I N V E R T EB R A T E

PA L A E O N T O L O G Y
AN I NT R O D U C T I O N T O T H E S T UD Y
OF F O SS I L S

H ER B E R T L EA D E R H A WKI N S
M sc
. .
, F G S. . .

LECT URER IN G EOL OGY, UN I VERS I TY COLL EGE , RE A DI N G

W ITH S I X TE E N I LL US TRATI ONS

M ET H UEN CO . LTD .

36 ESSEX STRE ET WC
. .

L ONDON
 PR E F A C E

H E fo ll o wi ng p a ge s are d ed ica t ed to the se rvice

o f tho s e Ad m irer s of F o s s il s who have n ot yet

e n tered i nto a s t rict exami natio n of the d is

ti nct ive character s of thes e i n tere s ti n g s ub s ta n ce s
”
.

Thu s alm o s t a cen tury a go J ame s Parki ns o n ope n ed

, ,

S tudy of F o s s i l
“
hi s Ou tli nes of Oryctology The
.

”
O r ga n ic R em ai ns h a s adva n ced s o far that the
mo n ograph s o f its pio neer s retai n l ittle m ore tha n
bibliographic a nd hi s torical val ue s The apo s tolic
.

s ucce s s io n of Palaeo ntologi s t s is one of progre ss ;

while the g reat n am es of old are rightly held i n
reveren ce the opi nio ns a nd philo s ophie s of early
,

worker s have n o better claim to authority tha n the

U p s tart here s ie s of th e curre n t ge n eratio n The .

truth s t h at th ey d i s covered m u st e n dure for all tim e

\

as imperi s habl e fou n datio ns for the S cie n ce ; their

theorie s mu s t be tried i n t h e fi re k i n dled by their
re search I f the tes t e n d s i n de s t ructio n of ol d beli efs
.
,

let it b e remembered that theo ry a nd hypothe s i s are

the fu e l witho ut which the fla me of S cie n ce would
“

s oo n grow dim ; a nd from th e fur n ace flow s a n ever

gr owi n g s trea m of pure k n owledge .

N everthe le ss the s e n te n ce quoted expre s s es the

,

purpos e a nd ded icatio n of thi s book F o ss il s are s till

.

objec t s of a dmiratio n ( in bo t h s ense s of the word ) ,

5
 IN VE R TE B R AT E PALAE ON T OL OGY

whether their beau ty captivates t he eye or their

complex ity bewilders the m i nd The brief chapter s .

that follow are de s ig n ed to e n courage th os e who s e

i n tere s t h a s bee n awake n ed by the form er s e nsatio n ,

a n d to s mooth the path for tho s e who require s om e

p alaeo n tological k n owle d ge for furthera n ce of other

s tudie s . The variou s a s pect s of the S cie nce are of
n ece ss ity touched o n but l ightly The grou n d -work
.

of s cie n tifi c i n quiry is n o m ore i n tere s ti n g a n d al m o s t ,

le s s i n s piri n g tha n the lear n i n g of the alph a bet A n

, .

average textb oo k m u s t al way s bear to the S cie n ce of

which it treats s omethi n g of the relatio n of a D ictio n a ry
to Lite rature ;it i s i n d i s pe n s able but l ac k s cohere nce
,

a nd co n ti n uity A m o n ograph that thrill s the s oul of

.

a s peciali s t leave s others col d if n ot i n d iffere n t At

,
.

“
the oppo s ite extrem e come t he s o cal l e d - Popul ar
treati s e s o n s cie n ti fi c topic s T oo ofte n accu racy i s
.

therei n s ubordi nated to s impl icity— who could i n te rpret

Mil to n i n w ord s of o ne s yl lable ?
A n attempt h as b ee n m ade he r e t o s teer a m iddle
cour s e The fi rs t part of t he book apart from s u nd ry
.
,

tech n ical detail s is i n te n ded to s how s omethi n g of

,

“ ”
what i s mea n t by P alaeo n tology I t i s a com m on
.

experie n ce to e n cou n ter pe ople who are horror -struck

whe n tol d that a n acquai n ta n ce is a Pal aeo n tologi s t .

Whe n it i s ex plai ned that he goes i n for fo s s il s

"
“ ”
,

they are s a t is fi e d t he word i s fam iliar a n d they

—
,
“
a s s um e t hat they k now al l about the s ubject But .

t ra n s latio n of Greek i n to Lati n add s n othi n g to u n der

s ta n d i n g o f the matt er co n cer n ed Palaeo n tol ogy ca n be
.

c alled O ryctology the s tudy of F o ss il s or An tediluvia n

, ,
”
Biology but the S cien ce rem ai ns th e s ame
, I t is .

hoped th a t the fi rs t s ect i o n of the book m ay s e rve to

 P REFACE

e nl igh t e n t hough with a fai n t gleam tho se wh o d es ire

, ,

s ome expla n atio n of the e n thu s i a s m wi th which P a la e

o n tologi s ts purs ue their re search .

The s eco n d pa rt i s des ign ed to give a n epitome of

the s eque n ce of evolutio n a s it h a s bee n un folded i n
geo logical time I n t he s mall comp as s of thi s I ntr o
.

ductio n to the S cie n ce the re h a s bee n n o s pace for the

biological diagn o se s th a t are n eces s ary to make s uch
a n accou n t i n telligible I n co n s eque n ce it h a s s eem ed
.
,

advi s able to fol low i n the m ai n th e cla s s ificati on

, ,

adopted i n the late s t a nd m o s t comprehe n s ive textbook

of Palaeo n tology available in th e Engl i s h l a n guage .

E x cept i n the ca s e O f E c hi n oderm ata ( which are d i s

’
cu s s ed o n the ba s i s s upplied in vol iii of L a nk e s te r s
. .

Tr ea tis e on Zoology) the variou s phyla are a rr a n ged

,
'

i n a cco r d with th e s cheme give n i n t he seco n d editio n

of Zittel -Ea s tm a n s Tex tbook of P a laeon tology vol i
’

,
. .

The s ectio n is in e ffect a ki n d o f co nde n sed guide to

, ,

that rem ark able work Although t he pre s ent autho r

.

ha s had occas io n rigorou s ly to s uppre s s h i s o wn

O pi nio n s i n thu s co n formi n g to the view s of other s ,

the s a cri fice i n volve d s eem s worth wh ile I n a m ul ti .

tude of cou ns el lor s there m ay be s afe ty bu t there is ,

s ure to be co n fu s io n F or a begi nn er a ny one s chem e

.
,

i s better tha n two .

A via med ia is ofte n recom me n ded a s a s afe cour s e ,

but i n t ru th s uch a path i s full of pitfall s a nd bes et with

thor ns S cyl la a nd Charybd i s a n d th e m ore fam iliar
.
,

d a n gers of t he D evil a nd the D eep S e a m ake the ,

attempt of s uch a pa s s age al mo s t foolhardy T o s uch .

fellow -workers as m ay compl ai n that their favourite

a s pect s of the S cie n ce are ign ored or s lighted I will ,

apologi z e whe n t hey h ave s uccee d ed w her e I h ave

viii IN VE R TE B RATE PALAEON TOLOGY

failed To begi nn er s who fi n d the bo ok too ab s tru s e

.

o r t oo d ul l I apologize herewith a n d hope that s om e ,

grai n of i n tere s t or i nst ructio n m ay have s urvived their

fi a i ls .

The ill u s t ra t io ns both photograph s a nd d rawi ngs

, ,

have bee n m a d e by my s elf Al l of the former a nd . ,

a l arge maj ority of the latter a re whol ly n e w— they ,

repre s e nt s pecime ns i n the col lectio n s of thi s D epart

m ent I n t he few c as e s where fi gu re s have bee n
.

adapted from d rawi ngs previou s ly publi s hed ack n ow ,

“ ”
ledgm e n t i s ma d e ;a nd s i nce s uch cop i es u s ually ,
.

s ho w s om e m odifi catio n ( apart from u n i n te n tio nal

clu m s i ne s s of draft s m a ns hip ) from their origi nal s ,

re s po n si bility for their accuracy m u s t fal l on m e .

