ATOLL HESEAnCH B U D T I N
NO. 25
Bryophytes from Arno A t o l l , Marshall Islands bs'
Harvey Alfred Miller and Maxwell S . Doty
Issued by
THE PACIFIC SCIENCE BOARD
National Aeadeqy of Sciences-National Research Council Washington,
D.C.
November 15,
1953.
by Harvey Alfred :lille$ and !.laxcell S. ~ o t ~ 3
There nppears to have been but a s i n g l e provious record of o bryophyte from t h e U a r s h d l s . This i s Calmmeres tene m a wldely d i s t r i b u t e d spe- c i e s in Oceania, c o l i e c t e d by
W. R.
Tayl-or-4
(12 i n t h e northern Marshall Is- lands i n 1946. Glassman (7), and o t h e r s (2,3,8,9,lO,ll), have reported on t h e bryophytes from t h e nearby Caroline Islands. Dixon (4) published a l i s t of mosses from the G i l b e r t Islands, south of the Marshalls. There i s l i t t l e o r no reference t o the bryophyta i n C a t a l a ' s "Report on the G i l b e r t Islandst1 prepared f o r t h e South P a c i f i c Commission, o r i n the vegetational and o t h e r r e p o r t s on t h e P a c i f i c Science Board's a t o l l studies.During t h e sumner of 1951, Leonard Horwitz c o l l e c t e d several species of mosses and l i v e r w o r t s on Arno Atoll i n the southern Marshall I s l a n d s in con- nection n i t h t h e Coral A t o l l P r o j e c t of t h e T a c i f i c Science Board. Horwits's
c o l l e c t i o n $ were turned over t o t h e authors a t the University of Hawaii f o r determination. I t i s these c o l l e c t i o n s which form t h e b a s i s f o r t h i s report.
The senior author made t h e determinations, produced the f i r s t d r a f t of t h e r e p o r t manuscript and drew up t h e s p e c i f i c conclusions i n reference t o t h e bryophyta. The junior author contributed the o r i g i n a l i l l u s t r a t i o n s , t h e key and t h e l a b o r h v o l v e d i n f i n i s h i n g t h e work in t h i s form.
The authors a r e g r e a t l y indebted to E. B. Bartram who made many of the o r i g i n a l determinations of t h e mosses. The specimens c i t e d herein a r e de- p o s i t e d i n t h e U. S. National Museum, Washington, D. C., and i n t h e herbarium of t h e B. P. Bishop Museum, Honolulu, n i t h duplicates, wherever possible, in t h e Chicago Museum of Natural History and t h e h e r b a r i a of E. B. B a r t r a m and t h e authors. The s e n i o r author wishes t o thank Prof. Harold St. John, of the University of Hawaii, f o r provision of the a s s i s t a n t s h i p and space i n t h e De- partment of Botany which have enabled him t o continue h i s s t u d i e s of Central P a c i f i c bryophytes. The authors a r e g r a t e f u l f o r the m a t e r i a l a s s i s t a n c e given t h i s p r o j e c t by M r . Harold J. Coolidge, Mrs. Lenore Smith, and Miss Ernestiae Akers, of t h e P a c i f i c Science Board.
For t h e convenience of f u t u r e a t o l l workers a t Arno who a r e n o t bryolo- g i s t s , t h e following key i s included. A s e t of f i g u r e s made from camera l u c i d a sketches of leaves has been included. !This has been done because t h e
'This work was done with the a i d of f i n a n c i a l a s s i s t a n c e granted by t h e P a c i f i c Science Board of t h e National Research Council under c o n t r a c t N7onr-291, Task Order I V , from t h e Office of Naval Research and with s i m i l a r a s s i s t a n c e from t h e P a c i f i c I s l a n d s Research Fund of t h e University of Hawaii.
2 ~ c h o o l of Biology, Stanford University, and Department of Botany, University of Hawaii.
3 ~ e p a r t m e n t of Botany, University of Hawaii.
leaves a r e so d i s t i n c t i v e t h a t they may be used to d i s t i n g u i s h r a t h e r posi- t i v e l y between t h e few know Amo mosses. mith the description of each f i g u r e t h e r e appears t h e nunber of t h e c o l l e c t i o n from which t h e drawing nas made and c e r t a i n remarks of a i d i n distinguishing t h e species. I l l u s t r a t i o n s have n o t been included f o r t h e l i v e r w o r t s as they a r e much more r a r e and t h e i r taxonomy i s a s y e t most incomplete in many respects.
