DEPA.RTMENT OF THE INTERIOR 

ALBERT B. FALL, Secretary 

UNIT.ED STATES GEOLOGICAL Su:irVEY 

GEORGE OTIS SMITH, Director 

Professional Paper 124 

THE INORGANIC CONSTir-fUENTS OF 

MARINE INVER~rEBRATES 

SECOND EDITION, REVISED AND ENLARGED 

BY 

FRANK WIGGLESWORTH CLARKE 

AND 

WALTER CALHOUN WHEELER 

WASHINGTON 

GOVERNMENT PRINTING OFFICE 

1922 
CONTENTS. 

Introduction ........................................................................................... . 

Fora1nini.fera .......................................................................................... . 

Page. 

1 

2 

Sponges: ................................. : ....................................... -o-.................... 3 

jfadreporarian corals ................................................................ ~ . . . . . . . . . . . . . . . . . . . . 6 

.Alcyonarian corals...................................................................................... 9 

llyclroids ................................. :............................................................. 14 

A.nnelids.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 

Echinoderm.s........... ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 

1. Crinoids.......................................................................................... 17 

2. Sea urchins ..................................... ·. . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 

3. Starfishes ...................................................................................... : . 25 

4. Ophiurans... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 

5. Holothurians..... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 

6. Summary ............... ·. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 

Bryozoa ........................ ~ ......................................... ~ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 

Brachiopods............................................................................................ 36 

l\follusks .............................. ~ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 

1. Pelecypods:. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 

2. Scaphopods and amphineurans. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 

3. Gastropods..... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 

4. Cephalopods ................ ,....... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 

Crustaceans. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 

Calcareous alg~e. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 

General discussion .............................................•.......... : . ........................ ~. . . 55 

II 
THB INORGANIC CONSTITUENTS 0}., MARINE INVERTE.BRAT:ES. 

By FRANK WIGGLl!:SWORTH CLARKE and WALTER CALHOUN WHEELER. 

IN':rRODUCTION. 

1'hat 1nany rocks were once 1narine sedin1ents and that anin1als and plants shared in their 

forn1ation is one of the c01nmonpla.ces of geology. Coralline limestones: shell limestones, and 

crinoidalllinestones are an1ong the most ·familiar illustrations of this statmnent. That corals 

and shells furnish calcium carbonate to the sediments, that radiolarians and diatoms are siliceous, 

and that vertebrate anin1als, so1ne crustaceans; and a few brachiopods are more or less phosphatic are also well-known facts, which, however, have been determined in a more or less desultory way. No systematic investigation to ascertain just what substances each class of organisms 

contributes. to the sedi1nents seen1s to have been made, and the present research is an attmnpt 

to cover the gi·ound a little more thoroughly than it has been covered heretofore. Conlpleteness is, of course, impossible, but son1ething of a foundation for future work we have tried to lay. 

Although 1nany analyses of corals and molluscan. shells are on record, son1e important 

groups of organisms have received only slight attention, and little has been done heretofore 

toward deter1nining the c01nposition of their tests or skeletons. The few published data, Inoreover, have been generally but not always incomplete in certain particulars, especially with 

regard to the localities fr01n which the speci1nens analyzed were obtained,. the depth of water 

in which each creature lived, and the ten1peratu1~e of its habitat. Even in our own wo'rk s01ne 

of these details are lacking, but their great significance when known is strikingly evident in. 

the study of such groups of animals as the echinoderms and alcyonarians. Furthennore, 

some of the older analyses are unsatisfactory for other reasons, for 1nany of the1n were made 

to detennine single facts, such, for instance, as the proportion ·of 1nagnesiun1 carbonate alone; 

and in others such essential <?Onstituents as organic matter were not taken into account,· an 

omission t4at is especially serious, for it renders the accurate comparison of the analyses with 

others in1possible. In the shells of 1nollusks the proportion of organic ~natter is small, but in 

the echinoderms, alcyonarians, and phosphatic brachiopods it is relatively large, ranging fr01n .10 

to even 40 per cent or n1ore. This state1nent, however, must not be n1isconstrued; it refers, 

of course, only to the specimens, dried or alcoholic, which were actually analyzed. In order to 

compare the analyses, therefoi·e, so as to determine the true con1position of the inorganic shells 

or skeletons, the very variable mnounts of organic n1atter n1ust be rejected, and the remainders 

recalculated to 100 per cent. Relations tl~en appea.r which are not recognizable when the crude 

um·educed analyses a.re c01npared. Any one of our tables of analyses will show this fact very 

clearly. 

In a strict sense com_Rleteness can not be claimed even for our analyses. Minor constituents 

which have been detected in 1narine organis1ns, such as barium, strontium, fluorine, n1anganese, 

copper, zinc, and lead, have been ignored. They occur, as a general rule, only in traces and have 

little or no sig:q.ificance with respect to the larger problems before us. What organisms tend 

to form limestones and what ones are notably 1nagnesian, phosphatic, or siliceous a.re the questions which we are atte1npting to answer. In 1nost of the analyses li1ne, 1nagnesia, phosphoric 

oxide, sulphur trioxide, silica, and loss on ignition were determined. Alumina and oxide of 

iron were also considered and weighed 'together. The loss on ignition covered carbon dioxide, 

1 
. 0 • 

2 THE INORGANIC CONSTITUENTS OF MARINE INVEi{T'EBRATES. 

organic 1natter, and water. The carbon dioxide was calculated to satisfy the bases, ·and the 

amount thus detern1ined, subtracted from the total loss on ignition, gave a fair but rough esti-. 

mate of organic matter plus water. Frmn the crude analyses thus obtained the reduced or 

rational analyses, recalculated to 100 per cent, were computed. These rational analyses are 

comparable, and give, closely enough, the essential composition of the material which goes 

to build up the limestones. More refined work would. add little toward the solution of our 

1nain problen1 al).d would require much tedious labor. In the following pages the detailed 

. analyses are given, group by group, together with some of the older published data. After 

the evidence has been presented a general discussion of its significance will be in order. 

In the former edition of this paper (publishe~ in 1917 as Professional Paper 102) the 

analyses, with few exceptions, were made by the .iunior author. In this edition we are able to 

present .a large number of new analyses, made in order to strengthen weak series or to clear up 

some outstanding uncertainties. These analyses, which were made by Messrs. A. A. Chambers, 

R. M.. Kamm, B. Salkover, and George Steiger, are credited to the respective analysts. All 

others were made by Wheeler. 

FORAMINIFERA. 

The Foraminifera, beca~se of their enormous abundance, are of great importance in the 

forn1ation of marine sediments. The globigerina ooze, for example, covers 49,520,000 square 

miles of the ocean floor, at a depth of 1,996 fathoms, or 3,653 meters. The analyses of it published in the· report of the Challenger expedition on deep-sea deposits show that it consists 

mainly of calcium carbonate, 'Yith very little magnesia. Foraminifera are also abundant on 

"coral reefs," and great masses of limestone are composed largely of their remains. 

These organisms, however, are so small, rarely larger than the head of a pin, that it is not 

easy to obtain enough material of any one species for chemical analysis. The difficulty, fortunately, is not insuperable, and with the help of others we were able to obtain representative 

samples of seven species, as follows: 

1. Pulvinulina men'ard1:i D'Orbigny. Albatross station 2573; latitude, 40° 34' 1811 N.; longitude, 66° 09' W.; in 

line from Long Island Sound to Cape Sable; depth of water, 3,188 meters; bottom temperature, 3° C. 

2. Sphwrodina dehiscens Parker and Jones. East coast of Mindanao, Philippine Islands; latitude, 8° 51' 4511 N., 

longitude, 126° 26' 5211 E.; depth of water, 804 meters. 

3. Tinoporus baculatus Carpenter. Murray Isla_nds, Torres Strait, Australia. 

4. Orbitolitcs marginat·is Lamarck.· South of Tortugas, Fla., at depth of 29.3 meters. 

5. Orbiculina adunca.Fichtel and Moll. Key West, Fla. 

6. Polytrema mineaccum Linne. Bahamas. 

7. Quinqueloculina auberiana D'Orbigny. Same locality as No.4; analysis by A. A. Chambers. 

Analyses of Foraminifera. 

1 2 4 5 6 I 

---~ ·- . ---------------1 --

Si02 • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 14. 70 : 8. 55 0. 03 0. 03 i 0. 11 } o. 02 } · (Al,Fe)20 3 ••••••••••••••••••••••••••••••••••••••••• 3. 81 1

. 4. 76 . 18 . 13 ,. . 09 

MgO ............... ·................................. 1. 68 . 82 · 5. 03 4 . .93 4. 64 5. 09 

CaO................................................ 41.36 45.48 47.35 48.92 1 48.79 47.35 

rg~ti~~:.-::: . .-:·:::.~:::::::::::::::,:::::::::::::::::: 3~~)12 3~~·~1 46:g~ r:.a~~-i r:.a~:· 4~~)24 l------·1-------:------l------------1-----

99. 67 98.62 I 99.16 99.48 I 99.19 98.70 

34. 41 36. 63 42. 14 43. 9o 1 43. 43 42. so s. n 2. 38 I 3. 73 1. 30 

1 

2. 13 3. 44 

C02 need eel .....••...•..........•................... Organic matter, etc ................................. . 

7 

0.54 

4.32 

49.02 

(?) 

45.54 

99.42 

43. 27 

2. 27 

Rejecting organic matter and water ("ignition") and recalcula:ting to 100 per cent, the 

analyses assume the following rational form: 
SPONGES. 

·Reduced analyses of foraminifera. 

1 2 

8. 89 ! 

4. 94 ; 

1. 79 I 

84.38 I 

(?) i 

3 4 5 6 7 

InN os. 1 and 2 the high percentages of silica and sesquioxides are evidently due to in1p1urities. If they are rejected, the percentages of magnesiun1 carbonate become .5.19 and 2.09, 

respectively. These two Formninifera are fron1 deep and presu1nably cold water. The others 

aro from shallow water in warm regions. A siinilar difference in the magne!:)ian content of the 

oL·ganisnls with regard to ten1perature appears, with Inuch regularity, in several other series 

of our analyses, notably in the series of crinoids and alcyona.rians. 

The only published analyses of Fora1ninifera that we have been ·able to find elsewhere 

n.ppear in If. B. Br~dy's volume 1 on the 1naterial collected b)~ the Ohallenge1· expedition. Four 

of these analyses relate to Orbitolites complanata var. laciniata, collected either at Fiji or at 

Tonga tabu, Friendly Islands. These analyses are as follows: 

1, 2. 'l'wo diffet·ent samples; analyses by C. R. A. Wright and .T. T. Dunn. 

3. Sn.mplo washed with boiling water. 

4. Same u.s No.3, unwashed; analyses 3 n.nd 4 by J. Gibson. 

Analyses of orbitolites. 

1 2 3 4 J ·I·-

Si02 ..................... 0. 58 0. 3 · 0.14 0.11 AI~03 a................... . . . . . . . . . . 2. 7 I ................... . (Al,Fo.)20 3 •••••••••••••••• • 68 .......... 

1

1 

•••••••••••••••••.••• 

MgC03 • • • • • • • • • • • • • • • • • • • 12. 52 8. 8 9. 55 10. 50 

CaC03 ..................... 86.46 88.2 88.74 87.91 

100. 24 100. o 1, 98. 43 98. 52 

a Including "phosphates of lime and magnesia." 

Other analyses, presented by Wright and Dunn, are of Amphistegina lessvnii, f1:on1 Cape 

Verde Islands, and Operculina complanata, from near Amboyna, in the Malay Archipelago. The 

analyses are obscurely stated but give from 4.8 to 4. 9 per cent of magnesiun1 carbonate. In 

a Tertiary fossil, N'l.tbecularia novorossica, 26 per cent of magnesium carbonate was found, an 

amount which probably represents a concentration by leaching. There are also an~lyses of 

several siliceous Foraminifera containing from 76.1 to 94.7 per cent of silica. I'heir tests appear 

to have been built up of sand grains and are of little or.no significance in relation to the n1ain 

problmn of the present investigation. Their ren1ains are not true additions to the sediments. 

SPONGES. 

Tho sponges are a rather unsatisfactory group of organisms for chen1ical analysis, on account 

of their very porous structure. They are liable to contain inclosed impurities, such as sand 

or n1ud, which are difficult to ren1ove entirely. S01ne sponges are siliceous, some calcareous, 

and others are composed almost wholly of organic matter, the proteid spongin. The i9,eal 

method of procedure for our purposes would obviously be to separate the clean siliceous or calcttreous spicules and to analyze then1 alone, but to do that a large amount of material would have 

to be gonQ over, and more tirne would be consumed than we could give to this part of our investiI Hmd~r, H. 13., Challenger Rcpt., vol. 9, pp. xvii-xxi, 1884. 
4 I' 

THE INOHGANJC CONSTITUJ!}NTS OF lVIAJ.UNE INVEHT'EBHATES. 

gation. As the sponges are of minor importance in relation to the marine sedin1ents, we have· 

followed our ustutl course, and so have obtained results which are of some significance, even if 

they are not all that might be desired. Twenty sponges have been examined, more or less 

eompletely, as follows: 

1. .Enplectella speciosa Quoy and Gaimard C' Venus's fiower basket"). Philippine Islands. 

2 . .Bathydorus uncifer F. E. Schulze. Galapagos Islands, Albat1·oss station 2818; latitude, 0° 291 S.; longitude, 

89° 54' 30'' \V.; depth. of water, 617.4 meters; bottom temperature, 9.9°·C.-

3. Halidwndria panicea Johnston. Alaska. 

4. Snberites snbcrea Johnston. Unga Island and Popof Strait, Alaska. 

5. 'l'clhya gral'ala Hyatt. Buzzards Bay, l\Jar;s. 

6. l'hcronema yray'i Kent. Oil southwest coast of Ireland, at depth of 1,098 meters. 

7. Geodia mesolriama var. pachana Lendenfeld. Oil southern California, Albatross station 2H09; latitude, 32° 22' 

~.;longitude, 120° 08' 33" \V.; depth 377 meters; bottom temperature, 7.:3° C. 

8. Phakell,ia grandis \1err.ill. Browns Bank, northeast of Cape Cod, between latitudes 40° 30' and 42° 08' N. and 

longitudes 66° W.; depth, 549 meters. 

!l. Chondr~cladia alaskensis I1ambe. On beach at Unalaska, Alaska. 

10. Gell-iodcs grandis Verrill. Gulf of Maine. 

ll. .Esperiopsis q11atsinoensis Lambe. Attu Island, .Alaska. 

1.2. Chalina arbuscula Verrill. Outer Island, Conn. 

13. Spinosella sororia var. crispa Duchassaing and Michelotti. Harrington Sound, Bermuda. 

H . .E':tspongia o.Oieinalis tnhulijem var. tun·ita Hyatt. Harrington Sound, Bermuda. 

15. lfircin·ia campana var. turr'ila Hyatt. Bermuda. 

16. Eusponyia o.fjicinalis var. r9tunda Hyatt. Nassau, Bahama Islands. 

17. Ilippospongia equina var. uwandr-iformis Duchassaing and Michelotti. Nassau, Bahamas. 

18. Hippospongia agarieina var. d'ura Hyatt. Off Duck Key, Fla., at depth of 4.6 meters. 

19. !Iippospongia canie'ulala var. gossypina Duchassaing and Michelotti. Off :Rock Island, Fla., at depth of 7.3 

meterR. 

20. Aplysina hirsnta Hyatt. Off St. Thomas, ·virgin Islands; depth of water, 37 to 42 meters. 

21. C:rantia ciliata, Fabricius. \Voods Hole, Mass. \Veight of sample only 0.0982 gram. 

22. Leucilla ci:trteri, var. homoraphis (Polejaeff). Off Culebra Island, Porto H,ico, Fish Jfawk station 6090 ;. bottom 

temperature, 26_° C. \Veight of sample, 0.1422 gram. ' 

Analyses 21 and 22 by R. 1\f. Kamm. 

The analyses were made by our usual methods; Nos. 21 and 22 were made on insufficient material and are therefore 

incomplete. 

Analyses of sponges. 

---- ·- ·-- --------~--~ --------.,------------,----:-------:-----,--------:--------

4 ~~~s~J 6 J _ 7 

.1 

9 10 ' I I 

~iO, ____ :: ______ ~:- =--:+;;_;- (Al,Fe)203.......................... 1.64 

~lf8 ... · .. _· _· _·: : ~ : : : : : : : : : : : : : : : : : : ~ : : : 1: ~~ P~0 ••••••••••••••••••••••••••••••• .31 so~............... . . . . . . . . . . . . . . . . (?) 

lgni tio n. . . . . . . . . . . . . . . . . . . . . . . . . . . . •U . 97 

99 .. 74 

52.2'1 

3.86 

.86 

.77 

.61 

.81 

34.64 

93.79 

2 3 

76.33 74.52 

1.05 6.19 

.48 1.43 

2.78 2.33 

.00 .17 

Trace. .08 

18.60 12.11 

99.24 96.83 

11 12 

------1 

51.94 

1.38 

.75 

1.77 

.46 

(?) 

42.29 

98.59 

I 

31.93 ,. 1.44 

.97 

.97 j 

.50 1 

.821 61.25 

97.88 I 

72.92 I 68.46 I 65.19 I 64.96 

1. 22 I 3 . 71 I 3 . 05 I . 21 

.55 1 .61 'l'race. \ .38 

.33 l .89 3.26 .29 

.49 j . 48 'l'race. 'frace. 

.39 . .40 .02 .21 

22.21 ! 23.02 ' 27.05 31.73 

98.11. 

13 

3.08 

.77 

.26 j 2.80 

.69 

(?) 

91.32 

U7.57 98.57 97.78 

14 1 15 21 

--j-- --·--

1.31 

.96 

.4:3 

7.58 

.77 

(?) 

89.75 

1 0.57 } ! 2 29 8 .45 

I 1..60 . ·1."'t7 

20.01 38.39 . 68 (?) 

(?) .00 

75.68 50.31' 

98.92 100.80 100.83 98.92 

8_ 

60.70 

2.30 

.23 

.51 

.46 

(?) 

35.84 

100.04 

22 

0.77 

5.50 

38.97 

(?) 

.00 

53.58 

98.82 
SPONGES. 5 

Sponges Nos. 16-20 were cmnposed nutinly ·of organic matter, and only loss on ignition 

wn.s determined. '.l.'he figures showing this loss are as follows: 

Per cent. 

16. Eusponyia o.Uidnalis . .......................................................... ·.... . . . • . . . . . 96.94 

17. lli]J]Jospongia eqnina . .... · ................... ·. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96. 98 

18. llippospongia agaricina ....................................................................... 95.55 

19. Hippospongia caniculata ................................................. ·.· .................. 94.72 

20. Aplysina hirsuta .................................. . · ......................................... 93.34 

In these sponges the inorganic matter is so small that it n1ay only represent i1npurities, 

a.n analysis of which would be meaningless. Euspongict and Flippospongia are the ordinary bath 

sponges of conunerce. 

In several of the ann.lyses, notably in Nos. 3, 5, 7, 10, and 12, the stnnmation is very low, 

on account of the presence of- soluble salts (sea salt), which were not deter1nined. Analyses 

1 to 12 represent siliceous sponges, and inN os. 2 and 6 ·the siliceous spicules were very conspicuous. 

No. 1, En1Jlectella., the beautiful "Venus's flower basket," n1ay be taken as the type of these 

sponges, and the analysis shows that its skeleton is con1posed of nearl)T pure opaline silica. 

Severn.l analyses of siliceous sponge spicules by other chemists lead to similar conclusions. In 

spicules fr01n Unnnmed species J. Tho~let 2 found fron1 12.88 to 13.18 per cent of water, F. E. 

Schulze 3 cites an analysis by Maly of spicules from Poliopogon a,ma,dO'I.L which contained 7.16 per 

cent, and in seven species of siliceous sponges W. J. Sollas 4 found water varying from 6.1 to 

7.34 per cent. In all these sponges the spicules or skeletons are composed.of an1orphous opaline 

silica. The other constituents shown in our ttnalyses, except organic Inatter, which is included 

in the loss on ignition, are probably but not certainly in1purities. To this statmnent one exception 1nay be 1nade·. ·The phosphoric oxide is perhaps a part of the organisms, which contain it 

not as such but as phosphorus in some organic compound. This, however,· i!:l by no 1neans 

·certain. That nearly all the sponges analyzed contain phosphorus in some form, although in 

s1nall a1nounts is clear. 

Analysis No. 13, of S1Jin'osella, is very high in organic matter, and its precise character is 

doubtful, at least so far as the chemical evidence goes. Nos. 14, 15, 21, and22, however, represent calcareous sponges. Reducing the analyses to standard fonn, rejecting organic n1atter, 

and recalculating to 1.00 per cent, we have the following figures for their inorganic portions: 

Reduced analyses of calcareous sponges. 

14 '15 21 22 

-------------

Si02 ••••••••••••••••••.••• 7.81 l. 36 } 10.47 0 94 (AI, Fc)z03 •••••••••••••••• 5. 72 5. 45 

Mg003 ••••••••••••••••••• 5. 37 8.00 4. Li1 14. 10 

Cn.C03 ••••••••••••••••••• 7i. 14 81.. 64 84. 92 84. 96 

3~~{d~~ ·. : : : : :: : : : : : : : : : : : 

9. 96 3. 55 (?) (?) 

(?) (?) . 00 . 00 

100.00 100.00 100.00 100.00 

-~ ~---~--~-------·_.:... ____ - -----~~ -~~----- ---

We can not assign 1nuch weight to these analyses, because the specin1ens analyzed evidently 

contained impurities-silica and sesquioxides. Even the s1na:ll amounts of phosphate shown in 

two of thmn may be clue to the inclosure in the sponges of minute crustaceans, such as copepods. 

The small percentages of P20 5 in the unreduced analyses are much magnified i1:1 the reduction. 

Some of the 1nagnesia also 1nay belong to inclosed sea water, and in this respect analysis N o.14 

is especially questionable. The presence of magnesia, however, in these sponges is somewhat 

2 'L'houlct;, J., Compt;. Rend., vol: 98, p. 1000, 1884. 

a Rc:hulze, I''· 1':., Challenger llept;., vol. 21, p. 28, lSSi. 1 Sollns, W. J., idem, vol. 2;i, p. xlvili, 1888. 

-- -~- -----~ -~--~ --- ---------------------
6 THE INORGANIC CONSTITUENT'S OF MARINJ<J INVERTEBHATJ~S. 

confirmed by an analysis by 0. Butschli 5 of ·spicules from a Mediterranean sponge, Leucandra 

aspera. His figures are as follo~s: 

CaC03 •••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••• 

J\fgC03 •••••••••••••••••••••••••••••••••••••••••••••• : •••••••••••••••••• ~ •••••••••••••••••••• 

Fe20 3 •••••••••••••••••.••••••••••••••••.••••••••••••••••••••••••••••••••••••.•••••••••••••••• 

CaSO.j-2H20 ............................... -.. ·- · · ·- · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · H20 ...................................................................................... . 

Organic matter ............................................................................ . 

86. 76 

6. 84 

. 26 

1. 97 

3. 14 

. 42 

99. 39 

If we reject w~'tter and organic 1natter and recalculate to 100 per cent the percentage of 

1nagnesium carbonate 'rises to 7.17. It seems probable, then, that the calcareous sponges are 

distinctly magnesian, although more analyses are needed to establish the fact. 6 

. 1\fADREPORARIAN CORALS. 

Madreporarian or stony corals have been repeatedly analyzed and with generally concordant results. Twenty-eight of these corals were studied in the course of this investigation, · 

and with the results two old analyses made in the laboratory of the United States Geological 

Survey may properly be included. The order of arrangement' here is regional, for reasons 

which will appear later, in our study of the ·alcyonarians. The list of corals analyzed is as 

follows: 

1. Balanophylliajloridana Pourtales. South of Key West, Fla.; depth of water, 165 meters. 

2. Paracyath'us defilipii Duchassaing and Michelotti. South of Key West; depth, 165 meters. 

3. Dendrophyllia cornucopia Pourtales. Fish Hawk station 7286, Gulf Stream, off Key West; latitude, 24° 18' 0011 

N.; longitude, 81° 4;71 4511 W.; depth, 243 meters; bottom temperaturj3, 11.9° C. · 

4. Siderastrea radians Pallas. Shoal water, Ragged Keys, Fla. 

5. Poritesjurcata Lamarck Shoal water, Ragged Keys. 

6. Faviajmgum Esper. Shoal water, Tortugas, Fla. 

7. Eusmilia aspera Dana. Shoal water, Tortugas. 

8. Oculina diffusa Lamarck. Shoal water, Tortugas. 

9. Cladocora arbuswla Lesueur. Tortugas; depth, 27~5 meters. 

10. Agaricia purpurea Lesueur. Reef, Loggerhead Key, Tortugas. 

11. Orbicella annularis Ellis and Solander. Reef, Loggerhead Key. 

12. Dasmosmilia lymani Pourtales. Albatross station 2087; latittide, 40° 061 50'1 N.; longitude, 70° 34' 1511 W.; 

.:epth, l19 meters: bottom temperature, 10° C. 

13. Flabellum alabastrwn Moseley. Albatross station 2677; latitude, 32° 39' N.; longitude 76° 50' 3011 W.; between 

Lhe Bahamas and Cape Fear; depth, 875 meters; bottom temperature, 4.5° C. 

14. Madrepora polifera Linne. Albatross stations 2662-2672, between the Bahamas and Cape Fear; depth, 512-894 

meters; bottom temperature, 6°-7° C. 

15. DendroJJhyllia JJrofunda Pourtales. Locality, etc., as under No. 14. 

16. Madracis decactis Lyman. Shoal water, Bermuda. 

17. Deltocyathus italicus Michelotti. Albatross station 2750; latitude, 18° 30' N.; longitude, 63° 311 W.; depth, 

!)08 meters; bottom temperature, 7° C. · ' 

lS. Ac1"opora cervicornis Lamarck. Shoal water, South Bight, Bahamas. 

19. Orb1:cella cavernosa Linne. Reef, Light House Point, Andros Island, Bahamas .. 

20.· Jlfmandra cli1;osa Ellis and Solander. I.ocaJity as under No. 19. 

21. Jfwandra labyrinthiforrnis Linne. Reef, Golding Cay, Andros Island, Bahamas. 

22. jl[ussa aff. Jf. dipsacea Dana. Reef, Golding Cay. · 

23. Porites cla·va.r-ia Lamarck. Reef, Golding Cay. 

24. Porites astreoides Lamarck. Reef, Golding Cay. 

25. Flabellum pm;onium var. pa1"ipavonimtrn Alcoc:k. Albalios~> station4080; north c:oast of Maui, Hawaiian Islands; 

depth, 326 to 369 meters; bottom temperature\ 13.5° C. 

26. Jlfadracis kauaiensis Vaughan. Albatross station 3982; .vicinity of Kauai, Hawaiian Islands; depth, 426-435 

meters; bottom temperature, 9.1° C. 

----------------------------- -- ----------------------

46 Butschli, 0., K. Gesell. '\\riss. Gottingcn Abh., No.3, 1908. 

6 On manganese and iron in sponges, sec Cotto, J ., Soc. biologic Compt. rend., vol. 55, p.139, 1903. '!'races ofmanaganese were noted in our own 

analyses. 
MADREPORAI~N CORALS. 7 

27. Flabellum sp. Albatross station 5273; latitude, 14° 03' N.; longitude.120° 27' 4511 E.; China Sea, near southern 

Luzon; depth, 102.5 meters. · ' . 

28. Desmophyllum ingens Moseley. Albatross station 2785; latitude, 48° 09' S.; longitude, 74° 36' W.; off coast 

of Chile; depth, 822 meters; bottom temperature, 8.3° C. 

The actual analyses appear in the following tables. Sulphates were rarely determined, 

but they are unimportant. 

Analyses of corals. 

1 l ·2 1 3 

-;------·-

SiOz ............................................... _i 

~N.~e!:~s:::::::::::::::::::::::::::::::::::::::::: 803 •••••••••••••••••••••••••••••••••••••••••••••••• 

rg~ti~n· .·: : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : 

C02 needed ........................................ . Organic matter, etc ................................. . 

I 

0.09 

. 69 

. 49 

51.46 

(?) 

Trace. 

47.32 

100.05 

39.97 

7.35 

8 

0.42 0.09 . 35 .11 . 39 .19 

52.03 51.71 . 00 (?) 

Trace. Trace. 

44.62 46.23 

97.81 98.33 

41.31 40.84 

3.31 5. 39 

9 10 

4 5 6 

I· -

0. 12 0. 11 0.28 

.11 .10 .12 0 21 . 36 .18 

51.23 51.61 53.69 

(?) (?) (?) 

·Trace . . Trace. Trace. 

47. 21 46.76 44.44 

98.88 98.94 98.71 

41.48 40.95 42. 38 

5.73 5. 81 2.06 

I 11 12 13 i --1- -.I. 

Wl" !:~~ ·; ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ 803 •••••••••••••••••••••••••••••••••••••••••••••••• 

rg~1i~n" .· .· : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : 

002 needed ........................................ . Organic matter, etc ................................. . 

0.07 

. 05 

.19 

53.90 

(?) 

Trace. 

44.57 

98.78 

42.56 

2.01 

15 

~;;·~·;·: ~ ~ :~·;-;·;· ~ ~ ~: ~ ·;-;·;· ~ ~ ~:: :;·;·;·i·j; ~-;·;·:· :: ~: ~ -";-;-:!.:: ii r:~~~·:·::: :-:·:·:·: ::::·:·:·:·::::::::: ;: ::::::::::::::: :! i~:9. 