To M r D M S Wat s o n I de s ire to expre ss my

. . . .

ex t rem e i n deb t ed n e s s for helpful critici s m of the m a nu

s cript a n d to D rs F A Bather a n d H Woodward my
, . . . .

a ppreciatio n of their ki n d ly i n tere s t a n d e n courageme n t

i n thi s work a s i n al l that I have do n e

,
To the s e .

frie n d s m uch of what i s good i n the book m u s t be

as cribed whe t her d ir e
,
ctly or i n dir ectly ;for i ts m a ny
s hortcom i n g s t hey ca n i n no way be held re s po n s ible .

H . L . H .

U NV I E R S ITY CO LL E GE , R EA D I N G,

[a n a a ry 1 9 20
 CONTENTS

PR E FACE

PA R T I .
— MATE R I ALS M ET H OD S A N D
,

P R I N C I PL E S

C H APTER I
“

T H E S CI E N C E O F PA L AE O N TO L O GY

CH A PTER I I
M AT E RIALS
(A) T H E N A T U R E OF F OSS I LS
D efi nition of rms F os sil an d Petrifaction
th e T e
”

Con d itions N e ces sary for F os s ilization

( )
a B i olog cal
i

( ) G e ologi cal
.

5 .

I mp e rfection of Palaeontologi cal E vidence

(B) RR E N CE O F F OSS I L S
T H E O CCU

I F oss iliferous a nd U nfoss iliferous R ock s

F os sils a s R ock F ormers
.

II .
-

III . F oss il B an d s
-

IV . N od ule s
V . D erived Fo s s ils
X I N VE R TE B R ATE PALAEON T OL OGY

( C) T H E P ESE R R VATI ON F OSS I LS

GE
OF
A
P

Complete or U nchanged Pre servation

II . Ca sts an d M oulds
III . Pe trifaction

(a ) B y I nfilling

( o) B y M ole cular R a ment

e pl ce

IV . D i stortion an d D i s fi gure me nt
I mp ression s an d T rack s

C H APTER III

T E CH N I QU E

(A ) CO LLE CT I N G
Purpos e in Vi ew

II . R ard
eg for S tratigr aph ical Pos ition
III . L arge a nd M inute Fo s s i ls

IV . F i eld Preparation
F ield La b elling nd Pack ing
a

( B) P R E PA RATI ON
N ee d for Patience
II . S e parati on Fo
of s s ils from M atrix
I II . D is s ection of F o ss ils

IV . S ta i ning
S t ora ge an d Pr es ervati on

( )
C T A XO N OM Y

C H APTER I V
G E O L O G I CA L PA L A E O N T O L O GY
(A ) ST R A TI GR A P H I CA L
Z on es and H eme ra e
Characteri stic F os s ils
 C ON TE N TS xi

(B) PH YS I O G R A P H I CA L
T e rre stri al F auna s

II . F lu iatil nd L a u tri n Fau a

v e a c s e n s

III . M ari n F auna

e s

IV . F il a E iden f Climat
os s s s v ce o e

C H APT ER V
B IOLO G I CA L PA L A E O N TO L O GY
Ph yloge n y a nd M orp h oge n y
Post -L arval O ntoge ny a n d M orph ogeny
S p e cializati on a nd R gr i e e s s on

A cce leration an d R tardati

e on

Th e L ife -Curv e

PA R T I I .
— H I S T OR I CAL B I O LO GY
C H APTER I
PR E -CA M B R IA N F A U N A S
C H APTER II

L O WE R PA L A E O Z O I C FAU NAS
G
W
P ral A ccounten e

Protoz oa
O Por ife ra
C Coele nte rata
E ch in od ermata
i
r
r

a
r Polyz oa
o Brach i op oda .

m M ollusca
a A rth rop od a

C H A P TE R I II

U PPE R PA L A E O Z O I C FA U NA S
; G
ral A ccount e ne

w Protozoa
o Porife ra
 xii I NVE R T E B R AT E PAL A E ON T OLO GY

U Coelenterata .

E ch inod e rmata
’

W Polyzoa
Q B rach iop od a
E M ollusca
E A rth ropod a

C H A PTER I V
M E S OZ O I C FAU NA S
G ral Account
e ne

Protoz oa
Porifera
Coele nte rata
E ch in od e rmata
Polyz oa
B rach iop od a
M ollu s ca
0 A rth rop od a

C H APTER V
CA I N OZ O IC FAU N A S
G eneral A ccount
Protozoa
Porife ra
Coelente rata
E ch in od e rmata
Polyz oa
B ra ch i op od a
M ollus ca
A rth ropo d a

C H APTER V I
CR I S E S I N E V O L U T I O N

I ND E x
 LIST OF PL A T E S
/

L
P A TE I

L
F I G.

k Car boniferous i mestone contai ning L z t/I ostr ah on

' '

I . Bloc of

i r r egu la re Port S t M a ry, I sle of M an

. . .

2 . Bloc k of Ca rbonif rous e Li mestone co mposed of colu mnals of

t
Cas leton, De rbyshir e .

B loc k of Ba th onia n Lim est one mposed of M z llen cr i rmr

‘ '

L
3 . co .

a n sd owne , nea r Bath .

L P ATE I I

I . Bloc k of Ludl ow( A ymes y i es tone tr ) L m composed of Cone/ti di u m

b u gI
/ i i . Weo Ed e , S hropsh i e g r .

2. F
N od ule of li n t fr om th e pp e r C h a l , cont ai ni ng , U k an d r
d e i ved
from, t r i eu lz tes Pep par d , Ox on
’

. .

3 Bloc k of Be m bri d g e L im e s one t ( O lig ocene ) la rge ly compose d Of

W
.
,

L z mnaea and Pla nor oz s B embri d g e , I s le of ig ht

’ '

. .

L
P AT E I I I

Tr zgom a wlla ta with th e liga men t preserved Corallian,

’

I cla

W ym
. .

e ou th .

2. M
’

a i a z nca r va . I nte rnal mould Port l ndian S wi nd on

p a ,

Di stort ed b y clea va g e Trema d oc sla tes

' ’

3 A ngeli na sed n eéz .

Wl
. . ,

Mi nfi ord , N
'

. a es .

is c u r mbling .

5 Ostr ea ed u lz s
.
'

. Plioce ne , S uffol k . Th e t
ca lci e sh ell i s in pe fec r t
x iv I N VE R T E B R ATE PALA E ON T OLOG Y

F AC N I G AG E
P

PLA TE IV
‘

F IG .

I . Or bi cu lozd ea
'

Coa l M easures, Coalbroo d ale

n i t zd a .
'

T he k .

g lossy, lar gely chitinous, shell is pe rfectly preserved in an

irons tone nod ule .

2 . Fr gm
a en t of Pelecypod Sh ell rom
f th e U pp r Gr e eensan d , nea r
S id mou h , D evon, t rep laced by cha lced ony i n th e fo rm of

D acty lzocer ar
’

mmu ne pper
co ia s, hi tby, Y ork s rom a
. U L W . F
n od ule of ce me nt -ston e Comp lete a nd uncompres sed .

Per onoeer ar a n n a la t u m (a clos e ly mi lar species)

si pper U
Li R F
.

a s, un swi c kB ay, Yo rk s . rom the sha le, flattened and

t elescop ed .

Clzon a
'

sp . Fli
t mould s of the cryp ts perfor ti ng the sh ell of
n a

I nocer mm U pp r Chalk S ussex

a . e , .

A t yp
f et u l r z r
a r Lud low Limestone West Kn wle Cr ven
’

zc a
’

.
,
o , a

A rms S lop , I ntern l mould ( ventra l si d e ) s h owi ng

a . a ,

mus cular a nd o th er i mpressions .

PL A T E v 0- :
4 4

I . Lioeer a: p cf. o a lz n u
'

m . B aj ocia n, B u rton Brad st ock , Do rse t .

I nterna l mould sh owing tr

su u es .

2 . A mmoni te wholly r t
e nc u s e d b y Py i te r and M arcasite . Oxford
Cla y, n ea r Oxford .

