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DEPARTMENT OF THE INTERIOR

ALBERT B. FALL, Secretary

UNITED STATES GEOLOGICAL SURVEY

GEORGE Otis SMITH, Director

Professional Paper 124

THE INORGANIC CONSTITUENTS OF
MARINE INVERTEBRATES /

SECOND EDITION, REVISED AND ENLARGED

BY

FRANK WIGGLESWORTH CLARKE

AND

WALTER CALHOUN WHEELER

; SF mew

WASHINGTON
GOVERNMENT PRINTING OFFICE
1922

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54
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Trve et. Sd San ee HOUSE OF REPRESENTATIVES 1 No. 884
7 c € t = a a — — = —

DEPARTMENT OF THE INTERIOR

ALBERT B. FALL, Secretary

UNITED STATES GEOLOGICAL SURVEY

GEORGE OtIs SMITH, Director

Professional Paper 124

THE LNORGANIC CONSTITUENTS OF
MARINE INVERTEBRATES /

SECOND EDITION, REVISED AND ENLARGED

FRANK WIGGLESWORTH CLARKE

AND

WALTER CALHOUN WHEELER

WASHINGTON
GOVERNMENT PRINTING OFFICE

1922

CONTENTS.

Introduction. >< cde cea aos: ee Ieee eee eee OR oe ee ae Sede Ue Sete pe ae eee ae
IM OVAMIN Oras oi ee ee ehaes oe dels cig ok a en 20 Seo pase mete Shows beeen Ree See
DON PCS. kee E ee SRE e eee ae anak euss f5s56 Je aeoase SUB 2 52 57 Se OSA PO ADRS aS Ootiae soc anoahS:
Ms dreporatianicoralesacn-5---0-no-< <a eeepiane BN AA oe ae 3 SR PEIN SE a aS aoe
Mleyonarian’ corals .,5 2/235) 2852 Solow wc St Sg yo ER 2
FV GTOldS Nt w= 8 Ser aos? win Als oes Ae ons eee Ce ee Pa bts araSeajn sa eee alos ee malas aa) tacleeee Meee ese
AMM eH AS repose a cits Berets a wie cre aitlara tearocnace Sack SERRE RMSE ee an Sr

aL OF ao) Ks |<) eh a eee heey Sh PP TE IP) © (8S SRI aa Slt pe BOR os See oo eG OE

2
SHO bATASDOS S253 Sat, saacc ein ue Meow eee See Oe eee EE ios Rene
4 NOpPHMITANG:: = on. cio ein Sosa wale onsale 5 2 SEER Re oe ee ee TRS oe 2 ee SRE
5, Holothurians:- «2% osu aaa oeiekt esicisin eee ee eee ee Se eee eee
6 ASuUMMAr ye isos oK x sin cc eee Oe Oe Ble Se SC oe RT. nn aR
BEV OZOB: =: <j<)<.2)= msm bins = o)Scie sia ays ela STS HI lesa 5d eae 2 oa eee re ence oe en
Brachiopods 21 crema: eta eee SAE ee es eet cen a2 et SG eee aoa Ciede base goose cen
ko) LV) eee ee eS Oe Se eRe ome TR De uN 2 Renee re oa Pes Ae cst Sy
I. WPelecypodee... ..< - oes dhs ac ees eeate reek Saket Li: ee ek 2) ee ee
2... Scaphopods'and amiphineurangs -'<.. ts fiy so. 1 Saas eye nee ere > a
Bs (Gastro pods! = 2:5 detec de fere eae hoo idl Re REE « RRO Se Ee eS = 2) lA nr a
4: Cephalopoda... tc. 5.jes tense ices els selerin = SRR tea ee ne ere ro) eI ee ae eee
G@rvistacean se. Sere se fate a cpatk ete sche Soh Uz ast elses <1 ae Tce vn AST Se hehe a
@alcareoustal pee. .2/; 2nis =k. Stet ct ceive ate ecy- Be 5 <n Saito: See
Generali discpissions = 4.8 22 eect ee ols oie ee ee Be Sear Sais ieee Eeftet oicte iia. of eee eae
I

THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES.

By Frank WiaeLeswortH CLARKE and Water CaLHouN WHEELER.

INTRODUCTION.

That many rocks were once marine sediments and that animals and plants shared in their
formation is one of the commonplaces of geology. Coralline limestones. shell limestones, and
crinoidal limestones are among the most familiar illustrations of this statement. That corals
and shells furnish calcium carbonate to the sediments, that radiolarians and diatoms are siliceous,
and that vertebrate animals, some crustaceans, and a few brachiopods are more or less phos-
phatic are also well-known facts, which, however, have been determined in a more or less desul-
tory way. Nosystematic investigation to ascertain just what substances each class of organisms
contributes to the sediments seems to have been made, and the present research is an attempt
to cover the ground a little more thoroughly than it has been covered heretofore. Complete-
ness is, of course, impossible, but something of a foundation for future work we have tried to lay.

Although many analyses of corals and molluscan shells are on record, some important
groups of organisms have received only slight attention, and little has been done heretofore
toward determining the composition of their tests or skeletons. The few published data, more-
over, have been generally but not always incomplete in certain particulars, especially with
regard to the localities from which the specimens analyzed were obtained, the depth of water
in which each creature lived, and the temperature of its habitat. ven in our own work some
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. Furthermore,
some of the older analyses are unsatisfactory for other reasons, for many of them were made
to determine single facts, such, for instance, as the proportion of magnesium carbonate alone;
and in others such essential constituents as organic matter were not taken into account, an
omission that is especially serious, for it renders the accurate comparison of the analyses with
others impossible. In the shells of mollusks the proportion of organic matter is small, but in
the echinoderms, alcyonarians, and phosphatic brachiopods it is relatively large, ranging from 10
to even 40 per cent or more. This statement, however, must not be misconstrued; it refers,
of course, only to the specimens, dried or alcoholic, which were actually analyzed. In order to
compare the analyses, therefore, so as to determine the true composition of the inorganic shells
or skeletons, the very variable amounts of organic matter must be rejected, and the remainders
recalculated to 100 per cent. Relations then appear which are not recognizable when the crude
unreduced analyses are compared. Any one of our tables of analyses will show this fact very
clearly.

In astrict sense completeness can not be claimed even for our analyses. Minor constituents
which have been detected in marine organisms, such as barium, strontium, fluorine, manganese,
copper, zine, and lead, have been ignored. They occur, as a general rule, only in traces and have
little or no significance with respect to the larger problems before us. What organisms tend
to form limestones and what ones are notably magnesian, phosphatic, or siliceous are the ques-
tions which we are attempting to answer. In most of the analyses lime, magnesia, 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

2 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES.

organic matter, and water. The carbon dioxide was calculated to satisfy the bases, and the
amount thus determined, subtracted from the total loss on ignition, gave a fair but rough esti-
mate of organic matter plus water. From 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
main problem and 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 (published in 1917 as Professional Paper 102) the
analyses, with few exceptions, were made by the junior 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, because of their enormous abundance, are of great importance in the
formation 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 pub-
lished in the report of the Challenger expedition on deep-sea deposits show that it consists
mainly of calcium carbonate, with 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, fortu-
nately, is not insuperable, and with the help of others we were able to obtain representative
samples of seven species, as follows:

1. Pulvinulina menardii D’Orbigny. Albatross station 2573; latitude, 40° 34’ 18’ 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. Spharodina dehiscens Parker and Jones. East coast of Mindanao, Philippine Islands; latitude, 8° 51’ 45” N.,
longitude, 126° 26’ 52’ E.; depth of water, 804 meters. .

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

4. Orbitolites marginatis Lamarck. South of Tortugas, Fla., at depth of 29.3 meters.

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

6. Polytrema mineaceum Linné. Bahamas.

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

Analyses of Foraminifera.

1 2 3 4 5 6 7

lias i |

ESOS BSG Se se Fs me \enT4e70)seeseooml em OsO3))||, 0:03) tame mil
IATA OnE tk, LUNGS SR SSRIS CL SERA 381+ 476 18° 238 golly 0:02 I O.i88
Reise S2iEk .aontas aed naeiaee Mee eile sae See sages sie | 1.68 82 5.08) 4.93) 4.64) 5.09} 4.32
GaGets. 5 at ae. oer De 2.2 41.36 | 45.48 | 47.35 | 48.92) 48.79] 47.35 | 49.02

LEAS} Songe Sarde aso ngcsennSsdodos aba sdcgdabeactonessS¢ He C2) (2) .00 | Trace.| Trace.| (?) | (?)

Ignition: Ao weeks: Rites eee. her eae s--| 38.12 39.01 46.57 | 45.20 45.56 | 46.24 | 45. 54
99.67 98.62) 99.16 99.48 99.19] 98.70} 99.42
COpneeded’ss kote rets ico e pe eee. seem 34.41 | 36.63 42.74! 43.90, 43.43 | 42.80| 43.27
Organic matter sete... cect ep a ey 2 eee sara 2. 38 3.73 1. 30 2.13 3. 44 | 2. 27

| ( 2 |

Rejecting organic matter and water (‘‘ignition”’) and recalculating to 100 per cent, the
analyses assume the following rational form:

SPONGES. 3

Reduced analyses of foraminifera.

1 2 3 4 5 6 7
Sy te eae cen ale Slates mie mis a aie sieienis ese ess 15. 33 8. 89 0.03 0.31; 0.11 5. |i: -
Tee, SS ee g ga\| 4. 94 Pe MeLONte y 1S. [e 09 } 0:02 |r) 0.56
NneT) 1 ie eyes 8 SCE SRS 3.67 1.79 11.08] 10.55) 10.04/ 11.22 9.33
ORG oy boa eee ae Aone 77.02 | 84.38 88.70| 89.01} 89.76) 88.76 90.11
(CELE Cs cece pace Sco OC OB CEC OSE te Oo mente (?) (2) - 00 | Trace.| Trace.| (7?) (?)
100.00 | 100.00 | 100,00 | 100. 00 | 100.00 100.00 100. 00

In Nos. 1 and 2 the high percentages of silica and sesquioxides are evidently due to impuri-
ties. If they are rejected, the percentages of magnesium carbonate become 5.19 and 2.09,
respectively. These two Foraminifera are from deep and presumably cold water. The others
are from shallow water in warm regions. A similar difference in the magnesian content of the
organisms with regard to temperature appears, with much regularity, in several other series
of our analyses, notably in the series of crinoids and alcyonarians.

The only published analyses of Foraminifera that we have been able to find elsewhere
appear in H. B. Brady’s volume * on the material collected by the Challenger expedition. Four
of these analyses relate to Orbitolites complanata var. laciniata, collected either at Fiji or at

’ Tongatabu, Friendly Islands. These analyses are as follows:
1, 2. Two different samples; analyses by C. R. A. Wright and J. T. Dunn.
3. Sample washed with boiling water.
4. Same as No. 3, unwashed; analyses 3 and 4 by J. Gibson.

Analyses of orbitolites.

1 2 3 4

Si OMe nee ese: 0.58 0.3 0.14 0.11 |
Wea MO eee oniateepte oer |29). F283 DIN | Poatte tae SA |
UME Ope: Oto. o2 5525 SOR steed oa] erecac oct oomeeemens |
6 00 ek eae See 12. 52 8.8 9. 55 10.50 |

CaCO fog. Sete ta. | 86. 46 88. 2 88. 74 87. 91

|
100. 24 100. 0 98. 43 98. 52

a Including “phosphates of lime and magnesia.”

Other analyses, presented by Wright and Dunn, are of Amphistegina lessonti, from 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 magnesium carbonate. In
a Tertiary fossil, Nubecularia novorossica, 26 per cent of magnesium carbonate was found, an
amount which probably represents a concentration by leaching. There are also analyses of
several siliceous Foraminifera containing from 76.1 to 94.7 per cent of silica. Their tests appeat
to have been built up of sand grains and are of little or no significance in relation to the main
problem of the present investigation. Their remains are not true additions to the sediments.

SPONGES.

The sponges are a rather unsatisfactory group of organisms for chemical analysis, on account
of their very porous structure. They are liable to contain inclosed impurities, such as sand
or mud, which are difficult to remove entirely. Some sponges are siliceous, some calcareous,
and others are composed almost wholly of organic matter, the proteid spongin. The ideal
method of procedure for our purposes would obviously be to separate the clean siliceous or cal-
careous spicules and to analyze them alone, but to do that a large amount of material would have
to be gone over, and more time would be consumed than we could give to this part of our investi-

1 Brady, H. B., Challenger Rept., vol. 9, pp. Xvii-xxi, 1884,

4 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES.

gation. As the sponges are of minor importance in relation to the marine sediments, we have
followed our usual 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
completely, as follows:

1. Euplectella speciosa Quoy and Gaimard (‘‘ Venus’s flower basket”). Philippine Islands.
2. Bathydorus uncifer F. E. Schulze. Galapagos Islands, Albatross station 2818; latitude, 0° 29’ S.; longitude,
89° 54’ 30’ W.; depth of water, 617.4 meters; bottom temperature, 9.9° C.
3. Halichondria panicea Johnston. Alaska.
4. Suberites suberea Johnston. Unga Island and Popot Strait, Alaska.
5. Tethya gravata Hyatt. Buzzards Bay, Mass.
6. Pheronema grayi Kent. Off southwest coast of Ireland, at depth of 1,098 meters.
7. Geodia mesotriena var. pachana Lendenfeld. Off southern California, Albatross station 2909; latitude, 32° 22/
N.; longitude, 120° 08’ 33’ W.; depth 377 meters; bottom temperature, 7.3° C.
8. Phakellia grandis Verrill. Browns Bank, northeast of Cape Cod, between latitudes 40° 30’ and 42° 08’ N. and
longitudes 66° W.; depth, 549 meters.
9. Chondrocladia alaskensis Lambe. On beach at Unalaska, Alaska.
10. Gelliodes grandis Verrill. Gulf of Maine.
ll. Esperiopsis quatsinoensis Lambe. Attu Island, Alaska.
12. Chalina arbuscula Verrill. Outer Island, Conn. 5
13. Spinosella sororia var. crispa Duchassaing and Michelotti. Harrington Sound, Bermuda.
14. Buspongia officinalis tubulifera var. turrita Wyatt. Harrington Sound, Bermuda.
15. Hireinia campana var. turrita Hyatt. Bermuda.
16. Euspongia officinalis var. rotunda Hyatt. Nassau, Bahama Islands.
17. Hippospongia equina var. meandriformis Duchassaing and Michelotti. Nassau, Bahamas.
18. Hippospongia agaricina var. dura Hyatt. Off Duck Key, Fla., at depth of 4.6 meters.
19. Hippospongia caniculata var. gossypina Duchassaing and Michelotti. Off Rock Island, Fla., at depth of 7.3
meters.
20. Aplysina hirsuta Hyatt. Off St. Thomas, Virgin Islands; depth of water, 37 to 42 meters.
21. Grantia ciliata, Fabricius. Woods Hole, Mass. Weight of sample only 0.0982 gram.
22. Leucilla carteri, var. homoraphis (Polejaeff). Off Culebra Island, Porto Rico, Fish Hawk station 6090; bottom
temperature, 26° C. Weight of sample, 0.1422 gram.
Analyses 21 and 22 by R. M. 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.

i !
| 1 2 a wig it 5 Cer diane, 8

SO) RRA RO Ro os aes can, Se a ee Ane | 88.56 | 76.33 | 74.52 | 72.92 68.46 65.19 | 64.96 60.70
GN LO OF eciam Sater morisc te) gain eee ae ee 32 1.05 6.19 1222) 3.71 3.05 -21 2.30
MOORS Sao «detect ne cee = eee eee ot Rae maior | -00 48 1.43 -55 61 Trace -38 23
CaO) Reece eee aes Mt on ae eee eee eee 22 2.78 2.33 -33 -89 3.26 29 -d1
REO) «Se RISES Ee ee: aE, -00 | -00 -17 -49 -48 | Trace. | Trace. -46
SOAS Se eecon tac Co ses ee eee Be nee tA oeeee = =| -00 Trace. -08 -39 -40 -02 PA (?)
WONTON. .2 ce eein oe sac hc eee ee see | - 10.26 | 18.60 | 12.11 | 22.21 © 23.02 27.05 31.78 35.84

99.36 |. 99-24 | 96.83} 98.11 97.57 98.57 97.78 | 100.04

|

9 10 iE 12 ae gees 4 15 21 22
Si Ogee ohn 4. DAsewraeie erect oe ee 53.96 52.24 51.94 31.93 3.08 1-31 0.57 AiR 0.77
CATR a) Obra teas hl Sess ees 1.64)" 3.86 91-88 | 1.44 Fe 196}. 2229 ; :
MeOke sea ease rete eee -58 86 75. -97 -26 -43 1.60 ed! 5.50
ORO SR CARs: 22! Eerie eee 1.28 ne S777 9 .97 2.80 7.58 20.01 38.39 38.97
1 ORE es Ae Oe OAs a eee tee 31 -61 -46 -50 .69 ie) -68 (?) (?)
SONS aeete Seen aaen wGSee Sao ssa (?) 81 (?) -82 (?) (?) (2) 00 -00
lenitioness%, 2.25: Les Se ee 41.97 | 34.64) 42.29 61.25 | 91-32 | 89.75 | 75.68 | 50.31 53.58 -
99.74 93.79 98.59 97.88 98.92 | 100.80 100.83 98 .92 98.82

SPONGES. 5)
Sponges Nos. 16-20 were composed mainly of organic matter, and only loss on ignition
was determined. The figures showing this loss are as follows:

Per cent.

16. Buspongia CRATER OER een ELSA /ns <b See ee enefeetern so ninicin oeca os 96.94
17. Hippospongia equind.........-22--. 220-2 eee e cence eee ee ee ee eee cee eee eee e tenn ence sete esee. 96.98
18, Hippospongia agaricina........----.-.-.- 02.22 e seen eee ee eee eee eee settee neers ect c cere 95.55
19. Hippospongia caniculata...........-.----2.- 202 cece eee eee cece cette nee te estes ees 94.72
DMPA pty sine Mirela. 2! a wins = wo Se ee eB es one inl ee wine nemecere senor aise 93.34

In these sponges the inorganic matter is so small that it may only represent impurities,
an analysis of which would be meaningless. Euspongia and Hippospongia are the ordinary bath
sponges of commerce.

In several of the analyses, notably in Nos. 3, 5, 7, 10, and 12, the summation is very low,
on account of the presence of soluble salts (sea salt), which were not determined. Analyses
1 to 12 represent siliceous sponges, and in Nos. 2 and 6 the siliceous spicules were very conspicuous.
No. 1, Euplectella, the beautiful “Venus’s flower basket,”’ may be taken as the type of these
sponges, and the analysis shows that its skeleton is composed of nearly pure opaline silica.
Several analyses of siliceous sponge spicules by other chemists lead to similar conclusions. In
spicules from unnamed species J. Thoulet * found from 12.88 to 13.18 per cent of water, F. E.
Schulze * cites an analysis by Maly of spicules from Poliopogon amadou which contained 7.16 per
cent, and in seven species of siliceous sponges W. J. Sollas ‘ found water varying from 6.1 to
7.34 per cent. In all these sponges the spicules or skeletons are composed of amorphous opaline
silica. The other constituents shown in our analyses, except organic matter, which is included
in the loss on ignition, are probably but not certainly impurities. To this statement one excep-
tion may be made. 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, is by no means
certain. Thatnearly all the sponges analyzed contain phosphorus in some form, although in
small amounts is clear.

Analysis No. 13, of Spirosella, 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, and 22, however, repre-
sent calcareous sponges. Reducing the analyses to standard form, rejecting organic matter,
and recalculating to 100 per cent, we have the following figures for their inorganic portions:

Reduced analyses of calcareous sponges.

SOs: oe Daas ee 7. 81 1.36 |) | ;

(il (0 1 aaa me Ba72i b> gbaoq imme De
NO see eee. ye 5.37 | 8.00 | 4.61 14. 10
CGO) ese Asean e oeaeee 71.14 81.64 | 84.92 84. 96
(CE JEN OM. On See See Cp aeeece 9. 96 3.55 | (2?) (?)
(OF S10}. Seas See See aeeee (?) (?) 00 00

100.00 | 100.00 100. 00 100. 00

t i

—— — .

We can not assign much weight to these analyses, because the specimens analyzed evidently
contained impurities—silica and sesquioxides. Even the small amounts of phosphate shown in
two of them may be due to the inclosure in the sponges of minute crustaceans, such as copepods.
The small percentages of P,O, in the unreduced analyses are much magnified in the reduction.
Some of the magnesia also may belong to inclosed sea water, and in this respect analysis No.14
is especially questionable. The presence of magnesia, however, in these sponges is somewhat

2 Thoulet, J., Compt. Rend., vol. 98, p. 1000, 1884.
3 Schulze, F. E., Challenger Rept., vol. 21, p. 28, 1887.
4Sollas, W. J., idem, vol. 25, p. xlviii, 1888.

6 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES.

confirmed by an analysis by O. Biitschli ° of spicules from a Mediterranean sponge, Leucandra
aspera. His figures are as follows:

(C0) sa eee eee ee OER Se Acinn Sc eecemce soon. asoneasm sr Iesdcs sze Fees Gres eS: 86. 76
MaQO 3-222. -$ o's 0 2520 oe oc So~ cleteinnle Ae ae eeetp ye C= Se ore ee mete el are ciate eee 6. 84
13 0) AO anos OCS e sere be cosas Ss Gas aacsaccctssnoph asSSE CoA Se. 26
GaSO}.2H.0). 2.2.2: 2222 shgscecc = oases meee ele wen lata eee telat iede n iedeeie tae 1.97
1: (0 hee eee ee Seer ASE ancien 4 eooe ESE eco esamad 5558Scs2sodesaser swe cae Toe 3. 14
Oreanic matter. = /. 5-7-2 5 - = > a seas ine ee ee ee aie oe eee een nine minim ae = cial 42

99. 39

If we reject water and organic matter and recalculate to 100 per cent the percentage of
magnesium 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.®

MADREPORARIAN CORALS.

Madreporarian or stony corals have been repeatedly analyzed and with generally concord-
ant 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 aleyonarians. The list of corals analyzed is as
follows:

1. Balanophyllia floridana Pourtalés. South of Key West, Fla.; depth of water, 165 meters.

2. Paracyathus defilipii Duchassaing and Michelotti. South of Key West; depth, 165 meters.

3. Dendrophyllia cornucopia Pourtalés. Fish Hawk station 7286, Gulf Stream, off Key West; latitude, 24° 18’ 00”
N.; longitude, 81° 47’ 45’ W.; depth, 243 meters; bottom temperature, 11.9° C.
Siderastrea radians Pallas. Shoal water, Ragged Keys, Fla.
Porites furcata Lamarck. Shoal water, Ragged Keys.
Favia fragum Esper. Shoal water, Tortugas, Fla.
. Eusmilia aspera Dana. Shoal water, Tortugas.

8. Oculina diffusa Lamarck. Shoal water, Tortugas.

9. Cladocora arbuscula 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 Pourtalés. Albatross station 2087; latitude, 40° 06’ 50’ N.; longitude, 70° 347 15 W.;
‘epth, 119 meters: bottom temperature, 10° C.

13. Flabellwm alabastrum Moseley. Albatross station 2677; latitude, 32° 39’ N.; longitude 76° 50’ 30’ W.; between
the Bahamas and Cape Fear; depth, 875 meters; bottom temperature, 4.5° C.

14. Madrepora polifera Linné. Albatross stations 2662-2672, between the Bahamas and Cape Fear; depth, 512-894
meters; bottom temperature, 6°-7° C.

15. Dendrophyllia profunda Pourtalés. Locality, ete., as under No. 14.

16. Madracis decactis Lyman. Shoal water, Bermuda.

17. Deltocyathus italicus Michelotti. Albatross station 2750; latitude, 18° 30’ N.; longitude, 63° 31’ W.; depth,
908 meters; bottom temperature, 7° C.

18. Acropora cervicornis Lamarck. Shoal water, South Bight, Bahamas.

19. Orbicella cavernosa Linné. Reef, Light House Point, Andros Island, Bahamas.

20. Meandra clivosa Ellis and Solander. Locality as under No. 19.
Mxandra labyrinthiformis Linné. Reef, Golding Cay, Andros Island, Bahamas.
. Mussa aff. M. dipsacea Dana. Reef, Golding Cay.
Porites clavaria Lamarck. Reef, Golding Cay.
. Porites astreoides Lamarck. Reef, Golding Cay.
5. Flabellum pavonium var. paripavoninum Alcock. Albatross station 4080; north coast of Maui, Hawaiian Islands;
depth, 326 to 369 meters; bottom temperature, 13.5° C.

26. Madracis kauaiensis Vaughan. Albatross station 3982; vicinity of Kauai, Hawaiian Islands; depth, 426-435
meters; bottom temperature, 9.1° C.

Nie

bo bo bo
w bo

es

tS bo
or

‘

5 Butschli, O., K. Gesell. Wiss. Gottingen Abh., No. 3, 1908.

6 On manganese and iron in sponges, see Cotte, J., Soc. biologie Compt. rend., vol. 55, p. 139, 1903. ‘Traces of managanese were noted in our own
analyses.

MADREPORARIAN CORALS. %

27. Flabellum sp. Albatross station 5273; latitude, 14° 03/ N.; longitude, 120° 277 45” E.; China Sea, near southern
Luzon; depth, 102.5 meters.

28. Desmophyllum ingens Moseley. Albatross station 2785; latitude, 48° 09’ S.; longitude, 74° 367 W.,; off coast
of Chile; depth, 822 meters; bottom temperature, 83°C:

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

but they are unimportant.
Analyses of corals.

1 2 3 4 5 6 7
E Vi See fe

CO ee ere ee icin oe ais ete teeta sale = = 2 =.= 0. 09 0. 42 0. 09 0.12 0.11 0. 28 0. 00
(Rie Ogee foot. ne. LL eeeeee cotaetet---. 69 35 iL ey 10 i112 04
oe WO) Ope Re OCR OE OB ON: af ae oe ISOS aero .49 39 19 ay Al . 36 | .18 21
(N01 Gta cere Boe Ben boat ooanacoc Gnna et aes ot 51.46 | -52.03 | 51.71) 51.23) 51.61 | 53. 69 53. 25

oye ee ge ee tearm. «sn (2) 00} (?) (2) @)- | _@) (2)
1D10 an aatlose DOSHE BDEEEDOEESP Ane obo cDOSaC a a ooog Trace. | Trace. | Trace. | Trace. | Trace. | Trace. | Trace
Wenition=.2 << are vicina. sive = = rien oor | 47.32 | 44.62 | 46.23 | 47.21 | 46.76 | 44.44 44, 90

|
100.05 | 97.81} 98.33 | 98.88 | 98.94) 98.71 98. 40
CO, needed ban roacec ue c 1onee> boone 39.97 | 41.31 | 40.84 | 41.48 | 40.95 | 42.38 42. 08
Organic matter, etc......---------++2-+ee rete ett Too Ne eotow 5390) petoe woe 2. 06 2. 82
|

8 9 10 11 12 13 14
Fo ioe list ety i ai pe ae oe eae aes ae 0.07| 0.30| 004, 0.00) 0.20; 0.09) 0.19
(ECE Oe, ae te ged SAE eae “05 -08 00 03 27 | i 07
hp AOR BS He eos obeat aie moore oor Do 719 . 05 . 28 wal «29 17 06
CaO Sree enes orale Sake arale eta inlaintsfe nja/= wieinteminy~)=lmuassinl= 538.90 | 53.31 53. 87 53. 58 53. 79 53. 54 53. 48
Skee a ARE ON ete a scise lave (2) Te) (2) 10} (2) 12
120 =e gn aan APR ESSE scacc cadences Seo sreonge Trace. | Trace . 00 00 Trace. | Trace. | Trace
TES in 0 Bo nee en odese eb >>> Sorc sce COpoRe OCR Ga | 44.57 44. 83 44, 62 44,76 44.47 44.80 | 44.50
| 98.78| 98.57 | 98.81) 98.64) 99.12 98.71 | 98.42
CO, needed... -2.. 222 -2- 2-22-2500 - 2222 meres 42.56 | 41.94| 42.64) 42.40 | 42.52) 42.26 | 42. 03
Organic matter, etc...-------------++-+-+--+---er5tt- | 2.01 2. 89 1.98 23 6e\ kage 2. 54 | 2.47

|
<3 | |
is 15 16 17 18 19 20° |) 821

98. . : 99.43 | 99.11
CO, needed....------------++---- 22-02-22 2 e erste ete: 42.35 | 41.84) 42.41 41.45 |) 42.09 42.56} 42.49
Organic matter, etc.....-.--------------+-++++2-0---> 2.14 3. 49 2. 28 4, 32 3.14 2.77 2. 61
|
22 23 24 25 26 27 28
SiO pee ac ase ae aay ete <meta (siecle ioe a= \S' 0.14) 0.04 0. 02 0. 40 0. 06 0. 07 0.13
(Al) Fe).03- ..------ 2-2 -2c 2-222 =a rere rent ete seo 05} . 09 . 02 .53 pp lit 15 . 06
RA) hele ce eee ea eteeemeitetate ala cinloine See idmle Sim(m =16/6 . 04 mali, .18 . 20 .15 Lt wok
(Gi 10) ene eee) Bee Se Reenter nee 53.34] 52.12) 53.84| 53.22 | 58.25 | (02.94 53. 69
Oi eRe Coen = aia eee (2) (?) (2) (2) (?) (2) (?)
TOKO Ren BOAR So cee qt Seni SaBSa oe oC cur ooo aan Trace. | Trace. Trace. | Trace. | Trace. Trace. Trace.
Tenition.§- ...- 2-22-2220 win nein tro 44.81) 45.84 44.98) 44.31 44,80 | 45.28 | 44. 64

98. 38 98. 26 99. 04 98. 66 98.37 98. 61 98. 79
CO, needed ....--.------------------++---- Rometa meee 41.95 41. 14 42.50 42. 04 42.81 41.79 42. 48
Organic matter, etc.......-.----+-----+++-2e6 see 2. 86 3.70 2. 48 D527 1.99 3. 49 2.16

8 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES.