The c h a r a c t e r i s t i c s t o which tiiis k& Tefers a r e those v i s i b l e in fresh.,"' water-soaked o r we*, r n a t u ~ e specimens. The 1 , e a d 04 a l l mosses should be . compared with the f i g u r e s a s to s i z e and other c h a r a c t e r i s t i c s , f o r juvenile
. . forms, in p a r t i c u l a r , , may n o t "key out" i n t h i s key. It i s our hope t h a t
these devices ~ ~ 1 1 ' f a c i l i t a t e g r e a t e r f a m i l i a r i t $ d t h t h e mosses i n t h e f i e l d and ~ r o m o t e a study o f t h e i r r o l e in the atolls. ..
KEX TO THE ERYOPHYTA K N O W FROM ARNO ATOLL
A. Axis with leaves borne in 3 planes and i n such a manner t h a t the moist f r o n s s a r e : n o t b i l a t e r a l ; leaves e l l i p t i c a l to l i n e a r lanceolate; branches u s u a l l y a l l e r e c t from t h e substratum. (mosses): . .
, . .
. . . , . . . . .
B.
Leaf bases, stems and oacasiondLysome leaves red chen. stems n o t densely clothed with leaves; growing, on the ground: Bryum nitens... .
. . , 5 . . . . . .
B. Leaf bases, stems and leaves never red (though often brom); grow- ing on -wood o r rocks o r occasionally on:the. ground:
. .
C. Growing on t r e e trunks o r oegetable,debris;
tufts
often over 1 cm. thick, dark brown i n s i d e s o r p a l e whitish-greenthroughout: , . . ,
. .
D.
Tufts pale whitish green throughout:. . Leucophanes fzlauculum..
D. Tufts .yellow a t surface, dark , b r o ~ i n t e r n a l l y : Calmperes mollucense.
C. Growing on various o t h e r s u b s t r a t a o r yellow green t o green throughout, o r i f i n dense t u f t s then yellow-green t o green:
E. Leaves with an area of l a r g e c e l l s conspicuous near l e a f base (Figs. 3, 4 ,
5 ) ;
midribs extending to apex in leaves, percurrent, o f t e n extending into macroscopically v i s i b l e propagulae (terminal lumps rvhich a r e asexual r e p k d u c t o r y bodies) ; no capsules known f o r t h e Arno material:. ., . . . . . . . ,
F.
P r o s t r a t e moss with green stems and often , . widely spaced green leaves; l e a f t i p s den-t i c u l a t e : Calwperes hyophs4laceum.
i .
F. Erect moss w i t h c l o s e l y placed brownish green leaves; l e a f t i p s n o t denticulate:
Calymperes tenerum.
E.
Leaves without an area of l a r g e c e l l s near t h e base (Figs. 2, 6 ) ; midribs lacking o r n o t extend- i n g completely t o t h e apex; l e a f t i p s n o t bearing propagulae; capsules often present in abundance:G. Leaves tapering quickly to a long point, o f t e n s i c k l e shaped; midrib absent o r ex- tremely short; capsules often abundant:
G. Leaves s t r a p s h a p e d , broadly rounded a t apex, almost symmetrical; midrib extending n e a r l y t.o l e a f apex; capsules n o t known in Arno ma- t e r i a l : Splachobrwm indicum.
A. Axis with leaves b o n e i n trio planes and in Such s manner t h a t f l a t o r hemispherical branches r e s u l t ; leaves e i t h e r overlapping and ovoid o r l i k e t h e f l a t t e n e d a x i s in s t r u c t u r e ; usually all branches (except in P t y c h o c o l ~ ) p r o s t r a t e on t h e substratum (liverworts).
B. Leaves and axis a l i k e in appearance, f l a t ; frond p r o s t r a t e : Riccardia multifida.
H. Leaves bladelike and a x i s t e r e t e ; frond f r e e o r p r o s t r a t e :
I
Leaves strongly overlapping so a s to form a frond which i s hemispherical in cross section when damp and more o r l e s s c y l i n d r i c a l when dry; fronds becoming f r e e from t h e sub- stratum, forming a moss-like coating on t r e e bases up t o one centimeter thick:Ptvchocoleus sp.
