islands. There*are four relatively natural sections
of the genus, characterized primarily by features
of the mature fruits and to a lesser extent by
leaf arrangement. Systematic treatment of sec-
tion Pelea , which includes the type species, Pelea
clusiaefolia, is completed. The following notes
apply to sections Apocarpa, Megacarpa, and
Cubicarpa, as defined by the writer (Stone in
Degener, 1962 ) . Related studies have recently
appeared or are now in press (Stone 1962^,
1962b, Stone in Degener, 1962). Casual collect-
ing of species of Pelea was begun by the author
in 1955; in 1958 and 1959 and summer, 1961,
intensive field and herbarium studies were car-
ried out. Through the courtesy of the U. S. Na-
tional Herbarium, Smithsonian Institution, loans
of historically important collections were ob-
tained, and I am grateful to Dr. H. R. Fletcher,
of the Royal Botanic Gardens, Edinburgh, Dr.
George Taylor of the Royal Botanic Gardens,
Kew, Dr. J. E. Dandy of the British Museum
(Natural History), Dr. Alicia Lourteig of the
Museum d’Histoire Naturelle, Paris, Dr. Richard
A. Howard of the Arnold Arboretum of Harvard
University, and Dr. A. J. Eames of Cornell Uni-
versity for their aid. I am particularly indebted
to the former Director of the Bishop Museum,
Dr. Alexander Spoehr, Botanist Marie C. Neal,
and Curator of Collections E. H. Bryan, Jr., for
their generous assistance.

The following comments are arranged by ge-
neric section.

1 The greater part of this work was accomplished at
the University of Hawaii and at the U. S. National
Herbarium. Manuscript received February 27, 1962.

2 College of Guam, Agana, Guam.

SECTION Apocarpa STONE

1. Pelea ovata St. John & Hume in Lloydia 7:

272, 1944.

P. Forbesii St. John & Hume, l.c. Syn. nov.

Examination of the holotype specimens shows
that the pubescence which is found on Forbes
369.K, the type of P. Forbesii, is not constant,
but is found only on one leaf, perhaps from an
abnormal cause. It has been noted that leaf-galls
of other species, even ordinarily glabrous ones,
will often be densely puberulent. Even the new
leaves and buds of P. Forbesii are found to be
glabrous. Since no other character or combina-
tion of characters appears to distinguish this
from Pelea ovata, it is concluded that but one
species is represented. Since the description of
P. ovata calls for glabrous leaves, and since there
is no question of priority involved, both descrip-
tions first appearing on the same page of the
publication, the name P. Forbesii is relegated to
synonymy.

Pelea ovata is a vinelike shrub endemic to the
island of Kauai.

2. Pelea hawaiensis Wawra in Flora 56:110.

1873.

P. cinerea var. y Hillebrand, FI. Haw. Ids. 69,
1888; Rock, Indig. Trees Haw. Ids. 239,
I913 (in part, excluding Oahu plants).

P. cinerea var. hawaiensis (Wawra) Rock in
Bot. Gaz. 65:265, 1918.

A tree with smooth light-brown bark, op-
posite leaves, and generally pubescent innova-
tions; the pubescence of fulvous or reddish hairs,
rarely pale; petioles, twigs, and leaves sometimes
soon glabrate, commonly with a more or less per-
sistent indument; inflorescences usually densely
puberulent, or tomentose, cymose, with generally
3 to 27 flowers, axillary; flowers with deltoid
acute thick tomentellous sepals; petals deltoid-

407

408

PACIFIC SCIENCE, Vol. XVII, October 1963

lanceolate, tomentellous, often reddish within;
stamens glabrous; ovary densely tawny or golden
tomentellous; style hirsutulous; stigma dark
maroon, glabrous; capsule commonly 16-34 mm
broad, often subtended by the persistent sepals,
the carpels with a dense persistent fulvous or
reddish pilosity; endocarp firm, thickly pilose
with pale hairs.

TYPE: Hawaii: Kawaihae-iuka, in 1862, Wm.
Hillebrand (in the Wawra Herbarium, Vienna).
A photo shows capsules 27 mm broad.

(a) var. hawaiensis

The typical variety, which is certainly distinct
from Pelea cinerea, as was stated (in herb.) by
the late C. N. Forbes, is nonetheless a southeast
counterpart of that species. The distribution of

the typical variety includes Hawaii and Maui.
Other varieties, some of them eminently distinct,
others only poorly differentiated, are found on
Hawaii, Maui, Lanai, and Molokai.

( b ) var. rubra ( Rock ) B. C. Stone, comb. nov.

P. cinerea var. rubra Rock in Bot. Gaz. 65:
264, 1918.

P. oblon^ifolia var. /3 Hillebrand, FI. Haw.
Ids. 65, 1888.

P. cinerea var. 8 Rock, Indig. Trees Haw. Ids.
239, 1913.

P. cinerea var. 8 Hillebrand, op. cit. 69. (as
to Kau specimen).

Hillebrand tells us that this is a glabrous
shrub with long rambling branches. The poorly

KEY TO VARIETIES OF Pelea hawaiensis

Capsules mostly 27-34 mm diameter (rarely smaller); inflorescences often few-flowered (3-9).
Leaves puberulent beneath, at least on the costa, at first ( and often persistently ) uniformly
tomentellous to pilose; inflorescence compact, with stout axes, peduncles ca. 2 mm thick,

densely tomentellous; capsular pubescence fulvous to brownish-orange. Hawaii

var. hawaiensis

Leaves glabrous, glossy; inflorescences elongate, the axes slender, peduncle to 1 mm thick,
glabrous or nearly so; capsular pubescence reddish. Hawaii var. rubra

Capsules smaller, mostly 16-22 mm diameter; inflorescences mostly compact and multiflorous
with 9-27 flowers, or rarely more open and with fewer flowers.

Blades densely shaggy-pilose beneath, short-lanceolate; branchlets puberulent; inflorescences
mostly 3-9-flowered.

Blades cordate at base; tomentum fulvous, the hairs extremely minute. Molokai

var. molokaiana

Blades subcuneate to rounded at base; tomentum brownish or olivaceous, the hairs up to

0.7 mm long. Maui, Lanai var. pilosa

Blades tomentellous to glabrate beneath, sometimes at last glabrous; inflorescence several-
to many-flowered (with up to 27 flowers), or less commonly only 3-7 -flowered.
Inflorescences mostly only 3-7-flowered.

Blades cordate or emarginate at base, slightly puberulent on the costa dorsally, elon-
gate and often acuminate. Lanai... var. sulfurea

Blades cuneate at base, glabrate to glabrous beneath, rather broad, ovate. Maui

var. Brighamii

Inflorescences mostly 9-27-flowered.

Blades cordate or subcordate at base, or sometimes merely emarginate; inflorescences
about 4-6 cm long.

Blades glabrate beneath; petioles mostly 20-45 mm long. Maui var. Remyana

Blades puberulent beneath; petioles mostly 7-18 mm long. Maui ..var. racemiflora

Blades cuneate or rarely slightly emarginate at base; inflorescences compact, 2-3 cm
long; costa puberulent or glabrous dorsally. Hawaii var. Gaudichaudii

Critical Species of Pelea—~$ TONE

409

developed capsule of the specimen which he
described (with a question mark) as a variety
of Pelea ohlongifolia (a species of sect. Mega -
carp a) misled him; on closer examination, it
can be seen that only one follicle is developed,
and it is thus difficult to ascertain whether the
fruit is apocarpous or syncarpous. However, the
tomentum of both the follicle and the endocarp,
as well as the leaf venation, suffice to establish
the relationship of this plant with P. hawaiensis ,
rather than with the similar P. cinerea of Oahu
(which has pubescent endocarp but glabrous
follicles), or the greatly different P. oblongifolia
of Hawaii ( with glabrous endocarp and sparsely
puberulent syncarpous capsules).

TYPE: Hawaii: North Kona; Huehue, lava-
beds, 6 June 1909, Rock 3565 (Bishop).

Distribution: Kona and Kau, Hawaii; Olo
walu, Maui.

SPECIMENS EXAMINED: Maui: Central ridge
of Olowalu valley, 12 May 1920, Forbes 2345. M.
(Bishop). Hawaii: North Kona, Puuwaawaa,
6 June 1909, Rock 3561 (Bishop). Kau, Hille-
brand 251 (Kew), 252 (Kew), without number
(Kew). Kau Desert, 2 August 1911, Forbes
385, H. (Bishop).

P. cinerea (3 var. Hillebrand, FI. Haw. Ids.
69, 1888 (as to Lanai specimens with
pilose blades).

This variety is limited to Lanai and Maui.
Hillebrand’s description is based mostly on the
Lanai specimen, although he cites Mann & Brig-
ham 371 from Makawao, Maui. This is a mis-
print for 377; Hillebrand wrote 377 on the label
of his own collection as a note citing the Mann
and Brigham specimen for comparison; but
their specimen is a different variety (var. Brig-
hamii) . Rock erred in calling his Maui speci-
mens the same, and, after coining the name
sulfurea, applied it to both the Hillebrand and
the Mann and Brigham specimens; var. sulfurea
is restricted to Lanai. Unfortunately, the type of
var. pilosa was destroyed at Berlin during World
War II.

SPECIMENS EXAMINED: Maui: South slope
of Haleakala, March 1920, Forbes s.n. (Bishop);
Auahi (Auwahi), March 1920, Forbes 2099 .M
2098M., 2097 M., 2115 M., and s.n. (Bishop).

(d) var. molokaiana B. C. Stone, var. nov.

Folia base cordata, infra fulvo-pilosa.

HOLOTYPE: Molokai: Ridge below Puu Ko-
lekole, July 1912, Forbes 126 Mo. (Bishop; iso-
type at US ) .

DISTRIBUTION: Restricted to Molokai.

The innovations are densely tawny-hirsutu-
lous; blades lance-ovate, rounded or acute at
apex, subcordate to cordate at base, up to 8 X
3.5 cm, with petioles densely hirsutulous, as are
the lower surfaces of the blades.

(e) var. sulfurea (Rock) B. C. Stone, comb,
nov.

P. cinerea var. sulfurea Rock in Bot. Gaz. 65 :
265, 1918.

P. sulfurea (Rock) St. John & Hume in
Lloydia 7:274, 1944.

TYPE: Lanai: Without locality, July 1870,
Hillebrand (Kew; isotype, Bishop).

DISTRIBUTION ; Restricted to Lanai. Abundant
collections are in the Bishop Museum Herba-
rium.

(f) var. Brighamii (St. John) B. C. Stone,
comb. nov.

P. Brighami St. John in Lloydia 7:271, 1944
(sub sect. Cubicarpae) .

TYPE: Maui: Makawao, Mann & Brigham
377 (Bishop, GH).

DISTRIBUTION: Maui.

Described as a species but, no doubt through
a typographic error, misplaced under section
Cubicarpae . This is the variety that Rock had in
mind when he discussed var. sulfurea (in Bot.
Gaz. 65:265, 1918), although he did not typify
that variety, but left Hillebrand s original intent
as applying to Lanai plants.

A number of specimens have been seen,
mostly from Makawao, Olinda, and Olowalu.

(g) var. Remyana B. C. Stone, var. nov.

Arbor, innovationibus dense fulvo hirtellis;
foiiis maturis supra glabris infra glabris costa
media sparse puberulo vel glabrato exceptis,
laminis 5-16 cm longis, 2-6.5 cm lads, suban-
guste ellipticis, apice rotundatis vel emarginatis,
base subcordatis vel cordatis ( vel emarginatis ) ;
inflorescentiis 15-21 (-27-) floriferis; capsulis

410

PACIFIC SCIENCE, Vol. XVII, October 1963

dense fulvo-puberulentibus, c. 20 mm diametro;
endocarpio hirtello.

HOLOTYPE: Hawaii: Without definite locality,
Jules Remy 626 (Paris).

DISTRIBUTION: Known only from Hawaii.

(h) var. racemiflora (Rock) St. John in Lloydia
7:272, 1944.

P. cinerea var. racemiflora Rock, Indig. Trees
Haw. Ids. 241, 1913.

type: Maui: Auahi, on aa lava, at about 1300
feet alt., November 1910, Rock 8676 (Bishop).

DISTRIBUTION: Maui.

SPECIMEN EXAMINED: Maui: Pakiloi forest,
March 1920, Forbes 2082.M. (Bishop).

(i) var. Gaudichaudii (St. John) B. C. Stone
in Pac. Sci. 16:369-371, 1962.

P. Gaudichaudii St. John in Lloydia 7:272,
1944.

brunelia sandivicensis Gaud. nom. nud. in
Bot. Freycinet Voy. in obs. 1826.

TYPE: 'Sandwich Islands,’ Gaudichaud (Paris;
fragment at Geneva ) .

DISTRIBUTION: Hawaii, especially around Ki-
lauea and vicinity.

3. Pglea rnakahae B. C. Stone, sp. nov.

Arbor; innovationibus minute puberulenti-
bus; foliis maturis infra dense cinereis, laminis
olivaceis, trichomiis ad 0.1 mm longis, cum
squamulis interspersis; petiolis 10-22 mm
longis, 1-2 mm crassis, glabrescentibus; laminis
anguste ovatis vel ellipticis, rare late lanceolatis,
coriaceis, ca. 3.5-12 cm longis, 1.5-5 cm latis,
base cuneatis, apice subacutis; inflorescentiis
axillaribus, cymosis unifloris (vel trifloris? ) ,
axibus cinereis, ad 12 mm longis, binodosis;
floris ignotis; sepalis caducis; capsulis 20-24
mm diametro lobis discretis follicularibus glabris
punctatis, apice minute mucronulatis; endocar-
pio glabro.

HOLOTYPE: Oahu: Waianae Mountains, Ma-
kaha Valley, near top of ridge, at about 4000 ft.
alt., 21 October 1954, Gordon A . Pearsall s.n.
( Bishop ) .

A characteristic species closely allied to and
simulating P. cinerea, but with glabrous endo-
carp; similar also to P. cinereops but with glab-

rous ovary. The disk may however be slightly
puberulent.

The cinereous indument is extremely fine,
composed of waxy scales intermingled with hairs
less than 0.1 mm long, obscuring the densely
glandular-punctate surface of the lamina. The
indument tends to persist on the younger peti-
oles and stems, also. The leaf margins are usually
very slightly inrolled; the lateral nerves are
ascending, parallel, and united by a sinuate
marginal nerve 1-8 mm or more in from the
edge.

SECTION Mega carp a stone

4. Pelea Balloui Rock, Indig. Trees Haw. Ids.
228, 1913.

P. ukuleleensis St. John (as to flowering
branch of type sheet), in Lloydia 7:267,
1944. Syn. nov.

The type specimen of P. ukuleleensis is a
mixture of two species: the flowering material
is from a plant of P. Ballouii; fruits of P. clusiae-
folia are mounted on the same sheet. Hence the
species is based on mixed material, of which
both elements already bear valid names, and the
name ukuleleensis is relegated to synonymy.

The type of P. Balloui is: Maui: Haleakala,
trail from Ukulele to Waikamoi Gulch at ca.
5000 ft. alt., 25 October 1910, Rock & von
Temp sky 8609 (Bishop).

The type sheet of P. ukuleleensis is: Maui:
Ukulele, July 1919, Forbes 749.M. (Bishop).

4. Pelea Hiiakae B. C. Stone, sp. nov.

Arbuscula; innovationibus minute puberu-
lentibus glabratis; foliis, petiolis, ramulisque
glabris; petiolis filiformibus 9-21 mm longis, 1
mm crassis; laminis ellipticis base cuneatis vel
subrotundatis apice obtusis vel rotundatis, 3.5-

6.5 cm longis et 2-3.5 cm latis, glabris, costa
media anguste dorsaliter salientis brunneis,
nervis lateralibus paralleliter ascendentibus, cum
nervio sublineare a margine distans 1 mm coali-
tis; inflorescentiis axillaribus 2-3-nodosis, ad

3.5 cm longis in fructu, axibus glabris, pedun-
culis 3-8 mm longis, in crassitudine petiolo
aequis, pedicellis clavoideis, sub calyce expansis;
cymis unifloriferis; floris ignotis; capsulis ca. 25
mm diametro profunde (2A-3A) lobatis lobis

Critical Species of Pelea — Stone

411

Fig. 1. Pelea kauaiensis H. Mann. Holotype specimen, Mann & Brigham, Cornell Univ. Herb. A leaf,
showing pubescence and venation of lower surface; a flower (the entire 1 -flowered inflorescence), showing
the glandular-punctate appearance; floral parts; immature capsule in side view; and reconstructed conjectural
top view of a mature capsule.

rotatis; exocarpiis ad basim sparse puberulenti-
bus glabrescentibus; endocarpio sparse hirtello
trichomiis ad suturas dorsales aggregatis; lobis
diseminatis; seminis ovoideis, ca. 5-6 mm
longis; testa Crustacea nigra nitida.

HOLOTYPE: Oahu: Koolau Range, Kipapa
Gulch, Waipio, on southerly ridge in woods at
1800 ft. alt., 16 October 1932, Edward Y. Ho-
saka 809 ( Bishop ) .

This species seems to be related to Pelea
descendens St. John, which however is charac-
terized by smaller capsules 13-17 mm in diame-
ter, with glabrous exocarp and quite glabrous
endocarp. Pelea Hiiakae is also related to P.
Wawraeana Rock, which differs in that the
pubescence of the endocarp is restricted to the
innermost part of the sutures (and may some-
times be lacking), and in the more compact,
shorter, multi-flowered pubescent inflorescences.

The species is named in honor of Hiiaka,
youngest sister of the goddess Pele, heroine of
the tales of Pele and Hiiaka.

It must be noted that this specific name was
first applied to certain specimens from the island

of Kauai which were thought to represent an
undescribed species ( Stone 1626, Faurie 225,
and Lydgate s.n. ) . Since doubt remains as to the
fruiting state of these plants and their specific
distinctness from Pelea anisata, they must re-
main undescribed. The name Pelea Hiiakae must
now be borne by the Oahu plant described
above, as this is the first publication of the name.
Should the Kauai plants ultimately prove dis-
tinct, another name will be given to the species.

5. Pelea kauaiensis H. Mann Jr. in Proc. Boston
Soc. Nat. Hist. 10:313, 1866.

Fig. 1

A small tree to 5 m high, with opposite
leaves; innovations minutely but densely tawny -
puberulent, but the branches quickly glabrate,
at the 3rd or 4th node quite glabrous. Leaves
distinctly petiolate, petioles 12-30 mm long,
flattened ventrally and narrowly and shallowly
channelled and nearly glabrous or with a few
scattered minute trichomes at the margins; the
rounded dorsal surface especially near the distal

412

PACIFIC SCIENCE, Vol. XVII, October 1963

end hirsute with trichomes 0.8- 1.0 mm long;
but the whole petiole glabrate and ultimately
glabrous. Blades 5-10 cm long, 3-6 cm broad, el-
liptic, usually rounded at base, broadly subacute
at apex or there minutely retuse, ventrally gla-
brous even in bud, densely villous beneath all
over the dorsal surface and densely hirsute along
the costa, with pale stramineous trichomes ca.
1 mm long, the indument more or less perma-
nent; costa raised and rounded dorsally; lateral
nerves 8-14 per side, nearly straight and parallel,
the connecting marginal nerve deeply arched,
with several minor veinlets intervening between
it and the margin; marginal nerve ca. 2-8 mm
from the edge. Inflorescence a highly reduced
cyme with usually 1 or rarely 2 to 3 flowers,
axillary among the leaves; peduncle and pedicel
sparsely and minutely puberulent in flower, later
subglabrate; peduncle reduced to a slight pro-
jection no higher than the 2 or 4 bractlets it
bears, and hidden by them; pedicel ca. 5 mm
long, 0. 3-0.4 mm thick, slightly flaring at calyx
to a breadth of 07-0.9 mm, with bractlets
tightly appressed at base of pedicel on the ob-
solete peduncle; bractlets deltoid, ca. 0. 3-0.4 mm
long, opposite, decussate; pedicels and perianth
segments copiously opaque-punctate. Pistillate
flowers ca. 3 mm high, sepals broadly ovate with
rounded summit and sharp apex, 1.3 mm long,
1.5 mm broad, glabrous except for the minutely
ciliolate margins. Petals glabrous, broadly ovate,
acute, 2-4 mm long, ca. 1.7 mm broad. Stamens
sterile, all shorter than the style, in 2 subequal
quartets, the shorter quartet just under 1 mm
long, the longer quartet bare more than 1 mm
long. Ovary 4-lobed, glabrous, ca. 1 mm high,
1.3 mm broad, on a glabrous shallow somewhat
reddish-speckled disk, the style 0.5 mm high,
stigma 4-lobed, ca. 0.7 mm broad, the lobes dark
purplish and minutely papillate. Staminate flow-
ers unknown. Immature capsule nearly 10 mm
diameter, the lobes (some abortive) still more
or less ascending, 5 mm long, deeply parted
(16-%), sepals caducous; exocarp glabrous, co-
piously glandular-punctate; endocarp glabrous;
lobes 2 -seeded.

TYPE: Kauai: Waimea, 2000-3000 ft. alt.,
Mann & Brigham s.n. (Cornell).

DISTRIBUTION: Kauai.

Only one further collection of this species is
known; it is from the general area of the type

locality (Waimea, Kauai) and was discovered
by Otto Degener. Unfortunately, it is a sterile
specimen.

This species has been consistently misinter-
preted since its first description. Hillebrand
(1888) described under the name Pelea kauai-
ensis a species superficially similar to that which
Mann had actually described, but clearly distinct;
Hillebrand’s material was correctly segregated
by Rock (1918) under the name Pelea recurvata
Rock. In 1897 Heller described Pelea cruciata,
which was however considered by later authors,
such as Rock, to be a synonym of Pelea kauai-
ensis Mann. Such was not the case; Heller’s
species was justly distinguished, and is quite
distinct from both Pelea kauaiensis or Pelea
recurvata. Skottsberg (1944) noted the discrep-
ancies between descriptions and specimens of
these three entities, and published in tabular
form some of their differences. Upon comparing
type specimens of the three species it becomes
quite apparent that they are distinct. The above
extended description of Pelea kauaiensis is based
on the original material, and should serve finally
to clarify this least understood member of a trio
of superficially similar species.

Pelea kauaiensis may be quickly distinguished
from the two other species formerly confused
with it as follows:

1. Endocarp glabrous.

2. Blades permanently pilose beneath;
capsule lobes ascending or rotate;
capsules 10-? (perhaps 20) mm di-
meter; inflorescence reduced, 1-3-

flowered kauaiensis

2. Blades ultimately slightly glabrescent;
capsule lobes recurved; capsules 21-
23 mm diameter; inflorescence stout,
3-9-flowered. recurvata

1. Endocarp pubescent; blades densely pu-
berulent cruciata

Mann’s original description indicated the
name as kavaiensis; since, however, the Latin v
is more commonly spelled out either as a true v
or as u in modern usage, and since the u is es-
sential to the meaning as well as the pronun-
ciation of the name, it is thought best to retain
with Hillebrand the name of the island in its
correct form, as kauaiensis. See art. 73, note 6,

Critical Species of Pelea — Stone

413

of the 1961 International Code of Botanical
Nomenclature.

Little need be added to the descriptions of P.
recurvata and P. cruciata, but it should be men-
tioned that Skottsherg 2863, commented upon
by Skottsberg (1944) as P. cruciata, is actually
a specimen of P. recurvata, as the endocarp is
quite glabrous.

6. Pelea Lakae B. C. Stone, sp. nov.

Arbor ad 5 m alta, ramis divaricatis, trunco
ad 5-10 cm diametro, innovationibus dense hir-
sutulis, trichomiis pallide albobrunneis adpressis
ad 1 mm longis; petiolis ramulisque glabratis;
petiolis 10-65 mm longis, 1.5-3 mm crassis;
laminis ellipticis vel subovatis vel subobovatis,
coriaceis, 4-25 cm longis, 2.5-11 cm lads, base
cuneatis vel obtusis, apice subacutis, obtusis vel
subrotundatis, supra glabris, infra puberulentibus
in pagine juvente glabrescentibus, costa media
salientis stramineis vel brunneis puberulentis
( trichomiis cum squamulis glaucis interspersis ) ,
nerviis lateralibus curvate ascendentibus in ex-
tremis cum nervo marginale arcuato 2-10 mm
intra margine coalitis, reticulis venulosis minori-
bus intervenantibus; inflorescentiis axillaribus
cymosis, cymis ad 2 cm longis axibus puberulen-
tibus glabratis, 2-4-nodosis, 3-5-floriferis, pe-
dunculis 2-7 mm longis pedicellis ca. 3 mm
longis; floris ignotis; bracteis deltoideis extus
puberulentibus; capsulis maturis 22-32 mm di-
ametro profunde (34) lobatis lobis recurvatis
pre- et postdehiscentionem et paullo reflexis,
uni- vel diseminatis, base 8 mm altis, lateraliter
compressis, apicem versus angustatis subfalcatis;
exocarpio minute denseque puberulento non-
ruguloso; endocarpio glabro in suturam basem
centralem excepto, trichomiis paucis aggregatis;
seminis ovoideis, 5-6 mm longis, testa Crustacea
nigra nitida.

HOLOTYPE: Oahu: Koolau Range, Pupukea, at
about 2000 ft. alt., at edge of small boggy region,
17 September 1961, B. C. Stone & A. K. Chock
3633 (Bishop; duplicates to be distributed).

A species reminiscent of Pelea recurvata of
Kauai, but with a puberulent rather than gla-
brous ovary, and a few hairs clustered near the
innermost angles of each suture on the endocarp.

The specific name honors Laka, goddess of
the hula.

7. Pelea Lohiauana B. C. Stone, sp. nov.

Fig. 2

Arbuscula? ramis virgatis elongatis, innova-
tionibus ramulis petiolisque dense tomentellis,
trichomiis stramineis val pallide brunneis ca.
0.8-0.9 mm longis; foliis maturis glabratis, ju-
venilis tomentellis; petiolis 20-30 mm longis,
glabrescentibus, supra canaliculatis glabris; lam-
inis ellipticis coriaceis 7-15 cm longis 3.5-7
cm latis base apiceque rotundatis vel paullo
emarginatis, marginibus paullo revolutis, costa
media prominente, nervis lateralis 5-9 (-12)
per latere unico, cum nervo marginale continue
vel paullo arcuato intra margine 1-2 (-5) mm
coalitis; inflorescentiis cymosis (3-) 5-7-flori-
feris longe-pedunculatis tomentellis, pedunculis
12-38 mm longis ad nodo primo, 1-2-nodosis,
bracteis oppositis lanceolatis 1.5-2. 5 mm longis
concavis; axibus majoribus 5-14 mm longis;
pedicellis 2-5 mm longis; floris masculis pro
genere magnis sepalis ad 5.5-6 mm longis et
ca. 4.5 mm latis, extus hirsutulo-ciliolulatis intus

Fig. 2. Pelea Lohiauana Stone. Staminate flowers
and floral parts of Degener 23984.

414

PACIFIC SCIENCE, Vol. XVII, October 1963

glabris, deltoideis, 7-9-nervatis; petalis toto gla-
bris sparse obscure punctatis ovato-lanceolatis
ca. 11 mm longis et 5 mm latis apice intus un-
guiculatis; staminibus fertilibus 8, (4 ad 10 mm
longis, 4 ad 7.5 mm longis, antheris 1.7— 1.8
mm longis, filamentis ligulatis ca. 1 mm latis);
ovario glabro ca. 2 mm alto, disco glabro ca.
1 mm alto; stylo ca. 2 mm longo, stigma ca. 1.4
mm lato lobis rufidis minute papillatis cruciatis;
floris foemeneis minoris sepalis ca. 3-5 mm
longis et 3 mm latis, petalis ca. 7-8 mm longis
et 3.5 mm latis, staminibus sterilibus ca. 2 mm
longis subaequitantibus; ovario discoque glabro,
sytlo glabro ca. 4 mm longo; capsulis quadrilo-
batis, glabris, ad 20-25 mm diametro, endo-
carpio glabro.

HOLOTYPE: Kauai: Kokee, August 1924,
Degener 8586 (Bishop; duplicates at Cath, NY).

DISTRIBUTION: Kauai, in the Waimea high-
lands.

SPECIMENS EXAMINED: Kauai: Near Lehua-
makanoe Bog, 18 September 1955, Degeners,
Hansen & Hanner 23984 (Bishop; staminate
fls. ). Kaholuamano, October 1895, Heller 2869
(formerly determined as P. resiniflora ) (GH,
edinb, US). Halemano, 14 February 1909, Rock
2294 (A). Waialeale, October 1911, Rock (a).
Kaholuamano, September 1909, Rock 5551 (A,
Bishop), 5552 (gh, Bishop).

This is by no means a complete citation of
specimens; more are extant, particularly in the
Bishop Museum, but many lack data, particularly
localities and collectors’ numbers. Most of these
specimens had been determined as Pelea macro-
pus Hillebr., which however is quite distinct in
having puberulent ovaries ( and hence capsules )
and clavoid pedicels. Because of this interpreta-
tion, this species has consistently been mistaken
for P. macropus, and this error has tended to
obscure knowledge of both of them. The gla-
brous endocarp and rotate capsular lobes place
P. Lohiauana in a group with P. oblanceolata
St. John (of Hawaii), P. manukaensis St. John
(of Hawaii), and P. pseudoanisata Rock (of
Hawaii ) .

The specific name honors Prince Lohiau of
Kauai, hero of the tales of Pele and Hiiaka, and
one of the memorable characters of Hawaiian
mythology.

8. Pelea macropus Hillebrand, FI. Haw. Ids. 65,

1888.

P. acutivalvata Leveille in Fedde Repert. Sp.
Nov. 10:153, 1911.

Innovations minutely and closely pale cinere-
ous-puberulent, hairs appressed, ca. 0.2 mm
long; branchlets at youngest nodes cinereous but
soon glabrate; leaves glabrous ventrally, at first
minutely cinereous dorsally (mostly on the
costa ) , but soon glabrate, the older leaves mostly
quite glabrous; petioles soon glabrate. Leaves
opposite, the petioles mostly 9-15 mm long;
blades thinly coriaceous, rounded or emarginate
at apex, contracted and subcuneate at base (the
margins sometimes slightly revolute); costa
ventrally pale, shallow-sulcate; dorsally pale and
prominent with a rather sparse and minute
ephemeral puberulence; lateral nerves at right
angles to costa or slightly ascending, about 7-9
per side, conspicuous, prominulent on both sur-
faces, connected at a distance of 2-10 mm from
the margin by a moderately to deeply arcuate
marginal nerve, with several sets of intervening
meshes. Inflorescences small, axillary or ramu-
line, mostly 1-3 -flowered, the peduncle very
short, about 1-5 mm long, bibracteolate at apex,
puberulent, the bractlets deltoid, rounded, 0.5
mm long, puberulent; axes minutely cinereous,
short ( to 5 mm ) in flower, somewhat elongated
in fruit; pedicels 2-5 mm long, flared at base
of calyx and clavoid, bibracteolate, cinereous;
pistillate flowers with broadly deltoid sepals
about 1.5 mm long and 1.7 mm broad, minutely
puberulent externally; petals about 2.5 mm long
and 1.7 mm broad, with a faint close puberu-
lence of few minute hairs along the midregion,
glabrous within; rudimentary stamens glabrous,
less than 1 mm long; disk reddish, glabrous,
shallow, ca. 1.7 mm broad; ovary puberulent,
ca. 1 mm high and 1 mm broad; style glabrous,
ca. 1.6 — 1.7 mm long (including stigma), stigma
lobes red, each 0.25 mm long, minutely papillate,
obovoid. Capsules 25-35 mm diameter, about
9 mm high, sparsely appressed-puberulent (es-
pecially near the base), the lobes deeply separated
( Y<$ ) , somewhat acuminate and laterally com-
pressed, often unequally developed; endocarp
glabrous, cartilaginous; seeds 5-6 mm long,
ovoid, the testa crustaceous, blackish.

Critical Species of Pelea— -Stone

415

HOLOTYPE: Kauai: Waimea, Knuds en 189
(Berlin; now destroyed).

DISTRIBUTION: Kauai.

SPECIMENS EXAMINED: Kauai: "Robinson’s
summer house,” February 1910, Faurie 112 (type
of P. acutivalvata; type at Edinburgh; isotype
at A). Waimea, Kaholuamano, October 1895,
Heller 2870 (bm, k. Paris, us).

The description given above sufficiently dis-
tinguishes Pelea macropus from the newly de-
scribed P. Lohiauana.

SECTION Cubicarpa stone

9- Pelea peduncularis Leveille in Fedde Repert.
Sp. Nov. 10:443, 1912.

Fig. 3

P. sandwicensis var. /3 sensu Hillebrand, FI.
Haw. Ids. 66, 1888 (not P. sandwicensis
[Hook. f. & Arn.] Gray).

P. sandwicensis var. macrocarpa Hillebr. ex
Rock in Bot. Gaz. 65:265, 1918.

P. nodosa Leveille in Fedde Repert. Sp. Nov.
10:443, 1912.

P. grandipetala Leveille l.c. (in part; specimen
typicum mixtum; in part P. honoluluensis) .
Syn. nov.

P. singuliflora Leveille l.c. (in part; see P.
Wawraeana ) . Syn. nov.

P. Rockii St. John in Lloydia 7:271, 1944.
Syn. nov.

A small or sometimes shrubby tree, the in-
novations finely scurfy and somewhat sparsely
cinereous-puberulent; trichomes soon caducous,
the mature branchlets and leaves glabrous, or
the petioles sometimes puberulent then glabrate.
Leaves petiolate, opposite; petioles 13-40 mm
long, lenticellate in age especially at the ex-
tremities; blades coriaceous, elliptic, rounded to
slightly emarginate at apex and at base, mostly
3-18 cm long and 2-11 cm broad (commonly
5-9 X 3-6 cm ) , darker above, the margin near
the base usually tightly revolute; midrib above
sulcate, raised beneath, often reddish or purplish;
major lateral nerves mostly about 7-10 per side,
united distally by a slightly sinuate marginal
nerve close to (about 1-5 mm from) the mar-
gin. Inflorescence a pedunculate usually 5—2 1-
flowered minutely cinereous glabrate cyme often

FIG. 3. Pelea peduncularis Levi. Inflorescences; at
left, a long inflorescence with a dehisced capsule, from
Faurie 189 (isotype); at right, inflorescences in flower
and in bud from Remy 621, with enlarged view of
one staminate flower.

longer than the adjacent petiole, the peduncle
up to 4 cm long and with as many as 8 nodes,
but usually about 3-4-nodose, stout ( 1-2 mm
thick), the pedicels short and stout, 3-4 mm
long, the bractlets lanceolate, the upper ones
deltoid, minutely ciliolulate and sparsely pu-
berulent, 1-3 mm long; pistillate flowers with
sparsely scurfy-puberulent deltoid-ovate sepals
about 2 mm long and 1.8 mm broad, lanceolate-
ovate glabrous punctate petals about 5 mm long,
reduced sterile stamens barely higher than the
glabrous 4-lobed ovary, the disk glabrous, the
style filiform reddish about 2 mm long, the
stigmas 0.5 mm long, slightly clavate and mi-
nutely papillate, rotate. Staminate flowers similar
but larger, the petals about 7 mm long, the
gynoecium much reduced, the fertile stamens
longer, in 2 subequal quartets, the longer sta-
mens about 8 mm long. Capsules mostly about

416

PACIFIC SCIENCE, Vol. XVII, October 1963

20—25 mm broad and 10 mm high, quadrate or
slightly lobed ( less than or barely halfway ) , the
exocarp glabrous, punctate, mostly dark green or
tinged with dull reddish-purple; endocarp pale
and glabrous; seeds mostly 2 per cell; carpels
after dehiscence reflexed in age, the axis per-
sistent only at the extreme base.