98.77 

C02 needed ......................................... 1 42. 35 

Organic matter, etc .................................. I 2. 14 

I 

22 

~~~~!:~~~::~~~:~::~~:::~~~:::~~~~~::::~:~~~:~:~~~ .:~~ r:~1~~-::::::::::::::::::::::::::::::::::::::::::: 'ril.~:i 

I 

0.30 0 04: 0. 00 1 o. 20 I 0.09 

. 08 . 00 i .. 03 . 27 .11 

. 05 . 28 ; . 27 : . 29 .17 

53.31 53.87 I 53.58 53. 79 53.54 

(?) (?) (?) .10 (?) 

Trace. . 00 . 00 Trace . Trace. 

44.83 44.62 1 44. 76 44.47 . 44.80 

98.57 98.81 1 98.64 99. 12 98.71 

41.94 42.641 42.40 42. 52 42. 26 

2. 89 1.98 2.36 1.95 2.54 

16 I 17 . I 18 19 I 20 

I I 

-0.21 I ·---'!'-

I 

0. 04 i 0.00 0. 09 : 0.02 

.13 I . 25 l . 06 o 00 I . 04 . 34: . 25 i . 20 . 35 . 34 

52. 78 I 53. 38 : 52. 47 53.07 53.70 

(?) I (?) : (?) (?) (?) 

Trace. I Trace. ; Trace. . 00 . 00 

45. 33 I 44. 49 i 45. 77 45,23 I 45. 33 

·---· 

98.621 98.58 I 98.50 98.74 : 99.43 

41.84 42,41 I 41.45 42.09 : 42. 56 

3. 49 I 2. 28 : 4.32 3.14 I 2. 77 

I 

23 24 25 26 27 

0.04 0.02 0.40. 0.06 0.07 

. 09 . 02 . 53 : .11 .15 

.17 .18 . 20 . 15 ; .17 

52.12 ' 53. 84 53. 22 53. 25 I 52. 94 

(?) (?) (?) ( ?) (?) 

Trace. \ Trace. Trace. Trace. J Trace. 

45. 84 _I 44. 98 44. 31 i 44. 80 45. 28 

98.66 ·1 98.37 ! 

, ___ , __ _ I 98.38 98.26 99.04 98. 61 

41.14 42.50 42.041 42.81 ; 41. 79 

3.70 2.48 2. 27 1. 99 1 3.49 

I 

C02 needed ........................................ ·1 41. 95 

Organic matter, etc. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. 86 

7 

0.00 

.04 

. 21 

53.25 

(?) 

Trace. 

44.90 

98.40 

42.08 

2.82 

14 

0. 19 

. 07 

. 06 

53.48 

.12 

Trace. 

44.50 

98.42 

42.03 

2.47 

21 

0.00 

. 04 

. 26 

53. 71 

(?) 

Trace. 

45. 10 

99.11 

42.49 

2. 61 

28 

0.13 

. 06 

. 27 

53. 69 

(?) 

Trace. 

44.64 

98.79 

42.48 

2. 16 
8 THE INORGANIC CONSTITUENT'S OF MARINE INVEHTEBHATES. 

The two other analyses of corals, which were made some years ago in the Survey laboratory, I . 

do not appear rn the foregoing table because they are stated in somewhat different form. They 

are, however, included in the following table of redQced analyses, in which the figures a:ferecalculated to 100 per cent and as definite salts, with organic matter and water rejected. The 

two in question are as follows: 

29. Siderastrea sp. Bermuda; analysis by L. G. Eakins. 

30. Coral, species undetermined. Panama; analysis by W. T. Schaller. 

Reduced analyses of corals. 

s:ioz .......... : ................... : ................. . ( 1\l, li'e ):.!0 3 .••••• '.•••••.•.•••.•.•••••.•••••.••.•••••••• 

lVlgC03 •••••••.•••••.•.•••••••.•••••••••••••••••.••••• 

Ca003 .••••.•••••.•••••••••••••.••.••.••••••• · •••••••• 

CaS<)., ............................................... . Ca3P20 8 ••••••.••••••••••••••••••••••••••••••••••••••• 

1 

0.10 i . 74 : 1. 11 

98. o5 1 

('?) 

Trace. i 

I 

2 3 4 i 5 

_: ___ J __ 

0. 44 I 0. 10 : 0. 13 I . 37 . 12 ; . 12 

. 87 : . 43 ' . 4.8 

98. 32 I 99. 35 i 99. 27 

. 00 ' ('?) : (?) 

Trace. , Trace. 1 Trace. 

0. 12 

.11 

. 82 

99. 95 

('?) 

Trace. 

100. oo 1 

I 

100. oo -~ 100. oo 1 100. oo 100. oo 1 

I 

6 

0. 29 

.12 

. :39 

99. 20 

(?) 

Trace. 

100.00 

7 8 11 12 

0. 00 0.07 . 04 . 05 

. 46 .. 41 

99. 50 99. 4.7 

(?) (?) 

Trace. Trace. 

100. oo 1 100. 00 

13 14 

. I_: rl?,i,~);o;·:.-.-.: :::.-.-.-.:::::::::::::::::::::::::::::::::I 

. ~!~L:::·.:·:·:---~::::.~::·:. ~:: ~:::::: ~ ~ ~: .. :::: ~:: ~ ~: ~ ~:: ~! 

0. 09 0. 20 . 

.12 . 07 

. 37 . 14 ' 

99. 42 

1

, 99. 38 ; 

(?) .. 21 ' 

15 

0. 34 : 

. 01 1 

.12 ' 

99.47 1 

(?) ' 

16 

0.04 

.14 

. 76 

99.06 

(?) 

Trace. 

17 

0. 22 

. 26 

. 54 

98. 98 

(?) 

Trace. 

18 

0.00 

. 06 

. 45 

99. 49 

(?) 

Trace. I Trace. Trace. 

100. oo !"1oo. oo [ 

Trace. i 

-------:------1-------

100. 00 I 10~. 00 I 100. 00 I 100.00 

~-1-9 _i 20 I 

----------------------------------, ; i 

!!irt\~T//.: ~ ~: (: [ [ ~ ~: ( [ [ ~ ~ [ ~ [ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ [ [ [ [ ~ ~-~~) !! · __ ~;_) :_~!_l_,r_t~-~-~f__ :--T-~~-~-~_-_T_~;_lc_!!____1~_<;_~~--~ I I 100. 00 100. 00 I 100. 00 'I 100. 00 100. 00 100. 00 

21 22 

-· ----- -----

23 24 

25 26 27 28 29 30 

---------------------------------------------·-------1-------l------lSi02 •••••••••••••••••••••••••••••••••••••••••••••••••• 0. 41 i 

(Al,Fe),o, ............................................ 

1 

. 55 , 

[fiL::.~·:·:-:::::·::: ~: ~ : ~ ~ ~ : ~ ~ ~ : ~ ~ ~ : ::: : ~ : ~ ~ ~ : ~ : ~ ~ ~ : : ~ : 1-'I-~:_;c_:~_. :-------:------\·-----

. · I 1oo. oo · 

0.06 0.07 

.11 .15 

. 33 . 37 

99. 50 99.41 

(?) (?) 

Trace. Trace. 

100. oo 1 1QO. 00 

0.14 0. 23 1. 25 

. 06 Trace . . 59 

. 59 . 42 ' . 59 

99.21 J,j 99. 35 97.57 

(?) (?) (?) 

Trace; (?) ('?) I 

100.00-1 100.00 100.00 
I 

ALCYONAlUAN CORALS. 9 

These analyses are rmnarkably concordant and show that the stony corals contain little 

besides calciun1 carbonate. Silica and sesquioxides are probably extraneous; 1nagriesia is altogether subordinate, although fairly regular in its mnount; phosphates occur only in traces. The 

older analyses all tell the same story, except that in six analyses by S. P. Sharples 7 of corals 

:fr01n the Gulf Strean1 frmn 0.27 to 0.84 per cent of calciun1 phosphate was detern1ined. In' 

:five analyses of Brazilian corals, L. R. Lenox 8 found frmn 0.42 to 0.54 per cent of 1nagnesimn 

carbonate, quantities like those appearing in our tables·. In short, the un.iformity of the data is 

so n1arked that it is unnecessary to reproduce all the published analyses here. 0 

.A.LCYONARIAN COU.ALS. 

The alcyon.arians, which include the red corals, the gorgonias, or sea fans, and other similar,. 

genm!ally branching forn1s, differ from the 1nadrepores in being notably n1agnesian. To this 

rule one exception is known, the genus Heliopota, which stands quite alone and rese1nbles the 

ordinru:y-_corals not only in structure but also in chemicnl con1position. 10 With this exception, 

the alcyouaria are chemically sim.ilar to the crinoids, but smne genera n.re nnteh richer in phosphates. In some respects the alcyonarians are diffLcult to deal with analytically, for 1nany of 

then1 hn.ve a horny axis, cmnposed chiefly of organic nuttter, surround~d by a cortex or envelope 

which is largely calcareous. These two portions .of the structure are so unlike in con1position 

tha.t itnperfections in a san1ple taken for analysis niay lead to very uncertain conclusions. 

The list of alcyonarians analyzed is as follows: 

l. .fleliOJ)Ora. cemlea. Pallas; blue coral. Soutbe~·n Philippine Islands. The. blue color consists of organic matter.ll 

2. 1."'ubipora. JnLrJntrea Lamarck. Singapore, Straits Settlements; latitude, 1° 201 N.; longitude, 103° 50' E. 

3. Comllium, elatior H.idley; a red coral. Murotsu, Tosa (Shikoku), Japan; latitude, about 33° N. 

4. Printnoa. reseda. Ven·ill. East of Nova Scotia; depth of water, 366 iueters; latitude, 42° 16' N.; longitude, G3° 15' \V. 

5. Le1)idis1:s caryOJ)hyllia Verrill. Off N autucket Shoals, Albatross station 2037; latitude, 38° 53' 0011 N.; longitude, 

(i\) 0 23' 3011 W.; depth, 3,108 meters; bottom temperature, 3.3° C. 

6. Pennatula aculeata Dana. St. Peters Bank and Banquereau; Albatross station 2470; latitude, 44° 47' N., 

-longitude, 56° 33' 4511 \V.; depth, 410 meters; bottom temperature, 4.5° C. · 

7. Pa.rmn·w·icea bo1·eal1:s Verrill. Southwest edge of the Grand Banks; depth, 641 to 732 meters. 

8. Paragorgia arborea Milne-Edwards and Haime. La Have Ridge, off Nova Scotia. 

9. Alcyonium ca~neum L. Agassiz. Albatross station 2468, off Newfoundland; latitude, 45° 11' 3011 N.; longitude, 

55° 511 30'1 W.; depth, 70.9 meters; bottom temperature, 0.5 °C. 'l'his analysis, of very pure material, replaces the 

unsu.tisfactot·y one given in the former edition of this paper. Analyst, H.. M. Kamm. 

10. Gorgonia s1~llmticosa Dana. Fiji Islands. Cortex and axis. together. 

11. Gorgonia acerosa Pallas. East end of Long Cay, Nassau, Bahamas; latitude, 25° 5' 611 N. Cortex and axis. 

12. G01·gonia. acerosa.. Caesars Creek, southern Florida; latitude, 23° 30' N., approximately. Cortex alone. 

13. 1lfuricea humilisl\Iilne-Edwards. Parahyba do Norte, Brazil; latitude, 7° to 8°S. 

14. M-uricea echinata Valenciennes. Cape San Lucas, Lower California; latitude, 22° 52' N. 

15. Plexaurella grandijlora Verrill. Mar Grande, Bahia, Brazil. 

16. Ctenocella J)ectinata Valenciennes. Torres Straits, Australia; latitude, about 10° S. 

17. X1:7Jlwgorgia anceps Pallas .. Caesars Creek, southern Florida; latitude, 23° 30' N., approximat-ely. 

18. Rh:i7Jidogorgia jlabelltmn Linne. Bermuda; latitude, about 32° N. 

19. Rhi])idogorgia .flabellum. :East side of Andros Island, Bahamas; latitude, about 25° N. 

20. Leplogorgia pulchra Verrill. La Paz, Gulf of California; latitude, 24° 2G' N. 

21. Leptogorgia rigida Verrill. Cape San Lucas, Lower California; latitude, 22° 52' N. . 

22. Phyllogo1·gia qv.ercifolia. Dana. Fernando de Noronha, Brazil; latitude, 3° 50'S.; longitude, 32° 25' W. 

T Slmrples, S. P., An;_, .Tour. Sci., 3d ser., vol. 1, p. 168, 1871. __ 

s .Lenox, :L. R., Harvard Coli. Mus. Comp. Zoo!. Bul1.1 vol. 44, p. 264, 1904. In a memoir by J. C. :Branner on sandstone reefs. 

o Sec also Silliman, 13., jr., Am. Jom. Sci., 2d ser., vol. l, p. 189, 184o, for 31 analyses of corals from the South Pacific and the Indian Ocean; 

Forchhn.mmer, G.", Neues Jahrb., 1852, p. 854, on magnesia in shells and corals; Nichols, H. W., Field Columbian Mus. l'ub. 111, p. 31, 1906; also,· 

for determinations of magnesia. In Gcr.laria drerlalea from Abyssinia Nichols found 0.35 per cent of MgCOa. A. G. Hogbom, (Ncues .lahrb., 1894, 

Hand 1, p. 262) in two Bermuda corals found 0.36 and 0.62 per cent of MgC03. 

1o Sec Moseley, H. ,V,, Philos. 'fmns., vol. 166, p. 91, 1876, on tho structural relations of Heliopom. 

II Sec Moseley, H. W., op. cit., and Bourne, G. C., Philos. Trans., vol. 186, p. 455,1895. 

'.:. 
10 THE INORGANIC CONSTITUENTS OF MARINE INVERT'EBRATES. 

Analyses of alcyonarians. 

1 2 3 4 5 6 .7 8 

+- ------ ----

Si02 ••••••••••••••••••••••••••••••••••••••• 0.14 1. 36 0.00 0.09 0. li 1. 21 0.26 0.13 

(.A..l,Fe)z03 .................................. . 07 . 55 .14 . 64 . 05 . 72 .18 . 02 

·~:3.-.-~:::::::::::::::::::::::::::::::::::: .16 5. 67 5.44 2. 15 3. 16 2. 62 2.27 3. 71 

54. 42 46. 60 48. 65 37. 76 49. 87 35. 67 29. 92 43.47 

803 •••• ~---······ ••••••••••••••••••••••••••• . 29 . 68 . 77 .71 . 37 . 35 1. 64 1. 10 

ig~i ti~;;_· .-.·. ·_ ~:::::::::::::::::::::::::::::: Trace. Trace. .18 . 28 Trace. 1. 02 . 39 . 22 

44. 33 44. 75 46.01 57. 58 45. 70 59. 26 77.32 50.48 

·99. 41 99. 61 101. 19 99. 21 99. 26 100.85 100.98 99. 13 

C02 needed ................................ 42. 78 42.48 42. 87 31.38 . 42.43 29.76 24.74 37.44 

Organic matter, etc ......................... 1. 55 2.27 3. 14 26. 20 3.27 29. 50 41. 58 \13.04 

9 10 11 12 13 14 15 

-- --- -·- ------ --- -

~~~?, :F~)i)a·-·_·_ ·. ~: ~ .-.-.·. ·_ ~:::::::::::::::::::::::::::::: 0. 88 0. 26 0. 13 0.03 0.44 0.09 0.38 

Trace. .13 .13 . 13 . 06 . 05 .13 

MgO ................................................. 1. 86 3.03 3.54 3. 51 5. 25 4. 58 5.61 

CaO ................................................ 29. 92 22. 32 28. 73 26.48 42. 14 38.04 40.94 

803 •••••••••••••••••••••••••••••••••••••••••••••••• . 74 1.51 . 68 1. 33 . 89 1. 35 Trace . 

ig~\i~;;.- _. _. _. _. _· _-.-_·_ ~ ~: ~::::::::::::::::::::: :·::::::::: 1. 43 . 10 . 99 .73 . 24 . 30 Trace . 64. 78 71. 46 62. 34 63. 49 51. 59 54.09 52. 49 

----

99. 61 100. 81 96.54 95. 70 100.61 98. 50 99. 55 

U02 needed ......................................... 23. 88 19.94 25. 17 23. 26 38. 13 33.91 38.34 

Organic matter, etc .................................. 40.90 51. 52 37.17 40. 23 13. 43 20. 18 14. 15 I 

The low su1nmation of Nos. 11 and 12 is due to inclosed or adherent sea salts. In No. 12 

the amount of water-soluble matter was determined and proved to·be 4.71 per cent, of which 

0.55 per cent was 803 • Applying these corrections to the analysis the summation becomes 

99.96. 

16 17 18 19 20" 21 22 

-~----- ---

Si02 •••••••••••••••••••••••••••••• ~ ••••••••••••••••• 0.17 0.08 0.12 0.16 0.06 0.21 0.26 

(Al,Fe)20 3 ••••••••••••••••••••••••••••••••••••••••• .10 . 04 .16 . 06 . 02 .16 .19 

~:3. ~ ~ .-.: : : : : : : : : : :': : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : 5. 90 3.52 3.42 4.22 4.60 5.14 5.59 

37.05 27.19 27.41 32.23 33.59 3,6.15 34.62 

803 ••••••••••••••••••••.••••••••••••••••••••••••••• . 79 1. 28 . 80 1.17 1. 21 .93 . 93 

Pz05··············· ....................... ·········· . 32 . 51 . 28 . 86 2.67 2. 78 2.97 

Ignition ........................................... 56.'!5 65.56 65.67 62. 10 55.89 55.39 55.28 

100.48 98.18 97.86 100.80 98.04 100. 76 99.84 

C02 needed ......................................... 34.87 24.07 25. 60 28.51 28.31 30.96 34. 14 

Organic matter, etc .................................. 21.28 41.49 40.07 33.59 27.58 24. 1:3 21. 14 

The reduced or rational analyses are as follows: 

Reduced analyses of alcyonarians. 

I i 5 6 7 

I 1 2 8 I 3 4 

I· 

I - .. 1·- ··-- - - -- ----------- --- I 

SiO~ ....................................... 0.15 1. 40 I 0.00 0.13 0. 11 1. 70 0.44 i 0.15 

(Al,Fe)20 3 •••••••••••••••••••••••••••••••••• . 07 . 57 

I 

.15 . 88 . 05 1.01 .30 . 03 

~f33aa: ." _- ~: :·: : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : . 35 12.23 11. 56.! 6.18 6.92 7. 71 8.03 9.05 

98.93 ! 84.61 86.57 I 90.39 92.24 85.62 85.11 ; 88.04 

CaS04 •••••••••••••••••••••••••••••••••••••• . 50 1.19 I 1. 32 I 1. 59 .-68 .84 4.69 2.17 

Ca3P:t08 ••••••• · •••••••••••••••••••••.••••••• Trace. Trace. . 40 . 83 Trace . 3.12 1.43 . 56 I 

100.00 100.00 I 100.00 I 100.00 100.00 100.00 100.00 100.00 

I I I 
ALCYON AlUAN CORALS. . 11 

Reduced analysis of alcyonarians-Continued. 

I 

9 10 11 I 12 13 I . 14 15 

--------- ----------~--- ---- . --j-· --------~--·-----

Si02................................................ 1. 50 I 0. 55 0. 22 0. 04 0. 50 0. 11 0. 45 (Al,Fe):.~0 • • • • • • • • • • • • • • • • • • • • • • • • • • • •.• • • • • • • • • • • • • Trace., . 28 . 22 . 24 . 07 . 06 . 15 

MgC03 •••• .......................................... 6. 66 13.43 121 52 13.29 12.64 12.28 13.79 

Cfi.C03 ••••••••••••••••••••••••••••••••••••••.••••••• 84. 50 

1 

79! 84 81. 45 79.48 8.4. 47 83. 79 85. 61 

CaS04 ••••••••••••••••••••• ~.. • • • • • • • • • • • • • • • • • • • • • 2: 15 5: 43 1. 95 4. 08 1. 73 2. 93 Trace. 

Ca3P20 8 ................................... .'......... 5.19 . 47 3. 64 2~ 87 . 54 1 • 83 Trace. 

• 100. 00 1100. Oil' 1100. 00 10Q. 00 100. 00 100. OQ 100. 00 

16 

-- -·-- ------ -------------1- I _I .. 