The two other analyses of corals, which were made some years ago in the Survey laboratory,
do not appear in the foregoing table because they are stated in somewhat different form. They
are, however, included in the follow ing table of reduced analyses, in which the figures are recal-
culated 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.

| |
| 1 2 3 4 5 | 6
be ee A Bh | es Bieta Jou hk
STORE Ree ee UE Mea cme she co ac Spee EAB Hoi cacao see 0. 10 0. 44 0. 10 0.18 0.12 | 0. 29
UAT NS) OS coi aetreee eien Ae Fe Sea A BAGS ieee 74 37 pa) v2 oti -12
TCL keh. SOR Rete a, MER So NI eis s 2 Teh 87 43 48 _ 82 | 89
(6/10, OR oat eeieiig Bomeaei ae So ictopaecaccomomoec -ormgEe= 98. 05 98. 32 99. 35 99.27, 99. 95 99. 20
WAS Ose | Ce eae ae nine oes erate are ee Been oh< SE oe (?) . 00 (?) (?) (2) (?)
(Gas P ORs ete not eee See se cee ore eee oot eee Trace Trace. | Trace.| Trace. | Trace Trace
100.00 100.00 100.00 100.00 100.00 100. 00
7 8 | 9 10 11 12
= | — = am =
SiOie ab 562 oe ese cere tes eos Sats See ana eee ree 0. 00 0. 07 0. 31 0. 04 0. 00 0. 21
(UNIKIEE)}( 0425 Sates SSotaes Sa UeSS re EOtes scree oe. cau | . 04 05 08 . 00 03 628,
INI OO Seton yn So aap Ran abaGobs coat auaranbeneEose 46 41 sali . 61 | 59 . 63
(CX 00 FE oe eae aaa sao Ieee AOR SES op see eeided cana | 99.50 99. 47 99. 50 99.35 | 99.38 98. 71
(CHIS\O), Sa Seles oet Sls BRR eee Sa cbbeaaric sscckya ceases a) (2) (?) (?) (?) allih
(CAP ONS Gne cae lo Se lobe Beers Padunme e nenao qedaas aos = | ‘Trace Trace Trace. 00 . 00 Trace.
/ 100.00 100.00 100.00 100.00 100.00 100. 00
13 14 15 16 17 18
SiO So areas Sete See Eee SBR as Sas hon one 36 0. 09 0. 20 0. 34 0. 04 0. 22 0. 00
CATEREO yo eee ae See Be aoe oe .12 07 07 14 | 26 | . 06
MglO, Faye oro soa conan ann So OC DOD Une eEaseae co soaee nc Ry) .14 12 . 76 | . 54 . 45
Ga@Osns 24: eet sees eee ee eee eee eee es ae eae 99. 42 99. 38 99. 47 99. 06) 98.98 99. 49
(OLNS\ OS eeia ae eh pg NRE Et A A ot I hth a 3 (?) 21 (?) (2) (?) (?)
CaP,’ 0. FES Sios Sones ao See ao sane eneenaanoseees = Trace Trace. | Trace. | Trace. | ‘Trace. Trace.
100.00 100.00 © 100.00 100.00 100.00 = 100. 00
‘ , |
19 20 Aik 22 23 24
SiO Hehe. Wb S ager Sas acoke door sero os Sa smocrinee acer ¢ 0. 10 0, 02 0. 00 0.15 0. 04 0. 02
(AI e Ose Soar et A) eae Seem eat oy eee 00 . 04 . 04 . 05 - 10 . 02
MoO Os ae ecco nr menete tee rele ie om cietetbe = eininieetnn tet e e eels hd .57 . 09 sot . 40
Cal 105. Be ao es aoe Oa OOS: OU Sc Ore ae Oo flamers 2S 99. 13 99. 21 99. 39 99. 71 99. 49 99. 56
OaS@ ateie ene ace ee ce eae een nce Seo rae eee (2) (?) (?) (2) (?) (2)
(OFS 0) = eeu gences ge anQn aber As aSeas4 some Berson eS - 00 .00 | Trace Trace Trace Trace.
100.00 100.00 100.00 | 100.00 100.00 100. 00
|
25 26 27 28 29 30
S| —
3) 0 ne Ae SEA ee ce er Ot IO Sac e asec 0. 41 0.06 | 0. 07 0.14 | 0. 23 1. 25
(Ale) 0529-28. 2a ee SR oe eee ae . 5d ally 15 .06 | Trace. . 59
MpGO,. os. fs. oe ok cast Le 2 eee 43 . 33 Ba7) _59 42 59
CAO O eee see oe costes wine Seiaietetelel= = lo ernie eee leer 98. 61 99. 50 99:41 | 99.21 99. 35 97. 57
CORSO foes ec Bets cooler ee RO Sc (2) (2) (2). 2) (2) (?)
(Of 55 20 Hes Es ea RA Sato! Soo oct aac ae TO aoseoe Trace Trace. | Trace. | Trace. (?) (?)
| _s_| =
100.00 100. 00 | 100. 00 | 100.00 100, 00 100. 00

ALCOYONARIAN CORALS. 9

These analyses are remarkably concordant and show that the stony corals contain little
besides calcium carbonate. Silica and sesquioxides are probably extraneous; magnesia is alto-
gether subordinate, although fairly regular in its amount; phosphates occur only in traces. The
older analyses all tell the same story, except that in six analyses by S. P. Sharples’ of corals
from the Gulf Stream from 0.27 to 0.84 per cent of calcium phosphate was determined. In
five analyses of Brazilian corals, L. R. Lenox * found from 0.42 to 0.54 per cent of magnesium
carbonate, quantities like those appearing in our tables. In short, the uniformity of the data is
so marked that it is unnecessary to reproduce all the published analyses here.’

ALCYONARIAN CORALS.

The aleyonarians, which include the red corals, the gorgonias, or sea fans, and other similar,
generally branching forms, differ from the madrepores in being notably magnesian. To this
rule one exception is known, the genus Heliopora, which stands quite alone and resembles the
ordinary corals not only in structure but also in chemical composition.” With this exception,
the alcyonaria are chemically similar to the crinoids, but some genera are much richer in phos-
phates. In some respects the aleyonarians are difficult to deal with analytically, for many of
them have a horny axis, composed chiefly of organic matter, surrounded by a cortex or envelope
which is largely calcareous. These two portions of the structure are so unlike in composition
that imperfections in a sample taken for analysis may lead to very uncertain conclusions.

The list of aleyonarians analyzed is as follows:

1. Heliopora cerulea Pallas; blue coral. Southern Philippine Islands. The blue color consists of organic matter.!!

2. Tubtpora purpurea Lamarck. Singapore, Straits Settlements; latitude, 1° 20’ N.; longitude, 103° 50’ E.

3. Corallium elatior Ridley; ared coral. Murotsu, Tosa (Shikoku), Japan; latitude, about 33° N.

4. Primnoa reseda Verrill. East of Nova Scotia; depth of water, 366 meters; latitude, 42° 16’ N.; lorigitude, 63° 15’ W.

5. Lepidisis caryophyllia Verrill. Off Nantucket Shoals, Albatross station 2037; latitude, 38° 53’ 00” N.; longitude,
69° 23’ 30” W.; depth, 3,168 meters; bottom temperature, 3.3° ©.

6. Pennatula aculeata Dana. St. Peters Bank and Banquereau; Albatross station 2470; latitude, 44° 47° N.,
longitude, 56° 33’ 45’ W.; depth, 410 meters; bottom temperature, 4.5° ©.

7. Paramuricea borealis 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 carneum L. Agassiz. Albatross station 2468, off Newfoundland; latitude, 45° 11’ 30” N.; longitude,
55° 51’ 30” W.; depth, 70.9 meters; bottom temperature, 0.5 °C. This analysis, of very pure material, replaces the
unsatisfactory one given in the former edition of this paper. Analyst, R. M. Kamm.

10. Gorgonia suffruticosa Dana. Fiji Islands. Cortex and axis together.

11. Gorgonia acerosa Pallas. East end of Long Cay, Nassau, Bahamas; latitude, 25° 5/ 6” N. Cortex and axis.

12. Gorgonia acerosa. Caesars Creek, southern Florida; latitude, : 23° 30’ N., approximately. Cortex alone.

13. Muricea humilis Milne-Edwards. Parahyba do Norte, Brazil; latitude, 7° to 8° S.

14. Muricea echinata Valenciennes. Cape San Lucas, Lower California; latitude, 22° 52’ N.

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

16. Ctenocella pectinata Valenciennes. Torres Straits, Australia; latitude, about 10° S.

17. Xiphogorgia anceps Pallas. Caesars Creek, southern Florida; latitude, 23° 30’ N., approximately.

18. Rhipidogorgia flabellum Linné. Bermuda; latitude, about 32° N.

19. Rhipidogorgia flabellum. East side of Andros Island, Bahamas; latitude, about 25° N.

20. Leptogorgia pulchra Verrill. La Paz, Gulf of California; latitude, 24° 26/ N.

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

22. Phyllogorgia quercifolia Dana. Fernando de Noronha, Brazil; latitude, 3° 50’ S.; ESS 2° 25! W.

7 Sharples, S. P., Am. Jour. Sci., 3d ser., vol. 1, p. 168, 1871.

§ Lenox, L. R., mecesed Coll. Mus! Gon: Zool. Bull., vol. 44, p. 264, 1904. In a memoir by J. C. Branner on sandstone reefs.

9 See also Silliman, B., jr., Am. Jour. Sci., 2d ser., Sai 1, p. 189, 1846, for 31 analyses of corals from the South Pacific and the Indian Ocean;
Forchhammer, G., Neues Jahrb., 1852, p. 854, on magnesia in shells and corals; Nichols, H. W., Field Columbian Mus. Pub. 111, p. 31, 1906; also,
for determinations of magnesia. In Calaria dadalea from Abyssinia Nichols found 0.35 per cent of MgCO,. A. G. Hégbom, (Neues Jahrb., 1894,
Band 1, p. 262) in two Bermuda corals found 0.36 and 0.62 per cent of MgCOsz.

10 See Moseley, H, W., Philos. Trans., vol. 166, p. 91, 1876, on the structural relations of Heliopora.

' See Moseley, H. W., op. cit., and Bourne, G. C., Philos. Trans., vel. 186, p. 455, 1895.

/

10

THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES.
.

Analyses of aleyonarians.

1 2 3 4 5 6 7 8
SOs tee Bie sae See. £5 ace aE 0. 14 1. 36 0. 00 0. 09 0.11 1, 21 0. 26 0.13
(GAIT 0) he eee ey CaO We SSE 07| 55 1 MoCte 205 |, 72 en 18 02
5, O een eae eee ene t aol Si jh ole 16 5. 67 5. 44 2.15 OG 2. 62 2020 Sui
CRORE SSE oe Ob cSt OE es say 8 ee A ea ed 54) 42 46. 60 48. 65 37. 76 49. 87 35. 67 29. 92 43. 47
SOjoot Jes ivstettas change se sa eRe ee neeaes 29 . 68 77 Rat eye . 35 1. 64 1.10
PIO recente <= seme icc ae «sce ete eee Trace. | Trace. 18 . 28 | Trace. 1.02 . 39 222
Peniion sos cc sce aeea es ee eee eee Ieee 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
CO; mead ed artec\-2eto= te a Ae ee eaten 42.78 42.48 42. 87 31. 38 42. 43 29. 76 24. 74 37. 44
Orzanic matter, ete... saceserep tess ae see 1.55 PRA | 3.14] 26. 20 3.27 | 29.50) 41.58 13. 04

9 10 11 12 13 14 15
SO pa s3ssac See: ate ee ae ate ose eb es 0. 88 0. 26 0.13 0. 03 0. 44 0. 09 0. 38
(AUR e) SO ge oan een ne se byte opis aeemt teeaeeie Trace. 3118} 13 13 - 06 - 05 -13
I OE acy ie seiaeot de, wh Male AES A ee eS le 1. 86 3. 03 3. 54 3.51 5. 25 4.58 5. 61
CaO ssa. ee FE RE su tie the get ee 29. 92 22. 32 28. 73 26. 48 42.14 38. 04 40, 94
BOrt oo is ee ee ie eo Wiehe mer tee 74,| 1.51 “68| 1.33|. .89| %35 | "Trace.
PIO: MCRAE, WEE VOR Nee At MT. Yat 1.43 -10 -99 273 124 30 | Trace.
Jomitions 5-25 eeeeian te ee ean Bao et SERA Ne 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
WOFmeed Cd paar sae onary Se pe: Be eed 23. 88 19. 94 25. 17 23. 26 38. 13 33. 91 38. 34
Orpanicimnaltersciesee sases cl: So Saas ate ee a kee 40.90 | 51. 52 iy td 40. 23 13. 43 20. 18 14.15
The low summation 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 SO,. Applying these corrections to the analysis the summation becomes

99.96.
16 17 18 19 20 21 22
ILO) pees ec te iataeise > o/s Benes OF Ce one 2t ee OC 0.17 0. 08 0.12 0. 16 0. 06 0. 21 0. 26
(CAUSE) 5 Onc eee Fs irs Seah ete RS TE. A agi) . 04 .16 . 06 - 02 l6 .19
No OAR Rey ae ee ete Le a RT el 5. 90 3. 52 3.42 4,22 4. 60 5. 14 5.59
CaO ari s5 J SOs eale cs ect Ce, eR R ae ne oo eee eae ae 37.05 | 27.19 | 27.41.) 32.23 | 33.59) 36.15 34. 62
SS ease a tere ie rel Se eee ee he een eee cle 79 1. 28 . 80 ieaea < TW 93 293
SO) at ae mR ROG El eek a. ge 32 51 . 28 286) O*67HamONTs 2.97
TRON ere he ok nr nt ee te ee ee 56.15 | 65.56 65.67 | 62.10 | 55.89] 55.39 55. 28
ss 100.48 | 98.18 | 97.86] 100.80 | 98.04 | 100.76 99.84
COnmeedediek -oi ec SoBe hoe ee ye ean ee 34. 87 24. 07 25. 60 28.51 28. 31 30. 96 34.14
Orsanie matter"etelst-sac.sascste ose ee ose eee 21.28 | 41.49 | 40.07} 33.59 | 27.58) 24. 43 21.14
The reduced or rational analyses are as follows:
Reduced analyses of aleyonarians.
at 2 3 4 5 6 7 8

DLO Ga eeakt ais hae See eee oe Renee 0.15 1. 40 0.00 0.13 0.11 1.70 0.44 0.15
GAISHG) SO poe Sons see Ree ee LA ee 07 .o7 15 88 | .05 1.01 - 30 | . 03
Me OO ie 2-355 Ri gtely thos. 2.sne nee ee $55} 12523) TIS6 61819 16.92 | 7a e803 9.05
CaGO et. tino cee ee eee tenn Coe ee 98.93:\| 84.61) 86.57) 90.39 | 92.24 | 85.62) 85.11 88. 04
(GETS | OF et ae echt em ofr ip ee ana ey ee ve | 50 1.19 1.32 1.59 | 68 | 84) 4.69 2a,
G16) (0 ee RN haere at aya UM Ua 3 We Trace. Trace. 40 .8a | Trace.| 3.12 1.43 - 56
| 100.00 100.00 100.00 100.00 | 100.00 100.00 100.00 100. 00

ALCYONARIAN CORALS. 11

Reduced analysis of aleyonarians—Continued.

9 10 1 12 13 AS eke
SLOT ae - SAAB nn he Sc Or ee 1.50 0.55 0. 22 0. 04 0. 50 0.11) 0.45
(@aWISING)),(0)+., Spa Oo SOE COE AOS See eee Trace. . 28 | 22 .24 “IY 06 | .15
IM os Onan eee ee Se need ne scan cle eaieesceie 6.66 | 13.48] 12.52] 13.29| 12.64] 12.28] 13.79
INGO) 5. 3 Caan SCS Sen OCB R Ee a ia 84.50 79: 84 81. 45 79.48 84.47 | 83.79 | 85.61
aS Oe oes ala Stctere Sie epee wiarcle.s <isfalecis ities occ eb cie ss 2.15 5.43 | 1.95 4.08 1.73 | 2.93 Trace.
(ORTHO) = Oud SSC Raa a YE Se Sento Se 5.19 .47 3.64 | 2.87 . 54 .83 | Trace.

100.00 100.00 | 100.00 | 100.00 | 100.00 | 100. 00 100, 00

16 7) LS TOAD 20 21 22
|

aes =e | =e —
SH Oe OS pee RP ret hE ee co Sone ae 5 ae |} 0.21 0.14 | 0. 21 0. 24 0.09 0. 28 | 0.34
(CRIARO) 1 O pegetece secides nade teeta Se eae cle sects a0 .13 .07 | ava} .07 . 03 aA - 26
IMG OO) a waet eas sas eins ee ain science eepeeereeicioncies oainis coms 15.65) 13:04 | 12.64) 13.19 | 13.71 | 14.18 15. 73
(GINO OR 47 PE pCR t Eg be ae clk Ute eee ee 81.44 80.96 83.38 | 80.75 | 74.99 75.36 72.99
CASO fae a laeicd an asiewese oe Cee se S-Bagascoese Berne | 1.69 3. 83 2.40 2.95 2.91 2.07 2.11
(CE DMO) ae es ma na Ge || aaa BB.f) 115 96) 115 09 | P.D8g08l|e leBeBT 4-795 8.57
| | —

100.00 100.00 | 100.00 | 100.00 | 100.00 100.00 | 100.00

Analysis No. 1, that of Heliopora, might be an analysis of an ordinary coral, being nearly
“nonmagnesian and hence different from 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 also studied
and to better advantage. The three analyses are as follows:

Gorgonia Rhipido-

-
| Gorgonia |
acerosa. suffruticosa. gorgia?
| Ev sees
[ie Si@imeeemedte. ° Agente ee ek 2c 0.04 0.39 0.07
f)) CAMHS Orete ts ALINE ay oo. pO! . 06 Trace
720) <i Sr page el a soak | “80 1.03
CCL. os ae ee ee O53 | 1.10 | 50
1F2(0)s oe ee ee nol Trace. 15
Sj: Je So Gages s be aeOe eee 94 1. 65 1.90
emi enie55 ce saeseces= 125 96. 45 94.39 95. 83
99.08 | 98.39 99. 48

The axes, evidently, are composed largely of organic matter, and the inorganic portions
differ in composition from the more abundant calcareous envelopes. In two of them magnesia
is in excess of lime, 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 specimen of Leptogorgia flammea Verrill, from 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 pro-
portion 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 and
calcium 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° 45’ N.; longitude 20° 12/ W., about 160 miles southwest of

the Canary Islands; depth 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 CONSTITUENTS OF MARINE INVERTEBRATES.

2. Aleyonium palmatum. Probably Mediterranean.
3. Corallium rubrum; the precious red coral. Mediterranean.
Analyses 2 and 3 by O. Biitschli in K. Gesell. Wiss. Géttingen Abh., No. 3, 1908.

Older analyses of alcyonarians.

it 2 3
CaGOst:...: eee ie eee 93. 39 86. 25 86. 78
ee © a sSle a Re aes Se 6. 00 9. 38 8. 97
"Phosphate 73-3. cet aes. yee 1.00 70
ag Ea Oe = ate emeee } “0a ee
Ce Se OOS Os te OPO Ed iE
CASOLOH.O 00. aes 1.15 1.55
NiO fonjsand= se. nse eee Trace WRS6 j|ccsstcosmee
Oke atc se aks Tene 30 82 1.46
Orpanic matters 25 arene clo: ee nou | . 06
Imbeluble ec: eaae;- sae Seeeeee Bec nlU/3) el he oe eee | S2ek: setae
99. 84 100.33 | 99.32
Ne roles) |
J The reduced analyses, rejecting obvious impurities, are as follows:
Reduced analyses of alcyonarians.
it! 2 3
S10, 233s. SA ee | Mracel.|eeece esse (es50set shes
Hes Oa eee sce eS- 2 ste estes (2) ohry eR: 1A cn ok ae
NIE CO eee are ap eer eee 6.03 9. 63 9.18
CaCOi 3. 0... ee tae Ot 93. 87 88. 42 88. 84
CaSO ponder emcee ees ee eee 93 1. 26
(OF SO Ba Aegis spn .10 1.02 72
100. 00 100. 00 100. 00

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

In a worked specimen of Corallium rubrum, H. W. Nichols found 9.32 per cent of magnesium
carbonate, not far from Biitschli’s figure. In Eunicea tourneforti and Plezaurella 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 reduced analyses. Three similar
analyses by G. Forchhammer ” are also recorded, of Tubipora musica, Corallium nobile (= C.
rubrum), and Isis hippuris, which gave, respectively, 3.83, 2.73, and 6.36 per cent of magnesium
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 Biitschli. A still earlier analysis of
red coral by Witting * gave 3.50 per cent of magnesium carbonate. The specimen analyzed,
however, contained about 12 per cent of impurities.

If, now, we arrange the alcyonarians in the order of ascending magnesium carbonate, a
remarkable relation connecting composition with temperature appears. Heliopora, being
anomalous, is not included in the table. The three analyses by Anderson and Biitschli 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.

% Forchhammer, G., Neues Jahrb., 1852, p. 854. 13 Witting, E., Annalen Chem. u. Pharm., vol. 1, p. 119, 1832.

ALCYONARIAN CORALS. US

Magnesium carbonate and calcium phosphate in alcyonarians.

Species. Locality. | Latitude. Ca,P,0,. | MgCO,.
Pleurocorallium johnsoni.......-.---------------- Atlanties!.: 2 (Sek sens 2b Da NESS ..2 0.10 | 6. 03
Primnoa reseda.......-.------.+---+--+--2+-+---- Noval Scotia. sc... 325 smiais ADAG fa oe . 83 6.18
Aleyonium carneum........--------------+---+--- Newfoundland.....-.... apo UUNne ene 5.19 6. 66
Lepidisis caryophyllia Off Nantucket.......... | 38° 33’ N.....-. Trace 6. 92
Pennatula aculeata.........-- Atlantic: <2 -...<jaep smote | Ado AION 5. -4- 3. 12 ra!
Paramuricea borealis... ...-. Grand ‘Banks: -- 22> -2s—<|=aeeeereee a=" r= 1. 43 8.03
Paragorgia arborea....-..-------------+----++++---- Nova) Scotias.- os ccc ace seamen raat . 56 9. 05
Corallium rubrum....---- Seas Caan Renan CoC Mediterranean........-. les sown saces 72 9.18
Alcyonium palmatum.........-..----------------- Mediterranean ?_ 3: 74-pec}i-<-=aeeeeer es 1. 02 9. 63
Gorallitim elatior... ....-:2----- 0562-2 ---25------- Japan teere - 5 6 faeeseas S3o NE eid. 3s: = . 40 11. 56
Tubipora purpurea. ......----.---+-----------+-+++- Singapore.....-..--...-- To-20GNigaeene. = Trace. 12. 23
Muricea echinata...........--.---2---------+----- Cape San Lucas. ..-..---. 22 °1b2ONE oo ase . 83 12. 28
Gorgomia acerosa.........-------02 +2222 eee eee eee Bahamasss-=<-<.2s-ee Ob OPH NaCe oars 3. 64 12. 52
Muricea humilis... ..0.-..-..2.-2:25<-2-+2-------2 Brazilgets shh aeae ses (ERs ase Poe 59 12. 64
Rhipidogorgia flabellum........-.---------------- (Bermiud ats. h. 4a sea See Nesee seeers 1.09 12. 64
Xiphogorgia anceps.....--.-----+-+-----+-+-++-+++-- Mloridatase= <= yee eee Doe OU AN aeeeter 1. 96 13. 04
Rhipidogorgia flabellum.....-..-.------------+---- Bahamas....---..------- 25 Ni axecaacee 2. 80 13. 19
Gorgonia acerosa....----------+--+---+--+2eeree eee lorid 93 - s2-- == 2 4 WPA RIEIVAIN SBS see 2. 87 13. 29
Gorgonia suffruticosa.......-----------+-+-----+---- thi) Seeee en Pepe ee conn Rees csatencctcee 47 13. 43
Leptogorgia pulchra....-....----------------+-+---- Lower California......- - 2AOEL BON ae ces 8. 27 is Hyall
Plexaurella grandiflora. .....-..-.----------------- iBiyvs I EER See OPS RG Aeris odeceoa sar Trace. 13. 79
Leptogorgia rigida........------.------++--++++--+-- Cape San Lucas. .....-.- 222120 Sesinces 7. 95 14.13
Ctenocella pectinata..-........-------+-++----++--- Torres Strait.......--.-- LO Se sseeeeses . 88 15. 65
Phyllogorgia quercifolia...........--------+------- 1g Beene seco ocr 331007 Nieee ee === 8. 57 | 15. 73

Since the foregoing table was compiled Prof. L. R. Cary, of Princeton University, has
kindly sent me a series of unpublished analyses of aleyonarian spicules from specimens collected
in shallow reefs around the Tortugas and southern Florida. The spicules were separated from
organic matter by treatment 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 aleyonarian spicules from the Tortugas and southern Florida.

l l

Species. CaCO;. | MgCO,. | Ca,P,0,.| Sum.
Plexaura flexuosa.....----- 80.87 | 16.90 0. 35 98.13
Plexaura homomalla. --.-. 86. 29 12.01 . 04 98. 34
Plexaurella dichotoma. -... 85. 90 14.12 09 100. 11
Eunecia crassa...----.----- 87. 27 11. 46 -09 98. 82
Eunecia rousseaui.....---- 86. 76 11. 45 12 98. 31
Plexaurella sp?......----- 86. 75 12. 08 15 98. 98
Briarium asbestinum. --.. - 85. 13 13. 21 Aas} 98. 47
Gorgonia flabellum......-.- 86. 04 13. 12 44 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 99. 02
Xiphigorgia anceps. - - ---- 84. 91 13. 23 . 26 98.40 |

These analyses admirably confirm the analyses, made in the Survey laboratory, of aleyo-
narians from the same region and the Bahamas. In addition, Prof. Phillips made a partial
analysis of spicules from Aleyonium rigidum from Samoa, in which he found 10.50 per cent of
magnesium carbonate.