TYPE: Oahu: Koolau Range, Kalihi, October
1909, Faurie 189 (Edinburgh; isotype at Paris).

DISTRIBUTION: Restricted to Oahu; found on
both the Waianae and Koolau mountain ranges.

This common Oahu species has been misin-
terpreted since it was first described. This is not
surprising, however, since the confusion involved
with the identity of the true Pelea sandwicensis
(H. & A.) Gray had obscured the concept
greatly, and since Leveille himself did nothing
to clarify the species, and presumably did not
realize that the other "species” so briefly and
inadequately described by him were either the
same or mixtures (the type specimens often
being composed of branchlets clearly from two
or three different species). In addition, some
rather clearly defined varieties of the species
occur in particular areas of Oahu, and some of
these have been described as distinct species.
Dozens of collections from various localities both
in the eastern and western mountain ranges are
known. A few representative specimens are cited
here; full citation of specimens will be presented
in the forthcoming monographic treatment.

SPECIMENS EXAMINED: Oahu: Koolau Range:
Pupukea, January 192 7, MacDaniels 547
(Bishop); February 1928, Degen er & Shear
8580 (NY, Catholic, US). Kalihi, October 1909,
Faurie 1 90 ( bm ) . Konahuanui, Famie 194 ( bm,
isolectotype of Pelea singuliflora ) . Manoa-Pauoa
hills, April 1861, Hillebrand 251 (Kew). Nuu-

anu, May 1861, Hillebrand 238 (Kew). Nuuanu-
Kalihi ridge, August 1922, Skottsberg 173
(Goth, Kew); Niu, Hillebrand 1797 (US, isotype
of Pelea molokaiensis (3 var. Hbd. ) . Waianae
Mountains: Puu Kaala, Mann & Brigham 600
(Cornell, Kew). Makaha Valley, Feb. 1909,
Forbes (Bishop). Puu Kanehoa, May I960,
Stone 3268 (Bishop, Goth, Kew, Leiden, us).
Without locality: Mann & Brigham 208, in part
(Cornell). Remy 621 (Paris, 3 sheets).

(a) var. peduncularis
Fig. 3

Found mostly along the southern stretches of
the Koolau Range, from Punaluu down through
Waialae and Niu valleys. The typical variety,
easily confused with Pelea oahuensis Levi, and
sometimes with Pelea Wawraeana Rock, but
differing in many respects from both; from the
former in the larger and flatter capsules, the
more ample and stouter cymes, and generally
larger vegetative parts, and the larger reddish
(not yellowish) flowers; and from the latter in
the larger capsules with glabrous endocarp, and
mostly glabrous petioles and branchlets as well
as cymes.

(b) var. niuensis (St. John) B. C. Stone, comb,
nov.

Pelea niuensis St. John in Lloydia 7:272,
1944.

Branchlets glabrate; petioles glabrate; blades
narrowly elliptic or oblong-elliptic, glabrous,
6-12 cm long and 2-5 cm broad; inflorescences
commonly 7-9-flower ed, glabrous; capsules
nearly quadrate when fresh, slightly lobate when
dry, 21-25 mm broad and about half as high,
glabrous; endocarp glabrous.

KEY TO VARIETIES OF Pelea peduncularis

1. Capsules slightly lobed (up to halfway), green, usually dull, sometimes reddish; blades

elliptic to elliptico-oblong.

2. Leaves opposite.

3. Blades very narrowly oblong-elliptic var. niuensis

3. Blades elliptic to broadly oblong-elliptic.

4. Cymes mostly 7-21 -flowered var. peduncularis

4. Cymes mostly 3-7-flowered. . var. pauciflora

2. Leaves whorled in fours - var. paloloensis

1. Capsules unlobed, quadrate, dark glossy green. var. quadrata

Critical Species of Pelea — Stone

417

HOLOTYPE: Oahu: Koolau Range; Niu, Oc-
tober 1940, St. John 20111 (Bishop).

DISTRIBUTION: Endemic to the southeastern-
most part of the Koolau Range.

This variety differs from the typical P. pedun-
cularis only in the much narrower leaves and
perhaps slightly fewer-flowered cymes.

SPECIMEN EXAMINED: Oahu: Northeast slope
of Puu Kumakalii, April 1936, Degener, Tam,
Takamoto & Martinez 10579 (Bishop, Catholic,
NY).

Pelea Rockii var. pauciflora St. John in
Lloydia 7:271, 1944.

Cymes mostly 3-7-flowered.

HOLOTYPE: Oahu: Koolau Range; Kaukona-
hua Gulch, Wahiawa, May 1909, Rock 3046
(Bishop).

DISTRIBUTION: Known only from the type
locality.

A poorly defined variety which may be only

an anomalous form of the species.

(d) var. paloloensis (St. John) B. C. Stone,
comb. nov.

Pelea paloloensis St. John in Lloydia 7:271,
1944.

Leaves whorled in fours at the nodes.
HOLOTYPE: Oahu: Koolau Range; Palolo,
Waialae-iki, Jan. 1911 , Forbes 2404.0. (Bishop).

DISTRIBUTION: Known only from the type
locality.

The variety, which probably is nothing more
than an anomalous form, is an example of the
occasional exception to the common taxonomic
criterion for distinguishing species which are
members of the section Pelea, with ordinarily
whorled leaves, and the other sections, with
generally opposite leaves.

(e) var. quadrata B. C. Stone, var. nov.

Fig. 4

Capsulis elobatis quadratis fere ca. 23 mm
latis.

HOLOTYPE: Oahu: Waianae Mountains; Puu
Hapapa, August 1932, Degener 8521 (Bishop).

DISTRIBUTION: Known from the northern
end of both ranges of Oahu.

A characteristic variety, found both in the
northern Waianae mountains and in the north-
ern Koolau Range (as for instance around Pu-
pukea), sometimes occurring in company with
the typical variety.

10. Pelea olowaluensis St. John in Lloydia 7:2 66.
1944.

A tree up to 3 m high, the branchlets at first
puberulent; leaves opposite; petioles glabrous;
blades 2.8-8 cm long, oval, subcoriaceous, gla-
brous; cymes mostly 3 -flowered, glabrous, the
peduncle ca. 9 mm long; capsules 16-20 mm
broad, flattened, lobed halfway or less, glabrous;
endocarp glabrous.

HOLOTYPE: Maui: Olowalu Valley, on the cen-
tral ridge, May 1920, Forbes 2326.M. (Bishop).

DISTRIBUTION: West and East Maui.

SPECIMEN EXAMINED: West Maui: Mauna
Huuma, May 1910, Forbes & Cooke 22.M.
(Bishop) .

This species is closely allied to Pelea pedun -
cularis of Oahu and falls properly into section
Cubicarpa . The characteristically revolute basal
leaf-margins are especially noticeable. When
fresh the capsules might perhaps be unlobed or
barely notched. The disjunction of the two col-
lections would imply that the species is, or was,
fairly widespread on Maui.

Fig. 4. Pelea peduncularis var. quadrata Stone.
Three views of a capsule in the fresh state, from
Stone 2832 ( Pupukea, Oahu ) .

418

PACIFIC SCIENCE, VoL XVII, October 1963

Fig. 5. Pelea oahuensis Levi, emend. Stone. Neotype, Stone 3282. Above, capsules and pistillate flowers;
below, staminate flowers.

11. Pelea oahuensis Leveille in Fedde, Repert.

Sp. Nov. 10:442, 1912, emend. B. C.

Stone.

Figs. 5, 6

A small to middle-size tree, pungent of anise,
with glabrous reddish branchlets and opposite
leaves; innovations scurfy, the waxy white
ephemeral scales mostly concealing an extremely
minute sparse puberulence of white hairs usually
less than 0.2 mm long, ephemeral also, the older
leaves and branchlets glabrous except for a few
rare hairs persisting in the axils or on the dorsal
midrib near the base; leaves petiolate, the peti-
oles mostly 6-20 mm long, reddish in age, gla-
brous, flattened ventrally, lenticellate; blades thin
coriaceous, mostly (3) -4-12 cm long and 2.5-7
cm broad, glabrous and moderately veiny, darker
above, moderately shining, mostly elliptico-
obovate, rounded to slightly emarginate at both
ends, the costa pale and shallowly canaliculate
toward the base ventrally, beyond prominulous,
beneath raised and reddish; lateral main nerves
slightly curvate-ascendent, joined marginally by
a lightly arched or nearly straight nerve about
1-5 (-9) mm from the edge. Inflorescence
cymose, mostly 3-5 -flowered (rarely with 7

flowers), rather short (less than 2 cm long),
on short glabrous greenish sometimes scurfy
peduncles 3-4 mm long, bibracteolate at apex
(the bracteoles minute, deltoid, minutely pu-
berulent or glabrous ) ; axes about 2-3 mm long,
similar to peduncles; pedicels about 3 mm long,
bibracteolate near the middle or above; flowers
functionally staminate or functionally pistillate,
or perfect, frequently with both staminate and
perfect flowers on the same plant; pistillate
flowers greenish, petals yellowish-green; sepals
deltoid-ovate, about 1.8 X 1.8 mm, the margins
minutely ciliolulate, otherwise glabrous; petals
4.5-5. 5 mm long and 2 mm broad, glabrous;
stamens rudimentary, about 1 mm long (some-
times in two slightly unequal quartets); ovary
and disk glabrous, punctate, the disk about 2.8
mm broad and nearly 1 mm high, the ovary
about 2 mm broad and 1 mm high; style slender,
glabrous, about 2 mm long including the 4-lobed
stigma; staminate and perfect flowers similar
but the stamens greatly elongated, the longer
quartet as long as the petals; petals slightly
longer than those in pistillate flowers, and the
ovary and disk greatly reduced, the style short,
only 1 mm long. Capsules subcuboid, 10-14 mm
broad, about 9 mm high, unlobed or nearly so.

Critical Species of Pelea— Stone

419

Fig. 6. Pelea oahuensis Levi, emend. Stone. Habit of fruiting specimen, from Stone 2805 ( Pupukea,
Oahu). At left, perfect flower from Lane 243 (Pupukea). At right, above, capsules and seed from Stone 3200
(Kaala, Oahu). At right, below, capsule and pistillate flower from Stone 2826 (Pupukea).

420

PACIFIC SCIENCE, Vol. XVII, October 1963

quite glabrous, often pale, greenish, or after
dehiscence brown, dehiscent along the upper
and lateral sutures; endocarp glabrous, pale, thin,
late detaching; seeds often only one per carpel;
testa crustaceous, shining black.

TYPE: Leveille cites three specimens, all from
Kalihi, Oahu, collected by Faurie: numbers 11,
217 , and 217 bis. No type is designated. A search
was made for these specimens in the herbaria of
the Bishop Museum, the British Museum, the
Royal Botanic Gardens at Kew and at Edin-
burgh, the Museum d’Histoire Naturelle, Paris,
Gothenburg Botanical Garden, U. S. National
Museum, Gray Herbarium, Arnold Arboretum,
Cornell University, etc., but without avail. Many
of Leveille s "types” are at Edinburgh or at the
British Museum, but none of the cited specimens
can be located. The original description is so
short and noncommittal that alone it is meaning-
less: "Affinis praecedenti a quo tantum differt
foliis longius 10-25 mm petiolatis, oppositis;
petiolo nec rugoso, nec vulnerato, corymbis 2-5
floris, pedicellis bis bibracteolatis, apice incras-
satis.”

Rock (1914) reduced Pelea oahuensis to P.
molokaiensis Hbd. (2 var. Hbd., on the basis of
Paurie 217, which perhaps was at the Berlin
Museum. The preceding species to which Le-
veille refers in the above description is his own
Pelea waianaiensis, which is more fully de-
scribed. It is based on a single number ( Faurie
215, isotype at Bishop), and is apparently a
form of Pelea peduncularis Levi. The reference,
however, is helpful in interpreting the present
species, as the two are similar in appearance.

In the absence of any type material, it seems
necessary to designate a neotype, which would
of course be replaced by original material should
any turn up. The species, as interpreted here,
is a very distinct and common one on Oahu,
and is the Oahuan counterpart of the "moki-
hana” or Pelea anisata Mann of Kauai; the same
strong anisate odor is present in all parts, the
flowers and fruits are very similar, and the
specific differences are mainly in characters of
the leaves and habit. There are a great many
collections of this species, most of which have
been determined as Pelea Wawreana Rock,
which is, however, a very different species of

Sect. Megacarpa. The following collection may
be selected as neotype:

NEOTYPE: Oahu: Waianae Mountains; Kunia
trail, 26 March I960, B. C. Stone & G. Pearsall
3282 (Bishop; duplicates to be distributed).

DISTRIBUTION: Endemic to Oahu. This is a
very common species on both the Waianae and
Koolau ranges. It may be identified in the field
by its strong anise fragrance, very short essen-
tially glabrous 3-5 -flowered cymes with small
greenish flowers, and small cuboid green or
whitish-green capsules. In the Waianae Moun-
tains it occurs in company with Pelea peduncu-
laris, P. kaalaensis , P. clusiaefolia var. crassiloba,
and P. elliptica.

REFERENCES

Heller, A. A. 1897. Observations on the ferns
and flowering plants of the Hawaiian Islands.
Minnesota Bot. Studies 9:760-922, pis. 42-69.

Hillebrand, W. 1888. Flora of the Hawaiian
Islands. Heidelberg, London, & New York,
i-xciv, 1-673.

Rock, Joseph F. 1914. Revisio plantarum Ha-
waiiensium a Leveille descriptarum. Repert.
Sp. Nov. ed. Fedde 13:352-361.

— — — 1918. Pelea and Platydesma. Bot. Gaz.
65:261-267.

Skottsberg, C. 1944. Vascular plants from the
Hawaiian Islands, IV. Phanerogams collected
during the Hawaiian Bog Survey 1938. Acta
Horti Gotob. 15:275-531.

Stone, B. C. 1962a. Studies in the Hawaiian
Rutaceae, I. Taxonomic and nomenclatural
notes on Platydesma (Hawaii) and Melicope
(Solomon Islands). Madrono 16:161-166.

1962 A Studies in the Hawaiian Ruta-
ceae, II. On the identity of Pelea sandwicensis.
Pacif. Sci. 16:366-373.

1962c Studies in the Hawaiian Ruta-
ceae, III. On the New Caledonian species of
Pelea, and a misunderstood species of Platy-
desma. Adansonia (Paris) : I, 2:94-99.

l%2d. Rutaceae: Genus Pelea. In: O.

Degener, Flora Hawaiiensis. Honolulu (pri-
vately printed).

A Prior Name for the Hawaiian Gouldia terminalis (Rubiaceae)

Robert L. Wilbur1

Among the most frequently encountered woody
plants in the wetter, forested portions of the
Hawaiian Islands are members of the extremely
variable genus Gouldia. Fosberg (1937) pre-
sented the results of his detailed study of this
baffling genus and concluded that the variability
could be properly categorized in not less than
three species composed of more than 90 varieties
and forms. However, even this number of for-
mally named taxa failed adequately to represent
the variability, for hybridization was so rampant
that at that time more than 50 hybrids were
also recognized and characterized. It is therefore
not surprising that Gouldia has acquired a repu-
tation, among botanists working on Hawaiian
plants, not unlike that of Crataegus and Rubus
in the eastern United States. Like those genera,
it is naturally felt that its taxonomy can now be
handled only by a specialist. The present note,
written far from Hawaii, is therefore merely
concerned with the nomenclature of the most
widespread and variable species of this endemic
genus.2

Although members of the genus undoubtedly
must have been collected by botanists on several
expeditions prior to that of the "Rurik” led by
Kotzebue, the first description of a species is
apparently the detailed analysis provided by
Chamisso and Schlechtendal (1829) of their
"Kaduae affinis.” Chamisso was the botanist on
Kotzebue’s voyage, and the original collection
apparently was made on the slopes of the Koolau
range of Oahu. Their account, as pointed out by
Heller ( 1897), Fosberg (1937:4,26) and Bul-

1 Department of Botany, Duke University, Durham,
North Carolina. Grateful acknowledgment is made to
the National Science Foundation for a grant of re-
search funds to Duke University (NSF-Grant 18799)
which made the present study possible.

Manuscript received March 29, 1962.

2 Dr. F. R. Fosberg’s thoughtful advice is here
acknowledged with appreciation, but this does not
imply that he is necessarily convinced of the change
proposed in this paper.

lock (1958), did not result in a published bi-
nomial at that time as was inferred by A. Gray
(I860) and the Index Kewensis (1895). In
spite of the unusually detailed analysis of the
sixth species appearing in their newly described
genus Kadua , Chamisso and Schlechtendal failed
to provide a binomial for this plant; they merely
indicated its close affinities to Kadua , from
whose species it differed in its indehiscent fruit
and toothed stipules.

The following year DeCandolle (1830) listed
each of Chamisso and Schlechtendal’s species
and condensed the original detailed accounts of
each into but a few lines. Their ”6. Kaduae
affinis ” appeared in DeCandolle’s Prodromus
in the form quoted below:

6. K? AFFINIS (Cham, et Schlecht. 1. c. p.
164.) ramis tetragonis transversim rugosis, foliis
elliptico-lanceolatis acutis basi obtusis breve
petiolatis, stipulis membranaceis utrinque sub-
dentatis deciduis, cyma thyrsoidea terminali,
drupa subglobosa, limbo calycis obliterato infra
apicem coronata, indehiscente. [Woody tree or
shrub} in insula O-Wahu. Flor. ignoti.

Fosberg (1936:4) dismissed DeCandolle’s
publication as a nomenclatural source in the
statement quoted below:

DeCandolle, in 1830, a year after the publication
of Chamisso and Schlechtendal’s work, credited
the latter with a f Kadua - affinis Cham, and
Schlecht.,’ appending a description which is an
obvious condensation of the description pub-
lished by Chamisso and Schlechtendal. DeCan-
dolle’s entire treatment of Kadua is based
directly on the original treatment of the genus
by Chamisso and Schlechtendal, with the same
species arranged in the same order and with
descriptions which are identical but somewhat
condensed ... It is obvious that f Kadua affinis’
is the result of a misinterpretation of the intent
of the original authors of the genus Kadua, as
DeCandolle added nothing to the descriptions
and no discussion. Therefore, it is evident that

421

422

PACIFIC SCIENCE, Vol. XVII, October 1963

DeCandolle did not have any intention of mak-
ing a new species 'Kadua affinis / making it pos-
sible to judge it on the basis of Chamisso and
Schlechtendal’s original intent. The Cambridge
Rules of 1930, Article 68, state that terms
which are merely words, not intended as names,
should be rejected; thus, fortunately, a means is
provided for disposing of this meaningless name.

It would seem, however, that Fosberg’s stated
reasons are not sufficient justification for reject-
ing the name. Chamisso and Schlechtendal con-
cluded in their final sentence concerning this
species that "the whole internal structure of the
fruit agrees therefore with Kadua.”3 Although
they did not publish a binomial, it certainly
would appear that DeCandolle did. Chamisso
and Schlechtendal expressed their doubts as to
generic position in one fashion and did not
provide the taxon with a name; DeCandolle was
no more certain as to the generic position of
the plants than the original authors, but he ex-
pressed his doubts as to generic position in a
different manner and did provide a binomial.
He was perfectly free to utilize the epithet
affinis, which he did. The question mark follow-
ing the abbreviation of the generic name merely
indicated that he too was uncertain that the
clearly described species actually was congeneric.
Chamisso and Schlechtendal more than ade-
quately described the species but failed to pro-
vide a binomial; DeCandolle provided a binomial
even though admitting that the plant might
eventually prove not to be a congener of the
other five species. Although DeCandolle "added
nothing to the description and no discussion,”
he did provide the binomial that the previous
detailed description of the species lacked. Fos-
berg’s conclusion that "it is evident that De-
Candolle did not have any intention of making
a new species Kadua affinis ” seems unwarranted.
DeCandolle provided a binomial, and there is
certainly no evidence that his intentions were at
variance with his accomplishment.

In the Pro dr omus DeCandolle did not cus-
tomarily place any authority at all after names
that he was proposing. Species and new combi-
nations being there published by him were not
followed by any authority or reference. Since

3 "Convenit ergo omnis interna fructus fabrica cum
Kadua.”

abbreviations for Chamisso and Schlechtendal.
together with a reference to the place of publi-
cation, were all included parenthetically by De-
Candolle after K? affinis , it might be argued
that he was merely accepting Chamisso and
Schlechtendal’s treatment and had no intention
of publishing a new name. This is impossible
to prove one way or the other, but certainly
Article 34, Note 2, of the International Code of
Botanical Nomenclature (1961) was never meant
to be applied in such a case, as is shown by the
examples provided. Furthermore, DeCandolle
employed the parenthetical citation to convey a
variety of information in addition to his custom-
ary indication of author and place of publication.
Other information characteristically conveyed
parenthetically by DeCandolle concerned authors
who had merely indicated a species as new on a
herbarium specimen (e.g., Eryngium Haenkei
Presl ex DC, Prodr. 4:94. 1830); or mention
that a description of a new species has been
provided in a letter by another (e.g., Eryngium
prostratum Nutt, ex DC., Prodr. 4:92. 1830);
or that a drawing had been seen in the herbar-
ium (e.g., Cornus disciflora Moc. & Sesse ex
DC., Prodr. 4:273. 1830). Certainly, since De-
Candolle had not seen specimens of this Ha-
waiian species, it was in accord with his rather
liberal usage of parenthetical citation to indicate
the source of his information. It is certainly not
necessary to conclude that DeCandolle was not
aware that Chamisso and Schlechtendal actually
had not published a binomial. The conclusion
therefore seems to me inescapable that DeCan-
dolle did originate a binomial that was the first
published for this species.

A combination based upon DeCandolle’s bi-
nomial, therefore, appears necessary for this
extremely common Hawaiian species. Combina-
tions for the multitudinous varieties and forms
within this species, of which more than 85 were
originally proposed by Fosberg, are not provided
here; it would seem most undesirable for anyone
not thoroughly familiar with these variants to
make the numerous transfers apparently re-
quired. It seems probable that a restudy of the
problem with the benefit of the numerous col- j
lections made during the past quarter of a

number of taxa worthy of recognition. However,

Gouldia terminalis — WlLBUR

423

Skottsberg (1944 a, b) has expressed the opinion
that there are at least several times as many
species as accepted by Fosberg, and he has pro-
vided binomials for a number of them by ele-
vating certain of Fosberg’s varieties and forms
in addition to accepting some binomials of
earlier authors. And, recently, the Degeners
(1961) have also expressed a different view
and, in so doing, provided 22 new combinations
largely as a result of elevating Fosberg’s taxa to
the next highest rank. Therefore, it would not
seem desirable to make wholesale transfers at
the present time.

Gouldia affinis (DC.) comb. nov.

"Kaduae affinis” Cham. & Schlecht., Linnaea 4:
164. 1829.

Kadua affinis [K? affinis] DC., Prodr. 4:431.
1830.

Petesia terminalis Hook. & Arn., Bot. Beechy’s
Voy. 85. 1832.

Petesia coriacea Hook. & Arn., Bot. Beechy’s
Voy. 85. 1832.

Gouldia sandwicensis A. Gray, Proc. Am. Acad.

4:310. I860, nom. illegit. Art. 63 and 67.
Gouldia terminalis (Hook. & Arn.) Hbd., FI.
Haw. Is. 168. 1888.

REFERENCES

Bullock, A. A. 1958. Nomenclatural notes:
VI. Type species of some generic names. Kew
Bull. 13:98 {Gouldia'].

Chamisso, L. C. A., von, and D. von Schlech-
TENDAL. 1829. De plantis in expeditione
speculatoria Romanzoffiana observatis. Lin-
naea 4:157-165 {Kadua] .

DeCandolle, A. P. 1830. Prodromus systematis
naturalis regni vegetabilis 4:430 + 431.
[Kadua].

Degener, Otto, and Isa Degener. 1961.
Gouldia in Hawaii. Phytologia 7:465-467.

Fosberg, F. Raymond. 1937. The genus Goul-
dia (Rubiaceae). Bishop Mus. Bull. 147:1-82.

Gray, A. 1860. Notes upon some Rubiaceae,
collected in the South Sea Exploring Expedi-
tion under Captain Wilkes. Proc. Am Acad.
4:310 + 311. [Gouldia].

Heller, A. A. 1897. Observations on the ferns
and flowering plants of the Hawaiian Islands.
Minnesota Bot. Stud. 1:896-899. [Gouldia].

Index Kewensis. 1895. 2:1.

International Code of Botanical No-
menclature. 1961. Utrecht. 372 pp.

Skottsberg, C. 1944^. Vascular plants from the
Hawaiian Islands. Acta Horti Gothoburg. 15:
466, 517. [Gouldia].

— 1944 A On the flower dimorphism in

Hawaiian Rubiaceae. Arkiv for Botanik 31A
(4): 11-14. [Gouldia].

Population Dynamics in a Sublittoral Epifauna

Willis E. Pequegnat1

At PRESENT we have little evidence that suc-
cessional changes following predictable patterns
occur among epifaunal communities living on
natural rock-reefs in the shallow sublittoral of
the open ocean. If this phenomenon does occur
here, it can be detected by sustained observa-
tions at a single study site. In the period from
1957 to I960 I conducted an intensive study of
the epifaunas of two submarine hogbacks lo-
cated at different depths off the coast of Corona
del Mar, California. Some observations made
during this period indicate that both gradual
and disruptive changes do occur in these epi-
faunal communities. Attention is called in this
paper to an abrupt change in population density
of a predominant species that occurred on part
of the shallower reef, and to the widespread
biotal adjustments that ensued.

The epifauna of the deeper reef (9.5-18.5 m)
remained quite stable throughout the study pe-
riod (Fig. 1, Reef 500). Nevertheless, this is
a dynamic stability with an appreciable flow of
individuals through the community structure.
For example, in the rock oyster, Chama pellucida,
which is dominant on the upper part of this
reef, natality and growth counterbalance mor-
tality from all causes and such erosive factors
as Lithophaga burrowings, resulting in a stable
community as determined by consecutive sam-
plings over a prolonged period. Some seasonal
population fluctuations were observed, especially
among sponges and ectoprocts, but I have not
detected there any changes that can be construed
as serial stages moving toward a climax differing
from any extant community.

But this stability appears not to be charac-
teristic of a significant part of the epifauna that

1 Formerly at National Science Foundation, Wash-
ington 25, D. C. Present address: Department of
Oceanography and Meteorology, Texas Agricultural
and Mechanical University, College Station, Texas.
Manuscript received May 2, 1962.

was present on the shallower reef (2-11.2 m)
when the first observations were made (Fig. 1,
Reef 200 ) . Although the temporal changes in
communities observed here cannot be presented
as a typical ecological succession, they have had
profound influences upon the composition of
the epifauna of this rock-reef. My purpose in
calling attention to these changes is to stimulate
other investigators to follow the course of simi-
lar population shifts that may occur in regions
under their purview. Additional data may make
it possible to delineate the role of successional
heterochrony in creating the complex epifauna
present on submarine reefs at any given point
in time.

The central species in the present study is the
common mussel, Mytilus edulis . Beginning in
late January, 1959, the Mytilus population on
the inshore reef began a decline in density,
which was not recognized as of singular im-
portance at the time. However, before this de-
cline leveled off it resulted in an almost complete
disappearance of this species from the reef. The
basic cause is unknown, although the early sharp-
ness of decline suggests that disease may have
been a contributing factor. Still, the decline did
not occur on the deeper reef, some 300 m distant,
although it must be pointed out that the maxi-
mum population densities here were only a
hundredth of those on the shallower reef. In
addition to the possibility of disease, it is known
that an unusually large population of Pisaster
giganteus was preying on the dense mussel popu-
lation during preceding weeks. Although the
decline extended through a year, the bulk of
the Mytilus population disappeared in half this
time. Unfortunately observations were not made
during part of this period, nor has it been pos-
sible to follow developments after Mytilus was
nearly wiped out. Nevertheless, sufficient data
are available to provide a picture of the biotal
shifts that accompanied the decline. The dis-

424

Population Dynamics— Pequegn AT

425

Fig. 1. Location of the two sublittoral rock-reefs
under study, in relation to the city of Corona Del Mar
and the entrance to Newport Harbor, Orange County,
California.

appearance of certain predominant species ap-
parently triggered increases and decreases in
associated species. And these changes proved to
be very extensive, involving some species that
would not have been expected to react to changes
in Mytilus populations.

STUDY SITE

The rock mass under study is composed of
siltstone that has been subjected to complex
folding. Gorsline (1962) advises that these
promontories may properly be called submarine
hogbacks. The dip of the strata ranges from a
few degrees to 90°, but the larger part exhibits
the latter dip. Because it is located approximately
200 m offshore, I shall refer to it hereinafter
as Reef 200.

Situated about 2.5 km southeast of the en-
trance to Newport Harbor, Reef 200 ( Fig. 1 )
rises from a sand bottom at a greatest depth of
11.2 m to a shallow point of 2 m at standard
mean sea level. It has a minimum surface area
of 1,200 m2 distributed over a length of 58 m
and an average width of 27 m (ranges from 8
to 44 m). Because its long axis parallels the
shore, the hogback lies athwart the direction
of surface wave propagation. This results in in-
terference with wave transmission and a high
degree of turbulence at shallow points. The
magnitude of turbulence is inversely related to

depth; hence it falls off rapidly down the walls
and is barely perceptible at the reef base on
days of average swell.

Most of the Mytilus population was confined
to the reef’s flat top, between depths of 2 and
4 m. The same was true of such brown algae as
Eisenia arborea , Egregia australis , and Laminaria
farlowi.

STUDY METHODS

Samples of the Mytilus populations of Reef
200 were obtained under water by using SCUBA.
Quadrats were established by placing metal rings
encompassing 0.1 m2 against the biota and re-
moving all components by hand. Where re-
quired, hammers, chisels, and. forceps were also
utilized. All specimens were placed in canvas
bags, which were sealed prior to ascent and re-
turned to the laboratory. Subsequently species
were identified, and individuals of numerable
species (noncolonial) were counted and meas-
ured.

Prior to the Mytilus decline, study sites were
selected by dropping metal rings onto the reef
from a moving boat. Samples were removed
from the sites where the rings came to rest.
During and after the decline, conscious attempts
were made to remove samples within a few
meters of predecline quadrats. This was possible
since chisel marks were still evident on the
rocks. The first samples were taken on October
29, 1958, and the last on February 22, I960.
The longest period without samples extended
from February 15 to August 19, 1959, when
the author was in Europe. For most purposes I
have divided the dates of sampling into three
periods of time: Predecline (October, 1958, to
January 15, 1959), Decline (January 16 to
October 15, 1959), Postdecline (October 16,
1959, to February, I960). Although Mytilus had
not disappeared completely from the reef at the
end of the study, its average population per 0.1
m2 quadrat was then only 0.5% of its original
value. Twelve 0.1 m2 quadrats were studied
during the above periods.

Large species, such as Pisaster giganteus,
Strongylocentrotns purpuratus, and Parasticho-
pus parvimensis, were sampled with metal rings
encompassing 1 m2. Specimens of these species
were not removed from the reef. Samples were

426

PACIFIC SCIENCE, VoL XVII, October 1963

taken at random distances along transverse and
longitudinal transects. Some 100 sq m quadrats
were established during this study.

RESULTS

During the decline, the Mytilus edulis popu-
lation at depths of 2 m dropped from an average
of about 1,100 to less than 10 individuals per
0.1 m2. Even more drastic reduction eventually
occurred at greater depths (Table 1). And as
Mytilus declined at specific sites, populations of
the red alga Corallina chilensis, which are stated
in terms of percentage of rock surface covered
in Table 1, increased on the same sites, particu-
larly above depths of 4 m. There was, however,
a considerable lag between Mytilus depletion
and the spread of Corallina. This lag permitted
rapid population increases of a few invertebrates.
But, as will be discussed later, some of these
increases were cut back after the encroachment
of Corallina got underway. In turn these popula-
tion shifts triggered other reciprocal changes
among less noticeable species, especially among
platyhelminths, nemerteans, mollusks, annelids,
and crustaceans (Table 2). Fortunately sufficient
samples had been taken prior to, or early in,
the Mytilus decline to establish a basis of com-
parison with subsequent unforeseen events. The
samples of November, 1958 (Table 1), reveal
( 1 ) that most of the Mytilus population lived
on horizontal surfaces at depths between 2 and
4 m, ( 2 ) that its density decreased sharply with
increasing depth to the 7-m level, and (3) that

it was absent from this point downward. These
relationships persisted unchanged as late as Jan-
uary 15, 1959. Soon thereafter, however, it was
noted that Mytilus shells were beginning to ac-
cumulate in unusual numbers on the shoreward
bottom of the reef. Unfortunately an impending
trip to Europe precluded additional sampling
for several months. But by August, 1959, the
Mytilus population at the 2-m level (samples
were taken only a few meters horizontally from
the predecline quadrats) had dropped about
80%. And, as can be seen in Table 1, by Febru-
ary, I960, considered here to be early post-
decline, Mytilus had almost disappeared from
the entire reef. Apparently at depths below 2
m the initial rate of decline was not as rapid,
but complete disappearance eventually occurred.
The later occurrence of small individuals at
depths of 8 and 9 m is probably explained by
dislodgment of small clumps from the reef-top.
As seen in Table 1, Corallina increased markedly
during this time, and came to form a uniform
mat, some 6 cm thick, over much of the reef’s
upper surface. This trend continued until at the
culmination of the study it covered 65% of the
shallowest part of the reef with a mat some 9
cm thick.

Meanwhile significant changes occurred in
molluscan populations. Whereas an average of
25 molluscan species per plot (among a total of
some 35 species on all plots) occurred with
Mytilus (at 2-m depths) prior to the decline,
this average dropped to 11 by September, 1959

TABLE 1

Changes in Population Densities, with Depth and Time, of Mytilus and Corallina
(Dashes indicate absence of data)

DEPTH

(m)

Mytilus edulis: IND /0.1 M2*

BY DATE

Corallina chilensis: % COVERf

BY DATE

11/58

9/59

10/59

2/60

11/58

9/59

10/59

2/60

1,086

176

180

* Individuals per quadrat,
f Per cent of quadrat surface covered by plant.