I. Leaves n o t strongly overlapping, i f a t all; frond usually f l a t t e n e d a g a i n s t t h e substratum:
J. Fronds green, about o r over 1.5 mm. broad:
Le.i eunea sp.
J. Fronds pale if green a t all, about o r l e s s than 0.5 mu. broad:
Microleleunea sp.
Leucophanes glauculum C. d11.
-
in Micholitz,I.
Philippines 163.(SW. Figure 1.)
Eastern end df I n e I s l a n d , H o m i t z 9669, %70B, August 19, 1951.
Distribution: Malaysia, Caroline Islands.
This p l a n t was c o l l e c t e d on decaying wood and coconu+,, Cocos nucifera, husks. It f o m e d dense t u f t s where found. E. B. Bertram (correspondaxe) re- p o r t s t h a t F. R. Fosberg h a s c o l l e c t e d t h i s - s p e c i e s i n abundance on s t r a n d coconuts in t h e Camlines.
C a l p p e r e s hyophyllaceurn C. M%i.
&
Besch., Essai.Calymp. Ann. Sci. Nat.287. 1895.
(SXE: Figure 3 . ) Malel I s l a n d , ' Horvsitz 91258.
D i s t r i t u t i o n : Sumatra, Java, P h i l i p p i n e s (1).
T6ixefi with RXcx;r2> @ i i f i d . a growing in t h e shade m d on t h e under s i d e of damp outer-:and h i r e r co.:onut nusks. This record i s based on a few s c a t t e r e d p l a n t s , b u t i t h e , species i s so d i s t i n c t t h a t t h e r e i s l i t t l e question of i t s
. ..
i d e n t i t y .
A t t h e apex t h e l e a v e s of t h i s genus may bear propagulae in varying s t a g e s o f development. To t h e naked eye t h e asexual reproductory lumps of c e l l s ap- pear a s l i t t l e b a l l s on the l e a f t i p s . In f i g u r e s 3,
4,
and 5 t h e v a r i a t i o n inr erence shape of t h e apex may be due to the degree of maturity, l a r g e l y i n re"
to t h e s%age of propngulum development. This genus of mosses i s the only one from t h e l a r s h a l l s t h a t produces such s t r u c h r e s . The presence of these
propagu1c.e i s a good f i e l d character, t h e presence of which on a moss from Arno can be taken a s i n d i c a t i n g t h i s genus, Calymperes.
Calym~eres mollucense Schwaegr., Suppl. IS, 11, p. 99, t. 127. 1824.
(SEE: Figure 5.)
Malel Island, Hormltz 9422, 9U6A, 9429B,
- -
A u g l s t 4 , 1951.Distribution: Moluccas, F i j i , Philippines (1).
Horwite c o l l e c t e d a l a r g e t u f t (#9Q2) of t h i s species from the base of one coconut t r e e b u t gave no i n d i c a t i o n of i t s abundance o r whether it occurs on more than one i s l a n d in t h e a t o l l . A few i s o l a t e d p l a n t s
(#9mB)
weres c a t t e r e d vdth t h e Ptychocoleus (#99~29~) reported here, a l s o from a coconut trunk.
Cdymperes senerum C.
~ h . ,
L b n a e a 37: 174. 1871-73.(SEE: Figure L.)
Ine Village, Ine Island, Hor~ritz 931OA, Augdst 3, 1951.
Distribution: India, Malaysia, Philippines, and P a c i f i c I s l a n d s t o Hawaii.
This i s a widely d i s t r i b u t e d species and t h e only one reported by Taylor (12) from t h e northern Marshalls. I t i s i n t e r e s t i n g t o note t h a t Taylor found it abundant on Rongerik, Rongelap, Bikini and Eniwetok. The c o l l e c t i o n from Arno consisted of s c a t t e r e d p l a n t s among l i c h e n s scraped from t h e trunk of a b r e a d f r u i t Artocarpus sp. t r e e .
Tie c o l l e c t o r noted t h e propagulae t o be p a r t i c u l a r l y conspicuous on t h i s moss when collected.
Splachnobmm indicum C.
~ 2 1 1 . ,
Linnaea 37: 174. 1871-73.(SEE: Figure 2.) Ine Village, Ine Island, Horwitz 9437B.