Although records of temperature and depths are available for only a few of these alcyo-
narians, the suggested relation is clearly evident. The organisms from cold, northern waters or
from very deep waters are low in magnesia, and those from warm regions are high. The same
relation appears in our analyses of echinoderms and is unmistakable. It is not rigorously exact,
but some apparent 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 CONSTITUENTS 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 aleyonarians may form
two or more distinct series that are not perfectly comparable in chemical composition. Coralliwm
and Tubipora, for example, are compact forms, with little organic matter and lower magnesia
than the genera with horny, organic axes, such as those whose names appear at the end 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 belong-
ing to the genera Millepora and Distichopora. The species and localities are as follows:

. Millepora alcicornis Linné. Shoal water, Tortugas, Fla.

. Millepora alcicornis. Bermuda.

Millepora braziliensis Verrill. Candeas, Pernambuco, Brazil.

Distichopora nitida Verrill. Micronesia, exact locality unknown.

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

. Distichopora sulcata Pourtalés. Off Habana, Cuba; depth of water, 143 to 179 meters.

Analyses of hydroids.

oor Wh re

1 2 3 4 5 6
SiOit Sees sae CoA tt BR Ee eee ~2 0234558 0. 23 0. 02 0.09 0.10 0.09 0. 07
(AUSBG) 50) ssp sees ee: = -sv eee eee an ee eee . 10 .07 . 06 . 20 .07 .05
Mio (See Fone Or ene ho. 2 nese Ree Lee Rete lew © oe e ee 43 ot) . 59 pill +12 -12
: 5 53. 14 53. 36 53. 07 53. 43 53. 90
04 1.03 . 69 65 61
Trace Trace Trace Trace Trace

GO; needed ').\- 2 20). vo SR OG eee 41.57| 41.85| 42.01| 41.44] 41.76 42, 15

1 2 3 4 5 6

on = ee 0 a |e pees | 8 22%
0. 02 0.09 0.11 0.09 0. 07

.07 . 06 $21 | .07 .05

P22) 1. 28 . 24 . 26 . 26

99. 63 96. 77 98. 22 98. 43 98. 56

. 06 1.80 a 22) 1.15 1.06

Trace. Trace. Trace. Trace. Trace.

| 100.00 100.00 100.00 100.00 100.00 100. 00

A few partial analyses and one fairly complete analysis 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. Hégbom “ found respectively 95.86 and 94.39
per cent of calcium carbonate, with 0.41 and 0.97 of magnesium carbonate. In Mullepora
alcicornis from the Gulf Stream S. P. Sharples ** found 97.45 per cent of calcium carbonate, 0.27
of calcium phosphate, 2.54 of organic matter and water, and only traces of iron and magnesia.
B. Silliman, jr. in one analysis of the same species reported no magnesia, which evidently

4 Hégbom, A. G., Neues Jahrb., 1894, Band 1, p. 262. 16 Silliman, B., jr., Am. Jour. Sci., 2d ser., vol. 1, p. 189, 1846.
18 Sharples, S. P., Am. Jour. Sci., 3d ser., vol. 1, p. 168, 1871.

ANNELIDS. 15

was not sought for. A single analysis of M. braziliensis by L. R. Lenox" gave the following
composition:

CACO empresas ncte ce wees ails: fl ts Fe ead e chs hehe ceneeeistes aleciimele se cc ceece 93. 80
MES Feerertatatete estate tetarats ota a/Siaiajalm ola c/aieiai= = as lals wis v0 So pls sata eeaelets elem arate ale)aiaeta minle/ainls «i='</=\o 0's 2.14
(QANO)...- cor eadn cog et ec dauecoGHo > Coe a BEB ESeEEBES 6 cobE ne aeRO E EO OSb ACCOR codop Cae GoCp po eAere 2.08
STO. ss se oer nan co dda be ped UDO SUC Sepa e DE BoeEae o Sade ooh thins eel aaee dae Geer e messes ence. 03
(ALLE) OF ca ncagn oS dee Don OSC COS Sete Be Ee aBEenCSrron canon cepesece cn: ae. Coane. eee ane 07

98. 12

It is interesting to note that the Brazilian species is the richest of all in magnesia, although
it is poor in comparison with the echinoderms and aleyonarians. As reef builders the mille-
pores contribute little to the coral rock but carbonate of lime.

In an analysis of Millepora cervicornis, by A. Damour,'* 8.51 per cent of magnesium carbonate
is reported. The name, however, belongs to a fossil species, and the specimen analyzed was at
least partly fossilized. Its locality is given as Bréhat, Manche, France.

ANNELIDS.

The curious tubes formed by marie 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. Filograna implexa Berkeley (Serpula complexa). Scarborough, England.

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

3. Hydroides dianthus Verrill. Vineyard Sound, Mass.

4. Leodice polybranchia Verrill. Off Marthas Vineyard, Mass.

5, 6. Hyalinecia artifex Verrill. Off Marthas Vineyard, Fish Hawk station 1025; depth of water, 384 meters;
bottom temperature, 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 worm tubes.

1 2 3 4 5 6
— == = 4 ———— = —
Tesco) Wed} ig bol 3 KG) eee ee St es eee ee COE Pe BEES ae 3 epeebose: NSE co acSe BScu ee sae Pee oes
BO OAR e <a Seem oc GUGR MEINE Sr Si tenet ae 1.12 0. 75 15. 45. 28. 05 0. 20 0. 24
GNIS Os ee. IE he aE SON 75 41 The Wi! BkBE 07 01
POMS RO: Ne i, Be os a gt 00 14) Shben lec 8.57 8.17
COON Sa OEE See MPR, te ERR Cy eR 35.98 | 50.89] 39.11 5.12] 5.35 5, 24
SO ea ie gn eae ela tooo) 2s SeEINS as eios es weccamcae cute Trace 07 | ‘Trace. 6.06 | 3. 47 4.33
TEL 0) es So DCE EA IOS OIE CO = COO REE OO oeaac eeee .29 | ‘Trace. 22) | 6.43} 20.72 20. 32
TG, 2 cA ee ae =, 5s a Re | 31.23| 45.99] 38.82] 46.91] 61.83 61. 41

|

(ees) 98.25 | 98.48 99.30 | 100.21 99. 72
OGreadagt amy ste... Pe Ns eel oted DESOO')|): +39. 95MIMAIS4! dqd| a mepemeel et rie) Alita
Qroanichmatten eter ...... sseeeers sss. <3 seaeeec ccs | 3. 23 6. 04 | 451361 | Saeeseee tal acie'e'ciaie:<ja) sates s 3

Tests for fluorine failed to show its presence in any of these tubes.

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 worm tubes.

1 2 | 3
i cl
Cy 0.00 0.32 | 9.72
COO} Barbee aaa 99.01 99.55 | 89. 66
(ChSO) 5 SaaS e ean toc eee eee Tracesni|m 13 Trace.
Wa be Oateenacaas se cdas caces «2 She) Trace. | 62
100. 00 100.00 100.00
Lenox, L. R., Harvard Coll. ieee) Comp. Zool. Bull., vol. 44, p. 264, 1904. 18 Damour, A., Compt. Rend., vol. 32, p. 253, 1851.

106135—22 2

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 aleyonarian or an echinoderm. In all three the organic matter is very low. Three other
partial analyses by Forchhammer are on record, which may be compared with these; although

only magnesium carbonate was determined. These are—
Per cent MgCOs.

Serpulasp: /Mediterrancan-:.\ <2 42 thenn sn 5: Gee 2 ee ene. ©. <3 See eee 7. 644
Serpula triquetra.. North Sea-i-lecn ete eee): =. ee oe eee eRe oe = 5 =e ee ceR eee 4.455
Serpula filograna es. .ssiaec a clos Bes Pee nee ae OEE ee. wo > 5 CoE ERE ere 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 Hyalinecia, being highly phosphatic, are difficult to
interpret chemically. The bases are insufficient to satisfy the acids if the phosphoric oxide is
assumed to represent the normal tribasic salts. Phosphorus may be present as 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 the reduction of analyses 4, 5, and 6 in abey-
ance until more evidence can be obtained. In this direction an analysis of Onuphis tubicola
Miiller, by Schmiedeberg,”! is suggestive. To the organic matter of the tube of Onuphis, “ onu-
phin,” he assigns the formula C,,H,,NO,,, and the tube itself he regards as a complex compound
of the composition

C,,H,,NO,, + CaHPO,+ 4MgHPO, + 22H,0.
The analyses given by Schmiedeberg are as follows:

Analyses of Onuphis tubicola.

Entire tube. Calcined ash.

Qnuphine seks eeeciee Ace. - eect e eet Seco e meer Bye lsh ||| JEN 0) alae PSR E Socata ccdcosaaesuassoe 57.20
Waterss 2S aeee ens Dee oe Sateen eee ZENO TANINCROME Sos. 2... 2 S242 Bae ee eee eee Ee eee eee 12.48
1X0) ae ete ee | aed 3 PATS) 20) ARERR 25. 02
MPO ez = Fe rere isals Semen a areca © ace eee oer O riGHl RG s oes foc cs... odacces seas se eae ease EEE ee 1.96
OS Ob net 2.2 Be Wc coe ea ee oe eee S4OWWNaws 2-22... .. . esas sare eee eee 50
Hy(acid) Settee een m2: epee eer ae eee eee 2a OOsgoLO>; CO,; anid Losd:a..-<ces seeneeee en eee ones PAE

100. 00 |, 99. 43

In a general way this analysis of Onuphis resembles our analysis of Hyalinewcia, 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.

19 Forchhammer, G., Neues Jahrb., 1852, p. 854.

20 In a later memoir (Philos. Trans., vol. 155, p. 203, 1865) Forchhammer gives the percentage of MgCOs; in Serpula filograna as 13.49 per cent.
W hich figure is correct?

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

THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. ity

ECHINODERMS.
1. CRINOIDS.
RECENT CRINOIDS.

In 1906 H. W. Nichols ® published a number of partial analyses of marine invertebrates,
and in a crinoid, Metacrinus rotundus from Japan, he found 11.72 per cent of magnesium
carbonate. This analysis attracted the attention of Austin H. Clark, and at his request two
other analyses of crinoids were made in the laboratory of the United States Geological Survey
by Chase Palmer, who also found that they contained abundant magnesia. These analyses,
which were published and discussed by Mr. Clark,®* 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 specimens 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
magnesium carbonate in notable proportions but varying in a most remarkable manner. The
data obtained are in detail as follows, beginning with the list of the specimens studied:

1. Ptilocrinus pinnatus A. H. Clark. Albatross station 3342, off the Queen Charlotte Islands, British Columbia;
latitude, 52° 39’ 30” N.; longitude, 132° 38’ W.; depth of water, 2,858 meters; bottom temperature, 1.83° C. Mean of
two analyses.

2. Florometra asperrima Clark Albatross station 3070, off the coast of Washington; latitude, 47° 29’ 30” N.;
longitude, 125° 43’ W.; depth, 1,145 meters; bottom temperature, 3.28° C.

3. Psathyrometra fragilis Clark. Albatross station 5032, Yezo Strait, Japan; latitude, 44° 05’ N.; longitude, 145°
30’ E.; depth, 540-959 meters; bottom temperature, 1.61° C.

4. Pentametrocrinus japonicus P. H. Carpenter. Albatross station 5083, 34.5 miles off Omai Saki Light, Japan;
latitude, 34° 04’ 20” N.; longitude, 137° 57’ 30” E.; depth, 1,123 meters; bottom temperature, 3-09> (Oe

5. Capillaster multiradiata Linné. Albatross station 5137, Philippine Islands near Jolo, 1.3 miles from Jolo Light;
latitude, 6° 04/ 25’ N.; longitude, 120° 58’ 30” E.; depth, 36 meters; no temperature record.

6. Pachylometra patula Carpenter. Albatross station 5036, Philippine Islands, North Balabac Strait, 15.5 miles
from Balabac Light; latitude, 8° 06’ 40’ N.; longitude, 117° 18’ 45” 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, 9° 37’ 05’ N.; longitude, 121° 12’ 37’ E.; depth. 612 meters; bottom temperature, 10.22° C.

9. Parametra granulata Clark. Albatross station 5536, Philippine Islands, between Negros and Siquijor, 11-8
miles from Apo Island; latitude, 9° 15’ 45’ N.; longitude, 123° 22’ E.; depth, 502 meters; bottom temperature, 11.95° C.

10. Craspedometra anceps Carpenter. Albatross station 5157, 3.3 miles from Tinakta Island, Tawi Tawi group,
Sulu Archipelago; latitude, 5° 12’ 30’ N.; longitude, 119° 55’ 50” E.; depth, 32 meters; no temperature record.

11. Ptilometra miilleri Clark. Sydney Harbor, New South Wales, Australia; latitude, 33° 15’ S.; longitude, 151°
12’ E., approximately.

12. Hathrometra dentata Say. Fish Hawk station 1033, off Marthas Vineyard, Mass.; latitude, 39° 56’ N.; longi-
tude, 69° 24’ W.; depth, 329 meters; bottom temperature, 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 temperature record.

14. Crinometra concinna Clark. Albatross station 2324, north of Cuba; latitude, 23° 10/ 35’ N.; longitude, 82° 207
24” W.; depth, 59 meters; bottom temperature, 26.17° C.

15. Isocrinus decorus Wyville Thomson, stem. Off Habana, Cuba; latitude, 24° N.; longitude, 82° W.; approxi-
mately.

16. Same as No. 15, arms.

17. Endoxocrinus parre Gervais, stem. Off Habana, 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 Antarctic 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. Anthometra adriani Bell. Same locality as No. 20. Nos. 20 and 21 were collected by the German South Polar
Expedition. :

22. Zygometra microdiscus Bell. Aru Islands, near the western point of New Guinea; latitude, 5°-6° 8.

23. Chlorometra ‘rugosa Clark. Near Rotti, Lesser Sunda Islands; latitude, 10° 39’ S.; longitude, 123° 40’ E.;
depth, 520 meters.

2 Nichols, H. W., Field Columbian Mus. Pub. 111, p. 31, 1906. 23 Clark, A. H., U. S. Nat. Mus. Proce., vol. 39, p. 487, 1911.

18

24. Tropiometra carinata, Lamarck. From Pigeon Point, Tobago, British West Indies.

THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES.

Shoal water, near shore.

temperature about 28° C. Received from Dr. Hubert L. Clark. According 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 | 2} 4 5 6
|
ws ae 3 Lf le | -
STON NOAA MC il ofa Ah eon ed ary h 8 AAR Wee eumonan | 1.11" | orgraereste 0.12
(Alan) Ossie ns miele ee See: ie Seren cee eo gE 1.07 Poa 10 a7 . 62 . 63
MoO oa. 2k )s Mi eieatlianes ame seers ouisia sas = wine oes ee eee 3. 08 3. 60 3.12 3.76 4.77 4.94
OE ORCHESTRA Cg Sprain el Oc oe ae ae Ee oars SoGne 40. 65 | 40.37 | 34.20 38. 50 38.12 41. 34
TP) Bie Se Re eg LEIS | ol «Pieper saul -21 | Trace? .40 | Trace 43
Uelpmiton Soe ses rape a ae eee nae tare Says ele a meee mic heres Cie arene 51.45 | 53.75 60. 04 55. 25 54. 61 51. 36
i 98.00 98.36 99. 48 98.99 98. 28 98. 82
COmeed edtt=- San. noe e poh soa ei SE apse emits 35.23 | 35.48 31. 37 34. 01 35.19 37.51
Organic matiterWetese.ctee = se ee tee nee eae een ere 16. 22 18. 27 28. 67 21. 24 19. 42 13. 85
alee 8 9 10 11 12
0.0] 0.07| 0.40! 018] 0.17 3.17
719 . 09 . 50 a19 gilt) .3l
4. 64 4. 44 4. 48 | 5.13 4.17 | 2.49
40.75 45, 86 41.79! — 42. 77, 38. 91 26.12
.33 | Trace. | Trace. eT eeliy a8)
51. 80 48. 32 51. 44 50. 28 o4. 61 65. 25
98.35 98.78 98. 61 98. 63 98. 22 97.57
CORneeded Messen sc cochec scence ccs cece cn nee ee 36. 81 41. 27 of. 0 Biel 34. 90 22.73.
Organic'matter; ete< .-- ey eee eee 14.99 | 7.05 13. 67 baal 19. 71 42.52
|
13 14 15 16 17 18
4 = |- —_
SIO fierce bes See oe Seto ee ae ee, eae a ee a 0. 40 0.04; 0.03 0.09 0. 04 0.15
(CAIN) SON e Soe eee ae. eee er eee a 58il 25 | .07 19 . 20 . 26
MG ONS 0) Sa ee kon: 1 ence Sag att aes eee 4.56 4.75 5. 08 4.70 5. 09 5. 04
CaO ie ae Ae Bem ci-d one sleaew ce woes ec ee Se 47.08 41.78 45. 67 42.77 45. 42 43. 41
REO ees Semeeecnses seem iaee aeias casts cae ee Trace. | Trace. | Trace. | Trace? | Trace. Trace.
omition 2 ccecctse: co cio ase omtcnre ons Meee oar eee 47.17 50.33 | 47.54 50. 59 48. 58 50. 00
99. 52 97.15 | 98. 39 98. 34 99. 33 98. 86
CO;meededsa- see: s teistoieceeee te fecal. eee ee 41.93 38.00 40.40 38. 71 41. 29 39. 65
Organic matterdetess sos. ect ner eo a ote ne ne eee 5. 24 12. 33 | 7.14 11. 88 7, 29 10. 35
|
i
19 | 20 21 22 23 24
— é = # | - a
SiQ yank -. us ee eee oe. Ste EU eek ed 0:02} 0.02|- 0.23 0. 04 0. 05 0. 40
CA, He)sOnsie Cee ae ene een tate eect eee: Une 35 45 | .37 48 23 .38
MeO ite Sa aseee aoe Rare er eae te Se oe aT 4.51 3. 02 BBVA 4.92 3.99 4. 87
GOO) E23 once Bee rapt icp a ee ee |, 39.57 40. 68 42.49 37.19 42.72 38. 35
oo 0 RE Seis oc 5 05 XS a SOR ee ods be -10| ‘Trace. | Trace. ey | “race ar
SOPs eceeo she dansodoSosseusscnscoccenemuosdoboesoscas (20 A Senge) (2) (?) (?) - 63
Tgnition..22.0Soveo< <2 te eee eee ee a ee eee 53.64 | 54.53 52. 22 55. 05 51. 69 56.14
98.19 98. 70 98. 58 97. 85 98. 68 97.99
CO, needed >). 227 ee ee ene eee e eee 36.05 | 35.18 37.08 34. 47 37.95 32. 58
Organic matter, ‘ete! 25--0-. 0 eae eee ok. eee 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, respectively.

ECHINODERMS. 19

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

to 100 per cent:
Reduced analyses of crinoid skeletons.

| |
1 2 ai" 4 5 6
SEO) geben coc BOER E ER Eee oe oe 2.01 0.05 1.57 0. 48 0.21 0.14
(Al Be),05 MES ie SA Me a ot 8 2 5 2. let 48 1.41 91 ATS a
MpGO gen - ee cee -s--- 200s ee ener eee enn sees e eee sooo 7.91 9. 44 9. 25 10.15 12. 69 12. 20
(GNC) oe ok cite odode Pap oube ee pe auelenel bone aeeaaeeaEeEe 98.48 | 89.45) 87.77 | 87.34] 86.32| + 85.81
“osc AU acing be ouGaMine (aap See '29| 58) Trace? | 1.12 ‘Trace. Lu
100. 00:| 100.00 | 100.00 | 100.00 | 100.00) 100.00
| |
7 8 9 10 ll 12
—s = =| = = so
een Ns eh) a ec ae eee RO eae 3 « 0.05] 0.08) 0.47] 0.24) 0.21 5.73
(GIRLY) 0 ee Re 2) 2 8 eee 95 | 10 a) 22 | 24 . 56
RS et ore sac een Sak menses -- 11.68| 10.16} 11.08] 12.34) 11.13} 9.36
CS) Sa aa > = ta I SS Bra | 96.46] 89.66|* 87.86| 86.93| 87.94) 83.47
sit aN RG a '86 | Trace. | Trace. Ae mee 88
100. 00 | 100.00 | 100.00 | 100:00| 100.00 100.00
| |
Ast we 15 16 17 18
2 ——|-— -|——_ i
BiG) Pee see. 28 1d cco emery sae Seee ee ees ee = 0. 42 0.05 | 0.03 0.10} 0.04 | 0.17
(UNI 08 BRINE Bee ono oacaceoeorepocoocCeOaHeS . 33 | . 30 | 08 2s| 21 | . 29
MeCOnt sen. -ic tics seat eon sates. || Av: | | 11V6S'| | (112 60) | poet haAd a) paBEe Tee
ARON os 2. 5 OE Ed Sota sae Bowel a2 2 87.16 | 87.96| 88.20| 88.27} 88.13] 87.58
Cas Ope cen ester -1n= een eens Beene tons seis Ses = = Trace. | Trace. | Trace. | Trace. | Trace. Trace.
2 100.00 | 100. 00 | 100.00 | 100. 00 100.00 100.00
| | |
|
| yeaa in 20 21 22 23 24
ve |= en ees aan ee
Sh O35 — Sy ee ne enna an 2050 50.3 3de 3aR = BbaDoaScERCe RoE | Too? i" > OF 02 0. 28 0.05 0. 06 0. 54
CREO) oot rere hen eeesece=-e a0 . 43 | 57 | 44 . 62 SPH) 251
116, Sepa aan legate So | Te Az 7. 86 ROE} || UBEey 9. $7 13.74
GROOMS os 5 EES Sitemeter rscre TT 87.51 | 91.55 | 91.05| 85.48 | 89.80 83.13
(Cae RLON Sol. = oon ae eee = acess =i so .27 | Trace. | ‘Trace. .48 | Trace. . 64
SSO); SSBB BBR neogie desc bis das ansBeadoce eee eeeaame (?) (?) (2) (?) (?) 1. 44
100.00 | 100.00 100.00; 100.00 100.00) 100.00

With these analyses the two made by Mr. Palmer may be advantageously compared,
although they were not quite so elaborate. The data are as follows:

25. Heliometra glacialis var. mavima. Iwanai Bay, northeastern part of the Sea of Japan; latitude, 43° 01’ 40” N.;
depth, 315 meters; temperature, surface, 20.5° C., bottom, WoC:

26. Metacrinus rotundus. Wastern Sea, off Kagoshima Gulf, southern Japan; latitude, 30° 58’ 30’ N.; depth, 278
meters; temperature, surface, 27.8° C., bottom, 13.3° C.

In No. 24, which contained much organic matter, Mr. Palmer found 2.68 per cent MgO
(=5.61 MgCO,) and 40.03 CaO (=71.48 CaCO,). In No. 25, with no organic matter, he found
4.89 MgO (=10.29 MgCO,) and 49.95 CaO (=89.19 CaCO,). Assuming that the crinoid skele-
tons consist essentially of carbonates, and recalculating to 100 per cent, we have as the content
of magnesium carbonate in these crinoids—

20 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES,

These figures fit in well with the others and even 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 Endozxocrinus are averaged together,
and so also are the two of Isocrinus.

Percentage of magnesium carbonate in crinoids.

Genus. Locality. Latitude. Depth. | Heuiper. | MgCO,.

Meters. °C: Per cent.
Heliometnas=- sss eeeeeeee = ces nae Northern Japan. ...--.. 43 SINISE no o's 315 1.5 7.28
Proms. chocrinsecese-seaecse = <1 oles Antarcticn-S-eeeee- see BS SAPEREEEE yi = - 375 —1.8 7. 86
Pinloenmus ssa essen <a 4 === British Columbia. ,-.--.| S20 39GNeeess. - =... 2, 858 1.8 7.91
Arto me Grater ae ee ere etei si ial Amtarct Caer erases sae CYA tS, 0: coe 375 —1.8 8. 23
Psathyrometra.....- See OEE Northern Japan. ..-.--.- Aq OUN Beepeda ss -2.6- (?) 1.6 9. 25
athrome@trataecsssse-2 mets =o oi Massachusetts. ......---- SOs G MN Ee as. > - = = 329 7.8 9. 36
Wlorome tries. settee = attr taislse Washington....-..------ ATED MN eee ss = > 1,145 345) 9. 44
Ghiorometrates se ---se-45- -\- 2-2 se Rottt Island’ee. ose LOSS seeres- == ---| 520 — (?) 9. 87
IB yiiho ChiaUs serra eee melee tae aoa Gulf of Mexico. .....--- Zo CRS SMNeeere (> = 255 (?) 10. 09
Pentametroernus:-.--=-2-<- «22 = Southern Japan. .....-- | QE ae NS... 1,123] 3.4 10.15
iy pal ocCnmnus ese er eects se erat Philippineslands=. -. -| 92 3i/@Neeeee------ - 612 | 10. 2 10.16
Mets erinisteeee oe ee ee a= cee Southern Japan=>=-----| SO°b8@Neeee-—...- < 278 13.3 10. 34
Leen es elon SOO SAE RG FoI EB OSS Philippine Islands. - - .- QO BGIN GS eee lo = =:-:2-- 502 | 12 11. 08
IPtilome tras sence een. sans eee AQIS LPAI As Se Se ees BpialGy|S\¢ 5, -Soaeeee (?) (?) 11.13
TSOCHINUAs S = sees sewers sor see WiGuba tcc esse sees DACAN Samson asc =< =: | (?) (2) 11. 56
Catoptometra eee --os a. epee | Philippine Wslands..- =|" (SoyNieee seems -\-.-- 104 (?) 11. 68
Grislome tras eeeeee as cs- cisicin eee Cuhatcecs tcteresnccre 23 cel OMe? «22: 59 26.2 11. 69
Mropiometra: 2s. --2es---- == eee Brazil eo. 2. Pee eee: Diane |S... S35 ee eee (2) (2) Terie
Bnd OXOCMINUB es Een so: see Ciba coe ccketee corse DACNN Eee a sc (?) (?) 11.79
Pach ylOmetita eee -i= lle = ae Philippine Islands. - - - - SONS Se ere eer at -.-1- 1, 044 (2) 12. 20
Grasped ometiare.. ~<a. see -sis-e ee Philippine Islands. . . ..) 5° 12’ N._.......-.. 32 (?) 12. 34
GCapillasters= 22245 .-< <.as3= 3 5 eee | Philippine-Islands=. —22|) G2" Neeeeeeeeee a= 36 | (2) 12. 69
Tiyeometrae eres van cs eee reopen \CArnglaland seen a eee Bee SL (?) (2) 13. 37

(Proplometrat a... 5-2 «ee tes- see Tobagor..--. += ape os 1d RDN Shoal. | 28 13.7

The percentage of magnesium carbonate in Chlorometra 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
dependent on latitude. Depth of water has also a distinct influence. The crinoids from rela-
tively 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 the 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 probable rating.

So far as we are aware such a peculiar relation between temperature and composition as
is here recorded 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 possibly 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 ermoids from colder regions. But tests by Meigen’s reaction proved that the organisms
were all calcitic, and so this supposition had to be abandoned.

Mr. A. H. Clark, who is an authority on the erinoids, informs us that those from warm
regions have the most compact skeletons, the compactness being in general proportional to
the temperature and to some extent dependent upon the size of the individual. Heliometra,
for example, is the largest of the ermoids, its skeleton is one of the least compact, and its mag-
nesian content is lowest among all the species examined. Structure as well as temperature

ECHINODERMS. ‘ 21

seems to be correlated with the proportion of magnesia in the crinoids, but the chemical ex-
planation of the facts is yet to be found. It may have connection with the gaseous content
of sea water, carbon dioxide, for example, being more soluble in warm than in cold waters,
but this is only a suggestion, which may or may not be fruitful. The same regularity as to
temperature also appears in our analyses of alcyonarians.

FOSSIL CRINOIDS.