Population Dynamics — PEQUEGNAT

427

TABLE 2

Data Gathered from 0.1 m2 Quadrats at 2-m Level Before, During, and
After Major Part of Mytilus Decline

NO. SPECIES IN PHYLA (Average)

PHYLA

Predecline

Decline

Postdecline

Porifera

2.5

1.5

Coelenterata

Platyhelminthes

Endoprocta

0.5

Ectoprocta

3.5

Sipunculoidea

Nemertea

Mollusca

Annelida

Arthropoda (Crustacea)

6.5

Echinodermata

2.5

Chordata (Ascidiacea)

TOTAL ANIMAL SPECIES

48.5

Numerable Species

Motile Species

TOTAL ANIMAL INDIVIDUALS/0.1 M2

1759

1754

841

(Table 2). Then, as the result of the encroach-
ment of different molluscan species, it increased
again to an average of 25 by February, I960.
Four types of population shifts were exhibited
by molluscan species during this time: (1) the
numbers of one species originally present in-
creased, (2) several species declined but per-
sisted, (3) others declined in numbers and
eventually disappeared, and (4) previously ab-
sent species invaded the Corallina mat. Chama
pellucida is the only species that increased and
retained its population gain, although, as would
be expected, the new individuals were very
small. Twelve of the original 35 species suffered
severe population drops, but were still present
in February, I960. Illustrative of these, in addi-
tion to Mytilus edulis, are Hiatella arctica , which
dropped from an average of 135 to 10 indi-
viduals / 0.1 m2, and Pterorytis nuttalli, which
dropped from an average of 16 to 2 / 0.1 m2.
Twenty species of mollusks that were originally
present disappeared completely. Among the
more prominent of these are Mytilus calif ornia-
nus, Modiolus capox, Tegula ligulata, Amphissa
hicolor, and Gians carpenteri . Among the 15
invading species are Anomia peruviana, Chlamys
hastatus, Flabellina iodinea, Doriopsilla fulva,

and Erato vitellina. The following three species
were unaffected by the other changes: Crepipa-
tella lingulata, Pholadidea penita, and P. ovoidea.

Similar changes took place among the poly-
chaetes. Whereas an average of 12 species oc-
curred on Mytilus quadrats (2-m depth) prior
to the decline, this dropped to 7 in October,
1959, and then rose sharply to 21 by February,
I960 (Table 2). The greatest increase was ex-
hibited by Nereis grubei, which shot from an
early average of 8 to 96 individuals / 0.1 m2
by October, 1959. It persisted at or above 80 /
0.1 m2 into February, I960. The populations of
Anaitides mucosa and Platynereis sp. more than
doubled in this time. Arabella iricolor declined
at first, but recovered its earlier population size
by February, I960. On the other hand, Eunice
rubra, Typosyllis pulchra, Polycirrus sp., and
Polydora sp. declined sharply and eventually dis-
appeared. Eleven species of polychaetes, not
previously present, invaded the Corallina mats.
Among the more interesting of these are Lum-
brinereis zonata, Anaitides sp., and Nainereis
dendritica. None of the invading species came
to be represented by many individuals.

Among the crustaceans, Balanus tintinnabu-
lum displayed a marked population surge from

428

PACIFIC SCIENCE, Vol. XVII, October 1963

a predecline average of 30 individuals to 800 /

O. 1 m2 during the decline. This dropped back
to an average of 350 living individuals per 0.1
m2 by February, I960. Curiously, Balanus tri-
gonus populations remained unchanged through-
out the period, as did those of Pachycheles rudis,

P. holosericus, and Crangon dentipes. Four spe-
cies of crustaceans disappeared during the My-
tilus decline: Paraxanthias taylori, Cirolana
harfordi , Lophopanopeus leucomanus, and L.
diegensis, the latter of which had been repre-
sented only by young. These losses were more
than offset by the appearance of six species,
among which are Pelia clausa and Pugettia dalli.

Among the smaller echinoderms, Ophiactis
simplex reacted quickly by nearly doubling its
population at the 2-m level, going from 180
individuals to 340/0.1 m2 by October, 1959.
This dropped back to an average of 154/0.1 m2
by February, I960. The plankton-feeding holo-
thuroid Cucumaria lubrica appeared on the 2-m
level for the first time.

While working with the above data I was
struck by the fact that throughout the entire
period from November, 1958, to February, I960,
over 72% of the numerable individuals at the
2-m level were found in only four species (Table
3). We see, however, that the relative percent-
ages that each contributed to the total shifted
considerably during the decline.

Now let us turn to an analysis of shifts among
the larger species existing on Reef 200. Not

only did populations of some of these decline
significantly, but there were also shifts of popu-
lation centers on the vertical axis. One might
easily have predicted some, but not all, of these
changes. Prior to the decline, 70% of the Pisas-
ter giganteus population was found on the upper
half of the reef in close association with Mytilus
(Table 4). Subsequently this percentage dropped
to 46. At the same time the Pisaster population
on the entire reef dropped about 45%, presum-
ably as a result of emigration to new sites. Most
of the Pisaster emigrants must have come from
the upper reef. Quite unexpectedly the sea urchin
Strongylocentrotus purpuratus appears to have
been markedly affected by the Mytilus loss. Its
population dropped 34% and data in Table 4
point also to a downward movement on the reef.
Strongylocentrotus franciscanus, on the other
hand, suffered only a small loss of total popula-
tion, and the center of population remained on
the lower reef as before. The quadrat densities
of the sea cucumber Parastichopus parvimensis
dropped 30% during the Mytilus decline, but
the center of population remained on the lower
half of the reef.

DISCUSSION

Four kinds of population changes occurred
during the Mytilus decline: (1) some species
previously present with the mussel increased in
numbers and held their gains; (2) some species

TABLE 3

Percentage of Total Individuals of Numerable Species on
Each Quadrat Provided by Only Four Species

SPECIES*

PERCENTAGES OF

TOTAL POPULATION BY DATEf

11/21/58

1/16/59

9/4/59

10/4/59

2/14/60

Mytilus edulis

0.2

Balanus tintinnahulum

Nereis grubei

0.2

0.7

Ophiactis simplex

PERCENTAGE OF POPULATION

74.2

74.7

72.2

NO. OF

OTHER NUMERABLE SPECIES

* Note that these four species accounted for an average of 75% of the total animal population, but their relative contribu-
tions to the total changed markedly during the decline.

t Data obtained prior to (11/21/58) and during the Mytilus decline on the dates heading columns. All quadrats situated
at depths of 2 m and within 3 m of the first.

Population Dynamics— -PEQUEGN AT

429

TABLE 4

Relative Population Densities of Large, Motile Species Before and
During the Decline of Mytilus Populations

SPECIES

INDIVIDUALS /M2*

Predecline

During Decline

depth (m) f

depth (m)f

2-7

7.1-11.2

2-7

7.1-11.2

Pisaster giganteus

1.7

1.2

1.5

Strongylocentrotus purpuratus

3.7

0.5

1.6

1.0

Strongylocentrotus franciscanus

0.5

5.8

1.1

4.8

Parastichopus parvimensis

0.1

0.6

0.5

* Average number of individuals per square meter.

f Reef is divided roughly into upper (2-7 m depth) and lower parts to show some redistribution of individuals of some
species with depth.

appeared only after Corallina became predomi-
nant; (3) others maintained their populations
at previous levels, though most underwent tem-
porary fluctuations; and (4) still others de-
creased and, in some instances, disappeared. Most
of the major population shifts occurred where
Mytilus was previously dominant, but a few
significant shifts were noted below the 4-m line.
The degree of these effects may provide some
measure of the closeness of relationship of in-
dividual species to the mussel (and to Corallina ),
both in positive and negative (inhibitory) ways.
Our knowledge of these relationships will be
increased significantly when careful studies are
made of the feeding habits of the many species
involved.

Some species increased in numbers simply
because space became available and they had the
reproductive capacity to spread out. Balanus
tintinnahulum exemplifies this; its larvae were
available to settle prior to the encroachment of
Corallina. It is apparent, also, that Corallina is
unable to invade an established Mytilus bed,
but when the latter is gone the alga moves in
after a few weeks’ lag. Other species increased
after this alga had become established, apparently
because it provided a more suitable milieu. Quite
possibly some species increased as a result of the
withdrawal of predators not adapted to living
in algal mats. The marked decrease in disaster
giganteus resulted from an abrupt loss of food.
The shift of remaining individuals to the lower

part of the reef is probably related to the pres-
ence there of Chama pellucida, an alternate food
source. The anemone Anthopleura sp. dropped
from an average of 40 individuals to 17/0.1 m2
because of a reduction- of substratum rather than
of food. Prior to the decline most anemone in-
dividuals lived on mussel valves, with a few
living in the rock niches. The number increased
on the rock as the mussel declined, but the total
available space was not sufficient to sustain the
previous populations. The reduction of Para-
stichopus parvimensis may be accounted for by
the greater production of detritus by Mytilus
than Corallina, but this is purely conjectural.
The different reactions of the sea urchins Stron-
gylocentrotus purpuratus and S. franciscanus to
the Mytilus decline indicate that broad generali-
zations are not appropriate. Whereas 5'. purpura-
tus suffered a severe decline, the other species
was unaffected. It is perhaps significant that S.
purpuratus existed primarily on the upper half
of the reef, whereas S. franciscanus held forth
on the lower half. The withdrawal of S. pur-
puratus may have resulted from the reduction
of niches provided by the Mytilus beds and the
fact that Corallina produced a very compact
cover over the rocky surface.

Some interesting relationships between the
numbers of species and individuals occurred
during this disruptive change in community
structure. These can be ascertained from the last
four line items of Table 2. Note, for example,

430

PACIFIC SCIENCE, VoL XVII, October 1963

that while the number of numerable (non-
colonial) species per quadrat dropped about
25% during the decline, the total of individuals
persisted unchanged for many months. But, as
can be seen, by the beginning of the postdecline
the total number of individuals in the numerable
species fell markedly, in spite of the marked
increase in numbers of species per quadrat. We
find part of the explanation of this in the fact
that when Mytilus was at its peak there was an
average of 31 individuals per numerable species
(aside from Mytilus) on each quadrat; that
during the decline this jumped to 41, and then
dropped to 12 in the early postdecline. These
phenomena are based upon the following oc-
currences: (1) the rapid population increases
of a few such species as Nereis grubei, Balanus
tintinnabulum, and Ophiactis simplex; while
(2) other species were leaving the sites; (3)
formerly absent species, principally motile ones
(Table 2), invaded the sites, resulting in an
increase of 30% over the original number of
species; (4) those species that underwent rapid
population increases overshot the mark and
dropped back; and (5) all of the invading
species, at least during the period of this study,
were represented by a small number of indi-
viduals. It is of interest that no single animal
species equalled the former population density
of Mytilus . Balanus came closest, but it is only
a fraction the size of the mussel. This indicates
that the animal contribution to the total biomass
in the Hyperbenthal Zone (Pequegnat, 1961)
was markedly reduced; the emigrations of the
echinoderms mentioned above also account for
a considerable loss. Reciprocally, the plant com-
ponent must have risen through the Corallina
increase.

For several reasons the appearance of the
Corallina mat and associated species appears to

comprise an unstable complex. Judging from the
population changes discussed above and from
observations made elsewhere in the subtidal and
intertidal, one gains the impression that ex-
tremely dense growths of Corallina are asso-
ciated with disturbed conditions. Though one
might expect this predominance of the corallines
to be temporary, Dawson (1959) points to the
distinct possibility that the corallines encroach
upon and later dominate areas subject to pol-
lution from human wastes. Hence interest is
heightened in obtaining additional samples in
the future.

REFERENCES

Dawson, E, Yale. 1959. A Primary Report
on the Benthic Marine Flora of Southern
California. Oceanogr. Survey of the Continen-
tal Shelf Area of S. Calif. Publ. No. 20. State
Water Pollution Control Board. Sacramento,
California.

Gorsline, Donn S. 1962. Personal communi-
cation.

HARTMAN, 0. 1944. Polychaetous annelids. Rep.
Allan Hancock Pacif. Exped. 3:1-33.

1950. Goniadidae, Glyceridae, Nephyti-

dae. Rep. Allan Hancock Pacif. Exped. 15:
1-181.

— 1951. The littoral marine annelids of

the Gulf of Mexico. Publ. Inst. Mar. Sci.,
Univ. Texas 2:7-124.

Pequegnat, Willis E. 1961. New world for
marine biologists. Nat. Hist. 70(4).

Preliminary Report on the Marquesan Sardine,
Harengula vittata, in Hawaii

Thomas S. Hida and Robert A. Morris1

The Marquesan sardine, Harengula vittata,

was introduced to Hawaiian waters in eight
plantings from 1955 through 1959 by the Bu-
reau of Commercial Fisheries in an attempt to
establish this species as a supplementary bait
fish for skipjack fishing. The details of the first
seven introductions have been reported by
Murphy (I960) and the eighth by Brock (I960).
The eighth introduction comprised an estimated
4,000 sardines ranging from 7.0 to 9.4 cm in
standard length2 and averaging 8.2 cm. This
brought the estimated total number of sardines
introduced to Hawaii to 144,000. All of the
releases have been made around the island of
Oahu (Fig. 1).

SOURCES OF INFORMATION

The introduction of the sardines was publi-
cized by means of posters and letters. Commer-
cial and sport fishermen and game wardens were
asked to cooperate in supplying information on
sightings and captures. Jars of formalin and
labels for recording data concerning captures
were supplied to the game wardens and the skip-
jack fishermen, whose bait nets were considered
a likely source of specimens. From 1956 through
I960, 54 samples comprising 336 fish were
turned in, the majority of them by skipjack fish-
ermen.

DISTRIBUTION

Recaptures of Marquesan sardines have been
reported from six of the eight major islands in

1 U. S. Bureau of Commercial Fisheries Biological
Laboratory, Honolulu, Hawaii. Manuscript received
May 21, 1962.

2 All of the length measurements appearing in this
report are expressed in standard length, which is the
distance from the tip of the snout to the end of the
hypural.

Hawaii ( Fig. 2 ) . The lack of reports from Lanai
and Niihau may be due to the fact that skipjack
fishermen rarely fish for bait in the waters
around those islands.

Tabic 1 lists all the sardine recoveries from
the Hawaiian Islands. The first recapture was
made in Keehi Lagoon, Honolulu, in 1956.
Subsequent recoveries were from Barber’s Point
and Kaneohe Bay, Oahu, in 1957. In 1958 sar-
dines were taken at the islands of Kauai and
Maui, where no releases had been made, and also
in Honolulu Harbor and Pearl Harbor, Oahu,
for the first time. In 1959 the sardine was re-
ported from the islands of Hawaii and Kahoo-
lawe, and for the first time from Haleiwa, Oahu.
The first specimens from Molokai and from Na-
wiliwili and Hanamaulu bays, Kauai, came in
I960. Sardines have been taken from both the
leeward and windward shores of Oahu and Kauai
but only from the leeward shores of Hawaii,
Maui, Kahoolawe, and Molokai ( Fig. 2 ) .

Neither our knowledge of the life history of
this species nor our observations of it in Ha-
waiian waters are adequate to tell us whether
the extension of its distribution from Oahu to
the other islands has come about through migra-
tion of the adults across the channels or through
the drifting of eggs or larvae with the currents.

The habitat occupied by the sardine seems to
coincide with that of the nehu, Stolephorus
purpureas, the most commonly used tuna bait
fish in Hawaii. This distribution pattern may be
only a sampling artifact, since most of the fishing
with gear likely to take sardines is done by
skipjack fishermen fishing for nehu. However,
because of the chronic shortage of tuna bait in
Hawaii, the fishermen are alert for reports of
bait supplies even in areas outside the usual
nehu fishing grounds. If sardines were present in
conspicuous abundance in any accessible area,
it appears highly probable that the fishermen

431

432

PACIFIC SCIENCE, Vol. XVII, October 1963

TABLE 1

Records of Marquesan Sardines Collected in Hawaii

LOCALITY

DATE OF

CAPTURE

METHOD OF

CAPTURE

NUMBER

CAUGHT

NUMBER

PRESERVED

STANDARD LENGTH
(cm)

Range

Mean

Oahu I.

Keehi Lagoon

9/27/56

night net1

9.97-

9.97

3/27/57

night net1

8.50-

8.50

10/4/59

night net1

est.

3.59-5.60

4.40

11/3/59

night net1

500

5.25-5.68

5.46

11/5/59

night net1

4.08-6 07

4.88

11/25/59

night net1

4.03-6.46

5.51

12/1/59

(?)

3.05-3.51

3.28

7/20/60

day seine

est.

2.92-

2.92

9/14/60

day seine

3.39-4.72

4.27

Kaneohe Bay

3/26/57

day seine

est.

8.75-

8.75

4/8/57

night net

8.07-8.75

8.46

6/4/57

night net

9.50-

9.50

6/30/58

(?)

9.75-

9.75

9/23/58

day seine

5.60-

5.60

11/3/58

day seine

3.46-

3.46

11/27/58

day seine

est.

3.53-4.30

3.96

7/20/59

day seine

4.30-13.21

7.66

9/4/59

day seine

4.90-8.56

6.73

9/6/59

day seine

3.04-3.22

3.13

9/11/59

day seine

8.30-

8.30

9/14/59

day seine

8.21-8.67

8.44

10/2/59

day seine

5.40-

5.40

2/26/60

day seine

12.01-

12.01

6/9/60

day seine

3.34-3.68

3.51

Barbers Point

9/9/57

gill net

11.20-13.50

12.55

Honolulu Harbor

2/28/58

night net

7.71-9.40

8.72

6/27/58

night net

9.54-

9.55

Pearl Harbor

4/29/58

day seine

10.05-10.71

10.45

Haleiwa

9/2/59

day seine

4.46-6.10

5.27

6/22/60

day seine

est.

3.98-5.14

4.54

7/22/60

day seine

4.71-8.64

6.67

Maui I.

Kihei

7/5/58

(?)

not exam.

10/7/58

day seine

3.49-5.32

4.67

11/3/58

day seine

6.32-7.00

6.59

5/19/59

day seine

est.

9.12-

9.12

5/28/59

day seine

9.10-10.91

9.91

5/29/59

day seine

9.29-10.83

10.06

6/2/59

day seine

8.56-

8.56

Maalaea Bay

8/13/59

day seine

5.62-6.00

5.87

1 The net used in Hawaii for taking tuna bait at night is a rectangular lift net pulled under the fish which are attracted to
an underwater light suspended alongside the fishing boat.

2 A "bucket” of bait in the Hawaiian skipjack fishery is estimated to represent about 7 lb of fish.

Marquesan Sardine — HlDA and MORRIS

433

TABLE 1 — Continued

LOCALITY

DATE OF

CAPTURE

METHOD OF

CAPTURE

NUMBER

CAUGHT

NUMBER

PRESERVED

STANDARD LENGTH
(cm)

Range

Mean

Kauai I.

Port Allen

9/10/58

night net

6.17-6.69

6.41

10/2/58

night net

3.90-7.90

6.60

5/24/60

day seine

est.

100

3.80-7.34

5.05

7/5/60

day seine

est.

500

4.00-6.87

5.45

7/9/60

day seine

est.

100

5.85-8.75

7.24

7/23/60

day seine

est.

4.67-5.99

5.54

Hanalei Bay

9/25/58

(?) .

6.90-7.90

7.50

Waimea Bay

l/P/59

pole-and-line

9.80-11.48

10.64

10/14/59

pole-and-line

14.62-

14.62

7/23/60

day seine

est.

11.34-13.30

12.32

Nawiliwili Harbor

6/2/60

night net

10.57-12.47

11.75

Hanamaulu Bay

9/6/60

gill net

12.21-

12.21

Hawaii I.

Kawaihae Bay

3/14/59

day seine

est.

3 bkts.2

7.56-10.05

8.73

Kahoolawe I.

SW Point

6/17/59

day seine

est.

Vl bkt.

9.76-12.08

10.83

Molokai I.

Laau Point

9/3/60

day seine

5.61-6.70

5.80

would find them. The main nehu baiting areas
are in the relatively protected waters of bays,
harbors, and canals where the water is brackish
and turbid owing to the influx of streams and
ground water. The Kihei area on the island of
Maui and Waimea on Kauai are exceptions, the
baiting grounds being exposed to the open ocean
and moderate surf action. Baiting, in general, is
done close to shore in shallow water, although
night-lighting for bait may be done in deeper
waters of channels and harbors. Since it appears
that the sardine has established itself in the
nehu grounds, it is important, from the stand-
point of the tuna fishery, to keep a close watch
on the population trend of the species in Hawaii
and the effect that it may have on the bait supply
in the future.

ABUNDANCE

Our only source of information on the abun-
dance of sardines in Hawaiian waters has been
the reports of observations by commercial and
sport fishermen. The numbers of specimens
turned in by fishermen are not a good index of
abundance, because often they are only a small
and arbitrarily selected fraction of the catch.
There have also been times when sardines were
seen but not caught.

The first observation of a large school of
sardines, 20 to 30 buckets,3 was made in Wai-
mea, Kauai, late in the summer of 1958 by skip-

3 A "bucket,” the unit commonly used by skipjack
fishermen for measuring bait, contains an average of
7 lb of fish.

434

PACIFIC SCIENCE, VoL XVII, October 1963

Fig. 1. Map of Oahu showing areas where sar-
dines have been released.

jack fishermen. The school remained in Waimea
Bay until January 1959; its disappearance from
the Bay coincided with a storm which occurred
in that month. In March, 1959, a skipjack boat
caught 3 buckets of sardines at Kawaihae, Ha-
waii, and reported catching 900 lb of skipjack
with them, a bait-catch ratio comparable to that
ordinarily obtained using nehu. The fishermen
commented that the sardines were an excellent
bait. In October, 1959, an estimated 500 sardines
were caught in Keehi Lagoon, Oahu, by skipjack
fishermen. These were used for bait, but no
report was submitted on the results. The latest
report of sizeable sardine catches came from
Port Allen, Kauai, where a catch of 100 sardines
was made in May, I960, and a catch of 500
sardines and another of 100 was made in July,
I960. Aside from these few instances of fairly
large catches, most of the reports from the fisher-
men have indicated that the sardine was taken
or sighted only in small numbers. Thus, although
the sardine appears to be well established in
Hawaii, it is apparently not abundant, at least
not in areas frequented by fishermen, and is not
as yet making any significant contribution as a
bait fish.

FOOD

We examined the stomach contents of 132
sardines caught in Hawaii to study their food

habits in their new environment. This examina-
tion covered samples representative of all areas
from which specimens had been turned in. Only
fish with identifiable food organisms were in-
cluded in the analysis. We did not consider as
food items nematodes, wood, and other foreign
matter, or material which we could not readily
identify. Thirty-nine (30%) of the stomachs
examined were considered to be empty. This
high percentage of empty stomachs was prob-
ably due to the fact that many of the fish had
been held captive for several days in baitwells.

In terms of the percentage frequency of oc-
currence of various organisms in the stomachs
of all samples (Table 2), copepods were first,
followed by gastropod larvae, adult and larval
shrimp, crab larvae, fish larvae, amphipods, and
polychaetes. Copepods were also observed in the
greatest numbers.

The following list of the numbers and kinds
of organisms found in the well-distended stom-
ach of a 9.4-cm sardine caught at Kihei, Maui,
represents the quantity and variety of food that
may be taken by this fish:

689 copepods 24 lucifers
34 amphipods 7 ostracods
30 shrimp larvae 3 crab megalops
27 crab zoeae 3 stomatopod larvae

1 unidentified fish larva

In contrast, the stomach of another 9.4-cm
sardine appeared filled to capacity with three
nehu larvae about 3 cm long.

Of the fish examined, 50% were between 3-5
and 6.0 cm long, the rest ranging up to 14.5 cm.

Fig. 2. Map of the Hawaiian Islands showing the
areas from which sardines have been recovered.

Percentage Frequency of Occurrence of Various Organisms in the
Stomachs of 93 Sardines

Marquesan Sardine — HlDA and MORRIS

435

vaaooaviD ^ i i • ! i I

vaodonaxd

: : : : : : : : o : :

: : : : : : : : <n : :

VHDVaiSAPY

3VAHVT XD3SNI

vaodosi

: oo vd o : : : : : :

: cn ; : : : : t-h :

vaoovHxso

: : : : : : : i — :

VHXVNDOX3VHD

>/N

3VA3VT <30dA33T3d

33313111

100

(•<Js pjuppy)
VCI0d033X3H

: : : : : : : : o : :

: : : : : : : : oo : :

3VA3V1

aodoxvwoxs

SIHdAD 313VN3V9

: : ■> — i : : r- : o o : :

: : m : : : cm *-i ; :

VXaVHOATOd

cot o */n : r — : cn : : : :

m v~\ cn : r-H : i— < : : : :

l/N

VdOdlHdDtV

: irs o : o o m:

: cn i-i (N t-=i : oo r-t rr> :

3VAHV1 HSI3

100

r-~-

3VA3V1 9V33

mi'— • o ro m : o o :

m:mooro »-4 : xj< ex vo :

avAHVT * snnav

dTtlHHS

3VA3V1 aOdORXSVO

: o 5— i o : m : : o m :

: m cn : m : : xp oo :

vaodadoo

100

LOCALITY
(No. of fish
examined )

Oahu I.

Honolulu Harbor
(3)

Haleiwa

(12)

Kaneohe Bay
(16)

Barbers Point

(5)

Keehi Lagoon
(12)

Maui I.

Kihei

(15)

Hawaii I.

Kawaihae Bay
(3)

Molokai I.

Laau Point

(5)

Kahoolawe I.

SW Point
(10)

Kauai I.

Port Allen

(6)

Waimea Bay
(1)

Ail areas combined

436

PACIFIC SCIENCE, VoL XVII, October 1963

The food habits of the larger and smaller sar-
dines appeared to be essentially similar, but
fish larvae were found only in sardines 8.0 cm
long or larger, and there seemed to be a size-
associated difference in the composition of the
copepod component of the diet. The smaller fish
fed more commonly on small cyclopoid cope-
pods, such as Corycaeus sp. and Oncaea sp.,
while the larger sardines had more often been

! :

119581

1959

1 19601

.1.

JAN. FEB. MAR. APR. MAY JUNE JULY AUG. SEPT. OCT. NOV. DEC.

Fig. 3. Size frequencies of sardines collected in
Hawaii in 1958, 1959, and I960.

feeding on larger calanoids, such as Candacia
sp., Labidocera sp., and Pleuromamma sp. The
samples were inadequate to carry on a detailed
study of food habits among the different areas.

As indicated in Table 2, 17% of the stomachs
with food had fish larvae in them. Of 34 fish
larvae found, 24 (71%) were identified as nehu.
The largest number of larval nehu found in a
stomach was 6, in a 10.8-cm specimen from
Kihei, Maui. The largest nehu found measured
3-3 cm, in a 9.4-cm sardine, also from Kihei,
Maui.

The food habits of H. vittata in its native
environment were extensively studied by Naka-
mura and Wilson (ms). They also found cope-
pods to be the most frequently occurring or-
ganism (79.1%), followed by pelecypods, gas-
tropods, barnacle cypris, pteropods, amphipods,
and megalops. Fish larvae occurred in only 1.6%
of the stomachs. The only distinct difference
between the feeding habits of the species in
Hawaii and in the Marquesas seems to be the
low percentage of occurrence of fish larvae in
the Marquesan fish.

The food habits of the sardine in Hawaii are
similar to those of the nehu, both species feed-
ing largely on the crustacean elements in the
plankton. Hiatt (1951) found that nehu feed
primarily on copepods* barnacle larvae, mysis
larvae of shrimps, ghost shrimp ( Lucifer sp.),
crab larvae, and palaemonid shrimps.

SPAWNING

The size of the first sardines taken at Maui
and Kauai in 1958 was such that, as pointed out
by Murphy (I960), it seemed more likely that
they were the products of spawning in Hawaiian
waters rather than members of the original
transplanted stock. A collection of 29 sardines
taken at Kihei, Maui, in October, 1958, con-
tained 7 measuring less than 4.3 cm, which was
the length of the smallest sardine measured in
the release of May, 1958. Subsequently, sardines
smaller than 4.3 cm were taken in Keehi Lagoon
and Kaneohe Bay in July, September, October,
November, and December, 1959 (Fig. 3). In
I960, small sardines were caught at Keehi La-
goon, Kaneohe Bay, and Haleiwa on Oahu, and
at Port Allen, Kauai, in May, June, July, and
September.

Marquesan Sardine— -HlDA and MORRIS

Nakamura and Wilson (ms) have reported
that in the Marquesas 8.4 cm, plus or minus
1.7 cm, is the mean length of females at sexual
maturity. They considered that year-round oc-
currence of ovarian eggs of 0.6 mm or larger
diameter indicated that sardines spawn through-
out the year in Marquesan waters. Although our
Hawaiian samples were inadequate for deter-
mination of the spawning season of the intro-
duced fish, the occurrence of ova 0.43 to 0.74
mm in diameter in sardines taken in May, June,
July, September, and October and the appear-
ance of small sardines from May through De-
cember lead us to believe that the species also
has a prolonged spawning period in Hawaii.

SUMMARY

Eight releases of the Marquesan sardine, Har-
engula vittata, have been made around the island
of Oahu in the hope that it would become estab-
lished and sufficiently abundant to increase the
tuna bait fish supply. An estimated 144,000
individuals were released from 1955 through
1959. Recoveries by skipjack fishermen have
been made from all of the major islands except
two. The habitat occupied by the sardine seems
to coincide with that of the most important
native bait fish, the nehu ( Stolephorus pur -

437

pureus ) . The occurrence of young sardines since
1958 indicates that they have spawned success-
fully, but the species is not yet making any
significant contribution as a bait fish. The sar-
dines feed primarily on copepods, and also eat
gastropod larvae, larval and adult shrimps, crab
larvae, fish larvae, amphipods, and polychaetes.
This diet is similar to that of the nehu.

REFERENCES

Brock, Vernon E. I960. The introduction of
aquatic animals into Hawaiian waters. Int.
Rev. Hydrobiol. 45(4) : 463-480.

Hiatt, Robert W. 1951. Food and feeding
habits of the nehu, Stolephorus purpureus
Fowler. Pacif. Sci. 5(4) .347-358.

Murphy, Garth I. I960. Introduction of the
Marquesan sardine, Harengula vittata (Cuvier
and Valenciennes) , to Hawaiian waters. Pacif.
Sci. 14(2): 185-187.

Nakamura, Eugene L., and Robert C. Wil-
son. ms. The distribution and biology of the
Marquesan sardine. Bu. of Comm. Fish. Bio.
Lab., Honolulu.

Hawaiian Records of Folliculinids (Protozoa) from Submerged Wood1

Donald C. Matthews2

ABSTRACT: To folliculinids in Hawaii, taken from naturally submerged objects
( Halofolliculina annulata, Ascobius simplex, Metafolliculina andrewsi) and from
submerged glass-plate panels ( Metafolliculina nordgardi, Parafolliculina violaceae) ,
are added those from Douglas fir panels riddled by subsistent teredines and
Limnoria: Eufolliculina lignicola, Mirofolliculina limnoriae, and Lagotia viridis.
Variations in loricae and moniliform nuclear components are illustrated and
discussed.

Thus far five species of folliculinids embracing
four genera have been recorded for Hawaii
(Matthews, 1962). Of these, Halofolliculina
annulata, Lagotia simplex, and Metafolliculina
andrewsi were taken from submerged naturally-
occurring objects, whereas Metafolliculina nord-
gardi and Parafolliculina violaceae were taken
from submerged glass-plate panels.

Douglas fir ( Pseudotsuga taxi folia?) frames
supporting these panels (Matthews, 1962, fig.
1), subsequently honeycombed by subsistent
teredines and Limnoria, were broken apart; the
tortuous burrows and their sequestered organ-
isms revealed the folliculinids recorded here.

Folliculina lignicola, Faure-Fremiet 1936, La
famille des Folliculinidae. Mem. Mus.
d’Hist. Nat. de Belg. (Ser. 2), 3: 1129-1175.

Eufolliculina lignicola, Hadzi 1951.

This slender folliculinid (reassigned by Hadzi
[1951] to Eufolliculina lignicola [Faure-Fre-
miet}) was abundant in tracheids opened by
burrowing gribbles. In riddled wood frames
examined November 21, 1961 (corroborated
January 11, 1962) E. lignicola was the most
abundant, although not the most sequestered,
folliculinid. In fact, just outside the burrows,

1 Contribution No. 196, Hawaii Marine Laboratory,
University of Hawaii, Honolulu. Manuscript received
March 28, 1962.

a Department of Zoology, University of Hawaii,
Honolulu.

attached especially to calcareous tubes of ser-
pulid worms ( Spirobis sp. and Mercierella sp.)
were numerous folliculinids whose sacs, necks,
and moniliform nuclei fell well within those
limits prescribed for E. lignicola. Thus, as sug-
gested by Mohr (1959: 86) E. lignicola is not
restricted to tracheids. Mention should be made,
however, that although they were abundant on
calcareous tubes of serpulids, none was attached
to calcareous walls of abandoned teredo burrows,
shells, or pallets, although these apparently af-
forded similar attachment potentials.

As frequently observed in other folliculinids,
sac length, width, and height often varied with
site conditions. In young colonies, composed of
few folliculinids, ample space resulted in normal
sac formation; whereas in old colonies, com-
posed of many folliculinids, limited space re-
sulted in "abnormal” sac formation. Thus, sacs
lying contiguously were usually longer, whereas
those laid one on another were usually shorter,
etc. Despite their occurrence on serpulid shells,
certain restrictive sites seemed preferred. A com-
mon restrictive site is illustrated in Figure 1.
Although the neck (c) was free and always ex-
tended at an angle from the opened portion of
a tracheid (b), the sac (/) usually was con-
fined, at least in part, in the unopened portion
(g). This condition, which resulted in a long
narrow sac (up to 1 66g), affected in no way
either the length of the neck (c) or the number
of its spiral whorls (d) . Thus, regardless of the

438

Folliculinids from Submerged Wood — Matthews

439'

165 p

Fig. 1. Eufolliculina lignicola, as viewed from the right side showing: a, extended peristomal lobes; h,
distal opening of neck; c, neck; d, spiral whorls; e, nuclear components; f, sac; g, unopened portion of tracheid;
h, opened portion of tracheid.

site chosen, neck lengths varied only between
132/x and l67g with the mean at approximately
159 g. Spiral whorls ( d ) usually numbered six,
although occasionally five and seven were ob-
served. The diameter of the neck (49g) was
approximately that of the unrestricted sac.

In reflected light the color of the extended
body resembled that of Parafolliculina violaceae
(Matthews, 1962), except that perhaps in E.
lignicola the wine color was somewhat more

intense. However, in transmitted light the body
and lorica appeared bottle-green, a characteristic
of most folliculinids. As previously reported
(Matthews, 1962), both size and number of
nuclear components varied in accordance with
body contraction resulting from fixation. Thus,
in contracted bodies, nuclear components were
large (5g) and often numbered as few as 6,
whereas in more relaxed bodies, nuclear com-
ponents (Fig. le) were small (3g) and often

440

PACIFIC SCIENCE, Vol. XVII, October 1963

numbered as many as 12. The peristomal lobes
(a) were long (approximately 11 6(x) and in
feeding specimens, were carried high above the
distal opening of the neck ( b ). No swimmers
were observed.