17 18 19 20 21 22 

~~~.:.~~::.~.~~ ~ ~ ~:: ~ ~ ~: ~:::: ~:::: ~ ~ ~ ~: ~ ~ ~ ~ ~ ~~: ~ ~ ~ ~ ~ ~;: H ~;: ~ ~;: ~ ~;ji ~a1 , ~;JI ~i~ U 

CaS0 4 ............................................. 1. 69 3. 83 2. 40 2. 95 2. 91 : 2. 07 2.11 

CaaP 

208 ............................................ ·l-1-0-0-: :-:-,!-1-0-~:-:-:-l-1-0~-:-:o- 9 

-l-1-0:-: -::-l--10-:-: :-:-~~·-1-0-.:·-. :-:-l--1-0:-: -:: 

Analysis No. 1, that of Heliopora, might be an analysis of an ordinary coral, being nearly 

nonmagnesian and hence different fron1 the others. 

In two of the Gorgonias the black wiry axis was separately analyzed, although the amount 

of material was very small. The axis of a Rhipidogorgia sp. from Bermuda was als<? studied 

and to better advantage. The three analyses are as follows: 

Gorgonia Gorgonia Rhipido- acerosa. suffruticosa. gorgia? 

SiO~.......... .. .. .. .. . .. . . .. . 0. 04 0. 39 0. 07 

(Al,J.<'eh03 • • ... .. .. .. .. • .. • .. .. • 10 . 06 Trace. 

~:3:. ~ .·. ·:. ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ : ~~ 1J8 1

: g~ 

§6~~ ~ .'. ~ ~ : : ~ : ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ : : ~! _Tr;~e5 ·1Jg Ign1t10n . .. .. .. .. . .. .. .. . . .. . . 96. 45 94. 39 95. 83 

1----------1----------1-------

99.08 98.39 99.48 

The axes, evidently,. are COlnposed largely of organic matter, and the inorganic portions 

differ in c01nposition fr01n the more abundant calcareous envelopes. In two of them magnesia 

is in excess of li1ne, and in all three the sulphates are conspicuous. To determine the exact 

chemical nature of these axes would require much material and a longer investigation than 

we are justified in attempting. 

A specin1en of Leptogorgia flam mea V en·ill, fron1 the Cape of Good Hope, was also examined, 

but it was not sufficiently perfect. Much· of the cortex had been broken away, leaving an 

excess of the axis. The partial· results obtained, however, showed an exceedingly high proportion of phosphates, and a complete study of the species is much to be desired.. Indeed, 

the genus Leptogorgia well deserves an exhaustive investigation, but perhaps more on biological 

than on ·geological grounds. The fact that it contributes both magnesium carbonate a·nd 

calciu1n phosphate to the marine sediments seems to be established. 

Several other analyses of alcyonarians, partial or complete, are on record, and three of 

them deserve reproduction here. The data are as follows: 

1. Pleurocorallium johnsoni Gray: From latitude 25° 451 N., longitude 20° 12' W., about 160 miles southwest of 

the Canary Islands; <;lepth_of water, 2,791 meters. At that depth the temperature must have been very low, probably 

not much above 0° C. Analysis by ,Anderson in Challenger Report, Deep-sea Deposits, p. 465, 1891. 
12 THE INORGANIC CONSTITU:l!:NTIS OJ? MAlUNE INVERTEBRATES. 

2. Alcyonium palmatttm. Probably Mediterranean. 

3. Corallium rubrum; the precious red coral. Mediterranean. 

Analyses 2 and 3 by 0. Biitschli in K. Gesell. Wiss. Gottingen Abh., No. 3, 1908. 

0 lder analyses of alcyonarians. 

- --------- ---;-----~-----

. I 

1 2 3 I 

___ ! _____________ -------

CaC03 • • • • • • • • • • • • • • • • • • . • • • • • • 93. 39 ; 86. 25 86. 78 

MgC03 • • • • • • • • • • • • • • • • • • • • • • • 6. 00 \ 9. 38 8. 97 

"Phosphate" ................. · ............ I 1. 00 . 70 

~i~:~i~;b:_:_::::::::::::::::: }_ ..... ·. ~~ __1- ..... ~--~~ ....... -~--~~ .. 

Si02 or sand................... . Trace. I 1. 36 ........... . H 20.......................... . . 30 'i .. 82 1. 46 

Organic matter ...... : . . . . . . . . . . . . . . . . . . . . . . 37 . 06 . Insoluble . . . . . . . . . . . . . . . . . . . . . . 05 I· ................ · . · . · · . 

99.84 1 100.33 99.32 

The reduced analyses, re.iecting obvious impurities, are as follows: 

Reduced analyses of alcyonarians. 

I. ~e~b;::::::.·::.::::::::::::::: I (~}ace. 1:: ~: :-:2:.::: :!~~::: ::::: ~~ I 

Mg003 ••••••••••••••••••••••••• 6. 03 9. 63 1 9. 18 

Ca003 ••• •• • • • • • • • • • • • • • • • • • • • • 93. 87 88. 42 1 88. 84 

caso •.................................... · . 93 ' 1. 26 1 Ca3P20 8 •••••••••••••••••••••••• • 10 i 1. 02 . 72 

I I 

I 1oo.oo 100.00 100.00 

That these analyses fit well into our own series is evident. 

In a worked specimen of Oorallium rubrrttm, H. W. Nichols found 9.32 per cent of magnesium 

carbonate, not far from Butschli's figure. In Eunicea tourneforti and Plexaurella dichotoma, both 

from the Bahamas, he found, respectively, 2.78 and 2.11 per cent. These percentages, how'" 

· ever, are based on crude material; that is, without rejection of organic matter and water. 

They are, therefore, unavailable for comparison with our reduped analyses. Three similar 

analyses by G. Forchhammer 12 are also recorded, of Tubipora musica, Oorallium nobile ( = 0. 

rubrnm), and Isis hippuris, which gave, respectively, 3.83, 2. 73, and 6.36 per cent of niagnesiun1 

carbonate. These data lack sufficiently explicit details to admit of comparison ·with ours. 

Why his figure for the red coral is so low is not clear. The material taken for analysis must 

have been quite different from that used by Nichols or Butschli. A still earlier analysis of 

red coral by Witting 13 gave 3.50 per cent of magnesium carbonate. The speci1nen -analyzed, 

however, contained about 12 per· cent of ill!purities. · 

If, now, we arrange the alcyonarians in the order of ascending 1nagnesium carbonate, a 

remarkable relation connecting composition with temperature appears. Heliopora, being 

anomalous, is not included in the table. The three analyses by Anderson and Butschli are, 

however, inserted. Localities and latitudes are abbreviated. The percentages of magnesium 

carbonate are from the reduced analyses, and those of calcium phosphate are also given. 

12 Forchhammer, G., Neues Jahrb., 18.52, p. 854; ' 

\ 

13Witting, E., Annalen Chern. n. Pharm., vol. 1, p. 119, 1832. 
ALCYON ARIAN CORALS . 13 

.~.lfagnesiunt carbonate and calcium phosphate in a.lcyonarians. 

Species. 

I 

Locality. Latitude. I CaaP20 8• MgC03• 

Atlantic... . . . . . . . • . . . . . 25° 45' N ..... ..1-. ~~- . 6. 03 

Nova Scotia. . . . . . . . . . . . 42° 16' N ..... ·' . 83 6. 18 

----------

Plcurocorallium jolmsoni ... _ ..................... . Primnoa t·eseda ................................. . Newfoundland........ .. 45° 11' N ....... : 5. 19 6. 66 

Off Nantucket .......... 38° 33' N .... · ... ! Trace. 6. 92 

Atlantic ................ 44° 471 N ....... ' 3.12 7. 71 

Alcyonium carncum. _ ........................... . Lcpidisis caryophyllia ........................... . Pennatula aculcata. _ .... _ .... _. _ ................. . Pn.ramuricea borealis ............................. . Grand Banks.... .. .. .. . .. .. .. .. .. .. .. . . 1. 43 8. 03 

Nova Scotia ............................ · . 56 9. 05 

Mediterranean ............. .' .... _ ....... I • 72 9. 18 

Parafio·?rgia arborea ..... _. _ .. __ . _ .. __ . ,1 •••••••••• _ •• 

Coralmm rubrum ............................... . 

Mediterranean? ......... _............... 1. 02 9. 63 

Japan .................. 33° N ........... : . 40 11.56 

Singapore............... 1° 20' N _ .... _ .. ' Trace. 12. 23 

Cape San Lucas ........ 22° 52' N .. __ .. _ . 83 12.28 

Bahamas.............. .. 25° 51 N ....... .' 3. 64 12. 52 Brazil~ .... _ ............ 7°-8° S..... .... . 59 12.64 

Alcyonium palmatum ..... _. _ .................... . Corallium elatior ...... _ .. _ ...................... .. Tubipora purpurca ............................... . :Muricea cchinata. _. _ .. _ ......................... . Gorgonia acerosa ............... _ ................. . 1\tur.icea humilis ..... _ .... _ ............ _. _ ......... . Bermuda .... _.......... . 32° N ........... _ 1. 09 12. 64 

Florida ... _ ............ 23° 30' N....... 1. 96 13.04 

Bahamas ................ 25° N............ 2. 80 13.19 

Florida ....... _ ........ 23° 30' N ....... ! 2. 87 13.29 

Fiji ..... -.. : .. ......... -....... -....... ' . 47 13. 43 

Lower California ....... _ 24 ° 161 N ....... , 8. 27 13. 71 

Brazil. .. _. _ ............................ ' Trace. 13. 79 

Cape San Lucas ........ 22° .52' N ....... 7. 95 14.13 

'forres Strait............ 10° S:.......... . 88 15. 65 

Brazil........ .. . . . . . . .. 3° 50' N ........ 1 8. 57 ~5. 73 

T~~lip.idogorgia flabellum ......................... . X1pho~org1a anceps ........... __ ................. . l~hipict?gorgia flabellum ........ __ ............... . <:: orgon~a ac~rosa: ................................. . borgoma suffrutlcosa ............................. . Leptogot·gia pulclU'a ....... _ ............... ~ ...... . Plexaurella grandifiora ........................ ~ .. . Leptogorgia rigid a .............. _ .... · ............. . Ctenocella pectinata ........................... _ .. Phyllogorgia quercifolia ... _ ... _ .......... : ....... . 

------------------------------------~----------------~--------------------~-------

Since the foregoing table was cmnpiled Prof. L. R. Cary, of Princeton University, has 

kindly sent n1e a series of unpublished analyses of alcyonarian spicules from specimens collected 

in shallow reefs around the Tortugas and southern Florida. The spicules were separated from 

organic n1atter by treat1nent with a strong solution of caustic soda and subsequent washing with 

rain water until they were freed from impurities. The analyses, by Prof. A. H. Phillips, are as 

follows: 

Analyses of alcyonarian spicules from the Tortugas and southern Florida. 

--- -- ------·--

Species. CaC0 I 3• MgC03• Ca3P20 8• Sum. I 

----

Plexaura flexuosa. _ ........ 80. 87 16. 90 0. 35 98. 13 

Plexama homoma.lla. _ . _ .. 86. 29 12.01 . 04 98. 34 ~lexat~r~lla dichotoma.- .. -~ 85. 90 14. 12 . 09 I 100. 11 

Eunec1a crassa ............. 87. 27 11. 46 . 09 I 98. 82 

Eunecia rousseaui. .... -'·. 1 86.76 11.45 .12 I 98. 31 

Plexaurella sp? .......... 86. 75 12.08 .15 I 98.98 

Briarium asbestinum ..... 85. 13 13.21 .13 I 

I 98. 47 

Gorgonia flabellum ... _ ... 86.04 13. 12 . 44 I 99. 60 

Gorgonia acerosa. _ ... _ ... 85.76. 13. 39 .44 99. 59 

Gorgonia citrina. _ ... _ ..... 85. 29 13.43 . 22 98.94 

Pseudoplexaura crassa ..... 86. 71 12. 12 .19 I 99.02 

Xiphigorgia anceps ....... 84. 91 13.23 . 26 i 98.40 

------------ -----'----

These annlyses ad1nirably confirn1 the analyses, made in the Survey laboratory, of alcyonarians frmn the same region and the Baha1nas. In addition, Prof. P~illips made a partial 

u.nrlysis of spicules from Alcyonium rigidum fron1 Samoa, in which he found 10:50 per cent of 

magnesitnn carbonate. _ 

Although records of ten1perature and depths are available for only a few of these alcyonarians, the suggested relation is clearly evident. T'he organisms from cold, northern waters or 

fron1 very deep \Vaters are low in 1nagnesia, and those frmn warm regions are high. T'he smne 

relation appears in our analyses of echinoderms and is unmistakable. It is not rigorously exact, 

buti some n.pparent irregularities are due to impurities, such as sand and mud, which appear in 

the analyses as silica and sesquioxides. If these were rejected the percentage of magnesia 
14 THE INORGANIC CONSTITUENT'S OF MARINE INVERTEBRATES. 

would be raised. Variations are also to be expected because of cold or warm currents and 

different depths of water. Very deep water, even under the Equator, is always cold, whereas 

shallow bays farther north may be relatively warm. Possibly, also, the alcyonarians may form 

two or more distinct series that are not perfectly comparable in chemical composition. Oorallium 

and Tubipora, for example, are compact forms, with littie organic matter and lower magnesia 

than the genera with horny, organic axes, such as those whose names appear at the ena of the 

table. It Is also noteworthy that the highest proportions of calcium phosphate are commonly 

associated with a high content of magnesia. 

HYDROIDS. 

In the course of this investigation six analyses have been made of coralline hydroids belonging to the genera Millepora and Distichopora. The species and localities are as follows: 

1. Millepora alcicornis Linne. Shoal water, Tortugas, Fla. 

2. Millepora alcicornis. Bermuda. · 

3. Millepora braziliensis Verrill. Candeas, Pernambuco, Brazil. 

4. Distichopora fl,itida Verrill. Micronesia, exact locality unknown. 

5. Distichopora coccinea Gray. South Sea Islands, exact locality unknown. 

6. Distichopora sulcata Pourtales. Off Habana, Cuba; depth of water, 143 to 179 metero. 

Analyses of hydroids. 

1 2 3 4 5 6 

-· . ---~· ---------------------·- ·-· ---1----1----1----1---~~ 

Si02 •••••••• · •••••••••••••••••••••••• ; •••••••••••••••• 

~8~~)·~-~3::: ::::::::::::::::::::::::::::::::::::::::: 

S03 ••••••••••••••••••••••••••••••••••••••••••••••••••• 

fg~i\i~~:: ::::.::::::::.: ::::::.·.::: ::::.::::::::::::::::: 

0 C02 needed ................... · ...................... . Organic matter, etc .................................. . 

0.23 

.10 

.43 

52.50 . 27 

Trace. 

44. 77 

98.30 

41.57 

3.20 

0.02 

. 07 

.10 

53.14 

.04 

Trace. 

45.16 

98.53 

41.85 

3.31 

Reduced analyses of hydroi(is. 

---------------·----·--

Si02 ••••••••••••••••••••••••••••••••••••••••••••••••• 

~~1::.?;::: ~ ~ ~: ~ ~ ~ ~ ~ ~: ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~: ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~: ~ Ca3P20 8 •••••••• : ••••••••••••••••••••••••••••••••••••• 

11 i 

O:ii II' 

. 95 

98.22 

. 481 Trace. 

100. oo 1 

2 

,. 

0. 02 : . 07 : . 22 1 

99.63 I 

. 06 : 

Trace. i 

100. oo 1 

0.09 

. 06 

. 59 

53.36 

1.03 

Trace. 

44.87 

100.00 

42.01 

2.86 

3 

0. 09' 

.06' 

1. 28 . 96.77 : 1. 80 ! 

Trace. I 

I 

I 100.00 i 

I 

0.10 0.09 0.07 . 20 . 07 .05 

.11 .12 .12 

53.07 53.43 53.90 . 69 . 65 . 61 

Trace. Trace. Trace. 

44.89 45.05 45.37 

1}9. 06 99.41 100.12 

41.44 41.76 42.15 3.~5 3.29 3.22 

4 5 6 

--1----i----

0.11 

. 21 

. 24 

98.22 

1. 22 

Trace. 

100.00 

1 

0.09 

.07 

. 26 

98.43 ! 

1.15 : Trace. i 

I 100.00 ; 

0.07 

.05 

. 26 

98.56 

1.06 

·Trace. 

100.00 

A few partial analyses and one fairly complete analysi~ of millepores have already been 

published, which in all important particulars agree with ours. The coralline structures consist 

essentially of calcium carbonate, with minor impurities, and resemble chemically the true 

corals. In two millepores from Bermuda A. G~ Hogbom 14 found respectively 95.86 and 94.39 

per cent of calciun1 carbonate, with 0.41 and 0.97 of ·magnesium carbonate. I;n Millepora 

alcicornis from the Gulf StreamS. P. Sharples 15 found 97.45 per cent of ·calcium carbonate, 0.27 

of ~alcium phosphate, 2.54 of organic matter and water, and only traces of iron and magnesia. 

B. Silliman, jr./6 in one analysis of the same species reported no magnesia, which evidently 

u Hogbom, A. G., Neues Jahrb., 1894, Band 1, p. 262. lG Sharples, S. P., Am. Jour. Sci., 3d ser., vol. 1, p. 168, 187L 16 Silliman, B., jr., Am. Jour. Sci., 2d scr., vol. 1, p. 189, 1846. 
(J 

ANNELIDS. 15 

was not soug~t for. A single analysis of M. bra~iliensis by L. R. Lenox 17 gave the following 

cmnposition: 

CaC03 • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 93. 80 

MgC03 •••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••• · •••••••••••••• 2.14 

CaS04 ••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••• 2. 08 

Si02 ••••••••••••••••••••••••••••••••••••• ·••••••••••••••••••••••••••••••••••••••••••••••••••• • 03 

(Al,Fe)20 3 •••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••• ~.............. • 07 

98.12 

It is interesting to note that the Brazilian species is the richest of all in magnesia, although 

it is poor in c01nparison with the echinoderms and alcyonarians. As ~·eef builders the millepores contribute little to the doral rock but carbonate of lime. 

In an analysis of ]!.illepota cervicornis, by A. Damour/8 8.51 per cent of magnesium carbonate 

is reported. The na1ne, however, belongs to a fossil species, and the specimen analyzed was at 

least partly fossilized. Its locality is given as Brehat, Manche, France. · 

ANNELIDS. 

The curious tubes formed by 1narine worms probably contribute little to the sediments. 

They are, however, of some interest in an investigation of this kind, and for that reason six 

'analyses of them have been made, as follows: · 

1. F·ilograna irnplexa Berkeley (Serpula cornplexa). Scarborough, England. 

2. Protula tabularia (Montagu) (Serpula tabularia). Locality unknown, probably British. 

3. Hydroides d-ianthus Verrill. Vineyard Sound, Mass. 

4. Leod·ice polybranch'ia Verrill. Off Marthas Vineyard, Mass. 

5, G. Hyalinrecia q.rtijex Verrill. Off Marthas Vineyard, Fish Hawk station 1025; depth of water, 384 meters; 

bottom temperatme, 7° C. Two analyses, of different samples; the tubes of this annelid, on ignition, gave an inorganic 

residue which fused to a white, porcelain-like mass. 

Analyses of worrn tubes. 

Insoluble in HCI. .................................... . 

~l~~)~~~::::::::: :::: :: ::::::::::::::::::::::::::::: r;~~~~::::::::::::::::::::::::::::: ::::::::::::::::: 

C02 needed ......................................... . Organic matter, etc ................................ · ... . 

1 

29.82 

1.12 

. 75 

. 00 

35.98 

Trace . . 29 

31.23 

99.19 

28.00 

3.23 

2 3 

. Tests for fluorine ~ailed to show its prese~ce in any of these tubes. 

4 5 6 

These analyses fall into two distinct groups, one low in sulphates and phosphates, the 

other unusually high. The first three are easily reducible to standard form if we reject the 

insoluble matter, the silica, and the sesquioxides, which represent inclusions of sand or mud. 

The reduced analyses then assume the following form: 

Reduced analyses of worrn tubes. 

1 

0.00 

99.01 

Trace. . 99 

100.00 

2 

0.32 

99.55 

.13 

Trace. 

100.00 

3 

9.72 

89.66 

Trace . . 62 

100.00 

1

7 Lenox, L. R., Harvard Coli. Mus. Comp. Zool. Bull., vol. 44, p. 264, 1904. 

106135-22-2 

1s Damour, A., Compt. Rend., vol. 32, p. 253, 1851. 
16 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. 

Two of these analyses resemble those of corals and mollusks; the third is more like one of 

an alcyonarian or an echinoderm. In all three the organic· matter is very low. Three other 

partial analyses by Forchhammer 19 are on record, which may be compared with these; although 

only magnesium carbonate was determined. These arePer cent MgC03. 

Serpula sp. Mediterranean .............................................................. 7. 644 

Serpula triquetra. North Sea .............................................................. 4. 455 

Serpula filograna 20 ••••••••••••••••••••••••••••••••••••••••••• _ •••••••••••••• _.. • • • • • • • • • • • • • 1. 349 

The tubes of Serpula are evidently quite variable in composition, at_least as regards their 

·magnesian content. 

The tubes formed by Leodice and Hyalinmcia, being highly phosphatic, are difficult to 

interpret chemically. The bases are insufficient to satisfy the acids if the phosphoric oxide is 

assumed to re-present the normal tribasic salts. Phosphorus may be present ~s part of the 

organic matter, and so, too, may a portion of the sulphur. .Metaphosphates, pyrophosphates, 

and acid orthophosphates are also possibly present, and between these alternatives it is not easy 

to decide. Acid salts are, however, improbable, for when boiled in water the tubes give faintly 

alkaline reactions. We prefer, therefore, to leave th~ reduction of analyses 4, 5, and 6 in abeyance until more evidence can be· obtained. In this· direction an analysis of Onuphis tubicola 

Muller, by Schmiedeberg,21 is suggestive. To the organic matter of the tube of Onuphis, "onuphin," he assigns the formula 024H43N018 , and the tube itself he regards as a complex compound 

of the composition · 

C24H43N018 + CaHP04 + 4MgHP04 + 22H20. 

The analyses given by Schmiedeberg are as follows: 

Analyses of Onuphis tubicola. 

Entire tube. Calcined ash. 

Onuphin ...................................... . Water ........................................ . 

·P20s. ··-----··------·-·-·-·-·-·----··-·--·----· 

~!8. --·~ ~ ~ _- _·--_ ·_ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ 

38.48 

24.07 

2i.58 

9.73 

3.40 

2.74 

6~~ -_ ~ ~ : _- _- _ -_ :_ ~ ~ ~: ~ ~ : : : : : : : : : : ~ : : : : : : : : : : : : : : : : : : r~: ;g ~~~ •• : : : : : ~ : : ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ : : : : : ~ ~ : : : : : : : : : ~e ~ : : : 2~: g~ Na ................................ : .......... ·... .50 

H (acid), etc ............................. ~ .... - S03, Si02, 002, and loss......... . . . . . . . . . . . . . . . . . 2. 27 

100.00 99.43 

In a general way this analysis of Onuphis resembles our analysis of Hyalinmcia, except 

" that the magnesia is much higher and the sulphates,·if present, are insignificant in amount. It 

will be noticed, moreover, that Schmiedeberg assumes the presence of acid phosphates in his 

interpretation of the data, which, as we have already stated, is improbable. 

None of the analyses of the phosphatic worm tubes can be regarded as wholly satisfactory, 

but it is clear that these structures add phosphorus to the sediments. If they decay in contact 

with calcareous sediments, the final product would probably be a tribasic phosphate, and that 

is as far as we now need to go. · 

1u Forchhammer, G., Neues Jahrb.-, 1852, p. 854. • 

20 In a later memoir (Philos. Trans., vol. 155, p. 203, 1865) Forchhammer gives the percentage of MgCOa in Serpula filograna as 13.49 per cent. 

Which figure is correct? · ' 

21 Schmiedeberg, 0., Zool. Sta. Naples Mitt., vol. 3, p. 373, 1882. 
ECHINODERMS. 

1. CRINOIDS. 

RECENT CRINOIDS. 

i7 

In 1906 I-I. W. Nichols 22 published a ·number of partial analyses of marine invertebrates, 

and in a crinoid, Metacrinus rotunclus frmn Japan, he found 11.72 per cent of n1agnesiun1 

carbo~nate. This analysis attracted the attention of Austin H. Clark, and at his request two 

other analyses of crinoids were n1ade in the laboratory of the United States Geological Survey 

by Chase Palmer,· who also found that they contained abundant n1agnesia. These analyses, 

which were published and discussed by. Mr. Clark/3 will be considered in detail later. They at 

once suggested that crinoids generally might be highly magnesian and so· play an important 

part in the formation of magnesian limestones. 

In order to settle this question Mr.· Clark supplied us with 24 specin1ens of recent crinoids, 

representing 21 genera and covering a wide range of localities. These were analyzed by Mr. 

Wheeler, and the analyses confirmed the original supposition. All the specimens contained 

n1agnesimn cm·bona~e in notable proportions but varying in a n1ost re1narkable manner. The 

data obtained are in detail as follows, beginning with the list of the specimens studied: 

1. Pt·ilocrinus pinnatus A. H. Clark. Albatross statp.on 3342, off the Queen Charlotte Islands, British Columbia; 

latitude, 52° 39' 3011 N.; longitude, 132° 38' W.; depth of water, 2,858 meters; bottom temperature, 1.83° C. Mean of 

two analyses. 

2 . .fi'lorom,etm asperrinw Clark Albatross station 3070, off the coast of Washington; latitude, 47° 29' 3011 N.; 

longHude,-125° 43' W.; depth, 1,145 meters; bottom temperature, 3.28° C. 

3. Psathyrometrajrag·ilis Clark. Albat1·oss station 5032, Y."ezo Strait, Japan; latitude, 44° 05' N.; long~tude, 145° 

30' K ; depth, 540-959 meters; bottom temperature, 1. 61° C. . 

4. Pentam,etrocrinus japon·icus P. H. Carpenter. Albatross station 5083, 34.5 miles off Omai Saki Light, Japan; 

latitude, 34° 04' 20" N.; longitude, 137° 57' 3011 E.; depth, 1,123 meters; bottom temperature, 3.39° C. 

5. Cap·illaster mult·imdiata Linne. Albatross station 5137, Philippine Islands nearJolo, 1.3 miles from Jolo Light; 

latitude, 6° 04' 2511 N.; longitude, 120° 58' 3011 E.; depth, 36 meters; no temperature record. 

6. Pachylometra patula Carpenter. Albatross station 5036, Philippine Islands, North Balabac Strait, 15.5 miles 

from Balabac Dight; latitude, 8° 061 4011 N.; longitude, 117° 18' 4511 E.; depth, 104 meters; no temperature record. 

7. Catoptometra ophiura Clark. Same locality as No. 6. 

8. Hypalocrinus naresianus Carpenter. Albatross station 5424, Philippine Islands, '3.4 miles off Cagayan Island, 

Jolo Sea; latitude, go 37' 05" N.; longitude, 121° 12' 3711 E.; depth: 612 meters; bottom temperature, 10.22° C. 

9. Parametra gramtlata Clark. Albatross statiQn 5536, Philippine Islands, between Negros and Siquifor, 11.8 

miles from Apo Island; latitude, go 15' 45" N.; longitude, 123° 22' E.; depth, 502 meters; bottom temperature, 11.95° C. 

10. Craspedmnetra an~eps Carpenter. Albatross station 5157, 3.3 miles from Tinakta Island, Tawi Tawi group, 

Sulu Archjpelago; latitude, 5° 12' 3011 N.; longitude, ll9° 55' 5011 E.; depth, 32 meters; no temperature record. 

11. Pt·ilom,etra m:illleri Qlark. Sydney Harbor, New South Wales, Australia; latitude, 33° 15' S.; longitude, 151° 

12' K, approximately. 

. 12. Ha~?~rometra dentata.Say. Fish Hawk station 1033, off Marthas Vineyard, Mass.; l~titudfl, 39° 56' N.; longitude, 69° 241 W.; depth, 329 meters; bottom tempemture, about 7.8° C. 

''13. Bythocrinus robustus Clark. Albatross station 2401, Gulf of Mexico, southeast of Pensacola; latitude, 28 ° 38' 30'' 

N.; longitude, 85° 52' 30". W.; depth, 255 meters; no temperatme record. 

14. Crinometra concinna Clark. Albatross station 2324, north of Cuba; latitude, 23° 10' 3511 N.; l,mgitudP., 82° 20' 

· 2411 W.; depth, 59 meters; bottom temperature, 26.17° C. 

15. Isocrinus decorus Wyville 'l'homson, stem. Off Habana, Cuba; latitude, 24° N.; longitude, 82° W.; approxi· 

mately. 

16. Same as No. 15, arms. 

17. Endoxocrinus parrre Gervais, stem. Off H~bana, Cuba. 

18. Same as No. 17, arms. 

19. Tropiometra picta Gay. Rio de Janeiro, Brazil; latitude, 25° 54'S.; longitude, 44° W., approximately. 

20. Promachocrinus kerguelensis Carpenter. Shores of the An.tarctic Continent in the vicinity of Gaussberg; 

latitude, 67° S.; longitude, 90° E., approximately; depth, 350-400 meters; bottom temperature, -1.85° C.; salinity of 

water, 3.3 per cent. 

21. Antho.rnetra adriani Bell. Same locality as No .. 20. Nos. 20 and 21 were collected by "the German South Polar 

l~xpedition. 

22. Zygometra microdiscus Bell. Aru Islands,,near the western point of New Guinea; latitude, 5°-6°S. 

23. Chlorometra rugosa Clark. Near Rotti, Lesser Sunda Islands; latitude, 10° 39'S.; longitude, 123° 40' E.; 

depth, 520 meters. 

22 Nichols, H. W., Field Columbian :M:us.'Pub. 111, p. 31, 1906. 2a Clark, A. H., U.S. Nat. Mus. Proc., vol. 39, p. 487, 1911. 
18 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. 

24. Tropiometra carinata, Lamarck. From Pigeon Point, Tobago, British West Indies. Shoal water, near shore: 

temperature about 28° C. Received from Dr. Hubert L. Clark. Accord.ing to A. H. Clark it is probably T. picta 

Analysis by R. M. Kamm. 

The actual analyses are given in the table below. SUlphates were determined in only one 

of them, because the material was insufficient. 

Analyses of crinoid skeletons. 

1 2 3 4 5 6 

~ .. t?,:F~ )~oa·:. ·.·: .· ~: ~ :: : : : : :: : : : : : : ~ : : : : : : : : : : : : : : : : : : : : : 1. 64 0.04 1.11 0.37 0.16 0.12 

1.07 . 34 1. 01 .71 . 62 . 63 

~:3:::::::::::::.-::: ~ ......... ::: ......... ::::::::::::: :·:::::::: 3.08 

I 

3.60 3.12 3.76 4.7"7 4.94 

40.65 40.37 34.20 38.50 38.12 41.34 

P20s··················· ............................. .. : .11 . 21 Trace? .40 Trace. . 43 

Ignition ............... ~ ..........................••.. 51.45 53.75 60.04 55.~5 54.61 51.36 

98.00 98.36 99.48 98.99 98.28 98.82 

C02 needed ............................................ 35.23 35.48 31.37 34.01 35.19 37.51 

Organic matter, etc.' .................................... 16.22 18.27 28.67 21.24 19.42 13.85 

7 8 9 10 11 12 

~N~ ~~~:::~:::.::~. ~- ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ : ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ : 

0.04 0.07 0.40 0.15 0.17 3.17 . 79 . 09 . 50 .19 .19 . 31 

4.64 4.44 4.48 5.13 4.17 2.49 

40. 75 45.86 41.79 42. 77 38.91 26.12 

f~ti~;;.-::::::::::::. ·. ~::::::::::::::::::::::::::::: ~:: . 33 Trace; Trace . .11 .17 . 23 51.80 48.32 51.44 50:28 54.61 65.~5. 

C02 needed ............ '................................ 98.'35 98.78 98.61 98.63 98.22 97.57 

36.81 41.27 37. 77 39.17 34.90 .22.73 

Organic matter, etc .............................. ~ ...... 14.99 7.05 13.67 11.11 19.71. 42.52 

-13 14 15 16 17 18 

~i?, F~ );o~::: ~ ~ ~: ~: ~ -~ ~ ~::::::::::: ~::::::::::::::: :::: 0.40 0.04 0.03 0.09 0.04 0.15 . 31 .25 . 07 .19 . 20 . 26 

~:8:::::::::::.·.- .. :::::::::::::::::::::: ·:.:: :::::::: ::: : 4.56 4.75 5.08 4.70 5.09 5.04 

47.08 41.78 45.67 42.77 45.42 43.41 

f~ti~~::: ~ ._._.:.:::: ·. ~---- .. :;:::::::::::::::: :·:::::::::::: Trace. Trace. Trace. Trace? Trace. Trace. 47.17 50.33 47.54 50.59 48.58 50.00 

99.52 97.15 98.39 98.34 99.33 98.86 

C0 2 n~eded ............................................ 41.93 38.00 40.40 38.71 41.29 39.65 

Organic matter, etc ..................................... 5.24 12.33 7.14 11.88 7.29 10.35 

1.9 20 21 22 23 24 -

Si02 ••••••••••••••••••••••• · •••••••••••••••••••••••••••• 0.02 0.02 0.23 0.04 0.05 0.40 

(Al,Fe)20 3 •••••••••••••••••••••••••••••• · •••••••••••••• . 35 .45 . 37 . 48 . 23 . 38 

~:8.-.-.-.-.-.-.-.-.-.-.-.- .~ .. ::::::::::::::::::::::::::::::::::::: 4.51 3.02 3.27 .4. 92 3.99 4.87 

39.57 40.68 42.49 37.19 42.72 35.35 

J~~~~:::: :·:·:·:·::::::::::::::::::::::::::::::::: :.:::: . 10 Trace. Trace . .17 Trace. . 22 

(?) (?) (?) (?) (?) . 63 

53.64 54.53 52.22 55.05 51. 69. 56.14 

98.19 98.70 98.58 97.85 98.68 97.99 

C02 needed ................. : ....... ~ .................. 36.05 35.18 37.08 34.47 37.95 32.58 

Organic matter, etc ..................................... 17.59 19.35 15.14 20.58 13.74 23.56 

The summation in most of these analyses is low. The deficiency is due mainly to inclosed 

or adherent sea salts, an inevitable impurity, as was proved in the analyses of two separate 

samples. In No. 15, 1.27 per cent of water-soluble salts was found, and in No .. _17; 0.21 'per 

cent. These additions raise the summations to 99.66 and 99.54 per cent, re~pectively. 
ECHINODERMS. 19 

The reduced analyses are as follows, reJecting organic matter and water and recalculating 

to 100 per cent: 

With these analyses· the two made by Mr. Palmer may be advantageously compared, 

although they were no.t quite so elaborate. The data are as follows: 

25. Heliometra glacialis var. maxima. Iwanai Bay, northeastern part of the Sea of Japan; latitude, 43° 01' 4011 N.; 

depth, 315 meters; temperature, surface, 20.5° C., bottom, 1.5° C. 

26. Metacrinus rotundus. Eastern Sea, off Kagoshima Gulf, southern Japan; latitude, 30° 58' 3011 N.; depth, 278 

meters; temperature, surface, 27.8° C., bottom, 13.3° C. 

In No. 24, which co~1tained much organic inatter, M!· Palmer found 2.68 per cent MgO 

(=5.61 MgCO~) and 40.03 CaO (=71.48 CaC03). In No. 25, with no organic matter, he found 

4.89 MgO (=10.29 MgC03) and 49.95 CaO (=='89.19. CaC03). Assuming that the crinoid skeletons consist essentially of carbonates, ·and recalculating to 100 per cent, we have as the content 

of 1nagnesium carbonate in these crinoidsPer cent. 

25. Heliometra glacili.lis var. maxima ............................... ~........................... 7. 28 

26. J.fetacrinus rotundus ..... ................... · .............. : ................................ 10. 34 
20 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. 

These figures fit in well with the others and eve:p. by themselves suggest a relation between 

temperature and the magnesia content of crinoids. In the following table the entire series is 

arranged in the order of ascending magnesium carbonate, with the accessory data as to latitude 

and locality abbreviated. In this table the two analyses of Endoxocrinus are averaged together, 

and so also· are the two of Isocrinus. 

Percentage of magnesium carbonate in crinoids. 

Genus. Locality'. Latitude. Depth. Temper- MgC03 . ature . 

. Meiers. oc. Pe1· cent. 

Heliometra ........................ Northetn Japan ........ 43° N .............. . Promachocrinus .................... Antarctic .............. 16. 7°S .............. . Ptilocrinus ........................ ·British Columbia ....... 52° 39' N .......... . Anthometra ........................ Antarctic .............. 67° S .............. . 

315 1.5 7.28 

375 -l..S 7:8() 

2,858 1.8 7.91 

375 -1.8 8.23 

Psa th vrometra. . . . . . . . . . . . . . . . . . . . Northern Japan . . . . . . . . 44 ° N .............. . Hathrometra .................... ,.. Massachusetts........... 39° 56' N .......... . (?) 1.6 9.25 

329 7.8 9.36 

Florometra ........................ Washington ........ · ..... 47° 29' N .......... . Chlorometra ........................ Rotti Island ............ 10° 39'S .......... . 1, 145, 3.3 9.44 

520 (?) 9.87 

Bythocrinus ....................... Gulf of Mexico ......... 28° 38' N .......... . Pentametrocrinus .................. Southern Ja-pan ........ 34° N .............. . Hypal~crinus. . . . . . . . .. . . . . . . . .. . . Philip-pine Islands ..... · 9° 37' N ........... . Metacrmus ........ : .......... , .... Southern Japan ........ 30° 58' N .......... . 

255 (?) 10.09 

1,123 . 3. 4 10.15 

612 10.2 10.16 

278 13.3 10.34 

Parametra ........................ Philippine Islands ..... 9° 15' N ........... . Ptilometra......................... Australia............... 33° 15' S .......... . 502 12 11. OR 

(?) (?) 11.1:{ 

Isocrinus .......................... Cuba .................. 24° N .............. . Catoptometra ...................... Philippine Islands..... 8° N ............... . Crinometra ......................... Cuba ..... : ............. 23° 10' N ........... · 

(?) (?) 11. 5fi 

104 (?) 11.68 

59 26.2 11.69 

Tropiometra ....................... Brazil. ................. 25° 54'S .......... . Endoxocrinus ...................... Cuba ...... -............. 24° N .............•. · Pachylometra ...................... Philippine Islands ..... 8° N ............... . Craspedometra .... · ................. Philippine Islands..... 5° 12' N ........ • .. .. Capillaster ......................... Philippine Islands ..... 6° N ............... . Zygometra ................... : . .... Aru Islands............. 5°-6° S ............ . Tropiometra ....... ,............... Tobago................. 11° 25' N .... ~ ..... . 

(?) (?) 11.77 

(?) (?) 11.79 

1,044 (?) 12.20 

32 (?) 12.34 

36 (?) 12.119 

(?) (?) 13.37 

Shoal. 28 13. 74 

The percentage of 1nagnesiu1n carbonate in Ohlorometra is low for the latitude of the 

locality, but that is doubtless due to the depth of the water (520 meters) in which the crinoid 

lived. The. probable temperature at that depth was between 7° and 10° C. 

From the foregoing table it is perfectly clear that the proportion of magnesium carbonate 

in crinoids is in some way dependent on temperature. Temperature, however, is not entirely 

dependenton latitude. Depth of water has also a distinct influence. The crinoids from relatively shallow depths in the Tropics are highest in their magnesium content; those from the 

Antarctic and the far north are lowest. The proportion given for No. 12, from the coast of 

Massachusetts, is probably too low, for th_e specimen as analyzed contained over 6 per cent of 

silica and sesquioxides-evident impurities, due to adherent mud from which the ·delicate 

structure could not be wholly freed. If these are rejected, the 'magnesium carbonate is raised 

from 9.36 to 10 per cent, which gives the crinoid a better and more p1'obable rating. 

So far as we are aware such a peculiar relation between te1nperature and composition as 

is here reco~ded has not been previously observed. To recognize it is one thing; to account for 

it is not so easy.. At first we supposed that it might po·ssibly be due to a difference in the form 

of the more abundant carbonate-the less stable aragonite in the warm-water forms and calcite 

in ·the crinoids from colder regions. But tests by Meigen's reaction proveq that the organisms 

were all calcitic, and so this supposition had to be abandoned. 

Mr. A. H. Clark, who is an ~uthority on the crinoids, infonns us that those from warn1 

regions have the most cmnpact skeletons, the compactness being in general proportional· to 

the te1nperature and to smne extent de,pendent upon the size of the individual. Heliometra, 

for exa1nple, is the largest of the crinoids, its skeleton is one of the least compact, and its nlagnesian content i$ lowest among ~:tll the specie$ exan1ined. Structure as well as temperature 
. ECHINODERMS. 21 

seems to 'be correlated with the proportion of magnesia in the crinoids, but the dhe1nical explanation of the facts is yet to be found. It may hav·e connection with the gaseous content 

of sea water, carbon dioxide, for example, being more soluble il). warm than in cold waters, 

but this is only a suggestion, which may or may not be fruitful. The same regularity as to 

temperattrre also appears in our analyses of alcyonarians. 

FOSSIL CRINOIDS. 

In order to make this investigation n1ore syste.matic it seemed desirH.ble to analyze a 

nmnber of fossil crinoids, so as to determine whether any definite and regular changes could be 

traced in passing from the recent to the ancient organisms. For the material studied we 

a.re in~ebted to the kindness of Mr. Frank Springer, .who selected the material with great care 

. so n.s to cover a range of horizons from the Lower Ordovician up to the Eocene. The 10 crinoids 

chosen are described in the list below, and t.he analyses which follow were nlade .in the same 

wny as those of the modern species: 

l. Pentacr1:n1ts decadactyhts D'Orbigny, stem. Eocene, Vincen:r.a., Italy. 

:2. jlfille1"icrinns mcspilij'ormis Gold fuss, stem. Upper Jma.ssic, Kelheirn, Bavaria. 

8 . .Pentacrinns basaltij'onnis :Miller, stetp. Middle Lias (Lower Jurassic) Breitenbach, Wurttemberg, Germany. 

4 . .Enc1··inus lili1j'o-nnis Lamarck, stem. Triassic, Braunschweig, Germany. 

5. GraJ)h·iocrimts nwgn1jicusl\filler and Gurley, complete crown. Pennsylvn.nian (upper Carboniferous), Kansas 

(:ity, Mo. 

tL Dorycrinus ·ttn·icorn·is Owen and Shumard, calyx and stem. Lower p<trt of Burlington limestone, Mississippian 

(lower Carboniferous), Burlington, Io,va. 

7. Jlfegi.~tocrinus nodosLts Barris, plates. Middle Devonian, Alpena, Mich. 

8. JJ)ncalyptocrim.ts crassus, plates. Silurian, western Tennessee. 

9. l\rinoid sp. '?,stem. Trenton limestone, Middle Ordovician, Kirkfield, Ca,nacht. 

10. D'i.0.1)Ql(){:rinu.s vespemlis White, plates and stem. Lower Ordovician, Tennessee. 

Analysc8 r~f fossil c1·inoids. 

1 2. I 3 L! 5 6 I 7 8 9 10 

---------· ------------

SiO ......................... 0.99 2.84 l. 55 0.24 3.07 6. 92 10.39 29. 11 2. 56 4.07 

(Al,l<'e)20 3 •••••••••••••••• 2.64 . 28 2.59 . 43. 2. 18 . 64 . 87 1.73 . 31 l. 95 

]<'cO ...................... 1.36 . 00 ................ . 00 l. 32 . 00 1. 19 . 27 . 00 . 88 

MnO ..................... . 1:3 . 00 .............. Trace. .15 Trace . .16 . 04 Trace. . 04 

~~;~_-_·.-.-.-.-.·.-.· .• ~::::::::::: ~ . 78 . 38 . 84 9.44 . 78 . 38 1. 21 . 58 . 91 . 79 

51. 22 53. 68 51.78 43.40 50.10 51. 20 46.57 37.43 53. 87 50.42 

P20 5 ••••••••••••••••••••••• . 00 Trace. . 09 Trace. Trace. Trace . Trace . . 00 Trace . Trace. 

Loss on ignition ........... 42. 80 42.93 43.21 45.95 41. 7l 41.00 39.20 30. 7l 42.45 41.53 

99.92 100. 11 100.06 99.46 99.32 100. 14 99.59 99.87 100. 10 99.68 

Reduced analyses of fossil c1'inoids. 

1 2 3 4 5 6 7 8 9 10 

----------------·------

SiO~-·-······· ............. l. 00 2. 85 1. 57 0.24 3. 11 6.94 10.48 29.30 2. 55 4.10 

.~;\·hl!'e)20a ................ 2.6fi . 28 2. 64 . 44 2. 22 . 64 . 88 l. 74 . :30 1. 97 .. ~e ·08 •••••••••••••••••• :. 2. 21 . 00 . 00 2. 16 . 00 l. 94 . 43 . 00 1. 52 

l\:ln003 ••••••••••••••••••• . 21 . 00 Trace. . 24 Trace. . 26 . 06 Trace . . 06 

Mg008 ••••••••••••••••••••• l. 66 . 80 1. 79 20.23 1. ()(i . so 2. 5() 1. 23 1.90 1. (:i7 

CaC03 ••••••••••••••••••••• 92.26 96.07 93. 80 79.09 90.61 91. 62 8:3.88 67.24 95. 25 90.78 

Ca3P:.!08 .•.••••••••••••••• . 00 'l'race . . 20 Trace. Trace. Trace. Trace. . 00 Trace. Trace . 

100.00 100.00 100.00 100.00 100.00 I 100.00 100.00 1.00.00 100.00 100.00 
22 . THE INORGANIC CONSTITUENT'S OF MARINE INVERTEBRATES. 

In some respects these analyses are unsatisfactory, for they show no regularities of .any 

kind. In only one of them, No. 4, is there exhibited a concentration of magnesium carbonate; 

in the others the percentage of this constituent is very low. The reason for this decrease of magnesia is by no means clear. It is conceivable that the ancient crinoids may have been deficient 

in magnesia, but it is more probable that the loss is due to alteration, perhaps to the infiltration 

of calcium carbonate. Such a change would obviously lower the apparent proportion of 

magnesium carbonate. Several of the crinoids contain noteworthy quantities of ferrous 

carbonate and manganese-constituents which did not appear in the analyses of the modern 

. species. In No. 8 there is a very strong silicification, 29.11 per cent; but the matrix of the 

specimen contained only 7.55 per cent of silica. Here the infiltration of the impurity seems 

to be very clea1·. Some of the deficiencies in magnesia may have been caused by solution and 

leaching, but calcium carbonate should then have been removed to a greater extent. In ·short, 

the fossil crinoids differ widely in composition from the still living species, and in a very irregular 

manner,· and it is worth noting that in three analyses of fossil algre reported by Hogbom 24 

a similar decrease of magnesia appears. It would be easy to speculate on the significance of 

these differences; but the conclusions so reached would not be entitled to much weight. That 

the recent crinoids are distinctly magnesian and that the proportion of magnesia is dependent 

in some way on temperature are two positive results of this investigation. 

2. SEA URCHINS. 

Twelve sea urchins selected for us by Mr. Austin H. Clark ·and two received from Dr. 

Hubert L: Clark were analyzed. The species' chosen were as follows: 

1. Strongylocentrotus d1·obachiensis 0. F. Muller. Upernivik, Greenland; latitude 72° 481 N. 

2. Strongylocentmtus jragilis Jackson. Albatross station 2946, off southern California; latitude, 33° 58' 0011 N.; 

longitude, 119° 30' 4511 W.; depth of water, 274.5 meters; bottom temperature, 13.6° G. 

3. Echinamchni1tS -parma Lamarck. Coast of New England. 

4. Encope californica Verrill. Galapagos Islands, on or near the Equator. 

5. Lytechinus anamesus H. L. Clark." Albatross station 2938, off Wilmington, Calif.; latitude, 33° 351 1511 N.; 

longitude, 118° 08' 3011 W.; depth, 86 meters; bottom temperature, 15° G. 

6. Loxechimts albtts Molina. Pqrt Otway, Patagonia; latltude, about 46° or 47° S. 

7. Tetrapygus niger Molina. Coast of Peru. 

8. Tretocirlaris a,(finis Philippi. Albatross stations 2316 and 2317, off Key \Vest, Fla.; latitude, 24° 25' N.; longitude, 8.1° 471 W.; depth, 85 meters; bottom temperature, 24° C. 

9. Heterocentrotus mammillatns Linne. Low or Tuamotu Archipelago, southern Pacific Ocean; latitude, between 

14° and ?4° S. " 

10. Encope micropora A. Agassiz. Puerto Viejo, Margarita Bay, Lower Califor!lia; latitude, 24° 301 N.; longitude,. 

112° W., approxima,tely. 

. 11. Clypeaster testudinariw~ Gray. Southern .Japan. 

12. Echimts affinis Mortensen. Albatross station 2206, between Hatteras and Nantucket; latitude, 39° 35' N.; 

·longitude, 71° 34' 3011 W.; depth, 1,919 meters; bottom temperature, 3.6° C. · 

Analyses 10-12 by B. Salkover. 

13. Ech1:nometra lucunter Linne. Pigeon Point, Tobago, British West Indies; latitude, 11° 25' N.; shoal water, 

near shore; temperature, 28° 0. 

14. Mellita sexiesperjoratus Leske. Pigeon Point, Tobagct, British West Indies; latitude, 11° 251 N.; shoal water, 

near shore; temperature, 28° C. 

Analyses 13 and 14 by R. M. Kamm. 

2i Hogbom, A. G., Neues Jahrb., 1894, Band 1, p. 252. 
0 

ECHINODERMS. 23 

Analyses of sea u-rchins. 

1 2 3 4 5 ·G 

- -----·- --·-·--- _:_ _______________ , ____ , ____ , ____ , ____ ,_;_ ___ , __ _ 8.i0 .•................................................... (AI, .. li'e):.!03 •••••••••••••••••••••••••••••••••••••••••••• 

~~ft?. ·.::::::::::: · .._._._.::: ·. ·.·: _·::: ~ ·. ·:: .-::::::::::::::::::: 

S03 ••••••••••••••••••••••••••••••••••••••••••••••••••• 

~~~~;i~~~::::::::: :.::::::::::::::::::::::::.:::::::::-::: 

CO., needed ........................................... . Organic mtttter, etc ................... ~ ................ . 

SiQ,, ...... ~ ............. .' ...................... · ....... 

(Al,~P.e) 0s- ............................ -............ -. 

t~f8:.::: .-.-:. ·.:::::::::::::::::::::::::::: :.:::::::::::: 

803 ••••••••••••••••••••••••••••••••••••••••••••••••••• 

~:~~;1ti~~: : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : 

CO:~ needed ............................................ Organic matter, etc ........................... · ........ 

0. 12 

. 34 

2.58 

47.34 . 20 

Trace. 

48.53 

99. 11 

39.92 

8.61 

7 

0.31 

. 30 

2.82 

48.86 

1. 43 

Trace. 

44.98 

98. 70 

40.69 

4. 29 

0.26 

. 65 

3.68 

41.08 

1. 15 

. 39 

52.21 

99.42 

34.20 

18.01 

10 

2. 38 

. 81 

5.45 

38.24 

Trace. 

Trace. 

51. 24 

98. 12 

36.64 

14.60 

0. 14 

. 27 

2.97 

49. 17 

·. 26 

. 05 

45. 74 

98.60 

41. 71 

4.03 

11 

0. 15 

.13 

3. 90 

49. 73 

Trace. 

'J'race. 

45.47 

99.38 

43. 36 

2. 11 

3.86 8.52 0. 05. 