L
P ATE V I

1. r
Pa t of a fossi l- band in U pp r L e ud low Fl g
a s, b
O ni ur y, S alo p .

rg t
T he i h -hand si d e was n ea r
th e qua rr
y
-fa ce a n d solut i on

F
,

has red uce d th e organisms to ca st s a nd mould s . ar th e r in

( to th e left of th e figure ) th e p re se v ed
sh ells a r e r in tact .

atella a n d Ca ma r otoeeéz a
” ' ’

Or tfi z s
'

lu na t a , C/zonetes stn
’

sp .

ar e r
th e resent ch i ef fo ms p .

2. Pa rt of a fo si l b nd i n C a d ocia n ( A cton S cott b ed s )

s - a ar ,

M arshbrook S lop Complete ly d eca lcifi ed

"
O t / us
'

, a . . r
 L I ST OF PLATES xv

FA I N GC PA G E

PLAT E V I I 1 04- 1 0 5

F IG .

I . D ia gram of contiguous h l es of a n inter mbula crum a nd a n a v a

a mbulacrum f Plex fi n us om a tu s showi ng loc lized

’ ‘
“
o z ee z ,
a

st a g es in d e velo me nt rm ”
p T he fi s t fo e d p es ( t th e
la t . r - a

bottom f the fi gure ) nd the las t formed pl tes ( t th e top )

o , a ~
a a

y uthful ch ra cters of orn ment

“
S h ow
”
o Th e full spe ci es a a .

ch racter is sh own a t th e mi d z one

a - .

M orp hogeny of s ome type s of a pical systems in th e Echin oi d ea .

F ig 2 D ia d e min e (e g Fi g 3 Ca lyci ne
,
. . . . .

F ig .
4 . F ig .
5 .
H garter .

Fig 6 H oleetyp us ( th e pos teri or

’

Fi g D z seozd ea
’

. . . .
7 .

ge nit al is a lwa ys rfor a ted ) F ig 8 Clypeas ter

L
n ot
p e . . . .

F ig 9 A nofl/z opygu n
. .
‘
Fig . 10 . Sp a l a ngu r ( a fte oven ) r .

F ig I I A d ul Spa ta ngu s
. . t ( a fter L ovén ) . F ig . 12 . Con ulu s .

F ig ms r expla nation
’

. 13. E ck z noc . F or furth e , see p .

94 .

L
P ATE VI I I 1 04— 1 0 5

PH YLOGERONT IC CH A RA CT E RS
mm z:
red uced ) B th ni n Burton ( much
'

Pa r k znson ia
'

M d ofi . a o a ,

Br d stock D orset
a T he i nner ( e rly) wh orls
, e str ng ly . a ar o

ribbe d Geronti c laevig ti on obliterates ornament in th e

. a

outer whorls .

2 Cl zzella longawa Eocene , Ba r ton H an ts Th e protoconch

ais followed
. . .
,

b y one who l sh o r wing F usi d cross orn a men t .

Th e t wo n ext wh o ls r a re s mooth ed b y gerontic la e viga ti on ,

th e rest bec o me ri ly spin us
secon d a o by exces sive calcifi ca

ti on .
( Base d on Grab a u s figure )
’
.

3 . C1a oella sola nd er z .

’

r ts A typ e simil r
E ocene , B a ton , H a n . a to
C longaeva ,
. b ut f i li ng to progress beyond la évi ga ti on
a .

4 . P/n llzjos z a gemomlzf er a

’ '
‘
'

( a ft er W ood war d ) . Car boni ferous i me L

s tone S howi n . g geronti c f e t res i n a u s i mp li fi ca ti on of the
‘

glab ell multiple pyg id i um a nd p ustular ornament

‘

a, ,
.
xvi I N VE R TE B R ATE PA LAEON T OLOGY

PL AT E IX

LOWE R LAEOZO I C FOSS I LS

PA

D z ay mogr aptm mu r c z z l som Lland ei l A ber id d y

' ' ’
’
o, e

L l nd o ry n r Gir an
. .

M onogr ap tus sodn k

’ ’

e z. a ve , ea v .

S tr omood e: typw Wenlock Limestone M uch Wenlock

L im
. .
,

F avori te: gotb la nd z e o a ( l ng itud in al secti on )

’

Wenlock . e

st one , P esth ope , S a lop r .

t
!
} H alysz ter
’

ea ten u la r z a .
’

W
k Limestone M uch Wenlock
enloc , .

D md r oqwtis ( fter B th er) Carad oci n

i '

C seot zea a a a

Wenlock Limestone M uch Wenl ck

. .

'
:
x Cyatfi ocr z nur sp .
,
o .

C mbri n Tremad oc (P rtly based

o '

p L z ngu lella a a , . a on

D a vid s on s fi
’
g ure ) .

Or ozcu loid ea Upper Lud low, O nibury, S a lop

'

r uga t a .
~
.

Of f/ Ms
”
D a lma ne lla ) tes tud z na r z a Ca ra d oci an , M arsh
' ' '

IO

R
. .

brook , S a lop . est o ed r f rom casts and mould s (see Pl . VI .

F ig .

I I. S tr op/zomm a Ca r ad oci a n, S oud ley, S alop

ex p m
es to ed
aa . . R r
fr om mould of ven al valve ( par ly ased on D avid son s
’
tr t b
fi u e) gr
U pp r L
.

Clzomter rtr z a tella ud low, Oni bury S alop

'

12 . . e , .

P AT E XL 1 36- 1 3 7

LAEOZO I C F OSSI LS LO WE R PA

A t ypr ti u l L ud low Lim tone Weo Ed ge S

a re c a r zs .
'

es a lop

Low r Lud l w Wh ttl t n S l p

, , .

O f/m m igid a
'

f I a r e o e e o a o

Wenl k Lim t n D ud ley

. .
, ,

Umpfi lot oefi us d

'

s a r z r eor . oc es o e, .

T nt Ca d oci n M r hbr k S l p
'

e l le aea z r. ra a a s oo a o

U pp r Lud low Onibury S l p

.
, ,

O tb
r ba ll l m
ocer as a u . e , , a o .

Pa r a bolz nella tr z a r t/z r a

' '

. S hi ne ton S hales , B e lswa rdi ne B rook ,

S alop ( S ligh tly restored )

Ll
. .

Og /gz oca ffs b ur /Ii

‘
'

. a n d e ilo, B uil h t .

mm tn eur Ca rad ocian, O nuy

'

Cryptolz t k us
’ ’

Tr z n ueleu s co

R rW
.

i ve , is ta nstow, S a lop
L
.

Cer a tzoea r z r Pleptoa a cty lu s

'

Lowe
‘
’
r ud low, M artin s Shell, ’

L
.

e int wa d i ne , H e efo d sh i e r r r r .
 L I ST OF PLATE S

L
P AT E X I

U PP ER PA LA EOZO I C FO S SI LS
F IG .

I . Ca lceola sa nd a li n a . M id dle D evoni a n, Gerols tei n , Eife l .

2 . L i t/zos tr ot i on ba r a lt sor me . Ca rboniferous Lim eston e, Poyl

V ai sh , I sle of M an .

3 . Or oitr enz i tes ellipt iow . Ca r boniferous Li m t

es on e , Cli th e roe .

4 . Pla ty or i n ur giga s . Ca rboniferous L imestone , Cli th e roe .

\5
M elon ee/
. z i n a r k ee i n i
p g . Car boniferous L im t
es on e . ( M od i fied
fr om Jac k s on .
)
6 . Pr od uetur semz r et i eu lat a s ma r ti n i
‘

. Car boni ferous L im es ton e ,

Castleton, D erbysh ire .

7 . Pugn ax acu mi na t us . Ca rboniferous Li m t

es one , D oved a le .

8 . D i ela sma b a s ta ta . Ca rbonifer o us Lm i es to ne , Ca st le ton .

9 . Sp i r i/er str i a t u s. Ca rboni ferous L im t es one . ( After D avi d son ) .

L
P AT E X I I 1 6 8-1 69

U PP ER PA LA EOZOI C FOSS I LS
I Pos ia on ongf a beolmi
’
. L wrC M
o e oa l r
easu es , nea r S hefii e ld .