Mirofolliculina limnoriae (Giard 1883) Dons
1927 {vid. Fig. 2 A and B)

Freya limnoriae Giard, Bull. Scientific, t.XV:
264-265, 1883.

Folliculina limnoriae (Giard) Fragments bi-
ologiques XIII. Sur les genres Folliculina
et Pebrilla. Bull. Sci. Nord. 3:310-317,
1888.

Since Giard s publication of Folliculina lim-
noriae, this bizarre folliculinid has been charac-
terized both by its habitat (dorsal surface of
Limnoria pleotelson or adjacent segments) and
by the lateral outpocketings of its lorica (Figs.
2 A,B; 3 A,B; 5 A, B; 6).

Although both Mohr (1959: 86), working
on Limnoria lignorum of the Pacific coast of
North America (Friday Harbor), and Giard
(1888:314), working on Limnoria (sp.) of the

a b

FIGS. 2 A, B. Tracings of Giard’s plates of Fol-
liculina limnoria taken at Wimereux showing: a,
spherical nucleus; b, extent and region of pouches.
Magnification not given.

I 1 , : 1

215 )i 1 80 p

FIGS. 3 A, B. Mirofolliculina limnoriae, showing:
a, empty loricae with lateral pouches.

Atlantic coast of France (Wimereux), consider
M. limnoriae an abundant species, rarely was it
observed on L. ( Paralimnoria ) andrewsi (Cole-
man) of Hawaii (Menzies, 1959:10). In fact,
not until November 29, 1961, after hundreds of
gribbles had been examined, were even the
empty loricae (Fig. 3 A, B) observed. This
paucity may in part be due to the fact that M.
limnoriae temporarily had lost its habitat to
another, as yet undetermined, organism; for,
almost covering the dorsal surface of the pleotel-
son of every third or fourth gribble examined,
was a large brown hemispherical cyst (Fig. 4a)
which, when opened, liberated two minute un-
pigmented dorsoventrally flattened worms. This
impression is further supported by the fact that
during January, 1962, while the numbers of
these cysts decreased, the numbers of M. lim-
noriae increased. However, even in old estab-
lished gribble colonies (such as those from the
riddled hull of the "Seth Parker" at Coconut
Island) only one M. limnoriae was taken for
every 21 L. (P.) andrewsi examined.

Of 25 M. limnoriae taken at random, the
length of the sac varied from 133ft to 21 6g,
with the mean at approximately 149 ft. The sac
at its greatest width, varied from 83 ft to 132g,
with the mean at approximately 11 6/x. The neck,
which was completely devoid of spiral thicken-
ings or proximal lip, varied in length from 49ft
to 66ft, with the mean at approximately 59g.

Folliculinids from Submerged Wood-— MATTHEWS

441

The diameter of the neck was almost uniformly
50 fiy except for those of certain empty loricae
whose necks were ellipsoidal; in such instances
the greater diameter ( at right angle to the longi-
tudinal axis) was roughly 58 /x and the lesser
diameter 2 5 /a.

The greatest variation was observed in the
number, arrangement, and size of the lateral
pouches (Figs. 2 A, B; 3 At B; 5 A, B; 6). Al-
though smaller loricae usually exhibited fewer
pouches, this was not always the case. For ex-
ample, a lorica (Fig. 5 A) with but 6 pouches
( the smallest number observed ) was larger than
a lorica (Fig. 5 B) with 16 pouches. (Nineteen
was the largest number observed. )

Although each lorica was somewhat swollen
at its base, the pouches did not necessarily ex-
tend, as did those shown by Giard (Fig. 2 A,
B ) , in finger-like protrusions from the swollen
base, nor were they bilaterally arranged. Ar-
rangement of pouches varied from as few as 2
pouches on one side and 4 on the other (Fig.
5 A) to as many as 9 on one side and 10 on the
other (Fig. 6). Figure 5 B is interesting for
not only does it show a bilaterally symmetrical
arrangement of the 16 pouches, but the anterior
2 (a) are dearly seen through the neck when
viewed dorsally; hence similar outpocketings (as

.66 mm.

Fig. 4. Pleoteison of L. ( Paralimnoria ) andrewsi,
showing: a, large hemispherical cyst on dorsal surface.

FIGS. 5 A, B. Mirofolliculina limnoriae as viewed
from the dorsal surface, showing: a, pouches visible
through neck; b, lobes of moniliform nucleus; c,
large, lateral pouch.

in Fig. 6a) may have been mistaken for the so-
called valves mentioned by earlier investigators.

Even more variable was the size of the lateral
pouches. These ranged from only a few microns
(Fig. 6b) to half the width of lorica (Fig. 5 A,
c ). And, although large pouches were usually
located posteriorly near the base of the lorica,
occasionally they were located anteriorly near
the neck. Because to my knowledge the process
of lorica formation in this species has not been
observed, neither pouches nor their variation in
number, arrangement, and size can at present
be explained.

Unfortunately, too many observers have been
concerned with the unique lorica and too few
with the living organism. Even actual descrip-
tions are meager. Giard '( 1888:315), who at-
tributed to this species a non-moniliform nu-
cleus (Fig. 2 By a) merely states . . [it] is a
little smaller than the preceeding species { Fol -
liculina abyssorum Giard], of a darker greenish
blue with the striation of the integument less
apparent. . . Kahl ( 1932 ) is concerned that

442

PACIFIC SCIENCE, Vol. XVII, October 1963

folliculinids of quite varied nuclear components
have been assigned to M. limnoriae. This is
understandable. Since many different species of
Limnoria have evolved in various regions of the
world (Menzies, 1959) one would not expect
the folliculinids, with which they are so inti-
mately associated, to have remained constant. In
M. limnoriae, as was previously pointed out for
E. lignicola, nuclear components varied with the
degree of body contraction following fixation.
With peristomal lobes well contracted into the
lorica, the nucleus appeared as little more than
a dark spherical mass near the point of the body
attachment. However, with peristomal lobes re-
laxed and carried high above the distal opening
of the neck, the nucleus appeared as a string of
beads whose size and numbers of components

-i

130 p

Fig. 6. Mirofolliculina limnoriae as viewed from
the dorsal surface, showing: a, pouch viewed through
neck; b, an extremely small pouch.

180 p

Fig. 7. Lagotia viridis, as viewed from the right
side, showing: a, neck; b, three spiral whorls; c, non-
flattened sac; d, spherical nucleus; e, non-spatulate foot.

varied. The largest number of nuclear compo-
nents (Fig. 5 A, a) observed was nine (and this
was not in a completely relaxed specimen). The
question of whether variations in nuclear com-
ponents and/or variations in number, arrange-
ment, and size of lateral pouches have specific
value must await further and more extensive
investigations. However, as mentioned by Mohr
(1959) some correlation seems to obtain be-
tween the latitude in which M. limnoriae is
found and the degree to which its lorica is
branched (compare Giard’s figure of M. lim-
noriae from France {Fig. 2 A, B] with my figure
of M. limnoriae from Hawaii {Fig. 6] ) .

Lagotia viridis, T. S. Wright 1858. Description
of New Protozoa. Edinb. New Phil. Journ,
New Series, 7:276-281.

While not directly attached either on or in
riddled wood panels, L. viridis was abundant on
and in the empty osseous valves of the prio-
nodesmacean mollusk Ostria virginica, which
was present on all panels observed. Although,
in an earlier publication (Matthews, 1953)
Lagotia simplex (Dons, 1917) (reassigned by
Hadzi, 1951, to Ascobius simplex ) was also
credited with occupying this habitat, there is no

Folliculinids from Submerged Wood-— MATTHEWS

443

good reason to confuse these related but quite
distinct species. Although both species are de-
void of vestibule and closing apparatus ( valves )
and possess spherical nuclei, Hadzi is justified
in his reassignment of L. simplex to Ascobius
simplex on the basis of its flattened lorica and
spatulate foot; whereas in Lagotia viridis , the
lorica (Fig. 7c) is not flattened nor is the foot
spatulate (e) . For representatives of L. viridis
in Hawaii an even more obvious difference is at
once apparent. Whereas in Ascobius simplex
the neck is exceedingly short (Matthews, 1952:
344), in Lagotia viridis (Fig. 7) the neck (a)
extends from the sac and is strengthened by
two or three spiral whorls (b). Moreover, while
A. simplex is comparatively small, L. viridis is
comparatively large, often possessing sac lengths
of over 180/a and neck lengths of over 100/a.

Thus, to date, folliculinids embracing eight
species and seven genera are known from Ha-
waii. Accepting Hadzi ’s reassignments, these
now stand as: Halofolliculina annulata (which
I have not yet taken), Ascobius simplex , Meta-
folliculina andrewsi, Metaf olliculina nordgardi
(which may prove to be a new species), Para-
f olliculina viola ceae, Euf olliculina lignicola, Mi -
rof olliculina limnoriae , and Lagotia viridis .

It is interesting to note that, whereas Fol-
liculinopsis gunneri Dons 1927, reassigned by
Hadzi (1951) to P achy f olliculina gunneri, is
present on the ventral surface, of Limnoria lig-
nomm from the Pacific coast of North America
(Mohr, 1959:86), not one specimen has been
observed on the hundreds of L. (P.) andrewsi
which I have examined.

There is every good reason to believe that
new records of folliculinids from Hawaii will
continue to appear. Re-examination of rather
an extensive pagurid collection made over the
past 10 years discloses many folliculinids which
previously had been overlooked. Although these
possess undamaged loricae, their still pigmented
but highly contracted bodies are not suitable
for specific determinations.

REFERENCES

Dons, G 1927. Neue und wenig bekannte Pro-
tozoen. KgL Norske Yidensk. Selsk. Skrifter.
1927(7): 1-17.

Faure-Fremiet, E. 1936. La famille des Fol-
liculinidae (Infusoria-Heterotricha). Mem. du
Mus. d’Hist. Nat. de Belg. (Ser. 2) Fasc. 3:
1129-1175.

Giard, A. 1888. Fragments biologiques XIII.
Sur les genres F olliculina et Pebrilla. Bull. Sci.
Nord. 3:310-317.

Hadzi, J. 1951. Studien fiber Follikulinider.
Academia Scientiarum et Artium. Slovenica
Biology. 2:1-390.

Kahl, A. 1932. Urtiere oder Protozoa. 1, Wimp-
ertiere oder Ciliata Spirotricha. P. 474 in
F. Dahl, Die Tierwelt Deutschlands, 25:399-
650.

Matthews, D. C. 1953. New Hawaiian Records
of Folliculinids (Protozoa). Trans. Amer.
Micros. Soc. 72:344.

1962. Additional Records of Folliculi-
nids (Protozoa) in Hawaii. Pacific Sci. 16(4):
429-433.

Menzies, R. T. 1959- The Identification and
Distribution of the Species of Limnoria. In:
Marine Boring and Fouling Organisms. Dixy
Lee Ray, ed. University of Washington Press,
Seattle. P. 14.

Mohr, J. L. 1959. On the Protozoan Associates
of Limnoria. In: Marine Boring and Fouling
Organisms. Dixy Lee Ray, ed. University of
Washington Press, Seattle. Pp. 84-9 L

Wright, T. S. 1858. Description of New Pro-
tozoa. Edinb. New Phil. Journ. New Series,
7:276-281.

Investigations of Rhinoceros Beetles in West Africa1

Charles P. Hoyt2

In April, 1959, the author was sent by the
South Pacific Commission to the countries of
Sierra Leone and Nigeria, West Africa, to con-
duct an investigation of the natural enemies of
the various species of Oryctes (Coleoptera,
Scarabaeidae, Dynastinae) occurring there. The
object of this study was to find and introduce
to the islands of the South Pacific suitable para-
sites and predators of these beetles in order to
establish a biological control over the introduced
Oryctes rhinoceros Linn, which has become a
serious pest of coconut palms in the area.

In Sierra Leone and Nigeria, coconut palms
are confined mostly to village sites; the groves
are from 15 to 50 palms, depending on the size
of the village. Because of this, nearly all the
work was carried out on oil palms ( Elaeis gui-
neensis Jacq. ) which, together with the Raphia
palms of the swamps, are the most important
hosts of the species of Oryctes found. The oil
palms occur in an extensive belt that follows the
coast of West Africa and extends inland nearly
100 miles in some places.

There are two distinct climatic seasons in
West Africa, and these greatly affect the insect
populations. The wet season begins generally
at the end of April and lasts until sometime in
September. By the end of November the dry
season has set in, and from then until April
little or no rain falls. At times the rains will
cease in August and commence again in Sep-
tember, but during I960 this type of rainfall
did not occur in Nigeria.

Insect activity begins with the coming of the
rains in April or May and gradually ends in
November or December. There are, of course,
many exceptions to this; some species seem to
be found only during the dry season and others
congregate along streams and in swamps, giving
a false impression of their absence.

1 Presented at the Tenth Pacific Science Congress,
Honolulu, Hawaii, 1961.

2 Technical Officer, Pe$ts & Diseases, South Pacific
Commission, Noumea, New Caledonia.

Manuscript received March 19, 1962.

During the dry season in Sierra Leone and in
parts of Nigeria, it is the practice to burn off
the bush to clear the land for planting. The oil
palms in these areas are usually not affected by
the fires, and the larvae of Oryctes and other
beetles within the standing rotten trunks are
not harmed.

The low-lying land of the southern part of
eastern Nigeria is divided by numerous slow-
moving streams which give rise to large swampy
areas as they wind their way towards the sea.
Because of this and the heavy rainfall, extensive
burning is not possible. In the swamps are dense
stands of Raphia palms which flower and die,
providing a continuous supply of breeding sites
for Oryctes. The higher ground between the
streams and swamps supports large stands of oil
palms, both cultivated and wild, from which
come the main export of the area in the form
of palm oil and kernels.

PREDATORS AND PARASITES ENCOUNTERED

Neochryopus savagei Hope (Coleoptera, Cara-
boidea, Scaritidae)

The large scaritid beetle Neochryopus savagei
was first found in Raphia palms growing in the
swamps at Umudike, near Umuahia, eastern
Nigeria. In April, I960, an adult beetle of this
species was found inside a standing rotten
Raphia palm trunk where it was actually en-
gaged in feeding on a larva of Oryctes ohausi
Minck. A search of other rotten palm trunks in
this area turned up two large larvae of Neo-
chryopus. Afterwards, additional adult speci-
mens were recovered from the debris which
normally collects in the old dead leaf bases just
below the crowns of oil palms left by workers
engaged in cutting the bunches of fruit in a
grove of palms on the Agricultural Station. This
accumulation of rotten organic matter provides
a breeding site for Oryctes sjostedti Koble, and
Neochryopus breeds here and preys on these
dynastid grubs.

444

Rhinoceros Beetles in West Africa — Hoyt

445

Data obtained from the examination of field-
collected larvae of N. sawagei indicated that
there were five instars. The length of the fourth
instar period of a single larva provided with
a constant supply of small cetonid and Oryctes
grubs was slightly over 2 weeks. The pupal
period was 10 days. Several of these scaritid
larvae lived 3 weeks without food. Considerable
difficulty was experienced in hatching the eggs
laid by captive adult beetles and rearing their
larvae in the laboratory because both stages were
frequently attacked by fungus. In the case of the
larvae this fungus appeared to be a Metar-
rhizium.

The adult beetles mated readily in small ciga-
rette tins (14 cubic inches in capacity), which
were partly filled with moist rotten palm fiber.
The eggs of these scaritids were found in the me-
dium, there being one egg per beetle. In a single
instance a second egg was laid 3 days after the
first had been discovered. The eggs were oval
in shape, about 2 mm long and V2 mm wide.
When laid they were sticky, and quickly became
covered with bits of frass. This made them hard
to find and, as no special attempts were made to
discover them, no doubt many were missed.
Adult female beetles which were dissected con-
tained a maximum of four large eggs.

Both the adults and larvae of N. savagei
readily attacked, killed, and consumed cetonid,
Oryctes, and other dynastid grubs of a size up
to 10 g in weight (the size of an early third
instar Oryctes larva). The larvae of the palm
weevil, Rhynchophorus phoenicis F., were of-
fered to the adult beetles and while these were
usually killed, the beetles did not seem to feed
on them to any extent.

The adult scaritids survived for long periods
without food, the maximum period noted being
about 4 weeks.

The adult beetles are winged and are strong
fliers. Occasionally they are attracted to lights
at night.

The sexes are readily distinguishable. The
mandibles of the male are elongated, and the
distal portion is without well-developed hori-
zontal teeth. Those of the female are slightly
broader and shorter, and the left mandible has
a broad, flattish, horizontal inner tooth which
forms a subapical notch. The mandibles of the

males are apparently adapted for grasping the
female around the back of the head during
copulation. However, both sexes feed readily on
scarabaeid grubs.

The sex ratio was found to be 1 to 1. Usually
one or two adult beetles were found in the
debris of an oil palm, but on one occasion four
were taken in a single site. No figures were
obtained on the sexes collected from individual
palms; the ratios were calculated from total
field collections.

The majority of the beetles were collected
from the oil palm debris; however, a fairly large
number of both adults and larvae were also
found in the rotten Raphia trunks. The fauna
of the oil palm trash and of the standing rotten
trunks was, with one exception, identical. The
exception was Oryctes sjostedti, which was only
found breeding in the oil palms. However, it
seems safe to regard this site as a sort of elevated
trash heap rather than an unique environment.

At Umudike the beetles were kept in round,
50-cigarette-size tins. At first only one adult
was put in each tin, but later, to conserve space,
two were placed in each container. Rotten palm
fibre was added to each tin to give the beetles
something in which to dig. After about 1,400
specimens had been collected at Umudike, they
were taken to Ibadan and there packed for ship-
ment. Plywood trays divided by partitions into
2 -inch cubes were used to hold the beetles in
transit. Each cube was partly filled with moist
wood wool. A single scaritid was placed in each
compartment, and the tray was covered with

-inch-mesh wire screen held down with
staples. Three or more trays were then packed
into a heavy polythene bag to prevent loss of
moisture. The trays in their bags were packed
into heavy cardboard boxes which were wrapped
with paper. From 4 to 6 boxes made up a ship-
ment. The insects were sent from the Ikeja air-
port via Paris to Fiji by air freight. Cigarette tins
were used as containers for the first two or
three shipments, but these were heavier than
the plywood trays.

During the shipments, there were always
more beetles than containers. Usually, - enough
males were discarded to bring the sex ratio to
1 male to 3 females instead of the normal 1 to 1.
An ideal container would have been an alumi-

446

num box of about 8 cubic inches capacity, and
had these been available in quantity many more
insects could have been sent.

An attempt was made to keep N. savagei
adults together in a large tin, but this was un-
successful due to cannibalism even when 8 cubic
inches were allowed per beetle.

At first it was thought desirable to provide
food for the insects during the time they were
held in Nigeria. Later it was found that they
fared much better if they were not fed. The
grubs put into the tins were only partly con-
sumed, and the rotting remains in the small
containers evidently created a most unfavorable
environment.

Dorylus ( Anomma ) nigricans 111. (Hymenoptera,
Formicidae, Dorylinae)

Throughout the search for parasites and pred-
ators of Oryctes in West Africa it was noted
that the larvae of the rhinoceros beetles occurred
in large numbers in standing rotten palm trunks
but not in fallen logs. In Sierra Leone this situa-
tion was often observed, but no experiments
were made to determine why the fallen logs
were not used by the beetles. It was thought
that possibly the principal species encountered,
Oryctes owariensis Beauv., unlike other mem-
bers of the genus, did not favour such sites.

On Moor Plantation, Ibadan, Nigeria, an
experimental grove of coconut palms was found
to be infested by a fair number of Oryctes
monoceros Ol. This species was found breeding
inside the bases of old coconut palm stumps and
in standing rotten coconut trunks, the products
of several severe lightning strikes in the grove.
However, in the numerous rotten trunks which
had been felled, there were no Oryctes larvae
nor, for that matter, any larvae of any of the
numerous species of cetonids and small dynastids
which usually occur with Oryctes in West Africa.
There were clear signs that, before the trunks
had been felled, a considerable amount of breed-
ing by Oryctes had taken place within them. It
was also noted that the hollowed interiors of
these fallen trunks contained much less frass
than would have been expected.

Specimens of adult rhinoceros beetles from
this grove were sent to the British Museum
(Natural History) with a request that they be

PACIFIC SCIENCE, Vol. XVII, October 1963

compared with specimens of Oryctes monoceros
from East Africa. This was done, and the Ibadan
species was found to be identical with the East
African 0. monoceros .

To determine whether the fallen trunks were
suitable as breeding sites, four sections were
selected and set upright. After these had been
in position for about 3 weeks, they were split
open. Seven adult beetles of 0 . monoceros and
14 eggs were collected from them.

Five sections of the fallen trunks were care-
fully split lengthwise, and new hollows were
either carved out by hand or the existing hollows
were utilized. These prepared sections were
stocked with Oryctes larvae, and frass was
packed into the hollows around them. The two
halves were then brought together and fastened
in place with bands of wire. Some sections were
left lying on the ground, others were raised
slightly at one end. In addition to the logs, a
sawdust pile was established and stocked with
larvae of O. boas F. and a few O. owariensis .
For all of these trials only third-instar larvae
were available.

The following events took place:

1. 15 November 1959. One log was entered,
and the larvae were eaten by a small greyish rat.

2. 30 November 1959. All the logs were en-
tered, the frass was removed, and the larvae were
eaten by a swarm of "driver” ants, Dorylus
( Anomma ) nigricans.

3. 14 December 1959. The logs, having been
restocked with Oryctes larvae, were again en-
tered, the frass was removed, and the larvae
were eaten by a very large swarm of "driver”
ants, Dorylus ( Anomma ) nigricans. The saw-
dust pile was also entered by the ants and about
one-third of the grubs was eaten.

4. 10 January I960. Additional logs were
added on 3, 4 January and a small section of
log which could be moved about with ease was
prepared. This was placed in the dry bed of a
stream, about 50 m distant, where "driver” ants
had been observed. On the evening of 10 Janu-
ary the ants invaded this log and also entered all
the other logs in the experimental area. One
of these logs was constructed so that the larvae
were in a central frass-packed hollow which had
no openings at either end. The only means of
access to this hollow were the holes left in the

Rhinoceros Beetles in West Africa- — Hoyt

447

trunk by the workings of xylocopid bees, a
natural feature of all of the rotten logs in the
grove. The ants entered through these holes,
removed the frass, and ate the Oryctes larvae,
leaving only the head capsules.

5. 11 January I960 . The logs which had been
invaded by the "driver” ants on the night of the
10th were broken up to determine if any of
the Oryctes larvae had escaped. In three of the
logs in the upper breeding site experimental
area a few intact larvae and pupae were found,
but in the log located in the stream bed nothing
remained. The following results were obtained
from the logs in the upper area: One log which
had been stocked with 7 larvae contained 1 liv-
ing larva and 1 pupa; one log which had been
stocked with 10 larvae contained 2 living larvae
and 1 pupa; one log with no access from either
end which had been stocked with 6 larvae con-
tained 1 living larva and 1 pupa.

The area was visited daily and the logs were
inspected at intervals of 4 to 5 days to determine
if the larvae were alive. The last stocking of the
logs with larvae was on 16 December 1959.

In November, the end of the wet season in
Nigeria, the ants were active throughout the
area. However, after the end of the rains the
ants moved into stream beds; by January, only
invasions of short duration were made into
nearby areas.

The invasion of the experimental site on 30
November lasted about 14 hr. The last sortie
on 10 January was only about 7 hr long.

Another species of Dorylus ( Anomma ) was
found in the coconut grove; this ant was never
observed to enter the logs. No attempts were
made by either species of Dorylus ( Anomma )
to enter the tops of the stumps or to climb up
the standing trunks. The ants were able to climb
vertical surfaces but did not seem to do any
work, such as removing frass, while on them.
When D. nigricans entered the rotten logs, they
did so by choosing holes which were more or
less horizontal. They always used these holes
to bring out the frass from the interior.

Ochryopus gigas Schio. (Coleoptera, Caraboidea,
Scaritidae)

Occasionally the very large scaritid Ochryopus
gigas was found both in the debris accumulated

below the crowns of oil palms and in standing
rotten palm trunks. During April and May,
I960, adults of this species were taken around
lights at night. However, this species was never
found to be common and its larva was not dis-
covered. In the laboratory O. gigas attacked,
killed, and consumed both larvae and adults of
Oryctes.

Scolia sp. ( Hymenoptera, Scoliidae)

In the oil palm debris and in rotten Raphia
trunks large Scolia cocoons were frequently
found. Many of those taken from these sites had
already hatched. Sometimes the head capsule of
the host grub would be found entangled in the
silk of the cocoon, and from this it was de-
termined to be parasitic on Oryctes ohausi and

O. sjostedti.

The adult wasps were never observed in the
field, but two males and two females were bred
out in the laboratory. The female specimens
were sent to the British Museum (Natural His-
tory), where they were identified only as Scolia
sp., "not in British Museum collection.”

Judging from the number of empty cocoons,
this parasite was thought to be rather common,
but large-scale collecting efforts resulted in only
about 65, nearly half of which contained dead
pupae. Nevertheless, the cocoons were sent to
Fiji where an attempt was made to rear the
wasps. Unfortunately, this has proved unsuccess-
ful, and the wasps which did hatch died in the
laboratory.

OTHER PREDATORS AND PARASITES

1. Alaus ? sp., Calais sp. (Coleoptera, Ela-
teridae)

2. Morio guineensis Imh. (Coleoptera,
Caraboidea, Carabidae)

3. Genus ?, species ? (Diptera, Tachinidae)

4. Platymerus higuttata Stal (Hemiptera,
Reduviidae )

5. Genus? species? (Diptera, Tabanidae)

6. Genus? species? (Araneida, Aviculari-
idae)

1. Large elaterid larvae were found inside the
standing rotten oil palm trunks in close proxim-
ity to Oryctes larvae in both Sierra Leone and

448

PACIFIC SCIENCE, Vol. XVII, October 1963

Nigeria. They were never common, and in Sierra
Leone only 24 were collected from over 500
rotten oil palm trunks. Specimens were de-
termined at the British Museum to be perhaps
a species of Alaus.

A smaller elaterid larva found in a rotten
oil palm was reared on Oryctes and cetonid
larvae, the adult beetle being identified as Calais
sp.

2. Occasionally larvae and adults of a small
carabid beetle, Morio guineensis, were found in
the frass in the rotten palm trunks. In the
laboratory these readily consumed first instar
Oryctes larvae.

3. Large numbers of the larvae of Oryctes
boas F., collected from a manure heap on the
farm of the University College, Ibadan, were
reared in captivity. These were dissected when-
ever they showed signs that something was
wrong with them. One, a prepupal stage larva,
died suddenly and was cut open. In the abdomen
was a very large larva of a tachinid fly which
had also perished. Following this discovery, some
1,000 Oryctes boas grubs were collected from
this manure heap, but no other parasites were
recovered.

4. Neanides and adults of the large reduviid,
Platymerus biguttata, were found fairly fre-
quently in the open hollow tops of the standing
rotten oil palm trunks in Sierra Leone during
April, May, and June. Laboratory trials were
conducted to see if this bug would prey on adult
Oryctes owariensis, but without result. Even
adults which had been held without food for
nearly 2 weeks showed no interest in this large
rhinoceros beetle. During May and June, 1959,
Platymerus biguttata came to lights at night on
several occasions. No other Coleoptera than
Oryctes owariensis were tried as prey for this
reduviid.2

5. In a standing rotten coconut palm trunk
on Moor Plantation, Ibadan, three tabanid larvae

2 The author is at present engaged in a study of the
allied species, Platymerus rhadamanthus Gerst., which
does attack the adults of Oryctes monoceros Ol. How-
ever, old adults which have accumulated large fat
deposits usually refuse to attack anything. At the
time Platymerus biguttata was investigated it was not
known that the fifth-instar neanides would also at-
tack adult rhinoceros beetles and, perhaps because of
their more rapid metabolism, are generally easier
subjects with which to work.

were found in the frass along with the grubs
of Oryctes monoceros. These readily attacked
and killed young scarabaeid larvae in captivity.
A week or so later all three pupated and after
2 weeks the adult flies emerged. Unfortunately,
the specimens sent to the British Museum were
badly broken in transit and were not able to be
identified. The tabanids were bright orange with
a large purple band across each wing.

6. In Sierra Leone, both inside the hollow
tops of standing rotten oil palm trunks and in
the crowns of living oil palms, a large aviculariid
spider was frequently encountered. This fear-
some-looking arachnid evidently attacked the
oil palm climbers, who had their own remedy
for the effects of its bite. Experiments were
made to see if this spider would kill adult
Oryctes owariensis, but the results were negative.

In Nigeria this palm tarantula seemed to be
unknown to the people and no specimens were
found.

Coleolaelaps sp. (Acarina, Laelaptidae)

Several investigators have reported that mites
feed on the eggs of Oryctes. Venkatraman re-
ported a Coleolaelaps sp. which "apparently fed
on Oryctes eggs” in Ceylon. Surany reported
several instances of mites feeding on Oryctes
eggs and stated that they were important under
certain circumstances in its control. Unfortu-
nately, Surany did not identify the mite in
question in his report.3

Clusters of mites around eggs of Oryctes
owariensis were frequently observed in Sierra
Leone from June onward throughout the rainy
season in 1959. These mites were found on eggs
enclosed in balls of frass formed by the adult
beetles as they oviposited in the rotten wood
inside the standing rotten oil palm trunks. It was
noted at the time that eggs which had mites on
them never hatched, while eggs in the vicinity
without mites hatched readily. These mites were
collected and sent to the British Museum where
they were identified as Coleolaelaps sp.

Adult mites of this species were common on
the larvae of Oryctes owariensis , O. boas, and
O. monoceros, but were encountered only rarely
on the adult beetles.

3 Diseases and Biological Control in Rhinoceros
Beetles. South Pacif. Com. Tech. Pap. 128. I960.

Rhinoceros Beetles in West Africa — Hoyt

449

In Nigeria, Coleolaelaps was again found on
the eggs of Oryctes boas in March, I960, follow-
ing a few early rains. The Oryctes eggs were
taken in a large manure heap on the farm of
the University College, Ibadan. Mite-infested
eggs were collected and placed for observation
in covered shallow glass dishes filled with a
manure and frass mixture. At the same time
newly laid eggs were obtained from captive
specimens of Oryctes boas. Mite eggs and im-
mature and adult mites were transferred from
the originally infested, field-collected eggs to the
newly laid eggs and to uninfested, field-collected
eggs.

It was noted that Oryctes eggs which had
mites or mite eggs on them were all slightly
discoloured and at times showed brownish spots.

When mite eggs were transferred to non-
discoloured, field-collected eggs, or to newly laid
eggs, the mites hatched and in a short time
made their way throughout the medium to the
original egg from which they had been trans-
ferred. Here they gathered and fed on the yolk
of the now partly collapsed egg.

The adult Coleolaelaps were never observed
to feed on either nondiscoloured, field-collected
eggs or on the newly laid eggs, but they fed
readily on the discoloured ones and on a non-
discoloured egg which was punctured with a
needle.

None of the discoloured eggs hatched and,
with the exception of the punctured, nondis-
coloured egg, both newly laid and field-collected
ones produced normal larvae.

No further tests were carried out with Coleo-
laelaps due to the discovery of other predators
which appeared to be more promising.

OTHER MITES ASSOCIATED WITH Oryctes

In all, some 14 species of mites (Acarina,
Uropodina, and Canestrinidae ) were found as-
sociated with the larvae and adults of Oryctes
in Nigeria. Two of these, a canestriniid and a
species of the Uropodina, were found commonly
on the adults of Oryctes monoceros and O.
owariensis. The canestriniid was found in large
numbers on the first three abdominal tergites
beneath the elytra. The uropodinid was taken
wandering about both on the venter of the

beetle and under the wing cases. At times both
species of mites were found heavily infested
with a fungus resembling closely some of the
Laboulbeniales. The fungus did not appear to
harm them, and gravid females were frequently
observed to produce normal eggs in spite of
heavy infestations.

The species of Uropodina found on Oryctes
was also collected in numbers from the adults
of Neochryopus savagei.

SPECIES OF Oryctes ENCOUNTERED
( Coleoptera, Scarabaeidae, Dynastinae)

1. Oryctes monoceros Olivier

Both in Sierra Leone and in Nigeria Oryctes
monoceros was always found in places where
there were coconut palms in the immediate vi-
cinity. On Moor Plantation, Ibadan, this species
was taken breeding in standing rotten coconut
trunks and in the bases of coconut stumps. With
the exception of Moor Plantation, however,
O. monoceros was rare in all of the places visited.

2. Oryctes boas Fabricius

In Sierra Leone Oryctes boas was commonly
encountered breeding in village compost heaps,
but in similar sites in Nigeria it was rare. How-
ever, in Nigeria a large manure pile was found
at the farm of the University College, Ibadan,
in which there were thousands of larvae, pupae,
and adults of this species. In March and early
April numerous adults and eggs were found in
this site, but at other times mainly larvae of
various instars were present. These findings
seemed to indicate that, while there was some
overlap of generations, nevertheless there was
a definite yearly cycle.

Recently an experimental planting of oil
palms in Northern Nigeria was reported to be
rather severely attacked by an Oryctes , and
specimens extracted from the crowns of these
palms proved to be O. boas.

3. Oryctes owariensis Palisot de Beauvois

Oryctes owariensis is a large species, the well-
developed adults being larger than the other
Oryctes species in West Africa and larger than
0. rhinoceros of Southeast Asia and the Pacific

450

region. Probably because of its size, the species
has been confused with O. gigas of Madagascar.

Oryctes owariensis breeds, as far as is known,
only in rotten wood and prefers rotten palm
logs. It was found commonly in Sierra Leone
and Nigeria in standing rotten oil palm trunks,
standing rotten coconut trunks, and in rotten
Raphia palm trunks. Usually this was the only
species of Oryctes present in these sites, but
sometimes the larvae of O. monoceros or O.
ohausi occurred with it. In Sierra Leone an adult
specimen was extracted from the crown of an
oil palm. Observations in both Sierra Leone and
Nigeria seemed to indicate that this beetle did
not prefer coconut palms as a host plant.

4. Oryctes sjostedti Kolbe

As has been stated previously, this species
was only taken breeding in the debris accumu-
lated in the dead leaf bases just below the
crowns of oil palms. In this site the maximum
number of larvae found was six, all third in-
stars. Pupae were not infrequently encountered,
usually in a cavity within one of the old leaf
bases; the adult beetles were also found both
within these and in the debris enclosed by them.
Adults were taken at mercury vapour lights
fairly7 frequently, but never in numbers; one or
two per night was the largest catch. This species
was most common at Benin, Nigeria, on the
West African Institute for Oil Palm Research
station and at Umudike in eastern Nigeria.