In order to make this investigation more systematic it seemed desirable to analyze a
number 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
are indebted to the kindness of Mr. Frank Springer, who selected the material with great care
so as 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 the analyses which follow were made in the same
way as those of the modern species:

1. Pentacrinus decadactylus D’Orbigny, stem. Eocene, Vincenza, ltaly.

2. Millericrinus mespiliformis Goldfuss, stem. Upper Jurassic, Kelheim, Bavaria.

3. Pentacrinus basaltiformis Miller, stem. Middle Lias (Lower Jurassic) Breitenbach, Wurttemberg, Germany.

4. Pncrinus liliiformis Lamarck, stem. Triassic, Braunschweig, Germany.

5. Graphiocrinus magnificus Miller and Gurley, complete crown. Pennsylvanian (upper Carboniferous), Kansas

City, Mo.

6. Dorycrinus unicornis Owen and Shumard, calyx and stem. Lower part of Burlington limestone, Mississippian
(lower Carboniferous), Burlington, Lowa.

7. Megistocrinus nodosus Barris, plates. Middle Devonian, Alpena, Mich.

8. Bucalyptocrinus crassus, plates. Silurian, western Tennessee.

9. Crinoid sp.?, stem. Trenton limestone, Middle Ordovician, Kirkfield, Canada.

10. Diabolocrinus vesperalis White, plates and stem. Lower Ordovician, Tennessee.

Analyses of fossil crinoids.

| b |

|e 2 2 Syn rs 4 5 6 7 8 9 10
fia) pal ewe ll =e Seek eee | Le ee
BIO Foose set see ciosia< 0.99 2. 84 155} 0.24 3. O7 6.92 10. 39 29. 1] 2.56 | 4. 07
(Awe lOs- oss Sess ccee. 2. 64 Sereno 243 1) 2° 18 . 64 aval abe il 1.95
NC(0).5 CL eee ete |) 836 FOO esos 5. H . 00 32 UA be ele lt] aT . 00 . 88
Min OM eee 25 hoo (Pmeres15 & O0n Sete | Trace. .15 | Trace.| .16] 04] Trace. | 04
Wh Oe. eee esse . 78 . 38 . 84 9. 44 EnS . 38 1.21 |} .o8 . 91 19
CaO...-.------------+-++--| 51.22 | 53.68 51.78 43.40 | 50.10 51. 20 46. 57 37. 43 53. 87 | . 50.42
P10 }..3 25 cSeeaeseeee Seem eS .00 | Trace. .09 | Trace. | Trace. | Trace. | Trace. .00 | Trace. | Trace.
Loss on ignition. ......-..- 42. 80 42.93 | 43.21 45. 95 41.71 41. 00 39. 20 30.71 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 erinoids.

1 2 3 1 5 6 7 8 | 9 10
SiO3obeawesee chee dcses cess | 1.00 2. 85 1.57 | 0. 24 Dap 6. 94 10. 48 29. 30 2.55 4.10
(Aljhe) Ose s- 22 .cee2c cae 2. 66 228 2. 64 | .44 PROM) . 64 . 88 1.74 a3] 1. 97
HaGO)s eee ects sae Daal S00} [eee <<: 00 2.16 00 1.94 - 43 00 1.52
MnGOs svc cers sokee ak} ROOM eameere- 5] Trace. .24 | Trace. - 26 .06 | Trace. 06
MeO Ses ae «1.66 80 1.79 | 20.23 1. 66 80 2.56 1.23 | - 1.90 1. 67
CaO 5. teehee seer an eeeeel 92.26 | 96.07] 93.80] 79.09 90.61] 91.62) 83.88) 67.24) 95.25] 90.78
Ca PiOe wos c= ae eee .00 | Trace. | .20 | Trace. | Trace. | Trace. | Trace. .00 | Trace. Trace.
| 100. 00 | 100.00 | 100. 00 / 100.00 100.00 | 100.00 100.00 | 100. 00 | 100. 00 100. 00

Die THE INORGANIC CONSTITUENTS 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 mag-
nesia 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 clear. 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 alge reported by Hégbom *4
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 mvestigation.

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 drébachiensis O. F. Miller. Upernivik, Greenland; latitude 72° 48’ N.

2. Strongylocentrotus fragilis Jackson. Albatross station 2946, off southern California; latitude, 33° 58’ 00’ N.;
longitude, 119° 30’ 45” W.; depth of water, 274.5 meters; bottom temperature, 13.6° C.

3. Echinarachnius 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° 35/ 15” N.;
longitude, 118° 08’ 30” W.; depth, 86 meters; bottom temperature, 15° C.

6. Loxechinus albus Molina. Port Otway, Patagonia; latitude, about 46° or 47° 8.

7. Tetrapygus niger Molina. Coast of Peru.

8. Tretocidaris afjinis Philippi. Albatross stations 2316 and 2317, off Key West, Fla.; latitude, 24° 25’ N.; longi-
tude, 81° 47’ W.; depth, 85 meters; bottom temperature, 24° C.

9. Heterocentrotus mammillatus Linné. Low or Tuamotu Archipelago, southern Pacific Ocean; latitude, between
14° and 24° S.

10. Encope micropora A. Agassiz. Puerto Viejo, Margarita Bay, Lower California; latitude, 24° 30’ N.; longitude,
112° W., approximately.

11. Clypeaster testudinarius Gray. Southern Japan:

12. Echinus affinis Mortensen. Albatross station. 2206, between Hatteras and Nantucket; latitude, 39° 35’ N.;
longitude, 71° 34” 30’ W.; depth, 1,919 meters; bottom temperature, 3.6° C.

Analyses 10-12 by B. Salkover.

13. Echinometra lucunter Linné. Pigeon Point, Tobago, British West Indies; latitude, 11° 25’ N.; shoal water,
near shore; temperature, 28° C.

14. Mellita sexiesperforatus Leske. Pigeon Point, Tobago, British West Indies; latitude, 11° 25’ N.; shoal water,
near shore; temperature, 28° C.

Analyses 13 and 14 by R. M. Kamm.

* Hogbom, A. G., Neues Jahrb., 1894, Band 1, p. 252,

ECHINODERMS,

Analyses of sea urchins.

23

i 2 3 4 5 6
SEO eerste aie aietnietetn cass Sivieietalsin e/<iciejee.n oo eels os cian a 0. 12 0. 26 0. 14 3. 86 8. 52 0. 05
ENR SOR: ey eS ee Aa 34 65 e1 5.03 3.01 W7
V0). Scsec Toone cudgel cb saaseCe= LOSc Rens 50 Ser cep EE eSooeEe 2.58 3. 68 2.97 4.75 3. 04 3. 07
(OO a tice DROS OC OUGGSS G0UISC COO SUC SUCRE OS eEo eee 47. 34 41.08 49.17 43. 42 37. 92 45. 87
NOM cocceSacbe cf asE PR onbeE CE Sene oriccs Dea Ca a Geese . 20 1.15 . 26 . 28 .37 . 3d
PO) 56,5 ipod SE BE OE EGE ERC C DURA OAnOde So eee eae Trace. 39 .05 | Trace. 19 Trace.
ISAT ee Soe. 7o 6 78D 005 SCOP OCC Leb obos 4c GOGO BEaeeoee | 48.53 52. 21 45.74 43. 01 45. 38 49. 47
- Web Mil 99. 42 98. 60 99. 35 98. 43 98.98
WOrmeed Cd ewe seca seo c te no onsen ee Cesc oe we oes 39.'92 34. 20 41.71 38. 99 32. 28 39. 34
Oreani@unattony etene..Us:32) Soe eee seme clea cs be 8. 61 18. 01 4.03 4. 02 13. 10 10. 13
7 10 11 12 13 14
SHO haste cearosdesonenrnos ceaodns oo. oct ene eB oe 545008 0. 31 2. 38 0. 15 3. O1 0. 10 0.14
(GAULT AES OP ete Bee PAF 2 AV Oe +A, eed 30 81 13 2,21 .28 36
IIS Oeecatiecuct peccnegieesunoneo= a8 SY acemeaan age Ieee 2. 82 5. 45 3.90 2.39 4.16 5. 23
(ORO)E sepscenosecammine ob ope On ce. bende + doco seu ge OSC Ieee 48. 86 38. 24 49.73 46.19 36. 97 44. 84
SO Haast concseas poco Ber saee es SERA ate Ne Ay 2S) o 0 av Sono 1.43 | Trace. | Trace Trace. 99 ey
12} 0 RS arto atin see pours aac esor daa: CAO Rees sees Trace. | Trace. | Trace Trace. . 64 Trace.
Tonition esos eee eee eee ene scar seiesic 44.98 51. 24 45. 47 46.90 51. 82 46.70
98.70 98. 12 99.38 | 100.70 94. 96 98. 64
GOxneeded! = 5s: . v.cseedaseee wecee ete ses sesesseecse 40. 69 36. 64 43. 36 38. 92 32. 49 39. 27
Oreanicimatter Cte: . meer aa pecs eaies cae task 4. 29 14. 60 2, 11 7.98 19. 33 7.43
Rejecting the very variable organic matter, etc., the reduced analyses assume the following
form:
Reduced analyses of sea urchins.
] 2 3 4 5 6
SiO 6 cubpoSoaebboedeoeSas 26 sn S6 DBL OOEEABOSOnaaee 0.13 0. 32 0.15 3299 9.93 0.05
GUE Oy cee ee AR a So 37 81 129 5. 20 3.51 18
IVS C10 AS ORB RB BORE rISE Se hing So.oc\nde Debs Bap See aOEe 5.99 | 6. 95 6.59 10. 38 7.44 7.38
CEC 0). Se depeucpeBe eee des: cad ct JaBSee Ac neACeneeeseanal 93.13 | 88.44 92.39 | 79.94 77.91 91.73
aS O eee ates he sect oe ere tato leks slaittcisoe ls isielstare aid 38 2. 42 . 46 | .49 .73 - 66
ORE (C5 SAME HSrnc cp ac.5 Cus bone Calis Benes eae Trace 1.06 > 12) |) ‘Erace: 48 Trace.
100.00 100.00 | 100. 00 | 100.00 | 100. 00 100. 00
7 10 11 12 13 14
ome = ee 3 | =
(3 2 Reo .2 2 2 a 0.33 2. 87 0.16| 3.25 0.13 0.15
(UNITING 0} 228 ES pee gericecc oro 5-405 55S ae epee Seer .32 | 98 14 2. 38 - 37 39
AO Sea eer eee «on cas emieee am sce seecisc cscs ccnes 6. 27 13.79 | 8. 41 5. 41 11. 56 11.91
(073010 sos Je DESERTED 6 doc Sn on OC BOOS Sete S Seen eaeeEe 90. 52 82.36 | 91.29 88. 96 83. 87 85. 02
CaS O eee ene nse s ns eeceeclene ee eee osimce ac dee 2.56 | Trace. | Trace. | Trace 2. 22 2.53
(OM LH Oe SOs Spero eee S oc SS cads CoCo See ee eee Trace Trace Trace. | ‘Trace. | 1. 85 Trace.
|
100.00 100.00 100.00) 100.00 100.00

100. 00 |

Sea urchins Nos. 8 and 9, Tretocidaris and Heterocentrotus, 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 purplish-

red spine. The large red spine analyzed was 15 centimeters long and weighed 13 grams.

24 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES.

Analyses of Heterocentrotus mammillatus.

: | Dental White Red

Dell ees : Se ae

| pyramid. | spines. spines.

DIO phen. = 3's Sodas ee | 0.02 0. 02 0. 05 0.05

(AU BeyOn. 0 eee ye eee Hie) 08 .13 | 26

MeO Use: age 28a afheeeea 5.21 | 5.50 3. 74 4,47
CaO 2. ph cctes oar ba seoes 43. 60 46.02 | 48. 26 47512 ~ ||
SOR (oe eee eee | 61 | 58 . 29 73. |
IPRA e4 Jee ht See | Trace. Trace. Trace, | Trace. |

Meaviney NG se hoes Gok Sse S-\- - 49. 62 47. 00 46. 46 45. 99

99595) 99. 20 98. 93 99. 27

GO; meeded =e eee ee 39.66} 42. 89 41.77 | 411. 49

Organic matter, etc. ...... 9.96 | 4.11 4. 69 4.50
east ee ser | i |

Reduced analyses of Heterocentrotus mammillatus.

| x

SiOy-k Skee eee 0: 02 | 0. 02 ORODi | eens

(GN) HOR sae so tscnsn on a . 14 | . 09 14 | 28

| UO eeescconansene sons 12526)) 12.27 8.32)| _ 19:86

GaCO3. 4 cise scet sees 86. 42 | 86. 57 90.97 | 88.43

| GzSO, eee eee on 1. 16 | 1.05 BERN Tsets

I Ga, PaOgeectenestee eee aes Trace. Trace. Trace. | Trace:
= |.
100.00 | 100.00 100.00 100. 00
E

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 can be
called so, would probably be somewhere near 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:
Analyses of Tretocidaris affinis.

Actual analyses. Reduced analyses.
Shell. Spines. | Shell. Spines.
| |
SiO pease sete taco eee 0. 11 OL SSMESLObe:---..-. ci eebee eee eee eee 0. 12 0. 56
(ATURE FO; eee ee nee 515 TEGO He), Os... snare ee eats eae ag 15
MaOe tt eee ek SS ee 4.02 ONO TAIN COs: =... ja eee ene 9. 30 4. 63
CAQRE sehen ees noo eee 45. 80 AO 7TOMIROACO gE 2. «= onc .zc eee see erecta 89. 35 94. 66
Soe ee Oe LEE NG © 57 (?) CHSC a ae cea a 1.07 (?)
BIO; eeceaaceac + atopic ke cs) hot Trace. Trace. || CagP203-------.-.-.----- ad beahese Trace. Trace.
Tipnitlonisentew sess oes ae 48. 32 46. 28
| 100. 00 100. 00

98. 97 | 98. 81
CO; meededs 5-2 5.) ee. 2 eee 40. 09 41. 40 °
Organic matter, etc...........-.--.- 8.23 | 4.88 | |

| j |

Here again the spines are lower in their content of magnesia than the shell.

In two of the analyses, Nos. 4 and 5, large percentages of silica and sesquioxides appear.
These 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 magnesium carbonate became 11.43 and 8.59, respectively. Similar corrections to
the other analyses are negligible. After 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.

ECHINODERMS.

Percentage of magnesium carbonate in sea urchins.

25

l

Locality. Latitude. | Depth pemiber: MgCO3.

| Meters. oO), © Per cent.

Bighinuge sae iscsi f= os ta Station: 22062:.-..2-2--.- 899. 354 IN) session 1,919 3.6 | 5. 41
Strongylocentrotus drébachiensis. . - - (Greenland: 5. 2-=--2-~ 1 72° 48° N (?) (ED, 5.99
Tetrapygus..:-.---.--------------- ISG oF So Son eneore se (td Beeaadee ee ecscc St (?) (?)
Echinarachnius........------------- New England. --.-......- A DOA BINS irtaanr= ote (?) (?) 6.59
Strongylocentrotus fragilis........--- @alifomya--=---.2...--- S35 BC INE aweeaeett 2, 745 13.6 6.95
lls Cicl nian: Gap ese amnode cpagorsct Baba ODE tee onto - 14-13 AUG cocUon: (?) (?) 7.38
Clypeaster. ..-..-.-.-------+----=- Joos. cob: soeeBeede| (GQ) Eee aes ses eee | (?) (?) 8. 41
Lytechinus.........---------------- Galittormigeee =e — =~ |). 33° S54 MG aaaee cee 85 15 8.59
Tretocidaris. -...-- KeyiWestea2=-:---.--- 24°: DB NE eee $5 24 9. 30
Encope californica Galapagos’: -\s2~-- --1--- Do MEN eoceoe node aa0- (?) (?) 11. 43
Bchinometra......-...----+------- AN ee <5 ge gape SDE W125 NGS eeeeree lm moose 28 11. 56
ie llitpesee seo mosses sas ae cee e MobgeOteesccece cai in! adS5 254 NS ce semrieneers Shoal. 28 11.91
Heterocentrotus...-'....----.-=----- IMWEniGlibsaaea=nesascenee 42) DAG S eeeeetopoteinte (?) (?) | 12. 26
Encope micropora...-.--..---+------ Galvtomiane= =e < <= - = DAES CN ic eats sentra (?) (rt 13. 79

These figures, like those for the crinoids, seem. to show a variation dependent upon tempera-
ture, at least so far as temperatures have been determined or can be inferred. The sea urchins
from cold regions are relatively low in magnesia; those from the Tropics are high. There are,
however, two apparent exceptions. The urchin from the coast of Peru probably owes its abnor-
mality to the cold Humboldt current, which flows northward from the Antarctic Ocean. Encope
micropora is also seemingly anomalous, and its high percentage of magnesia is difficult to ex-
plain. Unfortunately the actual temperature of its habitat is not given.

Two published analyses of sea urchins are worth reproducing. They are:

1. Echinus (Strongylocentrotus?) drébachiensis. North Sea; analysis by L.Schmelck. Norske Nordhavs Exped.,
No. 28, p. 129, 1901.
2. Echinus esculentus. Locality not stated, probably Mediterranean; analysis by O. Biitschli.

Gottingen Abh., No. 3, 1908.

K. Gesell. Wiss.

Older analyses of sea urchins.

Actual analyses. Reduced analyses.
cs -* = Blew = a
er, 2 1 2

— lee |
SilOlsenceos ly ‘Draces QHOSRISTO pease poe eee eerie Trace. | 0. 04
INIA Foon SEB ORE Ee epee cic bear “bere | ese oooces IRA OF pecnnde orooncareoce seca] Sheteeh || See copene
OO h 2... ene: ae A (i ol eee ee HG; cease os. Odes Cot eereee O36; esheets
COM ei ee. 50 cane aio eee SEO) || MeCOgi--o: 5 or esse sep casas co cee 6. 36 8. 84
MiG Osa ase --= = «oon © se seen oe 5. 30 | SRF CACO aoa ctceic a at entail win ms mimimwi 93. 28 89. 64
Te 0) ee ean aha awl =Falal heme tet ADrACel| saan ee I CaS Oj oe tee Se staeian wth == eerie Trace 1. 40
(Phosphatesescs sss /:< 2-1 ee iea Iter ain 2 BOS) Cas Pp Ope sener eee aes Stel Trace .08
(aS Oe ee anos eee | eo ee en Bee
(CHIS(0).. PACE Bees deapestacs soS6 | Soe» 1.70 100. 00 100. 00
18 {O)en¢ 22 2 eet Seperate a | Seeabecoe 2.54
Organic’ matter...----------------- 16.33 203 :

99.68 | 99.23

|

Although different in minor details these analyses are fairly comparable with ours. The
urchin analyzed by Schmelck was from the far north, and its composition is very near that of
our specimen from Greenland. Bittschli’s sea urchin is doubtless from warmer water and shows
the higher figure for magnesium carbonate.

3. STARFISHES.

In the former 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 ma-
terial is available, it is better to separate the two groups of organisms. Twenty-nine star-
fishes have been analyzed, of which two from Tobago were given us by Dr. Hubert L. Clark.

26 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES.

The other specimens were selected by Mr. Austin H. Clark from the collection of the United
States National Museum. The list is as follows:

1. Asterias vulgaris Packard. Eastport, Maine; latitude, 44° 55’ N.; longitude, 67° 00’ W.

2. Asterias tanneri Verrill. Albatross station 2309; latitude, 35° 43’ 30’ N.; longitude, 74° 52’ W.; depth, 102
meters; bottom temperature not given.

3. Asterina miniata Brandt. Pacific Grove, Calif.; latitude, 36° 36’ N.; longitude, 121° 55’ W.

4. Leptasterias compta Stimpson. Albatross station 2250; latitude, 40° 17/ 15” N.; longitude, 69° 51’ 45 W.;
depth, 86 meters; bottom temperature, 10.8° C.

5. Benthopecten spinosus Verrill. Albatross station 2568; latitude, 39° 15’ 00’ N.; longitude, 68° 08’ 00” W.;
depth, 3,249 meters; bottom temperature, 2.7° C. :

6. Luidia clathrata Say. Near Charleston, S. C.; latitude, 32° 47’ N.; longitude, 79° 57” W.; depth, between 2 and
22 meters.

7. Acanthaster planci Linné. Palmyra Island, in the Pacific Ocean, west of south from Hawaii, latitude, 5° 49’ N.

8. Asterina minuta Gray. Pigeon Point, Tobago, British West Indies; latitude, 11° 25’ N.; shoal water near shore;
temperature, 28° ©. Weight 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° 08’ N.; longitude, 50°
40’ W.; depth, 93 meters; bottom temperature, 1.1° C.

11. Odontaster hispidus Verrill. Off Marthas Vineyard, Mass.; depth, 245 meters; bottom temperature, 11.1° C.

12. Plutonaster agassizi Verrill. Off Marthas Vineyard, Mass.; depth, 584 meters; bottom temperature, 6.6° C.

13. Asterias forbesii Desor. Vineyard Sound, Mass.

14. Pontaster tenwispinus Verrill. Albatross station 2095, between Cape Hatteras and Nantucket; latitude, 39°
29’ N.; longitude, 70 ° 58’ 40” 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. Orthasterias tanneri (Verrill). Albatross station 2307, between Cape Hatteras and Nantucket; latitude, 35°
42’ N.; longitude, 74° 54’ 30’ W.; depth, 79 meters; bottom temperature, 14.5° ©.

17. Urasterias linckii (Miiller and Troschel). East of Nova Scotia; 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° 01’ S.; longitude, 73°
42’ 30” W.; depth, 675 meters; bottom temperature, 8.3° C.

24. Ctenodiscus australis Liitkin. Off Patagonia.

25. Marthasterias glacialis (O. F. Miiller). Horta Harbor, Azores; latitude, 38° 35’ N.; longitude, 28° 50’ W.
approximately.

26. Asterina pectinifera Miiller and Troschel. Otaru, Hokushu, Japan; latitude about 43° N.

27. Culeita novaeguineae Miiller and Troschel. An East Indian tropical species, locality unknown.

28. Linckia laevigata (Linné). An East Indian tropical species, locality unknown.

29. Coscinasterias calamaria Gray. New Zealand.

Analyses 16-29 by B. Salkover.

Analyses of starfishes.

1 2 3 4 5 6 7
SIO prea oe nce eh cae. eee ay ee ee Oraoaeemos7oy|) 0.035) Jezin25 06") Ose 0.19
CANE GO sees 85 ojo eee ee ee eee eee aL eee 21 55 20 48 79 36 14
MeO): Screen 6 yoo cet Aenea os ho ae DAO le SEB || SERS CSCing| SREB || 25:5) 4.36
CaOisio- Stee. a Soe EY Sal RE ees 35.71 | 38.51 | 36.85 | 30.35] 40.54] 41.30 33.18
SOgs ih: epee sce Sy eee ANC (?) (?) (?) (?) @) (?) (?)
PaO givens sitio cece eee ore ee a .07 24 .14 3 sill fal 07
Tpnition:... 225-52 2 Se ee OT 60.18 | 54.89 | 57.64] 63.91] 51.66] 52.03] 62.07
99.21] 98.81 | 98.84] 99.13] 99.09] 98.99] 100.01
GO) meeded!! het bec tc adh ents seek ae ee 30.84 | 34.30] 38.05] 27.08] 35.94] 37.70] 30.81
Orpanic mattervetent si aah Pe Lae ee 29.34 | 20.59 | 24.59] 36.83 | 15.72] 14.33 31. 26

ECHINODERMS.

Analyses of starfishes—Continued.

27

8 9 10 1 12 13 14
RO Re ibe ton Oona 1 RRO MERE See oe eee ee 0.49 0.16 0.31 0.51 0.31 0.10 | 0.27
(ANAS) Oh ou! Lvl oe Eee el oe 118 AW 10 16 48 39
WHO) oe ac sadansc SbSeb es Wo bEbous cae pee pO UU OR EeaeoS 4.16 4. 64 3.26 4.12 3.81 2.39 3. 67
(OSS Op reer re alee cin ate ae Seale semia ia a os feiciecn = wie 33.90 32.28 39.50 | 40.44 44.43 30.93 43.93
JO) ~ oc oS onind CACHE Gab OS nO SOLE ROB HOES? BoSHOsBEeBS Trace. | Trace. - 26 03 (?) 22 06
SAU ac os oo aeRO e ERSTE EE eee ae (2) 7 "75| 169] .62| 82 46
LSM AGT 5s. - secu PED ESEEBBOGHeE eho: No So- So peeeeEnoS 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
co, needed = £8 Ayreon Brno Bane S05 OH OSCE Te Deoaanaee 31.22 30.97 33.97 35.90 36.76 26.08 38. 28
Orpantenmatlens Cle sete teletoiajtereien nina =a = es ~ cle = 28.75 | 31.25] 21.7% 17.25 | 13.26 | 39.27 11.92
| 15 | 16 17 18 19 20 21
| |
RIO) SS0ES Soc AAgee ens A lee Bessie cite Bee eee EO! 278 |" 0268) x08 00" Us Ob45n Ie sOstSe | mets29 0. 66
CRIBS) Ooee One tke yh BOE mea 2 (20) 1.25] 1.38] 1.20 20} 6.89 1.16
INNO). a to net eee BAS GA DA ASCEND borne > OL Stee beeHOEHee 5.44 | 3.39 2.06 2.000} 2295 4.03 4.02
(CaO eeeen sobs case ae cee cee csiatccne e6 42.30 | 32.21 | 24.22 | 26.79) 30.50) 26.44) 29.13
1710 ae Cae aOR ie 25 Sn ae Trace. | Trace. | Trace. | Trace. | Trace. | Trace. | Trace.
SOR ince aoe saees conceals secteiemeer esas ss esl ae .74 | Trace. | Trace. | Trace. | 47 .75 .76
Uprin tone Sct artes srorqis = oeie microns oie wie ainla a aicin' st 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
CO bmeaded ee 5-1 sss «wicteeeeeemeeaemon.s ade sisnlae oie 38.81 | 29.04] 20.94] 23.80 | 26.96 | 24.17 26.89
Oreanicnmiattery OtC =. - co seem eee teeta nar sa )aie isle. 11.90 | 32.23 | 43.34] 40.57 | 35.95) 37.05 37.03
5 } \
22 23 24 25 26 27 28 | 29
== |
SiOz asso es Soeelas 5 Jeune mab ee er ace voce 0.05 | Trace. 0.70} 0.23 0. 24 0.08 0.49 | 0.00
(ARE) Ons - cconcoc ccs | ee 8 2.57| 0.79| 5.10| 1.64 .83 SO) ay 1.81
IMO roses fas ce kcl ae eee seo se 4,22 3.18 Sey) ZARB) 1.95 3.72 3.15 Ver itt
CaO BE bee «oo Sa adoewac eerie eaies aascese 29.44 | 29.32] 30.12] 27.75) 16.04] 25.59) 24.48 | 24.78
AOR Wao bocqataeccnckns eee eee Trace. | Trace. | Trace. | Trace. | Trace. | Trace. | Trace. | Trace
(SAAN eeeneenee RORBer pe brad occa sececansaene .81 | Trace. | Trace . 80 46 74 -71| Trace
Wernition: <2 015 cate ociceie a eres eae Sees e 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
OO snecded |. «5 ...55. ssn sktecinsiaics= sit eea 27,32)|' 26.54 || 27-889) 23.81 | 1450) )) 23579) S07 23. 52
Orpanie matter, etC:--.c00 seceeeeee ee. cee ee 33.59 | 37.61 | 31.09 | 43.46] 64.96 | 44.88 | 36.77 45.72
| |
Reduced analyses of starfishes.
1 2 3 4 5 6 7
S1\0)h oe ce eee a an BEBE Rea OSE cobtco AA SHE Tee See meer 0. 64 1.01 0.03 1.94 2.47 0.32 0.27
(CTSA SENS a eeae 30.) Serollie 297) | Sez7i| kgenleneiran “20
iat Ta) 2 Oy hc = Ed 1s 7.79 | 10.28| 11.24| 10.27] 9.88) 12.13] 13.83
(C010 Rees eee Nee so) RE Pe ee ee 91.06 | 87.44} 88.06 | 86.57 | 86.42 | 86.77 85. 99
ESO See a 2 bee RR a SD, (?) (?) (?) (?) (?) (?) (?)
Se) ie dee Ee a (21 57 40 45 .29| » 36 21
100.00 | 100.00 | 100.00 | 100.00 | 100.00 | 100.00 | 100.00
oe S5>50—_00—000N
8 Y) 10 ain 12 13 14
Serene Sk! o.70 { 0-24] 0:89) 0.62) 0.35] 0.17] 0.31
GEIR GNO sien as (pe 26 122 112 ats .78 145
LUPO 0 See eon e cas Soo: eo 12.53 | 14.31 8.78 | 10.58] 9.09) 8.24 8.86
CRO soos oes — sree eee ee eee. 86.77 | 83.42 | 88.48 | 87.16] 89.18) 88.19 89. 34
CagP,0s a hy EG SR eg is ote waa Trace. + Trace. 73 FOS Cae |) W728) 15
CaSO isso soe. =. Ses ee Ee Bos rns 5 coe. (?) 1.77 1.40 1.44| 1.20 1. 84 | 89
100.00 100.00 | 100.00 | 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
BiQj=2 2425 Fee et fois ces See Eee ee ee oe 0. 24 1.02 0.00} 0.82 0:29} 2.06 1.06
(Al Fe):O; See ene). Se eae ane FB -23) 1.88| 2.84) 2.19|/ .33| 11.04 1.86
MpCO) 2 <3: Soe eee ak oc ce ee 13.02 10. 69 8.91 9: 600 LOM SIS S06 13.52
CaCO) 2 RO ee ee as ee Es ene ere 85.08 | 86.41 | 88.25) 87.39| 87.93 | 71.28 81. 82
GCasP Os) cee ede ented an = ee ee ne ee Trace. | Trace. | Trace. | Trace. Trace. | Trace. Trace.
CaSO coerce gee tece facack ce eel eee eercee ee 1.43 | Trace. | Trace. | Trace. | 1.34 2.06 | 1.74
100.00 | 100.00 | 100.00 100. 00 | 100. 00 100. 00 100. 00
| |
————— I aa aI
22 23 | 24 25 26 27 28 29
(ik A meine oe 9S Ca ae | 0.06| Trace. | 1.05/ 0.41| 0.71| 0.15] 0.70] 0.00
(ATM 6) LO. eee kes 0S eee eee 3. 99 1.32 7.66} 2.90 2.44 | SAA fem yee | 3.49
MoCO seas: Sateen Peace re pores 13. 76 11.16 | 10.51 8.29 12.05.| 14.35] 9.32 11.22
GE CO 5 oc saa e ene ete cee See eee eee 80.04 | 87.52] 80.78} 85.99 | 82.51 | 82.27 | 85.63 85. 29
Gas P2 Og 25 pene Pee ERE eo cane Lee Trace. | Trace. | Trace. | Trace. Trace. | Trace. | Trace. | Trace.
CaS Oy: sce kira. emer es race aan Maemo 2.15 | Trace. | Trace. 2.41 2.29 253k Ua races
f |

100. 00 | 100.00 | 100.00 100.00 _ 100. 00 100.00 | 100.00 100.00

|

In the following table the analyses are arranged in the order of ascending magnesium
carbonate, like those of the crinoids and sea urchins.