2. Pter i n eop ecten papy raceus L wrC

. o e oa l M eas ures , Sta lybri d ge .

E uomp /za lus pen ta ngn lat us . Car bonif rous Li mestone
e , Cas le ton t .

B eller op k on sp . Carboniferous Lim estone , K e nd al, W es tmorl nd a .

Cy r tocer a s m M id d le D evoni an, Gerolstein

’
noaosu . .

Glyp lu ocer a:
’

cr en i str z a .
’

Ca rboni ferous Li mestone ,

Poyll V ai sh ,
I s le of M an .

7 Plz z llzjos i a
'

z wa ld i
ei e/ mu cr ona ta Ca rboni ferous Lim t
es one .

W
. .

( A fter ood wa d r .
)
8 . B elz n u r u s
’

r egi me . Coal M eas u res .

( Aft er W ood wa rd .
)
S li mon ia mmz nata Old R ed S and stone ( A fter La uri e )
'

9 . a . . .
xviii IN VE R TE B R AT E PALAE ON T OLOG Y

L
P AT E X I I I

M ESOZO I C F OSS I LS
F I G.

[ras tr aea ex p la na t a . Cora llian, S helli ngfo rd , B erks .

Api ocr i n m p a r ki nson i . B rad ford Clay, Bra d ford , Wilt s.

Pa r ad d a r i s flor igemma . Cora lli an, Stanford i n th e V ale .

A Pfiy mosoma éoen zg i p pe Chal , nea K i n scle e , H a n

. U r k r g r ts .

r r t
( Th e econst uc i on of th e a p ica l sys tem i s hypoth e ica l ) t .

M N ucleoli tes scu t a tu s . Cora lli a n, M archam, B er ks .

Q M i cr a ster eor a ngu i n u m . U pp r Ch lk e a , Ecch ins well, H a n ts .

Q Cy clot/zy r i s p li ca t i li s . U pp r Ch lk e a , nea r Newb ury .

O
O Ter eor a tu la p k i llzlt si . B a th onia n, B u ton B r rad stock ,
Do rset .

O Zei ller z a d zgon a

‘

B ad fo d Clay . r r . I nte i o r r of do rsal va lve .

tr
( Af e D a vi d son ) .

Sp i r zf er i na cf. wa lcotti . Lower Li as , T i ms bury R ad st ck

, o .

PL A T E x rv 1 4 1 85
8 —

M ES OZO I C F OS S I LS
Tr igon i a st r i a ta . B ath onia n, B u ton B r r a d s toc k , Do rset .

N Tr zgon i a ela oella ta . Co r a lli a n, M archa m .

»
9 M od i olu s s p . Bathonian ,
B u ton Br r a d stoc k .

P Pleu r ot oma r i a r et i cu la t a . Ki mm rid g e e Clay, W ym e outh .

M
“
N at i ca oj
a ooi en s z r
’

. B a th onia n , B u t on B r r
a d stoc k .

Q A rn a ltlzeur ma rga r i ta t u s . M i d d le L i a s, R un swi c kBy a ,

Yo rks .

S Cor mocer a r j a son . O xfo d Cla y, S t I ves r . .

O
O U pp r Ch lk
S eap fi i tes s p . e a .

O B l m i t ll m
ee n e t a U p p r Ch lk
ucr on a a . e a ,
Ar reton , I sle of Wight .

I O. P l o
a ae coite G ult F lk to
y p ra s s .
, o es ne .
 I N V E RT E B R A T E
PA L A E O N TOLOG Y

PA R T I .
— M A T ER I A L S , M ETH OD S
A ND P R I N C I PL E S
C H A PT E R I

T H E S C I E N CE OF PALAE ONT OL OG Y

a) s co p E

H E s cie nce of Palaeo ntology i s co ncer n ed with

n is ms that have exi s ted duri n g period s of
’
the E a rth s hi s to ry p rior to the s tage u s ually
“ ”
called the pre s e n t day I t is clear t hat there i s no
.

r igid li n e whereby Palaeo ntology ca n be s eparated

“
from Biology ; to day will be the pa st t o mo rrow
-
.

The two s cie nce s are i n tim ately rel ated a nd largely
‘

i n terdepe nde n t j u s t as Geology i s li n ked with Geography

, .

But Palaeo n tol ogy ca nn ot be treated a s a mere bra n ch

of Biol ogy ;not o nly doe s the nature O f it s m aterial s
lead to ad optio n of widely d iffere nt m ethod s but m a ny ,

of it s p rob le ms a re foreig n to t ho s e co n fro nti ng s tude nt s

of form s now livi ng I f a ny l ogical co nn exio n betwee n
.

the two s cie nces i s t o be m ai ntai n ed B iology mu st b e ,

s ubord i n ated a s a partic ular s ectio n al mo s t a n aft erm ath

, ,

of Palaeo n tology S uch degr a datio n would not meet

.

with approval b y Biologi s t s nor would it adequately ,

expre ss the relati o n be twee n t h e two l i n es of i n qui ry ;

s o th at bo t h mu s t retai n their i n depe n de nce .
2 IN VE RTE BRATE . PA LA E ON T OL OGY

A gai n Pal aeo nt ol ogy i s very clo s ely co nn ect ed with

, .

G e ol ogy e s pecially wi t h tha t sectio n k now n a s S t ra t i

,

graphy . The t race s of pa s t life are embed d ed i n

s t ra t ifie d rock s a n d ca n n o t be i n terpre t ed s ucce s s fully
,

un le ss t heir hi s torical s eque n ce i s proved B ut it would

.

be a s rea s onable to regard B iology a s a bra n ch of

Geography ( be cau s e d i s tributio n of exi sti ng life is a n
importa n t factor i n it s com prehe ns io n) a s to co ns ide r
Pal a eo n tol ogy a s ec t io n of geological s cie n ce I n deed .
,

the s t udy of S t ratigraphy though ultima t ely depe n di ng

,

“ ”
on t he law of t he order o f s upe rp o s itio n is prac t ical ly
,

im po s s ible without the aid of fos s il evide nce ;in m a ny

way s it i s a k i n d of applie d Palaeo nt ology .

H e n ce the phr as e with which t hi s chapte r ope ne d i s

ap t ;Palaeo nt ology is a d i s ti nct S cie nce with m aterial s
, ,

method s a nd aim s peculiar t o it I t i s largely depe n de nt

.

on a n d n o le s s co n tributo ry to the s i s ter S c ie nce s of

, ,

B iology a n d G eology . Thes e bra nche s of N a t u ral

H i s to ry have bee n explored m ore fully pa rtly be c a u s e
,

of t he l o n ge r appl icatio n of s t udy to them a nd pa rt ly ,

Si n ce Pal ae ontol ogy could hardly progre s s u n til thei r

pri n ciple s were fairly e s tabli s hed B ut Ci n d erella ha s
.

now reached the s tage of adole s ce n ce if not o f m a t uri ty

, ,

a n d m u s t take her place am o n g the other m ember s of

the fam ily T o pres s the s imile t oo far would be u nj u s t

.

t o t he elder s i s ter s—4 they a re far from u nlovely a nd i n ,

n o way s e n e s ce n t —bu t it may be claimed that i n

Palaeo nt ol ogy t here is to be fou n d th e peculiar fa s ci n a
tio n t hat s u rrou nd s the awake n i ng po ss ibilitie s of youth .

The s cope of Pal aeo n tology is wid e a nd h a s n o ,

d efi nite li m it s Theoretical ly i ts Oppo rtu nitie s would

.

com me nce with the fir s t a ppeara nce O f l ife on the E arth .