Distribution: India, Java (1).
S c a t t e r e d p l a n t s among Ectropothecium monumentorurn growing "on top, s i d e s , and even some on bottom of shady moist stones near 6 c i s t e r n . "
The P h i l i p p i n e s have r o t been included i n t h e d i s t r i b u t i o n of t h i s species because t n e r e i s considerable doubt t h a t t h e p l a n t which Bartram c i t e s (1: 126) i s
5.
indicum. If the Philippine p l a n t is s d i s t i n c t species, then t h i s record l e a v e s a l a r g e r gap i n d i s t r i b u t i o n than any of t h e o t h e r species c i t e d k,erein.Bryum
n i t e n s Hook,&
Wall., Cat. n. 7592 e t Icon. p l . r a r . t. 20, f i g . 6 . 1837.(SEE: Figure 7.)
On v e r t i c a l l a y e r of s o i l l i n i n g main path across J i l a n g I s l a n d i n open coco- n u t grove. Horwitz 9105, J u l y 17, 1951.
Main roadway of Ine Village, Ine Island. Horsitz 9312, August 3, 1951.
Distribution: Nepal, Ceylon, Java, B a l i ( 8 ) , F i j i .
The extension of range indicated by t h i s c o l l e c t i o n may n o t be a s g r e a t a s shorn here s i n c e t h e r e i s considerable question whether t h i s species and
ambi~uum Duby (known from the Philippines) a r e d i s t i n c t from one another. E. B.
Bartram (correspondence) suspects t h a t F l e i s c h e r ' s ( 5 ) remark about
g.
n i t e n s ,"Diese A r t z e i g t so geringe Unterscheide von voriger, d t s s i c h s i e nur als Abart auffassen kann. S i e i s s i c h e r nur e i n e k l e i n e r e V a r i e t a t von
&.
ambimum, von dem s i e s i c h mehr durch e i n e Summe k l e i n e r Merkmale, a l s wie s p e c i f i s c h unter- scheidet" was well j u s t i f i e d .Ectopothecium monumentorum ( D u t y ) Jaeg., Adumbr. 2 (1877-1878) 523.
(SEE: Figure 6.)
On ground i n a coconut grove, J i l a n g Island, Homitz 9103, . J u l y 17,' 1951. On damp stump and on o u t e r husk of coconut, Malel Island, Horsitz 9426B,
m,
m,
August 4, 1951. . , .. ., . .
On shady moist;. s t o n e s near a c i s t e r n , Ine Vill.age, 1 n e T k ) , ~ d , . . Hornitz 9L37A, August 9, 1951.
On decaying word a t eastern end of Ine Island, H o r ~ i t z 967OA, August 19, 1951.
Distribution: Sumatra, Java, Timor, Philippines, Carolines
(I),
G i l b e r t s ( 4 ) . This rnoss appears t o be common i n t h e southern Marshalls from :th& 'h&ikr..
of c o l l e c t i o n s Horwitz made. It was f r u i t i n g abundantly and was found i n a v a r i e t y of h a b i t a t s . This was t h e onkr species of noss i n t h e whole c o l l e c t i o n with sporophytes (capsules).
LIVERKORTS (HEPATICAE)
Riccardin m u l t i f i d a (L.) S. F. Gray, Nat. Arr. E r i t . P1. 1: 683, 1821.
Growing over damp coconut husks and occasionally a l s o on inner husks. These were growing i n t h e shade and on under surfaces. Malel Island, Horvtitz 9L25A, August 4, 1951.
Distribution: Vtorldwide (6).
Although cosmopolitan, t h i s i s t h e f i r s t record of t h e species from eastern Micronesia. See a l s o ' n e x t species.
Riccardia sp.
Found growing on a piece of o u t e r coconut husk by Horwitz. 9L26C;at Malel Island, August 4, 1951.
M i l e t h e l i t e r a t u r e i s n o t p r e s e n t l y a v a i l a b l e f o r a d e f i n i t e determina- t i o n , t h e material seems t o be placeable close to t h e widely d i s t r i b u t e d
&-
cardia s i n u a t a (Dicks.) Trev. Tne t h a l l i of t h i s material t h i s s?ecies ( 6 ) have no u n i c e l l u l a r margin more than one c e l l broad. R. m u l t i f i d a has a 2- to 3-celled u n i s t r a t o s e margin along with a d d i t i o n a l diffzrences i n the sporophyte.