Magnesium carbonate in starfishes.
a . Temper-
Locality. Latitude. Depth ature MgCO,
Meters. SC: Per cent.
Asteriasivuloanis:--.2--2s.- sien Hast port: << =-4so eee BAC OD CN ce ee (?) (?) 7.79
Astoriasiforbesi=2-- 32... .- goss te Vineyard Sound........ (?) Se eas = (?) (?) 8.24
Marthasterias glacialis............_. VAG) <2: ee a SS fesseigo” Ni... 2) ee (2) (2) 8.29
Ctenodiscus crispatus............... Newfoundland.......... ASci08. Ni. See 93 abt 8.78
Pontaster tenuispinus............_. Station) 22050). -eeeeeaes eoop.297 NS: eee 2, 456 (2) 8. 86
Urasterias linclai.. 2.222.222... _- Nova Scotia.~. 2 aan ase: [pasorg2” Ni... eee 403 (?) 8.91
Plutonaster agassizii-............... Marthas Vineyard...-.... (Cp eee 5... SR 584 6.6 9.09
ihinckialaevirata sss) oeeesseeen ee Hiastundiess-<-o) =eeee (ge) Bees 2 ee (2) (2) 9.32
ABteTias aAceLvatae sesso. seeee eee West’ Bank-3- soe (Qe. (?) (?) 9. 60
Benthopecten spinosus.............. Stationi2b68>.55 see SHS SVAN S a ee 3, 249 2 9. 88
Astropecten americanus.........._. Marthas Vineyard....... (G2 yaaa ene (?) (2) 10.11
Leptasterias compta................ Station 220025.) eee AOC N'.. cae 86 10.8 10. 27
sterias tanner sae see een et Station 2309............. BoeASZiNi . 22 eee 102 (?) 10. 28
Ctenodiscus australis Patagotiace:..540- = en (©) 5c Sc (?) (2) 10.51
Odontaster hispidus. ....... 2.22... Marthas Vineyard....... Ree Es < 2 -t le cioers 245 11.1 10.58
Orthasterias tanneri................ Station! 23070 ste BH GIAQAIN,. 5.45, ee 79 14.5 10. 69
Ctenodiscus procurator............- Off Chiled? = 22h IB" (US eer Bia) 8.3 | 11.16
Coscinasterias calamaria..........__. | New Zealand............ itacs oes (2) (?) 11.22
Asterina miniata.................. |kCaliforniat=.2. i aaa BHMEDOGING.. .. <wcsee (?) (2) 11.24
Asterina pectinifera................ Tapane 2 ey 2 hye MACRNGe ous (2) (2) 12.05
Tuidiaiclathrata 2-22) 7) Charleston’. =... 225544 BUCEAYEAN Gc aoa oc saat (?) (?) 12.13
Asterina minute... <2 225s-s. ee esee | <RODEPOS Sateen ee OO ee Shoal. . 28 12.53
Astropecten articulatus............- Blorida. 4s.Ger. | out sees D>)... (?) (2) 13. 02
Acanthaster planci...-............! Palmyra Island........- DUMOGN Es. = 2.3 (2?) (?) 18.33
Pharia pyramidata.-./....2...1.12. | Cape San Lucas. ....... ODONDOIEN MEE... 2 (2) (2) SPS
Phataria bifascialis................. | Cape San Lucas......-. eG ik) (?) (?) 13.56
Oreaster occidentalis... {Sart cious telandies seme oro is. (2) (2) 13.76
Linckia, guildingi1! = sees.) s22 os NeTiohagotaee een TIC Ob NGte es. =. Shoal... 28 14.31
Culcita novaeguineae..........____- | East Indies.-.......-... (7) (?) (?) 14. 35

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 ]owest in magnesia; those from warm regions are higher.
The regularity, however, is not absolute, and the variations from it may be real, or perhaps may

ECHINODERMS. 29

‘
be due to differences in habitat. The starfishes from Vineyard Sound and Marthas Vineyard,
for example, may have come from different depths of water, and so, too, may the two marked
as from the East Indies. Small impurities in the specimens analyzed may also account for
some irregularities. The analyses by L. Schmelck,” of starfishes from the North Sea, are, how-
ever, not so easy to explain. His data are as follows:
1. Arcaster tenuispinus. Station 10; latitude, 61° 41’ N.; longitude, 3° 19’ E.; depth, 402 meters; bottom tem-

perature, 6° C.
2. Arcaster tenuispinus. Station 25; latitude, 63° 10’ N.; longitude, 5° 25’ E.; depth, 179 meters: bottom tem-

perature, 6.9° C.
3. Astropecten andromeda. Station 10; details under No. 1.

Schmelck’s analyses of starfishes.

1 | 2 Ent sd
|
Trace. | Trace. 7.09 |
Trace. | Trace. | . 65
OF3Is | 0.60 | gad
| 9.36 | 9.24 8.55 |
| 74.11 78.92 64. 48
(?) 1.00 .83 |
Trace. | Trace. | Trace.
16.23 | 10. 24 13.83
100. 00 100. 00 95. 80

Schmelck’s analyses reduced.

|

| 1 2 x
SiO jae aeeetea= cc les aso sees | Trace. Trace 8. 66
ATS @ eee ons scsi dase s| Trace. Trace 79
Be, Opceerrasae aries sence tease es | 0. 36 0. 66 45
MeCOmme ct 1.18 | 10.29 10. 42
(AGIOS nde<5 BODES EROoCE oes 88. 46 87.92 78.67
CaSO Ee le | LO re ire 1.01
Ca,P,0, coc Snobeee eee eeee Trace Trace Trace

| 100.0 100.00 | 100. 00

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 are recalculated
to 100 per cent the figure for magnesium carbonate becomes 11.41, nearly that for No. 1. All
the percentages given for magnesium carbonate are higher than the temperatures 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.

. Gorgonocephalus arcticus Gray. Off Cape Cod, Mass.; about latitude 42° N.
. Gorgonocephalus caryi Lyman. Alaska.
. Ophioglypha sarsii Liitken. Albatross station 2176; latitude, 39° 32/ 30’ N.; longitude, 72° 21’ 30” W.; depth,
553 meters; bottom temperature, 5° C.

4. Ophioderma cinereum Miller and Troschel. Ensenada Honda, Culebra Island, east of Porto Rico; about lati-
tude 18° 20’ N.

5. Ophiomyxa flaccida Say. Pigeon Point, Tobago, British West Indies; latitude, 11° 25’ N.; shoal water, close in
shore; temperature, 28°C. Adult specimen.

wnme

2 Schmelck, L., Norske Nordhays Exped., No. 28, p. 129, 1901.

30 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES.

6. Like No. 5; half grown.

7. Ophiocoma pumila Liitken. Pigeon Point, Tobago. Like No. 5.

Analyses 5-7 by R. M. Kamm; specimens received from Dr. Hubert L. Clark.

8. Ophiocoma aethiops Liitken. Espiritu Santo Island, Gulf of California; about latitude 24° 30’ N.

9. Ophiothrix angulata Ayres. Cuba.

10. Gorgonocephalus eucnemis Miiller and Troschel. Albatross station 4912, 17} miles off Tsurikake Island,
southern tip of Japan; latitude, 31° 39’ 40” N.; longitude, 129° 20’ W.; depth, 715.5 meters.

ll. Ophiopholis aculeata japonica Lyman. Illiuluk, Unalaska; latitude, 53° 42’ N.; longitude, 166° 32/ W.

12. Ophioglypha lymani (Ljungman). Off coast of Chile; latitude, 48° 41’ S.; longitude, 74° 24” W.; depth, 355
meters; bottom temperature, 11.1° C.

13. Ophiomusium lynani W. Thomson. Albatross station 3407; off the Galapagos Islands; latitude, 0° 04’ S.; longi-
tude 90° 24’ 30’ W.; depth, 1,619 meters; bottom temperature, 2.9° C.

14. Ophiocamax fasciculata Lyman, Albatross station 2125, Caribbean Sea; latitude, 11° 43’ N.; longitude,
60° 09’ 30” W.; depth, 381 meters; bottom temperature, 10.4° C.

15. Ophionereis eurybrachiplax H. L. Clark. Albatross station 3702, off Honshu Island, Japan; depth, 57-75 meters.

16. Ophiocoma erinaceus Miiller 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
|

SI Op asi scinseteseee es sols 3 dae aebboee onc tet Re baie eee Uris 1.08 0.98 0) 21 0.11 0.43
CAMS OS Ee es ES Oe na a 62 72 53 09 salty 55
BOS ee eee Meso kd oe a ee oe 3. 36 | 2. 82 Shee) 5. 80 4. 67 4. 64
CEO SR eee Sie 5 Al ee aE, AMER DR NL Es 36. 13 29. 80 42. 14 41, 32 31. 57 30. 14
PaO ee cists cote ar icine’ s sisi Ses G Ue oeene ee ee er ee 22 noe 29 07 | Trace. Trace.
SOR eee Sie ke cn co alt (@ynn .69| (2) 16 1.42 1. 60
TENTION aes Mae Sec deen tee eee PRL eee 55.72} 63.37 50. 95 51.58 61.09 62.59
97. 82 98. 80 98. 88 99. 23 99. 03 99. 95
COs meeded teens foc 3.68 chases Posey eo ee 31. 65 25. 85 37. 29 38. 69 29.17 27.90
Oreanicmmatter etc; 2a2heseoeeee ecm cen ee ee 24. 07 37. 52 13. 66 12. 89 31. 92 34. 69

i 8 9 10 11 12
SIO Fea. Sete ae ee asa ache Mt Cent SALE oe 0.36 { 0. 10 0. 26 0.17 0.35 0. 00
(AIRS) .Of eo 2 Rt RE a Ce \ : . 67 . 67 1.99 40 1. 64
MeO rs atte ncacan ieee eee et a hs See 4.77 3. 06 4.99 202 3. 44 2.85
CaO esses aponoks sees Sideinle Giahei='n Soe aimee see) cig hie = AS 37. 20 47.12 42.18 33. 70 46. 21 36. 44
BOs: arias eee cen + isles PAROS ee OE eee ee .05 | Trace. (?) Trace. (?) Trace.
SO; Sirislereis sia unlalata\a\ais[«/<isicinis ste e stnfaiz elapaichaleisie arate isieie ee -90| ‘Trace. (2) Trace. (?) Trace.
TEMIMON ee shee sa ac co koe EME LA kd Se ea 56. 04 47.96 52. 11 61.51 49.74 58. 76
99. 32 98. 91 100. 21 100. 09 100. 14 99. 69
COxmecded Feo ees. ok eee, erect ee 33. 93 40. 39 38. 18 29. 47 39. 73 31.77
Organic matter, (ete. 2)/35 2 os tritide dase else eee eee 22, 11 7.57 13. 93 32. 04 10. 01 16.99

13 14 15 16 17
ro) LO Bae eee ity eT 505 6 nee tes BV a ee 0. 00 0. 00 0.00 0.00 | 0. 00
(ABR G)Ooseece. ceec a acto eeee BS aE a SOE Betas Asti dees 67 . 95 89 . 94 2.58
2 URS AAS Re ESSE airicits-) SS a Soda rcicas 2” 3. 04 | 3.18 5. 20 4. 83 3. 61
CaO Saree ee bo pecs See ee cle ee an tes 50.09 | 44. 80 39. 09 40.91 35. 95
bP 0 Pic ee eres ee aL eS ee Trace. Trace. Trace. Trace. Trace.
SOS oa iae cous oi aR ns Te oe ao eee eee ee nee ae Trace. | Trace. Trace. Trace. Trace.
TgnitOn's scons ccececa ne cee ne eee ee 46.05 | 50. 02 53. 87 51. 43 55. 21
99. 85 | 98. 95 99. 05 98.11 | 97. 35
CO; needed oh. ancitenana- 6.2 eee Ce eee See 42.79 | 36. 70 36. 43 37.45 32. 22
Organic matter vetoes: ssccss 22sec see ee eee 3. 26 | 13. 32 17. 44 13. 98 28. 99

ECHINODERMS.

Reduced analyses of ophiurans.

31

L 2 | 3 4 5 6
SHO} woth a2 sBRAs doco ¢ Gabe Spr ese bese ee 2.39 | 1. 76 IE a7) 0. 24 0.16 0. 66
(Al, eR) poeceretcneetale egsteta faye a afer <Iia saro/e ine ties tows eee es . 84 eae? . 62 aati . 70 | . 85
MegCO,. ..-- soctod: tse. “SAS Ags SEE scr 4a aa 9.53 | 9, 66 | 9, 84 14.08 | 14. 56 | 14. 95
CaCO wide fe Bp ac SOC n ee or aoe een Gee DS eer ees 86.60 | 84. 36 87. 65 85. 09 81. 02 | 79. 37
(CE AIEN Os Bo aReS ae asa aed SES enaeeee RAE eat fore oicte 64 1.14 74 .18 | Trace. Trace.
(CO): gos oa l45 Sea hse eo GRSS hoes specs Cpe eSeeee (?) 1. 91 (?) ~30) | 3.56 | 4.17

100. 00 100. 00 100. 00 100.00 | 100. 00 | 100. 00
7 8 9 10 ibility
——_ a al _
SOME ees a IAs coos S are ceclernae eeietey elcieite wisi statmeieie(S e/a. 0.47 0.11 0. 31 0.25. | 0.39 | 0. 00
(UNIAN) ORS oO es Fes Bee ome bans AE Heimer cee eee ts S78) ovlih 2. 92 . 44 2. 26
NERC Oho bok aoe ete BEBE EER Se Cayce o.ucp pean 12. 97 7.04 | 11.68 8. 39 8.01 8. 24
CEO sod aU Scone san NOBSA SB pneboechse.ccsedq eso ces aeeE 84. 44 92..12° | 87. 24 88: 44 91. 16 89. 50
Ca,P,0g bi Sc CO ORO CED CER SER Oo Cor oc Un6 HC He CA SaB REE . 14°) “Trace:.| (2) 5 |) “Brace: (?) Trace.
WES ye noacas onset aemietiss “aoe ee eee alias Se sos (ps 1-98 | Trace. (2) i dipacess |e 1( 2) Trace.
100.00 100.00, 100.00 100.00 100.00 100.00
| | |

| 13 14 15 16 17
SIO saeco eee siaks = 5 55 Ses Somes we eeees selec pieces 0.00 0. 00 0. 00 0. 00 0. 00
(Al, es (ORE A SOROS 22 cir) SEU RSM aed Oe ee ea . 69 | 1.08 1.09 1.12 | 3. 47
MESO eee yo, le Sel. et EE celta dees hess. 6. 61 7. 62 13. 38 | 12.05 | 10.19
CaCO, peolsegct Sa Cteonocaks Jocc 7 ol sess de seen re See eee 92. 70 91. 30 85. 53 86. 83 86. 34
Ca,P.0, Ae SERN EAR REE a0 2 ois So DEMO Ieee ne eas Trace. Trace. Trace. Trace. Trace.
(CES Oe ope te sane SCeBensG se sars- Coc cen SCGr Bee noee te eee | ‘Trace. Trace. Trace. Trace.| Trace.

| |

100. 00 | 100. 00 100. 00 100. 00 | 100. 00

|

Some of these analyses are obviously incomplete, for phosphates and sulphates were not

determined, because of insufficient material.

ments, and in that respect the analyses are not altogether satisfactory.

A number of the specimens analyzed were frag-
In the following table

the data are presented as in the tables for the other echinoderms; the figures for the two speci-
mens of Ophiomyzxa flaccida are averaged. The localities and latitudes are abbreviated as usual.

Percentage of magnesium carbonate in ophiurans.

Locality.

Ophiomusium lymani.....-....... (allapacoss 2.25: 23...
Ophiocoma aethiops. -..-.-........ Gulf of California.......-
Ophiocamax fasciculata. ............| Caribbean Sea. . -

Ophiopholis aculeata.............. (Uinalaskas 25%. 203 =. 22
Ophioglypha lymani........-...... (Clit 32 eae ae
Caagurics phalal euenemis J osse. 2 Papen sce tse. )i2 =~ 22 2.
Gorgonocephalus arcticus. ..-...._. Caz Cosls he eae
Gorgonocephalus caryi.............| Alaska...-..-..--------
Ophioglypha sarsii. . -......-..222.) Riche) 174 (i
Ophioglypha liitkemi.......-. 2.2... M@siiformigs = ..2~.-. 252
Ophiothnix angulata................ Vi ee i arr
Ophiocoma erinaceus. ............. Liye ee
Ophiccoma pumila:<-< 22 32-2. MOMS Ote cine, lete =~ Ja/s
Ophionereis eurybrachiplax.1......| Japan...........-.-...-
Ophioderma cinereum............. Ole brass sae st ss ses:
Ophiomyxa flaccida................ ANS Eon ys ee eee

Latitude.

Temper- 7

Depth. | “Stile. | MgCOs.

Meters SiGe Per cent.
5. 1, 619 2.9 6. 61
N. (?) (?) 7. 04
N. 381 10. 4 7. 62
N. (?) (?) 8.01
Ss. 355 lL 8. 24
Ot cae Od 8.39
N. (?) (2) 9.53
(2) (?) 9. 66
N 5d3, 5 9. 84
N. 113.5 (2) 10. 19
(?) (?) 11. 68
(2) (2) 12. 05
N. Shoal. 28 12.97
61 (?) 13. 38
N. (2?) (?) 14. 08
N. Shoal. 28 14. 75

In this table the relation between magnesium carbonate and temperature is less regular

than it was with the other echinoderms.

but the last six in the series show percentages characteristic of warm-water organisms.

106135—22——_3

Some of the low percentages are unaccountably low;

One

32 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES.

analysis of an ophiuran, Astrophyton, by L. Schmelck,” remains to be noticed. The accompany-
ing data are as follows: Locality, station 37 in the North Sea; latitude, 78° 48’ N.; longitude,
8° 57’ E.; depth, 199 meters; bottom temperature, 1.1° C.

Analysis of Astrophyton.

|
Actual Reduced
| analysis. . analysis.
SiOse. snc she See en = Ears SIS ee Mrace:|| (SiO sa mesweerer = s. =)... Se ee ee ree ree Trace.
AV O ek = ak. SS a ne eo i ee Are. cews| PANS Oeeer eee a, . 7... Soc eeeeees eee cee Trace.
Wer ss fe oateeeR an Beers esis 2 ees Ox31 Bet Ope heen... oe eee emery 0. 37
MoC OS at Nee me eee eke ja owes sess 7.00) (aM COke eS oo c.. ss. ss seats sates tes See 9.11
CaO) sates Peete eer A oiare notaries TALS Dr Ml CACO aaa F..-'. 5 2.0.5 Ae eer pete ieee a 89. 67
(OUYS| 0 Fees Sp ser a eet US Seer ee ee Sr 7 S@aSOleecc. 2.2. 5. Soe cane nce eee eee eee . 85
Bi Oee cae soe eee See: cele p ses oeenioee DracepileGanke@e. 2: - 2. Say See eee eee eee Trace.
Oreanie matter q. Sees ewe h in ok eee 17.57
100. 01 |) 100. 00
|

In this, as in Schmelck’s analyses of starfishes, the magnesium carbonate is high for the

temperature given. é
5. HOLOTHURIANS.

Although the holothurians are of slight 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, but their organic matter was so vastly in excess of their hard parts
that the analyses, with one exception, are far from satisfactory. However, the data, which are
not without some significance, are as follows:

. Holothuria floridana Pourtalés. Fajardo, Porto Rico.
. Cucumaria frondosa (Gunnerus). A northern cold-water species, precise locality unknown.
Trochostoma intermedium Ludwig. Off Cerros Island, Calif. Weight of sample, 0.2204 gram.

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 gram.

won

~

Analyses of holothurians.

|
| 1 2 3 4
SHO) aaa secentee eee 0. 08 0. 27 5. 56 6. 93
SG 0 aaa -18 25 8. 40 6.31
MEOH os eee se 3. 48 41 1.81 1.36
CRD Ee ae es ee oe eee 29. 19 1. 10 4.91 1. 44
1 Ch pe 3 oer de lege Trace 85 3.15 1. 86
SOnseewtes So eee 74 1.29 (2) (2)
WomVGOMS ie .=. 2 sips site cain 69. 93 95. 7 76. 71 82. 2]
99. 60 99. 88 100. 54 100. 11
CO; meededese- a5-- ~~ 28 23. 21 (?) | (?) (?)
Organic matter, ete.......| 46.72 (?) (2) (?)

In No. 2, 0.31 per cent of matter soluble in water was found, which raises the summation,
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 becomes—

SIO} -- ced. Ae Hedk Saws ark Se ees Se Se Se ao Se ao On 0 ee ee ne 0. 15
(ATF e)sOg. = 223 otarcins oie aboeits loin seis Se ee Sia l= Se See eee ae ashe o 3 cla = 108 eee . 34
Me@Qs. 2ig:cccb bce sBc waite tes Bae one bc ene ce Se ee ae a IERIE Ialers, 3,02 oo feeae ate anees 13. 84
CaGO3 ssee-ce-| SRE ESSE A SS SARE Se Sen 8 Oe © of eee 83. 29
(OS 0 Ree er ae aE St As RE Soa a naan Gan cannon SaOROS GEES OL oto seoe oS 2. 38
CaP Opi al ssc naw de teteeh Bete acne diend scence Beet eRe ee eae tn 50 eee eee Trace.

100. 00

26 Schmelck, L., Norske Nordhavs Exped., No. 28, p. 129, 1901.

ECHINODERMS. 33

This is very near the analysis of the ophiuran from Culebra Island, which is not far from the
locality of this holothurian. The coincidence is very striking.

In the specimen represented by analysis 2 the proportion of inorganic matter was so small
that good results could hardly be obtained. The other two analyses were also made with alto-
gether inadequate material. These analyses, however, show that the small inorganic portions
of the holothurians are relatively rich in phosphatic matter and in magnesia. The difficulties
of discussing them intelligently are like those of discussing the worm tubes and the axes of
aleyonarians. In Trochostoma there are brown spots, which were already known to be phos-
phatic, although quantitative analyses of them seem to be lacking. The true character of such
bodies can be determined only by an elaborate investigation upon abundant material, a task
which lies outside of our main problem.

Two analyses of holothurians are already on record, as follows:

1. Stichopus regalis. Locality not stated; analysis by O. Biitschli, K. Gesell. Wiss. Géttingen Abh., No. 3, 1908.

2. Ash of the epidermis (‘‘lederhaut”) of a large holothurian, species and locality not stated; analysis by Hilger.
Pfliiger’s Archiv f. Physiologie, vol. 10, p. 212, 1875.

Old analyses of holothurians.

|
ie J al 2 1 2

iN ASO) Sie 2 ee es om ls aaa ee | AAT | PAGO she. .8 2 aac ree soe ee 1. 02
REXOIE Sak 5 ee eae ee oe eee oe RSS 0h EO hc, S es oe See eee SHOT eee-
CRO eae Saw hs sees | 81.54 TEX CLOANSE GaGa ern gh es Bs 328s ae
MeC Ose 8 S.-i os. Sa 8. 10 ICLP SSO) pares eee ee eS Liles eee 57
LOA RSet SRS Cocaine MPT Gace aeeSe seen eee | TOAs Organics © Ui a25. 4. sesame mee eee MOSS |e eee
UasOPGHUOMe . 2 =. cs. seen DNOOy taaeeee sss |

a Or aaah 2 ne aie he 6 ee | 96 | | 99. 66 99. 95

| | 1 I |

These analyses, together with ours of Holothuria, show that the hard parts of these animals,
like those of the other echinoderms, are distinctly magnesian. Although definite data are
wanting, it seems probable that Hilger’s specimen came from warmer water than that analyzed
by Biitschli. The sodium salts are of course extraneous.

6. SUMMARY.

From the evidence now available it seems almost certain that the inorganic constituents
of any echinoderm will have the composition of a moderately magnesian limestone. There may
be exceptions, but none has yet been found. The five tables—for crinoids, sea urchins, star-
fishes, ophiurans, and holothurians—all tell the same story, and with remarkable unanimity.
Furthermore, the proportion of magnesium carbonate appears to be a function of temperature,
the organisms from warm 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 depths
of water account for nearly all irregularities. Schmelck’s analyses of starfishes from the North
Sea 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 sesquioxides are probably
altogether extraneous, although it is possible that small quantities of them may really belong
to the organisms. As shown by Meigen’s reaction, all the echinoderms studied are calcitic, and
no evidence of aragonite in them was found.

The temperature relations shown by the analyses offer an interesting biological problem,
with which we can not undertake to cope. It is not due to differences of composition in the
solid matter 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
proportion 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 warm water. Whether this fact has any relation to the phenomenon under discussion we
can not attempt to say. We can only report the facts and leave their biological discussion
to others.

84 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES.

BRYOZOA.