5. Oryctes ohausi Minck

Perhaps this species has been confused with
O. monoceros in the past; anyway when speci-
mens were sent to the British Museum for de-
termination, it was surprising to learn that no
examples of this species were in the collection.
0. ohausi was commonly found breeding in the
standing rotten trunks of Raphia palms. Adults,
larvae, and pupae were found in fair numbers
in these sites, and not infrequently in company
with O. owariensis. There was an average of
eight larvae per trunk during April, I960, at
Umudike in eastern Nigeria. A single adult
male was taken in a rotten oil palm stump on
Moor Plantation at Ibadan.

PACIFIC SCIENCE, Vol. XVII, October 1963

6. Oryctes erehus Burmeister

This species has been recorded a number of
times from West Africa but was not found
during this investigation.

NOTES CONCERNING ADULTS OF Oryctes

All of the investigations conducted in West
Africa on the adults of the various species of
Oryctes were carried out with the objective of
finding parasites of the mature beetles.

The main difficulty in searching for parasites
of the adults is to get large numbers of beetles
which have been exposed in nature. When ma-
ture beetles are taken from rotten logs, a fairly
high proportion of them are recently emerged
and have never been outside the breeding site.
If Oryctes flew readily to lights, the problem
would be simple. However, it is not particularly
attracted by light, although recently some suc-
cess has been reported with a special ultra-violet
lamp used in the Pacific.

In Western Samoa split logs laid flat on the
ground are used to trap adult rhinoceros beetles.
Each trap consists of four to eight logs each
about 4 ft long. A similar type of trap was tried
in Nigeria without success.

Another type of rotten log trap was devised
which gave fairly good results in Nigeria. This
was made of a foot-tong section of coconut
trunk through which a hole was bored from end
to end. This section was placed on top of a
No. 10 can which was either sunk into or rested
on a coconut stump. The beetles landed on the
top of the log, crawled down the hole and fell
into the can underneath. The trap caught quite
a few Oryctes monoceros , the largest single catch
being three.

Efforts were made to use captive beetles to
attract others. No positive results were obtained
along this line, but it was found that 0. owari-
ensis had two distinct flight times, one in the
evening and the other in the early morning.
Furthermore, beetles which did not fly in the
evening flew in the morning, and beetles which
did fly after sundown did not fly later on.

No parasites were found in the adults of
Oryctes, but the numbers examined were prob-
ably not high enough to give any reasonable
hope of success.

Rhinoceros Beetles in West Africa — HOYT

451

DISCUSSION AND CONCLUSIONS

In both Sierra Leone and western Nigeria
predators and parasites or Oryctes larvae were
not at all common. The only controlling factor
of this nature seemed to be periodic invasions
of the fallen logs by "driver” ants. Standing
rotten palm trunks contained large numbers of
Oryctes in all stages of development. It should
not be supposed, however, that there were nor-
mally large numbers of these breeding sites in
any one small area. Further, not all of the trunks
contained Oryctes. It was noted that rotten
trunks with long black fibers in their interiors
seldom supported anything at all. Perhaps the
form of decay, fungal or bacterial, made these
unattractive to the beetles.

In areas like one reported in western Nigeria
where an old oil palm plantation had been poi-
soned off and the dead palms left standing,
serious damage was caused to the young palms
planted between them by Oryctes, which bred
in the rotten trunks.

Parasites and predators were found readily
only in the regions of the deltas of the Niger

and Cross rivers of eastern Nigeria. While the
same species of these are wide-spread through-
out West Africa, they were common in this
particular area. It is thought that the practice
of burning off the bush, which is done on a
large scale in western Nigeria and Sierra Leone,
plays an important role in limiting the numbers
of predators and parasites. The land of the delta
areas is too swampy for burning. Oryctes is not
affected by bush fires on account of its habits.

It is, of course, logical to assume that the
long dry season, together with the absence of
numerous streams and swamps, probably also
plays a major role in limiting populations of
parasites and predators which have fairly short
life cycles. No doubt it does, but this and burn-
ing complement each other, and it is difficult to
say which is the more important.

Under West African conditions it appears
that many of the populations of Oryct^-%. re
limited by the number of available and suitable
breeding sites, and that their predators and
parasites are limited by unfavorable environ-
mental conditions which are determined largely
by cultural practices and climatic conditions.

Characteristic Features of the Volcanism of the Siberian Platform

V. S. Sobolev1

Widespread volcanism is characteristic of the
Siberian platform, and was especially intense
during uppermost Paleozoic and lower Mesozoic
time. The Siberian traps, which occupy an area
of more than 1,500,000 km2, are best developed
there. These are effusive and hypabyssal rocks
of basalt-dolerite type, closely resembling trap
rocks in other parts of the globe, especially the
Karroo dolerites of South Africa.

The rise of trap magma began in the upper
Paleozoic (Permian or even as early as Upper
Carboniferous) time, and reached its climax in
the Lower Triassic period. It was accompanied
by the ejection of much pyroclastic material,
which formed a thick series of tuffs. Lava sheets
and hypabyssal intrusions of various kinds and
sizes were formed.

The process of volcanism was rather com-
plicated, and at present M. L. Lurie and V. Z.
Masaitis distinguish five volcanic phases and 13
separate intrusive complexes, each having its
own specific features and pattern of develop-
ment in various parts of the platform. In spite
of this, however, the magma had some charac-
teristic features over the entire area, notably an
iron content somewhat higher than is usual and
an especially rapid increase of relative iron con-
tent during the process of crystallization dif-
ferentiation. The increase of the iron content of
the femic minerals in the process of crystalliza-
tion prevails over the conventional reaction
series of Bowen. For example, olivine of early
formation contains about 20% fayalite, that
characteristic of the usual type of traps contains
about 40% fayalite, and the iron content of
olivine in pegmatoid veins is as high as 80%.

The residue of the differentiation is as a rule
micropegmatite, either in the mesostasis, or in
some cases forming veinlets of granite compo-

1Siberian Division, Academy of Sciences, Novosi-
bizsk, U.S.S.R.

Manuscript received April 19, 1962.

sition. However, such veinlets are quantitatively
very small. Much rarer is the formation of alka-
line rocks, such as teschenite, in the last stages
of differentiation.

The above characteristics of crystallization
differentiation (rapid change of the iron con-
tent of femic minerals and the subordinate role
of the discontinuous reaction series ) are typical
not only of the trap formations, but also of the
deeper-seated magmatic complexes of the plat-
form. The gabbro-anorthosite-granite complex
of the margin of the Russian platform, with
its characteristic granites of Rapakivi type, be-
longs to this group. These characteristic features
distinguish complexes of this type from the
typical granodiorite complexes of orogenic
zones, in which crystallization largely corres-
ponds to the well-known Bowen series. Relative
increase of the iron content of femic minerals
is there much slower, as is clearly seen by com-
paring it with the change in piagioclase com-
position.

Along with the trap formation, in part simul-
taneously and in part a little later, another type
of volcanism developed widely on the margin of
the platform, with the formation of ultrabasic
and alkaline rocks. Differentiated effusive and
intrusive complexes were formed in some re-
gions, kimberlites in others.

A typical example of the differentiated com-
plexes can be seen in the northern part of the
Siberian platform in the area of the so-called
Gulinski intrusion. The effusives range from
meimechite, the closest extrusive analogue of
true intrusive ultrabasite, to different kinds of
alkaline basaltoids containing nepheline and
piagioclase. Among the intrusives are all kinds
of rocks from dunite to various alkaline rocks
rich in nepheline. Carbonatite is also present.
E. L. Butakova (1956) and Y. M. Sheinman
(1955) have shown that the volcanic rock for-
mation here was largely simultaneous with erup-

452

Volcanism of the Siberian Platform — Sobolev

tion of the trap magma. There are traps both
older and younger than the alkaline rocks.

Kimberlite has been formed extensively in
the northeastern border of the platform, more
than 100 pipes and dikes being known there at
present. For some kimberlite bodies the same
sort of relationship to the traps has been es-
tablished as for the alkaline basaltoids. Some of
the kimberlite is not younger than Permian in
age, since pyrope and diamonds from it occur in
the Upper Permian deposits. However, there
undoubtedly are younger kimberlites also, for
a fragment of a belemnite of Upper Jurassic or
Lower Cretaceous age was found in one of the
pipes.

Kimberlite, as a magmatic rock, belongs to
the ultrabasic group, its composition ranging
from a form nearly devoid of alumina and alka-
lies to one rather high in AI2O3 and especially
high in K20 in mica-rich varieties. The prin-
cipal mineral is always olivine, containing up
to 10% Fe2Si04, and is present in at least two
generations: large crystals, commonly partly re-
sorbed, and small idiomorphic microphenocrysts.
Phlogopite occurs in idiomorphic plates and
ranges widely in quantity. It is unquestionably
magmatic. Pseudomorphs of pyroxene microlites
are sometimes seen in the vitreous matrix. The
latter is always -altered. In the northern regions
fine-grained monticellite has been found in kim-
berlite for the first time, chiefly in dikes. Nephe-
line also is supposed to be present. Pyrope, and
probably picroilmenite and chrome spinel, com-
monly belong to the first generation of pheno-
crysts. Perovskite is a later accessory.

As in South Africa, the kimberlite has a brec-
ciated structure and is contaminated by frag-
ments of various sorts of rocks. There are, on
the one hand, typical pyroclastic rocks filling
explosion pipes, and, on the other, magmatic
breccias with various contents of xenoliths. Since
the rocks have been altered ( serpentinized and
carbonatized ) , it is not always possible to prove
the presence of magmatic cement.

The fragments of other rocks may be sub-
divided into:

1) Fragments of ultrabasites and eclogites
whose origin is in some way connected with the
origin of the kimberlite itself;

2) Fragments of rocks picked up by the

453

magma from (a) the crystalline basement for-
mations, and (b) the sedimentary cover.

Fragments of the first type include various
ultrabasites — olivinites, peridotites, and others
- — often containing pyrope as well as typical
eclogites. The discovery of diamond-bearing ec-
logites, resembling the well-known eclogite
xenolith found by Bonney (1899) in South
Africa, is of particular interest. Together with
such eclogites brought up from great depth,
there are eclogites and eclogite-like rocks (con-
taining plagioclase) picked up from the crystal-
line basement and formed by eclogitization of
hypersthene schists.

The fragments of rocks picked up by the
magma vary widely in quantity and composition.
Xenoliths of gneisses and schists are abundant
in several of the pipes. This can be taken as
proof that the "explosion” that formed the pipe
took place at a level lower than the base of the
sedimentary cover. Allowing for this, and taking
into consideration the geophysical data on the
depth of the crystalline basement in the area and
also the thickness of the rocks since removed
by erosion, we can say that the explosion took
place at a depth somewhere between 2 and 4
km. The depth is greater than in the case of
the formation of the trap necks, which was 0.5
to 1 km.

As in South Africa, xenoliths of rock forma-
tions that occur at much higher levels (some-
times several hundred meters higher ) are found
among the fragments, proving that there was
not only an ascending but also a descending
movement of the material in the pipe.

The synchroneity of their formation has led
to the hypothesis of a genetic connection be-
tween the ultrabasic rocks and kimberlites and
the trap magma of the Siberian platform. Petro-
graphic data, however, do not support this hy-
pothesis. The olivine of the kimberlites and the
meimechites contains only 10% fayalite, and
this is proof enough that these rocks could not
have originated as a result of differentiation of
the trap magma. The author quite agrees with
Y. M. Sheinman’s (1955) suggestion of the
formation, in this case, of magmatic chambers
at much deeper levels than those of the trap
magma.

The development of trap volcanism on such

454

PACIFIC SCIENCE, Vol. XVII, October 1963

an enormous scale leads us to the conclusion that
regional melting of the basalt layer took place
here, probably in its upper part. However, no-
where in the platform did the magma chambers
reach the sialic shell, and the small granitoid
veinlets were formed wholly as a result of local
crystallization differentiation. Taking into con-
sideration the existing data on the structure of
the earth’s crust in the platform, the depth of
such magma chambers appears to have been
about 25 km, and the geothermal gradient at the
time of volcanism appears to have reached 40 C
per kilometer.

Some differences in composition of the traps
might have resulted from differences in depth
of the magma chambers in different parts of
the platform and resultant differences in the
differentiation phenomena.

The presence of effusives of ultrabasic com-
position ( meimechites, kimberlites) is definite
proof of the existence of a corresponding
magma. This magma could be formed only by
the remelting of ultrabasic rocks, which in turn
is proof of the existence of rocks of corres-
ponding compositions below the Mohorovicic
discontinuity.

In the case of the formation of differentiated
complexes, there is no doubt of the presence of
big magma chambers and a relatively slow rise
of magma either to the earth’s surface or to the
corresponding intrusion chambers. A compli-
cated evolution of the rocks takes place as a
result of involvement of the higher levels of the
earth’s crust in melting and, perhaps, as a result
of assimilation and differentiation.

In the case of the kimberlites the quantity
of rising magma is very small. This can hardly
be the result of the low penetrability of the
earth’s crust. Rather, it is a proof of the forma-
tion of very small magma chambers in which
remelting was partial, and a magma which rose
very rapidly up the deep fissures containing
many suspended crystals that formed not only as
a result of crystallization in the chamber but also
that remained as a result of the partial melting.

Not only theoretical calculations but also ex-
perimental data now show that if diamonds
were formed at a temperature of about 1200 C,
the pressure must have been more than 40 kilo-
bars. The notion that diamonds were brought

by the magma from great depth, and not formed
at the time of explosions near the earth’s surface,
can be considered valid. Sometimes, on the basis
of the above data, an attempt is made to de-
termine the depth of the magma chamber from
the implied hydrostatic pressure of the overlying
rocks. This approach we cannot agree with, since
the pressure in the earth’s crust, within the zone
of metamorphism, can be as high as 15 kilobars,
which is several times the pressure resulting
from load at that depth, the difference between
the pressure at the time of the mineral forma-
tion and the calculated pressure due to the load
being as much as 10 kilobars. Such zones of
higher pressure may extend into the depths of
the mantle, and it is to them that the regions
of kimberlite development are likely to have
belonged. The depth of formation of the magma
chambers, in this case, may be less than calcu-
lated— that is, not 150 km but 70 to 100 km.
As a result, when magma rises to a higher level,
pressure still remains high, though it falls by
a quantity corresponding to the weight of the
vertical column of magma. In places at a depth
of 3-4 km (see above), a breaking of the earth’s
crust occurred, accompanied by the formation
of peculiar pipes and a sudden pressure decrease,
constituting a kind of explosion. A great quan-
tity of pyroclastic and xenogenic material rushed
into the pipe, part of it being thrown up and
then sucked back into the pipe again. The frag-
ments filling the pipe may later be cemented
by the rising magma.

The pressure before the explosion is not only
below that shown by the equilibrium curve of
diamonds but also below that shown by the
curve of pyrope, since the kelyphite rims around
grains of the latter must have formed before
the explosion. The fact that the diamonds are
neither completely resorbed nor graphitized is
due to the rapid rise of the magma and its com-
paratively low temperature. The temperature of
the magma must be lower than that shown by a
curve corresponding to the region of metastable
existence of diamonds (V. Sobolev, I960),
which begins at 1200 C at normal surface pres-
sure and rises to 2200 C at 30 kilobars. As
is known, the diamonds show only traces of
graphitization, which appear as graphitic ro-
settes near some inclusions.

Volcanism of the Siberian Platform — Sobolev

Recently, in connection with the discussion
of the composition of the subcrustal substratum,
great attention is once more being paid to the
eclogite problem. The suggestion has been made
(Fermor, 1914; Lovering, 1958) that the sub-
stratum below the Mohorovicic discontinuity is
eclogite. Some authors are of the opinion that
this eclogite layer extends to a depth of 900 km
(V. V. Belousov, I960), i.e., to the base of the
Galitzin layer. The only valid data available for
discussion of this matter have been derived from
study of the kimberlites — a fact that makes it
desirable to treat this problem here.

As is known, a great quantity of xenoliths of
ultrabasic rocks, in some cases directly related to
eclogites, are to be found in many kimberlite
pipes. This fact suggests that such xenoliths are
at least in part the remains of the partially-
melted substratum, the more so as the compo-
sition of the olivine in them resembles that of
the first-generation olivine of the kimberlites.
The question is, however, still open to discus-
sion. There is some probability that these rocks
were picked up by the kimberlites during their
rise toward the surface, not only in the sub-
stratum but also at much higher levels. The
ultrabasic magma chambers are likely to have
revived several times, and the formation of
ultrabasite intrusions may have taken place dur-
ing the first stages, further intrusions taking
place later with the rapid movement of new
portions of magma in new geologic conditions.
Such intrusive massifs could have consisted of
pyrope peridotite, such as that in Czechoslovakia.

The absence of diamonds (at least in appre-
ciable quantity) in the ultrabasite xenoliths
speaks against the supposition that the xenoliths
were brought directly from the deep magma
chamber. Although the presence of diamonds in
pyrope peridotites has been asserted by some
workers, neither a xenolith with a diamond, nor
its photograph, nor a detailed description of it,
can be found anywhere. Many attempts to obtain
diamonds by grinding and concentration of con-
siderable quantities of ultrabasite xenoliths and
eclogites have resulted in failure. This fact, how-
ever, cannot altogether disprove the hypothesis
of the subcrustal origin of these rocks, since the
distribution of diamonds in the substratum may
be nonuniform. Also, they may have, for the

455

most part, crystallized directly with the forma-
tion of kimberlite magma.

On the other hand, two findings of diamond-
bearing eclogites, which are subject to no doubt
and have been described in detail, are proof of
the existence of subcrustal eclogites. Diamond
formation in the zone of metamorphism is im-
possible. The pressure there has never reached
even that of coesite crystallization, which is
lower than that of diamond formation. It is in-
teresting to point out that the Jakutian diamond-
bearing eclogite in its ratio of FeO to MgO is
nearer to basic rocks than to ultrabasic rocks;
it undoubtedly was not brought directly from
the deep magma chamber, but was picked up
from higher levels in the substratum.

Comparing all the above-mentioned data, we
come to the following conclusions regarding the
constitution of the upper mantle and its rela-
tionship to the earth’s crust:

1. At comparatively small depths, probably
less than 50-70 km, the subcrustal substratum is
of peridotite composition, corresponding ap-
proximately to the composition of meimechite
or kimberlite, the latter being a magmatic rock.

2. In the region of kimberlite distribution,
higher than the peridotite layer but below the
Mohorovicic discontinuity, the substratum is
eclogite with chemical composition very near
that of basalt.

3. Pressures in the zone of metamorphism can
vary greatly at one and the same depth, reaching
at least 15 kilobars but never 20-25 kilobars.

4. There is some reason to believe that higher
pressures are characteristic of large parts of the
earth’s crust, especially the border of the plat-
forms. Pressure higher than simple hydrostatic
pressure is also characteristic of the upper part
of the subcrustal substratum in these same areas
(probably down to a depth of about 150-200
km).

5. Higher pressure persisted in certain zones
through considerable periods of geologic time,
as is proved by the finding of ancient eclogitized
schist in kimberlite of both Upper Paleozoic
and Mesozoic age.

Examining the above statements, we come to
the conclusion that the hypothesis of a subcrustal
eclogite layer (Fermor, 1914) has been con-
firmed, but only in part. The author quite agrees

456

PACIFIC SCIENCE, Vol. XVII, October 1963

with the opinion of J. F. Lovering ( 1958 ) , V. V.
Belousov ( 1960b), and others, that a change of
conditions (chiefly pressure, rather than tem-
perature) leads to a shift of the Mohorovicic
discontinuity, with the formation of eclogites at
the expense of gabbroic rocks of the "basalt”
layer. The total thickness of the basic layer,
however, is probably not more than 30-50 km.
Thus the eclogite layer is not to be found all
over the globe, but only in the zones of higher
pressure. In some cases, the basalt layer has been
fully eclogitized and has entirely disappeared,
and the sialic layer has come into direct contact
with the Mohorovicic discontinuity. We can ap-
proach the problem of distribution of the sub-
crustal eclogite layer by comparing geophysical
and geologic data: the distribution of kimberlite,
the appearance of eclogite inclusions in effusives,
and, partly, the distribution of rocks formed at
high pressures in the zone of metamorphism,
such as kyanite schists, eclogites, jadeite (taking
into consideration possible changes over periods
of time).

In the zones of normal or lower than normal
pressure, the pressure of about 15 kilobars,
which is necessary for the formation of eclogites,
is reached at a depth of about 60 km, which is,
as a rule, below the boundary separating the
basic and ultrabasic rocks. Garnet peridotites,
or some interlayers of eclogites that are close to
ultrabasic rocks in composition, may be present
there. In such cases the Mohorovicic discon-
tinuity evidently corresponds to the true com-
positional border between the basalt and the
peridotite layers, not to a phase transformation.

The isobar of the limit of possible diamond
formation (40 kilobars) is, of course, well below
that level. In the areas of normal or lower pres-
sures, it must be below 120 km. In the authors
opinion, the penetration of magma from such
a depth is unlikely. Still less likely is the preser-
vation of the diamond, even if magma chambers
have formed at such depths.

In connection with this problem it is inter-
esting to compare the data on the finding of
diamonds in meteorites. As far as is known, dia-
monds have been discovered in stone meteorites
of ureilite type (first in the Novourei meteorite)
and in some iron meteorites. The author quite
agrees with the opinion of Urey (1954, 1957)

that the presence of diamonds there is evidence
of the formation of these meteorites by the
breakdown of some celestial body, as big as the
moon or bigger, in which the pressure was high
enough for the formation of diamonds. Thus,
we cannot agree with A. P. Vinogradov’s (1959)
opinion that achondrites were formed by the
breakdown of small celestial bodies. The para-
genetic associations characteristic of eclogites,
and specifically pyrope itself, have not been
found in the meteorites, however. This shows
that in the basalt shell of the disintegrated body
pressures nowhere reached 15 kilobars. This
fact, together with the absence of meteorites of
granitic composition, suggests that the body
probably was smaller than the earth and less
differentiated. This agrees with ideas which have
already been developed on other grounds by A.
N. Zavaritski ( 1943 ) . The above-mentioned
data do not, however, prove that all meteorites
have had the same origin and resulted from the
breakdown of one planet.

Various meteorites are still being searched for
diamonds. This search is certainly of great in-
terest, but there is little likelihood that diamonds
will be found in other types of meteorites, par-
ticularly in chondrites. Without going into de-
tails on the hypotheses of the formation of
chondrites, we are quite certain that in the later
stages of existence of these meteorites the tem-
perature was high enough so that diamonds
would have turned into graphite even if they
had existed. If, however, we should succeed in
finding pseudomorphs of diamonds in chon-
drites, as we have in some iron meteorites, this
would be a direct proof of the formation of
chondrites by the breakdown of some big celes-
tial body.

On the other hand, the discovery of graphite
pseudomorphs of diamonds in iron meteorites
shows that, after the breakdown of the parent
planet, the temperatures of the meteorites were
greater than 1200 G The preservation of dia-
monds in the Canyon Diablo octahedrite sug-
gests that the temperature of that meteorite at
the time of the breakdown was below 1200 C,
which means that it was not melted.

The general questions discussed here are, of
course, still open to argument. Already, however,
on the basis of available petrographic and min-

Volcanism of the Siberian Platform— -Sobolev

eralogic data, we can be more certain of the
thermodynamic conditions in and the constitu-
tion of the upper mantle of the earth, and of the
conditions of meteorite formation. The forma-
tion and alteration of diamonds are of particular
importance in these considerations. There is no
doubt that further mineralogic investigations in
general, and the investigation of diamonds in
particular, will open new approaches to the
study of the composition of the earth and aid
in penetrating the secrets of the solar system.

REFERENCES

Belousov, V. Y 1960a. Development of the
earth and tectonics. Sovetskaja Geologia 7.

— — 19601c Development of the earth and
tectogenesis. J. Geophysical Res. 65:4127-
4146.

Bgbrievich, a. P, M. N. Bondazenko, et al.
1959'. Diamond deposits in Yakutiya. U.S.S.R.,
Min. Geol. i Okhrany Medv., Moscow. 527 pp.

— — G. J. Smirnov, and V. S. Sobolev.
1959. Eclogite xenoliths with diamond. Dok-
lady Akad. Mauk., U.S.S.R.: 126(3).

BONNEY, T. G. 1899. The parent-rock of the
diamond in South Africa. Roy. Soc. London
Proc. 65:235-236.

457

Butakova, E, L. 1956. On petrology of ultra-
basic and alkalic rocks of Meimecha-Kotuockii
complex. Tr. Niga 86:6.

Fermor, L. L. 1914. The relationship of isostasy,
earthquakes, and volcanicity to the earths
infraplutonic shell. Geol. Mag. 51:65-67.

Lebedev, A. P 1957. Some problems of the
petrology of original diamond rock in U.S.S.R.
Akad. Nauk., U.S.S.R., Izvestiya, ser. Geol. 11.

Lovering, J. F. 1958. The nature of the Mo-
horovicic discontinuity. Am. Geoph. Union
Tr. 39:947-955.

Sheinman, Y. M. 1955. Some petrologic char-
acteristics of ultrabasic rocks, and ultrabasic
activity in platforms. Zapiski Ysesoyuznogo
Mineralogicheskogo Obshchestva.

Sobolev, V. S. 1936. Petrology of traps in the
Siberian platform. Arctic Institute, Trans. 43 :
224.

— — — 1960a. Role of high pressure in meta-
morphism. Int. Geol. Cong., 21st Session,
Rep. 1:14.

— I960 A Conditions of the formation of

diamonds. Geol. and Geoph. 1.

Urey, H. C. 1954. Diamonds, meteorites, and
origin of the solar system. Astroph. 129.

— - 1957. Diamonds in stone meteorites.

Geochim. et Cosmochim. Acta 13: 1-4.

Gibsmithia hawaiiensis gen. n. et sp. n.1

Maxwell S. Doty2

ABSTRACT: A new genus, Gibsmithia, is described and tentatively placed in the
Dumontiaceae of the red algal order Cryptonemiales. Its diagnostic features are:
possession of auxiliary cells in specialized filaments separate from those bearing
the carpogonia; the known sexual structures occurring in sori at the tips of soft,
gelatinous branches which arise from perennial round stems so as to form a hemi-
spherical head; cruciate tetraspores borne on filaments protruding from the surface
of the branches and these same branch filaments often bearing terminal seirospores.
The type species of the genus is G. hawaiiensis , known only from the island of
Oahu in the Hawaiian Islands.

THE ALGA described here was first collected by
Dr. J. T. Conover in 1943, and from 1951 on
has been found by eight other individuals col-
lecting in Hawaii. It seems to comprise a new
genus of the Cryptonemiales in the Rhodo-
phyta, and we take pleasure in naming it in
honor of the late Prof. G. M. Smith, of Stan-
ford University, whose publications and teach-
ings have probably done more to facilitate work
with the algae in America than has the work
of any other phycologist.

Gibsmithia hawaiiensis genus et species nova
Figs. 1-17

DESCRIPTIO TYPI: Thalli ramis gelatinosis
fasciculatis, e stipite erecto fere cartilaginose
lignoso egredientibus, formam hemisphaericam
simulantes. Frondes e filamentis uniseriatis,
sicut in genere Callithamnio, compositae. Fila-
menta interdum pilis uniseriatis, multicellulari-
bus e superficie gelatinosa ramorum exstanti-
bus terminata. Tetrasporangia cruciformiter di-
visa, subsphaerica, in pilis supra descriptis uni-

1 Contribution No. 190 from the Hawaii Marine
Laboratory. Received for publication February 26,
1962. This study was made possible by funds from the
U. S. Atomic Energy Commission (contract AT-f04-
31-15) and the Graduate Research Committee of the
University of Hawaii. Dr. Hannah Crossdale prepared
the Latin diagnosis.

2 Department of Botany, University of Hawaii,

Honolulu 14, Hawaii.

lateraliter in cymulis fortasse velut in genere
Spermothamnio producta, Cystocarpia mo do
Cryptonemialium simili crescentia.

The holotype (Fig. 1) was found drifting
in the water at Waikiki, Honolulu, Oahu, Ha-
waii (21° 161 min N, 157° 49.5 min W), on
January 12, I960; 19263.3 It is cystocarpic. The
type material, consisting of a liquid-preserved
isotype specimen and a dried holotype her-
barium specimen, is being deposited in the Ber-
nice P. Bishop Museum in Honolulu, Hawaii.
Tetrasporic material has been collected from
the vertical surface of ■ a dredge-cut in the reef
at Waikiki, first on January 5, 1961, by Dr.
D. W. Strasburg. This specimen (8525) is also
being deposited in the Bishop Museum her-
barium. Other collections are listed below.

DESCRIPTION OF GENUS AND SPECIES: Thalli
(Figs. 1, 2), as generally collected, consisting
of 'woody’ stems and a number of soft, but not
slippery, gelatinous ultimate branches. Forming
bright-pink, hemispherical tufts often 8 cm
high, the stems are up to 3 mm in diameter,
blackish when dried, and they bear rings be-
lieved to be indicative of successive crops of
the ultimate branches. The stems are cylindrical
or, when large, compressed toward the apex
and sometimes branched once or twice. When
the stems are fresh they are colored approxi-
mately Eugenia Red or darker to Acajou Red

3 Such 4- and 5 -digit numbers are the collection or
herbarium numbers of the author.

458

Gibsmithia hawaiiensis — Doty

459

FIG. I

CVI

FIGS. 1-2. Gross aspects of Gibsmithia hawaiiensis (type specimen 19263). I, Whole thallus illustrating
the rounded stem with growth rings. 2, A few of the soft gelatinous branches enlarged to show asymmetrical
placement of the sexual sori and the superficial filaments which, in tetrasporic thalli, would bear the tetra-
spores and seirospores. Below is shown some of the stem material of very different, almost woody, nature and
the origins and rebranching of the ultimate branches of the thallus.

(Ridgway, 1913). The soft, gelatinous, ultimate
branches are lighter red in the same series or
their bases are lighter and their apical portions
darker. The branches are often up to 4 cm
long, and they gradually enlarge apically until
they are club-shaped and up to 1 cm in diam-
eter near their tips. These ultimate branches
rarely arise singly from the parent stem; more
often (Fig. 2) they arise from a disk that is
a flattened expanse of the same gelatinous na-
ture and which divides somewhat dichoto-
mously into the branches.

The gelatinous branches are made up of cal-
lithamnioid filaments (Fig. 3). The exact na-
ture of the central axis of the thallus has not
been determined, but the alga is believed to
be multiaxial in structure. The major axes in
the medulla bear, most often, opposite branches
which seem to arise near the apical cell. Toward
the exterior the branching of the filaments is
pseudodichotomous, and the surface of a gela-
tinous branch is generally covered densely, espe-
cially toward the tip, with free tufts of such
filaments.

Branches of the axial filaments in the medulla

often give rise to recurrent branches (A in
Figs. 3-5) or rhizoids, which anastomose with
a particular cell of another branch (Fig. 3),
the supporting cell of the branch (Fig. 4), or
with the next cell below in the same branch
(Fig. 5). Pit connections appear between all
cells concerned in these cases. In content, the
cells of such anastomosing branches, like those
reported and figured by Feldmann-Mazoyer
(1940: 142, 393, fig. 48) in Pleonosporium
horreri, are not particularly different from the
cells of similar nearby filaments.

Reproductive structures of three kinds have
been found in the collections studied. These
are interpreted as seirospores, as tetrasporangia,
or as various stages and structures leading
toward carpospore production. The same thal-
lus may produce both tetraspores and seiro-
spores, a situation I have seen in Seirospora,
and which is reported ( Feldmann-Mazoyer,
1940) in Dohrniella. The evidences of sexuality
have been seen positively in only the type and
two other collections. Unfortunately the critical
stages involving the primary and secondary
connecting filaments between the carpogonia

460

PACIFIC SCIENCE, Vol. XVII, October 1963

and auxiliary cells have not been found. Thus,
though for the present Gibsmithia seems cer-
tainly to be a member of the Dumontiaceae as
this family is circumscribed by Kylin (1956),
the description of the sexual reproduction and
discussion of the systematic disposition within
the family are of necessity brief.

The seirospores are borne in unbranched
chains at the tips of the exterior filaments in
the asexual material. At some place along a
branch bearing seirospores (B in Fig. 5) the
chromatophore and central vacuole disappear
rather abruptly and the cytoplasm ( after killing
in Karpetchenko’s seawater fluid) is then more
densely granular and with a refractory body
becoming conspicuous as the cells age. The
spores, which are about 9 ^ in diameter, appear
to become more spherical with maturity, as in
Seirospora. Their ultimate fate is unknown, but
they are broken off easily in preserved material
and presumably in nature.

The subspherical tetrasporangia appear on
the more basal portions of scattered external
filaments similar to those bearing the seiro-
spores. They are, for the most part, in acro-
petalous series (Fig. 6) on the adaxial side
of the filaments on which they are produced.
In a few cases immature tetrasporangia were
seen below an otherwise acropetalous series. In
some instances what appeared to be chromo-
some groups undergoing meiosis were distin-
guished within immature tetrasporangia. The
tetrasporangia are apically dehiscent, as in
Antith amnion, and at least the terminal sporan-
gia were pedicellate. The first division of the
cytoplast in the tetrasporangium (Fig. 7) is
near the equatorial plane. Usually the tetra-
spore pairs are at right angles to each other,
i. e., decussately cruciate (Figs. 8, 9); only
rarely is a somewhat tetrahedral (Fig. 10) or
other ( Fig. 11) arrangement seen. The tetra-
spore groups in the collection 12343 are often
as long as 25 or 30 /i in their major dimension.

The maturing cystoearps, when stained, ap-
pear as dots in raised asymmetrically placed
welts (Figs. 1, 2) near the tips of the branches.
The largest thalli collected, 10386, were be-
lieved to be sterile until smears made of the
branch tips were examined. When this was
done what are accepted as carpogonial branches

(Figs. 12-14) bearing nonfertilized carpogonia
were found.

The carpogonial branch of Gibsmithia is the
terminal portion of a modified member of the
two lateral branch systems arising (Fig. 13)
from a cell in a medullary filament. Upon oc-
casion both lateral branch systems bear carpo-
gonia; less frequently only one of the mem-
bers is carpogonial, and the other is vegetative
but reduced in its development. Only mono-
carpogonial branch systems have been seen.
Branching from the cells below the special
cells of the carpogonial branch is variable. In
some cases there are no such branches (Fig.
12), and in others (e. g., Figs. 13, 14) up to
three branches of one to six small cells each
occur. The cells of these branches, except for
size, are of the same appearance as the homol-
ogous cells in vegetative branches nearby.

The carpogonial branch itself is consistently
composed of five cells ( cp, a, b, c, and d in Figs.
12-14), though the terminal carpogonium is
(Fig. 14) not always present. The antepenulti-
mate cell (b in Figs. 12, 13) is smaller than
the other hypogynous cells. Its contents are
often of different appearance and it is flattened,
so that its axial length is noticeably less than
its diameter. The other three hypogynous cells
are more spherical, similar in content, and the
largest one (c in Figs. 12, 13) is usually about
9 to 13 v in diameter. The remaining two
(a, d in Figs. 12, 13) are usually similar in size.