5.03 3.01 . 17 

4.75 3.04 3.07 

43.42 37.92 45.87 •. 28 . 37 . 35 

Trace. .19 Trace. 

43.01 45.38 49.47 

99.35 98. 43• 98.98 

38.99 32. 28 39. 34 

4.02 13. 10 10. 13 

12 13 ]4 

3.01 0. 10 0. 14 

2. 21 . 28 . 36 

2. 39 4. 16 5. 23 

46. 19 36.97 44. 84 

Trace. . 99 1. 37 

Trace. . 64 Trace. 

46.90 51.82 46. 70 

100. 70 94.96 98. 64 

38.92 32.49 39. 27 

7.98 19. 33 7.43 

Rejecting the very variable organic matter, etc., the reduced analyses assume the following 

form: · Reduced analyses of sea 1.trchins. 

1 2 3 4' 5 6 

Si02 •• --~---········································· 0. 13 0.32 0. 15 3.99 9. 93· 0.05 

~~3.~-:?:·: _:. :_:: : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : . 37 . 81 . 29 5. 20 3.51 .18 

5.99 6.95 6.59 10. 38 7.44 7.38 

93. 13 88.44 92. 39· 79.94 77.91 91.73 

CnS04 •••••••••••••••••••••••••••••••••••••••••••••••• . 38 2.42 . 46 . 49 . 73 . 66 

Ca3P:s0s- ............................•................ Trace. .1.06 .12 Tt;ace. .48 Trace. 

100.00 100.00 100.00 100.00 100.00 100.00 

7 10 11 12 13 14 

fit~:~-:_:_:~~H//--//H//H- .0. 33 2.87 0. 16 3.25 0.13 0.15 . 32 .9~ .14 2. 38 . 37 . 39 

6.27 13. 79 8.41 5.41 11.56 11.91 

90.52 82.36 91. 29 88.96 83.87 85.02 

2.56 Trace. Trace. Trace. 2.22 2.53 

Ca3P20 8 •••••••••••••••••••••••••••••••••••••••••.••••• Trace. Trace. Trace. Trace. 1.85 Trace. 

100.00 100.00 100.00 100.00 100.00 100.00 

Sea prchins Nos. 8 and 9, Tretocidaris and Heterocentrot1)ft, must be considered separately 

from the others. No. 9, a giant form, was the subject of four analyses, the shell or test, the 

dental pyramid, the small white spines on the border of the peristome, and the large purplishred spine. The large red spine analyzed was 15 centimeters long and weighed 13 grams. 
24 THE INORGANIC CONSTITUENT'S OF MARINE INVERTEBRATES. 

Analyses of Heterocentrutus mammillahts. 

Dental White 

pyramid. spines. Shell. 

Si02 ••••••••••••• :·· •••••• 0.02 0.02 0.05 

(Al,Fe)20 3 •••••••••••••••• .13 . OS .13 

~;g_ ·. ~ ~: .· .·. ·. ~ ~ ~: ~ ~ ~ ~ ~ ~ ~ ~ -~ ~ 5.21 5. 50 3.74 

"13. 60 46.02 48. 26 . S03 •••••• · ••••••••. ~ ••••••. . 61 . 58 . 29 ;p~os· .. ~ ................. Trace. Trace. Trace, 

Ignition .................. 49. 62 47.00 46.46 

99. 19 99. 20 I 98.93 

C02 needed ................ 39.()6 42. 89 41. 77 

Organic matter, etc ....... 9.96 1·11 I 4.69 

Reduced analyses of Iieterocentrotus mam:m·Watus. 

8i02··········· ........... . (Al,Fe)20 3 •••••••••••••••• 

MgC03 • .......... : .•• · .•.. 

CaC03 •••••••..•••..•.•.•••• 

Ca.S0 4 •••••••••••••••••••• 

Ca3P20 8 •••• ~ ••••••••••••• 

0.02 

.14 

]2. 26 

86.42 

.1. 16 

Trace. 

100.00 

0.02 

. 09 

12. 27 

SS.57 

l. 05 

Trace. 

0. 051 .14 

8 -~J') 

. ·J-

!)0. H7 

. 52 

Trace. 

100. oo I· 100. oo 

Red 

spines. 

0.05 

. 26 

4.47 

47.72 . 78 

Trace. 

45.H9 

99. 27 

41.49 

4. 50 

0.05 

. 28 

9.86 

88.43 

1.38 

Trace_. 

100.00 

From these analyses we see that the inorganic ·constituents of Heterocentrotus are not 

uniformly distributed. The shell and teeth are alike and are rich in magnesium carbonate; the 

coarser spines are much less magnesian. The composition of the entire skeleton, if it c~n be 

?alled. so, :would probably be somew~ere ne~r that of the red spines alone, only a little higher 

In magnesia. 

A similar example is offered by No. 8, Tretocidaris. In the specimen analyzed the shell 

and. spines were taken separately, but the spines were dead when. the urchin was collected. 

The analyses are as follows: 

A9tual analY,ses. 

Si02 ••••••••••••••••••••.••••••••••• 

(Al,Fe)20 3 ••••••••••••••••• -.-.--. 

~3. ·. : ~::-·. ·. ~ : : : ~ : ~ : ~ : : : : : ~ : ~ :. : : : ~ S03 •••••••••••• · ••••••••••••••••••• 

rg~iti~Ii: : : : : ~ : : ~ : ~ : ~ ~ : : : : ~ : : : : : :· : 

C02 needed .. : ... : ............. _ ... ··· Orga'iric matter, etc ................ . 

Analyses. of Tretocidaris a.Oinis. 

Shell.· 

0.11 

.15 

4.02 

45.80 . 57 

Tra.ce. 

48 .. 32 

98.97-

40.09 

8,23 

Spines. 

0. 53 

.14 

2.07 

49. 79 

(?) 

Trace. 

46.28 

98.81. 

41.40 

4.88 

j 

Reduced analyses. 

Si02 ••••••••••••••••••••••••••••• ·• 

(Al,Fe)20 3 ••••••••••••••••••••••• 

t~53a •• •• -_-_ •• •• • ••• • .·: : : : : ~ : : : : : : : : : : : ~ CaS0.1 ••••••••••••••••••••••••••• 

Ca3P20 8 ... : ...................... . 

Shell. 

0. 12 

.16 

9.30 

89.35 

l. 07 

Trace. 

100.00 

Here again the spines are lower in their content of magnesia than the shell. 

Spines. 

0.56 

.15 

4.63 

94.66 

(?) 

Trace. 

100.00 

In two of the analyses, Nos .. 4 ,and 5, large percentages of silica and sesquioxides appear. 

'l'hese are due to inclosed or adherent sand. and mud, which were visible in the specimens but 

not readily removable. On rejecting these impurities and recalculating to 100 per cent, the· 

percentages of.magneE?ium carbonate beca:r;ne 11.43 and 8.59, resp.ecti~ely. Similar corrections to 

the other analyses are negligfble. Mter making these corrections and assuming the percentages 

'found for the shell rather than the spines in Nos. 8 and 9, the next table has been constructed. 

If, however, the composition of the entire Heterocentrotus should be taken, it would fall below 

Encope californica. · 

0 
ECHINODERMS. 

Percentag.e of nwgnesit~m cw·bonate in sea ·urchins. 

Locality. Latitude. 

Echinus........................... Station 2206 ............. 39° 351 N .......... . AtrongyJoccntrotus drobachiensis.... Greenland.............. 72° 48' N .......... . Tctrapygus ........................ Pent. .................. (?) ................. . 

Echinnraehnius ..................... ~e~v. En~la,nd .......... 42° 451 N .......... . Strongylocentrotus fragilis.. .... . . . . . C ahtorma.............. 33° 581 N .......... . .Loxcchinus. . . . . . . . . . . . . . . . . . . . . . . Patagoni<1................ 46° 471 S .......... . 

Clypeaster. ........................ r,a.p.a.n .. : ............... ('?) ................. . Lytcchimrs......................... (.ali forma.............. 33° ~~51 M ......... . '.rretocidaris........................ Key West. ............. 24° 251 N .......... . En<"ope r:tlifornica .................. Galapagos .............. Equator ............ . 

Ech.inomctra ...................... Tobago ................. 11° 25' N-.......... . Mellita ................ ~ ............ Tobago ................. 11° 25' N .......... . 

.llcteroccntrot.us ..... , .............. Tuamotu .............. -~14° 24' S ... : ...... . Encope rnicropora ...... "· .......... California ........ ; ..... 24° 301 N .......... . 

··-- ---------------

Depth. 

Meters. 

1,919 

(?) 

(?) 

(?) 

2, 745 

(?) 

(?) 

85 

85 

(?) 

Shoal. 

Shoal. 

(?) 

(~) 

Temperature. 

oc. 

3. 6 

('!) 

(?) 

(?) 

13.G 

(?) 

(?) 

15 

24 

(?) 

· 28 

28 

(?) 

(?) 

25 

l\fgC03 . 

Per cent. 

5.41 

5.99 

6. 27 

6.59 

6.95 

7.38 

8.41 

8. 59 

!). 30 

11.43 

Jl. 56 

11.91 

12. 26 

13. 79 

These :figures, like those for the crinoids, seem to show a variation dependent upon temperature, n,t least so far as temperatures have been determined or can be inferred. The sea urchins 

fr01n cold regions are relatively low in magnesia; those from the Tropics are high. There are, 

however, two apparent exceptions. The urchin fr01n the coast of Peru probably owes its abnormality to the cold I-Iumboldt current, which flows northward from the Antarctic Ocean. Encope 

· ?nic?·opo1·a, is also seemingly anomalous, and its high pe1~centage of magnesia is difficult to explain. Unfortunately the actual temperature of its habitat is not giYen. 

· Two ·puhlished analyses of sea urchins n.re worth reproducing. They are: 

l. Eehintts (Stronyylocentrotus?) drobachiensis. North Sea.; analysis by L. Schmelck. Norske Nordhavs Exped., 

No. 2R, p. l2!J, 1901. 

2. Ech-t:nus esct.dentu.<;. Locality not stated, probably Mediterranean; analysis by 0. Biitschli. K. Gesell. Wiss. 

GiH;i,ingen .Abh., No. 3, 1908. 

Older andlyses of 'sea 't/.rcldns. 

Actual analyses. Reduced analyses. 

1 2 1 2 

·--------·----------1----l----lll--------------l-----l----

S.iO~............... . . . . . . . . . . . . . . . . Trace. 0. 04 

Al:-!03 •••••••••• :............. •• • • • • Trace .......... . 

g~c?o;::::::::::::.·: ~ ~- ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ 1~: ~~ · · · · s6: 4o · Mg003 • • • • • • • • • • • • • • • • • • • • • • • • • • • 5. 30 8. 53 

P20 5 ••••••••••••••••••••••••••••• Trace .......... . 

~~.~sct ~:~~ ~: ·_ ·_·_·: ~: ~:::::::::::::: · · ·1,.r~1.~~-. · ...... ·_ ~~. CnSO.,. 21T:.!O.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . l. 70 

:n:lo........................................ 2. 54 

Or·ga.nic uuttter·.................... 16. 33 . 03 

99.68 99.23 

Si02 •••••••••••••••••••••••••••••• 

Al20 3 ••••••••••••••••••••••••••••• 

Pe20 3 ••••••••••••••••••••••••••••• 

~;883~---.-.-.-.-.·::: ~:::::::: ~ ~::: ~ ~ ~ ~ CaS04 ••••• : • •••••••••••••••••••• 

C~aP20s·· ........................ . 

Trace. 

Trace. 

0.36 

6.36 

93.28 

Trace. 

Trace. 

100.00 

0.04 

8.84 

89.64 

1. 40 

. 08 

100.00 

Although different in minor details these analyses are fairly comparable with ours. The 

u:l-chin analyzed by Schmelek was from the far north, and its composition is very near that of 

our specin1en from Greenland. Bi.itschli's sea urchin is doubtless from warmer water and shows 

the higher figure "for 1nagnesiurri carbonate. 

3. STARFISHES. 

In the fo:nnor edition of this paper the. starfishes and brittle stars were treated as one 

series, and only eleven analyses of them were made. Now, when a much larger mass of material is available, it is better to separate the two groups of organisms. Twenty-nine starfishes haYe been analyzed, of which two from Tobago were given us by Dr. Hubert L. Clark. 
26 THE INORGANIC CO'NSTITUJ~NTS OF MARINE INVERTEBRATES. 

The other specimens were selected by Mr. Austin H. Clark from the collection of the United 

States N ational• Museum. The list is as follows: · ' . 1. Asterias vulgaris Packard. 'Eastport, Maine; latitude, 44° 551 N.; longitude, 67° 001 W. 

2. Asterias tanneri Verrill. Albatmss station 2309; latitude, 35° 431 3011 N.; longitude, 74° 52' W.; depth, 102 

meters; bottom temperature not given. 

3. Asterina miniata Brandt. Pacific Grove, Calif.; latitude, 36° 361 N.; longitude, 121° 551 W. 

4. Leptasterias compta Stimpson. Albatross station 2250; latitude, 40° 171 1511 N.; longitude, 69° 511 4511 W.; 

depth, 86 meters; bottom temperature, 10.8° C. 

5. Benthopecten spinosus VerrilL Albatross station 2568; latitude, 39°151 0011 N.; longitud~, 68° 081 0011 W.; 

depth, 3,249 meters; bottom temperature, 2.7° C. 

6. Luidia clathmta Say. Near Charleston, S.C.; latitude, 32° 47' N.; longitude, 79° 571 W.; depth, between2 and 

22 meters. 

7. Acanthaster planci Linne. Palmyra Island, in the Pacific Ocean, west of south from Hawaii, latitude, 5° 49' N. 

8. · Asterina minuta {hay. Pigeon Point, 'fobago, British West Indies; latitude, 11° 25' N.; shoal water near shore; 

teniperature, 28° C. 'Veight of specimen, ''a large adult," only 0.1009 gram; a quantity insufficient for complete 

analysis. · . 

· 9. Linckia guildingii Gray. Pigeon Point, Tobago, British West Indies; latitude, 11° 25' N.; shoal water near 

shore; temperature, 28° C. · 

10. Ctenodiscus crispatus Retzius. Albatross station 2434; off Newfoundland; latitude, 43° 081 N.; longitude, 50° 

401 W.; depth, 93 meters; bottom temperature, 1.1° C. 

11. Odontaste1· hispidus Verrill. Off Marthas Vineyard, Mass.; depth, 245 meters; bottom temperature, 11.1° C. 

12. Plutonaster agassizii Verrill. Off Marthas Vineyard, Mass.; depth, 584 meters; bottom temperature, 6.6° C. 

13. Asterias forbesii Desor. Vineyard Sound, ·Mass. 

14. Pontaster tenuispinus Verrill. Albatross station 2095, between Qape Hatteras and Nantucket; latitude, 39° 

29' N.; longitude, 70 ° 581 4011 W.; depth, 2,456 meters; bottom temperature not given. 

15. Astropecten articulatus Say. West coast'of Florida. 

Analyses 8-15 by R. :M. Kamm. _ 

16; 01·thaste1·ias tanneri (Verrill). Albatross station 2307, between Cape Hatteras and Nantucket; latitude, 35° 

421 N.; longitude, 74° 541 3011 W.; depth, 79 meters; bottom temperature, 14.5° C. 

, 17. Urasterias linckii (Muller and Troschep1

• East of Nova Scoti~; latitude, 44° 32' N.; longitude, 57° 09' W.; depth, 

· 403 meters. Weight of sample, 0.6226 gram. · · 

18. Asterias acervata borealis Perrier. Western Bank, east of Nova Scotia. 

19. Astropecten americanus Verrill.· Off Marthas Vineyard, Mass. 

20. Phataria bifascialis Gray. Cape San Lucas, Lower California; latitude, 22° 52' N. 

21. Pharia pyramidata Gray. Cape San Lucas. 

22. Oreaster occidentalis Verrill. San Jose Island, Gulf of California; latitude about'25° N. . 

23. Ctenodiscus procurator Sladen. Albatross station.2780, off coast of Chile; latitude, 53° 0~1 S.; longitude, 73° 

42' 3011 W.; depth, 675 meters; bottom temperature, 8,.3° C. · 

24. Ctenodiscus australis I:utkin. Off Patagonia. 

25. Marthasterias glacialis (0. F. Muller): Horta Harbor, Azores; latitude, 38° 35' N.; longitude, 28° 50' W. 

approximately. 

26. As.terina pectinifera Muller and Troschel. Otaru, Hokushu, Japan; latitude about 43° N. 

27. Culcita novaeguineae Muller and Troschel. An East Indian tropical species, locality unknown. 

28. Linckia laevigata (Linne). An East Indian tropical species, locality unknown. 

29. Coscinasterias calamaria Gray. New Zealand. 

Analyses 16-29 by B. Salkover. 

Analyses of starfishes. 

1 2 

---

Si02 •••••••••••••••••••••••••••••••••••••••••••••••• 0.45 0.79 

~8~.~~~~:·::: ': ·:·:·:::: :·:·:·:·:::: :·:·:·:·:::: ':':":':':::: ':":":':' :':: .21 . 55 

2.59 8.83 

35.71 38.51 

S03 •••••••••••••••••••••••••••••••••••••••••••••••••• (?) (?) 

rg~ti~~~:: .-.:: : : : : : :,: : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : .07 . 24 

60.18 54.S9 

99.21 98.81 

C02 needed ............................. · ............ 30.84 34.30 

Organjc matter, etc .................................. 29.34 20.59 

3 4 

------

0.03 1. 21 

.20 .48 

3.98 3.05 

36.85 30.35 

(?) (?) 

.14 .13 

57.64 63.91 

98.84 99.13 

33.05 27.08 

24.59 36.83 

5 6 

---

2.06 0.27 

. 79 . 36 

3.93 4.89 

40.54 41. 30' 

(?) (?) 

.11 .14 

51.66 52.03 

99.09 98.99 

'35. 94 37.70 

15.72 14.33 

7 

0.19 

.14 

4.36 

33.18 

('?) .07 

62.07 

100.01 

30.81 

31.26 
ECHINODERMS. 27 

Analyses of starfishes-Continued. 

8 9 10 11 12 13 14 

--------- ---

SiO~ .................. · ......................... : .. } 0.49 { 0.16 0.31 0.51 0.31 0.10 0.27 

(Al, ~eh03 •••••••••••••••••••••••••••••••••••••••••• .18 .17. .10 .16 .48 .39 

~:8 ...... ~ : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : 4.16 4.64 3:26 4.12 3.81 2.39 3.67 

33.90 32.28 39.50 40.44 44.43 30.93 43.93 

f~~;;~~: :_: _::: : : : : : : : : : : : : : :: : : : : : : : : : : : : : : : : : : : : : : : Trace. Trace. . 26 .03 (?) . 22 .06 

(?) .71 .75 . 69 . 62 .82 .46 

59.97 61.32 55.70 53.15 50.02 65.35 50.20 

98.52 99.29 99.95 99.04 99.35 100.29 98.98 

C02 needed ......................................... 31.22 30.97 33.97 35.90 36.76 26.08 38.28 

Organic matter, etc .................... ·: ............ 28. 75 31.25 21.73 17.25 13.26 39. 27 11.92 

15 16 17 18 19 20 21 

SiO~ .... · ................... · ........................ 0.21 0.68 0.00 0.45 0.18 1. 29 0.66 

(Al, ~e) 3 ••••••••••••••••••••• · •••••••••••••••••••••• .20 1.25 1. 38 1.20 .20 6.89 1.16 

~f8: .............. : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : 5.44 3.39 2.06 2.50 2.95 4.03 4.02 

42.30 32.21 24.22 26.79 30.50 26.44 29.13 

~:~~-.-::·:: _:_:_:_ :_::::::::::::::::::::::::::::::::::: 

Trace. Trace. Trace. Trace. Trace. Trace. Trace. 

.74 Trace. Trace. Trace. .47 .75 .76 

50.71 .61. 27 70.28 64.37 62.91 61.22 63.92 

99.60 98.80 97.94 95.31 97.21 100.62 99.65 

C02 needed ..................... · .................... 38.81 29.04 20.94 23.80 -26.96 24.17 26.89 

Organic matter, etc .................................. 11.90 32.23 43.34 40.57 35.95 37.05 37.03 

22 23 24 25 26 27 28 . 29 

------- -· 

SiO~······································ 0.05 Trace. 0.70 0.23 0.24 0.08 0.49 0.00 

(Al, e)20 3 •••••••••••••••••••••••••••.••••••• 2·. 57 0.79 5.10 1. 64 .83 .50 1. 87 1. 81 

~:8: .. ::: : : : : : : : : : : : : : : : : : : : : : : : : : : : : : :: : : : 4.22 3.18 3.33 2.23 1. 95 3.72 3.15· 2.77 

29.44 29.32 30.12 27.75 16.04 25.59 24.48 24.78 

J~~:i~~: _:_:_:::::: :: : : : : : : : : : : : : : : : : : : : : : : : : : Trace. Trace. Trace. Trace. Trace. Trace. Trace. Trace. . 81 Trace . Trace. .80 .46 . ~4 .71 Trace . 60.91 64.15 58.42 67.27 79.46 68.67 66.94 69.29 

98.00 97.44 97.67 99.92 98.98 99. 30. 97.64 98.65 

C02 needed ................. ~ .............. 27.32 26.54 27.33. 23.81 14.50 l 23.79 30.17 23.52 

Organic matter, etc .......................... 33.59 37.61 31.09 43.46 64.96 44.88 S6.77 45.72 

Reduced analyses of star.fis'/l£s. 

1 2 3 4 5 6 7 

------------

SiO~··············································· 0.64 1.01 0.03 1. 94 2.47 0.32 0.27 

(Al, e)20 3 ••••••••••••••••••••• · •••••••••••••••••••••• .30 .70 .27 .. 77 .94 .42 .20 

8~8:: ":": .: ·:·:·:· ....... :_ :_: : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : 7.79 10.28 11.24 ·10.27 9.88 12.13 13.33 

91.06 87.44 8K06 86.57 86.42 86.77 85.99 

(?)" (?) (?) (?) (?) (?) (?) 

Ca3P20 11 ••••••••••••••••••••••••••••••••••••••••••••• .21 .57 .40 .45 .29 .36 .21 

100.00 100.00 100.00 100.00 100.00 100.00 100.00 

8 9 10 11 12 13 14 

---- ' 

!~~t;:.:;:-)_~_t.·;·in ~ ~ l /: ~ ~ u ~ ·;. l ~ ~ l;:; ~ ~ ·. 

} 0.70 { 0.24 0.39 0.62 0.35 0.17 0.31 

.26 . 22 .12 .18 .78 .45 

12.53 14.31 . 8. 78 10.58 9.09 8.24 8.86 

86.77 83.42 88.48 87.16 89.18 88.19 89.34 

Trace. Trace~ .73 .08 (?) .78 .15 

(?) 1. 77 1.40 1.44 1.20 1. 84 .89 

100.00 100.00 100.00 I 100.00 100.00 100.00 100.00 
28 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. 

Reduced analyses of starfishes -Continued. 