A l though rece n t re search h a s revealed traces O f organ

i s ms i n rock -sys tem s far Older tha n m a ny lo ng regar d ed
as
“
Azoic it h a s t hereby r e ndered pai n ful ly evi d ent
,
”
 T H E S C IE N C E OF PALA EO NT OL OG Y 3

the rel ative m odern ity of s uch evide nce O n bo th .

geolog ical a nd biologi ca l grou nd s it i s po ss ible to

,

argue t h a t the record s of l ife may cover but hal f or ,

les s t ha n hal f of the hi s tory of l ife i t s el f I t i s thu s

,
.

with all form s of hi s torical i n qu iry ; k n owledge pa s se s

back i nto u ncertai n ty u n ce rtai n ty i nto lege nd a n d
, ,

l egen d i nto mythology ; while beyo n d is a regio n

wher e i n fere nce a n d fi nal ly i m agi n atio n ar e freed from
, ,

the i n co nve n ie n t co ntrol of fact s or eve n of verifi able

hypothe s es I t i s not the purpo s e of the p res e n t work
.

to roam through still le s s to Cultivate tho se E ly s ian

, ,

field s ; s o that the ulti mate l im it of palaeo ntological

i nve stigatio n m ay be take n at the oldes t k n ow n fo ss il
who s e orga n ic qualitie s are biologi ca lly demo ns trable .

The oppo s ite bou n d ary of the provi nce o f Pa lae o n

t o logy i s eve n le s s defi n ite A t what di s ta nce from the
.

feet of a Palaeo n tologi s t do the dom ai n s of Biology ( in

which t er rito ry he i s a lifelong pris o n er ) come to a n
end ? The carca ss of the l a s t member of a race n ow
exti nc t would not s erve a s a la nd m ark I t is at l eas t
.

poss ible th a t s ome s pecie s o r variety that i s i n exi s te nce

while the s e word s are bei n g writte n ma y have died out
com pletely before they are read The arrival O f M a n
.

as a n i n tellige n t i n habita n t of the E arth a fford s n o

be tt e r cri t erio n if o nly for the rea s o n that the time of

,

hi s a ppea ra n ce i n that capacity i s u n k n own The .

Palaeo ntologi s t m u s t n eed s s tudy the fau n a a nd flora

O f t he curre nt geol ogical period s hari n g with the
,

Biolog is t th e evide n ce thereby a fforded a nd applyi n g ,

it t o the s pecial problem s with which he h a s t o d eal .

I n ord i na ry practice the divi s io n betwe e n B iology a nd

,

Palaeo ntology may be take n at the l i ne where the

Plei s t oce ne is s uc ceeded by the H oloce ne ;but thi s i s
‘

at be s t a n ill -d e fi ned d atum

, .

The in d efi ni te cha racter of i ts bou nd arie s ( a quality

4 I NVE R T E B R AT E PALAEON T OL OGY

s h a re d with a ll othe r bra nche s of S cie nce) c anno t

ob s cure t he m ai n fiel d of s tudy t ha t lie s ope n to
P a laeo nt ology Be t wee n the d im outli ne s of Pre
.

Cambria n fau na s a nd the cleare r but b a ffli ngly com ,

pl ex detail of pre s e nt day l ife there ex t e nd s a l on g

, s -
,

ra n ge of fo ss il remai n s which s upply certai n though ,

fragme nt ary evid e n ce of the cours e of orga n ic evolutio n

, .

Thes e fo s s il s are the m aterial s tha t form the ba si s of

palaeo n tological s cie n ce Before procee d i ng to d is
.

cus s i on of the varied co nditio ns u n der which the

evide n ce m ay occur a n d the method s employed i n i ts
,

i nt erpretatio n it m ay be wel l to co nsider briefly the

,

replie s t hat a Palaeo ntol ogi st could m ake t o the qu es

tio ns so ofte n a s ked of s tude n t s of N atural S cie nce : a
va a i s Cn i oon o ?
’
? a nd

( )
I I A I MS

S i n ce Palaeo nt ol ogy is i n i t s e ss e ntial a s pec t s H istori

, ,

cal Biol ogy i t s aim s m u s t agree i n l arge m ea s ure with

,

tho s e of Zoology a n d Bota ny M a ny of the co nditio ns

.

a n d m aterial s avail able to the latter S c ie n ce s are lack i n g

i n the evide n ce with which P alaeo ntology deal s s o that ,

i n;s om e way s i t s s cope i s cu rta iled ;but the i ntroductio n

of t he hi s torical eleme n t give s com pe n s ati n g exte ns io n
i n other d irectio ns The fu n dam e n tal a n d eleme nta ry
.

aim s m ay be s u m marize d a s the d i s covery de s criptio n ,

a nd cl a s s ificatio n of or ga n ic bei n g s a n d s ub sequent

,

ded uctio n O f the law s whi ch have determ i ned a nd s till ,

gover n their life a n d rel atio n s P al aeo n tology the .

,

d is cuss i on of pa s t orga n i s m s m u s t alway s be preceded

,

by Palaeo ntography thei r d escr ip ti on Thi s i s m an i

, .

fe s t ly im po ss ible u n til they ha ve bee n d i s covere d ; s o

tha t the fi rs t a nd perpetual duty Of a Palaeo n tol ogi s t
i s to collect fo s s il s The he althy out d oor s por t ( for
.
,
 T H E S C I E N CE OF PALA E ON T OL OGY 5

s uch i t m ay a nd s hould be) of fo i l hu nti ng n ot o nly ss -

provide s grateful relaxatio n from the relati vely s ede n

tary occupatio ns of de s criptio n a n d s tudy but e n ables ,

the worker t o purs ue the l atter with e nha n ced i n tere s t

a nd u n d e rs ta n di ng N o on e ca n b e qu al ified to d is
.

cours e upo n palaeo ntological topic s who h a s not

per s o n ally exper ie n ced the thrill s a nd di s appoi nt me n t s
a s s ociated with work in the field .

Whe n the three prelim i n arie s have bee n at tai ned the ,

aim s peculiar to Palaeon tology rem ai n The hi s torical .

s eque nce i n which l ivi n g form s h ave ari s e n a nd d i s

appeared i s s till a m atter i n which dire ct ob s ervati o n

play s a domi na n t part although it i s of s uch a n ature
, ,

a nd o fte n s o imperfe ctly d i s pl ayed that s ome O p p or

,

t unit ie s ari s e fo r the exerci s e of argume n t T he d i s .

t r ib ut ion o f form s of life at the pre s e n t day ( which s till

a fford s m uch s cope for biological re s earch) i s no le ss
i mporta nt though le ss appare n t whe n pa s t period s are
, ,

i n review ; m a ny d ata of great geological i mpo rta n ce

ca n b e determi ned by s uch phe n ome n a .

F i nally whe n the s ucce s s io n of orga n i s m s h as bee n

,

s u fficie n tly e s tabli s hed the ultim ate phil o s ophic aim of
,

the s cie nce appear s The relatio n s betwee n the fau na s

.

a nd fl o ra s of the s everal period s which ca n hardly be

,

proved a nd yet m ay be c o nfi de n tly s urm i s ed s upply ‘

i n dicatio ns of the cour s e a nd to s ome degree of the

,

proce ss of E volutio n Palaeo n tology i n thi s s e ns e may

, .
, ,

be defi ned a s the s cie nce that decipher s the tor n a n d

blotted s cript i n which the hi s to ry of orga n ic evol utio n
h a s bee n writte n . Biol ogy ( u s i n g the term i n a narrow
a nd fortu n ately ob s oles ce n t s e ns e) ca n s u s pect a n d ,

eve n experime n t upo n the pri n cipl e o f evolutio n ;but

,

Palaeo ntology alo n e ca n trace the cour s e of it s progre s s ,

a nd di s play the broad view s that are n eeded for a

p
precia t io of a proce s s so s low a n d o far reachi ng
n s - .
6 IN VE R TE B R AT E PALAE ON T OLOGY

( )
I I I U S ES
Pal aeo ntol ogy occupie s a n al mo s t u n ique po s itio n
a m o n g the s cie nce s i n that it h a s n ot a nd ca n har dly be
,

expected to acqu i re a n eco n om ic s ide Thi s q ual ity

,
.

of purity will preve n t tho s e wh o value s cie n ce for i ts

fi na n cial yiel d from devoti n g their e nergie s to s o p ro fi t
”
le s s a s tudy To other s the ab s e n ce of the tai n t of
.
,

com mercial i s m i s not i ts l ea s t attrac t ive featu re I t is .