The more dependable sporophyte characters were n o t a v a i l a b l e i n our s t e r i l e ma- t e r i a l .
Microleleunea sp.
On an o l d inner coconut husk found lying in a damp s i t u a t i o n , Malel Island, Horwitz 9L260,-, August 4, 1951.
This i s a l a r g e and widespread t r o p i c a l genus, t h e previous n e a r e s t r e p o r t being from Ponape by Glassman ( 7 ) .
Lej eunea sp.
Growing on t h e trunk of
Cocas
nucifera, J i l a n g Island, with various o t h e r bryophytes and l i c h e n s , & ~ r w i t z 9102B, J u l y 17, 1951.I n abundance but only on dampest p a r t s of the same b r e a d f r u i t t r e e trunk which bore t h e Calym~eres tenerum collection, 9310, Ine Village, HorvcLtz 9316, August 3, 1951.
The c l o s e s t l o c a l i t y knovm f o r t h i s genus i s Yap, from which Stephani (10) reported tno species.
Ptvchocoleus sp.
Crowing on upfacing s i d e of a s l a n t i n g coconut t r e e from the ground l e v e l up t o about t h e L o r 6 ft. l e v e l where it thinned out, Male1 Island, Hornitz
m,
August 4, 1951. On t h e same i s l a n d and date on a coconut husk, Horwitzm.
This may prove t o be a species d i s t i n c t from a l l s o f a r described; a s a search of t h e s v s i l a b l e l i t e r a t u r e has revealed nothing c l o s e t o it. However, here a s u&+h the pr+viously c i t e d lejeuneoid species, t h e l i m i t e d l i b r a r y sources immediately a v a i l a b l e have n e c e s s i t a t e d a temporary reservation of judgment con- cerning t h e s p e c i f i c i d e n t i t y .
DISCUSSION
The only bryophytes knom t o be common t o both t h e G i l b e r t and t h e Marshall I s l a n d s a r e Calmperes tenerum and E c t r o ~ t h e c i u m monumentorum. These two a r e species of v:ide Indomalaysian d i s t r i b u t i o n .
It i s p a r t i c u l a r l y i n t e r e s t i n g t o note t h a t although a goodly number of in- v e s t i g a t o r s have reported on moss c o l l e c t i o n s from the Carolines, most of t h e
s p e c i e s found on Arno Atoll have n o t been reported from there. Perhaps t h e rea- son t h a t these species have n o t been reported from t h e Carolines l i e s i n t h e f a c t t h a t t h e g r e a t e s t i n t e r e s t i n these i s l a n d s has been centered on the r a i n f o r e s t s of t h e higher i s l a n d s and t h e moss s t r a n d vegetation, which i s similar t o t h a t of an a t o l l , has been grossly neglected.
There a r e several good reasons f o r believing t h a t t h e Carolines a r e t h e r o u t e of migration f o r t h e Marshallese species. Nearly all of t h e mosses c i t e d here a r e found i n t h e Philippines. The strong storm winds of the f a l l and w i n - t e r months a r e widely experienced coming from t h e west and southwest and it seems reasonable
to
assume t h a t these winds might a t some time o r o t h e r carry spores a s f a r a s t h e Marshalls. There i s a p o s s i b i l i t y , too, t h a t t h e e a r l y in- h a b i t a n t s of these i s l a n d s unwittingly c a r r i e d spores o r propagulae them during t h e i r migrations.,6
There i s a p o s s i b i l i t y of comparatively recent introduction by t r a d e r s end copra b06ts, but i f t h i s i s so then one i s l e d t o :;onder why these mosses a r e n o t found i n t h e Gilberts which a r e i s o l a t e d by the winds and c u r r e n t s from t h e Marshalls. I t seems l o g i c a l t h a t some boats would v i s i t both groups.
U n t i l more i s known of the d i s t r i b u t i o n of bryophytes in the P a c i f i c one can propose only such broad hypothesee,to be l a t e r subqtantiated, modified o r r e j e c t e d on the basis of more complete information.
.