The carpogonium is borne on the first4 hypo-
gynous cell excentrically, often by a displace-
ment from the axis of the carpogonial branch
of about 45 degrees. The carpogonium {cp in
Figs. 12, 13) is usually relatively small and
does not stain strongly in aniline-blue, in con-
trast to the other cells of the carpogonial branch.

In the material available, stages were not
found which would enable a satisfactorily com-
plete description of sexual reproduction. How-
ever, my impression is that the heavily aniline-
blue stained condition of almost all carpogonial
branches seen is a result of development be-
yond the fertilizable point. Most such branches
consist of either only the cells a through d ( Fig.
14) or have in addition what are interpreted
as remnants of a carpogonium. Such carpo-
gonial branches are relatively straight and not

Gi bsmithia hawaiiensis — Doty

461

FIGS. 3—11. Vegetative and asexual reproductive microscopic structural details of Gibsmithia hawaiiensis
(collection No. 12343). 3, An axial filament from the medullary region of a gelatinous branch of the thallus
showing the opposite ramification of the medullary filaments and, A , characteristic reconnecting rhizoidal
branches. 4, A more simple reconnecting rhizoidal branch, A, from one of the subdichotomous cortical fila-
ments. 5, A seirospore-bearing ultimate branchlet on a subdichotomous cortical filament with, A, a very simple
reconnecting filament or rhizoid. Such lateral nonrebranched filaments, B, with a terminal series of seirospores
were found arising from almost every second cell in the vegetative axis of this part of the filament system.
The origins of but three such seirosporic branches are shown here. 6, A tetrasporangium-bearing cortical fila-
ment system. Divisions within the tetrasporangia are not shown and no attempt was made to represent all pit
connections. 7—11, Tetrasporangia in various stages of maturity. The arrangement of spores is always at first
cruciate but, as indicated by Figure 10, in age they may not be in a strictly cruciate arrangement.

IQOjj 5QJJ

FIG. 16 FIGS. 12-15,17

FIGS. 12-17. Microscopic sexual reproductive features of Gibsmithia hawaiiensis. The specialized cells,

a-d, are similar in nature whether in a carpogonial or auxiliary-cell branch. On the latter, the piliform exten-
sion, h, shows as a multicellular hair, the cells of which are in staining reaction and appearance, rather like
vegetative cells of similar size and relative terminal position elsewhere. 12-14 , Carpogonial branches with or
without nonfertilized carpogonia, cp. The hypogynous cells a-d were always densely granular and their contents
took up aniline-blue from aqueous solution much more than did the other nearby cells. Cell b was usually
smaller and the contents were usually different from the rest. Cell c was almost always the largest. The num-
bered cells, 1, 2, 3, etc., represent cells that looked like the strictly vegetative cells of similar size and position
nearby and connected the carpogonial branch to a medullary cell, med. The cytologically similar cells of the
sterile branchlets are indicated as la, lb, lc, etc. (Collection No. 10386.) 15, A nonfertilized auxiliary-cell
branch arising from one of two opposite branch systems of the same medullary cell, med, the one vegetative
branch system, veg, being of only three cells, the other more extensive and bearing an auxiliary-cell branch.

Gibsmithia hawaUensis—DoTY

463

strongly curved apically, as is usual in the Du-
montiaceae. In a very few cases carpogonium-
bearing branches that are more curved are
found with hypogynous cells that are less highly
differentiated in staining quality and in form.
In these cases the hypogynous and carpogonial
cells are more nearly similar in staining quality
and are smaller; such (Fig. 12) are judged to
be more nearly normal.

Primary connecting filaments4 were not seen,
and no clues as to the origin of the abundant,
presumed secondary, connecting filaments that
connect to the auxiliary cells were found. In a
very few cases the carpogonium as well as the
cell beneath it were seen to have some indica-
tion of protuberances extending toward one
another or toward the smallest of the hypogy-
nous cells. This could have been the effect of
smearing the material for microscope examina-
tion.

The auxiliary-cell branches (Fig. 15) develop

4 The author is attempting to apply terms to the
post-fertilization morphological phenomena in such a
way that they are consistent with general biological
usage, and so that the frequent special cases and varia-
tions from the normal do not require a special no-
menclature. Thus, the term "zygote” is used for the
post-fertilization carpogonium and its contents, and
"primary connecting filament” for the connection be-
tween the zygote and the auxiliary cell without dis-
tinguishing whether a "carpogonial” or "generative”
auxiliary cell is concerned. "Secondary connecting
filament” is used to refer to any connecting filament
("ooblast” of the older nomenclature) arising from an
auxiliary cell. Similarly, the term "hypogynous” is
used for any of the specialized cells of the carpogonial
branch other than the carpogonium itself; these are
referred to as a, b, c, etc., or the "first,” "second,” etc.,
proceeding away from the carpogonium.

farther from the apex than the carpogonial
branches. It may be that initiation, or at least
maturation, of an auxiliary-cell branch system
is dependent upon fertilization, for in 10386,
where many nonfertilized carpogonial branches
were found, especially of the forms illustrated
in Figures 13 and 14, branches bearing mature
auxiliary cells were not observed. They are
usually in the same positions as carpogonial
branches, or, at times, appear to have developed
(Fig. 16) after the lateral branch system had
grown further.

While the complement of cells in the auxil-
iary-cell branches is similar in number, form,
arrangement, and appearance to the hypogynous
cells in a carpogonial branch, there are two
major differences: in place of the carpogonium
there is a reduced terminal branch or row of
vegetative cells, and there are more sterile fila-
ments developed laterally from and near the
base. The auxiliary cell itself is an intercalary
cell, and, contrary to the condition illustrated
in Figures 15 and 16, is usually the cell there
labeled as b.

The presumed secondary, connecting fila-
ments ( cn in Fig. 16) were seen fused with
the auxiliary cells (c in Fig. 16). The tissue
of the weltlike sori (Fig. 2) in which cysto-
carps develop is the toughest of all parts of
the branch surface and this toughness seems to
be related to the great abundance of sterile
branches appearing in the fertile system, as well
as to numerous, presumed secondary, connect-
ing filaments. In addition, multicellular rhizoi-
dal branches from the bases of the branch
filaments (rh in Figs. 15, 16, and possibly A

This particular auxiliary-cell branch would seem to be peculiar, for usually one of the two cells labeled b
and c is about half the size of the cells labeled a and d, which are similar in size. The other cell, c or b, is
usually distinctly the largest of the four. A rhizoidal branch, rh, arises from the single vegetative cell, l, con-
necting this auxiliary-cell branch system to the medullary system. (Collection No. 19263.) 16, A lateral branch
system arising as one of a pair, B and C, from a single medullary cell, med, and producing vegetative filaments,
fil, and rhizoidal filaments, rh, in a normal manner, and showing (at c) a fertilized auxiliary cell. Two other
types of filaments are illustrated here: what may be labeled as an accessory rhizoid, arh, of multicellular
nature arising near a fertilized auxiliary cell and, ac, an accessory connecting filament. The auxiliary cell, c, is
shown with a connecting filament, cn, associated. The actual relationships of cells, c, cn, and ac, to one another
were not entirely clear at the highest magnifications. (Collection No. 19263.) 17, A young gonimoblast

showing the apparent carposporogenic cells heavily stippled. Cell 1, seemed to correspond to cells B and C in
Figure 16. Cell d bore three vegetative branches in addition to the auxiliary-cell branch. No connecting fila-
ment could be distinguished. Seemingly, the auxiliary cell, b, had produced the gonimoblast to the left at the
tip of the expansion, p, from but one gonimoblast initial. The six cells to the right seemed unattached and
were interpreted as having been broken free from the mass at the left. (Collection No. 19263.)

464

PACIFIC SCIENCE, Vol. XVII, October 1963

in Fig, 3) are abundant, and undoubtedly con-
tribute to the strength of the otherwise weak,
gelatinous, branch substance.

The surface of the auxiliary cell (b in Fig.
17), beneath the connecting filament, appar-
ently develops a protrusion that moves the con-
necting filament away from the auxiliary-cell
branch axis. From this protuberance of the
auxiliary cell (p in Fig. 17) gonimoblast ini-
tials arise. While the gonimoblast, upon smear-
ing, reveals its fundamentally filamentous na-
ture (Fig. 17), in the undisturbed condition it
is a dense almost spherical mass. Older gonimo-
blasts have one (or a few) very large cells at
the center: all the other cells appear to be uni-
form, as though all could become carposporan-
gia in time.

MATERIALS EXAMINED (all from the island
of Oahu, Hawaii; the collection numbers are
the authors, except the first): J. T. Conover,
200 Berkeley,5 tetrasporic, found about 550
ft along a line extending about 113° from the
old marine laboratory at Waikiki on underside
of a coral head about halfway out on the reef
at a depth of 1.2 m, VIII-43; D. W. Strasburg ,
8525 (bishop) tetrasporic and seirosporic,
University of Hawaii old beach laboratory, Wai-
kiki, Honolulu (21° 16.2' N, 157° 49.6' W),
1-51; M. Doty, 8562 (Stanford), another
branch of the same thallus as 8525, but col-
lected 25 days later; George Ikeda, 10385
(AUSTRALIA) tetrasporic, Lae o Kaoio, Kua-
loa, Kaaawa, one of two clumps seen in 5 ft
of water, 11-52; George Ikeda, 10386 (Stan-
ford, BERKELEY) carpogonial, in 4-5 ft of
water on limestone bottom (same area as Ikeda
collection of 11-52, above), V-52; Jack Randall,
17020 (BERKELEY) seirosporic, drifting freely
below the surface off Oahu, Hawaii, VI-52;

6 Names appearing in this position refer to the fol-
lowing repositories of the particular material : bishop,
B. P. Bishop Museum, Honolulu, Hawaii; STANFORD,
Hopkins Marine Station of Stanford University, Pa-
cific Grove, Calif.; BERKELEY, Herbarium of the
University of California, Berkeley, Calif.; PARIS,
Laboratoire de Cryptogamie du Museum National
d’Histoire Naturelle; AUSTRALIA, Herbarium of the
University of Adelaide, Adelaide, S. Australia; HOK-
KAIDO, Herbarium of the Faculty of Science, Hokkaido
University, Sapporo, Japan; VANCOUVER, Herbarium
of the University of British Columbia, Vancouver,
British Columbia.

M. Doty, 12343 (Stanford) tetrasporic,
floating freely at the surface just north of the
mouth of Koloa stream south of Laie, 1-54; M.
Doty, 12286 (hokkaido) first of a series
of collections from the same tetrasporic thallus
growing about 7 ft below the surface along the
inshore edge of the swimming channel in front
of the new University of Hawaii beach labora-
tory at Waikiki, Honolulu, XII-52; M. Doty,
12652 and 12647 (PARIS) tetrasporic, both
same thallus as 12286, about X-54; 17019
(bishop) sterile, same thallus as 12286, IH-
55; Jan New house and Malvern Gilmartin,
12671 (VANCOUVER) tetrasporic, beyond the
reef, Waikiki, Honolulu, XI-54; HOLOTYPE, M.
Doty and B. C. Stone, TYPE 19263 (BISHOP)
cystocarpic, drifting at the surface, Waikiki,
Honolulu, (near 21° 16' N, 157° 49.5' W),
1-60; Elvin Fong, 19478 (Stanford) cysto-
carpic and tetrasporic, on a 3 by 4 ft rock in
15 to 20 ft of water 150 yd off Haleiwa Army
Beach near the channel at Haleiwa, VI-55.

This is one of the less common algae occur-
ring along the shores of Oahu in Hawaii. It has
been found adrift in a few cases, but for the
most part it is found growing L to 2 m below
low tide on reef flats. The hemispherical pale
thalli were often found growing in a concavity
or on a vertical surface. One individual thallus,
near the University’s aquarium and beach lab-
oratory at Waikiki in Honolulu, was watched
and pieces were harvested from it at irregular
intervals for about 3 yr. The deepest collection
was that made by Jan Newhouse and Dr. Mal-
vern Gilmartin from the base of a coral patch
on rock bottom in 7 m of water about 1000 m
beyond the reef opposite the Natatorium at
Waikiki.

The stem, the perennial nature of the thallus
and some of the reproductory features of Gib-
smithia show some resemblance to the much
recopied small portion of Gmelin’s specimen
(1768: pi. 5, fig. 2) of Constantinea 6 (e. g.,
Kylin, 1956: fig. 102).

It is to be noted that at least three collections
of Gibsmithia, including the type, were of free-
floating branches. It would appear that the

w The figure given by Okamura (1912: pi. 77, figs.
1, 2) conveys a rather different conception of this
alga— a more correct one it would seem.

Gibsmithia hawaiiensis — Doty

465

branches are deciduous at maturity, as is the
disklike peltate blade of Constantinea simplex.
However, the many quite different features of
Constantinea, especially as revealed by Masaki
(1952), indicate that Constantinea could be,
at best, but remotely related to Gibsmithia.

Some of the anatomical features, the seiro-
spores, and tetrasporangia are reminiscent of
the Ceramiaceae. Yet I believe Gibsmithia
should be placed among the primitive Crypto-
nemiales for several reasons, most notable of
which is the occurrence of the carpogonia and
the auxiliary cells in separate but similar spe-
cial branch systems. In their general morphology
and in having a small differentiated cell among
the enlarged hypogynous cells, the carpogonial
branches of Gibsmithia (if, indeed, we have
seen normal carpogonial branches) recall both
the carpogonial and auxiliary-cell branches of
Dudresnaya crassa , as I have seen them in ma-
terial collected by Dr. A. J. Bernatowicz in
Bermuda (his number 51-580), and as they
are illustrated by Taylor (1950: figs. 5-7, 38)
in his splendid account of the reproduction of
that species.

The terminal row of sterile cells distinguishes
an auxiliary-cell filament from a carpogonial
branch with its terminal trichogyne. While the
terminal row of sterile cells is distinct from
the terminal part of a normal vegetative branch,
it is less specialized than the "hair” to be seen
in Dudresnaya crassa and which Taylor ( 1950:
figs. 36, 37 ) refers to as a "piliform extension.”
In other features Gibsmithia also recalls such
a genus as Dudresnaya rather than any of the
polycarpogonial forms discussed by Norris
(1957), which are far more complex. It is like
Thuretellopsis (Kylin, 1925: 14) in that the
carpogonial and auxiliary-cell branches are
alike; but in Gibsmithia they are less compact
and less specialized. In having a terminal row
of sterile cells and in having a larger number
of cells, the auxiliary-cell filaments of Gib-
smithia would seem to be more primitive than
those of Thuretellopsis.

A point of some systematic interest is the
development of low, wartlike sori containing
the cystocarps. The tetrasporangia are borne
quite differently, being exposed as in the Cera-
miaceae. These two features seem to set this
genus apart from most of the other genera now

placed in the Dumontiaceae. The possession of
cruciately divided tetrasporangia further distin-
guishes Gibsmithia from the type genus of the
Dumontiaceae and likens it to families such as
the Squamariaceae of the Cryptonemiales which
have cruciate tetrasporangia and the reproduc-
tive structures in nemathecia. Until the simpler
cryptonemiaceous families are better known, it
seems best to place Gibsmithia tentatively
among the Dumontiaceae.

REFERENCES

Feldmann-Mazoyer, G. 1940. Recherches sur
les Ceram iacees de la Mediterranee Occiden-
tale. Imprimerie Minerva, Alger. 540 pp.

Gmelin, S. M. 1768. Historia fucorum. Typo-
graphy Academiae Scientiarum, Petropoli.
12 + 239 + 6 pp. + illus.

Hollenberg, G. J. 1959. Smith ora, an inter-
esting new algal genus in the Erythropelti-
daceae. Pacif. Natural. 1(8): 3-11.

Kylin, H. 1925. The marine red algae in the
vicinity of the biological station at Friday
Harbor, Washington. Lunds Universitets
Arsskrift, n. f. Avd. 2, Bd 21., Nr 9., 87 pp.

1956. Die Gattungen der Rhodophy-

ceen. Gleerup, Lund. Xv + 673 pp.

Masaki, T. 1952. Studies on the reproductive
organs of the red algae. I. Constantinea rosa-
marina (Gmel.) Post, et Rupr. and C. subu-
lifera Setchell. Bull. Jap. Soc. Sci. Fish. 18:
30-38.

Norris, R. E. 1957. Morphological studies on
the Kallymeniaceae. Univ. Calif. Publ. Bot.
28:251-333.

Okamura, K. 1912. leones of Japanese algae.
Privately published by the author, Tokyo.
Vol. II, 191 pp. + illus. + index.

R IDG WAY, R. 1913. Color standards and color
nomenclature. Private publication by the au-
thor, Washington, D. C. 43 pp. and 53 color
pi. of 115 different colors. (Though the
book is dated 1912, according to D. H. Hamly
[Science 109:605], it appeared in early
1913.)

Taylor, W. R. 1950. Reproduction of Dudres-
naya crassa Howe. Biol. Bull. 99:272-284.

Revision of the Genus Pandanus Stickman, Part 16
Species Discovered in Thailand and Vietnam

Harold St. John1

The following new species are all but one
based upon specimens collected by others and
preserved in Bangkok, either in the herbarium
of the Department of Agriculture at Kasetsart
University or in the herbarium of the Royal
Forestry Department.

SECTION Aero stigma

Pandanus biplicatus sp. nov. (sect. Aero stigma)
Fig. 192

DIAGNOSIS HOLOTYPI: Licet acaulescens, foliis
1.01 m longis 3.6 cm latis parte lA basali sub-
coriacea parte ultima crassiter chartacea supra
viridibus infra pallidis et licet glaucis 1-sulcatis
acute 2-plicatis in sectione mediali M-formatis
et cum 41 nervis parallelis secundariis in quoque
dimidio, nervis tertilis in parte 2A ultima con-
spicuis et reticulis anguste oblongis formantibus,
laminis ligulatis subiter in apice trigono subu-
late 3 cm longo 1-2 mm lato diminuentibus
basi amplexicauli et inermi sed ex 14-15 cm
marginibus cum aculeis 1.5-2 mm longis 2-8
mm separatis subulato-serratis complanatis lu-
teis, midnervo infra ex 20 cm cum aculeis 2-3.5
mm longis 8-15 mm separatis subulatis reflexis,
in sectione mediali marginibus cum aculeis
1.5-2 mm longis 5-10 mm separatis subulatis
proxime adscendentibus, midnervo infra cum
aculeis 2.5-3 mm longis 15—22 mm separatis
subulatis adscendentibus, in apice subulato mar-
ginibus et midnervo infra cum serrulis 0.2-0.3
mm longis 1.5-3 mm separatis, in pagina superi-
ori ad apicem plicis cum serrulis 0.2-0. 3 mm
longis 1-4 mm separatis, inflorescentia foeminea
erecta et recta, pedunculo 22-30 cm longo gracili
clavato 9-12 mm diametro in apice folioso brac-
teato, bracteis herbaceis coloratis sed in sicco

1 B. P. Bishop Museum, Honolulu 17, Hawaii,

U.S.A. Manuscript received September 28, 1961.

nigro-brunneis infima 19 cm longa 23 mm lata
lineari-lanceolata ad basem integra parte XA ul-
tima proxime subulato-serrata, syncarpio 9.5 cm
longo 7.2 cm diametro obovoideo-ellipsoideo
cum drupis numerosis, eis 32-34 mm longis
6-7.5 mm latis 4-6 mm crassis anguste oblanceo-
fusiformibus 1-carpelatis parte lA supera libera
corpore 24-26 mm longo oblanceoloideo lateri-
bus planis, pileo 13-14 mm longo cartilagineo
basi 5-6 mm alta rotundato-pyramidali 6-angu-
lata, stylo 6-8 mm longo crassiter subulato
anguloso divergent! subsinuoso, stigmate 3-4
mm longo distali lineari brunneo sulcato papil-
loso in apice extenso, endocarpio in parte }A>
infera obovoideo cartilagineo stramineo in apice
truncato lateribus 0.1 mm crassis, semine 8-9
mm longo obovoideo, mesocarpio apicali 7 mm
longo cum medulla alba delicata completa, meso-
carpio basali 5-6 mm longo fibroso et carnoso.

diagnosis of holotype: Perhaps acaules-
cent; leaves about 1.01 m long, 3.6 cm wide,
the lower lA subcoriaceous, the outer 2A thick
chartaceous, green above, pale and apparently
glaucous below, 1 -furrowed above the midrib,
sharply 2 -pleated, at midsection M-shaped in
section and with 41 parallel secondary veins in
each half, tertiary cross veins conspicuous in
outer 2A’s , forming narrow oblong meshes, the
blade ligulate, abruptly narrowed to a trigonous
subulate apex 3 cm long, 1-2 mm wide, the
base amplexicaul and unarmed, but beginning at
14-15 cm the margins with prickles 1.5-2 mm
long, 2-8 mm apart, subulate-serrate, flat, yel-
lowish; the midrib below beginning at 20 cm
with prickles 2-3.5 mm long, 8-15 mm apart,
subulate, reflexed; at midsection the margins
with prickles 1.5-2 mm long, 5-10 mm apart,
subulate, closely ascending; the midrib below
with prickles 2.5-3 mm long, 15-22 mm apart,
subulate, ascending; on the subulate apex the
margins and midrib below with serrulations
0.2-0. 3 mm long, 1.5-3 mm apart; on upper

466

Fig. 192. Pandanus biplicatus St. John, from holotype. a, Syncarp profile, X 1; b, drupe, lateral view, X 1;
c, drupe, longitudinal median section, X 1; ^ drupe, apical view, X 1; e, drupe, lateral view, X 4; j, drupe,
longitudinal median section, X 4; g, style and stigma, distal view, X 4; h, leaf base, lower side, X 1; h leaf
middle, lower side, X 1; j, leaf apex, lower side, X 1; k, leaf apex, upper side, XI \ U young pistillate
inflorescence, lateral view, X Vl-

468

PACIFIC SCIENCE, Vol. XVII, October 1963

side towards the apex the pleats with serrula-
tions 0.2-0. 3 mm long, 1-4 mm apart; pistillate
inflorescence erect, straight; peduncle 22-30 cm
long, slender clavate, 9-12 mm in diameter at
apex, leafy bracted; floral bracts herbaceous, col-
ored, drying blackish brown, the lowest one 19
cm long, 23 mm wide, linear-lanceolate, entire
below, the outer p3 closely subulate-serrate; syn-
carp 9.5 cm long, 7.2 cm in diameter, obovoid-
ellipsoid, with numerous drupes, these 32-34
mm long, 6-7.5 mm wide, 4-6 mm thick, nar-
rowly oblance-fusiform, 1 -celled, upper free,
the body 24-26 mm long, oblanceoloid, the sides
flat; pileus 13-14 mm long, cartilaginous, the
base 5-6 mm high, rounded pyramidal, 6-angled,
tapering upwards; style 6-8 mm long, stout
subulate, angled, divergent, slightly sinuous, but
nearly straight; stigma 3-4 mm long, distal,
linear, brown, creased, papillose, running to the
apex; endocarp in lower obovoid, the apex
truncate, cartilaginous, stramineous, the walls
0.1 mm thick; seed 8-9 mm long, obovoid; api-
cal mesocarp 7 mm long, filled with a delicate
white pith; basal mesocarp 5-6 mm long, fi-
brous and fleshy.

HOLOTYPUS: Thailand, Lower Siam, Kopah,
Jaujau, 9 Dec. 1917, Md. Haniff & Md. Nur
2,703 (SING).

discussion : P. biplicatus is a member of the
section Aero stigma, as is its closest relative, P.
pseudofoetidus Martelli of Burma, a species
which has the syncarp subglobose; drupes 20
mm long, 5 mm wide; endocarp 10 mm long;
and the apical mesocarp 4 mm long. P. bipli-
catus has the syncarp obovoid-ellipsoid; drupes
32-34 mm long, 6-7.5 mm wide; endocarp 13
mm long; and the apical mesocarp 7 mm long.

The new epithet is from the Latin, bi, twice,
plicatus, folded, in reference to the two longi-
tudinal pleats of the leaves.

Pandanus monotheca Martelli, Soc. Bot. Ital.,

Bui. 303, 1904; Webbia 4(2): pi. 41, fig.

18-20, 1914.

THAILAND: Bachaw, Pattani, evergreen forest,
600 m. alt., July 14, 1923, A. F. G. Kerr 7J91
(bk). This species, originally described from
Malaya, is here recorded for the first time for
Thailand.

Pandanus Toei sp. nov. (sect. Acrostigma)
Figs. 193, 194

NOM. vern.: "toei.”

DIAGNOSIS HOLOTYPI: Acaulescens, foliis 1.3
m longis in media 20 mm latis proxima basem
15-18 mm latis supra midervum sulcatis
2-plicatis chartaceis supra viridibus infra palli-
dioribus ligulatis in apice 9 cm longo 0. 3-0.8
mm lato subulato trigono in sectione mediali
cum 26-29 nervis secundariis parallelis in quo-
que latere, nervis tertilis non evidentis basi
amplexicauli et inermi sed ex 4. 5-6. 5 cm mar-
ginibus cum aculeis 1.3-2 mm longis 3-7 mm
separatis subulatis adscendentibus luteis, mid-
nervo infra ex 6-8.5 cm cum aculeis 1.5-2. 5 mm
longis 6-15 mm separatis crassiter subulatis
arcuatis reflexis, in sectione mediali marginibus
cum serrulis 0.2-0. 6 mm longis 3-7 mm sepa-
ratis, midnervo infra cum aculeis 2 mm longis
remotis subulatis arcuatis reflexis vel adscenden-
tibus, in apice subulato marginibus et midnervo
infra cum serrulis 0.2 mm longis 0.5-2 mm se-
paratis, inflorescentia foeminea terminali erecta
cum syncarpio unico, pedunculo 15 cm longo
clavato trigono in apice 5 mm diametro in di-
midia ultima cum bracteis foliosis paucis eis su-
peris 40-50 cm longis 15-18 mm latis, syncarpio
3.5 cm diametro globoso viridi cum circa 160
drupis, eis 15-18 mm longis 4-5 mm latis 3-4
mm crassis subfusiformibus parte 2A supera li-
bera corpore oblanceo-ellipsoideo obtuso 10-12
mm longo, pileo 9-11 mm longo basi 2-3 mm
alto oblato-pyramidali-hemispherico laevi 5-7-
anguloso angulis in stylo continuis, stylo 5-7
mm longo valido 5-7-anguloso subulato proxime
arcuato parte exteriori osseosa lucida lutea, stig-
mate 2.5-3 mm longo distali sublineari sulcato
brunneo papilloso paene ad apicem extento, en-
docarpio in tertia infera cartilagineo luteo muris
0.1 mm crassis intra lucidis, semine 6-7 mm
longo 2-2.5 mm diametro oblanceoloideo trun-
cato, mesocarpio apicali 2 mm longo hemisphe-
rico vel oblato-hemispherico et cum membranis
medullosis albis, mesocarpio basali fibroso et car-
noso.

DESCRIPTION OF ALL SPECIMENS EXAMINED:
Acaulescent, or decumbent with the stem 20
cm tall, 12-22 mm in diameter, brown, smooth;
leaves 0.96-1.3 m long, 20 mm wide near the
middle, 15-18 mm wide near the base, 1-fur-

Fig. 193. Pandanus Toei St. John, from holotype. a , Peduncle and syncarp, X 1; ^ drupe, lateral view, X
1; c, drape, apical view, X 1; ^ drape, longitudinal median section, X 1; drupe, lateral view, X 4; f,
drape, longitudinal median section, X 4; g, style and stigma, distal view, X 10; h, leaf base, lower side, X 1;
i, leaf middle, lower side, X 1; h leaf apex, upper side, X 1; k, leaf apex, lower side, X 1.

5 »nnv.

FIG. 194. Pandanus Toei St. John, from Smitinand 1,391. a, Staminate inflorescence, X 1; b, axis and
stamens, X 10; c, leaf base, lower side, X 1; ^ leaf middle, lower side, X 1; e> leaf apex, upper side, X 1;
f, leaf apex, lower side, X 1; g> secondary longitudinal veins and tertiary cross veins, at midsection, lower
side, X 4.

Page 309: Revision of Pandanus, 16. Thailand, Vietnam— St. John

471

rowed over the midrib, 2 -pleated, chartaceous,
green above, paler green below, ligulate, taper-
ing to a 9 cm subulate, trigonous apex 0.3-0.8
mm wide, at midsection with 2,6 -29 secondary
parallel veins in each half, the base unarmed,
but beginning at 4. 5-6, 5 cm the margins with
prickles 1.3-2 mm long, 3-7 mm apart, subulate,
ascending, yellowish; the midrib below begin-
ning 6-8.5 cm up with prickles 1.5-2. 5 mm
long, 6-15 mm apart, stout subulate, fully re-
flexed arcuate; at midsection the margins with
serrulations 0.2-0.6 mm long, 3-7 mm apart,
the midrib below with remote prickles 2 mm
long, subulate, arcuate, reflexed or ascending; on
the subulate tip the margins and midrib below
with serrulations 0.2 mm long, 0.5-2 mm apart;
pistillate inflorescence terminal, erect, with 1
syncarp or rarely with a small secondary one;
peduncle 15 cm long, clavate, trigonous, 5 mm
in diameter at apex, with a few leafy bracts on
upper half, the upper bracts 40-50 cm long,
15-18 mm wide; syncarp 2. 5-4.5 cm in diame-
ter, globose to ellipsoid, green, bearing about
160 drapes, these 15-18 mm long, 4-5 mm
wide, 3-4 mm thick, subfusiform, upper 2A free,
the body oblance-ellipsoid, obtuse, 10-12 mm
long; pileus 9-11 mm long, the base 2-3 mm
high, oblate pyramidal-hemispheric, smooth,
5-7-angled, the angles continuing well up the
style; style 5-7 mm long, stout, 5-7-angled, su-
bulate, proximally arcuate, the outer part bony,
shining, yellowish; stigma 2.5-3 mm long, distal,
sublinear, creased, brown, papillose, running al-
most to the apex; endocarp in lower 14, carti-
laginous, yellowish, the walls 0.1 mm thick, the
inside shining; seed 6-7 mm long, 2-2.5 mm in
diameter, oblanceoloid, truncate; apical meso-
carp 2 mm long, hemispheric or depressedly so,
containing white, medullary membranes; basal
mesocarp fibrous and fleshy.

STAMINATE PLANTS: Prostrate, stems 7-11
mm in diameter, very short; leaves 80-90 cm
long, 18 mm wide at the middle and base, the
subulate apex 3-4 cm long, at midsection with
24-27 parallel secondary veins in each half;
staminate inflorescence almost erect, 18 cm long,
fragrant; peduncle 7 cm long; flowering part
11 cm long, with conspicuous yellow bracts, the
lowest one 9.5 cm long, 16 mm wide, linear

lanceolate, chartaceous, serrulate only on the
short subulate tip; median bract 6 cm long, 14
mm wide, lanceolate, the margins of the outer
half and the midrib below near the tip with
pale, fragile serrulations 0.1 mm long; spikes
about 7 and 1.5-2 cm long, 8-15 mm in diame-
ter, dense; stamens numerous, attached singly,
directly to the axis; filaments 0. 1-0.2 mm long;
anthers 6-7 mm long, linear, bearing at the apex
a projection of the connective 0.2-0.3 mm long,
sometimes subulate but more commonly flat,
lanceolate, obtuse.

HOLOTYPUS: Thailand, Peninsular, Krabi, 83
km. from Huay Yawt, very common in moist
localities by roadside, 100 m. alt., 28 Jan. 1958,
T em Smitinand 4,1 43 ( BKF ) .

SPECIMENS EXAMINED: Thailand, Trat, Koh
Chang, shrub in evergreen forest by stream, 40
m. alt., 19 Feb. 1954, ( BKF ) ; erect shrub 40 cm.
tall, common in swampy ground in evergreen
forest, 1 June 1952, Bunnak ( Sang Kachand )
333 (BKF); Trat, Huayraent, Tern Smitinand
1,392 (BKF); Peninsula, Trang, Kantang, Khuan
nang hong, less than 100 m. alt., common in
evergreen jungle, 26 Aug. 1955, Tem Smitinand
3,012 (BKF); S. E. Part, Trat, Kaw Kut, Ao
Salat, common in evergreen forests, 5 m. alt., 5
April 1959, Tem Smitinand 3,702 , staminate,
( BKF ) ; S. E. Part, Trat, Huang Raeng, Dong
Tapit, prostrate, common in swamp by path in
evergreen jungle, under 50 m. alt., 21 June 1952,
Tem Smitinand 1,391, staminate, (bkf no.
7,276); Kasvon, Nov. 1896, C. Curtis 3,247
(SING).

MALAYA: Penang, near Tanjong Toking, July
1889, C. Curtis 1,821 (SING); Perak, Lahat,
Ipoh, Oct. 1904, H. K.^Riidley'], (sing),, one
of the original paratypes of P. perakensis RidL

DISCUSSION: P. Toei is a member of the sec-
tion Aero stigma, as is its closest relative, P, Kin-
gianus Mar tell i of Malaya, a species which has
the drupes 11-13 mm long; style 5 mm long;
pileus rounded; and the leaves 40-50 cm long,
14 mm wide. P. Toei has the drupes 15-18 mm
long; style 5-7 mm long; pileus prominently
angled; and the leaves 96-130 cm long, 20 mm
wide.

The new epithet is the vernacular name of
the plant in its native region in Thailand.

Fig. 195. Pandanus calcis St. John, a-e, h-j, from holotype; f—g from Kerr 13,358. a, Syncarp, X 1;
drupe, lateral view, X 1; c, drupe, longitudinal median section, X 1; ^ drupe, apical view, X 1; ^ drupe,
median transverse section through horizontal plate, X 4; f, drupe apex and stigma, proximal view, X 4; g, i
drupe apex and stigma, lateral view, X 4; h, leaf base, lower side, X 1; i, leaf middle, lower side, X 1; j,
leaf apex, lower side, X 1.

Page 311: Revision of Pandanus , 1 6. Thailand, Vietnam -St, John

473

section Micro stigma

Pandanus catch sp. nov. (sect. Micro stigma)
Fig. 195

nom. verm.: "lang kai.”