n ot tha t the s cie nce doe s n o t co ntribute d irectly a n d ,

i n directly to hu m a n progre s s ( s uch barre n d i s ti nctio n

,

pertai n s to s ophi s try not to true philo s ophy) i n deed it

, , ,

m ay ofte n po s se ss that lowe s t of al l attribute s a mo n ey ,

“ ”
va lue The l ocal expert who s in k s a s haft for coal
.

through graptolitic s hale s m ight s ave h i s own and

h i s s upporter s m o ney b y the use o f elem e ntary
’

palaeo n tol ogical k nowledge N everthele s s it m u s t be

.

co n fe s s ed that di s covery of a ne w fo s s il i s not l ikely to

prod uc e the s am e ki n d of re t ur n that follow s sy n the s is
of a n e w dye fabricatio n o f a n e w machi n e or e ve n s tudy
, , ,
_

of the l ife hi s t ory of i nsect s or fu ngi A s a re s ul t it is

- .
.

pe cul iarly ea s y for s hall ow critic s to di s m i s s fo s s il

hu n ti n g as a futile hobby s uited to the requireme n t s of
,

a s chool b oy who h a s tired of s tamp s a n d m arble s or to ,

the po n derou s dotage of a retired cra n k who h as the

m ea n s a n d lei s ure to dally with the few year s rem ai n i ng
to him S uch critic s woul d not be appea s ed by the
.

re ply that Palaeo ntology h a s a charm that i ns pire s i ts

devotee s to perpetual a nd i n crea s i ng s ervice they m ight
—

j u s tly argue th a t the s am e plea s a n t thraldom i s exe rted

by gambli n g a n d o t her pur s uit s of doubtful profi t a nd
m orality .But a P alaeo ntologi s t s houl d n ot be a n
a p ologi s t for h is s cie n ce whe n the voca t io n s O f prophe t
,

a n d eva n geli s t are ope n to him .

F ir s tly Palaeo ntol ogy i s a s cie nce a nd wo ul d have

, ,
8 I NVE R TEB R AT E PALA EON T OL OGY

of t he t re n d of evol u t io n ge n erally a nd i n detail lea d s

, ,

autom atically t o i mp rove d preci s io n i n the i nte rpretatio n

of the record s of S tratigraphy I t is s u ffi cie nt t o refe r
.

to the chao s o f u n certai n ty that e ns hrou d s the Pre

Cambria n era a nd to com pare that ig n ora n ce with the
,

relatively clear u nd er s ta n di ng of s uch a pe riod a s t he

J u ra s s ic to s how that H i s torical Geology i s practically
,

d epe n de n t upo n fo s s il evide n ce O f l ate year s detailed

.
,

s tudy of the palaeo n tol ogical charac t er s of G raptolite s ,

Coral s a nd Am mo n ite s h a s revealed feature s of im me ns e

theoretical a nd p ractical importa nce to S tratigrapher s .

The lo n g a nd too ofte n m i s lead i n g l i s t s of fo ss il s that

, ,

occur i n m o st s tratigraphical paper s are te s timo n ie s t o

the fu ndam e ntal val ue of P al aeo n tology i n s uch re search .

N ot o n ly d o fo s s il s s upply evide nce for comparative

“
dati n g of rock -sy s t em s but by the m odificatio ns a nd
,

propor t io n s s how n by fau n al group s they i nd icate th e ,

co nditio n s prevale n t d uri n g their exi s tence Attem p t s .

to re s tore the phy s iography of pa s t period s gai n m ore

reliable s upport from Pa laeo ntology tha n from s tudy of
the l ithology o f s edime nt s or eve n de s criptio n of t he
k n ow n di s tributio n of s t rata I t is n ot for a Pa lae o nt o
.

logis t to write a n apology for the s tudy of Geology but ,

he would be a n u n rea s o n i n g s ceptic who doubted th e

practical val ue of that S cie nce A n d S tratigraphy the
.
,

cul m i n atio n of Geological i n quiry depe nd s al m o s t ,

whol ly upo n the evide nce that Palaeo n tology alo ne ca n

provide .

Thi rdly the archaic s chool O f B iology th at was con

,

te n t to re s trict i ts a ctivitie s to the s tudy of exi s ti n g

orga n i s m s regardi n g fo s s il s a s i ntere s ti n g though s om e
, ,
”
tim e i co ve ie t extra s s now well nigh exti nct
s n n n n ,
“
i ,
- .

A [s tude n t of hu ma n hi s t ory wh o s trove t o u nd er s ta nd

a nd explai n the i n t ricacie s of diplomacy that led u t
p o
-

the rece nt worl d war withou t refere nce t o a ny event s

 T H E S C I E N C E OF PALA EON T O L OGY 9

p r i o r Augu s t 1 9 1 4 woul d be in no wor s e pligh t tha n

to ,

a Biologi s t who s tud ied pre s e nt fau na s or flora s withou t

taki ng i nto accou nt tho s e that preced e d a nd ge nerated
th em There yet rem ai n biologi st s who di s s ipate their
.

e nergies i n e ffort s to fabricate:s cheme s of evol utio n i n

defi a nce of palaeo ntological evide nce ;they have s till t o
learn that the pict ure s que m i s t s of hypothes i s however ,

val u able where d ire ct obs ervatio n i s l acki ng m u s t yield ,

before the s u ns hi ne of truth a s revealed by fact s .

The co ncep tio n of E volutio n h a s bee n th e guidi n g

s tar of all rece n t biological s tudy however i t s i nt erpreta
,

tio n m ay vary I t wa s n ot by accide nt tha t the two

.

protago nis t s i n the i ntrod uctio n of that doctri ne ,

Lamarck a nd D a rwi n were Palae on tologi s t s of wide

,

exp erie n ce E volu t io n m ight well have ari s e n a s a

.

theory am on g p ure Biologi s t s ;but to Pa laeonto

“

logis t s it i s a m a n ife s t a nd prove n pri n cipl e A s to t he .

determi na nts a nd expre s s io ns of thi s all pervadi ng law -

there i s s till ample S cope for di s cu s s io n a nd re s earch .

R ece n t palaeo n tological work h a s n ot o nly co n tributed

m uch to the s ol utio n of the s e p roblem s but bid s fair to ,

s upply more evide n ce to that e n d The Biol ogi s t ca n

.

s ugge s t that certai n m etho d s of evol utio n m ight have

bee n em ployed a nd that certai n cour s e s m ay have bee n

,

followed ; it res t s with the Palaeo ntologi s t to s ho w

whether or not s uch thi ngs have happe n ed Whe n the .

exte n t t o which bel ief i n E vol utio n pervade s all pha s e s

o f huma n thought i s real ized the value of the s tudy of
,

fo s sil s is appa re nt I n d ire ctly b ut n o n e the le s s s urely

.
, ,

it h a s e n larged the outlook of rece n t ge neratio n s whil e ,

'

yet in i ts i n fa ncy a s a S cie nce I f s uch re s ult s h ave

.

bee n achi eved whe n the pio neeri n g traver s es are

hardly com pleted what m ay be expected from detailed
,

s u rvey of the territory compri s ed withi n the provi nce of

Palaeo ntology
 C H A PT E R I I

MAT E R I ALS

( )
A T H E N ATU R E O F F O S S I L S

(I) D E F I NI T I O N OF T ERM S F OS S I L A ND
P ET R I FA CTI O N
“
HE am e fo s s il w h ich mean s a thi n g dug Up ,
n ,
“

was applied by early Geol ogi s t s t o objects that coul d

be extrac t ed from rock s bu t t hat were not s trictly
,

s a mpl e s of the rock s I t thu s com pri s ed s eparable

.

m i ner a l s ( particularly tho s e i n vei ns or geod es ) a s well

“ ”
a s the curiou s ly s haped s to n e s of orga ni c ori g i n
, ,

as t o who s e character s m uch co ntrover s y exi s ted .

The s e two k i nd s of fo s s il s were d i s ti ngui s hed by the

” “
adj ective s n ative a nd ex tra n eou s re s pectively the ,

l atter types be i n g clearly di s ti n ct from a nd largely ,

i n depe nde nt of the rock s e n clo s i n g them Whe n con

, .

t i nue d s tudy had plai n ly d emo n s trated the biological

qual itie s of extra n eou s fo s s il s that n on-com m i t tal
,

“ ”
adjec t ive wa s replaced by the word orga n ized a nd
'

the exp re ss io n orga n ic remai ns bec am e ge nerally

“
em ploye d A t a bou t the s am e tim e the term petri
.