\ 

15 16 17 18 19 20 21 

~~~-:-~::::: :::: ~ ~: -~ -~. ~- ~ ~ ~ ~ ~ ~ ~ : : ~ ~ ~ ~ ~ ~ ~ : ~ ~ : : ~ ~ ~ ~ ~ ~ ~ ~ : ~ . 

0.24 1. 02 0.00 0.82 0.29 2.06 1. 06 . 23 1. 88 2.84 2.19 . 33 11.04 1. 86 

13.02 10.69 8.91 9.60 10.11 13.56 13.52 

85.08 86.41 88.25 87.39 87.93 71.28 81.82 

8:s~~~ ·. ·_ ·_ ·_ ·_ ·_ ·_ ·_ ·_ ·_ ~ ~ : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : Trace. Trace. Trace. Trace. Trace. Trace. Trace. 

1. 43 Trace. Trace. Trace. 1. 34 2.06 1. 74. 

100.00 I 100.00 I 100.00 I 100.00 I 100.00 100.00 I 100.00 

22 23 24 25 26 27 28 29 

------

~J~:.:-~~~ :·;-~-~ ~ ~ ~ ~ ~: ~ ~ ~: ~ ~ ~ ~ ~: ~: ~ ~ ~:: ~:: ~ ~ 0.06· Trace. 1. 05 0.41 0. 71 0.15 0.70 0.00 

3.99 1. 32 7.66 2.90' 2.44 . 92 2.64 3.49 

13.76 11.16 10.51 8.29 12.05 14.35 9.32 11.22 

80.04 87.'52 80. 78 85.99 82.51 82.27 85.63 85.29 

8:s~~~ ~ ~ ·_ ~ ~ : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : Trace. Trace. Trace. Trace. Trace. Trace. Trace. Trace. 

2.15 Trace. Trace. 2.41 2.29 2.31 1.71 Trace. 

100.00 I 100.00 I 100.00 100.00 I 100.00 I 100.00 100.00 I 100.00 

I~ the following table the analyses are arranged 1n the order of ascending magnesium 

carbonate, like those of the crinoids and sea urchins. 

Magne.~ium carbonate in starfishes. 

Locality. Latitude. 

Eastport.' .............. . Vineyard Sound ....... . . Azores ..•............... Newfoundland ......... . Station 2205 ............ . Nova Scotia ........... . 

Asterias vulgaris ................... . Asterias forbesi .................... . Marthasterias glacialis ~ ............ . Ctenodiscus crispatus .............. . Pontaster tenuispinus ............. . Urasteriaslinckii. ................ . 

44° 55' N .......... . (?) ............... . 

38° 351 N ......... .. 43° 08' N .......... . 39° 291 N .......... . 44° 32' N ......... .. Marthas Vineyard.:: .... . East Indies ........... .. West Bank ............ . Station 2568 ........... :. Marthas Vineyard ...... . Station 2250 ............ . Station 2309 ............ . 

Plutonaster agassizii. .............. . Linckia laevigata ................. . Asterias acervata ................. . Benthopecten spinosus ............. . Astropecten americanus ........... . Lep,tasterias compta ............... . Asterias tanneri ....... ~ ....... : ... . 

(?) ................ . 

(?) ................ . 

(?) ................. . 

39°151 N .......... . (?) ................. . 

"40° 17' N .......... . 35° 43' N .... · ...... . Patagonia .............. . Marthas Vineyard ...... . Station 2307 ............ . 

Ctenodiscus australis ............... . Odontaster hispidus .............. . Orthasterias tanneri. ....... : . ..... . 

(?) .... · ............. . (?) ................ . 35° 42' N .......... . Off Chile .... ·, ......... . N e:v Z~~land .... :· ....... . Califorma ............. . Japan ......... .': .. .- ... . Charleston ............ . Tobago ............. , ... . Florida ............... . Palmyra Island ........ . Cape San Lucas ....... . Cape San Lucas ....... . San Jose Island ........ . 

Cte.nodiscus procurator ...... ~ ..... . Cosci1;1aster~a~ calamaria ........... · .. Asterma ;mm1ata. · ................ . Asterina pectinifera .............. ·. ~ 

Luidi_a clat~ata ................... . Asterma mmuta ................... . Astropecten articulatus ............ . Acanthaster planci ................ . Pharia pyramidata ........ · ........ . Phataria bifascialis ....•............ Oreaster occidentalis ............... . 

53° 01' S .......... . (?) ................. . 

36° 38' N ......... .. .43° N: ............. . 32° 47' N .......... . 11 o 25' N .......... . (?) ...... ~ .......... . 

5° 49' N ........... . 

22° 52' N .......... . 22° 52' N .... .' ..... . 25° N ............ · .. . 

Tobago ................ . East Indies ............ . Linckia guildingii. ................ . Culcita novaeguineae .............. . 11° 25( N .......... . (?) ................. . 

·Depth. 

Meters. 

(?) . 

(?) 

(?) 

. 93 

2,456 

403 

584. 

(?) 

(?) 

3, 249 

(?) 

86 

102 

(?) 

245 

79 

675 

(?) 

(?) 

(?) 

(?) 

ShoaL ... . (?) 

(?) 

(?) 

(?) 

(?) 

ShoaL ... (?) 

Temper- MgC03

. 

ature. 

oc. 

(?) 

(?) 

(?) 

1.1 

(?) 

(?) 

6.6 

(?) 

(?) 

2.7 

(?) 

10.8 

(?) 

(?) 

11.1 

14.5 

8.3 

(?) 

(?) 

(?) 

(?) 

28 

(?) 

(?) 

(?) 

(?) 

(?) 

28 

(?) 

Per cent. 

7. 79 

8.24 

8.29 

8. 78 

8.86 

8 .. 91· 

9.09 

9.32 

9.60 

9.88 

10.11 

10.27 

10.28 

10.51 

10.58 

10.69 

11.16 

11.22 

11.24 

12.05 

12.13 

12.5:3 

13.02 

13.33 

13.52 

13.56 

13.76 

14.31 

14. 3.5 

Here again we meet the same coordination of temperature with content of magnesium 

carbonate that was noted in the series of crinoids, sea urchins, and alcyonarians. As a general 

rule the species from cold regions are lowest in magnesia; those from warm regions are higher. 

The regularity, however, is ~ot absolute, and the variations from it may be real, or perhaps may 
ECHINODERMS. 29 

be due to differences in habitat. The star:fisheA from Vineyard Sound and Marthas Vineyard, 

for example, may have come from different depths of water, and so; too, ~ay the two marked 

as from the East Indies. Small impurities· in the specimens analyzed may also account for 

some irregularities. The analyses hy.L. Schmelck,25 ofstar:fish~s froin ·the North Sea, are, 4ow-. 

ever, not so easy to explain. . His data. are as follows: 

1. Atcaster tenuispinus. 

perature, 6° C. ' 

Station 10; latitude, 61° 411 N.; longitude, 3° 191 E.; depth, 402'meters; bottom tem- , . 

2. Atcaster tenuispinus. 

perature, 6.9° C. 

Station 25; latitude, 63° 10' N.; longitude, 5° 25' E.; depth, 179 meters; bottom tem3. Astmpecten andromeda. Station 10; details under No. 1. 

Schrnelck' s analyses of starfishes. 

Si02 ••••••••••••••••••••••••••• 

f!l~-: (/\;!! [ [ [ [)~) [ [).. [ Organic matter .............. .. 

1 

I' 

.Trace. 

Trace. 

0.31 

9.36 

74.11 

(?) 

Trace. 

16.23 

100.00 

_ Schmelck' s analyses ?'educed. 

1 

Trace. 

Trace .. o:36 

11.18 

88.46 

(?) 

Trace. 

2 

Trace. 

Trace. 

0.60 

9.24 

78.92 

,1.00 

Trace. 

10.24 

100.00 

2 

Trace. 

Trace. 

0.66 

10.29 

87.92 

1.13 

Trace. 

3 

7.09 

. 65 

. 37 

8.55 

64.48 

. 83 

Trace. 

13.83 

~5.80 

3 

8.66 

. 79 

.45 

10.42 

78.67. 

1. 01 

.Trace. 

Nos.' 1 and 3 of this group of analyses represent the same locality and temperature .. If: 

the excessive and surely extraneous silica in No. 3 is rejected and the analyses ~re r_ecalculafea 

to 100 per cent the figure for magnesium carbonate becomes 11.41, nearly that for No. 1. All 

the percentages g~ven for magnesium ca;rbonate are higher than the te~peratur~s would lead 

us to expect, but they still fall far below the highest ten in our table. A general tendency .to 

regularity as regards temperature is evident, but it is perhaps not invariable. 

4. ·oPHIURANS OR BRITTLE STARS. 

The following data relate to the composition of the ophiurans, as distinguished from the 

ordinary starfishes. · · · · 

1. Gorgonocephalus a1'cticus Gray. Off Cape Cod, Mass.; about latitude 42° N. 

2. Gorgonocephalus canJi Lyman. Alaska. < 

3. Ophioglypha sarsii L\it.ken. Albatross station 2176; latitude, 39° 321 3011 N:; longitude, 72° 211 3011 W.; depth, 

553 meters; bottom temperature, 5° C. 

4. Ophioderma cinereum Muller and Troschel. Ensenada Honda, Culebra Island, east of Porto Rico; about latitude 18° 20' N. 

5. Ophiomyxajlaccida Say. Pigeon Point, Tobago, British'West Indies; latitude, 11° 25' N.; shoal water, close in 

shore; temperature, 28° C. Adult specimen. ' 

2~ Sclm1olck, L., Norsko Nordhavs Exped., No. 28, p.l29;1901. 
30 THE INORGANIC CONSTITVENTS OF MARINE INVERTEBRATES. 

6. Like No. 5; half grown. _ 

7. Ophiocoma pumila Lutken. Pigeon Point, Tobago. Like No.5. 

Analyses 5-7 by R. M. Kamm; specimens received from Dr. Hubert L. Clark. 

8. Ophiocoma aethiops Lutken. Espiritu Santo Island, Gulf of California; about latitude 24° 30' N.' 

9. Ophiothrix angulata Ayres. Cuba. · 

10. Gorgonocephalus eucnemis Muller and Troschel. Albatross station 4912, 17! miles off Tsurikake Island, 

southern tip of Japan; latitude, 31° 39' 4011 N·.; longitude, 129° 20' W.; depth, 715.5 meters. 

11.· Ophiopholis aculeata japonica Lyman. Illiuluk, Unalaska; latitude, 53° 42' N.; longitude, 166° 32' W. 

12. Ophioglypha lymani (Ljungman). Off coast of Chile; latitude, 48° 411 S.; longitude, 74° 241 W.; depth, 355 

meters; bottom temperature, 11.1° C. 

13 .. Ophiomusium lymani W: Thomson. Albatross station 3407; off the Galapagos Islands; latitude, 0° 04' S.; longitude 90° 24' 3011 W.; depth, 1,619 ·meters; bottom temperature, 2.9° C. ' 

14. OphiocamaX jasciculata Lyman, Albatross station 2125, Caribbean Sea; latitude, 11° 43' N.; longitude, 

60° 09' 3011 W.; depth, 381 meters; bo}tom temperature, 10.4° C. 

15. Ophionereis eurybrachiplax H. L. Clark. Albatross station 3702, off Honshu Island, Japan; depth, 57-75 meters. 

16. Ophiocoma e:rinaceus Muller and Troschel. Hawaiian Islands. 

17. Ophioglypha liitkeni Lyman. Albatross station 3114, off central California; latitude, 37° 06' N.; longitude, 

122° 32' W.; depth,.113.5 meters. ' 

Analyses 8-17 by B. Salkover. 

Analyses of ophiurans. 

1 2 3 4 5 6 

~~-~; !:~~ ~ ~ ·: ·:. :·: ~ ~ ~ ~ ~ : ~ : ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ : : ~ : ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ 1. 77 1. 08 0.98 0.21 0.11 0.43 . 62 . 72 . 53 . 09 .17 . 55 

3.36 2. 82 3.99 5.80 4.67 '4. 64 

36.13. 29.80 42. 14 41.32 31.57 30. 14 

f~~~~L: :_:_:: :: _: _:_ :_:: : :: _: _:_ :_::: _:_:: : : : : : : : : : : : : : : : : : : : . 22 . 32 . 29 . 07 Trace. 'frace . (?) . 69 (?) .16 .• 1. 42 l. 60 

55. 72 63.37 50.95 51.58 61.09 ' 62.59 

97.82 98.80 98 .. 88 99.23 99.03 99.95 

C02 needed ....................................... ; .... 31.65 25.85 37. 29 38. 69 29. 17 27.90 

Organic m!J.tter. etc ............................. : ...... 24.07 37.52 13.66 12. 89 31.92 34.69 

7 8 9 10 11 12 

i~~~ j~~~ ~-;-;-;-~: : : : : ~ ~ ~ ~ ~ : : : : : ~ : : ~ ~ ~ ~ : : : : : ~ ~ : : : : : : ~ ~ ~ } 

0.36 { 0. 10 0.26 0. 17 0.·35 0.00 

. 67 . 67 l. 99 .~40 1 .. 64 

4. 77 3.06 4.99' 2. 72 3.44 2.'85 

37.20 47.12 42.18 33. 70 46.21 36.44 

f~~~i~:: : : : : : : : : : : : : : : : : : : : : : : : : :,: :: : : : : : : : : : : : : : : : : . 05 Trace. (?) Trace. (?) Trace. 

. 90 Trace. (?) Trace. (?) Trace . 

56.04 47.96 52.11 61.51 49.74 58.76 

99.32 98.91"\ 100.21 100.09 100.14 99.69 

C02 needed ............... · ...... ' ................ · ....... 33.93 40.39 38.18 29.47 39. 73 31. 77 

Organic matter, etc ... ~ .............................. , 22. 11 7.57 13.93 32.04 10.01 16.99 

13 14 15 16 17 

Si02 •••••••••••••••••••••••••••••••••••••••••• : • ••••••••••• 0.00 0.00 0.00 0.00 0.00 

~0~~~~~3 .. ::: ·.·.·:::::::::::::::::::::::::::::::::::::::::: . 67 . 95 . 89 . 94 2.58 

3.04 3.18 5.20 4:83 3.61 

CaO ......................................................... 50.09 44.80 39.09 40.91 35.95 

~b~~: :. : : : : : : : : : : : : : : : : : : : : : : : : : ": : : : : : : : : : : : : : : : : : : : : : : : : : 

Trace. Trace. Trace. Trace. Trace. 

Trace. Trace. Trace. Trace. Trace. 

Ignition .................................................. 46.05 50.02 53.87 51.43 55.21 

99.85 98.95 99.05 98. 11 97.35 

C02 needed .................... " ............................ 42. 79 36. 70 36.43 37.45 32. 22 

Organic matter, etc .......................................... 3.26 13. 32 17.44 13 .. 98 28.99 
0 

BCHINODERMS. 

Reduced analyses of ophiurans. 

1 

Si02 •••••••••••••••••••••••••••••••••••••••••. • ••••••• 2. 39 

~~gt_:~:·:·:::::::::::::::::::::::::: ~::::::::::::::: . 84 

9. 53 

86. 60 

g~~d~~ ~ : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : . 64 

(?) 

100.00 

--------------------------------------- ~---1 ~l?, :i4'~)i>~---~:::::::::::::::::::::::::::::::::::::::::} 

~lf83as: .": : : : : : : : : : : : : : : : : : : : : : : : : : : : : :·: : : : : : : : : : : : : : : 8~s~~~ ~ : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : 

0. 47 { 

12. 97 

84.441 .14 

l. 98 

I 

2 

l. 76 

l. 17 

9. 66 

84. 36 

l. 14 

L 91 

100.00 

8 

o.n 

. 73 

7.04 

92. 12 

Trace. 

Trace. 

3 

1. 15 

. 62 

9. 84 

87. 65 

. 74 

(?) 

100.00 

9 

0. 31 

. 77 

11. 68 

87.24 

(?) 

(?) 

100. oo I 100.00. 100.00 

13 

0.00 

. 69 

6. 61 

92. 70 

'l'race. 

Trace. 

100.00 

14 

0.00 

l. 08 

7. 62 

91. 30 

Trace. 

Trace. 

100.00 

4 

0.24 

.n 

14. 08 

85.09 

.18 

. 30 

100.00 

10 

0. 25 

2. 92 

8. 39 

88.44 

Trace. 

Trace. 

100.00 

15 

0. 00 

1.09 

13. 38 

85. 53 

Trace. 

Trace. 

100.00 

5 

0. 16 

. 70 

14. 56 

81.02 

Trace . ·3. 56 

100.00 

,n 

0. 39 

. 44 

8.01 

91. 16 

(?) 

(?) 

100.00 

16 

0.00 

1. 12 

12.05 

86. 83 

Trace. 

Trace. 

100 .. 00 

31 

6 

0. 66 

. 85 

14. 95 

79.37 

Trace . 4. 17 

100.00 

12 

0.00 

2. 26 

8. 24 

89.50 

Trace. 

Trace. 

100.00 

17 

0.00 

3. 47 

10. 19 

86. 34 

Trace. 

Trace. 

100.00 

Some of these analyses are obviously incomplete, for phosphates and sulphates were not 

determined, because of insufficient material. A number of the specimens analyzed were fragments, and in that respect the analyses are not altogether satisfactory. In the following table 

the data are presented as in the tables for the other echinoderms; the figures for the. two specimens of Ophiomyxafiaccida are averaged. The localities and latitudes are abbreviated as usual. 

Percentage of magnesiunt Carbonate in ophiurans. 

I 

Locality. Latitude. Depth. Temper- MgC03 • ature. 

Meters. oc. Per cent. 

Ophiomusium lymaui. ·.:.-· ......... Galapagos .............. 0° 041 s. 1, 619 2. 9 6. 61 

Ophiocoma aethiops ............... Gulf of Califomia ........ 24° 30' N. (?) (?) 7.04 

Ophiocamax fasciculata ............. Caribbean Sea ......... 11° 43' N. 381 10.4 7. 62 

Ophiopholis aculeata .............. Unalaska .............. 53° 52' N. (?) (?) 8.01 

Ophioglypha lymani. .............. Chile ................... 48° 411 s. 355 n. 1 8. 24 

Gorgonocephalus eucnemis ......... Japan ................. ·. 31° 39' N. 715. 5 (?) 8. 39 

Gorgonocephalus arctic us .......... Cape Cod .............. 42° ± N. ??) (?) 9. 53 Gorgonoce~halus caryi. ............ Alaska ................. · (?) ?) (?) 9. 66 

Ophioglyp a sarsii .......... · ...... Station 2176 ...... · ..... 39° 32' N. 553 5 9. 84 Oph~o~pha lutkeni ............... California .............. 37° 061 N. 113. 5 (?) 10. 19 

Oph10t IX augulata ................ Cuba .................. (?) (?) (?) n. 68 

Ophiocoma erinaceus .............. Hawaii ................. (?) (?) (?) 12.05 

Ophiocoma pumila ... ~ .......•..... Tobago ................. no 25' N. Shoal. 28 12.97 

Ophionereis eurybrachiplax ........ . Japan .................. (?) 61 (?) 13. 38 

Ophioderma cinereum ............. Culebra.· ............... 18° 20' N. (?) (?) 14.08 

Ophiomyxa flaccida ................ Tobago ................. no 25' N. Shoal. 28 14. 75 

In this table the relation between magnesium ca1;bonate and temperature is less regular 

than it was with the other echinoderms. Some of the low percentages are unaccountably low; 

but th_e last si...~ in the series show percentages characteristic of warm-water organisms. One 

106185-22--3 
32 THE INORGANIC CONS1'ITUENTS OF MARINE INVERTEBRATES. 

analysis of an ophiuran, Astrophyton, by L. Schmelck, 26 remains to be noticed .. The accompanying data are as follows: Locality, station 37 in the North Sea; latitude, 78° 48' N.; longitude,· 

8° 57' E.; depth, 19~ 1neters; bottom temperature, 1.1° C. 

Analysis of Astrophyton. 

Actual 

analysis. 

Reduced 

analysis. 

) 

Si02 • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • Trace. Al~03 ••••• ~ ••••••••• , • • • • • • • • • • • • • • • • • • • • • Trace. 

~!§g,;_:::::::: :.:::::::::::::::::::::::::::: 7~ ~g (iaS04 • •• • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 71 

P20 5·••••••••••••••••••••••••••••••••••••••• Trace. 

Organic matter ........... _................... 17 57 1-----11 

. ~i02 ...................... · ............... . Al 20 3 •••••••••••••••••••••••••••••••••••••• 

·Fe20 3 ••••••••••••••••••••••••••••••••••••• 

~:88a3. ·. : .·. ·. ~: : : : : : : : : : : : : : : : : : : : : : : : : :·: : : : : 

CaS04 •••••••••••••••••••••••••••••••••.•••• 

Ca3P20 8 ••••••••••••••••••••••••••••.••••••• 

Trace. 

Trace~ 

0. 37 

9.11 

89. 67 

. 85 

Trace. 

100.01. 100.00 

In this, as in Schmelck's analyses of starfishes, the magnesium carbonate is high for the . 

temperature given. 

5. HOLOTHURIANS. 

Although the holothurians are of siight importance as contributors to the marine sediments, 

it seemed desirable to compare a few of them with the other echinoderms. Four holothurians 

were therefore analyzed, bu.t their organic matter was so vastly in exces.s of their hard parts 

that the analyses, with one exception, are far from satisfactory. However, the data, which are 

. not without s~me significance, are as follows: 

1. Ffolothuria floridana Pourtales. Fajardo, Porto Rico. 

, 2. · Cucumariafrondosa (Gunnerus). A northern cold-wat~r species, precise locality unknown. 

3. Trochostoma intermedium Ludwig. Off Cerros Island, Calif. Weight of sample, 0.2204 gram. 

4. ·Trochostoma intermedium. Albatross station 3307, in Bering Sea; latitude, 56° 12' N.; longitude, 172° 55' W. 

depth of water, 130 meters; bottom temperature, 3.3° C. Weight of sample, 0:1175 gra,m. 

Analyses' of holothurians\ 

1 2 3. 4 

SiD2 •••••••••••••••• · ••••• 0.08 0. 27 5. 56 6.93 

(.Al,Fe)20 3 •••••••• : ••••••• . 18 . 25 8.40 6. 31 

MgO ..................... 3.48 . 41 . l. 81 1. 36 

CaO ...... · ............... 29. 19 1. 10 4. 91 1.44 

PzOs· ···········' . .' ...... Trace. . 85 3. 15 1. 86 

803 •••• , ................... . 74 1. 29 (?) (?) 

Ignition· .................. 69. 93 95. 71 76. 71 82. 21 

99. 60 99. 88 100. 54 100. 11 

C02 needed ............... 23. 21 (?) (?) ~?) o~;ganic matter, etc ....... 46. 72 . (?) (?) ?) 

In No. 2, 0~31 per cent of matter soluble in water was· found, which raises the summa:tion; 

to 100.19. 

Of these analyses only the first is reducible with any certainty to standard form. The 

percentage composition of the inorganic portion of the specimen then becomesSi02 •••••••• ." •••••••••••••••••••••••••••••••••• : ••••••••••••••••• .-••••••••••••••••••••••••• 

(Al,Fe)z03 •· •••••••••••••••••••••••••••••••••••••••••••••.•••••••••••••••.••••••••••••••••••• 

MgC03 •••••••••••••••••••••••• •.• •• ~ •••••••••••••••••••••••••••••••••••••••••••••••••• .' ••••• 

z:~g: ~ .· .· :·. ·. ~ ~ ~· ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ : ~ ~· ~ ~ ~ ~ ~ ~ ~ : : : : : : : 

·ca3P20 8 ••••••••.•••••••••••••••••••••••• · •••••• : ••••••••••••••••••••• • .. :.· ••••••••••••••••••••• 

1', .' 

26 Schmelck, L., Norske Nordpavs Exped., No. 28, p.129, 1901. 

0. 15 

. 34 

13. 84 

83.29 

2.38 

Trace. 

100.00 

• 
ECHINODERMS. 33 

This is very near the.analysis of the ophiuran from Culebra Island, which is not far frmn the 

locality of this holothurian. The coincidence is very striking. 

In the specin1en represented by analysis 2 the proportion of inorganic 1natter was .so small 

that good results could hardly be obtained. The other two analyses were also made with altogether inadequate n1aterial. These analyses, however, show that the s1nall inorganic portions 

of the holothurians .are relatively rich in phosphatic matter and in n1agnesia. The difficulties 

of discussing them intelligently 'are like those of discussing the worn1 tubes and the axes of 

alcyonarians. In Trochostoma there are brown spots, which were already known to be phosphatic, although quantitative analyses of them seem to be lacking. The true character of such 

bodies can· be deter1nined only by an elaborate investigation upon abundant 1naterial, a task 

which lies outside of our main problem. 

Two analyses of holothurians are already on record, as follows: 

l. Stich'Jvus ·regalis. Locality not stated; analysis by 0. Btitschli, K. Gesell. \Viss. Gottingen Abh., No. 3, 1908. 

2. Ash of the epidermis ("lederha.ut") of a large holothurian, species and locality not stated; analysis by Hilger. 

l'flttger's Archiv f. Physiologic, vol. 10, p. 212, 1875. · 

Old analijses of lwlothurians. 

1 

. ~ ~~~ ~ · ..... · ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ Ca003 • • • • • • • • • • • • • • • • • • • • • • • • • • 81. 54 

Mg003 •••••••• : •••••••••••• .-.... 8. 10 

CaS04 •••••••••••••••••••••••• : • ••••••••••• 

2 

4.47 

. 83 

78. 96 

12. 10 

1.04 

. UaS04 .2ll20...................... 2. 09 .......... . Ca3P20 8 ••••••••••••••••••••• ~ ••••••••••••• ·• . 96 

1 . 2 

~:8?. ~ • .... : ~ ·.·:: : : : : : : : : ~ : : : : : : : : : ~ ••••• 3.' 67 ....... ~·. ~~ Sand ............................. · 3. 23 ......... . 

Si02 ••••••••••••••••••••••••••••••••••••• ·• • 57 

Organic . . . . . . . . . . . . . . . . . . . . . . . . . l. 03 ......... . 

99. 66 99. 95 

These analyses, tog~ther with ours of Holothuria, show that the hard parts of these animals, 

like those ~f the other echinoder1ns, are distinctly magnesian. Although definite data are 

wanting, it seems probable that Hilger's specimen came from warmer wa.ter than that analyzed 

by Biitschli. The sodium salts are of course extraneous. · 

6. SUMMARY. 

FrOin the evidence now available it seems ahnost certain that the inorganic constituents 

of any echinoderm will have the composition of a moderately magnesian iimestone. There n1ay 

be exceptions, but none has yet been found. The five tables-for crinoids, sea urchins,. stt'trfishes, ophiurans, and· holothurians-all tell the same story, and with remarkable unanimity. 

Furthermore, the proportion of magnesium car:bonate appears to be a function of tmnperature, 

the organisms from warn1 waters being richer in it than those from cold waters. The exceptions 

to this rule are few and may be only apparent, for cold or warm currents and varying deptl;.ls 

of water account for nearly all irregularities.· Schmelck's analyses of starfishes frmn the North 

Se.a are the most .troublesome to explain. 

The sea urchins seem to be a little poorer in magnesia than either of the other groups, but 

the analyses are 'fewer and therefore less conclusive. Silica and sesquiox:ides are probably 

altogether extraneous, although it is possible that small quantities of thmn rnay really belong 

to the organisn1s. As shown by Meigen's reaction, all the echinodenns studied are calcitic, and 

no evidence of aragonite in them was found. 

The te1hp,erature relations shown by the analyses offer an interesting biological problem, 

with which we can not undertake to cope. It is not clue to differences of comp.osition in the 

solid 1natter of sea water, for that is practically uniform the world over. In all the great oceans, 

and even in minor bodies of water like the Mediterranean, the Baltic, and the Black seas, the 

pr.oportion of magnesia to lime is very nearly if not actually constant. In gaseous contents 

and especially in carbon dioxide the waters vary; the gases being more soluble in cold than 

in wa~·1n water. 'Whether this fact has any relation to the phenomenon under discussion we 

can not att~~pt to say. We can only report the facts and leave their biological discussion 

to others. · 
34 THE INOHGANTC CONSTITUENT'S O:F MARINg INVERT:I<;BHATES. 

BRYOZOA ... 

Thirteen Bryozoa have been analyzed in the course of this research, as follows: 

1. Schizoporella' unicorn1:s Johnston. Vineyard Sound, Mass.; depth· of water, 15.5 meters; bottom temperature, 

20.5° c. 

2. Schizoporella unicornis Johnston. Between John~ Pass and Pass a Grille Fla. 

3. Microporella grisea Lamouroux. Australia. 

4. Cellepora incrassata Lamarck. Northeast part of the Grand Banks. 

5; Flustra membranacea tru'(l,cata Smith. St. Paul Island, Pribilof group, Alaska; latitude, 57° N.; longitude, 

170° W. Only 0.1165 gram available for analysis. 

6. Lepralia pallasiana Moller. Off Gloucester, Mass. A thin coating on a granite pebble; weight of organism, 

0.4078 gram. 

7. Lepralia rosterigera (Smitt). Cedar Keys, Fla. 

8. Holoporella albirostris (Smitt). Cedar Ra.rids, Fla. 

9. Frondipora 1Jerrucosa Lamouroux. From the Naples Zoological Station, without definite statement as to locality. 

10. Amathia spira'lis Lamouroux. Albatross station 2619, between Cape Hatteras and Charleston, S. C.; latitude, 

33° 38' N.; lon&ritude, 7,7° 36' W.; depth, 27.5 meters. Only 0.3 gram w~s available for analysis. 

Analyses 7-10 by R. M. Kamm. 

11. Catenicella margaritacea Busk. Australia. 

12. Bugula turrita 'Desor. Georges Bank, off coast of Massachusetts; Albatross station 2578; latitude, 41° 20' 3011 

N., longitude 68° 34' 3011 W.; depth of water, 67.7meters; bottom temperature, 12.4° C. 

13. Bugula neritina Linne. Florida. · 

The actual analyses are as follows: 

Analyses of Bryo~oa. 

Si02 ••••••••••••••••••••••••••••••••••••••••••••••••• 

(Al,Fe)20 3 ••••••••••••••••••••••••••••••••••••••••••••• 

~'~!3.· _.: : : : : : : : : : : : : : : : : : : : : : : : .: : : : : : : : : : : : : : : : : : : : : : : 

P205 ................................................ . 

~~iiti~~: : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : 

C02 needed .......................... ~ .............. , _ Organic matter, etc ................. : ................ . 

1 

1. 65 

. 29 

. 28 

50.90 

Trace. 

7 

.73 

45. 64 

99.49 

39. 92 

5. 72 

8 

2 

2.49 

. 37 

2.04 

48. 17 

Trace. . 77 

45. 70 

99. 54 

40. 16 

5. 54 

9 

3 

0. 17 

.11 

. 49 

50. 83 

. 10 

. 79 

46. 80 

99. 29 

39. 94 

6. 86 

10 

4 

0.19 } . 19 

2. 74 

49.46 

Trace . . 98 

46.03 

99.49 

41. 26 

5.06 

11 

5 

3.00 

2.06 

30. 81 

. 09 

(?) 

61.45 

97.41 

26. 39 

35.06 

12 

6 

{ 4. 78 

. 13 

2. 32 

43.08 

.11 

(?) .............. 

50.42 

36. 29 

13. 29 

13 

-------------------------1------1------- -, 

Si02 • · •••••••• ~ •••.••••••••••••••••••••••••••••••••••• 

(Al,Fe)20 3 •••••••••••••••.•••••••••••••••••••••••••••• 

MgO .. -............................................. . CaO ............................................... . 

. P205································· .............. . 

~~ii"ti~~-- ·_ ~: ~ _· .-.-. ·. ~-~:: :: ::: : ::: : :: : : ::::: : :::: : _: : :': : : 

C02 needed ........................................ . Organic matter, etc ..... · ........................... . 

0.36 } 'o. 55 { . 39 

l. 25 1.16 

50.54 . 50. 9'4 

Trace. . 80 

45. 96 

99. 30 

40. 64 

5. 32 

Trace . .86 

45. 87 

99. 38 

40. 83 

5.04 

1. 27 }. 5. 75 . 64 

. 07 . 31 

48. 51 3.44 

Trace. (?) 

l. 49 (?) 

44.. 00 18. 79 

95.98 98.29 

3'7. -381 3. 04 

6. 62 85. 75 

Reduced analyses of Bryozoa. 

1 2 

2. 66 

. 39 

4. 58 

90. 90 

Trace. 

1.40 

100.00 

3 

0. 18 

. 12 

1. 11 

96. 90 

. 24 

1. 45 

100.00 

{ 0.95 

. 41 

3. 50 

40. 83 

.16 

1. 00 . 52.66 

4 

99. 51 

35.23 

17.43 

0. 20 } . 20 

6.07 

91. 77 

Trace . 1. 76 

100.00 

8. 77 

1. 18 

2. 55 

20.45 

. 38 

1. 47 

61. 51 

96.31 

17. 69 

43. 82 

5 

4. 82 { 

6. 94 

87. 92 

. 32 

(?) 

100.00 

6. 31 

. 75 

2. 57 

19. 69 

. 60 

2.43 

59.47 

91.82 

16. 41 

43.06 

6 

5.52 

.15 

5. 62 

88.44 

. 27 

(?) 

100.00 
BRYOZOA. 35 

Reduced analyses of Bryozoa-Continued. 

7 8 9 10 11 12 13 

---------------------1----1----1-----------------

~J~i·:-~~:: :::: ~ ~ ~ :: ·;· ;· ~ ~ ~ ~ : ~ : ~ ~ ~ ~ : : ~ ~ : : ~ : ~ . ~ . ~ : ~ : ~ ~ ~ ~ ~ 0. 38 } 0. 59 { l. 42 l. 15 16. 71 12. 94 . 41 . 72 . 50 2. 25 l. 54 

2. 80 2. 59 . 17 9. 57 8. 96 10. 19 11.08 

94. 96 95.2R 94. 86 90.43 86. 89 64. 51 63. 29 

8:~~~~ ·. ·. ·. ·. ·. ·. ·. ·. ~ ... ·. ~ ~ ~ : ~ ~ ~ ~ ~ ~ ~ : ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ : : : :. ~ : ~ ~ ~ ~ :I Trace. Trace. 'l'race. (?) . 43 l. 58 2. 68 

l. 45 1. 54 2. 83 (?) 2. 07 4. 76 8.47 

I 100.00 

I 

100.00 100. 00 I 100.00 

I 

100.00 .100.00 

! 

100.00 

I 

Reduced analysis No. 10, of Amathia, requires explanation. This bryozoan was one of the 

1nossy or fernlike forms and contained inclusions of sand and n1inute shells. Its organic matter 

was very large, and the entire material available was insufficient for a good analysis. The 

figures given for it in this table merely represent. the lime and magnesia alone, calculated as 

carbonates and to 100 per cent. The analysis is obviously of small sigJ+i,ficance, except in so far 

as it gives the ratio between magnesia and lime. To that extent it has some value. 

BryozoaN os. 1 to 5 and 7 to 9 were massive coralline forms and were easily handled. No. 6, 

Lepralia, was an incrustation upon a peb~le, from which it could not well be separated mechanically. The entire specimen was therefore weighed, the bryozoan was then dissolved by hydrochloric acid, and the pebble was weighed again. The solution only was available for analysis, 

and the usual loss on ignition could not be determined but was necessarily taken by difference. 

Such' an analysis is obviously unsatisfactory but not entirely worthless. Bryozoa Nos. 10 to 

13 were delicate mossy or fernlike organisms, and the poor summations of the last two indicate 

the presence of undetermined saline matter. The high silica in them is evidently due to 

inclosed sand. If silica and sesquioxides are rejected and the remainders recalculated to 100 

per cent the percentages of magnesium carbonate become 12.51 and "12.95, respectively. The 

magnesia, however, shows no such regularity as regards temperature as appears in our series of 

alcyonarians and echinoderms. The difference between the magnesian . content of the two 

specimens of Schizoporella is very striking. What this difference may signify is yet to be determined. 

Several other analyses of Bryozoa, complete or partial, are recorded in the literature. Two 

of the1n, fairly complete, were made by A. Schwager for J. Walther. 27 They are as follows, 

both specimens being from the Bay of Naples: 

] . Eschar a. foliacea. 

2. Lepralia sp. 

SiO:t: ........................... . (AI,lre)20 3 •••••••••••••••••••••••• 

t

1

ag3 .... ." : : : : : : ~ : : : ~ : : ~ ~ ~ ~ : : : : ~ ~ : : ;. 

002 ............. ~ ........... : ... . Organic matter+H20 .............. . 

Sch'Wage1''s analyses of Bryozoa. 

Actual analyses. 

1 

0.29 

. 32 

1. 20 

50.12 

41.06 

6.88 

2 

2.39 

1. 47 

2.22 

47.18 

39. 51 

7. 53 

99.87 100.00 

Reduced analyses. 

1 2 

SiO:t: . . . . . . . . . . . .. . . . . . . . . . . . . . . 0. 31 2. 58 

(Aldj'e)20 3 ••••••••••••••• ,....... • 34 1. 58 

J t~Cg • •• ~ ••• •• ~: ~ : ~ ~ ~ ~ ~ : : : ~ ~ ~ : : ~ ~ ~ : : 9~: ~~ 9g: ~~ 1----1·----

100.00 100.00 

27 Walther, J., Deutsch. gool. Ooscll. Zcitschr., vol. 37, p. 3381 1885. 
36 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. 

The reduced analyses were computed by us from Schwager's data. 

In Flustra foliacea from California H. W. Nichols 28 found 1.23 per cent of magnesium carbonate, and in a bryozoan from Bermuda 5.35 per cent. Several similar determinations by G. 

Forchhammer 29 on other Bryozoa gave Jess than 0.6 per cent. The other constituents of the 

organisms were not determined, and the figures given for magnesium carbonate therefore have 

little present value. . 

From the evidence now at hand no broad general-conclusions can be drawn. That the 

magnesian content of the Bryozoa varies widely, however, is clear, being lowest in the com-' 

pact coralline forms and highest in the fernlike varieties. Even this conclusion needs to be 

verified by a much large;r series of analyses. 

BRACHIOPODS. 

A few analyses of brachiopod shells already on record show that they fall into' two chemi-

·cally distinct groups-one calcareous, the other highly phosphatic: This conclusion is sv.pported and emphasized by the new data .obtained by us, which also bring out some minor 

peculiarities that seem·not to have been previously observed. For our material we are indebted 

to Dr. W. H. Dall, who selected typical specimens from among the duplicates in the United 

States National Museum. First in order come five brachiopods representing as many genera 

in the calcareous group. The analyses are as follows: 

1. Terebratula cubensis Pourtales. Coast of Florida. 

2. Terebratulina septentrionalis Gray. Eastport, Maine. 

3 .. Laqueus californicus Koch. Esteros Bay, Cal] f. 

4. Rhynchonella psittacea Gmelin. Shetland Islands. 

5. Crania anomala Muller. Coast of Norway. 

Analyses of calcareous brachiopods. 

~l?,~~ )~0~ ~ : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : 

~:3.·.:.·.·-~:::::::::::::::::::::::::::::::::::::::::::·:::·:: 803 ••••••••••••••••••••••••••••••••••••••••.••••.••••••• : • • 

P20s- .. ········ ...................... ·······.·· .......... . Loss on ignition ........................................... . 

C02 needed ............................................... . Organic matter, etc ............... , ...................... . 

1 

0.06 

. 04 

. 44 

54.96 

. 21 

Trace. 

44.35 

100.06 

43.42 

.-93 

2. 

0.50 

.14 

. 62 

51. 79 

. 66 

Trace. 

45.28 

98.99' 

40.55 

4. 73 

3 4 5 

0.18 0.14 0.21 . 47 . 23 . 26 . 32 . 23 3.90 

54.48 53.76 48.67 . 21 . 31 . 97 

Trace. .17 . 25 

44.46 44! 81' 45.38 

100.12 99.65 99.64 

42.91 42.16 40.88 

l. 55 2.65 3.52 

Rejecting organic matter and recalculating to 100 per cent, the analyses assume the following rational form: 

Reduced analyses of calcareous brachiopods. 

1 2 3 4_ 

Si02 •••••••••••••••••••••• · .•••••••••••••••••••••••••••••• 

~?,:.~.:-~~~ ~ ~ ~ ~ ~ ~ ~: ~:: ~ ~ ~ ~: ~ ~ ~: ~: ~ ~ ~::: ~ ~ ~::::: ~ ~ ~ ~ ~ ~: ~: 

0.06 0.52 . 04 .15 . 93 l. 37 

98.61 96. 78 . 36 1.18 

0.18 ' 0.15 

. 48 . 23 . 68 . 49 

98.30 98.20 . 36 . 55 

Ca3P20 8 •••••••••••••••••••••••••••••••••••••••••••••••••• Trace. Trace. Trace. . 38 

100. oo 1 100.00 100.00 100.00 

28 Nichols, H. W., Field Columbian Mus. Pub. 111, .p. 31, 1906. 29 Forchhammer, G., Neues Jahrb., 1852, p. 854. 

5 

0.22 

. 27 

8 .. 63 

88.59 

l. 72 

. 57 

100.00 
BRACIDOPODS • 37 

. For comparison the following ·analyses of calcareous brachiopods, made elsewhere, are 

sig.nifican t: 

6. Terebratula sp. Collected by Pourtales between Florida and Cuba; S. P. Sharples, analyst. 30 

7. Terebratulina caput serpentis. Locality not given; F. KunckeU, analyst. 31 

8. Orania anomala MUller. Locality not given; F. Kunckell, analyst. 

9. Waldhei?nia craniu:n1 M tiller. Locality not given; F. Kunckell, analyst. 

10. Waldheimia cranium. Collected by the Norwegian North Sea Expedition, station 255; latitude, 68° 12' N.; 

longltude, 15° 401 ]!}. ; depth, 624 meters; bottom temperature, 6.5° C. 

· 11. Waldheimia cranium. Lofoten Islands. 

Analyses 10 and 11 by L. Sclunelck. 32 

Older analyses of calcareous brachiopods. 

6 7 8 · I 9 10 11 

100.00 100.00 99. 731" 100. 18 100. 49 99.64 

With these analyses ours agree in a broad, general way, although the oldP-r ones vary 

much as regards completeness. Kunckell's analyses, showing free lime and magnesia, are 

suspicious, but only in this ·detail; otherwise they have confirmatory value. All the analyses 

show that brachiopods of this group have shells in which calcium carbonate is the principal 

constituent and that the proportion of organic matter is low. The only aberrant one is Orania, 

which is noteworthy on account of its high percentage of magnesia. In this respect, Kunckell's 

analysis, if recalculated to a common basis, .agrees approximately with ours. Rhynchonella 

is also interesting for the reason that an analysis by I-Iilger of shells supposed to. belong to this 

genus indicates that they are· phosphatic and practically identical in compoaition with those 

of Lingula. The authenticity of Hilger's 1naterial is qu~stionable, and his analysis will not 

be reproduced here. 

Four analyses of shells of phosphatic brachiopods have b.een 1nade by us. As these shells 

contain a large amount of organic 1natter, which possibly varies with the age or 1naturity of 

the animal, we prefer to report our results, as others, before us have done, in proximate rather 

than ultimate form. The analyses are as follo~s: 

1. Lingula anatina Gmelin. Coast of Higo Province, Japan. Organic matter, rejected, 40 per cent. 

2. Lingula anatina. Iloilo, Philippine Islands. Organic matter, rejected, 39.5 per cent. 

3. Discinisca lamellosa Broderip. Coast of Peru. Organic matter, rejected, 25 per cent. 

4. Glottidia (formerly Lingula) JJyramidata Stimpson, coast of North Carolina. Organic matter, about 37 per cent; 

analysis incomplete for lack of sufficient material. 

Analyses of phosphatic brachiopods. 

SiO ................. . (Al,Fe)20 3 ••••••••••• 

MgC03 •••••••••••••• 

CaC03 •••••••••••••••• 

CaS04 ••••••••••••••• 

Ca3P20 8 ••••••••••••• 

........ 

-----

..... 

....... 

....... 

..... 

1 

0.91 

.54 

2.70 

1.18 

2.93 

91.74 

100.00 

2 

0.50 

. 29 

. 79 

4.25 

4.18 

89.99 

100.00 I 

3 4 

0.85 0.49 .. 58 1.16 

6.68 1.71 

8.35 ~?) 8.37 _?) 

75.17 74.73 

100.00 1······ .. ·· 30 Sharples, S. P., Am. Jour. Sci., 3d sor., vol. 1, p. 168, 1871. 32 Schmelck1 L.1 Nor~k~ Non;lhavs Exved.! No. 28, ll· 129, 19Ql, 

31 Kunckoll, F., Jour. _prakt. Chomic, 2d sor.-1 vol. 59, p. 102, 18Q9, 
38 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. 

These analyses are noteworthy on account of the unusual proportion of calcium sulphate 

reported in them. Discinisca is especially remarkable in this respect and also in its percentage 

of magnesium carbonate. Small amounts of sulphates have been found in many mollusks and 

corals as well as in the calcareous brachiopods but in nothing like the proportion given here. 

A new analysis of Discinisca. made upon fresh material is much to be desired. · 

In the older analyses of this group the sulphate seems to have been ignored, or at least 

to have escaped attention. The figures are as follows: 

5. Lingula o11alis. Hawaiian Islands; T. S. Hunt, analyst.33 

6, 7. Lingula ovalis. Locality not given; A. Hilger, analyst.34 

8. Lingula anatina. S. Cloez, analyst;35 recalculated to 100 per cent after rejecting 42.6 per cent of organic matter. 

Older analyses of phosphatic brachiopods. 

·1 SiO, .......................... 5..... , :18 :17 ~ce. I MgC03 .......................... -~. 2. 94 3.13 ......... . I CaC03 • • • • • • • • • • • • • • • • • • • 11. 75 10. 76 10. 86 12. 19 

I 

Ca3P20 8 •••••••••••••••••• 85. 79 84.94 85.24 77.17 

I 

Mg3P20 8 •••••••••••••••••• • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • . • 7. 03 

FeP04 • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 77 . 76 3. 61 

MgO?..... .. .. . . . . .. .. . . . 2. 80 ............................. . 

100. 34 99. 59 100. oo 1 100. oo 

The relatively high figures for calcium carbonate shown in this table are doubtless due to 

the neglect to determine sulphate. The analysis by Cloez differs from the others principally in 

form-that is, in its rp.ode of calculation. If the phosphoric oxide in it is assigned entirely to 

the lime, then the proportion of calcium phosphate becomes 88.6 per cent, which is well in line 

with the other figures. The amount of calcium carbonate would be correspondingly reduced. 