Thirteen Bryozoa have been analyzed in the course of this research, as follows:

1. Schizoporella unicornis Johnston. Vineyard Sound, Mass.; depth of water, 15.5 meters; bottom temperature,
20.5° C.

2. Schizoporella unicornis Johnston. Between Johns Pass and Passa Grille Fla.

3. Microporella grisea Lamouroux. Australia.

4. Cellepora incrassata Lamarck. Northeast part of the Grand Banks. i

5. Flustra membranacea truncata 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 Rapids, Fla.

9. Frondipora verrucosa Lamouroux. From the Naples Zoological Station, without definite statement as to locality.

10. Amathia spiralis Lamouroux. Albatross station 2619, between Cape Hatteras and Charleston, S. C.; latitude,
33° 38’ N.; longitude, 77° 36’ W.; depth, 27.5 meters. Only 0.3 gram was 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’ 30”
N., longitude 68° 34’ 30’ W.; depth of water, 67.7 meters; bottom temperature, 12.4° ©.

13. Bugula neritina Linné. Florida.

The actual analyses are as follows:
Analyses of Bryozoa.

il 2 3 4 5 6
SIO}. ose meso ses ae aie ood o Seimei ee SeclomeGles nae ae eee 1. 65 2. 49 0.17 0.19 3.00 4.78
(NEL (O) 5 <a eI ge Cer me ACE rae ser 29 37 pal 19 .13
MeQuih leben Fa... te atin Sette coins sik 2. SE ee : . 28 2. 04 49 2.74 2.06 2, 32
CaO e aerate eens coos tapas tet Sep ae eae eee 50. 90 48. 17 50. 83 49. 46 30. 81 43. 08
DO 5 ciccre ie mine wien isnis vials Hale's sas sini hie aiesis'= ss) 5 Trace Trace. .10} Trace. |- - 09 11
SOE EE welt. Soo: cme ents bt Meer eee oe eee 73 mili sik) - 98 (2) (?)
PEMPONes- 2a. = 2 sha = sesh wees ae seeececer es osteo 45. 64 45. 70 46. 80 46. 03 GI4n | stereos
99. 49 99. 54 99. 29 99. 49 97. 41 50. 42
GO sniceded Se sce vases mete bet cen eee =. eee 39. 92 40. 16 39. 94 41. 26 26. 39 36. 29
QOrganicimatiter, etc. << 5-20. 452 seas teresa a5 ee 5. 72 5. 54 6. 86 5. 06 35. 06 13. 29
7 8 9 10 11 12 13

\ 0.55 |{ teat 5.75 | 0.95) 877) 631

. 64 41 1.18 75

1.16 .07 oil 3. 50 2. 55 2. 57
50.94 | 48.51 3.44] 40.83] 20.45 19. 69
Trace. | Trace. (?) . 16 . 38 . 60

- 86 1.49 (2) 1.00 1. 47 2. 43
45.87 | 44.00} 18.79] 52.66] 61.51 59. 47

99. 38

: 95.98 | 98.29} 99.51 | 96.31 91.82
GO; needed -5e..2. Seis eae ee LG. Pee Re 40.64 | 40.83 | 37.38 3.04] 35.23] 17.69 16. 41
Wrpanicimatiter, ete. 2-- sa.) Macnee asigs cee 5. 32 5. 04 6.62 | 85.75 | 17.43] 43. 82 43. 06

Reduced analyses of Bryozoa.

] | | |
oa | 2 3 4 5 6
| | 7
A NE ke TR eee ELOY 1.77| 266| 0.18} 020 5.52
Cit Ss ca Maat apie) oo Sher Pee eke ole ee ee “12 “39 |} 482 115
MeO e805. 05)5.. 05. . Sean Sige is 1 63) 4.58 Lu 6.07} 6.94 5. 62
OaGOg i 8 he 0) i, ee eS eT | 95.97] 90.90| 96.90| 91.77] 87.92 88. 44
Gaatg0s Se Se eee os ee ee oe Ae eE oe ease | Trace. | Trace. . 24 | Trace. ~ 32 BPX!
CaSOg is. ao SAG. Sa SR 08 eee 132} 2140] 145/ 176| @) (2)
100.00 100.00 100.00 100.00, 100.00! 100.00
|

BRYOZOA. 35)

Reduced analyses of Bryozoa—Continued.

7 By Meg | 10 u 12 13
| |
SiO rae ee meme na AIL REN 2 0: 3811) kG { es EE 154 16:71 12. 94
(AER Orem ee NS re ee 41 gy SP .50| 2.25 1. 54
INFRCIO), 1. = 2 Sa aa ap Bie Sea et ee 2.80 | 2.59 LUA Oban 8296 | LOI. 17508
CR CORPE Eee eet Ok Shoe NTE ER 94.96 | 95.28 | 94.86] 90.43] 86.89!) 64.51| 63.29
(OS O eee eee arene ig Upc teeing ee aries ew's oacent- Trace.| Trace. | Trace. (?) 43 1.58 2. 68
CRSO) Bae oe. Sisko oS el ee ee 1.45 |) 1540) NS80 le wey | nee2: Ove | eae 76 8.47
100.00 | 100°00 100.00 100.00 | 100.00 . 100.00 100. 00

Reduced analysis No. 10, of Amathia, requires explanation. This bryozoan was one of the
mossy or fernlike forms and contained inclusions of sand and minute 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 significance, except in so far
as 1t gives the ratio between magnesia and lime. To that extent it has some value.

Bryozoa Nos. 1 to 5 and 7 to 9 were massive coralline forms and were easily handled. No. 6,
Lepralia, was an incrustation upon a pebble, from which it could not well be separated mechan-
ically. The entire specimen was therefore weighed, the bryozoan was then dissolved by hydro-
chloric 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 sesquiodxides 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
aleyonarians 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 de-
termined.

Several other analyses of Bryozoa, complete or partial, are recorded in the literature. Two
of them, fairly complete, were made by A. Schwager for J. Walther.*’? They are as follows,
both specimens being from the Bay of Naples:

1. Eschara foliacea.

2. Lepralia sp.

Schwager’s analyses of Bryozoa.

| Actual analyses. Reduced analyses.

2 ata ie 1 2

| ia Se) a , ss
SiO et eee ee ess oe ee 0. 29 D:'30i| SiQs- ¢.ceeuls Skoucasc acetone 0. 31 2.58
CAT We) ,0 5.25 a2 ean ss enna ee 32 1 A7\| CANS Re) Osscs sae = Anerson . 34 1.58
MpOsises pee oe gs eee 1.20 9} 99 Il Me GOs. peak aie (oes ae cea eee 2.71 5. 02
CAO. .i4 Sep. oc. eee eae 50.12 RIS: f CaCO. ke gs eee mes 96. 64 90. 82
GO}: Shon .c thee. eee eee 41. 06 39. 51 _—$—$____}_—_____——
Organic matter+-H,O.............. 6. 88 7259 100. 00 100. 00

99. 87 100. 00

7 Walther, J., Deutsch. geol. Gesell. Zeitschr., vol. 37, p. 338, 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 * found 1.23 per cent of magnesium car-
bonate, and in a bryozoan from Bermuda 5.35 per cent. Several similar determinations by G.
Forchhammer*’ on other Bryozoa gave less 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 larger series of analyses.

BRACHIOPODS.

A few analyses of brachiopod shelis already on record show that they fall into two chemi-
cally distinct groups—one calcareous, the other highly phosphatic. This conclusion is sup-
ported 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 Pourtalés. Coast of Florida.

2. Terebratulina septentrionalis Gray. Eastport, Maine.
- . - a od * ~
3. Laqueus californicus Koch. Esteros Bay, Calif.
4. Rhynchonella psittacea Gmelin. Shetland Islands.
5. Crania anomala Miiller. Coast of Norway.
‘
Analyses of calcareous brachiopods.
1 2 3 4 5
= — =: > — Z
SiO; Ark 2. Mitel 52 (Qua. as ee 0. 06 0. 50 0.18 0.14 0. 21
CAT HO), O 8 goo = ote et lage he Con eee . 04 .14 Beh 528) . 26
FO ae Aaa A dD Gp MEN hepa bao eee 44 62 32 RO 3.90
CHO Pee es Set a Rehm. ae ese Ce ea eee ce oe eee 54. 96 ol. 7§ 54. 48 53. 76 48. 67
SONS Ce ashes So Ne £2 SN a et 21 . 66 21 31 .97
leAO}Rs aes 5 Set bciseawot secs Seedadeeacesesadsosses 2522562 Trace. Trace. Trace. -17 .20
Losiondgnitions . s/t. ST Se A Fi ee ee 44.35 45. 28 44. 46 44. 81 45. 38
f 100. 06 98. 99 100. 12 99. 65 99. 64
(CO pmeed eden 5. =~ teenie erate eisates aa eee 43. 42 40. 55 42.91 42.16 40. 88
QOroanic) matter, /ote.!: =. Sager mcs. te .s eee . 93 4.73 1.55 2. 65 3. 52

Rejecting organic matter and recalculating to 100 per cent, the analyses assume the follow-
ing rational form:
Reduced analyses of calcareous brachiopods.

oe at 2 By || eee! 5
SOs 2s oh eee ss oa heck eee Seen eE Re ts eee eee RE, 0. 06 0. 52 0.18 | 0.15 0. 22
CAB e) O52 Se one ah eee ee See se arch, . 04 Bal ks) . 48 | +23 | aya e
MoCOs SIN One A: ke CRs eaten. | ao a “93 1.37 .68 49 8. 63
CAO, eh ences =e orate ee eee toe th ee ee 98. 61 96.78 98. 30 98. 20 88. 59
CASO pod 2 BANAT as aso ee oe eee ae . 36 1.18 . 36 -55 | 1.72
CagP sO geri mia de cerca ss bet Oe aero en he eee eee es Trace. Trace. Trace. . 38 -57
100. 00 100. 00 100.00} 100.00 | 100. 00

8 Nichols, H. W., Field Columbian Mus. Pub. 111, p. 31, 1906. 22 Forchhammer, G., Neues Jahrb., 1852, p. 854.

BRACHIOPODS. 37

For comparison the following analyses of calcareous brachiopods, made elsewhere, are
significant:

6. Terebratula sp. Collected by Pourtalés between Florida and Cuba; 8. P. Sharples, analyst.”

7. Terebratulina caput serpentis. Locality not given; F. Kunckell, analyst.*!

8. Crania anomala Miller. Locality not given; I’. Kunckell, analyst.

9. Waldheimia cranium Miller. Locality not given; I’. Kunckell, analyst.
10. Waldheimia cranium. Collected by the Norwegian North Sea Expedition, station 255; latitude, 68° 12’ N.;

longitude, 15° 40’ E.; depth, 624 meters; bottom temperature, 6.5° C.

11. Waldheimia craniwm. Lofoten Islands.

Analyses 10 and 11 by L. Schmelck.**

Older analyses of calcareous brachiopods.

6 | 7 8 7) ai) 11
See as
SHO lst cael ee Ree pe gl ae os SO ee [gta ered ten Pech | re | 0.60 (2)
Fe,0 Sot seo EdorSoBeaBbeae Se sc oka: ce SnhessseaDrsoee ACE: | 2.24 = see see oe eee ae 40 0.15
Mec0, = OC ACRE GOCE REO eR EE cou onc oqocd see sen ae trace.) |" eat 05 3 Ae ete 1. 20 1. 40
(OCLC ORNS, SEP SAREE soe ees 9A OSs 2 98. 39 94. 6 87.8 96.2 | 96.20 95. 98
(CaS Olt stew mbes tal Soom ee eh Boel lL et 2.4 a5 9 | 85 | (2)
(Gi OSE ee une ney mer ae VON a Hl. sy Ge teas ae ee G1 Leese 28 | DSU Meee | seater
DU ertes crane iat cite scan ee roan s egw ate EPS Soot e es a es Sale ets | eerie) Se I ro | tS eee yeas Trace
(C1 OY oe ee eS See ee ECE. Sos Gonos aoe ae Se [oe Sas eeilee. jeer 0 Ae ie iO LE OR pe
WHO) SAR SS OS SOCS ISS Aka SU OO Ro COURSES SOSA UGE COS CHe] AEECEterse |--------- 1.8 6 | SRE Rad heey ahcor
PO ke c ccassbeegobehootes see ewr pe esha se Posoo oe See Se Ban Beee GPMOmre od BAPESE bed an ennsne Trace 12
(Oye een Ey zie eee Bee De Re” Soh ce See Aa Der OS eee AEE 1. 00 2.55 | 4.3 2.0 | 1. 24 1.99
100.00 | 100.00} 99.73 100.18 | 100.49} 99. 64
| |

With these analyses ours agree in a broad, general way, although the older 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 Crania,
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 Hilger of shells supposed to belong to this
genus indicates that they are phosphatic and practically identical in composition with those
of Lingula. The authenticity of Hilger’s material is questionable, and his analysis will not
be reproduced here.

Four analyses of shells of phosphatic brachiopods have been made by us. As these shells
contain a large amount of organic matter, which possibly varies with the age or maturity 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 follows:

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) pyramidata Stimpson, coast of North Carolina. Organic matter, about 37 per cent;
analysis incomplete for lack of sufficient material.

Analyses of phosphatic brachiopods.

if 2 3 4

DIOR see oan Aceh cess a 0. 91 0.50 0.85 0.49

(VITAE) (0) a Re Sepa 54 29 ics ili paallgg ls

pCO sere ete eos. 8. | 2.70 .79 6.68 eval
(COO aga * ee eae 1.18 4.25 8.35 | (?)
CAS OSPR ees) ss ecce 2.93 4.18 8.37 (2)

Ca,P,0; >t Cea eae STRATE! 89.99 75.17 74.73

100. 00 100. 00 LOO! OOM iE sane s<
30 Sharples, S. P., Am. Jour. Sci., 3d ser., vol. 1, p. 168, 1871. = Sehmelek, L., Norske Nordhavs Exped., No. 28, p. 129, 1901,

3 Kunckell, F., Jour. prakt. Chemie, 2d ser., vol. 59, p. 102, 1899.

38 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES.

These analyses are noteworthy on account of the unusual proportion of caleitum 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 ovalis. Hawaiian Islands; T. S. Hunt, analyst.**
6, 7. Lingula ovalis. Locality not given; A. Hilger, analyst.
8. Lingula anatina. S$. Cloéz, analyst:*° recalculated to 100 per cent after rejecting 42.6 per cent of organic matter.

Older analyses of phosphatic brachiopods.

5 6 ji 8

BIO3s shawnee estore eee cette ee eee 0.18 0.17 Trace
MoCO gsi. ook ae eee 2.94 Bp epee ea
CaCO ie ee toe ieee Dice 11.75 10. 76 10. 86 12.19
Gas Pesci sie Ta fea oe 85. 79 84.94 85. 24 77.17
WAH OF 6on os Soo esecocro|-Serscccon|ossse- fet eeeeeeec 7.03
PePO pp estes sae Wace | Soe ee | iia) 76 3.61
MeO 23 ieee eee ene PEGI | acs Aaa Bee Eo Se cee

100. 34 99.59 | 100.00 100. 00

The relatively high figures for calcium carbonate shown in this table are doubtless due to
the neglect to determine sulphate. The analysis by Cloéz differs from the others principally in
form—that is, in its mode 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 matter. 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.

Numerous analyses of molluscan shells have been published, and they show remarkable
uniformity of composition. It has nevertheless seemed desirable to make a liberal series of
new analyses, which are best classified into groups. Sulphates were not determined, except
in four analyses.

1. PELECYPODS.

The pelecypod shells analyzed are as follows:

Astarte crenata Gray. Off Marthas Vineyard, Mass.; depth of water, 668 meters; bottom temperature, 7.2° C.
Callista convexa Say. Vineyard Sound, Mass.
- Macoma sabulosa Spengler. Massachusetts Bay; depth, 825 meters; bottom temperature, 5.5° C.
Fecten dislocatus Say. Charlotte Harbor, Fla.
. Pecten ventricosus Sowerby. Head of Concepcion Bay, Lower California.

6. Venericardia ventricosa Gould. Off Point Conception, southern California; latitude, 34° 25’ N.; depth, 447
meters; bottom temperature, 7.3° C.

7. Cardium substriatum Conrad. Long Beach, Calif.

8. Calyptogena pacifica Dall. Clarence Strait, Alaska; latitude, 55° 46’ N.; depth, 589 meters; bottom tempera-
ture, 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.

oR wo to

33 Logan, W. E., and Hunt, T.S., Am. Jour. Sci., 2d ser., vol. 17, p. 237, 1854.
“ Hilger, A., Jour. prakt. Chemie, vol. 102, p- 418, 1867.
% Cloéz. S., Jahresb. Chemie, 1859, p. 642; from L’Institut, 1859, p- 240.

MOLLUSKS,

Analyses of pelecypod shells.

39

|
1 2 3 4 5 CAR ep 8 9 10 i
SiQsse e poece eee 0.25 | 0.18 0.29) 0.31 | 0.14 0.12 0.11 0.08 0.33 | 0.09 0.00
(Al, Fe),0;- ------- .08 -ll » 22°) . 08 15 08 09 04 46 | . 08 . 08
Mo Owe ae eieere 2.5 - .00 | Trace. 00 | . 46 . 34 .00 | Trace. .00 | Trace. | 00 32
(C0) -ec oSowesaunace 53.92 | 53.67 | 43.77 | 53.68) 54.13 | 54.16 | 53.99 53.95 | 51.18 | 53.36 | 53.80
1220 Fer sane Sob be See Trace .03 | Trace. | Trace. | Trace. | Trace. | Trace. | Trace. -17 | Trace. | Trace.
Ignition........---- 44.79 | 44.77 | 44.62) 44.34) 44.36) 44.65) 44. 85 | 44.93 | 46.67 | 45.59 44.10
99.04 | 98.76 | 98.90 | 98.87 | 99.12 | 99.01 99.04) 99.00) 98.81 | 99.12 | 98.30
CO, needed.....-.-- 42.37 | 42.10 | 42.25 | 42.69 | 42.90 | 42.55 | 42.41 | 42.39 | 39.86 | 41.93 41.73
Organic matter, ete.| 2.42 | 2.67 2.37 1.65 1.46 | 2.10) 2.44 2.54 6.81 3. 66 2.37
} |
Reduced analyses of pelecypod shells.
1 2 3 4 5 6 7 8 9 10 11
— | = = ——- ——_— > —
| |
HO psiseeecte ee apoo 0. 26 0.19 0. 30 0. 32 0.14 0.13 0. 11 0.09} 0.36} 0.10) 0.00
(ANGRC).O; = see 09 a2 eS 08 cals G3 09 VO43i/men 250!) 08 08
IMiGO eeaeeceae cae .00 | Trace. 00 1. 00 avRil .00 | Trace. .00 | Trace. | 00 . 70
MaGOysessarteee == 99.65 | 99.62] 99.47 | 98.60] 98.98 | 99.79 | 99.80 | 99. 87 | 98.74 | 99.82) 99.22
Wank, O75 gees Trace .07 | Trace. | Trace. | Trace. | Trace. | Trace. | Trace. | .40 | Trace. | Trace.
100.00 100.00 | 100.00 | 100.00 | 100.00 | 100.00 | 100.0 | 100. 00 | 100. 00 | 100. 00 | 100. 00

2. SCAPHOPODS AND AMPHINEURANS.

Under this heading we have only two analyses to offer, as follows:

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 of scaphopods and amphineurans.

Analyses. Reduced analyses.

12 13 12 13
C11 0)5 1, Sena RneHEpoEaneec Seciacened 0. 39 (aI ASO )s pemect eecinqccessocccosseHne oe 0. 40 0. 61
(RU MANOn. : i. cc cc. eee - 26 oe | CAT Hate ncaa eae meey A eae ees 27 22
CONSE <r. Se osc eee eee 09 POS TEC One mete Saute ne See 1 Ra . 20 45
DAO Poe ana wa a ns = ole wfeierelaeefermet=iae 53. 89 BY AP" RORT OO 5 Hat E ES ASG id gape eas Ase 99. 13 98. 37
SO Meee io AR ot ee (2) DON || CaS Opes nie dttec cima ot cna tees (2) » 35
P,0; 5 SOSC Ro BERet mS agbere ae cueb0See Trace. braces ||lcCasbsO gcse = eee acbieniaie = Seree eeees Trace Trace.
ORTON esse = oles aces eee eee 44. 48 44.74

100. 00 100. 00

Cha tat 99. 41 -
CORNeEd ad: -< «2 -- noes ieee 42. 44 42.13
Organic matter, etc.....-..........- 2. 04 2. 61

3. GASTROPODS.

14. Purpura lapillus Linné. Eastport, Maine.
15. Aporrhais occidentalis Beck. Off Marthas Vineyard, Mass.; depth of water, 435.5 meters; bottom tempera-

ture, 5.5° C.
16.
Le:
18.
19.
20.
21.
temperature, 9° ©.

Buccinum undatum Linné. Narragansett Bay, R. I.; depth, 53.3 meters; bottom temperature, 9° C.
Natica duplicata Say. Cape Lookout, N. C.
Olivia literata Lamarck. Sarasota, Fla.
Fasciolaria distans Lamarck. Sarasota, Fla.
Crepidula onyx Sowerby. San Pedro, Calif.
Antiplanes perversa Gabb. Off Bodega Head, Calif.; latitude, 38° 23/ 35’’ N.; depth, 112.5 meters; bottom

22. Nassa californiana Conrad. Monterey Bay, Calif.; latitude, 36° 47’ 50’ N.; depth, 67.7 meters; bottom tem-

perature, 11.3° C,

40

23. Nassa tegula Reeve. Musgrove Island, Magdalena Bay, Lower California.
24. Nassa insculpta Carpenter. Cortez Bank, Calif.; latitude, 32° 20’ 30’ N.; depth, 165 meters; bottom tem-

perature, 9.5° C.
25.
26.
ture, 4.5°
21.
28.
29.
30.

Cc.

Pyrolofusus harpa Morch.
Plicifusus dirus Reeve.

Tachyrhynchus erosa Couthouy.
Ranella pulchra Gray. China Sea, north of Prabas Island; depth, 256 meters; bottom temperature, 14.2° C.

Turritella gonostoma Valenciennes.
Volutometra alaskana Dall.

Unalaska, Alaska.
Sitka Harbor, Alaska.
Avatcha Bay, Kamchatka.

Mulege, Gulf of California.
Off Unalaska, Alaska; latitude,

55

THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES.

° N.; depth, 143 meters; bottom tempera-

31. Cerithium aluco Linné. Reef opposite Cebu, Philippine Islands.
32. Strombus canarium Linné. 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
SMO 6 Se8as" Sec seeene sso 0.14 1. 27 0. 94 0. 00 0. 05 0. 33 0.15 0. 46 0.27 2.07
(AlRe),@3------.----------- -15 22 - 41 . 05 . 04 . 04 eylllt cat! 12 oe
Win Oe aeasovkooseer sees os .19 | Trace. . 12 | Trace. . 00 .06 | Trace. | Trace. sty) . 20
(GO aa ese a senctboseaem 53.34] 52.84] 52.77 | 54.83} 54.38 | 53.87) 53.63] 54.06] 53.20 51. 30
LO eeosae ee con Aceonebee (?) 08 (?) (?) (?) (?) - 00 (@2) (?) (?)
12 Oat eek ane Sino dap oamnse Trace. | Trace. | Trace. | Trace. | Trace. | Trace. | Trace. | Trace. | Trace 08
Ignition ...--.-------------- 45.33 | 44.09 | 44.49] 44.22) 44.46| 4449] 44.40] 4415] 44.84) 44.76
99.15 | 98.50] 98.73] 99.10 | 98.93] 98.79} 98.29] 98.78 | 98.60 | 98.74
@Opmecdedsece=—- =e 42.12] 41.52] 41.59] 43.08| 42.73] 42.39 | 42.14 | 42.48) 41.99 40. 37
Organic matter, ete... .--- BEA 2. 57 2. 90 1.14 1.73 2.10 2.26 | * 167 2. 85 4. 39
24 25 26 27 28 29 30 31 32 33
S10 FaSaoee sss saeaeoqsarc 0. 47 0. 25 0. 07 0.08 0.16 0. 23 2. 20 0. 00 0. 08 0.71
(UAIING) WO). Seo sot eSeoeenese 47 1. 83 17 08 . 22 . 81 61 09 . 20 <3)
INTs OA Sone core docsaS sonar ult, . 20 | Trace. | Trace. slit 44 . 23 | Trace. | Trace. - 09
GO eke Sascugooue eS ssesee 49.32 | 52.57] 53.88] 54.23 | 52.66 | 49.30] 52.63 | 54.47 | 54.04 53. 73
SOj.-..------------------- (?) i (?) (?) (?) (?) (?) (?) (?) (?)
TEXO KS 5 Sain es dgsectsseas .04 | Trace. | Trace. | Trace. | Trace. . 36 | Trace. | Trace. | Trace. - 38
Weniioness-s.=-e—-=-- 4 =~ 48.00 | 43.51 | 44.60] 44.42 | 45.83] 47.60] 43.31 | 44.37 | 4431 43. 82
99.07 | 98.47] 98.72] 98.81] 98.98] 98.74] 98.98] 98.93 98.63 99. 12
GO} needed ese seen eee ee 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.
len. 15 16 17 18 OM ZO 21 22 23
F z ai |
0.15 1. 32 0. 98 0. 00 0. 05 0. 34 0.16 0. 48 0. 28 2.19
16 | - 23 43 | .05 -04) . 04 suki = itil 21K . 35
. 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
OES coos coesseeeacseaze | (?) PLD) bat@?) 0) Sate) (COP Ved) - 00 (?) (?) (2)
Ca3P,0, .| Trace. | Trace. | Trace. | Trace. | Trace. | Trace. | Trace. Trace. | Trace -18
| =e | | | E =
100.00 100.00 | 100.00 | 100.00 | 100.00 | 100.00 100.00 | 100.00 | 100. 00 | 100. 00
|
24 25 26 27 28 29 30 31 32 33
| 0.07 0. 08 0.17} 0.26 2. 26 0. 00 0. 09 0. 73
18 - 08 208. SAY ay ea TD aA! . 40
| Trace. | Trace. -24) 1.02 | .51 | Trace. Trace. . 20
99.75 | 99:84] 99.36 | 97.00 | 96:60] 99.90 99.70 97. 82
aC?) (?) (?) Osi od (2) (?) (?) (?)
Trace. | Trace. | Trace. | . 82 | Trace. | Trace. | Trace . 85
- 100.00 100.00 | 100.00 | 100.00 100.00 | 100.00 100.00 | 100. 00

MOLLUSKS. 41

4, CEPHALOPODS.

34. Nautilus pompilius Linné. Mindanao, Philippine Islands.

35. Argonauta argo Linné; the paper nautilus. High seas, Pacific Ocean.

36. Sepia sp.; cuttlefish bone. Port Tataan, Tawi, Tawi group, Philippine Islands.
Analyses of cephalopods.

34 35 36
SIO seo ass s eee aciee cles ise 0.18 0.08 0. 00
(MIAN Oh Sewak, Ce ses Bae 14 oy . 06
IN OS Eo a SOO eE EO eS .07 2.58 66 |
ar (V0 Tes 2A eae 2h ae oan 52. 44 46.78 47. 42
1 ON: At Ghat. Bee spo SEeooeees Trace. Trace. Trace.
Tenitonse eres one es. 46. 65 49.16 51. 63
| 99. 48 98.72 99.77
CO;meededse rete See. 4-2 41. 28 42.30 38. 65
Organic matter, etc.........-.-- 37 6. 86 2.98

Reduced analyses of cephalopods.

| 34 35 36
tape al a ee | 0.19 0.09 | 0.00
(Ane ae a Te | 115 “13 06
MeCO,..-.- bates ea “16 6.02 1.62
ETOH heen ene see 99.50 93.76 | 98. 32
(tinal 15 Ooo Soe as Gerace dees Trace. Trace. | Trace.
100, 00 100.00 | 100.00

For comparison a series of four analyses of cephalopod shells by O. Biitschli * is worth
citing in full. They are as follows:

1. Nautilus pompilius. 2. Argonauta argo. 3. Spirula peronii. 4. Sepia officinalis.

Analyses of cephalopods by Biitschli.

| |
Tl iP ly, etal ae 4

| |
(0/:10)0 Hohe ee ee eee 94.75 85.07 | 90.43 87. 36
MOCO see Sasa cass scs os 16 5.08 | 46 21
Phoapuaieseeerc se secs | aa nas 3.06) 4 3. 20 2.26
GaSOR2HAOR = 5-20: 20M ele 48 1S
RW Pee een c's wis es a3 2.62 2.92 Zils 4.89
) Orpantenmmativer---<--.--5: 2.03 2.43 2.32 5. 36

WSL ate cy Se ee Sel (a ea eres as ae 230) |Joea =~ =n

99.76 100.31 | 99.39 100. 86

Exactly what is meant by “phosphate” is not clear. Assuming it to mean calcium phos-
phate and reducing the analyses to uniformity with ours, they take the following form:

Biitschli’s analyses reduced.