”
factio n cam e i n to freque n t use i n dicati n g tha t fo s s il s
,

ha d ofte n becom e m i n eral ized du ri n g e n tomb me nt i n

“ ”
the rock s Gradu al ly applicatio n of the word fo s s il
.

to purely m i neral s p eci me ns dropped out a nd with i ts ,

“
re s trictio n to the orga n ized k i nd s n eed for qualifying,

adj ec t ive s d i s appeared I n i ts m ode rn u sage t he wor d

.
 MAT E R I ALS 1 1

fo s s il m ay be take n t o refer to taos o re ma i ns or tr aces

’
ofp as t life tha t occu r i n tk e r ock s o f tile E a r tlt s cr us t , a nd

afi or d oi ologica l f
evi d ence o t/zei r
The orga nic or i gi n .

la s t phra s e i n thi s defi nitio n exclude s s uch phen ome n a

a s be d s of graphite or m a s s e s of m arbl e whi ch retai n

n o ve s tige s of orga n ic s tructure a n d give o nly pre ,

s u mp t ive evide n ce of orga n ic origi n .

The defi n itio n s how s a certai n l ack of preci s io n ,

for rea s o ns s im ilar t o tho s e that m ade limitatio n of the

s cope of Palaeo n tology im practicable A n e mp ty s he ll .

lyi ng on the bea ch covered with s a nd by the was h of

,

o n e wave a n d laid bare by the s cou r of the n ext h as

, ,

fulfilled the co nditio ns r e quired to a s s ume the titl e of

fo ss il ;a n d the s ame q ual ificatio ns wou ld appert ai n to
a body exhu med by order of the H om e S ecreta r y If .

the s hel l were d ug up from a prehi st oric kitche n -m idde n ,

or the body di s i n terred from a tu m ul u s applicatio n ,

of t he t erm fo s s il would be o nly le s s i nappropriate .

S i nce orga nic s tructure s may re ma i n practically u n

cha n ged from very remote period s it i s u seles s to ,
“
i nvoke the word petri factio n a s i n dic ati n g a cri terio n
fo r the recogn itio n of fo s s il s The u ngai nly phra s e
.

“ ”
s ub fo ss il
- h a s s ometi me s bee n appl ied to tho s e
orga nic rem ai n s t hat occur o n the border -li ne betwee n
pa s t a n d pre s e nt ;b ut s i n ce the ety mology of that
term i s grote s que a nd it s S cope devoid of preci s io n
,

i n both directio n s i ts us e rather aggravate s the d i ffi

,

culty I n practice it i s co nve nie nt to exclude orga n i s m s

.

of the H oloce ne peri od from tho se that are called fo ss il s ,

but s uch a d ivi s io n is purely arbitrary a nd at time s ,

u nte nable I n s ome a spect s of h is work a Pa la eo n

.
,

t ologi s t i s com pelled to regard him s el f a n d the livi n g ,

creature s t hat s urrou n d him as pote ntial fo s s il s

,
.

The i n t roductio n of the word petrifactio n i nto the

prece d i ng paragraph dema nd s a furt her defi n it io n .
 12 IN VE R T E B R ATE PALA E ON T OL OG Y
“
J u s t as the n am e fo ss il h as com e t o be u s ed s ome
what loo s ely ( for example a s a m ild term of abu s e )
, ,
“ ”
so petrifactio n both a s a proce ss a nd a s a re s ul t
, ,

i s ofte n appl ie d i n ca s e s where it i s i n appropriate .

The two word s are freque n tly treate d a s s y n onym s ,

bu t n either fact s n or etym ology s upport s uch id e nt i

ficatio n Pa s s i n g over the obviou s ly i nacc urate u se of
‘

“
the word i n co nn exio n with t he prod uct s of a petr ify
”
i n g s pri n g ( where obj ect s become coa t ed with fi lm s
“ ”
of calciu m carbo n ate precipitated from hard water ) ,

it m ay be s ta t ed that while all petrifactio n s are fo ss il s

, ,

al l fo s s il s are n ot petrifactio ns The co nvers io n i nto

.

s to n e
( or n e w m i n eral m atter ) that i s impl ied by the
l atter term i n volve s a co ns iderable cha n ge in the s ub
’
s ta n ce petrified whether i t be achieved by Med u s a s
,

head or by le s s heroic method s Petrifactio n i s believed

.

to b e carried ou t by m olecul ar replacem e nt a nd will ,

be di s cu ssed i n a later s ectio n of thi s chapter ;but a

l arge proportio n of fo s s il s have u n dergo ne no S uch
radical cha n ge I mpreg n atio n of the s pace s of origi n
.

ally porou s s hell s with t h e s ame or s om e o t her m i neral

m ay po s s ibly ra n k a s p etrifactio n s i n ce the proce s s is
,

s im ilar to that whereby s a nd -grai ns becom e u n ited

“ ”
i n to the rocky s tate of s a n d s to n e But i n preci s e .
,

term i n ology the word petrifactio n m u s t be re s tricted

,

to tb ose f oss ils w/zi clt na ve u nd ergone c/ea nge ei ther by

com p le t e r ea r r a ng e m en t of tb c molecu le s o f tb c m i ner a l
of wl z i clt ta y
e w er e com p os ed ( whe n all m i n ute s tructure
i s lo s t ) or by s u bs ti tu ti on of a d if er en t mi ner a l ( whe n
, ,

if replaceme n t h a s bee n grad ual de t ail s of s tructure ,

m ay be retai n ed ) .
 MATE R I ALS

( )
I I C O N D I T I O N S N E CE SS A R Y F OR FO S S I L I Z ATI ON

( )
a B i ologica l

Although i n exceptio n al ca s e s impre s s io n s a nd eve n ,

pet rifactio ns of s oft ti s s ue s have bee n fou n d fo s s iliza

,
1
,

tio n i s not likely to follow the death of a n a n imal that

failed t o s ecrete mi neral or othe r che mica lly s tabl e
,
.
,

s truc t ure s d uri n g life Mo s t purely orga n ic ti s s ue s '

be com e di ss ip a ted by decompo s itio n very s oo n afte r

dea th eve n if they e s cape d ige s tio n i n the alimen tary
,

t ract of a co n temporary S i nce a large propo rtio n of

.

I nvertebrate s now livi n g part ,

a n d s maller type s are without

,

exi s ti ng fau na c a nnot be e x p e c

m ately complete repr e s e ntatio n for s tu dy by future
palaeo ntologi s t s There i s no rea s o n t o bel ieve tha t
-
.

the proportio n of n ormally peri s hable form s was ever

r

i n ferior t o tha t at the pre s e nt day— probably it wa s

greate r in m ore rem ote pe riod s Palaeo ntology i s thu s .

practic a lly cut o ff from the s tudy of a very co n s iderable

n u mber of group s of a n i m al s ;a n d a s a rule ca n deal , ,

directly on ly with the hard part s of s uch cla s s e s as are

capable of fo s s E ve n am o n g s uch s tructure s ,

m uc h d iver s ity ili ty exi s t s The hard p a rt s .

of I nvertebrate s m ay be roughly cl a s s ified a s cohere n t

or di sj u nct a nd as siliceou s calca reou s or chiti nou s
, , .

Cohere n t s hell s or s keleto n s ( s uch a s tho s e of typical

G a s tropoda or E chi noidea ) will retai n their s hape after
di s s ol utio n of the a ss ociated s oft ti s s ue s a n d s o m ay ,

provide s ati s factory evide nce of the form a n d character s

1 roz en ca rcasses of anim ls preserv ed b y cold storage in Arctic
Th e f a -

regions since the Pleistocene peri od a e geologi ca lly consid ered transie nt
r , , ,

an d could not survive a ny cli mati c cha ng e s th a t ra i sed the te mperature

of their surround i ng s .
14 IN VE R T EB R AT E PAL AE ON TOL OGY

of the a n im al s t hat built them ;bu t th o

‘

se th at are
made u p of loo s e s cale s Or o s s icle s ( e g m a ny S po n ges . .
(
,

H olothuria n s a n d Arthropod s) held i n place by m u s cle s

,

or ligam en t s wil l coll ap s e i n to fragme n t s on the death

,

of their ow ne r s a nd r eco n s t r uctio n will be a ma tt er of

,

di ffi cul ty a n d u n certai nty .