The brachiopods, as ·stated at the beginning of this section, fall into two. distinct groups; 

the shells of one consist mainly of calcium carbonate, with little organic matter, and those of 

the other predominatingly of calcium phosphate, with much organic matt~r. The two groups, 

although they may be alike structurally, are physiologically quite dissimilar, the chemical 

reactions involved in building the shells being of two different orders. Such a distinction 

ought to be significant to biologists, and it is for them to determine what it means. Geologically, 

however, we· can see that the phosphatic brachiopods have probably played some part in the 

formation of phosphatic sediments, a function which is shared by vertebrate animals and some 

crustaceans. 

MOLLUSKS. 

Nun1erous n,nalyses of .molluscan shells have been published, and ·they show rmnarkable 

uniformity of composition. It has nevertheless semned desirable to make a liberal series of 

·new analyses, which are best classified into groups. Sulphates were not determined, except 

in four analyses. 

1. PE~ECYPODS. 

' The pelecypod shells analyzed are as follows: 

1. Astarte crenata Gray. Off Marthas Vineyard, Mass.; depth of water, 668 meters; bottom temperature, 7.2° C. 

2 Callista convexa Say. Vineyard Sound, Mass. · 

3. Macoma sabulosa Spengler. ;Massachusetts Bay; depth, 825 meters; bottom temperature, 5.5° C. 

4. Pt!-cten dislocatus Say. Charlotte Harbor, Fla. 

5. Pecten ventric_osus Sowerby. Head of Concepcion Bay, Lower California. 

6. Venericardia ventricosa Gould. Off Point Conception, southern California; latitude, 34° 251 N.; depth, 447 

meters; bottom temperature, 7.3° C. · · 

7. Cardium s.ubstriatum Conrad. Long Beach, Calif. 

8. Calyptogena pacifica Dall. Clarence Strait, Alaska; latitude, 55° 46' N.; depth, 589 meters; bottom temperature, 5.8° C. · 

9. Nucula expanser Hancock. North of Bering Strait. 

10. Acila mirabilis Adams and Reeve. Japan· Sea, off the coast of Chosen (Korea); depth,. 128 meters; bottom 

temperature, 16° C. · 

11. Placuna orbicularis Retzius. Off Luzon, Philippine Islands. 

aa Logan, W. E., and Hunt, T. S., Am. Jour. Sci., 2d ser., vol. 17, p. 237, 18.54. 

a• Hilger, A., Jour. prakt. Chemie, vol. 102, p. 418, 1867. · 

a& Cloez. S .. Jahresb. Chemie, 1859, p. 642; from L'lnstitut, 1859, p. 24(). 
MOLLUSKS. 

Analyses of pelecypod shells. 

1 2 3 4 5 6 7 8 

-~------- ------------ ---

~1?,1?~)~0~~ -.-.-::::: 

0.25 0.18 0.29 0.31 0.14 0.12 0.11 0.08 . 08 .11 . 22 .08 .15 .08 .09 .04 

~:3.·.·: _.::::: :: ::: : .00 Trace. .00 .46 .34 . 00 Trace . .00 

53.92 53.67 43.77 53.68 54.13 54.16 53.99 53.95 

P20s······· ........ Trace. . 03 Trace. Trace . Trace. Trace. Trace. Trace. 

Ignition ............ 44.79 44.77 44.62 44.34 44.36 44.65 44.85 44.93 

99.04 98.76 98.90 98.87 99.12 99.01 99.04 99.00 

C02 needed ........ 42.37 42.10 42 .. 25 42.69 42.90 42.55 42.41 42.39 

Organic matter, etc. 2.42 2.67 2.37 1. 65 1. 46 2.10 2.44 2.54 

Reduced analyses of pelecypod shells. 

). 2 3 4 5 6 7 8 

------·---

SiO~---····· ....... 0.26 0. 19 0.30 0. 32 0. 14 0. 13 0.11 0.09 

(Al6Fe)20 3 •••••••••• . 09 .12 . 23 . 08 .15 . 08 . 09 .04 

Mg 0 3 ••••••••••••• . 00 Trace. . 00 1. 00 . 73 . 00 Trace. . 00 

CaC03 ••••••••••••• 99. 65 99. 62 99.47 98. 60 98. 98 99. 79 99.80 99. 87 

Ca3P20 8 •••••••••••• Trace. . 07 Trace. Trace. Trace . Trace. Trace. Trace. 

100.00 100.00 100.00 100.00 100.00 100.00 100.0 1100.00 

2. SCAPHOPODS AND AMPHINEUR.ANS. 

Under this heading we have only two analyses to offer, as follows:. 

9 10 

0.33 0.09 

.46 .08 

Trace. .00 

51.18 53.36 . 17 Trace. 

46.67 45.59 

98.81 99.12 

39.86 41.93 

6.81 3.66 

9 10 

---

0.36 0. 10 . 50 . 08 

Trace. . 00 

98. 74 99. 82 . 40 Trace. 

---- 100.00 100.00 

39 

11 

0.0 0 

.08 

. 32 

53.8 

Trace . 

0 

0 

0 

3 

7 

44.1 

98.3 

41.7 

2.3 

11 

---

0.00 

. 08 

. 70 

99.22 

Trace . --

100.0 0 

12. Dentalium solidum Verrill. Off Georges Bank, east of Cape Cod, Mass.; depth of water, 2,361 meters; bottom 

temperature, 4.5° C. 

13. Mopalia muscosa Gould. Santa Barbara, Calif. A chiton. 

Analyses. 

SiO~~- ................... -.- · · · · · · · (AI0

t e)20 3 •••••••••••••••••••••••• 

~fo .".:::: :::. ~::::::::::: : :: :-: : : :: 

S03 ••••••••••••••••••••••••••••••• 

Analyses of scaphopods and amphineurans. 

12 

0:39 

. 26 

. 09 

53. 89 

(?) 

13 

Reduced analyses. 

Si02 •••••••••••••••••••••••••••••• 

(Al,Fe)20 3 ••••••••••••••••••••••• 

2~&:--:-:-:-:::·:·:·:·:::::::::::::::::: 

12 

0.40 

. 27 

. 20 

99. 13 

(?) 

13 

ig~%i~~::: : : : : : : : : : : : : : : : : : : : : : ·: : . Trace. 

44.48 

0.59 

. 21 

. 25 

53.42 

. 20 

Trace. 

44. 74 

Ca3P20 8 ••••••••••••••••••••••••••• 

1------·1-------

Trace. 

0. 61 

.·22 

. 45 

98. 37 

. 35 

Trace. 

C02 needed ....................... . Organic matter, etc .............. , .. 

99. 11 

42.44 

2.04 

99.41 

42.13 

2.61 

3. GASTROPODS. 

14. Purpura lapillus Linne. Eastport, Maine. 

100.00 100.00 

15. Aporrhais occidentalis Beck. Off Marthas Vineyard, Mass.; depth of water, 435.5 meters; bottom temperature, 5.5° C. 

i6. Buccinum undatum Linne. Narragansett Bay, R. I.; depth, 53.3 meters; bottom temperature, go C. 

17. Natica duplicata Say. Cape Lookout, N.C. 

18. Olivia literata Lamarck. Sarasota, Fla. 

Jg. Fasciolaria dis tans Lamarck. Sarasota, Fla. 

20. Crepidula onyx Sowerby. San Pedro, Calif. ... 

21. Antiplanes perversa Gabb. Off Bodega Head, Calif.; latitude, 38° 23' 35" N.; depth, 112.5 meters; bottom 

temperature, go C. 

22. Nassa calijorniana Conrad. Monterey Bay, Calif.; latitude, 36° 47' 50" N.; depth, 67.7 meters; bottom temperature1 11.3° C. 
40 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. 

23. Nassa tegula Reeve. Musgrove Island, Magdalena Bay, Lower California. 

24. Nassa insculpta Carpenter. Cortez Bank, Calif.; latitude, 32° 20' 3011 N.; depth, 165 meters; bottom temperature, 9.5° c. 

25. Turritella gonostorna. Valenciennes. Mulege, Gulf of California. 

26. Volutornetra alaskana· Dall. Off Un~laska, Alaska; latitude, 55° N.; depth, 143 meters; bottom temperature, 4.5° C. · 

27. Pyrolofusus harpa Morch. Unalaska, Alaska. 

28. Plicifusus dirus Reeve. Sitka Harbor, Alaska. 

29. Tachyrhynchus erosa Couthouy. Avatcha Bay, I<amchatka. 

30. Ranella pulchra Gray. China Sea; north of Prabas Island; depth, 256 meters; bottom temperature, 14.2° C. 

31. Cerithiurn aluco Linne. Reef opposite Cebu, Philippine Islands. 

32. Strornbus canariurn Linne. Subig Bay, Luzon, Philippine Islands. 

33. Cavolina longirostris Lesueur. Off Adyagan Island, Philippine Islands; depth, 247 meters. 

Analyses of gastropods. 

14 15 16 17 18 19 20 21 22 23 

------------------

Si02 •••••••••••• ------.---- 0. 14 1. 27 0. 94 0.00 0.05 0. 33 0. 15 0.46 0. 27 2.07 

(Al,Fe) 20 3 .••••• ------------ . 15 -22 . . 41 . 05 . 04 . 04 .11 .11 .12 . 33 

~!8_-_-_-_-_-_-_-_-_·:::::::::::::: . 19 Trace . . 12 Trace. . 00 . 06 Trace . Trace . . 17 -20 

53. 34 52. 84 52. 77 54.83 54.38 53. 87 53. 63 54.06 53. 20 51. 30 

SOa .......... --- .......... (?) . . 08 (?) (?) (?) (?) . 00 (?) (?) (?) 

fg~Ai~~_-_-_-_-_-_-_-_-~·-'_-_.-_::::::: Trace. Trace. Trace. Trace. Trace. Trace. Trace. Trace. Trace. . 08 

45. 33 44.09 44. 49 44. 22 44.46 44. 49 44.40 44. 15 44. 84 44.76 

99. 15 98. 50 98. 73 99. 10 98. 93 98. 79 \ 98. 29 98. 78 98. 60 98. 74 

C02 needed ................ 42. 12 41.52 41.59 43.08 42. 73 42. 39 42. 14 42.48 41.99 40. 37 

Organic matter, etc ....... 3.21 2. 57 2. 90 1.14 1. 73 2. 10 2. 26 1. 67 2. 85 4.39 

. 24 25 26 27 28 29 30 31 32 33 

---

Si02 •••••••••••••• • ••• --- ••• 0.47 0.25 0.07 0.08 0. 16 0. 23 2. 20 0.00 0.08 0. 71 

(Al,Fe) 20 3 .................. . 47. 1. 83 .17 . 08 . 22 . 81 . 61 . 09 . 20 . 39 

~!8::::::::::::::::::::::: . 77 . 20 Trace. Trace. .11 . 44 . 23 Trace. Trace. . 09 

49. 32 52. 57 53. 88 54.23 52. 66 49. 30. 52. 63 54.47 54.04 53. 73 so3·······-------- •••••••• (?) .11 (?) (?) (?) (?) (?) (?) (?) (?) 

fg~Ai~~_-_-_-_-_-_-_-_-_-_-_-_·_~:::::: . 04 Trace. Trace. Trace. Trace. . 36 Trace . Trace . Trace. . 38 

48.00 43. 51 44. 60 44.42 45. 83 47. 60 43. 31 44. 37 44. 31 43.82 -

99. 07 98. 47 98. 72 98. 81 98. 98 98. 74 98. 98 98. 93 98. 63 99. 12 

C02 needed .................. 39. 51 41. 46 42. 33 42. 61 41. 50 38.54 41. 64 42.80 42.46 41.96 

Organic matter, etc ....... - 8. 49 2.05 2. 27 1. 81 4. 33 9.06 1. 67 1. 57 1. 85 1. 86 

Reduced analyses of gastropods. 

14 15 16 17 18 19 20 . 21 22 23 

----

Si02 ••••••••••••••••••••••• 0. 15 1. 32 0.98 0.00 0.05 0.34 0.16 0.48 0. 28 2. 19 

(Al,Fe)20 3 •••••••••••••••••• .16 . 23 . 43 . 05 . 04 . 04 .11 .11 .13 . 35 

~!883 _-_-_-_-_-_-_-_-_-_-::::::::::: . 41 Trace. . 26 Trace. . 00 .14 Trace. Trace. . 37 . 44 

99. 28 98. 30 98. 33 99. 95 99. 91 99.48 -99. 73 99.41 99. 22 96. 84 

CaS04 ••••• ~ ••••••••••••••• (?) .15 (?) (?) (?) (?) . 00 (?) (?), ('!) 

Ca3P20 8 .............. - •• · ••• Trace. Trace. Trace. Trace.

1 

TrlUOe. Trace. Trace. Trace. Trace. .18 

100.00, 100.00 100:00 100. 00 100. 00 100.00 100.00 100.00 100. 00 1100. 00 

24 25 26 27 28 29 30. 31 

I 

32 33 

SiO}' .....••.•............. 0. 52 0.26 0.07 0.08 0. 17 0.26 2.26 0. 00 0.09 0.73 

(AI, e)20 3 ••••••••••••••• --. . 52 1. 89 .18 . 08 .• 23 . 90 . 63 .10 . 21 . 40 

MgC03 ..................... 1. 78 . 44 Trace. Trace. . 24 1.02 . 51 Trace. Trace. . 20 

CaC03 ..................... 97.08 97.21 99. 75 99.84 99. 36 97.00 96. 60 99. 90 99. 70 97. 82 

CaS04 •••••••••••••••••••••• (?) . 20 (?) (?) (?) (?) (?) (?) (?) (?) 

Ca3P20 8 •••••••• ---- • ·- • • • • • . 10 Trace: Trace. Trace. Trace . . 82 Trace. Trace. Trace. . 85 

100.00 100.00 100.00 100.00 100.00 100.00 100.00 100. 00 1100. 00 100.00 
MOLLUSKS. 

4. CEPHALOPODS. 

34. Nautilus pompilius Linn~. Mindanao, Philippine Islands. 

~5. Argoria.utct argo Linn~; the paper nautilus. High seas, Pacific Ocean. 

R6. 8e7Ji't sp.; cuttlefish bone. Port Tataan, Tawi, Tawi group, Philippine Islands. 

An·:tlyses of cephalopods. · 

Si02 ••••••••••••••••••••••••••• 

(Al,.Fe):Pa· .................. . 

~!8. .. · .·.· ... · .·.· ... · ~ ~ : : : : : : : : : : : : : : : rg~Iti~~.-.-.-.-.-.-.-.-.-.·.·. ·.::::::::::: 

C02 needed ................... . Organic matter, etc ............ . 

34 

0.18 

.14 

. 07 

52.44 

Trace. 

46.65 

99.48 

41.28 

5.37 

35 

0.08 

.12 

2.58 

46. 78 

Trace. 

49. 16 

98.72 

42.30 

6.86 

Reduced analyses of cephalopods. 

Si02 ••••••••••••••••••••••••••• 

(Al,F'e) 20 3 •••••••••••••••••••• 

Mg003 •••••••••••••••..•••••••• 

CaC03 ••••••••••••••••••••••••• 

Ca3P208 •••••••••••••••••••••••• 

34 

0.19 

.15 

.16 

99.50 

Trace. 

100.00 

35 

0.09 

.13 

6.02 

93.76 

Trace. 

100.00 

36 

0.00 

.06 

. 66 

47.42 

Trace. 

51.63 

99. 77 

38.65 

2.98 

36 

0.00 

.06 

1. 62 

98.32 , Trace. 

100.00 

41 

For comparison a series of four analyses of cephalopod shells by 0. Butschli 36 is. worth 

citing in full. They are as follows: 

1. Nm.ttilus pompiliW:. 2. Argonauta argo. 3. Spirula pemnii. 4. Sepia officinalis. 

Analyses of cephalopods by BiUschli. 

1 2 3 4 

CaC03 ••••••••••••••••••••• 94.75 85.07 ~fgCO~. .. .. .. .. .. .. .. .. . . . 16 5. 08 

Phosphate................. . . . .. .. .. . 3. 06 

90.43 87.36 . 46 . 21 

3.20 2.26 

CaS04 .2H20............... . 20 1. 75 

H 20...................... 2. 62 2. 92 

Organic matter.. . . .. . .. .. .. 2. 03 2. 43 

Sand ......................................... . 

. 48 .78 

2.17 4.89 

2.32 5.36 . 33 .................... 

. 99. 76 100.31 99. 39 100.86 

Exactly. what is meant by "phosphate" is not clear. Assuming it to mean calcium phosphate and reducing the analyses to uniformity with ours, they take the following form: 

Butschl,i's analyses reduced. 

1· 2 3 4 

CaC03 ••••••••••••••••••••• 99. 66 89. 93 95. 75 96. 58 

1\fgC01 ................... .17 5. 37 . 48 .13 

Ca3P2U 8 ............................ 3. 24 3. 39 2. 50 

CaS04 .................... .17 1. 46 . 38 . 69 

1--------·1---------1--------1--------1 

100.00 100.00 100.00 100.00 

ao BUtschli, 0., K.Gesell. Wiss. Gottingen Abh., No. 3, 1908. Butschli also gives analyses of three other mollusks, Pinna japonica, Pat,ella 

vu.lgala, and Purpura lapillWJ, but they show no unusual features. 
42 THE INORGANIC CONSTI'l;'UENTS OF MARINE INVERTEBRATES. 

The unusual proportion of magnesium carbonate in· Argonauta is confirmed by our analysis, 

but the high phosphate in three of Biitschli's analyses is unlike anything in our series. fossibly 

the calcium sulphate is really present in the hydrated form, that is, as gypsum, but that is 

uncertain. However, it is not necessary to make that assumption, and to do so would complicate the comparison of analyses. 

From the evidence now at hand and from many older data it is clear that molluscan shells 

consist almost entirely of calcium carbonate with quite insignificant impurities. The only 

notable exception is Argonauta, which contains 6 per cent of magnesium carbonate, but the 

paper nautilus has no importance as a contributor to the marine sediments. The mollusks 

generally are. of immense importance, a fact to which their fossil remains abundantly testify. 

In composition they resemble the corals and millepores, so much so that an analysis from one 

group might readily pass for an analysis from anotherY 

CRUSTACEANS. 

The .19 crustaceans analyzed in the course of this investigation fall into two distinct groups. 

First, the barnacles, which have shells composed mainly of calcium carbonate with very little 

organic matter. Second, a group of the more familiar crustaceans, such as crabs, shrimps, and 

lobsters. These have shells containing notable amounts of calcium phosphate and much ( 

organic matter. . The analyses of the first group cover the following species: 

1. Lepas anatijera Linne. Florida. 

2. Lepas ansmfera Linne. Woods Hole, ·Mass. 

3. Mitella polymcrus Sowerby. Pacific Grove, Calif.; latitude, 36° 36' N.; longitude, 121° 55' W. 

4. Balanus harneri Ascanius. Georges Bank, east of Cape Cod, Mass. 

5. Balanus amph1:tritc niveus Darwin. Cape May, N.J. 

6. Balanus eburneus Gould. Smiths Creek, Potomac River, M d. 

7. Scalpellum regium Wyville Thomson and Hoek. _Albatross station 3342; off Queen Charlotte Islands; latitude, 

52° 39' 3011 N.; longitude. 132° 38' W.; depth of water, 2,906 meters; bottom temperature, 1.8° Q. 

The analyses are as follows: 

Analyses of barnacles. 

~l?}~)~o3···-·.: ~ ~ ~: .·.·_·_ ·_ ~ ~:::::::::::::::::::::::::::: 1\f.gO .......•.•............................•......... CaO ................................................ . 

ig~~i~~ .· .· .· .· _· .· .· .-. ·. ~ -~ : : : : : : : : : : : : : : : : : : : : :·: : : : : : : : : : : : 

C02 needed ........................................ " 

Organic, etc ....... :.· ............................... . 

1 

0.04 

.19 

l. 14 

52.33 

Trace. 

44.50 

98.20 

42.37 

2.13 

I 

2 

0.04 

. 56 

. 86 

52.53 . 34 

43.90 

98.23 

41.82 

2.08 

Reduced analyses of barnacles. 

1 2 

100. 00 100. 00 

3 

0.08 

. 30 

. 94 

50.83 

Trace. 

-16.18 

98.33 

40.97 

5.21 

3 

4 

0.03 

.14 

. 35 

53.57 

.00 

<14. 39 

98.48 

42.47 

l. 92 

4 

5 

1. 99 

. 68 

.73 

50.09 

Trace. 

44.84 

98.33 

40.16 

4.68 

5 

2.12 

. 72 

1. 63 

95.53 

Trace. 

100.00 

6 

0.39 

. 22 

. 76 

53.23 

Trace. 

44.28 

98.88 

42.67 

1. 61 

6 

0.40 

. 22 

1. 65 

97.73 

Trace. 

100.00 

7 

} 0.73 

1. 03 

52.65 

Trace. 

4·1-. 79 

} 

99.20 

42.48 

2.31 

7 

0.75 

2.23 

97.02 

Trace. 

100.00 

37 For additional data see Schmelck, L., Norske Nordhavs Exped., No. 28, p. 129, 1901. Fourteen analyses of molluscan shells are given, of 

the genera Buccinum, Astarte, Neptunea, and Pecten, all from the North Sea. The highest percentage of magnesium carbonate foimd was 0.78, 

the lowest 0.26. For ten complete analyses of oyster shells see Chatin, A., and Muntz, A., Com pt. Rend., vol. 120, p. 531, 1895. For eleven analyses 

of land shells see an inaugural dissertation by A. Doring, Gottingen, 1872. Several determinations of magnesium carbonate, maximum 1 per cent, 

are given by G. Forchhammer in Neues Jahrb., 1852, p. 854. 
CRUSTAC~ANS. 43 

Sulphates and soluble salts were not determined. Except for the slightly higher magnesia, 

these analyses closely resemble those of mollusks. 

One analysis of a J:>arnacle, Balanus tintinnabulus, by Butschli, 38 is essentially similar to ours. 

I-Iis figures areCaC03 ..................................... ·. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94. 44 

MgC03 •••••••••••••••••••••••••••••••••••••••••••••••••• ~................................... 1. 20 

Ca3P20 8 ••••••••••••••••••••••••••••••••••••••••••••••••••• ~- •••••••••••••••• _.................. • 36 

CaS04 .2H20................................................................................. 2. 06 

98.06 

In all the analyses calcium carbonate is the main constituent, nnd there are only minor 

impurities. We have found ·no other analysis of a barnacle recorded. 

In the second group of crustaceans the following species were anfllyzed: 

1. Tryphosa pingu1:s Boeck; sand flea. Woods Hole, Mass. 

2. Pagurus rathbuni Benedict; hermit crab. Bering Sea, Albatross station 3531; latitude, 50° 551 N.; longitude, 

174° 171 W.; depth of water, 105 meters; bottom temperature, 1.7° C. 

3. Callinectes sapidus M. J. Rathbun;. blue crab. Ranges from Cape Cod to Florida. 

4. Homarus arnericanus Milne-Edwards; c.ommon lobster. Vineyard Sound, Mass. 

5. Pandalus platyceras Brandt; shrimp. Off Monterey Bay, Calif.; Albatross station 3129; latitude, 36° 39' 4011 

N.; longitude, 122° 01' W.; depth, 49.4 meters; bottom temperature, 6.5° C. 

6. Lithodes maia Linne; spider crab. North Atlantic coast, precise locality of specimen unknown. 

7. Libinia emarginata Leach; spider crab. Vineyard Sound, Mass. 

8. Munida iris Milne-Edwards; long-armed munida. Off Chesapeake Bay; Albatross station 2420; latitude, 37° 

03' 20'' N.; longitude, 74° 3~1 4011 W.; depth, 190.3 meters; bottom temperature·, 8.7° C. 

9. Crago dalli Rathbun;· a shrimp. Off Cape Strogonof, Alaska; Albatross station 3294; latitude, 57°161 4511 N.; 

longitude, 159° 03' 3011 W.; depth, 54.9 meters; bottom temperature, 5° C. 

10. Panulirus argus Latreille; spiny lobster. West Indies. 

11. Chloridella empusa Say; mantis shrim:t>. Ranges from Cape Cod to Flo~!da. 

12. Eriphia sebana (Shaw). Chance Island, Mergui Archipelago; off coast of Tenasserim, Lower Burma. Latitude 

about 10° N. A crab. 

~3. Grapsus grapsus tenuicrustatus (Herbst). Aldabra Islands, Indian Ocean; latitude, 9° 261 S.; longitude,. 46° 

35' E. A crab. 

Analyses 12, 13 by B. Salkover. 

14. Penaeus brasiliensis Latreille; edible shrimp. Ranges from New Yo~k to Brazil. 

A'f!-alyses of crustaceans. 

1 2 3 4 5 6 7 

~]l~ ):~~: ~ ~ ~ ~ : ~ ·; ·;· ;· ;· ~ ~ : ~ ~ ~ : ~ : ~ : : : : : ~ : ~ : : : : : : : : : : : : : 

0.62 0.16 0.04 0.00 0.32 0.00 2.61 . 37 .15 .04 .21 .26 .15 . 78 

1. 27 1. 55 2.06 2.35 1.44 1. 90 2.81 

28.63 29.11 33.42 31.35 18.56 24.15 32.79 

J~~:i~~-:-:::: : :: _: ·'· :_ :_: : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : 

4.56 3.25 4.26 3.04 4.60 3.51 2.73 . 23 . 75 . 23 .45 .54 . 63 .49 

64.73 64.50 58.89 61.82 73.81 69.00 57.07 

100.41 99.47 98.94 '99.22 99.53 99.34 99.28 

C02 needed .................................... _ .. _ . _ . 19.53 21.14 24.43 24.08 11.58 17.46 26.05 

Organic matter, etc .................................. 45.20 43.36 34.46 37.74 62.23 51.54 31.02 

8 9 10 11 12 13 

~ll~ ~~~;:: ~:~ ::: ~ ~ ~ ~ ~ : ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ : ~ ~ : ~ ~ ~ ~ : ~ ~ : ~ ~ ~ ~ ~ ~ : ~ ~ : 

0.32 0.62 0.16 0.02 Trace. Trace. 

.16 .20 .13 .17' 4.82 4.73 

2.58 1. 01 3.15 2.60 1.19 1. 57 

31.28 9.95 25.36 15.38 34.20 25.30 

~6~~:::::::: ~ ·_·_-_._. _.:::::::::::::::::::::::::::::::::::: 1. 87 2.65 2.09 7.75 3.38 2.98 . 47 .47 .41 1.07 Trace. '!'race . Ignition .............................................. 62.29 83.70 66.69 72.35 53.92 64.87 

C02 needed ...................... · ...................... 98.97 98.60 97.99 99.34 97.51 99.45 

25.41 6.20 21.23 7.15 25.04 18.83 

Organic matter, etc .................................... 36.88 77.50 45.46 65.20 28.88 47.79 

18 BUtschli, 0., K. Gesell. Wiss. Gottingen Abh., No.3, 1908. 
44 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. 

In No. 14, the common edible shrimp of the Washington (D. C.) markets, the shell was so 

thin that the available material was too smaii for complete analysis. Only·P20 5, 3.07 per cent, 

was determined. · . Reduced analyses of crustaceans. 

1 2 3 4 5 6 7 

Si02 ••••••••••••••••••••••••••••••••••••••••••••••• 1.12 0.28 0.06 0.00 0.87 0.00 3.82 

~8&!.:-~~-:_:_:_:_:_:_:_:_-:·:·:·_:_:_:_ :_::::::::::::::::::::::::::: 

. 67 . 27 . 06 . 34 . 70 . 31 1.14 

4.84 5.80 6.69 80.2 8.09 8.35 8.65 

74.64 78.03 78.14 79.50 60.94 73.07 76.44 

g:~la?~ -_ ~ -_ -_ ·_ -_ ·_ -_ ~ -_ ~ ~ ~ ~ ~ ~ ~ .~ ~ ~ ~ ~ ~ ~ ~ : ~ ~ ~ ~ ~ ~ : ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ : 18.02 13.55 14.45 10.91 26.94 16.03 8.73 

.71 2.07 . 60 l. 23 2.46 2.24 l. 22 

100.00 100.00 100.00 I 100.00 100.00 100.00 100.00 

8 .9 10 11 12 13 

~J6~}_;~,:·:·:·:·:· ::::::::::::::::::::::::::::::::: 

0.52 2.94 0.31 0.06 Trace. Trace. . 27 . 95 . 25 . 50 7.02 8.86 

8. 71 10.05 12.58 15.99 3.65 6.18 

CaCOa-············ .............................. 82. 64· 54.83 76.87 28.56 78.58 72. 77 

Ca3P20 8 ••••••••• · ••••••••.••••••••••••••••••••••• 6.57 27.44 8.68 49.56 10.75 12.19 

CaS04 ••••••••••••••.••••••••••••••••••••••• ~ •••• l. 29 3. 79 l. 31 5.33 Trace. Trace. 

100.00 100.00 100.00 100.00 100.00 100.00 

In addition to the foregoing analyses two more were made by Mr. Salkover of very minute 

crustaceans, such as form a large part of ·the marine plankton.. Two samples, each rriade up 

of hundreds of individuals, were obtained from 'the U. S. National Museum, as follows: 

1. Temora longicornis (0. F. Muller), from the coast of .New England. Weight of dried sample, 0.6105. gram. 

A copepod. . . - · 

2. Thysanoessa inermis (Kroyer), from Balena, Newfoundland. Weight of dried sample, 1.5973 grams. A small 

shrimp. 

As the amount of material was insufficient for a thorough analysis, only three determina-· 

tions were .made on each sample. They were: Loss on ignition, mainly organic matter and 

water; phosphoric oxide; and residue insoluble in acid. The phosphoric oxide, P 20 5, was recalculated into the form of tricalcic phosphate, Ca3P 20 8 , and with that adjustment the analyses 

assume the following shape: 

Analyses of minute crustru:enns, 

i 

I 

1 2 

-~--1-: 

Loss.on 1gmt1on .......... ~.: .... ·. ~ · .......... 

1 

96.05 ·92 .. 08 

Tricalcic phosphate......................... 2:77 7:68 

Insoluble.................................. 0. fl2 0. 22 

·._ ,--99 ___ 7_4-l--99 ___ 9_8_1 

These analyses show that the inorganic matter of these minute creatures consists almost 

.entirely of calcium phosphate, although more refined analyses on larger quantities of material 

would doubtless' show small percentages of other things. So far, however, it seems that these 

very small.organisms effect what is perhaps a primary concentration of the traces of phosphorus 

that exist in sea water,. and so, as food for the larger animals, they furnish the material from 

which the skeletons of marine vertebrates are built. It is a familiar fact that vertebrate skeletons consist largely, although not exclusively; of calcium phosphate. 
CRUSTACEANS. 45 

With the exception of the barnacles the analyses of crustaceans show that they are definitely 

phosphatic, a fact that was already well known. They are also magnesian, another fact that in 

nearly all previous analyses had either been neglected or overlooked. The magnesia, however, 

shows no such regularity with regard to temperature as has been established by om· analyses of 

alcyonarians and echinoderms. Eriphia, for example, a tropical form, is abnormally low in n1agnesia; whereas Ohloridella, also from warm water, is remarkably high. In other analyses the pro-

. portion of n1agnesium cm~bonate appears. to be about normal, but unfortunately the records for 

son1e specimens are defective as to precise localities and temperatures. Furthermore, the large 

an1ounts of organic matter and water render the Teduced analyses to son1e extent unsatisfactory. 

This is especially true of analysis No. 9, in :which the inorganic portion amounted to only 21 

per cent. In reducing such an analysis the unavoidable analytical errors are multiplied, and 

t.he percentages of lllagnesiunl carbonate, calcium carponate; and calcium phosphate may be 

uncertain by as 1nuch as 1 per cent each. In no.case is the order of magnitude seriously changed, 

hut the n.ccuracy of the figures is not what:we should wish it to be. 

The irregularities in the magnesian content of the crustaceans led to a suspicion that they 

might be partly due to differences in the age or maturity of the specimens that were an~lyzed. 

In order to test this supposition we obtained, through the kindness of Dr. I-I. M. Smith, Director 

uf the United States Bureau of Fisheries, the large claws of two lobsters (Homarus) taken at 

U1e station at Boothbay Harbor, Maine. One specimen was from a small lobster, the other 

from a large one, but the actual size of each lobster was not given. The analyses, by George 

St~igQr, were as follows: 

Actual analyses. 

Small. 

Si02 •••••••••••••••••••••••••••••• } 

(AI, Fe)203 ..................... .. 

6'!3. ·. : ~ : ....... ·. : : ~ : : : ~ : : : : : : : : : : : : : : 

0.11 

l. 68 

30 .. 60 

f~~i~~~:-::: : : : : :: : : :::::::::: : : : • 3:22 

. 45 

63.42 

99.48 

C02 needed ....................... . Organic matter) etc ................ . 22.66 

40. 76 

Analyses of lobster claws. 

} 

Large. 

0 .. 43 

2.90 

23.81 

9.03 

. 60 

62.44 

99.21 

18. 17 

44.27 

Reduced analyses. 

Si02 ••••••••.•••••••••••• : •••••••• } 

(AI, Fe)20a ...................... . }{fg003 ••.••••••.•.•••••••••••••••• 

CaC03 •••••••••••••••••••••.•••••• · 

g:{o~s-·: ~:: ·.·.·.·.·::::::::::::::::: 

Small. 

. 0.19 

6.02 

80.52 

11.98 

l. 29 

100.00 

} 

Large. 

0.81 

ll. 51 

64.37 

21.46 

1. 85 

100.00 

The differences between these two analyses are very striking and confirmed our first 

susplCwn. In order to make the investigation more precise, Dr. Smith had sent us from the 

san1e locality,. Boothbay Harbor, parts of three lobsters; one small, one medium, and one large. 

The actual figures for each entire lobster were as follows: 

1. Small lobster: Length, 8! inches; weight, 10 ounces. 

2. Medium lobster:. Length, 11! inches; weight, 2 pounds. 

3. Large lobster: Length, 16! inches; weight, 5! pounds. 

Further1nore, for each lobster a large claw and pai:t ot the carapace was supplied, so that 

for each ani1nal two analyses could b.e made. This last precaution was taken because it was 

reme1nbered that in two sea urchiQs it was found that the spines and the shell differed in 

co1nposition. Is an analogous difference between two parts of the san1e animal to be found 

n.Inong. the crustacea~s ~ · · · 
I . 

46 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. 

The analyses, by Mr. Steiger, are as follows: 

Analyses of lobster shells. 

1. Small lobster. 2. Medium lobster. 3. Large lobster. 

Claw. Carapace. Claw. Carapace. Claw. Carapace. 

SiOt.: · · ·----------- ·---------- ·--------- ~ · · ·--- ·- ·- · ·} 0 17 0 30 0. 14 0. 24 (Al,.l!'e)20 3 •

7 ••••••••••••••••••••••••••••••••••••••••••• • • 0.18 0.18 

~!8 _·: ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ : ~ : : : : : : : : : : : : : 2~: g~ 2~: ~g 2~: ~~ 2i: ~g 2.35 1.96 

27.04 26.55 

P20 5 .·----······-·····-··---·-·-·······----··.•····-·-- 6. 70 4.07 6.06 4.48 

~~niti~;;.-:: ~:::::::::::::::::::::::::::::::::::::: :::: 64: ~~ 68: i~ 66: ~~ 69: ~g 

3.82 2. 79 

.40 . 37 

63.60 64. 90 

l-------:------l-------·l-------l-------1·------

97.67 98.85 97.37 97.64 

14.39 16.45 13.81 14.03 

55. 19 52. 57 52. 51 55. 46 

002 needed ............ _ .......... __ .... ___ .. _ ..... __ 

Organic matter, etc .................................. . 

97.391 95. 75 

17.48 18.06 

46.12 ! 46.84 

I 

Reduced analyses of lobster shells. 

1 2 3 

Claw. Cara- Claw. Cara- Claw. Carapace. pace. pace. 

SiOC+(Al,Feh03 ••••••• ~ ••••••••••••••••• ____ • ________ 0.33 0.35 0.36 0.66 0.31 0.57 

~!c8:. -_::::: .·. ·. ~:: : : : : : : : : : : : : : ~ : : :: ~ ~ ~ : : :: ~ ~ ~ ~ : ~ : ~ : ~ ~ 10.81 7. 74 11.28 8.12 10.99 8. 77 

72.41 78.98 55.46 70.58 56.89 65.14 

8:~~~~ : : ~ ~ ~ ~ ~ : : : : : : : : : : : : : ~ : : : : : : : : : : : : : : : : : : : ~ : : : : : 15.21 11. 70 30. 78 19.06 29.49 23.20 

1.24 1.23 2.12 1.58 2.32 2.32 

100.00 100.00 100.00 100.00 100.00 100.00 

Here again our suspicions were verified. In each case the large claw is more highly magnesian and phosphatic than the carapace, and the increase in magnesia and phosphorus in· 

passing from youth to age is manifested. This last variation is more clearly shown by averaging 

together the two analyses for each lobster, as follows: 

Average analyses of lobster shells. 

1. Small. 2. Medium. 3. Large. 

9.27 

75.69 

13.45 

' 1. 24 

9. 70 

68.02 

24.92 

1.85 

9.88 

61.01 

26.35 

2.32 

. It is also worth noting that the proportion of calcium sulphate increases regularly, whiCh 

indicates that it is a definite constituent of the lobster shells and not a mere impurity. 

From these variations in the composition of lobster shells, all from a single locality, it 

seems clear that in any future investigation of the same sort relative to crustaceans samples 

of the entire· shells should be analyzed and <;>nly adult specimens should be studied. Only 

under such conditions will it be possible to determine whether regularities exist like those which 

have been observed in other series of organisms. 
CRUSTACEANS. 47 

The older analyses of the shells of crustaceans are more or less unsatisfactory, but they all 

agree as to the phosphatic character of these organisms. C. Schmidt,39 for example, gives three 

analyses, as follows: 

1. A.~tacus .fluviatilis; fresh-water crawfish. 53.27 per cent inorganic. 

2. Lobste.r; probably either Hmnarm vu~garis or Palinurus. 77.06 per cent inorganic. 

3. Sq·uilla. mant1:s: a shrimp. 37.17 per cent inorganic. 

Schm,idt's ana.lys~s of crustaceans. 

1 2 3 

8:o~~~·::.·:.·.·.·:::::::::::::::: 13. 17 12.06 47. 52 

86. 83 87. 94 52.08 

100.00 100.00 100.00 

The figures given here are for the inorganic matter alone. Schmidt mentions magnesium 

phosphate as present but gives no determinations of it. 

Two analyses of crustacean shells by E. Fremy 40 are also worth citing: 

1. "Langouste"; Pa.linurm vulgaris. 

2. '' Ecrevisse"; probably the crawfish, Astacus .fluviatilis. 

FremY_' s analyses ojcrmtacear~,s. 

8:{6~~·::::::::::::::::::::::::::::::::::: Organic matter ............ : .............. . 

1 

6. 7 

49.0 

44. 3 

2 

6. 7 

56. 8 

36.5 

100.0 100.0 

IIere, again, the determination of magnesia has been neglected. If we reject the organic 

matter, the percentages of calcium phosphate in the inorgn,nic part of the shells become 12.03 

and 10.55, respective~y. 

Astac'lts.fluviatilis seems to have received more attention from che1nists than any other crustacean. fl.· Weiske 41 found in fresh shells 4.97 to 5.31 per cent of calcium phosphate and in 

old shells, partly cast off, 9.16 to 9.21 per cent. In "Krebsteine," the concretions found in the 

shells, the percentages ran from 10.73 to 11.28 per cent. The total inorganic matter varied 

between 61 and 67 per cent. 

Astacus has also beeP. studied by Agnes Kelly, 42 who also analyzed a myriapod, Julus 

(lulus). IIer figures are as follows: 

Analyses by Agnes ~elly. 

CaO ...................................... . C02 ••••• · •••••••••••••••••••••• : ••••• ••••••• 

P20s··········· ···························· 

BP Schmidt, C., Annalen Chemio u. Pharm., vol. 54, p. 303, 1845. 

Astacus. 

30.44 

21.23 

2. 79 

.Julus.· 

36. 29; 

21. 60 

3. 37· 

54. 52 . 56. 26 

40 Fro~ny, E., Annales chimie et pi1ys., 3d ser., vol. 43, p. 94, 1855. Fremy also gives many analyses of vertebrate bones, both recent and fosRil. 

41 Wolske, H., Landwirthschaftliche Versuchsstationen; vol. 20, p. 45, 1877. 

42 Kolly, Agnos, Janaisches Zeitschr., vol. 35, p. 429, 1901 

106135-22-4 

(j) 
48 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. 

In an analysis of Astacus fluviatilis by 0. · Biitschli 43 n1agnesia was actuaily determined. 

We append his analysis, together with our own reduction of it: 

BiUschli' s analysis of Astacus. 

Actual analysis. 

CaC03 •••••••••••••••••••• 47. 51 

MgC03 •••••••••••••••••••• l. 38 

Phosphate................ 6. 77 

CaS04 .2H20 . . . . .. .. .. .. .. l. G6 

H 20........ .. . . . ... .. .. . l. 34 

Organic matter... . . . . . . . . 40. 60 

99.26 

Reduced analysis. 

CaC03 • • • • • • • • • • • • • • • • • • • 83. 40 

MgC03 •••••••••••••••••• '. 2. 42 

Ca3P20 8?................. 11. 88 

CaS04 ••••••••••••• ·••••• 2. 30 

100.00 

In this analysis the low percentage of magnesia is very significant. Astacus is a f!·esh-water 

crustacean, whereas the analyses in our series are all of marine forms. In fresh water-the 

average river water-calcium is 6 times as abundant as ·magnesimn, but in ocean water nlagnesium is 3! times as abundant as calcium. This difference in the environtnent 1nay possibly 

explain the difference· between the fluviatile and the marine shells, 2.42 per cent of MgC03 in 

one and 4.84 per cent in the lowest of our determinations. 

One more determination of phosphoric oxide in a crustacean ren1ains to be noted. In the 

shell of a lobster, Homarus vulgaris, W. H. Hudleston 44 found 3.26 per cent of P20 5 • This is 

'equivalent to 7.12 per cent of Ca3P20 8 , or, if the organic inatter was about the same in arnount 

as in our analysis of the A1nerican lobster, 11.44 per cent in the inorganic portion alone. This 

is not far from the figure given in our reduced analysis No.4, nan1ely, 10.91 per cent. 

Although the crustaceans are not of great importance as contributors to the 1narine sedi-

·ments, they are .more important than appears at a casual glance. A crab or lobster sheds its 

shell annually and grows a new one, so that an old individual has contributed many times. A 

single shell might count for little, but when multiplied by a dozen or more the contributions 

Qecome significant. How significant they may be is for zoologists to detennine. · 

CALCAREOUS ALG.lE. 

The calcareous algro are so important as reef builders that, although they are not Inarine 

invertebrates in the ordinary acceptance of the term; it semnecl eminently proper to include 

them in this investigation. In many places they far outrank the corals in importance,. and 

of late years much attention has been p~tid to them.' To Dr. Marshall A. Howe, of the New 

York Botanieal Garden, we are indebted for a fine series of algro, and to hi1n we express our 

thanks. ' · 

For the purposes of this research the calcareous algro fall into two groups. One of these, 

of which Lithothamnium is the most familiar example, is highly n1agnesian; the other, represented by Halimeda, is almost free from magnesia. These groups are best considered ~eparately, and ~nder the first one we have the following species: · 

l. Lithothamnium glaciale Kjellman. Topsail, Conception Bay, Newfotmdland; latitude, 48° N.; longitude, 

53° w. 

2. Lithothamnium erubescens Foslie. Raingsisi, near Timor, East Indian Archipelago. 

3. Archceolithothamnium episporum Howe. Point Toro, near Colon, Isthmus of Panama. 

4. Lithophyll~m craspedium Foslie. Palmyra Island, in the Pacific Ocean, west of south from Haw·aii; latitude, 

5° 491 N. 

5. Lithophyllum pallescens Foslie. Bay of La Paz, Gulf of California; latitude, 24° 161 N. 

6. Lithophyllum dcedaleum Foslie and Howe. Salinas Bay, near Guanica, Porto Rico; latitude, about 18° N. 

7. Lithophyllum antillarum Foslie and Howe. Culebra Island, Porto Rico; latitnde, about 18°.201 N. 

8. Lithophyllum oncodes Heydrich. Coetivy Island, in the Indian Ocean, northeast of Madagascar; latitude, 7° 

61 S.; longitude, 56° 301 E. 

9. Lithophyllum intermedium Foslie. Fort Clarence, near Kingston, Jamaica. 

10. Lithopliyllum pachydermum Foslie. Dollar Harbor, South Cat Cay, Bahamas; latitude, about 25° N. 

•3 Htitschli, .0., K. Gesell. Wiss. Gottingen Abh., No.3, 1908. H Hudleston, W. H., Geol. Soc. Quart. Jour., vol. 31, p. 376, 1875. 
CALCAREOUS ALGlE. 49 

11. Lithophyllum tamiense fleydrich. New Guinea. 

12. Aniphiroajragilissinta Lamourotix. Lemon Bay, near Guanica, Porto Rico. 

13. Phyniatolithon cmnpactum Foslie: Torbay, Newfoundland. 

:t.<!. Goniol-itlwn acrqpetum, Foslie and Ho.we. Culebra Island, Porto Rico. 

15. Goriiolithon strictum Foslie. Bernini Harbor, Bahamas. 

16. Goniolithon strictum,. Soldiers Key, Fla .. 

The actual analyses are as follows: 

Analyses of algm. 

1 2 3 4 5 6 7 8 

-------------------'-------·1---· ----------------------

98. 97. 

002 needed· ................................. 

1 

40. 86 

Organic matt~r. etc......................... 7.14 

9 10 

---

Si02 ••••••••••••••••••••••••••••••••••••••• 0.27 0.04 

(A1,.Fo) 20 3 •••••••••••••••••••••••••••••••••• . 21 . 08 

~~f8.-: .·: ~ : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : 7.25 6.47 

42.55 42.60 

P20 11 ••••••••••••••••••••••••••••••••••••••• Trace. .00 

S03 ••••••••••••••••••••••••••••••••••••••• .02 . 59 

Ignition .................................... 48.63 49:55 

98.93 99.33 

C02 needed ................................ 11.89 40.24 

Organic matter, etc .. ·: ..................... 6. 74 9.31 

l. 47 

.-89 

5. 93 

44.80 

Trace. . 53 

45.69 

99.31 

41.43 

4.26· 

11 

---

0.12 

. i6 

8.81 

41.07 

Trace~-

. 69 

48. 77 

99.62 

41.58 

7.19 

0.03 

. 09 

8.64. 

41.56 

.03 

.15 

48.37 

98.87 

42.04 

6.33 

12 

---

2.82 

l. 31 

6. 71 

34.82 

Trace. . 56 

48. 78 

95.00 

34.43 

14.35 

l. 01 0.18 0.04 O.OB . 22 . 08 .10 . 12 

.6.42 8. 1.4 7.24 8.07 

40.39 40.48 43.34 42.57 . 11 Trace. Trace . . 08 . .71 . 44 . 57 . 27 

49.94 49.99 48. ll 4-7.96 

98.80 99.31 99.40 99.1(; 

38.31 40.51. 41.70 42. 1.l 

l.l. 63 9.48 6.41 5.85 

1.3 14 15 16 

------------

0.10 0.06 0.02 } 0.05 . 23 .04 . 01 

4. 02 8.27 10.39 10.n3 

38.19 40.60 38.54 38.03 

.10 .18 Trace. Trace . . 52 . 62 . 60 . 61 

55.42 49.56 49.91 49.91 

98.58 99.33 91-l. ·17 99.53 

35.05 40.50 41.38 41.56 

20.37 9.06 8.53 8.35 

The low smn1nation' of No. ~2, a very fragile form, is due to inclosed salts, mainly alkaline 

chlorides and sulphates. These ~ere detennined as 5.40 per cent, which brings the summation 

up to 100.40. 

Reducing the analyses, as usual, to standard .form, we have the .following table:_ 

Reduced analyses of algm. 

1 I 2 i 3 4 5 (I 6 7 8 

------ ---

Si02 •••••••••••••••••••••••••••••• · ••••••••• 0.45 0.20 ]. 55 0.03 1.16 0.20 0~04 0. 10 

(A1,.I"e)20 9 •••••••••••••••••••••••••••••••••• . 25 . 1.3 . 94 .10 . 25 . 09 .ll .13 