DIAGNOSIS HO'LOTYPI : Truncus 3 m alms ra-
mosus eo ramisqne cum "spinis brevibus arma-
tis,” foliis 75 cm longis 2-2.5 cm latis tenuibus
sed subcoriaceis infra subpallidioribus supra sub
catis marginibus subrevolutis in sectione rnediali
cum 32 nervis secumdariis in quoque latere ligu-
latis sed sensim in apice subulate longe diminu-
entibus eo 10 cm ex apice 4 mm lato basi
exarmatis sed ex 4 cm marginibus cum serris
fortibus arcuatis subulatis 1-2 mm longis 5-15
mm distantibus toto obscure brunneis, midnervo
infra inermi, in sectione medialis marginibus
cum dentibus 0.2-0. 5 mm longis 6-8 mm dis-
tantibus subulatis toto adpressis adscendentibus
brunneis, midnervo infra cum dentibus simu-
lantibus 3-8 mm distantibus, circa apicem mar-
ginibus cum serris 0.2-0.3 mm longis 4-9 mm
distantibus brunneis, midnervo infra cum ser-
rulis simulantibus 13-15 mm distantibus,
pedunculo 7 cm longo 7 mm diametro trigono
folioso, syncarpio subgloboso 6 cm diametro ter-
minali solitario erecto cum 26 drupis eis 3. 2-3. 4
cm longis 1.7-2 cm latis 1.7- 1.8 cm crassis (sed
in gemino 2.6 cm latis) ellipsoideis in sicco
brunneis obovoideis sed basi cuneata lateribus
laevibus sublucidis 5-anguiatis angulis subrotun-
datis parte Vz supera libera, apice rotundato sed
biangulatis, stigmate 3 mm longo excentrico
obdeltoideo cordato sulcato obscure brunneo pa-
pillose valde oblique et proximo, endocarpio
supramediali osseoso obscure brunneo 12-13
mm longo lateribus 2 mm crassis, semine 6 mm
longo adamantine sed trilobate, mesocarpio
apical i plerumque medulloso sicco, mesocarpio
basali fibroso et medulloso.

DESCRIPTION OF ALL SPECIMENS EXAMINED:
Trunk 3-5 m tall, branching, it and the branches
"armed with short spines”; leaves 75-84 cm
long, 2-2.7 cm wide, rather thin but subcoria-
ceous, slightly paler beneath, above with a cen-
tral furrow and the margins gently down-rolled,
at midsection with 32 secondary veins in each
half, ligulate but gradually long tapering to the
subulate apex which 10 cm down is 2-4 mm
wide, the base unarmed for 4 cm then the mar-

gins with heavy arcuate subulate serrations 1-2
mm long, 5-15 mm apart, wholly dark brown,
the nearby midrib below unarmed; at the mid-
section the margins with the' teeth 0.2-0. 5 mm
long, 6-8 mm apart, subulate, fully appressed
ascending, brown; the midrib below with simi-
lar teeth 3-8 mm apart; near the tip the margins
with brown serrations 0.2-0.3 mm long, 4-9
mm apart, the midrib below with similar ser-
rations 13-15 mm apart; peduncle 7 cm long,
7 mm in diameter, 3 -angled, leafy bracted; lower
bracts 25-33 cm long, 15 mm wide, distended
and paler at base; syncarp subglobose 6-7 cm
in diameter, terminal, solitary, erect, bearing
20-26 drupes, these 3.2-3 .8 cm long, 1. 7-2.1 cm
wide, 1.7—1. 8 cm thick, (but in a twinned one
2.6 cm wide) , ellipsoid, when dried brown, obo-
void but the base cuneate, the sides smooth,
somewhat shiny, 5-6-angled, the angles more
rounded than sharp, upper Vz free, the apex
rounded, but two lateral angles produced into
the sharp ridges curving and leading to the ex-
centric stigma; stigma 2. 5-3. 5 mm long, obdel-
toid cordate or oblong cordate or lance-cordate,
creased, dark brown, papillose, sharply oblique
and proximal; endocarp supramedian, bony, dark
brown, with two vertical columns near the cen-
ter, 12-13 mm long, a narrow transverse median
plate connecting to the marginal vertical flanges
2 mm thick; seed 6 mm long, diamond-shaped
but with 3 ascending lobes; apical mesocarp -
mostly of dry, membranous pith; basal mesocarp
fibrous and pithy.

HOLOTYPUS: Thailand (Siam): Kao Tala,
Ranawng, on limestone rocks, 200 m. alt., Feb.
3, 1927, A. F. G. Kerr 11,796 (bk).

SPECIMENS EXAMINED: Thailand, Kaw
Wieng, Chumpawn, limestone rocks, 50 m. alt.,
Jan. 11, 1927, A. F. G, Kerr 11,373 , nom. vern.
"chan deng,” (BK); Ban Kawp Kep, Surat, form-
ing pure growth on top of limestone hill, 100
200 m. alt., Aug. 16, 1927, Kerr 13,338 (bk);
Sapli, Chumpawn, Sept. 8, 1927, Put, A. F. G.
Kerr 1,018 (bk) ; Kantuli, Surat, Sept. 10, 1931,
Put, A. F. G. Kerr 4,189 (bk).

DISCUSSION: P. edicts is a member of the
section Microstigma, as is its nearest relative,
P. utilissimus Elmer, of the Philippines, a species
which has the syncarp 60 cm long, 20 cm in
diameter; drupes 4.5-7 cm long, linear-oblan-
ceoloid, the sides semiviscid, the upper VS free;

FIG. 196. Pandanus bifdus St. John, from holotype. a, Drupe, proximal view, X 1; b, drupe, longitudinal
median section, X 1; c, d, lateral drupes, apical view, X 1; e, lateral drupe and stigma, proximal view, X 4;
/. apical drupe, lateral view, X 1; g, h, apical drupes, apical view, XI \ h apex of apical drupe and stigma,
oblique view, X 4; j, leaf base, lower side, X 1; ^ leaf middle, lower side, X 1; h leaf apex, lower side,
X 1;». margins of base of mature leaf, of Kerr 20,084, X 1.

Page 313: Revision of Pandanus , 16. Thailand, Vietnam- St. John

475

endocarp inframedian, lacking distinct marginal
flanges; bark grayish, unarmed; and the leaves
5 m long, 8-20 cm wide, the apex abruptly short
acuminate. P. calcis has the syncarp 6-7 cm in
diameter, subglobose, drupes 3. 2=3. 8 cm long,
not viscid, the upper Vl free; endocarp supra-
median, with large lateral flanges; bark armed
with short spines; and the leaves 75-84 cm
long, 2=2.7 cm wide, gradually long tapering to
the subulate apex.

The new epithet is from the Latin, calx , lime,
and is given in reference to the habitat of the
species, the side cliffs or summits of precipitous
limestone ridges.

SECTION Rykia

Pandanus bifidus sp. nov. (sect. Rykia)

Fig. 196

nom. vern.: "t6i noi.”

DIAGNOSIS HOLOTYPI: Planta subacaulis, fo-
lds 1.3 m longis 3.3 cm latis subcoriaceis
gladiatis in sectione mediali cum 32 nervis
secundariis parallelis in quoque latere in apice
subulato breve diminuentibus et 10 cm ex apice
14 mm latis proxima basem marginibus cum
aculeis 3.5=4 mm longis 25-35 mm distantibus
subulatis adpressi-adscendentibus stramineis,
midnervo infra cum aculeis 2.5-3 mm longis
50-58 mm distantibus simulantibus sed valde
reflexis, in sectio mediali ; marginibus et mid-
nervo infra cum aculeis 2.5-3 mm longis 15-48
mm distantibus adscendentibus, proxima apicern
marginibus et midnervo infra cum serris 0.8-1. 5
mm longis 2=7 mm distantibus, syncarpio soli-
tario terminal! 12 cm longo 10 cm diametro ap-
parente ellipsoideo cum drupis numerosis eis
3-3.2 cm longis 8-10 mm latis 8-11 mm crassis
oblongo-oblanceoloideis 5-7-angulosis lateribus
subplanatis, parte Va supera libera, pileo 4-5
mm alto semiorbiculari 5-7-anguloso sublaevi
tarde deciduo, stylo 3.5-5 mm longo valido
osseoso corniformato eis lateralibus proximo-
curvatis irregulariter bifidis vel in eis apicalibus
integris et subulatis, stigma d 3-4 mm longo
brunneo papilloso proximo lanceolate vel bifido,
endocarpio mediali osseoso obovoideo acumi-
nate pallido sed intra obscure brunneo lucido
lateribus 1 mm crassis, seminibus 12 mm longis
obovoideis, mesocarpio apicale cavernoso, meso-

carpio basali fibroso et carnoso.

DESCRIPTION OF ALL SPECIMENS EXAMINED:
Trunk from almost none to 3 m tall; leaves
1.3-3 m long, 3.3-5 cm wide, subcoriaceous,
gladiate, at midsection with 32 secondary veins
in each half, tapering to a short subulate apex
which 10 cm down is 14 mm wide, near the
base the margins with prickles 3.5-4 mm long,
20-35 mm apart, subulate, appressed ascending,
stramineous; the nearby midrib below with
prickles 2.5-3 mm long, 50-58 mm apart,
similar but sharply reflexed; at the middle the
margins and midrib below with prickles 2.5-3
mm long, 15-48 mm apart, ascending; near
the apex below finely reticulate, and the margins
and midrib below with serrae 0.8-2 mm long,
2-17 mm apart; syncarp solitary, terminal, 12
cm long, 10 cm in diameter, apparently ellip-
soid, bearing very numerous drupes, these 3-3.2
cm long, 8-10 mm wide, 8-11 mm thick, ob-
long-oblanceoloid, 5-7-angled, the sides almost
plane, upper Va free; pileus 4-5 mm high, semi-
orbicular, 5-7-angled, nearly smooth, finally de-
ciduous; style 3.5-5 mm long, stout, bony, horn-
like, the lateral ones sharply curved towards
apex of syncarp, irregularly bifid, or on the
apical ones entire and subulate; stigma 3-4 mm
long, brown, papillose, proximal, lanceolate or
bifid; endocarp median, bony, obovoid, with an
apical acumination, pale except for the dark
brown, shiny inner surface, the lateral walls 1
mm thick; seed 12 mm long, obovoid; apical
mesocarp cavernous; basal mesocarp fibrous and
fleshy.

HOLOTYPUS: Thailand (Siam): Pak Tong
Chai, Nakawn Rachsima (=Korat), marshy
ground, 200 m. alt., Dec. 27, 1923, A. F. G. Kerr
8,132 (bk).

SPECIMENS EXAMINED: Thailand (Siam),
Kao Krading, Loi, evergreen forest, 1,200 m.
alt., Feb. 11, 1931, A. F. G. Kerr 20,084 (bk).

DISCUSSION : P. bifidus St. John is a member
of the section Rykia. Its closest relative is P.
furcatellus Martelli from Tonkin. This has the
endocarp in the lower Vdl apical mesocarp cav-
ernous, 10-11 mm long; and the style forks
ovate, acute. In contrast P. bifdus has the endo-
carp median; apical mesocarp cavernous, 4 mm
long; and the style forks subulate.

The new epithet is the Latin, bifdus, two-
cleft, in reference to the cleft style.

“T

to cm.

0 3 cm.

1 » i t_ i i

Fig. 197. Pandanus bipollicaris St. John, from holotype. a, Drupe, lateral view, X 1; b, drupe, longitudinal
median section, X 1; c~i> lateral drupes, apical view, X 1; k—n, apical drupes, apical view, X 1; o, drupe,
lateral view, X 4; p, drupe, longitudinal median section, X 4; q, apex and stigma of lateral drupe, proximal
view, X 4; r, apex of lateral drupe, apical view, X 4; s, apex and stigma of apical drupe, proximal view, X
4; t, leaf base, lower side, X 1; u, leaf middle, lower side, X 1; ^ leaf apex, lower side, X 1; w, in upper
third of leaf, lower side, secondary and tertiary veins, X 4; z, in upper third, secondary and tertiary veins,
upper side, X 4.

Page 315: Revision of Pandmus , 16. Thailand, Vietnam— St. John

477

Pandanus bipollicaris sp. nov. (sect. Rykia )
Figs. 197, 205 a, b

DIAGNOSIS HOLOTYPI: Acaulescens caespi-
tosus 80- cm diametro, foliis 2-3 m longis 4-5
cm latis supra obscure viridibus lucidisque infra
viridibus vel luteo-viridibus supra sulcatis 2-
plicatis in sectione mediali cum 36 nervis
secundariis parallelis in quoque latere, nervis
tertiis transversis multis et in sicco conspicuis
et reticulis quadratis vel oblongis formantibus,
laminis ligulatis sensim per 40 cm ultimas in
10 cm apice subulato trigono contract is eo
circa 10 cm ex apice 2 mm la, to, basi amplexi-
cauli inermf luclda cupracea et ex 12-15 cm
marginibus cum aculeis 3-6 mm longis 10-
20 mm separatis crassiter subulatis subarcuato-
adscendentibus stramineis vel nigris, midnervo
infra ex 14-17 cm cum aculeis 4-6.5 mm longis
10-20 mm separatis crassiter subulatis arcuatis
reflexis, in sectione mediali marginibus cum
aculeis 2.5-3 mm longis 14-30 mm separatis
arcuato-subulatis valde adscendentibus basibus
incrassatis, in apice contractescentibus margini-
bus undulatis et cum subulato-serris 1.5-2 mm
longis 1.5-5 mm separatis, midnervo infra an-
gusto et cum serris simulantibus, in apice mar-
ginibus et midnervo infra cum serrulis 0. 5-0.8
mm longis 1.5-3 mm separatis, inflorescentia
foeminea terminali erecta cum syncarpio solita-
rio, pedunculo 20 cm longo 12 mm diametro
trigono folioso-bracteato, syncarpio ex bracteis
tandem siccis et papyraceis subclauso, syncarpio
11.5 cm longo 6.7 cm diametro ellipsoideo sub-
trigono viridi cum circa 672 drupis, eis 20-25
mm longis 5-9 mm latis 4-8 mm crassis anguste
cuneatis apice hemisphaerico vel pyramidali-
hemisphaerico 5-7-anguloso, corpore 19-22 mm
longo, basi pile! 5-6 mm alto viridi glauco,
stylo 3-4 mm longo brunneo lucido osseoso
proxime curvato lato compresso cornuformi bi-
fido lobis 0.5-2 mm longis deltoideis (raro inte-
gro), stigmatibus 1.5-2. 5 mm longis late ovato-
lanceolatis proximis paene ad apicem productis
papUlosis brunneis, endocarpio in parte lA in-
feta oblongo-ellipsoideo apice truncate intra
pallidi lucido lateribus lateralibus 0.2 mm cras-
sis osseosis brunneis exteriori sillonati, sernini-
bus 7-8 mm longis 3-3.5 mm diametro, meso-
carpio apicali albi medullosi in media sine fibris.

mesocarpio basali 2-3 mm longo in lateribus
fibroso in media carnoso.

DIAGNOSIS OF holotype: Stemless herb with
several shoots, forming a clump 80 cm in di-
ameter; leaves 2-3 m long, 4-5 cm wide, above
dark olive green and shining, below green to
yellowish green, above with V-shaped furrow
over the midrib and two lateral pleats, at mid-
section with 36 secondary parallel veins in each
half, tertiary cross veins numerous and visible,
but on drying conspicuous, forming squarish or
oblong meshes, blade ligulate, gradually con-
tracted in the last 40' cm to a 10 cm subulate,
trigonous apex, this about 10 cm down 2 mm
wide, the base amplexicaul, unarmed, shining,
dull copper-colored, beginning at 12-15 cm the
margins with spines 3-6 mm long, 10-20 mm
apart, thick subulate, slightly upcurved, stramin-
eous to blackish; the midrib below beginning
at 14-17 cm with spines 4-6.5 mm long, 10-20
mm apart, heavy subulate, arcuate, reflexed; at
midsection the margins with prickles 2.5-3 mm
long, 14-30 mm apart, arcuate subulate from a
thickened base, strongly ascending; the midrib
below unarmed; in the contracting part near the
tip the margins undulate and with subulate-
serrae 1.5-2 mm long, 1.5-5 mm apart; the nar-
row midrib below with similar serrae; on the
apex the margins and midrib below with ser-
rulations 0. 5-0.8 mm long, 1.5-3 mm apart;
pistillate inflorescence terminal, erect, bearing
one syncarp; peduncle 20 cm long, 12 mm in
diameter, 3 -sided, leafy bracted; syncarp sur-
rounded and nearly hidden by bracts that be-
come dry and papery; syncarp immature but ap-
parently about full sized, 11.5 cm long, 6.7 cm
in diameter, ellipsoid, slightly 3 -sided, green,
bearing about 672 drupes, these 20-25 mm long,
5-9 mm wide, 4-8 mm -thick, narrowly euneate,
the apex hemispheric or pyramidal-hemispheric,
5-7 -angled, the body 19-22 mm long; p ileus
with its base 5-6 mm high hemispheric or pyra-
midal-hemispheric, green, glaucous (immature);
style 3-4 mm long, brown, shining, bony, proxi-
mally curved, broad, flattened hornlike, equally
or unequally bifid, the lobes 0.5-2 mm long,
deltoid (rarely unbranched this mostly on ter-
minal drupes); stigmas 1.5-2. 5 mm long,
broadly ovate-lanceolate, proximal, following
each lobe almost to the tip, papillose, brown;
endocarp in lower oblong-ellipsoid, the apex

478

PACIFIC SCIENCE, Vol. XVII, October 1963

truncate, the inner surface pale, shining, the
walls 0.2 mm thick, bony, brown, and the outer
surface with sharp longitudinal ridges; seed 7-8
mm long, 3-3.5 mm in diameter; apical meso-
carp of white pith, without fibers except up the
sides; basal mesocarp 2-3 mm long, fibrous up
the sides, fleshy within.

HOLOTYPUS: Vietnam, Bach Ma, Province de
Thua-Thien, on forested stream bank, 1,400 m.
alt., Jan. 26, 1961, H. St. John 26,379 ( BISH ) .

DISCUSSION: P. bipollicaris is a member of
the section Rykia , as is its closest relative, P.
bicornis Ridl. of Malaya, a species which has the
peduncle 5-10 cm long; syncarp 7-8 cm long,
5.6 cm in diameter, bearing 100-120 drupes,
these 28-34 mm long, 8-10 mm wide; style 2-3
mm long; endocarp central, with a central apical
prolongation, the lateral walls 2 mm thick;
leaves 0.7-2 m long, at midsection with 24 sec-
ondary veins in each half, the blade quickly
tapering into a 5 cm subulate apex, and near the
base the margins with prickles 2.5-4 mm long
and 2-7 mm apart. P. bipollicaris has the pe-
duncle 20 cm long; syncarp 11.5 cm long, 6.7 or
more cm in diameter, bearing about 672 drupes,
these 20-25 mm long, 5-9 mm wide; style 3-4
mm long; endocarp in lower V3, the apex trun-
cate, the lateral walls 0.2 mm thick; leaves 2-3
m long, at midsection with 36 secondary veins
on each side, very gradually tapering into a 10
cm subulate apex, the margins near the base
with spines 3-6 mm long, 10-20 mm apart.

The epithet is the Latin adjective, bipollicaris,
length of two thumbs or two inches, and is
chosen with reference to the leaves which are
about two inches wide.

Pandanus magnifibrosus sp. nov. (sect. Rykia)

Fig. 198

DIAGNOSIS HOLOTYPI: Planta sine caule, foliis
"3.65 m. longis,” 4.8 cm latis coriaceis ligulatis
in sectione mediali cum 34 nervis secundariis in
quoque latere in apice subulato longe diminuen-
tibus et 10 cm ex apice solum 16 mm latis triner-
vatis et in sectione M-f ormatis, in regione
mediali marginibus cum aculeis 2-2.2 mm
longis 8-15 mm distantibus arcuato-subulatis
subluteis, midnervo infra cum aculeis 2-2.5 mm
longis 9-17 mm distantibus salientibus sed api-

cibus reflexis, proxima apicem marginibus et
midnervo infra cum serris 0. 5-0.8 mm longis
1-4 mm distantibus et nervulis secundariis supra
cum serris simulantibus 3-8 mm distantibus,
"pedunculo 0.38 m. longo, syncarpio 0.34 m. am-
bitu,” drupis numerosissimis 27-30 mm longis
3-4 mm latis et crassis anguste oblanceo-fusi-
formibus 5-6-angulatis, parte 3/7 supera libera,
pileo 12-13 mm longo arcuato lineari-lanceo-
loideo firmo papilloso rugosoque stramineo, stylo
5-6 mm longo subulato arcuato ad apicem vel
ad basem syncarpii cartilagineo laevi et lucido,
stigmate 6-7 mm longo sublineari brunneo pa-
pilloso proximo, endocarpio in parte 2A infera
et 12 mm longo pallido, seminibus non evidentis
forsan sterilibus, mesocarpio apicali et laterali
conjunctis valde fibrosis et medullosis, mesocar-
pio basali fibroso et carnoso.

DIAGNOSIS OF HOLOTYPE: Stemless; leaves
"3.65 m. long,” 4.8 cm wide, coriaceous, ligulate,
at midsection with 34 secondary veins in each
half, long tapering to a heavy subulate apex
which 10 cm down is 16 mm wide; the base not
seen; 3 -nerved and in section M-shaped; at the
middle the margins with prickles 2-2.2 mm
long, 8-15 mm apart, arcuate-subulate, yellow-
ish; the nearby midrib below with prickles 2-2.5
mm long, 9—17 mm apart, salient but the apex
reflexed; near the apex the margins and midrib
below with serrae 0.5-0. 8 mm long, 1-4 mm
apart, and the secondary nerves above with simi-
lar serrae 3-8 mm apart; "peduncle 0.38 m. long;
syncarp 0.34 m. in circumference”; drupes very
numerous, 27-30 mm long, 3-4 mm wide and
thick, narrowly oblance-fusiform, 5-6-angled,
the upper 3/7 free; pileus 12-13 mm long, ar-
cuate linear-lanceoloid, firm papillose and ru-
gose, stramineous; style 5-6 mm long, subulate,
curved and often recurved, cartilaginous, smooth
and shining; stigma 6-7 mm long, almost linear,
brown papillose, proximal; endocarp in lower
7 6 and 12 mm long, pale; seeds not seen, per-
haps sterile, though apparently mature; apical
and lateral mesocarp continuous, strongly fibrous
and pithy; basal mesocarp fibrous and fleshy.

HOLOTYPUS: Thailand (Siam), Kao Soi Dao,
Patalung, evergreen forest, 300 m. alt., April 29,
1930, A. F. G. Kerr 19,227 (bk).

DISCUSSION : Pandanus magnifibrosus St. John
is a member of the section Rykia. Its closest

FIG. 198. Pandanus magnifihrosus St. John, from holotype. a, b, Drupe, lateral view, X 1; c, drupe, longi-
tudinal median section, X I; d, drupe, lateral view, X 4; e, drupe, longitudinal median section, X 4; f, drupe,
apical view, X 4; g, drupe apex and stigma, proximal view, X 4; h, drupe, median transverse section, X 10;
i, leaf middle, lower side, X 1; j, leaf apex, lower side, X 1; 4 leaf apex, upper side, X 1-

5 cm..

Fig. 199. Pandanus obconicus St. John, from the holotype. a, Syncarp and bracts, X 1; b, drupe, distal
view, X 1; c, drupe, apical view, X 1; ^ drupe, longitudinal median section, X 1; lateral drupes, apical

view, X 1; o, apical drupe, lateral view, XI \ P, drupe, proximal view, X 4; q, drupe, longitudinal median
section, X 4; r, apex of apical drupe and stigma, distal view, X 4; s, stigma of apical drupe, proximal view,
X 4; t, apex of lateral drupe, distal view, X 4; u, apex of lateral drupe and stigma, proximal view, X 4; v,
leaf base, lower side, X 1; ^ leaf middle, lower side, X 1; x, leaf apex, lower side, X l;^ lower side of leaf
in outer third, with secondary and tertiary veins, X 4.

Page 319: Revision of Pandanus, 16. Thailand, Vietnam — St. John

481

relative is P. Sarasinorum Warb. of the Celebes
and Minhassa, a species having the syncarps 3.5
cm in diameter, racemose; drupes 12-14 mm
long, 2 mm in diameter; style 4 mm long; and
the leaf spines becoming black. In contrast, P.
magnifibrosus has the syncarps 11.3 cm in di-
ameter, solitary; drupes 27-30 mm long, 3-4
mm in diameter; style 5-6 mm long; and the
leaf spines yellowish.

The new epithet is from the Latin, magnus,
large, fibrosus, with fibers, in allusion to the very
large, longitudinal fibers in the drupes.

Pandanus obconicus sp. nov. (sect. Rykia )

Fig. 199

DIAGNOSIS HOLOTYPI: Planta acaulescens, fo-
liis 96 cm longis in media 3.6 cm latis proxima
basem 2.7 cm latis coriaceis late sulcatis 2-
plicatis in sectione mediali AA-formatis et cum
37-39 nervis secundariis parallelis in quoque
dimidio nervis tertiis ad basem et ad apicem
conspicuis et reticulos breviter oblongos for-
mantibus laminis gladiformatis et in apice 8-12
cm longo trigono subulato eo 10 cm ex apice 3
mm lato basi amplexicauli et inermi sed ex 5-13
cm marginibus cum aculeis 2.5-5 mm longis

7- 15 mm separatis crassiter arcuatis subulatis
stramineis adscendentibus, midnervo infra ex

8- 15 cm cum aculeis 5-6 mm longis 12-25 mm
separatis subulatis subcurvatis valde reflexis in
sectione mediali marginibus cum aculeis 2-3
mm longis 6-9 mm separatis crasse subulatis
adpresse adscendentibus, midnervo infra cum
aculeis 3-4 mm longis remotis subulatis arcuatis
basi incrassati reflexis vel adscendentibus, in
apice subulato marginibus cum serris 0.5-1 mm
longis 2-4 mm separatis, midnervo infra cum
aculeis simulantibus 4-6 mm separatis, infructe-
scentia terminali erecta cum syncarpio unico,
pedunculo 17 cm longo 1 cm diametro trigono
folioso-bracteato, bracteis superis 9-10 cm longis
4 cm latis minute serrulatis dentibus 0.2 mm
longis, syncarpio 7.5 cm longo 4.5 cm diametro
ellipsoideo cum circa 496 drupis eis 16-19 mm
longis 7-9 mm latis 4-7 mm crassis eis termi-
nalibus majoribus crassiter cuneiformatis com-
pressis 5-7-angulatis parte Ve supera libera
corpore 14-17 mm longo lateribus planis, pileo
3-4 mm alto basi depresse obtuso 5-7-anguloso
minute papilloso, stylo 2-3 mm longo osseoso

brunneo lucido compresso proxime curvato
(vel V5-V2 ) bifido lobis plerumque divergen-
tibus, stigmatibus 1.5-2 mm longis apicalibus
vel in parte proximalibus brunneis papillosis,
endocarpio infra mediali osseoso pallide brun-
neo lateribus 0.6-0.7 mm crassis intra obscure
brunneis lucidis apice cum acumine fragili, se-
mine 7 mm longo anguste obconico, mesocarpio
apicali grandi cavernoso cum membranis medul-
losis albis, mesocarpio basali fibroso et carnoso.

DIAGNOSIS OF HOLOTYPE: Plant acaulescent;
leaves 96 cm long, 3.6 cm wide at the middle,
2.7 cm wide near the base, coriaceous, broadly
furrowed above the midrib, 2 -pleated, in section
AA-shaped, at midsection with 37-39 parallel
secondary veins in each half, the tertiary cross
veins marked near the base and the tip, forming
short oblongs, and from these the outer third of
the blade has a reticulate appearance, the blade
sword-shaped, narrowed to an 8-12 cm trigonous
subulate apex which 10 cm down is 3 mm wide,
the base amplexicaul, unarmed, but beginning
at 5-13 cm the margins with prickles 2.5-5 mm
long, 7-15 mm apart, stout arcuate subulate, as-
cending, stramineous; the midrib below begin-
ning at 8-15 cm with prickles 5-6 mm long,
12-25 mm apart, subulate, slightly curved,
sharply reflexed; at the midsection the margins
with prickles 2-3 mm long, 6-9 mm apart, stout
subulate, appressed ascending; the midrib below
with remote prickles 3-4 mm long, subulate,
arcuate, with an incrassate base, reflexed or as-
cending as here the change-over occurs; on the
subulate apex the margins with serrations 0.5-1
mm long, 2-4 mm apart; those of the midrib
similar but 4-6 mm apart; infructescence termi-
nal, erect, bearing one syncarp; penduncle 17
cm long, 1 cm in diameter, trigonous, leafy
bracted, the upper bracts 9-10 cm long, 4 cm
wide, minutely serrulate, the teeth 0.2 mm long;
syncarp 7.5 cm long, 4.5 cm in diameter, ellip-
soid, bearing about 496 drupes, these 16-19
mm long, 7-9 mm wide, 4-7 mm thick, the
apical ones perceptibly the larger, thick cunei-
form, compressed, 5-7-angled, upper Ve free,
the body 14-17 mm long, the sides flat; pileus
3-4 mm high, the base low obtuse, 5-7 -angled,
minutely papillose; style 2-3 mm long, bony,
brown, shining, flattened, curved proximally,
bifid usually 2A way, rarely ]/$ or V2 way, the
lobes usually divergent; stigma 1.5-2 mm long,

482

PACIFIC SCIENCE, Vol. XVII, October 1963

on the apex or partly on the proximal side of
each lobe, brown, papillose; endocarp subme-
dian, bony, pale brown, the lateral walls 0.6-0. 7
mm thick, the inside surface dark brown, shiny,
crowned by a weak apical acumination; seed 7
mm long, narrowly obconic; apical mesocarp a
large cavern divided by white medullary mem-
branes; basal mesocarp fibrous and fleshy.

HOLOTYPUS: Thailand, s. e. part, Chanburi,
Makham, 100 m. alt., common along edge of
savannah in scrub forest, 13 Jan. 1958, Tem
Smitinand 4,054 (SING).

DISCUSSION: P. obconicus is a member of the
section Rykia, as is its closest relative, P. hori-
zontalis St. John, of Vietnam, a species with
peduncle 45 cm long; syncarp 10.5 cm long, 7-8
cm in diameter, with about 720 drupes, these
26-31 mm long, 9-11 mm wide; pileus base
high rounded conic; endocarp in lower P3; seed
12 mm long; stem 1.5 m tall; leaves 2.7 m long,
at midsection with 33 secondary veins in each
half, and near the base the midrib below with
thorns 3-4 mm long, 17-35 mm apart. P. ob-
conicus has the peduncle 17 cm long; syncarp
7.5 cm long, 4.5 cm in diameter, with about 496
drupes, these 16-19 mm long, 7-9 mm wide;
pileus base low obtuse; endocarp submedian;
seed 7 mm long; acaulescent; leaves 96 cm long,
at midsection with 37-39 secondary veins in
each half, and near the base the midrib below
with prickles 5-6 mm long, 12-25 mm apart.

The new epithet is Latin, ob, inversed, coni-
cus, conical, given in reference to the obconic
shape of the endocarp.

Pandanus obovatus sp. nov. (sect. Rykia )

Fig. 200

DIAGNOSIS HOLOTYPI: Arbor ”6 m. alta ra-
mosa, trunco cum spinis parvis,” foliis 1.1 m
plus longis 3.1 cm latis subcoriaceis in sectione
mediali cum 39 nervis secundariis in quoque di-
midio gradatim ad apicem longe diminuentibus
supra secundum midnervum canaliculatis mar-
ginibus reflexis, basi integra amplexicaulique
sed ex 5-7 cm marginibus cum aculeis pallidis
2-2.5 mm longis 5-9 mm distantibus subulatis
adscendentibus, midnervo infra per 15 cm
inermi deinde cum aculeis reflexis 0.5-1 mm
longis 15-18 mm distantibus subulatis, in sec-
tione mediali marginibus cum aculeis 0.8-0. 9

mm longis 7-10 mm distantibus subulatis ad-
scendentibus, midnervo infra cum aculeis 0.3-
0.5 mm longis 8-18 mm distantibus simulanti-
bus adscendentibus, ad apicem marginibus mid-
nervoque infra cum serrulis 0.2 mm longis 5-15
mm distantibus, syncarpio "cum 15-18 carpelis,”
drupis 7. 5-8. 3 cm longis 4.9 cm latis 3-7 cm
rrassis obovoideis deltoideis laevissimis, lateri-
bus superis laevibus brunneo-lucidis quasi cera-
tis et pallide multirimosis deltoideis, parte
supera libera, parte V2 infera pallide brunnea et
cum dorsis et valleculis angustis longitudinalibus
intercbseratis, apice rotundato sed cum dorsis
duobis ad stigmatem connectis, stylo cornicu-
lato compresso divergento, stigmate 8 mm
longo late obovato exsulcato subverticale brun-
neo, endocarpio submediali parte centrali 4.3
cm longa osseosa obscure brunnea ellipsoidea
muris lateralibus 2-3 mm crassis et cum humeris
fortibus pallidis lateralibus submediale affixis
eis oris 3. 5-4. 5 cm longis marginalibus crassis,
semine unico 25 mm longo 9 mm diametro
ellipsoideo mesocarpio laterali apicalique con-
tinuo sicco et medulloso sed cum fibris longitu-
dinalibus, mesocarpio basali dimidio supero
simulanti, dimidio infero fibroso et carnoso.

DIAGNOSIS OF HOLOTYPE: Tree ”6 m. tall,
branching; stem with small spines”; leaves more
than 1.1 m long, 3.1 cm wide, subcoriaceous,
at midsection with 39 secondary veins in each
half, gradually long tapering towards the apex,
(the tip missing in ours), channeled above the
midrib, the margins reflexing, the base am-
plexicaul and entire, but beginning 5-7 cm
up the margins with pale prickles 2-2.5 mm
long, 5-9 mm apart, subulate, ascending; the
midrib below unarmed for 15 cm, then with
reflexed prickles 0.5-1 mm long, 15-18 mm
apart, subulate; at midsection the margins with
prickles 0. 8-0.9 mm long, 7-10 mm apart, su-
bulate, ascending; the midrib below with similar
appressed ascending prickles 0.3-0. 5 mm long,
8-18 mm apart; towards the apex the margins
and midrib below with serrations 0.2 mm long,
5-15 mm apart; syncarp "with 15-18 carpels”;
drupes 7. 5-8. 3 cm long, 4.9 cm wide, 3.7 cm
thick, obovoid, 3 -sided, very light, the upper
sides smooth, brown and shiny as if varnished
and with many pale longitudinal cracks, 3-
angled, upper lA free, lower half pale brown
and with several sharp longitudinal ridges and

FIG. 200. Pandanus ohovatus St. John, from holotype. a, Drupe, lateral view, X 1; b, drupe, longitudinal
median section, XI \ c, drupe, apical view, X 1; 4 stigma, proximal view, X 1; e, apex of drupe, lateral
view, X 1; /> leaf base, lower side, X 1; g, leaf middle, lower side, X 1; h, leaf apex, lower side, X 1-

484

PACIFIC SCIENCE, Vol. XVII, October 1963

valleys like mortising angles, the apex rounded
but from each of two angles a ridge mounts to
support the stigma on a flattened, overhanging
process or style; stigma 8 mm long, broadly obo-
vate, without crease, nearly vertical, brown; en-
docarp submedian, the central part 4.3 cm long,
bony, dark brown, and ellipsoid, its lateral walls
2-3 mm thick, a little below the middle of this
central endocarp are heavy, pale, lateral shoul-
ders bearing heavy marginal flanges 3. 5-4. 5 cm
long; seed solitary, 25 mm long, 9 mm in di-
ameter, ellipsoid; lateral and apical mesocarp
continuous, dry and pithy but with longitudinal
fibers, and this pithy mesocarp also occupies the
upper half of the area below the endocarp, the
remaining basal area fibrous and fleshy.