1 2 3 4

eat | |

ONCE eo 99.66 | 89.93 95.75 96.58 |

IVENCIOE ES ee ea Sir ee Ne 48 Tig} |

Teo). Uae Me 3.24 3. 39 2.50 |

Chino. Si alaaernee 17 | 1.46 38 | 69 |
100.00 | 100.00 | 100.00 | 100.00

36 Biitschli, O., K. Gesell. Wiss. Géttingen Abh., No. 3, 1908. Butschli also gives analyses of three other mollusks, Pinna japonica, Patella
vulgata, and Purpura lapillus, but they show no unusual features.

42 THE INORGANIC CONSTITUENTS 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. Possibly
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 com-
plicate 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 another.*?

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:

. Lepas anatifera Linné. Florida.
. Lepas anserifera lanné. Woods Hole, Mass.
. Mitella polymerus Sowerby. Pacific Grove. Calif.; latitude, 36° 36’ N.; longitude, 121° 55’ W.
. Balanus hameri Ascanius. Georges Bank, east of Cape Cod, Mass.
. Balanus amphitrite niveus Darwin. Cape May, N. J.
. Balanus eburneus Gould. Smiths Creek. Potomac River, Md.
7. Scalpellum regium Wyville Thomson and Hoek. Albatross station 3342; off Queen Charlotte Islands; latitude,
52° 39/ 30” N.; longitude, 132° 38’ W.; depth of water, 2,906 meters; bottom temperature, 1.8° C.

DO em wD

The analyses are as follows:
Analyses of barnacles.

1 2 3 4 | Be lhe) ale 7
: = im
SLO ses a scas aie BAER Cree seen ie era eek ee 0.04 0.04 0.08 0.03) 1.99) 0.39). 9 7s
UNTO 0 See as ae aa: ee a eee a3 19 .56 .30 9145|" ee 68 meee a
MG()Er <= tase tecucls Vad egeteme ean: ae cee eee ara ae 1.14 . 86 Oda SSS Mioaen TS .76 1.03
CAO Sexson gen oc See eee eee oe aap ee ae 52533:| 52.53 | 50.83.) 53)57 | 50.091) 753223) 52565

EOP eacs ode Se cts tas Sta eae ete eeetoc oie Bera er Trace. . 34 | Trace. 00
VENOM) hoe a. bk cee SA ceite wena oe ok reset 44.50 43.90 46.18 | 44.39

WOymaededseinay 2 Lace 7 hepa eter as See eRe 49.37 | 41.82| 40.97 | 42.47
Organic fete eas sete ion 2G eee ee ee ee ee ee PAyil3) 2.08.) 5.21 1.92

98.20) 98.23] 98.33 | -98. 48 | 98.33 | 98.88 99. 20
|

| 1 2 3 | 4 5 6 7
2 | i |. |
BiQ,. 2.05-caeiecees Ree hres 2 eee Seren eae 0.04 0. 04 0. 09 0.03 2.12 0. 40 \ 0.75
(AT, Pe): Ogs2 ee te ne oa ED |. 20 58 .33 15 Tt eaoe, :
MaGO;2.-. cote. cae hee ee |) (or49) eegre | 7.2810 75) sce 63el mele 65 OR
CaCO. 522s ee ee ee eee 3 Pete apictelets, ons Sa he eae a | 97.27 96.74 | 97.47 | 99.07 95.53 | 97.73 97.02
CagPaOg- oasis ons sabia inn = MOREE ioe nee eee Trace. .77 | Trace. .00 | Trace. | Trace. | Trace.
| 100.00 100.00 | 100.00 100.00 | 100.00 100.00 | 100.00
i

%7 For additional data see Schmelck, L., Norske Nordhays 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 found was 0.78,
the lowest 0.26. For ten complete analyses of oyster shells see Chatin, A., and Muntz, A., Compt. Rend., vol. 120, p. 531, 1895. For eleven analyses
of land shells see an inaugural dissertation by A. Déring, Géttingen, 1872. Several determinations of magnesium carbonate, maximum 1 per cent,
are given by G. Forchhammer in Neues Jahrb., 1852, p. 854.

CRUSTACEANS. 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 barnacle, Balanus tintinnabulus, by Biitschli,* is essentially similar to ours.
His figures are—

(CERO ho - Qolnoed cOGnEIS GCSE ic SOON BES eae eee SBS SSS SES ra Se ea ho Smee aecoas 94.44
Rat COMME CEE M8 wae. eg I 1.20
ee te od owns. «wie an kine coeronta neta Ene ee eS ae 36
(CHRO) EOS ei heb 3d 21d se ok BSS ae Oe eee SRE Binear Seseeecc cee Race oaeencmacnece 2.06

98. 06

In all the analyses calcium carbonate is the main constituent, and 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 analyzed:

1. Tryphosa pinguis Boeck; sand flea. Woods Hole, Mass.

2. Pagurus rathbuni Benedict; hermit crab. Bering Sea, Albatross station 3531; latitude, 50° 55’ N.: longitude,
174° 17’ 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 americanus Milne-Edwards; common lobster. Vineyard Sound, Mass.

5. Pandalus platyceros Brandt; shrimp. Off Monterey Bay, Calif.; Albatross station 3129; latitude, 36° 39’ 40’
N.; longitude, 122° 01/ W.; depth, 49.4 meters; bottom temperature, 6.5° C.

6. Lithodes maia Linné; 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° 31’ 40’ W.; depth, 190.3 meters; bottom temperature, 8.7° C.

9. Crago dalli Rathbun; a shrimp. Off Cape Strogonoi, Alaska; Albatross station 3294; latitude, 57° 16’ 45” N.;
longitude, 159° 03’ 30’ W.; depth, 54.9 meters: bottont temperature, 5° C.

10. Panulirus argus Latreille; spiny lobster. West Indies.

11. Chloridella empusa Say; mantis shrimp. Ranges from Cape Cod to Florida.

12. Eriphia sebana (Shaw). Chance Island, Mergui Archipelago, off coast of Tenasserim, Lower Burma. Latitude
about 10° N. A crab. be

13. Grapsus grapsus tenuicrustatus (Herbst). Aldabra Islands, Indian Ocean; latitude, 9° 26’ S.; longitude, 46°
35’ E. A crab.

Analyses 12, 13 by B. Salkover.

14. Penaeus brasiliensis Latreille; edible shrimp. Ranges from New York to Brazil.

Analyses of crustaceans.

l
1 2 ad ge 5 6 7
as este eau cei. ok aS tae
RAO g - ae St Ss 0.62) 0.16) 0.04 | 0.00; 0.32) 0.00 2.61
MINS OS 2. oo a RS AO ectee: AM ae By 115 04 | 21 26 15 .78
TPO ec SE HS np OE Vi27 | Us BBe | 12006)" 295 1e44h) a0 s 2ST
(O50). soe eo ae BO ae eae 28.63 | 29.11 | 33.42 | 31.35 18.56) 24.15 | 32. 79
12( 0) = eR Soe cr i, 2) BF oP 4.56 | 3.25| 4.26| 3.04] 4.60| -3.51 2.73
Someone rt) Giameamarmerenc ey 6 60795)" | 93| .75| .293| —.45 54 63 | 49
Tprigiipniteennc te. + nota eee ng 7 | 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
COS eee © cc. a rr 19.53 | 21.14| 24.43| 24.08| 11.58| 17.46 | 26.05
Orpanichnattornete + 2 een et 45.20 | 43.36 | 34.46| 37.74 | 62.23| 51.54] 31.02
8 | 9° | —20 Wy | de 13
= py = | = SS = hers a ee

por 6 Feet eer tre SAR) 300-2 SL a 1) S80232) 9 10:62 0.16] 0.02) Trace. Trace.
(NAYS ON ESTE chi: ee 16 | 20 | 13 | ili 4. 82 4.73
IPO pea eee es La ET. a rele ee 2.58| 1.01 3.15 | 2.60 1.19 1.57
ONO pean era Eo te i fit ak eee 31.28 9.95 | 25.36 | 15.38] 34.20 25. 30
PMO Re aL Pee sy as Sa ee 1.87 2. 65 2.09| 7.75 3.38 2.98
S10 Pee Rees Ae See AT | 47 Al 1.07) Trace. Trace.
WECV TG geese eee te, ee en | 62.29} 83.70) 66.69 72.35 53. 92 64. 87
98.97 | 98.60| 97.99| 99.34] 97.51 99. 45
(Os Ecol eke ST 2) ee 25.41| 6.20] 21.23 7.15 | 25.04 18. 83
(Grpansicunatter, ete eee ele eed 36.88 | 77.50) 45.46 65.20 | 28. 88 47.79

8 Biitschli, O., K. Gesell. Wiss. Géttingen 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 small for complete analysis. Only P,O,, 3.07 per cent,

was determined.
Reduced analyses of crustaceans.

|
pe Wee 3 4 5 6 7
Rigo or oe 5 Se Ra < Reet ee 1.12]} 0.28] 0.06| 0.00! 0.87] 0.00 3.82
(ADR 6 kOschc oo cos no tee ee Ra oe 67 27 06 184/70 31 1.14
MpUOs St. 2 167. -eta. eee en. eee seek 4.84| 5.80] 6.69! 80.2| 8.09| 8.35 8.65
aCOg > sds ui conde Daa, Bide Cb erat BO aie eee 74.64 | 78.03| 78.14| 79.50| 60.94| 73.07! 76.44
Cas Bs pale: k cee end Soe ey ee or eee 18.02 | 13.55 | 14.45 10.91 26.94) 16.03 8.738
CaSOEs. ee eee at | aig Nasa 60} 1.23] 2.46] 2.94) 1.22
| |
100.00 | 100.00 | 100.00 100.00 100.00. 100.00 100.00
| j | |
l
bere g BIS ofl iO 11 12 13
| | |
(= O82 2.94} — 0.31 0.06 Trace. Trace
alee S95 25 “50 7.02 | 8. 86
8.71 10.05 | 12.58 15.99 3. 65 | 6.18
82. 64 54, 83 76. 87 28.56 | . 78.58| 72.7
| 657| 27.44] 8.68 49. 56 10. 75 12.19
| 1.29 3.79 | 1.31 5. 33 Trace.) —‘ Trace.
| 100.00, 100.00 100.00 100.00} 100.00 100.00
| | | }
i } I |

Tn 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 made up
of hundreds of individuals, were obtained from the U. S. National Museum, as follows:

1. Temora longicornis (O. F. Miiller), 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,O,, was re-
calculated into the form of tricalcic phosphate, Ca,P,O,, and with that adjustment the analyses
assume the following shape:

Analyses of minute crustacecns.

l
1 2

| '

| ae ae 4 =r
InossTONASNIONe =: +2 eee ee eee 96.05 | O9NORMS

| Tricalcic “phosphate. . oes Sa es QUE i 7.68

| ieee ee eee Siicc Ra OR 0.92 | 0.22

| |

| | 99. 74 99. 98

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 skele-
tons 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 our analyses of
aleyonarians and echinoderms. Eriphia, for example, a tropical form, is abnormally low in mag-
nesia; whereas Chloridella, also from warm water, isremarkably high. In other analyses the pro-
portion of magnesium carbonate appears to be about normal, but unfortunately the records for
some specimens are defective as to precise localities and temperatures. Furthermore, the large
amounts of organic matter and water render the reduced analyses to some 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
the percentages of magnesium carbonate, calcium carbonate, and calcium phosphate may be
uncertain by as much as 1 percent each. In no case is the order of magnitude seriously changed,
but the accuracy 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 analyzed.
In order to test this supposition we obtained, through the kindness of Dr. H. M. Smith, Director
of the United States Bureau of Fisheries, the large claws of two lobsters (Homarus) taken at
the 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
Steiger, were as follows:

Analyses of lobster claws.

Actual analyses. Reduced analyses.
Small. Large. Small. Large.
Si@aees oe ae ee ae BolGiOLaxae eek Wane DR | a
GRIGRE)Og0 02 5 cee oe } Boe lf (AU e),O ee noc ee eae ae i Hele) \ oe
MeORs Be ett So, ! Ee 1.68 2800 MoCO2 ues acd a aS eet joer | 6. 02 | 11.51
CaO he 35s. 5 eee. 2. a 30. 60 2380 ll CaCO ska seek see oases eee 80. 52 | 64. 37
120 1 ee aR ot 3.29 DROS. | Cas PiO ge oo ee one a: 11.98 | 21. 46
SO sent a5. 5 So cee ee eee 1.45 RBOs'||' CAS Ose cee en Aomec ter eer ts era 129" | 1. 85
RENTON 2). oth wire Saree eee 63. 42 62. 44
100.00 | 100. 00
99: 48°) 99°21
WOimeeded!:-- 5: -s-aseet. ok eee 22. 66 18.17
Organic matter, ete. ..-2-.. 2222. s<< 40.76 44. 27

The differences between these two analyses are very striking and confirmed our first
suspicion. In order to make the investigation more precise, Dr. Smith had sent us from the
same 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, 114 inches; weight, 2 pounds.
3. Large lobster: Length, 164 inches; weight, 5} pounds.

Furthermore, for each lobster a large claw and part of the carapace was supplied, so that
for each animal two analyses could be made. This last precaution was taken because it was
remembered that in two sea urchins it was found that the spines and the shell differed in
composition. Is an analogous difference between two parts of the same animal to be found
among the crustaceans ?

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.

| Cara- Cara- Cara-

Claw. pace. Claw. pace. Claw. pace.
0.18 | 0.18 0.17 0. 30 0.14 0. 24
2. 35 1.96 2. 55 1.80 2.35 1. 76
27. 04 | 26. 55 23. 08 23. 53 21. 89 21.09
3. 82 2.79 6. 70 4.07 6. 06 4.48
. 40 PBY/ . 59 43 . 61 .58

63. 60 64. 90 64. 58 68. 72 66. 32 69. 49

97.39 95. 75 97.67 | 98.85 EB 97. 64
CO; meeded'e 28 ee ase cc's wine cyarstn atelelereieee ee ere ate 17. 48 18. 06 14. 39 16. 45 13. 81 14. 03
Oreanicimatierreteey 2 le Se ee eee 46.12 | 46.84 55. 19 52.57 52. 51 55. 46

Reduced analyses of lobster shells.

1 2 3
Cara- Cara- A Cara-
Claw. pace. Claw. pace. Claw. pace.
= > i

SiO. (Ad Ro),Os «tows 2 ena 0.33 0.35 0. 36 0. 66 0.31 0.57
M CO, Sapte eee tere rete nee tate aerate ie siete re Meet ee 10. 81 7.74 11. 28 8.12 10.99 8.77
CaCO gost acess = sec esaabelac Slseis sae eisisinis saree orelsole eee 72.41 78.98 55. 46 70. 58 56. 89. 65. 14
(ins Pi Ose eee Se Foon 82 ST Ae 15.21| 11.70| 30.78| 19.06] 29.49 23. 20
CaS Ope nasec oe wa te eect ce ecsemic ste aeeceewe cee aaa 1. 24 23 Zhe 1.58 py Sy) 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 mag-
nesian 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. la Medium.}| 3. Large.

MoCOs fh jceeseae ss ee neat 9.27 9.70 9. 88
ORCO. os ke eee ER 75. 69 68. 02 61.01
CRPLOM Eo. eee aires 13. 45 24. 92 26. 35

ANOpeeee oc shes eee aw 1. 24 1.85 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 only 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,*® for example, gives three
analyses, as follows:

1. Astacus fluviatilis; fresh-water crawfish. 53.27 per cent inorganic.

2. Lobster; probably either Homarus vulgaris or Palinurus. 77.06 per cent inorganic.

3. Squilla mantis; ashrimp. 37.17 per cent inorganic.

Schmidt's analyses of crustaceans.

1 x Willie ct Sele |
4

CaP, On cere eS: ese: 13.17 12. 06 | 47. 52
CACO Ree eee es... 86. 83 7.94 | 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 * are also worth citing:

1. ‘“‘Langouste”; Palinurus vulgaris.

2. ‘“Eerevisse”; probably the crawfish, Astacus fluviatilis.

Fremy’s analyses of crustaceans.

1 2
(GEYEL( Os. ceadoags6gn SGC Cob Gn Ge ene ae eine GH 6.7
CRC ORIN e808 cock siete 49.0. | 56.8
Orrani@nnaitene etsy oe \= re cee oo: < = =aiain'ernin's 44.3 | 36.5

100.0 | 100. 0

Here, again, the determination of magnesia has been neglected. If we reject the organic
matter, the percentages of calcium phosphate in the inorganic part of the shells become 12.03
and 10.55, respectively.

Astacus fluviatilis seems to have received more attention from chemists than any other crus-
tacean. H. Weiske *' 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 been studied by Agnes Kelly,*? who also analyzed a myriapod, Julus

(ulus). Her figures are as follows:
Analyses by Agnes Kelly.

Astacus. | Julus. |

|

ONON ae a ee 30.44 | 36.29 |
CIC. See 21.23 | 21.60 |
ly Oheacct cer eae 2.79 Bar|
54.52 56.26 |

49 Schmidt, C., Annalen Chemie u. Pharm., vol. 54, p. 303, 1845.
Fremy, E., Annales chimie et phys., 3d ser., vol. 43, p. 94, 1855. Fremy also gives many analyses of vertebrate bones, both recent and fossil.
41 Weiske, H., Landwirthschaftliche Versuchsstationen, vol. 20, p. 45, 1877. 0

Kelly, Agnes, Jenaisches Zeitschr., vol. 35, p. 429, 1901

106135—22——_4

48 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES. E

In an analysis of Astacus fluviatilis by O. Biitschli * magnesia was actually determined.
We append his analysis, together with our own reduction of it:

Biitschli’s analysis of Astacus.

|

Actual analysis. Reduced analysis.

|

CaCOn.. 2 eee #275108 OaGOse ees... 83.40 |

MEUOr. 2. =. 2 eee 1.196, || MeCOMMeernS. . 2.42 |
Phosphate..-.-<222 === Git, {Oath O pileeeeer a =e 55ni5, 11. 88
CENO WOE Osseo 45 58 e 1466; || CaS Opes. oe 2. 30

17 0 eget eae ae 1.34 a
Organic matter_..-.......- 40. 60 100. 00

99. 26

In this analysis the low percentage of magnesia is very significant. Astacus is a fresh-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 magnesium, but in ocean water mag-
nesium is 34 times as abundant as calcium. ‘This difference in the environment may possibly
explain the difference between the fluviatile and the marine shells, 2.42 per cent of MgCO, in
one and 4.84 per cent in the lowest of our determinations.

One more determination of phosphoric oxide in a crustacean remains to be noted. In the
shell of a lobster, Homarus vulgaris, W. H. Hudleston * found 3.26 per cent of P,O;. This is
equivalent to 7.12 per cent of Ca,P,O,, or, if the organic matter was about the same in amount
as in our analysis of the American lobster, 11.44 per cent in the inorganic portion alone. This
is not far from the figure given in our reduced analysis No. 4, namely, 10.91 per cent.

Although the crustaceans are not of great importance as contributors to the marine 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
become significant. How significant they may be is for zoologists to determine.

CALCAREOUS ALG.

The calcareous alge are so important as reef builders that, although they are not marine
invertebrates in the ordinary acceptance of the term, it seemed eminently proper to include
them in this investigation. Im many places they far outrank the corals in importance, and
of late years much attention has been paid to them. To Dr. Marshall A. Howe, of the New
York Botanical Garden, we are indebted for a fine series of alge, and to him we express our
thanks.

For the purposes of this research the calcareous algie fall into two groups. One of these,
of which Lithothamnium is the most familiar example, is highly magnesian; the other, repre-
sented by Halimeda, is almost free from magnesia. These groups are best considered sepa-
rately, and under the first one we have the following species:

1. Lithothamnium glaciale Kjellman. Topsail, Conception Bay, Newfoundland; latitude, 48° N.; longitude,
53° W.

Ss)

Lithothamnium erubescens Foslie. Haingsisi, near Timor, East Indian Archipelago.
. Archxolithothamnium episporum Howe. Point Toro, near Colon, Isthmus of Panama.
Lithophyllum craspedium Foslie. Palmyra Island, in the Pacific Ocean, west of south from Hawaii; latitude,
5° 49’ N.
5. Lithophyllum pallescens Foslie. Bay of La Paz, Gulf of California; latitude, 24° 16’ N.
6. Lithophyllum dedaleum Foslie and Howe. Salinas Bay, near Guanica, Porto Rico; latitude, about 18° N.
7. Lithophyllum antillarum Foslie and Howe. Culebra Island, Porto Rico; latitude, about 18° 20’ N.
8. Lithophyllum oncodes Heydrich. Coetivy Island, in the Indian Ocean, northeast of Madagascar; latitude, 7°
6’ S.; longitude, 56° 30’ EB.
9. Lithophyllum intermedium Foslie. Fort Clarence, near Kingston, Jamaica.
10. Lithophyllum pachydermum Foslie. Dollar Harbor, South Cat Cay, Bahamas; latitude, about 25° N.

— cot

43 Biitschli, O., K. Gesell. Wiss. G6ttingen Abh., No. 3, 1908. “ Hudleston, W. H., Geol. Soc. Quart, Jour., vol. 31, p. 376, 1875.

CALCAREOUS ALG. 49

11. Lithophyllum tamiense Heydrich. New Guinea.
12. Amphiroa fragilissima Lamouroux. Lemon Bay, near Guanica, Porto Rico.
13. Phymatolithon compactum Foslie. Torbay, Newfoundland.
14. Goniolithon acropetum Foslie and Howe. Culebra Island, Porto Rico.
15. Goniolithon strictum Foslie. Bemini Harbor, Bahamas.
* 16. Goniolithon strictum. Soldiers Key, Fla.

The actual analyses are as follows:
Analyses of alge.

1 2 3 4 5 6 7 8

DLO Sewer at Femen oe Onase ociteene sicie eas a 0.41 0.19 1. 47 0.03 1.01 0.18 0. 04 0. 09
(CAINWG) Ose ee sce ike a oleate eters 20 12 . 89 09 322 08 10 12
MONS A aee aati os feast eee ois RC OE tate 4.78 7.52 5.93 8. 64 6. 42 8.14 7,24 8.07
(CEOs RO eS ane Ae Seg ee ng SS ae 45. 4) 42.96 44.80 | 41.56} 40.39 | 40.48 | 43.34 42.57
PO pees e es eas te Aals = sect tee este ae ters Trace.| Trace. | Trace. . 03 .11 | Trace. | Trace 08
NSO SRO ee ae ee eRe Pe 14 61 .53 .15 71 WW 57 27
l2qa0 mG) 1 ames ee ne nee Eee mene 32a on ae 48.00 | 47.89 | 45.69 | 48.37 49. 94 49.99 | 48.1] $7. 96

98:97 | 99.29) 99.31 98.87 | 98.80} 99.31 99. 40 99.16
CO; mesdedi as =23522 sage emacs 40.86 | 41.69] 41.43 | 42.04] 38.31 | 40.51] 41.70 42.11
Oreanicsmatter, 6ies jason ee ens tee re eee 7.14 6. 20 4, 26 6233: alilnoe 9.48 6. 41 5. 85

9 10 me 12 13 14 15 16

IO aap coo econ Enmoe eocic ce cotan cometee 0. 27 0. 04 0.12 2. 82 0.10 0. 06 0.02 0.05
CATR GO Bye: eee meee meee oJ Oy 08 FG least 23 .04 01 a
Mig OR ee eee 2 ae sR ee ee 7.25 6. 47 8.81 6.7 4.02 8.27 10. 39 10. 93
CaO «50h cers. sa 2 sie's + bts sere Se ets nrn we 42.55 | 42.60] 41.07] 34.82] 38.19] 40.60 | 38.54 38. 03
17210) elle SRE Ses Reet fre aoo Ba geen Trace. .00 | Trace. | Trace. 10 .18 | Trace. | Trace.
DOR ee ame ee Sse cod eae eee iat eer. . 02 09 . 69 36 .52 62 . 60 61
pn tions eo. ae slaw Sti eae eee ee ae 48.63 | 49.55 | 48.77 | 48.78 | 55.42 49.56 | 49.91 49.91

98.93 | 99.33 | 99.62] 95.00] 98.58] 99.33] 99.47 99.53
GO; meededat2. -t.'50). eee eae seer ets 41.89 | 40.24] 41.58 | 34.43] 35.05 | 40.50] 41.38 41. 56
Organic'matter, eic:A-: 28... =p eae eee = = 6.74 9.31 TAS, 14.35 | 20.37 9.06 8.53 8. 35

The low summation of No. 12, a very fragile form, is due to inclosed salts, mainly alkaline
chlorides and sulphates. These were determined 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 algx.

|
1 2 3 4 SANE agi 8
A | uP, lee
SDE SRR Soe moe SI ie 0.45| 0.20) 1.55| 0.03] 1.16] -0.20] 0.04] 0.10
ud 53 0 Se i epee AS te RE 125 1S 94 “10 25 | “09 u 13
Mp CO estes 205 aes SS ee aS 10. 93 16.96 | 13.09 19. 60 15. 46 19.03 16.35 18.17
CACO ss Soe plac os, 2 ctecechiee Ose erate. 88.11 81.59 83. 47 79. 92 81.48 | 79. 85 82.46 80.93
Cane Ogee cect ans ost. ee rep ene eee Trace. | Trace. | Trace. 08 26 | Trace. | Trace 18
CaSO sttes: .Jesa2202. oe. SE eee - 26 tee: | . 95 wat 1.39 .83 1. 04 | 49
100.00 | 100.00 | 100.00 | 100.00 | 100.00 | 100.00 | 100.00 | 100.00
|
9 10 ll 12 "|, als 14 15 16
SIO Sih sot eas Lc ae EE 0.30| 0.04] 0.13) 3.50) 0.13] 0.07] 0.02 \ 0.06
CAIRB IS) Of: 2 base ok See eR OMS 2. = 275K | 220 . 09 Ld, 1. 62 | . 30 | .05 | .O1 A
MpCO sin asosi=<icceccce eee eee Oe ..| 16.59 | 15.08 20.02 17.47 | 10.93 19. 24 24. 00 25.17
WaCO nab. s sa.2 fos cee ee EE oases 82.85 | 83.68 78. 43 |; 76.23 87. 21 79.05 74. 85 73. 63
(CEA 4 0 PA Reet Godan cos aenCenine Trace. | Trace. | Trace. | Trace. . 29 | .43 | Trace. | Trace.
WAS Ole. soo ccies =o cee es Ieee ae ee cas cies oe . 03 1.11 1. 25 nea alae! 1.16 1, 2 1.14
100.00 | 100.00 | 100.00 100.00 100.00 | 100.00 | 100.00 | 100.00

50 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES.

In No. 12, Amphiroa, the high silica and sesquioxides are evidently impurities. If they
are rejected, the percentage of magnesium carbonate rises to 18.41.
Of Halimeda, four species were analyzed, as follows:
17. Halimeda opuntia Lamouroux. Key West, Fla.
18. Halimeda simulans Howe. East of Guanica Harbor, Porto Rico.

19. Halimeda tridens Lamouroux. Cayo Maria Langa, Bay of Guayanilla, Porto Rico.
20. Halimeda monile Lamouroux. Same locality as No. 19.