~~~;-:::: ~: ~ ~ ~ ~ ~ ~ ~ : ~ ~ ~ : :: : : : : : : : : : ~ : : : : : : : 

10.93 16.96 l3 .. ·Q9 19.60 15.46 .1·9. 03 1.6.·35 18.17 

88.ll 81. fi9 83.-47 79.92 81.48 79.85 82.46 80.93 

Trace. Trace. Trac~i .08 . 26 Trace. Trace. . 18 . 26 1..12 . 95 . 27 1.. 39 ! . 83 l. 04 I .49 

1.00.00 100.00 100.00 100.00 100.d0 I 100.00 100.00 I 100.00 

9 10 11 12 13 14 15 i6 .. 

---· ------------

Si01 .................................. .' ... 0.30 0.04 0.13 3.50 0.13 0.07 0.02 } 0.06 (Al, ~0):.~0 3 •••••••••••••••••••••••••••••••••• . 23 . 09 .17 1. 62 . 30 .05 . 01 

~f!,~;-:_~~ ~ ~: : : : : : : ~ : : : : : : : : ~ :·: ~ ~ ~ : : : : : : ~ ~ : 

16.59 15.08 20.02 17.47 10.93 19.24 .24.00 25.17 

82.85 83.68 78.43 76.23 87.21 79. 05. 74.85 73.63 

Trace. '!'race. Trace. Trace. . 29 .43 Trace. Tr·ace. 

.03 - 1.11 1. 25 1.18 1. 14 1.16 1.12 1.14 ·--- 100. oo 1 100.00 100.00 100.00 100.00 100.00 100.00 ·100.00 

·' 
50 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. 

In No. 12, Amphiro~, the high silica and sesquioxides are evidently impurities. If they 

are rejected, the percentage of Inagnesium carbonate rises to 18.41. 

Of Halimeda, four species were analyz~d, as· follows: 

17. Halimeda opuntia Lamouroux. Key West, Fla.· 

18. Halimeda simulans Howe. East of Guanica Hn.rbor, Porto Rico. 

19. Halimeda tridens Lamouroux. Cayo Maria Langa, Bay of Guayanilla, Porto Rico. 

20. Halim,eda monile Lamouroux. Same locality as No. 19. 

Analyses of Halimeda. 

Si02 ••••••••• ~ ••••••••••• 

(Al,Fe) 20 3 ............... . ·MgO .................... . 

CaO .. -·----·-··------·-- Pz05-- .. : .•.... --·--- ... -·- S03 ________________ . ---- .. Ignition. ____ .,_. ___ .. ___ , 

17 

0.37 

. 20 . .01 

50.32 

Trace. . 07 

48.20 

18 

0.44 

.. 18 

.19 

50.20 

Trace. . 23 

48.38. 

19 

0.88 

. 58 

. 44 

45.32 

Trace. . 49 

51.94 

I 

20 

l. 90 

1.10 

. 45 

48.21 

Trace. . 02 

47.56 1-~--1--------··-----

002 needed ...... _ . _ ... _ . Organic matter, etc ..... __ 

·99.17 

39.47 

8.73 

99.62 

38.53 

8.85 

Reduced analyses of Halimeda. 

17 18 

Si02 : •••••••••••••••••• -.- 0.41 0.48 

(Al,Feh03 .. -............. - . 22 . 20 

MgC03 .. - . -. -.... - -. - -... .02 .44 

Ca003 ......... : ......... 99.21 98.45 

Ca3P 20 8 •••••••.••.•••••.. Trace. ··Trace. 

CaS04 ••••••••••••••••••• _· .14 . 43 

100.00 100.00 

99.65 

35.83 

16.]1 

19 

l. 04 

. 68 

l. 09 

96.21 

Trace. 

.98 

100.00 

99.24 

38.35 

9.21 

20 

2.11 

l. 22 

]. 04 

95.58 

Trace. 

.05 

100.00 

After allowance is made for obvious impurities the essential identity of these ·four analyses 

becomes clear. The inorganic portion of Halim(3da consists of calcium carbonate and an insignificant amount of magnesia. In the first group of alg::e, Nos. 1 to 16, magnesium carbonate 

is conspicuou~, more so than in ·any other class ~f organisms so far analyzed. In Goniolithon 

strictum especially it ranges from 24 to over 25 per cent, or more tha~ halfway to dolomite. 

These alg::e are probably the largest contributors of magnesia to the marine limestones. 

It is also ~oteworthy that the two alg::e of the first group, Nos. 1 and 13, are from the cold 

waters of N ewfoundlan·d, and that the others are from tropical or subtropical regions. Several, 

if not all of them, are shoal-water organisms and. were collected on reefs, or rocks, or on sands 

near low.,..tide lev~ls. It is de.sirable that Arctic species shoul<;l be carefully studied-at least 

more carefully than hitherto. · The older published data leave much to be desired, especially 

as to definiteness of species and--localities. 

In addition. to the material received from Dr. Howe, three more calcareous alg::e were 

submitted for an~lysis by T. Wayland Vaughan. The species, with analyses by A. A. Chambers, 

are as follow.s: 

I. Goniolithon .frqtescens 1 Foslie. Cocos-K:e~ing. Islands, in the Indian Ocean; latitude, 12.5° S.; longitude, 

96.53° E . . : ~~ 

2. Goniolithon: orthoblastum (Heydrich) Howe; Murray Island, Torres Straits, Australia. 

3. Lithophyllum kaiseri Heydrich. Co~os-Ke~l~ng r.sla~ds. 
CALCAHEOUS ALGlE. 

Analyses of algx. 

Si02,Al20 3 )i'e20 3 •••••••••••••• 

~1!3::: ....... · ...... : ~ : : : : : : : : : : : : : : : ~0~~:.::: ~ · .. ·:::::::::::::::::::: 

Igmtwn ....................... . 

C02 needed ................... . Organic matter,· etc ............ . 

1 

0.07 

6. 29 

46. 16 

Trace. . 00 

46. 70 

99. 22 

43. 19 

3. 51 

Reduced analyses of algce. . 

2 

0. 11 

5. 71 

42. 39 

Trace. . 00 

50.97 

99. 18 

39. 59 

11. 38 

3 

0. 28 

7.09 

45. 92 

Trace . . 00 

45. 72 

99.01 

43. 88 

1. 84 

------·-·-- --- ·- ---- ---·--------;-·-----:-----------, 

1 

0.07 

13. 80 

86. 13 

Trace. . 00 

100.00 

2 

0. 12 

13. 66 

86. 22 

Trace. . 00 

100.00 

3 

0. 29 

15. 33 

84. 38 

Trace . .00 

100.00 

51 

These algre are much lower in· magnesia than our sp~cimens from similarly warm regions. 

The two species of Goniolithon contain little more than half as much magnesium carbonate as 

the species from Florida and the West Indies. That this difference might be due to differences 

of age or 1naturity seen1ed to be possible, and therefore two more algre were obtained from 

Dr. I-Iowe and analyzed by R. M. ICamn1 with the following results: 

l. Litho7Jhyll-wn pachydennunt, Dollar Harbor, South Cat Bay, Bahamas. 

la. Upper or younger layer, solid. · 

lb. Lower or older layer, porous and somewhat discolored. 

2. Goniolithon strictum, Ben~ini Harbor, Bahamas. 

2a. Upper or younger branches. 

2b. Lower or olcH.lr branches. 

Act·u.al ar:alyses of algc:e. 

1a 1b 

Si02 •••••• : •••••••••••••••• 0.04 0.09 

(Al,Feh03· •••••••••••••••• .11 .14 ~1g0 ....................... 10. 69 6.54 

CaO ....................... 37.66 44. 88 

P205······················· Trace. Trace. 

·S03 ••••••••••••••••••••••• . 79 .71 

Ignition .................. 50.64 47. 51 

99. 93 99. 87 

CO., needed ................ 41.42 42.08 

Organic, etc ................ 8. 72 5.43 

Reduced analyses of algm. 

1a 1b 

Si02 ....................... 0.04 0.09 

(Al, F'e):Pa- ............... .12 .15 

:NfgC03 ••••••••••••••••••••• 24. 95 15. 43 

CaC03 •••••••••••••••• " •••• 73. 65 83.06 

8~8~~~---.---.-:~:~---------:::::: Trace. Trace. 1. 24 1. 27 

100.00 100.00 

2a 

0.02 

. 32 

9.64 

37. 64 

. 00 

. 63 

51. 14 

99. 39 

39. 83 

11. 31 

2a 

0.02 

. 36 

22.98 

75.42 

. 00 

1. 22 

100.00 

2b 

0.08 

. 57 

10. 26 

38. 25 

. 00 

. 67 

48. 76 

98. 59 

40. 97 

7. 79 

2b 

0.09 

. 63 

23. 74 

74. 29 

. 00 

1.25 

I 100.00 

I 
52 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. 

From these analyses no positive conclusions can be drawn. In No. 1 the older layer is 

much impoverished in magnesium carbonate, and its porous character indicates an alteration 

by leaching. In No. 2 the difference is very slight, but the older branches are a little richer in 

magnesia than the younger ones. That differences of composition may exist in different parts 

of the same specimen is, however, manifested. 

To Prof .. L. R. Cary, of Princeton University, we are indebted for two unpublished analyses 

of calcareous algre by Prof. A. H. Phillips. In one, from Samoa, he found 36.36 per cent of 

magnesium carbon_ate, an astonishingly high figure. In tlw other, from southern Florida 

or the Tortugas he found, in perce~tages, CaC03 , 73.23; MgC03 , 25.32; and Ca3P20 8 , 0.35; 

total, 98·.90. This analysis agrees well with ours of Goniolithon from the same region. · 

Of the earlier analyses of calcareous · algre those by A. Damour 45 are the most complete 

and most nearly comparable with ours. Five species were analyzed, as follows: 

1. Lithophyllum sp. Mediterranean Sea. 

2. Melobesia sp. Coast of Algeria. 

3. Amphiroa tribulus. Antilles. · 

4. Halimeda opuntia. Red Sea. 

5. Galaxaurafragilis. Antilles. 

Damour's " Millepora cervicornis," of which he gives an analysis, is not included here on 

account of its doubtful character~ (See p. 15.) 

Damour's analyses of algre. 

1 2 3 4 5 

·-----·--·--···-------------------·l----1----l----l----l----

Ca003 • _-.--.-............................................. 77. 36 72. 78 70. 83 86. 17 72. 56 

Mg003 ........................................... ·.......... 11.32 12.32 16.99 .56 .86 

Na20 ....................................................... 55 1.75 .89 1.13 .73 

~~~3-- ~: ~: .-_-_-_ -_:: ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~:::: ~ ~ ~ ~:::: ~:::::::::::::::: :·a~ : ~g ______ ·_ ~~ ______ . -_ ~~ ___ . ___ ~-- ~~ S03 ..... ---- .... -- ........... ---- ..... -. . .... -.--.-.·----- . 95 1. 25 . 93 (?) (? 

P20 5 ••••••••••••••••••••••••••••••••••••••••••••••••••••••• • ?2 . 38 . 27 (?) (?) 

01. - - --. - - - -- -- . -.. --.. - -... - - -.. -....•.............. -. -. . . .60 . 34 . 53 . 84 1. 17 . CaS04 •••••••••••••••.•••••••••••••••••• :...................................... .20 .55 1.80 

Organic matter............................................ 4:70 · 3. 95 6. 40 8. 30 17. 50 

H 20. .. .. .. .. .. .. .. .. .. .. . .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. 1. 46 1. 40 . 1. ·38 .· 90 . 95 

Si02 •••••••••••••••••••••••••••••••• ·••••••••••••••••••••••• • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 2. 20 

Sand. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. 36 4. 28 . . . . . . . . . . . ..... ·. . . . . ........ . 

98. 97 99.30 98.81 98.99 98. 79 

In these analyses Na20, K20, Cl, and SO~ evidently represent sea salts. R~jecting them 

as impurities, together with the organic matter, water, and sand, and recalculating to 100 per 

cent, we have the following reduced analyses, which are similar to ours: . 

Damour's analyses reduced. 

1 2 3 4 5 

~~~o;.·. -_-_._._._._._ ·_::: ~ ~ ~: ~ ~ ~ ~ ~: ~ ~ ~:::: :·::: :::::: :: :':::: : ::::: : · · · · · o: o9 · · · · · · o: 23 · :::::::::: :: :::::: : : ...... ~-- ~: Mg0u3 ........ : ........................... ~'--.............. 12. 76 14.44 19. 29 0. 64 1.11 

Ca003 ••••••••• • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 86. 36 84. 36 79. 81 9_8. 73 93. 73 

Ca,iP20 8.................................................... _.79 .97 .67 (?) (?) 

CaSO 4 : ••••••••••••••••••••••• • ••••••••••••••••••••••••••• ·• • • • • • • • • • • • • • • • • • • • • • 23 . 63 2. 32 

100.00 100.00 1.00.00 100.00 100.00 

.,, Damour, A., Compt. Rend., vol. 32, p. 253, 1851. 
CALCAREOUS ALGJE. 53 

Three of these algrn are highly magnesi~n; Halimeda and Galaxaura are not. 

Two analyses of algrn by A. Schwager were published by J. Walther 46 iri 1885. ·There is 

also one by C. W. Glin1bel, 47 and these three may be combined in one table, as follows: 

l. Lithothmnni'wn sp. Bay of Naples; A. Schwager, analyst. 

2. Lithothamnium 1·antulostt?n. Bay of Naples; A. Schwager, analyst. 

a. Lithothamnittm nodosum. Locality not stated; C. W. Gi.imbel, analyst. 

Actual analyses of algre. 

1 2 3 

~~k.:::_:;::_:_:::::: :::::::::: qi q} ) ..... -~.-~~-. J\{n0. .... ... . . .. .. . . . . . . . . . . . . . Trace. Trace. 

~~3:.·.·.·.·.·:::: ~:::::::::::::::: 4~: ~~ 4~: ~~ 4~: ~~ C02 ............................ 39. 87 39. 41 40. 06 

P20 5••••••••••••••••••••••••••• • • • • • • • • • • • • • • • • • • • • • • • • • 06 

Organic matter+ H20. . . . . . . . . . 5. 06 5. 57 ........... . Insoluble .............................................. . H 20 and loss .................. · ............ · ........... . 

100.15 99.85 

4.96 

2.57 

1oo.oo· 

. Reducing these to uniformity with our analyses we have the following table: 

Red'l.f-ced analyses of algre. 

1 2 

SiO,........................... l. 68 · 2. 03 

(Al,Fe)20 3•••••••••••••••••••••• 3. 83 4. 26 · MgC03 •• ••••• •• ••• • • • • • • • • • • • • • 4.19 6. 81 

CaC03 ••••••••••••••••••••••••• 90. 30 86. 90 

Ca3P20 8 ••••••••••. • ••••••••••••••••••••••••••••••••••••• 

100.00 100.00 

3 

2.76 

6.06 

91.04 

.14 

100.00 

In these an11,lyses, all of Lithothamnium, the percentage of magnesium carbonate is remarkably low. Why this should be so is not clear, but it may be due· to differences in the age of 

the plants; that is, young specimens n1ay have secreted less magnesia proportionally to lime 

than older, more mature examples. This suggestion might well be tested by special analyses 

of properly selected material. Until that has been done the suggestion can carry little weight. 

Eleven partial analyses of Lithothamnium, made by A. G. Hogbom,48 assisted by R. Mauzelius, N. Sahlbom, and J. Guinchard, are curiously irregular. The percentages of magnesium 

carbonate vary enormously, .and with no apparent or even probable relation to temperatures. 

'rhe data given are as follows: 

CaC03 • 

Lithothamnium sp. Java Sea ............... Lithothamnium sp. Galapagos Islands ....... Lithothamnium sp. Spitzbergen ............. Lithothamnium polymorphum. Kattegat ..... Lithothamnium sp. Honolulu ............... Lithothamnium ramulosum. Naples ......... Lithothamnium sorijerum. Arctic Ocean ..... Lithothamnium sp. Bering Islands .... , ...... Lithothamnium racemus. Naples ............ Lithothamnium sp. Bermuda ............... Lithothamnium glaciale. Arctic Ocean ....... 

iO Wnlthor, J., Deutsche. goo!. Oosoll. Zoitschr., p. 338, 1885. 

i7 OUmbol, C. W., K. bayor. Akad. Wiss. Abh., ?>fnth.:phys. Classo, vol. 11, p. 26, 1871. 

iB Hogbom, A. 0., Nones .lahrb., 1894, Band I, p. 262. 

72.03 

83. 60 

84.83 

74. 22 

84.01 

63.00 

80. 90 

74. 24 

77.39 

82.44 

83.10 

MgC03• 

3.76 

6.53 

8.67 

9.10 

9.39 

9.46 

9.56 

.9.94 

]1. 33 

12.37 I 

13.19 
54 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. 

In this series the percentages of magnesium carbonate are much lower than those found by 

us, and there is uncertainty in six examples as to the exact species. Hogbom, however, points 

out the importance of these algre in respect to the formation of dolomite, which was the real 

subject of his investigation. In Ha~imeda sp., from Labuan, only a trace of magnesia was ~ound. 