A search was made f o r l i t e r a t u r e report'& e t h n i c uses of bryophitesin
~ i c r b n e s i a . None was found. It i s hoped t h a t f u t u r e workers tit Arno will in- q u i r e i n t o t h e ethnot.otany of the d i f f e r e n t bryophytes and t h a t such work w i l l be f a c i l i t a t e d by t h e treqtment which has been presented here.
BIBLIOGRAPHY
1. Bartram, E. B. 1939. Mosses of t h e Philippines. Philipp. Journ. Sci.
68: 1-423, 29 p l a t e s ,
2.
-
1945. P a c i f i c Outpost Mosses. The Bryologist 48: 45-53.3. Brotherus, V. F. 1902. Lauhoose G . Volken, Die Vegetation d e r Karolinen.
. . En&.
Bot. Jahr. 31: 1~50-453.4. Dixon, H. N. 1927. G i l b e r t I s l a n d s Mosses. Journ. Ebt. 65: 254-257.
5. F l e i s c h e r , M. 1902-1922. Die Musci der Flora von Buitenzorg. 1729 pp.,
266 p l a t e s , Leiden.
6. Frye, T. C., and Lois Clark. 1937-1947. Hepaticae of North America.
Univ. &ash. Publ., ( 1 - 1-1018, i l l u s t r a t i o n s .
7. Glassman, S. F. 1952. L i s t of Non-vascular P l a n t s from Ponape, Caroline Islands. Amer. Midland Nat. 48(3) : 735-740.
8. Horikana, Y. 19L9. Extension of Range f o r some Species of Bryophytes.
Journ. Jap. Bot. 24: 140-144.
9. P a r i s , E. G. 1904-1906. Index Bryologicus Ed. 2. Paris.
10. Stephani, F. 1902. Lebermoose
&
G. Volken, Die Vegetation der Karolinen..
Ehgl. Rot. Jahrb. 31: 450.
11. Sukari, K. 1943. Bryoflora von Micronesia I & 11. Bot. Mag. Tokyo 30: 1-14; 249-257.
12. Taylor, Pi. R. 1950. P l a n t s of Bikini. 218 pp., 78 p l a t e s . Univ. o f Mich. Press, Ann Arbor, Michigan.
Figure 1 )
Figure 2)
Figure 3)
Figure
4)
Figure 5)
Figure 6)
Figure 7)
The p a l e color, hardly green a t all, and t h e l e a f s i z e d i s t i n - guish t h i s species from o t h e r Arno species.
S v h c h n o ~ indicum
(9437B).
.
.iNote t h a t the midrib does n o t q u i t e reach the l e a f t i p , t h e r e o f t e n b e i n g a group of s p e c i d i z e d ce?.ls between the l e a f apex and t h e t i p of t h e midrib.
. . .
. , ....
Calymperes hyophyllaceum
(9425B).
. . . . ,
,The. l e i f t i p shapes a r e v a r i a b l e and of r e l a t i v e l y l i t t l e taxonomic value. The l e a f bases with t h e i r c b n s p i ~ u o u s a r e a s of l a r g e c e l l s , , c a l l e d cancellineae, and the n a t u r e of t h e l e a f . margins a r e much more dependable taxonomic characters. .In t h i s p a r t t c u l a r species, opposite the c w c e l l i n e a e and again near t h e apex, t h e margin o f t h e l e a f tends t o have a l i t t l e tooth opposite each marginal c e l l
(denticulat,ions)
.
. .Calvmperes tenerum (9310A).
-
See notes under f i g u r e 3 and in t e x t under t h e o t h e r species of Calwperes. The margins of t h e l e a v e s i n t h i s species a r e nowhere d e n t i c u l a t e . Note the d i f f e r e n c e i n shape of the cancellineae.
, , . .
Calymperes mollucensis (9L22).
Arno species.
Ectropothecium monumentorurn (9103).
Most of t h e l e a v e s of t h i s species a r e sickle-shaped. There a r e only very small t r a c e s of midrib thickenings n e a r t h e base of the leaves.
Bryum
n i t e n s (9712).The midrib extends right t o t h e apex o f these u s u a l l y strongly con- cave, boat-shaped, green leaves.
See notes under f i g u r e 3 and i n t e x t under t h e , o t h e r species of Calmperes. The dotted l i n e s j u s t within t h e l e a f margin i n d i c a t e the e x t e n t of a thickened s t r i p . This i s absent in t h e o t h e r two
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