HOLOTYPUS: Thailand (Siam): Menam Pasak,
Keng Koi, on rocky limestone hill, Dec. 9, 1923,
A. F. G. Kerr 7,669 (bk).

DISCUSSION: P. ohovatus is a member of the
section Rykia, though with its large, dry, pithy
drupes it is unique and without a close relative.
It can be compared to P. borneensis Warb.
which has the syncarps spicate, 6-8 cm long;
drupe apex 10-12 mm broad; and the style 2
mm long. In P. obovatus the syncarp is solitary,
about 23 cm long; drupe apex 4.9 cm broad;
and the style 1 1 mm long, horn-like, flattened.

The type specimen consists of a leaf minus
the tip, and three fruit halves, yet so unique is
the species that this much suffices. There are no
really close relatives known.

The new epithet is the Latin obovatus, refer-
ring to the obovate profile of the drupes.

Pandanus reticulosus sp. nov. (sect. Rykia )
Fig. 201

NOM. VERN.: "toei yai” (— Pandanus big).

diagnosis holotypi: Licet acaulescens, foliis
2 m longis 5 cm latis in medio 3.8 cm latis circa
basem coriaceis reticulatis in sectione mediale
cum 33 nervis secundariis parallelis fortibus in
quoque dimidio, nervis tertiis transversis fortibus
elevatis et reticulam quadratorum vel oblongo-
rum brevium formantibus, laminis ligulatis sed
subabrupte in apice circa 10 cm longo caudato
trigono subulato diminuentibus eo circa 10 cm
ex apice 5 mm lato, basi amplexicauli et inermi
pallida sed ex 7-15 cm marginibus cum aculeis

3-5 mm longis 12-22 mm separatis subulatis
curvatis fortibus pallidis adscendentibus, mid-
nervo infra ex 12.5 cm cum aculeis 3-6 mm
longis 32-65 mm separatis arcuato-subulatis
reflexis, in sectione mediali marginibus cum
aculeis 2.8-3 mm longis 14-24 mm separatis
arcuatis subulatis adscendentibus, midnervo infra
inermi, in apice subulato marginibus cum ser-
rulis 0.6-1 mm longis 1-3 mm separatis, mid-
nervo infra cum serrulis 0. 3-0.8 mm longis 3-4
mm separatis, pedunculo "brevi,” syncarpio soli-
tario 7 cm longo 5.5 cm diametro latiter ellip-
soideo purpurascenti cum 113 drupis, eis 22-26
mm longis 9-14 mm latis 9-11 mm crassis ob-
lanceoloideis basi truncata 6-angulosis parte Va
supera libera, corpore 18-22 mm longo later ibus
planis, pileo 6-8 mm alto basi 4-5 mm alto
semiorbiculari vel depresse semiorbiculari rugosa
exlaevi minime 6-angulosa, stylo 2.5-5 mm
longo osseoso brunneo lucido compresso proxime
curvato eis druparum lateralium plerumque
breviter bifidis sed aliquis basalibus subulatis,
stigmate 1-1.5 mm longo cordato vel late cor-
dato brunneo papilloso in apice proximali caver -
noso affix!, endocarpio mediali 11 mm longo
osseoso extus pallid! lateribus 0.8 mm crassis
intra obscure brunneis et lucidis, seminibus 10
mm longis 8 mm diametro obovoideis, meso-
carpio apicali aerenchymati cum membranis latis
transversis albis, mesocarpio basali fibroso et
carnoso.

DIAGNOSIS OF holotype: Apparently acau-
lescent, gregarious on ground among under-
shrubs; leaves 2 m long, 5 cm wide at the mid-
dle, 3.8 cm wide near the base, coriaceous, reti-
culately veined and conspicuously so in outer
half, at midsection with 33 parallel secondary
veins in each half, the tertiary cross veins heavy,
raised, forming a reticulum of squares or short
oblongs, blade ligulate, rather abruptly narrowed
to a caudate trigonous subulate apex about 10
cm long, this about 10 cm down 5 mm wide, the
base amplexicaul and unarmed, pale, but be-
ginning 7-15 cm up the margins with spines
3-5 mm long, 12-22 mm apart, stout curved
subulate, ascending, pale; the midrib below be-
ginning at 12.5 cm with spines 3-6 mm long,
32-65 mm apart, arcuate subulate, reflexed; at
midsection the margins with prickles 2.8-3 mm
long, 14-24 mm apart, arcuate subulate, ascend-

1 o cm/.

Fig. 201. Pandanus reticulosus St. John, from holotype. a, Drupe, lateral view, X 1; b, drupe, longitudinal
median section, X 1; c— h drupe, apical view, X 1; (d and / are the most common kinds; e is basal, rarely
lateral); m, apex of apical drupe and stigma, lateral view, X 4; n, apex of lateral drupe, oblique view, X 4;
o, apex of lateral drupe and stigma, proximal view, X 4; p, leaf base, lower side, X 1; leaf middle, lower
side, XI \ leaf apex, lower side, X 1; s, leaf middle, upper side, secondary and tertiary veins, X 4; t, leaf
middle, lower side, secondary and tertiary veins, X 4.

486

PACIFIC SCIENCE, Vol. XVII, October 1963

ing; the midrib below unarmed; on the subulate
apex the margins with serrations 0.6-1 mm
long, 1-3 mm apart; the midrib below with
serrations 0. 3-0.8 mm long, 3-4 mm apart; pe-
duncle "short”; syncarp single, 7 cm long, 5.5
cm in diameter, broadly ellipsoid, purplish, bear-
ing 113 drupes, these 22-26 mm long, 9-14 mm
wide, 9-1 1 mm thick, oblanceoloid from a trun-
cate base, 6-angled, upper Va free, the body
18-22 mm long, with flat sides; pileus 6-8 mm
high, its base 4-5 mm high, semiorbicular or
oblately so, the surface roughened, dull, weakly
6-angled; style 2.5-5 mm long, bony, brown,
shining, flattened, proximally curved, the lateral
ones mostly shortly bifid, but some of the basal
ones with a single, subulate tip; stigma 1-1.5
mm long, cordate or broadly so, brown, papil-
lose, in the hollow proximal tip; endocarp me-
dian, 1 1 mm long, bony, pale without, the walls
0.8 mm thick, within dark brown and shining;
seed 10 mm long, 8 mm in diameter, obovoid;
apical mesocarp an aerenchyma with broad
transverse, white membranes; basal mesocarp fi-
brous and fleshy.

HOLOTYPUS: Thailand, n. e. region, Loei, Phu
Krading, Tham Saw, common in evergreen for-
est, 1,300 m. alt., Tem Smitinand 406 (bkf no.
11,999).

DISCUSSION: P. reticulosus is a member of
the section Rykia, as is its closest relative, P.
bicornis Ridl. of Malaya, a species which has
the syncarp 15 cm long; drupes 34 mm long;
style forks spreading and with the tips recurved;
seed 6 mm long, cylindric; and the leaf margin
near the base with spines 2.5-4 mm long, and
2-7 mm apart. P. reticulosus has the syncarp 7
cm long; drupes 22-26 mm long; style forks as-
cending or diverging; seed 10 mm long, obo-
void; and the leaf margin near the base with
spines 3-5 mm long, and 12-22 mm apart.

The new epithet is the Latin adjective, reticu-
losus, with a network, and is given in reference
to the reticulate venation of the leaves.

Pandanus thailandicus sp. nov. (sect. Rykia )
Figs. 202, 203

nom. vern.: "toei nam.”

diagnosis holotypi: Frutex 1.5 m altus
ramosus, foliis 35-37 cm longis 6-8 mm latis
subcoriaceis ligulatis plana tis vel marginibus

paene involutis in sectione mediali cum 13
nervis secundariis parallelis in quoque dimidio
parte Vi supera sensim in apice subulato dimi-
nuenti 10 cm ex apice 4 mm lata in sicco palli-
dis et infra subglaucis in basi amplexicaulibus
et integris proxima basem marginibus cum acu-
leis 1.8-2. 2 mm longis 8-15 mm separatis ad-
scendentibus arcuato-subulatis stramineis, mid-
nervo infra cum aculeis 2.2-2. 5 mm longis 8-25
mm distantibus simulantibus adscendentibus, in
sectione mediali marginibus cum aculeis 1. 3-1.8
mm longis 4-7 mm separatis simulantibus sed
subadpressis, midnervo infra cum aculeis 1.7-2
mm longis 6-15 mm separatis simulantibus ad-
scendentibus, proxima apicem marginibus cum
serris 0. 1-0.3 mm longis 1-4 mm separatis ad-
pressis cum apicibus subulatis, midnervo infra
cum serris 0.2-0.6 mm longis 2.5-6 mm sepa-
ratis simulantibus, pedunculo terminali 7 cm
longo 3-4 mm diametro folioso-bracteato, syn-
carpio solitario pendenti 36 mm longo, 35 mm
diametro elliptico-subgloboso cum circa 230
drupis, eis 14-1 6 mm longis 4-4.2 mm latis 3
mm crassis anguste oblanceoloideis 5-7-angu-
losis lateribus subcurvatis parte Va supera libera,
pileo conico sed in sicco cum rugis longitudi-
nalibus, stylo 2.5-3 mm longo crasse subulato
cartilagineo pallido proximo-curvato, stigmate
2-2.5 mm longo lanceolato proximo brunneo
papillose, endocarpio centrali 6 mm longo an-
guste obovoideo cartilagineo obscure brunneo
intra lucido lateribus 0.1 mm crassis, mesocar-
pio apicali lateralique conjuncto sicco et medul-
loso, mesocarpio basali fibroso et carnoso.

DESCRIPTION OF ALL SPECIMENS EXAMINED:
Branching shrub 1.5 m tall; stem 8-10 mm in
diameter, yellowish, unarmed; prop roots not
mentioned; leaves 35-50 cm long, 6-10 mm
wide, subcoriaceous, ligulate, flat or the margins
slightly involute, at midsection with 13 parallel
secondary veins in each half, the upper Vi gradu-
ally tapering to the acute tip, this 10 cm from
the apex 4 mm wide, when dried pale and below
somewhat glaucous, the very base amplexicaul
and unarmed, near the base the margins with
prickles 1.8-2 .2 mm long, 8-15 mm apart,
ascending, arcuate subulate, stramineous; the
nearby midrib below with similar ascending
prickles 2. 2-2. 5 mm long, 8-25 mm apart; at
midsection the margins with prickles 1.3-1. 8
mm long, 4-7 mm apart, similar but subap-

Fig. 202. Pandanus thailandicus St. John, from holotype. a, Syncarp, XI \ b, drupe, lateral view, X 1;
c, drupe, longitudinal median section, XI \ d, drupe, lateral view, X 4; e, drupe, longitudinal median section,
X 4; f, drupe, apical view, X 4; g, b, apex of drupe and stigma, proximal view, X 10; /, leaf base, lower
side, >< 1; j, leaf middle, lower side, X 1; ^ leaf apex, lower side, X 1*

Fig. 203. Pandanus thailandicus St. John, from Dee Bun Phong 914. a, Staminate inflorescence, X 1;
b, column and anthers, X 10; c, leaf base, lower side, X 1; d, leaf middle, lower side, X 1; e, leaf apex,
lower side, X 1*

Page 327: Revision of Pandanus, 16. Thailand, Vietnam — St. John

489

pressed; the midrib below with similar prickles
1.7-2 mm long, 6-15 mm apart, ascending; near
the apex the margins with subulate-tipped ap-
pressed serrations 0. 1-0.3 mm long, 1-4 mm
apart; those of the midrib below similar but
0.2-0.6 mm long, 2.5-6 mm apart; peduncle
terminal, 7-12 cm long, 3-4 mm in diameter,
with many leafy bracts, the median ones 21 cm
long, 8 mm wide, the upper ones 7 cm long, 17
mm wide, foliaceous, white; syncarp single,
pendent, elliptic-subglobose, 36-55 mm long,
35-42 mm in diameter, bearing 230-592
drupes, these 14-19 mm long, 4-4.2 mm wide,
3 mm thick, narrowly oblanceoloid, 5-7 -angled,
the sides gently curving, upper lA free; pileus
conic, but when dry with longitudinal wrinkles,
the base connate with adjacent ones, leaving no
visible suture; style 2-3.5 mm long, stout su-
bulate, pale, cartilaginous, curved upward to-
wards apex of syncarp ( or the basal ones curved
distally); stigma 1.5-2. 5 mm long, lanceolate,
proximal, brown papillose; endocarp central, 6
mm long, narrowly obovoid, cartilaginous,
brown, shiny within, the walls 0.1 mm thick;
apical and lateral mesocarps continuous, dry
pithy; basal mesocarp fibrous and fleshy.

STAMINATE PLANT: Stem 8 cm or a little
more in height, 7-10 mm in diameter, dark
brown, striate, shining; leaves numerous in
terminal tuft, 46-48 cm long, just above the
base 12-13 mm wide, at the middle 7-8 mm
wide, green, secondary parallel veins prominent
throughout and at midsection 14 in each half,
the base amplexicaul, unarmed, softer, beginning
3.5-4 cm up the margins with prickles 2. 3-3. 5
mm long, 5-15 mm apart, arcuate, heavy subu-
late, ascending, yellow; the nearby midrib below
beginning at 5-5.5 cm with prickles 2-2.5 mm
long, 6-15 mm apart, similar but reflexed, at
the point of reversal of direction often paired;
at midsection the margin and midrib below with
similar prickles 1-2.5 mm long, 4-6 mm apart,
ascending; staminate inflorescence terminal; pe-
duncle 6 cm long, 1.5 mm in diameter, leafy
bracted; floral part 11 cm long; lowest floral
bract about 45 cm long, the base 15 mm wide,
soft, white, veiny, unarmed, narrowly lanceolate,
the upper part 6 mm wide, green, subcoriaceous,
at midsection the margins with prickles 0.5-1
mm long, 2-3 mm apart, arcuate subulate, as-
cending, yellow; the midrib below with similar

ones 0.3-0. 5 mm long, 1.5-3 mm apart; the 7
spikes fragrant, 2.3-3 cm long, 6-8 mm in di-
ameter, narrowly ellipsoid, densely flowered;
stamens numerous, in fascicles; staminal column
2-2.5 mm long, stout, but tapering upwards;
free filament tips 0.5 mm long; anthers 0.7-0.8
mm long, lanceolate, bearing on its apex a su-
bulate projection of the connective 0.2 mm long.

HOLOTYPUS: Thailand (Siam): Pii Wieng,
Kawn Ken, by dry stream bed, 300 m. alt., Feb.
7, 1931, A. F. G. Kerr 20,016 (bk).

specimens EXAMINED: Thailand (Siam),
Bung, Ubon, by stream in open forest, 100 m.
alt., Jan. 29, 1924, A. F. G. Kerr 8,365 (bk);
n. e. part, Loei, Phu Luang, by stream, 1,000 m.
alt., 24 Sept. 1957, Dee Bun Pheng 914, stami-
nate and pistillate (bkf no. 16,154).

DISCUSSION: P. thailandicus is a member of
the section Rykia, as is its closest relative, P.
immersus Ridl. of Malaya, a species with the
syncarp (immature) 9.5 cm long, 6 cm in di-
ameter, ellipsoid, with about 1,440 drupes, these
5-6 mm wide; pileus 7-8 mm long; style 4-6
mm long; plant aquatic; leaves "many feet long,"
10.2 cm wide, caudate subulate apex 3.0. , cm
long, at midsection with 70 secondary veins in
each half, and the margins with prickles 2.5-4
mm long, 9-18 mm apart. P. thailandicus has the
syncarp 3.6-5. 5 cm long, 3. 5-4.2 cm in diame-
ter, elliptic-subglobose, with 230-592 drupes,
these 4-4.2 mm wide; pileus 3.5-6 mm long;
style 2-3.5 mm long; plant terrestrial; leaves
35-50 cm long, 6- 10 mm wide, the subulate
apex 7 cm long, at midsection with 13 veins in
each half and the margins with prickles 1.3-1. 8
mm long and 4-7 mm apart.

The new epithet is geographic, referring to
the native country of the species.

Pandanus unicornutus sp. nov. (sect. Rykia)
Fig. 204

NOM. VERN.: "chawng 11”; "toi yai.”

DIAGNOSIS HOLOTYPI: Arbor ad 8 m alta
ramosa, foliis "ad 3.5 m. longis” 10-11 cm latis
proxima basem crassiter coriaceis in apice subu-
lato 8 cm longo abrupte contractis eo 10 cm ex
apice 7 mm lato in sectio late M-formatis in
puncto 30 cm ex basi cum 70 nervis secundariis
in quoque latere, basi integra et amplexicauli sed
ex 15 cm marginibus cum spinis 4.5-6 mm longis

Fig. 204. Pandanus unicornutus St. John, from holotype. a, Infructescence, X Vdl h, drupe, lateral view,
X 1; c, drupe, longitudinal median section, XI \ d, drupe, apical view, X 1; e> drupe apex and stigma,
proximal view, X 4; f, rare drape apex with bifid stigma, apical view, X 1; S> apex of rare drupe with bifid
stigma, proximal view, X 4; h, leaf base, lower side, X h leaf middle, lower side, X 1» is lea^ apex, lower

side, X 1-

Page 329: Revision of Pandanus, 16. Thailand, Vietnam — St. John

491

Fig. 205 a, Pandanus hipollicaris St. John,
from holotype. Branch, leaves, and syncarp.

7- 15 mm distantibus basi crassa apice subulato
subarcuatis divergentibus vel subadscendentibus
pallide brunneis, midnervo infra cum spinis 4-5
mm longis 10-41 mm distantibus crasse conico-
subulatis reflexis, in sectio mediali foliis 12.5-13
cm latis in paginis ambis rugoso-reticulatis
marginibus cum spinis 4-6 mm longis 21—38
mm distantibus adscendentibus basi crassa apice
crassiter subulato, midnervo inermi, proxima
apicem marginibus et midnervo infra cum serris
subulatis 0. 5-0.8 mm longis 2-4 mm distantibus
paginis ambis rugoso-reticulatis, laminis in sec-
tio M-formatis fere ad apicem, spicis cum 7
syncarpiis, "syncarpio maximo 15 cm. longo” 11
cm diametro, drupis multis 3. 5-3. 9 cm longis

8- 10 mm latis crassisque anguste ellipsoideis
lateribus laevibus planis vel subcurvatis 5-angu-
latis parte 1/7 supera libera, pileo oblato-semi-
orbiculari firmo quoque solum dehiscenti, stylo
6-8 mm longo osseoso corniformi integri ( rari-
ter bifido) proxime curvato pallide brunneo,
stigmate 3-4 mm longo lanceolate brunneo pa-
pillose, endocarpio submediali 25 mm longo la-
teribus 0.1 mm crassis cartilagineo stramineo
pagina interiori lucido apice lanceoloideo pro-
ducto, seminibus 15-16 mm longis 5-6 mm

FlG. 205 b. Pandanus hipollicaris St. John,
from holotype. Branch, leaves, and syncarp,
and longitudinal median section of syncarp-.

diametro ellipsoideo, mesocarpio apicali in mar-
ginibus fibroso intra cavernoso, mesocarpio
basali fibroso et carnoso.

DESCRIPTION OF ALL SPECIMENS EXAMINED:
Tree 7-15 m tall, 17-20 cm in diameter, branch-
ing; bark pale, with short spines; "prop roots
to 1 m. long, 5-8 cm. in diameter, armed with
short spines”; leaves '"3.2-3.85 m. long,” 8.4-11
cm wide near the base, thick coriaceous, abruptly
acuminate to a subulate apex 8 cm long, the tip
10 cm down from the apex 7 mm wide, very
wide M-shaped in section, at 30 cm from the
base with 70 secondary veins in each half, the
base entire and amplexicaul, but beginning 15
cm up the margins with thorns 4.5-7 mm long,
7-15 mm apart, heavy based, subulate-tipped,
somewhat arcuate, divergent or slightly ascend-
ing, pale brownish; the midrib below with
thorns 4-5 mm long, 10-14 mm apart, heavy
conic-subulate, reflexed; at midsection 12.5-13
cm wide, both surfaces rugose reticulate, the
margins with thorns 4-6 mm long, 21-38 mm
apart, ascending, the base heavy, the tip thick
subulate; the nearby midrib prominent but un-
armed; near the apex the margins and midrib
below subulate-serrate, the teeth 0. 5-0.8 mm

492

PACIFIC SCIENCE, Vol. XVII, October 1963

long, 2-4 mm apart, both the upper and lower
surfaces rugose-reticulate from the heavy cross
veinlets, the blade M-shaped in section and the
pleats running almost to the tip; peduncle 70
cm long; spike with 5-7 syncarps, the largest
14-18 cm long, 11-12 cm in diameter, ovoid
but distinctly 3 -sided; drupes numerous, these
3. 5-4.3 cm long, 8-10 mm wide and thick, nar-
rowly ellipsoid, the sides smooth, plane or gently
curved, 5 -angled, upper 1/7 free; pileus oblate-
semiorbicular, firm, shedding singly, ending in
a style 6-8 mm long, bony, horn-like, entire
( rarely bifid ) , strongly curved towards the apex
of the syncarp, pale brownish; stigma 3-4 mm
long, lanceolate, brown papillose; endocarp
slightly submedian, 25 mm long, the lateral
walls 0.1 mm thick, cartilaginous, stramineous,
the inner surface shining, the apex produced
upwards into a long lanceoloid tip; seeds 15-16
mm long, 5-6 mm in diameter, ellipsoid; apical
mesocarp fibrous around the margin, cavernous
within; basal mesocarp fibrous and fleshy.

holotypus: Thailand (Siam), Kaw Tao,
Surat, in high evergreen forest, under 5 m. alt.,
April 16, 1927, A. F . G. Kerr 12,772 (bk).

specimens EXAMINED: Thailand (Siam),
Kaw Tao, Surat, evergreen forest, 300 m. alt.,
Sept. 21, 1928, A. F. G. Kerr 16,052 (bk);
Wangka, Kanburi, by stream in evergreen forest,
700 m. alt., Feb. 3, 1926, A. F . G. Kerr 10,432
( BK ) ; Ta Ngaw, Chumpawn, common along
streams in savannah evergreen {forest}, Jan. 22,
1927, A. F. G. Kerr 11,601 (bk).

DISCUSSION : Pandanus unicornutus is a mem-
ber of the section Rykia, as is its closest relative,
P. penangensis Ridley, a Malayan species which
has the drupes 42-47 mm long, 6-1 mm thick;
endocarp 32-33 mm long, the lateral walls 1-1.5
mm thick; and the leaves 12 cm wide, near the
base the margins with prickles 1.3-1. 5 mm long,
4-7 mm apart. P. unicornutus has the drupes
35-43 mm long, 8-10 mm thick; endocarp 25
mm long, the lateral walls 0.1 mm thick; and
the leaves 10-11 cm wide, near the base the
margins with spines 4.5-6 mm long, 7-15 mm
apart.

The epithet is from the Latin unus, one, cor-
nutus, with a horn, in allusion to the usual single
hornlike style.

Index to Volume XVII

Author Index

Banner, A. H. :

On Malayan Shores, (review) , 374

Barlow, George W. :

Species Structure of the Gobiid Fish Gillichthys
mirabilis from Coastal Sloughs of the Eastern
Pacific, 47-72

Bonham, Kelshaw, and Edward E. Held :

Ecological Observations on the Sea Cucumbers

H olothuri&'atra and H. leucospilota at Rongelap
Atoll, Marshall Islands, 305-314

Bowers, Darl E. :

Field Identification of Five Species of Californian
Beach Hoppers (Crustacea: Amphipoda), 315-
320

Carlquist, Sherwin, and Martin L. Grant:

Studies in Fitchia (Compositae) : Novelties from
the Society Islands; Anatomical Studies, 282-298

Chang, Jen-hu:

Role of Climatology in the Hawaiian Sugar-Cane
Industry: An Example of Applied Agricultural
Climatology in the Tropics, 379-397

CHUANG, S. H. :

On Malayan Shores, (review of) , 374-375

Doty, Maxwell S.:

Gihsmithia haw alien sis, gen. n. et sp. n., 458-465

Emery, K. O. :

Aerial Study of Hawaiian Wave Patterns, 255—260

Gosline, William A.:

Notes on the Osteology and Systematic Position of
Hypoptychus dybowskii Steindachner and Other
Elongate Perciform Fishes, 90-101

Hida, Thomas S., and Robert A. Morris:

Preliminary Report on the Marquesan Sardine,
Harengula vittata, in Hawaii, 431-437

Hobson, Edmund S. :

Feeding Behavior in Three Species of Sharks, 171-
194

Hoyt, Charles P. :

Investigations of Rhinoceros Beetles in West Africa,
444-451

Hsiao, Sidney C, Walter K. Fujii, and Helen H.
Fine :

Simple Device for Making Successive Photomicro-
graphic Records of Large Groups of Developing
Organisms, 321-328

Inman, D. L., W. R. Gayman, and D. C. Cox:

Littoral Sedimentary Processes on Kauai, a Sub-
tropical High Island, 106-130

Lamoureux, Charles H. :

Additional Plants from the Midway Islands, 374

Lewis, Alan G. :

Life History of the Caligid Copepod Lepeophtheirus
dissimulatus Wilson, 1905 (Crustacea: Caligoida),
195-242

Martin, Edgar J.:

Toxicity of Dialyzed Extracts of Some California
Anemones ( Coelenterata ) , 302—304

Matthews, Donald C. :

Hawaiian Records of Folliculinids (Protozoa) from
Submerged Wood, 438— 443

Nizamuddin, Mohammed :

Studies on the Green Alga, Udotea indica A. & E. S.
Gepp, 1911, 243-245

Okutani, Takashi :

Preliminary Notes on Molluscan Assemblages of
the Submarine Banks Around the Izu Islands,
73-89

Paramonov, S. J. :

Lord Howe Island, A Riddle of the Pacific, Part III,
361-373

Pemberton, C E. :

Important Pacific Insect Pests of Sugar Cane, 251-
252

Pequegnat, Willis E. :

Population Dynamics in a Sublittoral Epifauna,
424-430

PLUCKNETT, D. L., J. C. Moomaw, and
C. H. Lamoureux:

Root Development in Aluminous Hawaiian Soils,
398-406

St. John, Harold :

Revision of the Genus Pandanus Stickman, Part 14.
New Species from Malaya and Singapore, 3-46

Part 15. Malayan Species Described by H. N. Rid-
ley, 329-360

Part 16. Species Discovered in Thailand and Viet-
nam, 466-492

Schafer, Rita D. :

Effects of Pollution on the Amino Acid Content of
Mytilus edulis, 246-250

SCHWABL, MATHILDE :

Solenogaster Mollusks from Southern California,
261-281

Sobolev, V. S.:

Characteristic Features of the Volcanism of the
Siberian Platform, 452-457

493

494

Stone, Benjamin C. :

Studies in the Hawaiian Rutaceae, IV. New and
Critical Species of Pelea A. Gray, 407-420

Tester, Albert:

The Role of Olfaction in Shark Predation, 145-170
Tomlinson, Jack T. :

Lithoglyptes hirsutus (Cirripedia: Acrothoracica) ,
A New Burrowing Barnacle from Hawaii, 299—
301

PACIFIC SCIENCE, Vol. XVII, October 1963

Wilbur, Robert L. :

A Prior Name for the Hawaiian Gouldia terminalis
(Rubiaceae) , 421-423

Winkler, Lindsay R., and E. Yale Dawson:

Observations and Experiments on the Food Habits
of California Sea Hares of the Genus Aplysia,
102-105

Yoshida, Tadao, Takeo Saw ada, and
Masahiro Higaki:

Sargassum Vegetation Growing in the Sea around
Tsuyasaki, North Kyushu, Japan, 135-144

Subject Index

adoption of the metric system and Celsius scale, 131

aerial study of wave patterns, 255-260

aluminous Hawaiian soils, root development in, 398—

406

amino acid content of Mytilus edulis, effects of pollu-
tion on, 246—250

amphipoda, Californian, identification of, 315-320
anemones, toxicity of, 302—304
Aplysia, food habits of, 102-105

applied agricultural climatology in the tropics, 379-
397

California anemones, toxicity of, 302-304
beach hoppers, identification of, 315-320
sea hares of the genus Aplysia, 102-105
solenogaster mollusks from, 261—281
Celsius scale, adoption of, 131

characteristic features of the volcanism of the Siberian
platform, 452-457

cirriped, new species of, from Hawaii, 299—301
climatology in Hawaiian sugar-cane industry, 379-397
copepod, caligid, life history of, 195-242

device for making successive photomicrographic rec-
ords of developing organisms, 321-328

eastern Pacific, species structure of Gillichthys mirabilis
from, 47-72

effects of pollution on the amino acid content of
Mytilus edulis , 246—250

feeding behavior in three species of sharks, 171-194
Fitchia, studies in, 282—298
folliculinids from submerged wood, 438-443
food habits of California sea hares of the genus
Aplysia, 102-105

Gibsmithia hawaiiensis, gen. n. et sp. n., 458-465
Gillichthys mirabilis, species structure of, 47-72
Gouldia terminalis, a prior name for, 421-423

Harengula vittata in Hawaii, 431—437

Hawaii, new species of burrowing barnacle from, 299-
301

records of folliculinids from submerged wood, 438-
443

Rutaceae, studies in, 407-420
wave patterns, aerial study of, 255-260
Holothuria atra, 305-314
Holothuria leucospilota, 305-314
Hypoptychus dybowskii, osteology and systematic posi-
tion of, 90—101

identification of Californian beach hoppers, 315—320
important Pacific insect pests of sugar cane, 251-252
insect pests of sugar cane, 251-252
investigations of rhinoceros beetles in West Africa,

444-451

Izu Islands, molluscan assemblages of the submarine
banks around, 72-89

Japan, Sargassum vegetation in the sea around, 135—
144

Kauai, littoral sedimentary processes on, 106—130

Lepeophtheirus dissimulatus, life history of, 195-242
life history of Lepeophtheirus dissimulatus, 195-242
Lithoglyptes hirsutus n. sp., 299—301
littoral sedimentary processes on Kauai, 106-130
Lord Howe Island, Part III, 361-373

Malaya, new species of Pandanus from, 3-46
Malayan species of Pandanus described by H. N. Rid-
ley, 329-360

Marquesan sardine in Hawaii, 431—437
Marshall Islands, sea cucumbers at Rongelap Atoll,
305-314

metric system, adoption of, 131
Midway Islands, additional plants from, 374
molluscan assemblages of submarine banks around the
Izu Islands, 73-89

mollusks, solenogaster, from southern California, 261-
281

Index to Volume XVII

495

Mytilus edulis, effects of pollution on the amino acid
content of, 246—250

Notes :

Additional Plants from the Midway Islands, 374
Adoption of the Metric System and Celsius Scale,
131

On Malayan Shores: A Review, 374

olfaction, role of, in shark predation, 145-170
osteology and systematic position of elongate perci-
form fishes, 90—101

Pacific insect pests of sugar cane, 251-252
Pandanus, Malayan species described by H. N. Ridley,
329-360

new species from Malaya and Singapore, 3-46
revision of genus, 3-46; 329—360; 466— 492
species discovered in Thailand and Vietnam, 466-
492

Pelea, new and critical species of, 407-420
perciform fishes, osteology and systematic position of,
90-101

photomicrographic records, device for, 321-328
pollution, effects of, upon amino acid content of
Mytilus edulis, 246-250

population dynamics in a sublittoral epifauna, 424-

430

predation by sharks, role of olfaction in, 145—170
prior name for the Hawaiian Gouldia terminals, 421-
423

rhinoceros beetles in West Africa, 444-451
Rongelap atoll, sea cucumbers at, 305-314

root development in aluminous Hawaiian soils, 398—

406

Sargassum vegetation around Tsuyazaki, Japan, 135-
144

sea cucumbers at Rongelap Atoll, 305-314
sedimentary processes on Kauai, 106-131
sharks, feeding behavior in three species of, 171-194
role of olfaction in predation, 145—170
Siberian platform, volcanism of, 452-457
Singapore, new species of Pandanus from, 3-46
Society Islands, Pitchia of, 282—298
soils, aluminous, and root development, 398—406
solenogaster mollusks from southern California, 261-
281

studies on the green alga, Udotea indica, 243—245
sublittoral epifauna, population dynamics in, 424-430
submarine banks around Izu Islands, molluscan as-
semblages of, 73-89

sugar-cane industry, role of climatology in, 379-397
sugar cane, Pacific insect pests of, 251-252

Thailand, Pandanus species from, 466-492
toxicity of California anemones, 302-304

Udotea indica, 243-245

Vietnam, Pandanus species from, 466-492
volcanism of the Siberian platform, features of, 452-
457

wave patterns, aerial study of, 255-260

West Africa, investigations of rhinoceros beetles in,

444-451

introduction and Summary. It is desirable to state the
purpose and scope of the paper in an introductory
paragraph and to give a summary of results at the end
of the paper.

Batzo, Roderick L., and J. K. Ripkin. 1849. A
Treatise on Pacific Gastropods. Rice and Shipley,
Boston, vii + 326 pp., 8 figs., 1 map.

Crawford, David L. 1920 a. New or interesting
Psyllidae of the Pacific Coast (Homop.). Proc.
Hawaii. Ent. Soc. 4(1): 12-14.

1920&. The sandalwoods of Hawaii. Proc.

Hawaii. Ent. Soc. 4(2): 374-375, 13 pis.

In the text, sources should be referred to by author,
date, and page, as follows: "It was noted (Rock,
1916: 18) that . . or "Rock (1916: 21-24)
says . . .”

ILLUSTRATIVE MATTER

Artwork for illustrations should be 8 V2 x 11 inches
or smaller, and it should accompany manuscript, on
separate sheets. Often more valuable than a photo-
graph is a good line drawing.

It is strongly urged that an indication of scale be
incorporated as a part of all drawings in which mag-
nification and size are critical considerations.

Illustrations will be returned to the author.

Tables. Tabular matter should be kept to a minimum.
Each table, prepared to conform with Pacific Science
style, should be typed on a separate page, and its posi-
tion indicated on the manuscript.

Mathematical Formulas. Complicated formulas cannot
be set by the printers. Authors should submit them
as illustrations.

Captions. Readily identifiable captions for figures,
graphs, photographs, and other illustrative matter
should be supplied on a separate page.

PROOF

All queries on proof should be answered. If cor-
rected proof is not received within four weeks after
being sent to the author, author’s changes cannot be
accepted.

REPRINTS

Internet Archive Audio

Featured

Top

Images

Featured

Top

Software

Featured

Top

Books

Featured

Top

Video

Featured

Top

Mobile Apps

Browser Extensions

Archive-It Subscription

Save Page Now

Full text of "Pacific science"

See other formats