In Lithothamnium racemus, from the Bahamas, H. W. Nichols 49 found 5.35 per cent of 

magnesium carbonate, but without rejection of organic matter, etc., undetermined. This 

percentage is remarkably low. A similar low figure for magnesium carbonate-5.85 per centwas found by E. W. Skeats 50 in Lithothamnium phillipi var. funafutiensis Foslie. The speci- 0 

men was taken on the atoll of Funafuti. 

In a very fresh Halimeda opuntia Skeats found 0.60 per cent of MgC03 and 86.50 of CaC03 , 

but in a mass of fronds dredged up from a depth between 50 and 60 fathoms the percentages 

were 4.0 MgC03 to 93.59 CaC03 • The increase in magnesia may have been due to concentration by leaching .. These data are given by J. W. Judd,S1 who also cites an analysis of dead 

Halimeda fronds made for him by C. G. Cullis, who found in them 1.39 per cent of MgC03 and 

98.32 of CaC03 • Judd also quotes an analysis of Halimeda, cited _by Payen in his Flora, which 

'contained 5.50 per cent of MgC03 and 90.16 of CaC03 • The character of the specimen represented by the last analysis, whether fresh or old, is uncertain. The best analyses agree In 

assigning little magnesium to Halimeda. 

In an important memoir on Melobesia Madame P. Lemoine 52 gives several analyses of 

calcareous algre. The analyses, made for her by M. Chari, are not very complete, but they 

are, nevertheless, of distinct value. The data are as follows: 

1. Lithothamnium calcareum. St.-Vaast-la-Hougue, Manche, France. 

2. Lithothamnium calcareum. Isle Glenan, Finistere, France. 

3. Lithothamniumjornicatum. Norway. 

4. Lithophyllum incrustans. Gatteville, Manche, France. 

5. Lithophyllum incrustans. Mazagan, Morocco. 

6. Lithophyllum tortuosum. Genoa. 

7. Lithophyllum craspedium. Tahiti. 

To the analyses as reproduced in the following table we venture to add a line for reduced 

or corrected magnesium carbonate, computed on the basis of 100 per cent for the two carbonates alone. 

Lemoine's analyses of alga;. 

1 2 3 4 5 6 7 

---------

Soluble 'in H20 ............ ~ ............... · ... -...... 2.3 3.0 1.2 1.7 3.8 5. 7 3.2 

Insoluble in HCl ................................... .0 . 0 .0 . 0 . 0 

6. 21 .0 

~~~~-~~i~;: : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : : 

80.9 75.2 81. 6 84.4 77.7 73.~ 75.4 

11.8 10. 7 9.3 10.8 9.8 9.3 15.9 

5.0 11.1 7.9 3.1 8. 7 5.4 5.5 

100.0 100.0 I 100.0 

I 

100.0 

I 

100.0 100.0 

I 

100.0 

MgC03 corrected .................................... _ 12. 7 12.5 10.2 11. 3 11.2 11.2 17.4 

In this series the highest magnesia is in the alga from Tahiti and the lowest in that from 

Norway. ·This tendency toward 'increased magnesia in algre from warm regions, as cbmpared 

with those from cold waters, wfl:s noticed by Madame Lemoine but only incidentally. The 

subject was not given any detailed consideration by her. 

Madame Lemoine 'also cites the. older analyses of algre, including three by J. Chalon, as 

follows: 

1. Lithothamnium calcareum. Roscoff, Finistere, France. 

2. Lithophy-llum incrustans. Banyuls, France, on the Mediterranean. 

3. Lithophyllum tortuosum. Naples. 

t9 Nichols, H. W., Field Columbian Mus. Pub. 111, p. 31, 1906. 

r,o Skeats, E. W., The atoll of Funafuti, pp. 376, 377, London, The Royal Society, 1904. 

61 Judd, J. W., idem. 

t~Lemoine, P ., Inst. oceanographique Monaco Annales, vol. 2, fasc. 2, 1911. 
GENERAL DISCUSSION. 55 

Chalon's analyses of algre. 

' 

~{~~?;1·::::::::. ·. ~:: : : : : : : : : : : : : . 82. 4i. 76.06 82.20 

11.80 14.38 11.57 

Organic matter .... · ............ 4.30 7. 38. 5.26 

Water ............ · .............. . 86 1. 72 . 80 

99.37 '99.54 

I 

99.83 

MgC03 corrected ................ 12.52 15.90 12.35 

These recent analyses, fr01n the 1nemoir of Madame Lemoine, agree well with ours and, 

notwithstanding IIogb01n's div-ergent data, strengthen the suggestion that the proportion of 

. magnesia in the algrn is influenced by temperature. · 

The material studied by us was carefully chosen by Dr. IIowe, with direct reference to the 

pul'pose of our investigation. Every species was thoroughly identified, its locality was definitely 

stated, and the specimens were remarkably clean aud free from misleading impurities. The . 

results obtained by us are therefore as nearly trustworthy as it is practicable for us to make 

then1. The significance of the algrn in reference to dolomite was already well established by 

pr.evwus workers, but our Jlew data strengthen the conclusions which our predecessors had 

drawn. 53 

GENERAL DISCUSSION. 

In the foregoing pages we have reported 322 new analyses oi marine invertebrates and have . 

cited many other analyses mn,de elsewhere. These data shed much light upon the chemistry 

of the marine sedin1ents, and they also suggest various problems, some of them biological, 

which are yet to be solved. The limitations of our research have been pointed out in the introduction to this mCinoir and are taken for granted in the following general discussion of the 

results that we have obtained. First in order we may consider the disti·ibution of the essential 

constituents of the invertebrate skeletons, taking each one separately. 

SiZica.-The skeletons of radiolarians and diatoms and the spicules of siliceous sponges 

consist almost entirely of opaline silica. The radiolarian and diatom oozes of the Challenger 

expedition show the importance of these organisms. In our own work we have studied only 

the sponges, and our results show nothing new. Our analysis of Euplectella, however, is 

probably more complete than any previous analysis of a siliceous sponge. We have found 

recorded in the literature only partial analyses of sponge. spicules. 

In nearly all our analyses, in every group of organisms, silica appears, generally in small 

but exceptionally in rather large proportions. Some of this may be essential, but in most cases 

it is an impurity. In fact, sand grains were distinctly visible in some of the specimens analyzed, 

but were not readily removable. · 

Alumina and iron oxide.-In most of our analyses alumina and iron oxide were usually 

determined, but they are to be regarded generally as impurities due to adherent silt or mud. 

Iron is doubtless a normal constituent in small amounts. 

Lime.-The most in1portant base in nearly all marine shells or skeletons, whether vertebrate or invertebrate, is lime. Only the siliceous organisms are free from it. Molluscan 

sh.ells, the stony corals, the hard parts of millepores, some brachiopods, and the barnacles are 

composed almost entirely of calcium carbonate and contain only minor impurities. In the 

other series of 1narine invertebrates, with few exceptions, it is the dominant inorganic constituent. Calcium phosphate and sulphate were also determined in most of our analyses, but 

they will be considered in other paragraphs. 

Magnesia.-One of the most interesting results of our investigation is the discovery that 

magnesium carbonate is much more widely distributed as an essential constituent of marine 

invertebrates than it has hitherto been supposed to be. In the Foraminifera, alcyonarians, 

echinoderms, crustaceans, and coralline algrn it is especially important, and some other organisms contain it in notable proportions. Its 'peculiar relations to temperatur~ have been noted 

in several sections of tllis work and will be discussed more fully later. 

r.a On tho importance of algm as reef builders·, see an interesting paper by Dr. Howe in Science, new ser., vol. 35, p. 837, 1912. He cites much 

other literatme. 
56 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. 

Our determinations of magnesia, however, are subject to at least one small correction. 

Many of the specimens analyzed contained inclosed or adherent sea salts, and in a few of them 

they could not be estimated. They rarely amounted to more than 2 per cent, but in one analysis 

5 per emit was found .. Sea salts pontain magnesium, and its equivalent in magnesium carbonate· 

must therefore be deducted from the percentages of.magnesium carbonate given in our reduced 

analyses: The maximum correction to be thus applied is about 0.4 per cent, but 0.1 per cent 

would be the more common amount. In our work such a correction is negligible, for the proportion of magnesium carbonate in our important magnesian series ranges from 5 to 25 per 

cent. The small quantities of magnesia found iri most mollusks and corals, however, may be 

due in part-, if not entirely, to saline impurities. . 

Phosphorus.-In nearly all our analyses phosphoric oxide appears, but generally in trifling 

quantities. It is abundant, however, in the series of phosphatic brachiopods, the crustaceans,' and the alcyonarians. Some worm tubes also are notably phosphatic. In reducing the 

analyses ·to standard form we have assumed that the phosphoric oxide is best represented 

in combination as tricalcium phosphate, although the assumption is not absolutely proved. 

It is a pure convention, adopted for the sake of uniformity and to simplify the comparison 

of analyses. It is of course possible that magnesium phosphates may exist in some of the 

organisms and that a part of the phosphorus may be contained in their organic matter .. Magnesium phosphates, ho'Yever, are very rare as minerals, whereas calcium phosphate is 

extremely common. The organic matter decomposes after the death of the animals, and its 

phosphorus would doubtless appear !n the se.;liments as a phosphate. In any case the dead 

organisms are likely to be buried among calcareous sediments, where calcium phosphate should 

· i>e formed. Even the phosphatic worm tubes, in which the calcium is insufficient to form a 

·tribasic salt, would probably ~ollow this rule. Lime from the sediments would supply the 

deficiency. · 

Sulphur.-In many of our analyses sulphur was determined as sulphur trioxide and recal-

. culated into the form of calcium sulphate. Part of the sulphur may really exist in organic 

combination, especially in the phosphatic brachiopods, and another part may be derived from 

sea salts, but this part is extraneous and should not be considered as ~ontributory to the sediments. A correction for it would be like that which we have regarded as applicable to the 

magnesia and of the same order of magnitude. ·In the marine sediments generally calcium 

sulphate is of minor importance. \ 

Other constituents.-Among the inorganic constituents of invertebrates there are other 

elements than those which we have determined. The most important one of these is fluorine, 

which is probably present in small amount in all living organisms. P. Carles, 54 for example, 

has detected fluorine in the shells of mollusks-as much as 0.012 per cent in oyster shells. In 

combination with calcium phosphate fluorine may .form a compound ana.logous to or identical 

with apatite. lts presence in vertebrate bones is well known. Boron also is widely distributed in the animal kingdom. G. Bertrand and H. Agulhon 55 detected it in. crustaceans, 

'mollusks, and echinoderms, as well as in vertebrate animals. Trac~s of barium have been 

detected in various organisms, and in the soft part of certain rh}zopods .granules of barium 

sulphate have been found. 56 Strontium is reported by 0. Vogel 57 in corals and molluscan 

shells, and according to 0. Butschli 58 the skeleton of a radiolarian, ~odecanelius, consists 

almost entirely of ·strontium sulphate. , Iron and manganese are of common if not of general 

occurrence in marine organisms, and copper, lead, zinc, cobalt, and nickel have also been 

found. 59 The presence of copper in oysters has long been known. Silver has been detected 

f>4 Carles, P., Compt. Rend., vol. 144, pp. 437, 1240, 1907. 

55 Bertrand, G., and Agulhon, H., idem, vol. 156, p. 732, 1913. 

ae Cited by Samoilov in Mineralog. Mag., Jw1e, 1917. 

>7 Vogel, 0., Zeitschr. anorg. Chern., vol. 5, p~ 55, 1894. 

58 Btitschli, 0., Deutsche Stidpolar Exped., vol. 9, p. 237, 1908. 

w See Forchhammer, G., Philos. Trans., vol. 155, p. 203, 1865. 

On manganese see Cotte, J., Soc. biologic Com pt. rend., vol. 55, p. 139, 1903; Bradley, H. C., Jour. Bioi. Chern., vol. 3, p. 151, 1907, and vol. 8, 

p. 2.37, 1910; Boycott, A. E., Naturalist, 1917, p. 69, and Phillips, A. H., Carnegie lust. Washington Pub. 151, p. 89, 1917. Phillips also fow1d iron, 

copper, and zinc in the soft parts of invertebrates, and, rarely, lead. See also Mendel, L. B., and Bradley, H. C., Am. Jour. PhysiolQgy, vol. 14, 

p. 313, 1905. F9r copper, .see Rose, W. C., and Bodaiisky, M., Jour. Bioi. Chern., vol. 44, p. w; 1920. 
GENERAL DISCUSSION. 57 

in oyster shells by A. Lhrersidge, 60 and vanadium has been reported in the blood of an ascidian 

by M. I-Ienze, 61 and in a holothurian by A. H. Phillips. 62 In short, a systematic search for 

minor metallic constituents in marine invertebrates· would probably show that they contain 

many other elements. This subject, however, lies outside the scope of our investigation, and 

these few citations are enough for present purposes. 

For the intensive ·study of coral reefs the analyses furnished by us together ·with those 

cited frmn others are of great significance. The limestone immediately below the zone of 

living forn1s owes its co1nposition to all the organisms that flourished on the reef. Algre, corals, 

alcyonarians, Foraminifemt, and other fonns of less importance contribute their remains to the 

building of the lilnestone, which may vary in cmnposition as the life upon it varies. Corals 

1nay, pred01ninate in one place, algre in another. Each reef must therefore be studied on its 

indi\Tidual merits if its chemical character is to be· understood. Precipitated carbonates, 

whether of bacterial origin or not, must also be taken into account, and their quantity may 

be large. At Funafuti, where the limestone has been studied with unusual thoroughness, 

the order of importance of· the leading organisms is estimated by A. E. Finckh 63 as follows: 

1, Lithot7uimnion; 6

'

1 2, Hctlimeda; 3, Foraminifera; 4, the corals, including Heliopora and other 

·alcyonaria.ns and the 1nillepores. Here the corals are subordinate to the algre, and even the 

Formninifera outrank thetn. To call the Funafuti rock a coralline limestone would therefore 

be somewhat misleading. 

At other localities the relative ·abundance of the marine organisn1s· is different from that at 

Funafuti. A careful analysis of smnples from reefs at Murray Island, Australia, conducted by 

T. Wayland Vaughan, 65 gave the following results: "1,600 feet frmn shore, madreporarian 

corals, 41.9 per cent; calcareous algre, 32.6 per cent; Foraminifera, 12.4 per cent; Mollusca, 10.2 

per cent. .At 200 feet from the shore the order is: Calcareous algre, 42.5 per c~nt; 1nadreporarian 

corals, 34.6 per cent; Mollusca, 34.6 per cent; Fora1ninifera, 4.1 per cent." Around the Tortugas, 

according to L. R. Cary, 66 the alcyonarian fauna is the 1nost in1portant contributor to the formation of reef li1nestones. lie estimates the quantity of alcyonarian spicules at this locality to 

avernge 5.28 tons to the acre; and at least one-fifth of this amount is added to the reefs annually. 

· In the Murray Island samples studied by Vaughan the alcyonarian remains were lacking. 

Chmnical analysis, however, is not the only factor of importance 1n determming the composition of a marine li1nestone. The crystalline character of 'the shells and skeletons, whether 

cnlcitic or aragonitic, must also be. considered. For this purpose the well-known reaction 

with cobalt nitrate, the "~1eigen ~·eaction," is conunonly mnployed, especially by W. Meigen 

himself, who has studied a considerable nurnber of organisrns, both recent and fossil, and some 

of his determinations 67 relate to genf'.ra examined by us. For these, excluding fossil forms, 

the data are as follows: 

Calcite. 

Lithothamnium. Alga. 

Lithophyllum. Alga. 

Polytrema. Foraminifer. 

Cora.Uium. Alcyonarian. 

Tubipora. Alcyonarian. 

Serpula. Annelid. 

Terebratula. Brachiopod. 

Argonauta. Cephalopod. 

BalanuR. Crustacean. 

GO Livorsidgo, A., Jour. Chern. Soc., vol. 71, p. 298, 1897. 

Aragonite. 

Halimeda. Alga.. 

Galaxaura. Alga. 

Millepora. Hydromedusa. 

Distichopora. HyclromeduHa. 

Heliopora. Alcyonarian. 

Spirula. Cephalopod. 

Sepia. Cephalopod. 

et Ronzo, M., Zoitschr. physiol. Chern., vol. 72, p. 401, 19ll,.a.nd vol. 86, p. 340, 1913. Henze also found copper in tho liver of cephalopods (idem, 

vol. 33, p. 417, 1901.) · 

02 Phillips, A. R., Am. Jour. Sci., 4th scr., vol. 46, p. 473, 1918. 

oa 'l'ho atoll of Funafuti, pp. 125-150, London, Tho Royal Society, 1904. 

o• Tho term Litl10thamnion as used in the Funafuti report is general and includes not only Lithothwrnnium but also Litho]Jhyllum, Goniolithon, 

uud perhaps othor genera. Soo 'l'ho atoll of Funafuti, p. 332, London, The Royal Society, 1904. 

· o~ Vaughan, '1'. W., Cool. Soc. Americu llull., vol. 2S,•p. 942, 1917. 

oo Cury, .L. R., Curnogic Inst. Washington Pub. 213, 1918, p. 356. 

oT Molgou, W., Naturf. Gesell. Frciburg Bor., vol. VI, p. 13, 1903. 
58 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. 

Of these genera, so far as they have been studied chemically, all in the aragonite column 

. are almost completely nonmagnesian. The trifling amounts of magnesia which they contain 

may be due to impurity or to alteration. Two in the calcite column, Terebratula and Balanus, 

are also nonmagnesian, and the others are all rich in magnesium carbonate. Meigen also tested 

twenty zoantharian corals, all aragonitic and nonmagnesian, and a considerable number of 

mollusks. Some of the molluscan shells were aragonitic and some were ·calcitic, but all except 

Argonauta were nearly free from magnesia. One echinoderm in Meigen's list, of a genus not 

represented in our series of analyses, was calcitic, and so too were ours. All the echinoderms, 

so far as we know, are distinctly magnesian. In short, it seems proboole, in the light of existing 

evidence, that all aragonitic organisms are essentially nonmagnesian; and that those characterized by the presence of much magnesia are calcitic. Many calcitic forms, however, are 

practi"cally free from magnesia. The general relation thus brought out is very suggestive. 

Magnesium carbonate associates itself only with calcite, with which it is isomorphous, rather 

than with aragonite, of different crystalline form, but why some organisms should secrete calcite 

and others aragonite in building their shells or skeletons is as yet unexplained, although the 

difference may be of physiologic origin, and may perhaps be correlated with differenc~s of 

structure. 

The considerations presented in the preceding pages bear directly upon the problem of 

the origin of marine dolomite. We now know what classes of organisms supply magnesia 'to 

the limestones and something also of what may be called their mineralogic nature. The dolomite ratio between the two carbonates is, however, never directly reached; there is always at 

first a large excess of calcium over magnesium, and a mixture is formed instead of the true 

double salt. To pro.duce dolomite the original limestone must either be eirriched by magnesia. 

derived from sea water or else concentrated by l~aching away of lime; furthermore, its two 

component carbonates must be somehow forced to combine. These processes may be operative 

simultaneously, but it is more probable that the change from magnesian limestone or dolomite 

is brought about by a series of steps, taken one at a time. 

In this connection the report on Funafuti, already cited, is remarkably suggestive. On 

. that atoll or island a committee of the Royal Society put down a boring 1,114 feet, in limestbne- all the way. The rock throughout cont'ained organic remains and was studied both 

microscopically and chemically. At a depth of 4 feet from the surface it contained 4.23 per 

cent of magnesium carbonate, and at 15 feet 16.4 per cent.· To this point a concentration 

by leaching is indicated, even if not absolutely proved, and it is probable that the relatively 

soluble nonmagnesian aragonitic structures had been in part at least dissolved away. The 

unstable aragonite is more easily dissolved than calcite, a relation so well established 'that it does 

·not need to be discussed here. 68 The fact that many sections of the core are described as "cavernous" in structure is additional evidence that solution had occurred. Solution is also aided 

by the carbonic acid generated during the decomposition of-the organic matter of the organisms, 

and through its agency calcite would be dissolved also. Magnesium carbonate is much less 

readily removed. 

At a dep'th of 25 feet the core contained 16 per cent of magnesium carbonate, but the 

specimen examined consisted largely of Lithothamnion remains, which accounts for its composition. On the other hand, the core at 40 feet carried only 5.85 per cent of magnesium carbonate and was in great part composed of Heliopora and Millepora, both originally nonmagnesian. 

The core secti<;>ns evi~ently varied in composition according to the variations in the organisms 

from which they were formed. A sample taken only a few feet away from the boring might 

have had a different composition. Some fluctuations in the series of analyses may be accounted 

for in this_ way. 

Below 40 feet the magnesian content of the rock diminished rapidly, falling at one 

point to 0.79 per cent of magnesium carbonate. Between 40 and 637 feet the composition of 

the rock was about that of an ordinary limestone, but at the latter depth crystals of dolomite 

68 This leaching at Funafuti has already been pointed out by J. W. Judd in The atoll of Funafuti, p. 384. 
GENERAL DISCUSSION. 59 

began to appear. 60 At 640 feet the percentage of magnesium carbonate was 26.33, and it 

increased, with some fluctuations, to the final depth of 1,114 feet. At 950 feet it reached 43 

per ce~t and at the lowest depth it was 41.05 per cent. From 640 feet downward the rock was 

essentially dolomite, although it contained an excess of calcite. Pure dolomite contains 45.65 

per cent of magnesium carbonate, a figure that was very nearly approached. 

The column of rock represented by the Funafuti boring thus appears to be divisible into 

three fairly definite zenes. The surface layer is about 25 feet thick, and its composition is 

directly detertnined by the organisms living above it. In this zone the evidence of concentration by leaching is quite clear. From 25 feet down' to 640 feet the rock is essentially a 

lin1estone, with very little magnesia. The lowest zone, from 640 feet downward, is dolomite, 

aJld the dividing line between this and the limestone above is very distinct. Within 3 feet 

the proportion of magnesium carbonate in the rock rises from 2.44 to 26.33 per cent. 

To account for all the differences in the Funafuti column does no,t fall within the scope 

of this investigation, even if it were possible to explain them. Possibly the limestone of the 

middle zone was laid down during a period when nonmagnesian organisms were relatively 

much n1ore abundant than they are now. This supposition, however, does not account for 

the sudden change frmn limestone to dolomite in passing from the second to the lowest zone. 

In order to partly explain these changes we venture to offer some· purely speculative suggestions, believing that speculation is legitimate if it tends to stimulate investigation or to 

provoke a closer scrutiny of existing evidence. 

The lowest portion of the Funafuti rock is, of course, the oldest, and it contains fragments 

of Lithothamnion and other organisms which flourish abundantly only at moderate depths. 

Magnesia was then concentrated in the rock, in part directly from living forms and in part 

by leaching, as at present. The thickness of the rock shows that it must have been deposited 

during a long period of depression, which may have submerged the island to a depth at which 

few of the magnesian organisms, especially the alg::e, could thrive. A prolonged rest, a period 

of equilibrium, then followed, during which very little roek was formed, and in this period 

much of the dolomitization took place. 

The period of rest was succeeded by one of elevation, which brought the dolomitic rock 

again to the surface, when reef building began anew, but with relatively fewer magnesian 

organisms than formerly. Between ·552 and 660 feet the nonmagnesian Halimeda is the main 

constituent of the cores. The new rock, then, was less magnesian than the older, and the 

sharp break between the two zones becomes intelligible. Magnesian organisms were not extinct, for their remains appear throughout the Funafuti cores, but they were much less abundant than at first. Whether this supposition is true or not might be determined by a quantitative study of the thin sections of the rock, which ought to be still in existence. The published 

records of them seem to be purely qualitative, except in so far asothey indicate the frequency 

of occurrence of the different organisms. They do not show their relative quantities.70 At 

present magnesian organisms predominate, and their composition is.reflected in the composition 

of the surface limestone. 

Our assumptions regarding changes of sea level at Funafuti are not altogether imaginary. 

In their-report upon the geology of the island T. W. E. David and G. Sweet n assert that 

the surface geological evidence collected by us proves, in our opinion, that several oscillating vertical movements 

of the above have taken place in the immediate past at Funafuti, and we should not. therefore, be surprised if the 

evidence gained ft·om the cor~ .shows that movements of the shore line in both directions have occurred at Funafuti 

at earlier epochs. 

The chemical and algal evidence reinforce the physiographic evidence. On the formation 

of "coral reefs" during subsidence or elevation there is an abundant literature, which we can not 

attempt to summarize. We are dealing with a specific instance from a single point of view. 

The subject is one over which there has been much controversy. Our principal assumption, 

69 See the petrographic report by C. G. Cullis in The atoll of Funafuti, pp. 392-420. 

70 The atoll of Funafuti, tables on pp. 336-361. 

n David, '1'. W. E., and Sweet, G., idem, p. 88. 
60 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. 

which may or· may not be sustained, is that the dolomitic ~one at Funafuti represent~ an old 

reef upon which the present reef is superimposed. A similar basing of new reefs upon older 

ones has already been pointed out by T. Wayland Vaughan. 72 

On many other islands in the South Pacific dolomitized limestones which were originally 

reefs are found at elevations hundreds of feet above sea level. A number of these rocks have 

been described by E. W. Skeats,73 who made good series of chemical analyses of them and also 

discussed their origm~ Organic and especially algal remains were common in the rocks and 

were clearly recognizable in spite of the fact that in m·any places they had been much altered 

by the process of dolomitization; In these islands an elevation of the land had clearly taken 

place. Similar limestones from the Fiji, Tonga, Tuamotu, and Ladrone islands have been. 

studied by R. L. Sherlock,74 who found them to be composed in great part of algal and foraininiferal remains. Among 4 7 thin sections which he examined, "Lithothamnion" was found 

in 35, Polytrema in 21, echinoderm fragments in 17, and corals in 15. In short, all the evidence 

goes to prove the importance of the algre as limeston~ builders and the subordinate character 

of the corals. This importance is now fully recognized by students of marine limestones and 

by paleontologists generally. 

It is not our purpose to discuss the origin of dolomite in general, for probably the rock 

originates in mo:re than one way.75 At Funafuti, however, and at other similar localities marine 

organisms have much to do with its origin, and that phase of the dolomite problem may appropriately be considered here. The first step in the process,. concentration by leaching, has already 

been described, but that is only a beginning. The living organisms, plant or animal, contain 

much less magnesia than is required to form dolomite, and its quantity must be increased 

from some outer source. The source, or at least· the only source which we can discover, is 

found in ocean water, in which magnesium is much more abundant than calcium. This source 

has beeri recognized by many authorities, and it is generally assumed that an exchange may 

occur between the magnesium of the water and the calcium of the limestones, the one 'replacing 

the other. This assumption is due_ to J. D. Dana/6 who sought to explain the dolomitization 

of a limestone on tht~ island of Makatea. · 

That an enl'ichment in magnesia from sea water is possible is shown by some experiments 

made by C. Klement/7 who found that a concentrated solution of magnesium sulphate and 

sodium chloride, at 90° C., attacked aragonite and corals strongly, yielding a mixture of carbonates containing as high as 41.9 per cent of magnesium ca:r:bonate. Calcite, on the other 

hand, was but slightly altered. In the light afforded by these experiments the nonmagnesian 

aragonitic organisms assume new importance and are perhaps more influential in the production of dolomite than the distinctly magnesian species. Their solubility is evident in the 

first stage of :magnesian concentration; their alterability is effective in the second. 

Klement's experiments, however, were performed with concentrated solutions and at a 

rather high temperature. Under natural conditi"ons, with less concentration and at lower temperatures, the same reaction may take place very slowly but be equally complete in time. Years, 

or even decades, may be needed to effect such changes as were produced in the.laboratory 

within· 48 hours. In the study of geochemical processes the time factor must always be taken. 

into account. 

So far, according to Klement, a mixture of the two carbonates has been formed. To 

convert them into the double carbonate, dolomite, another step must be taken, and here again 

time may be important. A porous rock has been produced, which is saturated with water and 

buried at a depth which subjects it ·to considerable pressure. If the "two carbonates of calcium 

and magnesium were perfectly dry, pressure alone would probably not.effect their union, but 

12 Vaughan, T. W ., Papers from the Tortugas laboratory of the Carnegie Institution of Washington, vol. 5, p. 66, 1914. 

78 Skeats, E. W., Harvard Coli. Mus. Comp. Zoology Bull., vol. 42, p. 51, 1903. 

1~ Sherlock, ~. L., idem., vol. 38, p. 349, 1903. . 

76 For a rather full summary of the subject see Clarke, F. W., The data of geochemistry, 4th ed., U.S. Geol. Survey Bull. 695, pp. 557-572, 1920. 

10 Dana, J. D.,. Corals and coral islands, 3d. ed., p. 393, 1890. 

11 Klement, C., Min. pet. Mitt., vol. 14, p. 526, 1894. See also Pfaff, F .. W., Centralbl. Mineralogie, 1903, p. 659, and Neues Jahrb., Beilage Band 9, 

p. 485, 1894. Two separate papers. · 
GENERAL DISCUSSION. 61 

under the influence of moisture, with slight solution going on at the surfaces of the solid particles, there would be a degree o( molecular mobility which might bring about combination. 

'fhis is probable, although so far as we know it has not been actually proved. The establish- · 

ment of the facts ought not to be beyond the range of experimental investigation. Views 

similar to ours relative to the formation of dolomite have already been advanced by General 

E. Jourdy, who especially recognizes the importance of the algffi, of aragonite, and of time. 

Much remains to be done, however, before the problem of dolomitization can be completely 

solved.78 

On the subject of phosphate rock we have little to .say. We have shown that several 

groups of organisms are rich in phosphates, but the extent of their contributions to the sedinlents is uncertain. At best they can at first form beds of only moderately phosphatic limestone, which may perhaps be concentrated by the leaching away of the excess of carbonates. 

Vertebrate skeletons are also phosphatic and may possibly be more important additions to the 

sediments than invertebrate remains. In some localities limestones have been phosphatized 

by percolations from beds of guano/9 but that process is not one which needs to be considered 

here. It has no relation to the present research. 

In the course of our investigation we have made one very curious discovery, to which 

we have repeatedly called attention, namely, the fact that in certain grQups of organisms the 

proportion of magnesium carbonate is dependent upon or determined by temperature. The 

crinoids and alcyonarians show this relation very clearly, and it is also suggested by our analyses 

of foraminifera, crustaceans, and algaL As a rule the organisms from warm waters are much 

richer in magnesia than those from cold waters, and the observed differences are often strikingly 

conspicuous. Tlus rule, or rather tendency, we are inclined to believe is general, although we · 

1nust admit that there are probably exceptions to it. Whether the increase in magnesia in 

passing fro1n cold to warm regions is absolutely regular or not we do not venture to say, but 

apparent irregularities may be due to any one of several different causes. Slight analytical 

errors, uncertainty as to exact temperatures, impurities in the specimens analyzed, and differences in the concentration of sea water may all help to produce irregularities, which, however, 

are not likely to be large. The salinity of ocean water is very variable; it is 3.5 per cent in 

·the great ocean, 4 per cent on the southern shores of the Mediterranean, and less than 1 per 

cent in the Baltic, diff.erences that are great enough to exert some influence upon the vital 

processes of marine animals. Although the ratio between calcium and magnesium is practically constant in all oceanic waters, a concentrated water would contain more magnesilJ.m, · 

volume for volume, than a water that was more dilute. Whether an organism living in a concentrated water would assimilate more magnesia because of its enriched environment no one 

can say, but conceivably it might do so. The influence of temperature might in that way be 

slightly modified. This is only a suggestion, not a staten1ent of established fact. That warmth 

favors the assimilation of magnesia by marine invertebrates seems to be reasonably certain, 

but why it shoul~ be so is not clear. The relation is definite but as yet unexplained. We 

hope it is not inexplicable. · . 

Attempts will probably be made to use our data in studies of climatology, but are such 

atten1pts likely to be fruitful~ The question is not easy to answer. At a first glance it would 

seem as if warm. regions should be more favorable to the formation 9f magnesian limestones 

than cold regions, but the evidence is by no means conclusive. A. dense population of coldwater organisms might add more magnesia to the sediments than a sparse population of warmwater forms. The massiveness of the animals must also be considered. The alcyonarian 

Paragorgia arborea, as its specific name indicates, grows to treelike dimensions, but its skeleton 

contains only about 9 per cent of magnesium carbonate. Tropical alcyonarians are much richer 

individually in magnesia, but they are not so· large. One Paragorgia would therefore count 

78 Jourdy, K, Soc. gaol. France Bull., 4th ser., vol. 14, p. 279, 1914. 7° Clarke, F. W., 'l'he data of geochemistry: u.S. Geol. Survey Bull. 695, pp. 515-526, 1920; gives a summary of our knowledge or phosphate rock. 
62 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATJ<JS. 

for more than many smaller alcyonarians in the formation of magnesian limestone. If, however, the warm-water organisms are as abundant as the cold-water forms, and if their ~ggregate mass is as great, then the tropical limestones of marine origin should b~ moreTichly magnesian than those from higher latitudes. The determination of the facts we must leave to 

zoologists. 

In conclusion we must express our thanks to the officers of the United States National 

Museum and the United States Geological Survey, who have aided us by furnishing authentic 

material for our investigations. Messrs. Paul Bartsch, Austin H. Clark, William H. Dall, 

W. L. Schmitt, T. Wayland Vaughan, and 'Frank Springer and Miss M.G. Rathbun have all been 

most generous with their services. Dr. Marshall A. Howe, of the New York Botanical Garden, 

has also been most kind in supplying us with algrn, and Prof .. L. R. Cary has kindly furnished 

us with valuable unpublished analyses. Without the help of these friends our research would 

have been impossible.· 

Addendum. -Since the manuscript of this paper was prepared Prof. A. G. Mayor 80 has 

published an important paper on Rose Atoll, near Samoa. The reef here consists chiefly of 

Lithothamnium remains, with very little coral. Analyses of the reef rock by A. H. Phillips 

gave percentages of magnesium carbonate ranging from 14.36 to 19.47. 

so Am. Philos. Soc. Proc., vol. 60, p. 62, 1921. 

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