Analyses of Halimeda.

| 17 18 19 20 |
| |
SiOf4. 2. RP 0.37 0.44 | 0.88 | 1.90
14m (CA Bile) EOss see is eee . 20 .18 D8 | 1.10
MoO! i... cabal aad aaneee ‘01 19 age oe A
CaO! ..cc2e tek ee a. eee. 50. 32 50. 20 45.32 | 48.91
POs. Mies See RE ie ee Trace. Trace. | Trace. | Trace
SOs. csi ties pentane! 07 23 49 02
Tonitione..2--c-4e t-te ee 48.20 | 48.38 51.94 | 47.56
99.17 | 99.62 | 99.65 | 99.24
CO, needed’. --<....2-=--=- 39.47 | 38.53 35.83 | 38.35
Organic matter, etc. ....-- 8.73 | $85} T6017 | grat

Reduced analyses of Halimeda.

ge |). 18 19 20
e | |
SiOR att ee | SS 0.41 | 0.48 1. 04 2.11
(AM e)O;) Se ah seen | 22 . 20 . 68 122)
Mig COR mene Same: oe 2 eens | (re 44 1.09 1.04
CaCO, ase FOR 5 Lea oh eal 98.45 | 96.21 95. 58
Gash Opa bancsseoeete ee | ‘Trace. Trace. | Trace. Trace.
CaSO fea Vee: Sie A .14 . 43 .98 .05
100.00 | 100.00 | 100.00 | 100.00
!

After allowance is made for obvious impurities the essential identity of these four analyses
becomes clear. The inorganic portion of Halimeda consists of calcium carbonate and an insig-
nificant amount of magnesia. In the first group of alge, Nos. 1 to 16, magnesium carbonate
is conspicuous, more so than in any other class of organisms so far analyzed. In Goniolithon
strictum especially it ranges from 24 to over 25 per cent, or more than halfway to dolomite.
These alge are probably the largest contributors of magnesia to the marine limestones.

It is also noteworthy that the two alge of the first group, Nos. 1 and 13, are from the cold
waters of Newfoundland, 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 levels. It is desirable that Arctic species should 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 alge were
submitted for analysis by T. Wayland Vaughan. The species, with analyses by A. A. Chambers,
are as follows:

1. Goniolithon fratescens Foslie. Cocos-Keeling 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. Cocos-Keeling Islands.

CALCAREOUS ALGAE,

Analyses of algx.

1 | 2 | 3
| o Bei |i rl sa
Si0,,Al 20./F e. 20s- doce 0. 07 0.11 0. 28
MgO. ie 6. 29 5.71 7.09
NOM ence arenes 2k. 2.5. 46. 16 42. 39 45. 92
EMO eine cactepra cic Trace. Trace. Trace.
S(O) esas eee er 00 00 00
Renrtlonyatst esse ee cklt.,-- 5532) 46. 70 50. 97 45. 72
99. 22 99. 18 99. O1
Opmecdedae Uareum scans). St). 43. 19 39.59 | 43. 88
Organic matter, etc............- 3.51 | 11. 38 | 1. 84
Ratieed gists ae CEES

1 | 2 3
SiOS ATO Mne Os neees. 0.07 | 0.12 | 0.29
WOO en acoscen ce SSE ae eBeOes 13.80 | 13. 66 15. 33
CaCOg, digcttns¢ gosh seaaweoonede 86. 13 | 86.22 | 84. 38
Ca,P, Os Sere ace tees ee ly a Drace: yy “Traces | Trace.
CaS Orme clr daicrs\orsecss -00 | .00 | . 00
100. 00 | 100. 00 100. 00

These alge are much lower in magnesia than our specimens from similarly warm regions.
The two species of Goniolithon contain little more than half as much magnesium carbonate as
That this difference might be due to differences
of age or maturity seemed to be possible, and therefore two more alge were obtained from
Dr. Howe and analyzed by R. M. Kamm with the following results:

the species from Florida and the West Indies.

1. Lithophyllum pachydermum, Dollar Harbor, South Cat Bay, Bahamas.
la. Upper or younger layer, solid.
1b. Lower or older layer, porous and somewhat discolored.
2. Goniolithon strictum, Bemini Harbor, Bahamas.

2a. Upper or younger branches.

2b. Lower or older branches.

Actual analyses of alge.

la 1b 2a 2b

SiO peste eee acee = 0. 04 0.09 0. 02 0.08
(CRIGPOOMS Bie. 2 inl 14 32 L537
Mo@ Senet ree eck. sd 10. 69 6. 54 9. 64 10, 26
Ca@Peeeeeericerics Secs. =1- ~ 37. 66 44, 88 37. 64 38. 25
PiOpeetenon a see etioass cee Trace. Trace. 00 00
Sma aes Th 1 Ny 7 63 .67
Tomiloue acres = esc is1te= 50. 64 47.51 51.14 48. 76
99. 93 99. 87 9939 98. 59

CQ; meeded)..-<-2-.----=--- 41. 42 42.08 39. 83 40. 97
Organictetesess nese. 2-2. - 8. 72 5. 43 iil, Bi) 7.79

Reduced analyses 3; alge
|
la lb | 2a 2b

SiO,S=2- fio. 0.04 | 0.09 0. 02 0. 09
(Al; Fe),Ofieet.-..52.0... 12 Bt _36 63
MEGO eget .55.-... | 24.95 15.43 | 22.98 23.74
(07010 eo Eee 73. 65 83. 06 . 75. 42 74. 29
CasP.O setae: o2--ses-2=--- Trace. Trace. | 00 00
CAS eae se ca alas Se oo 3.0 1. 24 eR | 1. 22 1. 25
| 100. 00 100. 00 100. 00 100. 00

51

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 alge by Prof. A. H. Phillips. In one, from Samoa, he found 36.36 per cent of
magnesium carbonate, an astonishingly high figure. In the other, from southern Florida
or the Tortugas he found, in percentages, CaCO,, 73.28; MgCO,, 25.32; and Ca,P,O,, 0.35;
total, 98.90. This analysis agrees well with ours of Goniolithon from the same region.

Of the earlier analyses of calcareous alge those by A. Damour * are the most complete
and most nearly comparable with ours. Five species were analyzed, as follows:

. Lithophyllum sp. Mediterranean Sea.
. Melobesia sp. Coast of Algeria.

. Amphiroa tribulus. Antilles.

. Halimeda opuntia. Red Sea.

- Galaxaura fragilis. Antilles.

om wh Ee

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 algz.

1 2 3 4 5
CaCO Ree ccs ae strc ore Soe ite hee hic wie ROR RC CE eee 77. 36 72.78 70. 83 86.17 72. 56
MoG Ober Hse etie. oka. Se8el eT a8 S08 Aas ee ee NES 2 12. 32 | 16.99 . 56 . 86
Nay Orie diate cccehe cas nee ek bis teens at eee oe .d0 1.75 | 89 1.13 73
1) Ee a SCE TERE Leah PT Mac) il sii . 65 | 39 5 1.02
FOE Ooo a ee secre ee Cee ea 08 ETO Ieee alg cl, ie
Oe eRe re Swe aia ncicie Sie a ea eo cS wok 2 .95 IGP) || 93 (2) (?
PROC Eee te Gatti ck Aen oe ene, ae a 32 38 | pal lee Go) (2)
0) St aE Seis Se ee eine SCPC CACO Ea SEM OM Mena Ss Seco Sen 60 . 34 53 . 84 ilealy/
CASON Soe cots curoouesceliene vogue tibenete), oan |e eee | 20 55 1.80
Oreanieimatiterss = seccccetie solace ceo ee ee cine ee 4.70 3.95 | 6. 40 8. 30 17. 50
1 OS ASG ee Ten Acre aE oe eile cake 1.46 1.40 | 1.38 . 90 95
SiOgidc Boscubas bocce coh eh dum. t. «ce Tees ene | So eSNG Mire eh tl | ree 2. 20
Dandie eee Soe shee a tee ons ore see ere oe 1.36 BS eo cocecon|soccassa05|saadoocese

98. 97 99. 30 | 98. 81 98. 99 98. 79

In these analyses Na,O, K,O, Cl, and SO, evidently represent sea salts. Rejecting 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.

100. 00 100. 00 100. 00 100. 00 100. 00

4° Damour, A., Compt. Rend., vol. 32, p. 253, 1851.

CALCAREOUS ALG. 53

Three of these algw are highly magnesian; Halimeda and Galaxaura are not.
Two analyses of algw by A. Schwager were published by J. Walther * in 1885. There is
also one by C. W. Giimbel,"” and these three may be combined in one table, as follows:

1. Lithothamnium sp. Bay of Naples; A. Schwager, analyst.
2. Lithothamnium ramulosum. Bay of Naples; A. Schwager, analyst.
3. Lithothamnium nodosum. Locality not stated; C. W. Giimbel, analyst.

Actual analyses of alg.

|
i | 2 3
= : _|
|
Se Ce ee 1,50) ihn" paclSBaa eaele eases |
/ NWO PSE SP eRe Se oe cee eee eoor 3.36 | 3. 61 |
1 Oa) OA Pee eae scm 41 2.55
MnO.... REP tst(s- -- Trace. | Trace. |
MO pees eee sete = == 1.90 | 3. 06 2.66, |
CaQiew. Fosse eeee ane e snes =e 48.09 45. 88 47.14
COR eee ceere eerie teenie = c-' 39.87 | 39.41 | 40,06 |
1240) aa eae schcae bes) nen EBOAeOd PROOenO Ont Meroe s ZOG ea
Organic matter+H,O...------- 5.06 | BED ip ose mer aetein
msolubles sees -eetesss ees --12|4=- == yr eee en soar 4.96 |
HH Ojand| losseee a2 22 -2<-- ------|---- 2 ee gh ne ea 2.57
100.15 99.85 | ° 100.00 |

Reducing these to uniformity with our analyses we have the following table:

Reduced analyses of alge.

a dba'|
| 1 | 2 | 3
|
SHO hss. 8eer CU She REA ee eee ane 1. 68 O03: tse eeee ae

UNIAN) 00a eee 3. 83, 4.26 | 2.76
WI C(0) 4 Sao caseauseee ede sees 4.19 6.81 | 6. 06
(ORKC(0)t J Jas eabeannooec Beeson | 90. 30 86.90 | 91.04
CaeD Oe i ES ok Weare leek Uae [OE eee | 14
190. 00 100.00 | 100.00

In these analyses, all of Lithothamnium, the percentage of magnesium carbonate is remark-
ably 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 may 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,* assisted by R. Mauze-
lius, 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.
The data given are as follows:

| | CaCO3. MeCO.
| qNewresnrie) oti) bevva |, ntact
Lithothamnium sp. Java Sea...------------ 72.03 3.76
Lithothamnium sp. Galapagos Islands.. - - ---| 83. 60 6. 53
Lithothamnium sp. Spitzbergen.----------- 84. 83 8. 67
Lithothamnium polymorphum. Kattegat..--- 74. 22 9.10
Lithothamnium sp. Honolulu.-.-.----------- 84. 01 9.39
Lithothamnium ramulosum. Naples... ------ 63. 00 9. 46
Lithothamnium soriferum. Arctic Ocean..--- 80. 90 9. 56
Lithothamnium sp. Bering Islands...-------- 74. 24 9. 94
Lithothamnium racemus. Naples------------ 77.39 11.33 |
Lithothamnium sp. Bermuda...------------ . 82.44 12.37 |}
= Lithothamnium glaciale. Arctic Ocean....--- 83. 10 13.19 |

46 Walther, J., Deutsche. geol. Gesell. Zeitschr., p. 338, 1885.
« Giimbel, C. W., K. bayer. Akad. Wiss. Abh., Math.-phys. Classe, vol. 11, p. 26, 1871.
#8 Hégbom, A. G., Neues Jahrb., 1894, Band 1, p. 262.

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. Hégbom, however, points
out the importance of these alg in respect to the formation of dolomite, which was the real
subject of his investigation. In Halimeda sp., from Labuan, only a trace of magnesia was found.

In Lithothamnium racemus, from the Bahamas, H. W. Nichols ® 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 cent—
was found by E. W. Skeats °° in Lithothamnium phillipi var. funafutiensis Fosle. The speci-
men was taken on the atoll of Funafuti.

In a very fresh Halimeda opuntia Skeats found 0.60 per cent of MgCO, and 86.50 of CaCO,,
but in a mass of fronds dredged up from a depth between 50 and 60 fathoms the percentages
were 4.0 MgCO, to 93.59 CaCO,. The increase in magnesia may have been due to concentra-
tion by leaching. These data are given by J. W. Judd,*! 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 MgCO, and
98.32 of CaCO,. Judd also quotes an analysis of Halimeda, cited by Payen in his Flora, which
contained 5.50 per cent of MgCO, and 90.16 of CaCO,. The character of the specimen repre-
sented 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 ® gives several analyses of
calcareous alge. The analyses, made for her by M. Charf, are not very complete, but they
are, nevertheless, of distinct value. The data are as follows:

Lithothamnium calcareum. St.-Vaast-la-Hougue, Manche, France.
. Lithothamnium caleareum. Isle Glenan, Finistere, France.
Lithothamnium fornicatum. Norway.

. Lithophyllum incrustans. Gatteville, Manche, France.

. Lithophyllum incrustans. Mazagan, Morocco.

. Lithophyllum tortuosum. Genoa.

. Lithophyllum craspedium. Tahiti.

ISD or WOW E

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 carbon-
ates alone.
Lemoine’s analyses of alge.

| |
At 2 3 4 5 6 | 7
Poluble mH Ol es. sen cee ee eey ete ne eee ee cee 2.3 3.0 1.2 ai 3.8 5.7 | 3.2
Insolublejin tH Ola 3422/55 427 eee eee esis Acee wee eoen 0 a) 0 | 0 0 CZ 0
(GE OL0) oc cede cu euanoquescs cou. ase a sae asacnes ace 80.9 | 75.2 81.6 84. 4 ewe 73.4 75. 4
MeO Ogee eee SEE RRS earn ete aterctets hoy pts ean 11.8 10.7 9.3} 10.8 9.8 9.3 | 15.9
Organieimatiiere ae. c> eest eet eee soe eee 5.0 1.1 7.9 3.1 8.7 5.4 5.5
100.0 100.0 | 100.0 100.0 100.0} 100.0} 100.0
Mg C@;.corrected sccm -eac-cn seis nee eee e eee ee MD TE |) PR) LON2i jes Les 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 alge from warm regions, as compared
with those from cold waters, was 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 alge, including three by J. Chalon, as
follows:

1. Lithothamnium caleareum. Roscoff, Finistere, France.

2. Lithophyllum incrustans. Banyuls, France, on the Mediterranean.
3. Lithophyllum tortuosum. Naples.

#7 Nichols, H. W., Field Columbian Mus. Pub. 111, p. 31, 1906.

0 Skeats, E. W., The atoll of Funafuti, pp. 376, 377, London, The Royal Society, 1904.
51 Judd, J. W., idem.

‘Lemoine, P., Inst. océanographique Monaco Annales, vol. 2, fase. 2, 1911.

GENERAL DISCUSSION. 55

Chalon’s analyses of algz.

at de lias S|

CRO Ore sano oe ltciajesis(a an—eins.4 05.5 82.41 76. 06 82. 20 |
WHO) eemncecner cc ccce sacse ces a 11. 80 14. 38 11. 57
Organic matter...........--... 4. 30 7.38 5. 26
Woh et ee eee . 86 ON . 80
99.37 | 99.54 | 99.83

Mo@O; corrected=.....-.-.....-< | 12.52 | 15. 90 | 12. 35 |

i

notwithstanding Hégbom’s divergent data, strengthen the suggestion that the proportion of
magnesia in the alge is influenced by temperature.

The material studied by us was carefully chosen by Dr. Howe, with direct reference to the
purpose of our investigation. Every species was thoroughly identified, its locality was definitely
stated, and the specimens were remarkably clean and free from misleading impurities. The
results obtained by us are therefore as nearly trustworthy as it is practicable for us to make
them. The significance of the algzw in reference to dolomite was already well established by
previous workers, but our new data strengthen the conclusions which our predecessors had

drawn.®*
GENERAL DISCUSSION.

In the foregoing pages we have reported 322 new analyses of marine invertebrates and have
cited many other analyses made elsewhere. These data shed much light upon the chemistry
of the marine sediments, 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 intro-
duction to this memoir and are taken for granted in the following general discussion of the
results that we have obtained. First in order we may consider the distribution of the essential
constituents of the invertebrate skeletons, taking each one separately.

Silica.—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 Huplectella, 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
itis animpurity. 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 important base in nearly all marine shells or skeletons, whether verte-
brate or invertebrate, is lime. Only the siliceous organisms are free from it. Molluscan
shells, 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 marine invertebrates, with few exceptions, it is the dominant inorganic con-
stituent. 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 algx it is especially important, and some other organ-
isms contain it in notable proportions. Its peculiar relations to temperature have been noted
in several sections of this work and will be discussed more fully later.

83 On the importance of alge as reef builders, see an interesting paper by Dr. Howe in Science, new ser., vol. 35, p. 837, 1912. He cites much
other literature.

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 cent was found. Sea salts contain 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 pro-
portion of magnesium carbonate in our important magnesian series ranges from 5 to 25 per
cent. The small quantities of magnesia found in 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 crusta-
ceans, 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. Mag-
nesium phosphates, however, 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 in the sediments as a phosphate. In any case the dead
organisms are likely to be buried among calcareous sediments, where calcium phosphate should
be formed. Even the phosphatic worm tubes, in which the calcium is insufficient to form a
tribasic salt, would probably follow 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 contributory to the sedi-
ments. 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, 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 analogous to or identical
with apatite. Its presence in vertebrate bones is well known. Boron also is widely distrib-
uted in the animal kingdom. G. Bertrand and H. Agulhon * detected it in crustaceans,
mollusks, and echinoderms, as well as in vertebrate animals. Traces of barium have been
detected in various organisms, and in the soft part of certain rhizopods granules of barium
sulphate have been found. Strontium is reported by O. Vogel * in corals and molluscan
shells, and according to O. Biitschli® the skeleton of a radiolarian, Podecanelius, consists
almost entirely of strontium sulphate. Iron and manganese are of common if not of general
occurrence in marine organisms, and copper, lead, zine, cobalt, and nickel have also been
found.* The presence of copper in oysters has long been known. Silver has been detected

5 Carles, P., Compt. Rend., vol. 144, pp. 437, 1240, 1907.

56 Bertrand, G., and Agulhon, H., idem, vol. 156, p. 732, 1913.

56 Cited by Samoilov in Mineralog. Mag., June, 1917.

57 Vogel, O., Zeitschr. anorg. Chem., vol. 5, p. 55, 1894.

8 Bitschli, O., Deutsche Siidpolar Exped., vol. 9, p. 237, 1908.

89 See Forchhammer, G., Philos. Trans., vol. 155, p. 203, 1865.

On manganese see Cotte, J., Soe. biologie Compt. rend., vol. 55, p. 139, 1903; Bradley, H. C., Jour. Biol. Chem‘, vol. 3, p. 151, 1907, and vol. 8,
p. 237, 1910; Boycott, A. E., Naturalist, 1917, p. 69, and Phillips, A. H., Carnegie Inst. Washington Pub. 151, p. 89, 1917. Phillips also found iron,
copper, and zinc in the soft parts of invertebrates, and, rarely, lead. See also Mendel, L. B., and Bradley, H. C., Am. Jour. Physiology, vol. 14,
p. 313, 1905. For copper, see Rose, W. C., and Bodansky, M., Jour. Biol. Chem., vol. 44, p. 99, 1920.

GENERAL DISCUSSION. 57

in oyster shells by A. Liversidge,®’ and vanadium has been reported in the blood of an ascidian
by M. Henze," and in a holothurian by A. H. Phillips. 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 from others are of great significance. The limestone immediately below the zone of
living forms owes its composition to all the organisms that flourished on the reef. Alger, corals,
aleyonarians; Foraminifera, and other forms of less importance contribute their remains to the
building of the limestone, which may vary in composition as the life upon it varies. Corals
may predominate in one place, alge in another. Each reef must therefore be studied on its
individual 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® as follows:
1, Lithothamnion;™* 2, Halimeda; 3, Foraminifera; 4, the corals, including Heliopora and other
aleyonarians and the millepores. Here the corals are subordinate to the alge, and even the
Foraminifera outrank them. To call the Funafuti rock a coralline limestone would therefore
be somewhat misleading.

At other localities the relative abundance of the marine organisms is different from that at
Funafuti. A careful analysis of samples from reefs at Murray Island, Australia, conducted by
T. Wayland Vaughan,” gave the following results: ‘‘1,600 feet from shore, madreporarian
corals, 41.9 per cent; calcareous alge, 32.6 per cent; Foraminifera, 12.4 per cent; Mollusca, 10.2
percent. At 200 feet from the shore the order is: Calcareous alge, 42.5 per cent; madreporarian
corals, 34.6 per cent; Mollusca, 34.6 per cent; Foraminifera, 4.1 percent.” Around the Tortugas,
according to L. R. Cary, the aleyonarian fauna is the most important contributor to the forma-
tion of reef limestones. He estimates the quantity of aleyonarian spicules at this locality to
average 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 aleyonarian remains were lacking.

Chemical analysis, however, is not the only factor of importance in determining the com-
position of a marine limestone. The crystalline character of the shells and skeletons, whether
calcitic or aragonitic, must also be considered. For this purpose the well-known reaction
with cobalt nitrate, the “Meigen reaction,” is commonly employed, especially by W. Meigen
himself, who has studied a considerable number of organisms, both recent and fossil, and some
of his determinations ” relate to genera examined by us. For these, excluding fossil forms,
the data are as follows:

Calcite. Aragonite.
Lithothamnium. Alga. Halimeda. Alga.
Lithophyllum. Alga. Galaxaura. Alga.

Polytrema. Foraminifer. Millepora. Hydromedusa.
Corallium, Alcyonarian, Distichopora. Hydromedusa.
Tubipora. Alcyonarian. Heliopora, Alcyonarian.
Serpula. Annelid. Spirula. Cephalopod.
Terebratula. Brachiopod. Sepia. Cephalopod.

Argonauta. Cephalopod.
Balanus. Crustacean.

® Liversidge, A., Jour. Chem. Soe., vol. 71, p. 298, 1897.

6 Henze, M., Zeitschr. physiol. Chem., vol. 72, p. 401, 1911, and vol. 86, p. 340, 1913. Henze also found copper in the liver of cephalopods (idem,
vol. 33, p. 417, 1901.)

62 Phillips, A. H., Am. Jour. Sci., 4th ser., vol. 46, p. 473, 1918.

63 The atoll of Funafuti, pp- 125-150, L onGtay The Royal Society, 1904.

64 The term Lithothamnion as used in the Funafuti report is general and includes not only Tithothamntum but also Lithophyllum, Goniolithon,
and perhaps other genera. See The atoll of Funafuti, p. 332, London, The Royal Society, 1904.

6 Vaughan, T. W., Geol. Soc. America Bull., vol. 28, p. 942, 1917.

6 Cary, L. R., Garrenis Inst. Washington Pub. 213, 1918, p. 356.

67 Meigen, W., Naturf. Gesell. Freiburg Ber., vol. 13, 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 probable, in the light of existing
evidence, that all aragonitic organisms are essentially nonmagnesian; and that those char-
acterized by the presence of much magnesia are calcitic. Many calcitic forms, however, are
practically 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 differences 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 dolo-
mite 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 produce dolomite the original limestone must either be enriched by magnesia
derived from sea water or else concentrated by leaching 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 lime-
stone all the way. The rock throughout contained 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."* The fact that many sections of the core are described as ‘“ cav-
ernous”’ 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 depth 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 compo-
sition. On the other hand, the core at 40 feet carried only 5.85 per cent of magnesium carbon-
ate and was in great part composed of Heliopora and Millepora, both originally nonmagnesian.
The core sections evidently 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

® 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.” 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 cent 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 zones. The surface layer is about 25 feet thick, and its composition is
directly determined by the organisms living above it. In this zone the evidence of concen-
tration by leaching is quite clear. From 25 feet down to 640 feet the rock is essentially a
limestone, with very little magnesia. The lowest zone, from 640 feet downward, is dolomite,
and 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 not 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 more abundant than they are now. This supposition, however, does not account for
the sudden change from 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 sug-
gestions, 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 alge, could thrive. <A prolonged rest, a period
of equilibrium, then followed, during which very little rock 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 dolomitice 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 ex-
tinct, for their remains appear throughout the Funafuti cores, but they were much less abun-
dant than at first. Whether this supposition is true or not might be determined by a quanti-
tative 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 as they indicate the frequency
of occurrence of the different organisms. They do not show their relative quantities.” 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” 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 from the core 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,

® See the petrographic report by C. G. Cullis in The atoll of Funafuti, pp. 392-420.
7 The atoll of Funafuti, tables on pp. 336-361.
7 David, T. 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 zone at Funafuti represents 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.”

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,”* who made good series of chemical analyses of them and also
discussed their origin. Organic and especially algal remains were common in the rocks and
were clearly recognizable in spite of the fact that in many 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,’ who found them to be composed in great part of algal and forami-
niferal remains. Among 47 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 alge as limestone 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 more than one way.” 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 appro-
priately 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 been 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,’* who sought to explain the dolomitization
of a limestone on the island of Makatea.

That an enrichment in magnesia from sea water is possible is shown by some experiments
made by C. Klement,” who found that a concentrated solution of magnesium sulphate and
sodium chloride, at 90° C., attacked aragonite and corals strongly, yielding a mixture of car-
bonates containing as high as 41.9 per cent of magnesium carbonate. 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 produc-
tion 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 conditions, with less concentration and at lower tem-
peratures, 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

7” Vaughan, T. W., Papers from the Tortugas laboratory of the Carnegie Institution of Washington, vol. 5, p. 66, 1914.

73 Skeats, E. W., Harvard Coll. Mus. Comp. Zoology Bull., vol. 42, p. 51, 1903.

7 Sherlock, R. L., idem., vol. 38, p. 349, 1903.

7 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.

76 Dana, J. D., Corals and coral islands, 3d. ed., p. 393, 1890.

7 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 par-
ticles, there would be a degree of molecular mobility which might bring about combination.
This 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 Général
E. Jourdy, who especially recognizes the importance of the alge, of aragonite, and of time.
Much remains to be done, however, before the problem of dolomitization can be completely
solved.’

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 sedi-
ments is uncertain. At best they can at first form beds of only moderately phosphatic lime-
stone, 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,” 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 groups of organisms the
proportion of magnesium carbonate is dependent upon or determined by temperature. The
crinoids and aleyonarians show this relation very clearly, and it is also suggested by our analyses
of foraminifera, crustaceans, and alge. 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. This rule, or rather tendency, we are inclined to believe is general, although we
must admit that there are probably exceptions to it. Whether the increase in magnesia in
passing from 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 differ-
ences 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, differences that are great enough to exert some influence upon the vital
processes of marine animals. Although the ratio between calclum and magnesium is prac-
tically constant in all oceanic waters, a concentrated water would contain more magnesium,
volume for volume, than a water that was more dilute. Whether an organism living in a con-
centrated 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 statement of established fact. That warmth
favors the assimilation of magnesia by marine invertebrates seems to be reasonably certain,
but why it should 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
attempts 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 of magnesian limestones
than cold regions, but the evidence is by no means conclusive. A dense population of cold-
water organisms might add more magnesia to the sediments than a sparse population of warm-
water 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 aleyonarians are much richer
individually in magnesia, but they are not so large. One Paragorgia would therefore count

78 Jourdy, E., Soc. géol. France Bull., 4th ser., vol. 14, p. 279, 1914. :
79 Clarke, F. W., The data of geochemistry: U.S. Geol. Survey Bull. 695, pp. 515-526, 1920; gives a summary of our knowledge of phosphate rock.

62 THE INORGANIC CONSTITUENTS OF MARINE INVERTEBRATES.

for more than many smaller alcyonarians in the formation of magnesian limestone. If, how-
ever, the warm-water organisms are as abundant as the cold-water forms, and if their aggre-
gate mass is as great, then the tropical limestones of marine origin should be more richly magne-
sian 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 alge, 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 ® 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.

80 Am. Philos. Soc. Proc., vol. 60, p. 62, 1921.

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