The few a n im al s th at build s kele t al s t ru c t ure s of

s il ica ( chiefly R adiol aria a n d S po n ge s) s ecrete that
m i neral i n the opali n e or a morphou s co n ditio n— i t s lea s t
d urable form — liable t o rem oval by s ol utio n in alk al i n e
water Cal careou s s hell s or s keleto n s are s ometime s
.

m ade of calcite a form o f calci um carbo nate which i s

,

rela t ively s table ( s ee Pl iii fi g

. . bu t n o n e the le s s
.

readily s ol uble in a cid ul ated water ;tho s e t ha t a re co m

po s e d of the rhombi c Arago n ite are par t icul arly p ro n e
to s olu t io n a nd decay ( s ee Pl iii fi g . Chiti n which
. .
,

i s a n orga n ic s ub s ta n ce far m o re s table tha n ordin ary

s oft t i s s ue s ( com parabl e i n thi s re s pect with the wo ody

fi bre of pl a n t s) ca n be pre s erved o nly if hermetically

,

s e a led up s oo n after death ( s ee Pl iv fi g 1 ) eve n the n

. . .

it o fte n pa s s e s i nto a carbo n ized a nd p artly collap s ed

co n ditio n .E ve n whe n m ixed with variou s s alt s of
l ime ( a s i n m a ny l arge Cru s tacea ) it i s s till tra ns ie nt
,

if expo s ed to atmo s pheric a ctio n H e n c e the po ss es s io n

.

of S hel ly s truc t ure s i n n o way e n s ure s the fo ss ilizatio n

of a n orga n i s m al t hough it i s n ece s s ary i f s uch a con
,

s um m a tio n i s to be n orm ally attai n a ble .

Agai n s ecretio n of m i neral m atter i s t he e s pecial

,

a tt ribute o f a n im al s I n ad ult s tage s ;i t i s ra rely or but

feebly developed i n you ng or larval forms H e nce a .

large proportio n of the m aterial avail able for pa la e onto d

logical s tudy co n s i s t s of fully gro wn s pecim e ns ;early

developm e ntal s tage s have rarely left r eco gn izable
t race s S i nce the hi s t o ry of a n i n dividual life s eem s
.

t o pre s e nt a n epitom e of t he evol u t io n of the rac e to

16 IN V E R TEB R AT E PALA EON T OL OG Y

much of t he We al d e n a nd t he R e a d i n g cl ay s need not ,

be a s cribe d to a paucity of fau n a ;it is more p robably

d ue t o t he c a u s e s m e nt io ned abo ve .

The se a i s t he chief recep t acle for t he p r od uc ts of

rock -wa s t age a nd it i s a fo rtu n ate coi n cide nce tha t t he
,

m aj o ri ty of s h ell -beari ng I nver t ebrate s ( excludi n g the

I ns ec t s ) are m ari ne D epo s it s accu m ula t e i n gre a t est
.

“ ”
bulk a nd m os t rapidly i n the de t rital zon e which
, , ,

exte nd s over t he u s ually shallow part s of the s e a arou n d

co a s t -li n es A gai n the s ati s facto ry phe nome no n ca n be
.

rec orded t ha t the riche s t z o ne s of I n vertebrate m a ri ne

l i fe ( perhap s excl udi n g the Protozoa) coi ncid e ve ry
n e a rly wi t h the bel t of coa s tal wa t er where p rompt

burial i s m o s t likely to follo w death F u rther rock s of

f

.
,

t he de t ri t al zo n e a re the m os t favou rably s i t uated for

s ub s eque n t elevatio n s o t hat the fulle s t record s O f pa s t
,
-

life available for palaeo ntological s tudy a re n ormally

tho s e of the d e n ize ns of the m argi n al tract s of the s ea .

O rga n i s m s left upo n or drifted to the beach will al m os t

, ,

certai n ly s u ffer from the abradi n g i n flue nce o f wave s ,

a n d m ay becom e expo s ed to a tm o spheri c weatheri n g .

The l arger I nvertebrate s of deep water are for the m o s t ,

pa r t i ne ffi cie n tly clad in m i n eral arm our a nd he nce are

, ,

n ot wel l adapted for pre s ervatio n i n tact The ideal .

s ituatio n for fo s s il izatio n i s i n the s hallow wat er be low

‘

t he level of l ow tide s where the s cou r of curre nt s

,

a nd wave s i s i n s u fficie n t to preve n t depo s itio n of fi n e

s ilt. Coral s S e a Urchi ns B rachiopod s M ollu s c s a nd

,
-
, ,

Cru s tacea n s are the group s that abou nd or have

'

flouri shed ;m o s t i n s uch s urrou n di n gs ;their rem ai ns

co ns titute the chief part of the palaeo n tological evide nce
a vailable f or s tudy of the larger I nve rtebr a tes .
 M ATE R I A LS

Po sse ss io n of d urabl e s tructure s a nd s elec t i on of a

,
'

l ocal ity favourable for b urial i n n o s e ns e en s ure fo ss iliza

,

t io n The n ature of the m aterial s in which s hell s are

.

em be d ded a nd the geo logical i n flue nce s that m ay

,

operate on t hem m u s t i nevitably a ffec t the pre s e rvatio n

,

of fo ss il s The three type s of s ed im e n ta ry m atri x which

.

m o s t frequ ently co ntai n fo ss il s are t ho se k n ow n a s

a re n aceou s ( s a n dy or gritt y
) argillaceou s (
,
cl ayey ) nd
a ,

calcareou s ( ooze or s hell -s a n d ) Whe n s uch depo s it s

.

are form ed u nder m ari ne co nditio ns it is u s ual for the ,

fi rs t -named type to occur on or near the bea ch for the ,

s eco n d t o accum ulate i n the outer part of the det rit al

zo ne a nd for the third to be s pre a d over the s hallowe r

,

p art s of the ope n ocea n S ome calcareou s rock s s uch

.
,

as ool ite s a nd s h e ll -l i me s to nes are developed i n the

,

pos itio n n orm ally occ upied by s a nd A lthough n o .

defi nite l i ne ca n be draw n betwee n the th ree types o f

sedime nt owi n g t o the freque n t i n term i n gli ng of thei r
,

i ngred ie n t s wh e n rel atively pure they afford very

,

di ffere nt m atrice s .

S a n d i s largely compo se d of wor n grai ns of qua rtz ,

the harde s t a nd m o s t d urable form of S ilica a nd is ,

likely to e nclo se abu nda n t remai n s of s hell beari ng -

I nvertebrate s owi ng t o it s littoral di s trib utio n But it .

I s e mI ne nt l porous a n d percolati ng water which ca n n ot

l y , ,

attack quartz will fi nd i n the c alcareou s s he ll s mate rial

,

relatively ea s y to di ss olve S a nd bu rial thu s a fford s

-
.

but poor protect io n from the wea t he r The e n clo s ed .

rem ai ns may occa s io nally become petrified by repla ce

ment if the water that di ss olve s them hol d s s om e le ss
,

s oluble s al t s i n s oluti o n ; but m ore ofte n they a re

e nti rely removed I f the ar enaceou s rock h as bee n
.

compa cted before the di ss olu t ion of t he S hell s c as ts a nd ,

2
18 I NVER T E BR A TE PALA EON TOL OGY

m oul d s will be left r epre s e nti ng the fo s s il s i n n ega t ive

,

( see Pl
. Bu t if s ol utio n t ake s place befo re the
s a n d h a s becom e co ns olidated the grai n s clo s e i n to
,

fi ll up the gap s a nd no trace s of fo s s il s m ay remai n

, .

The s oluble m i n eral m atter is carried away by perco

l ati n g water to be red epo s ited el sewhere a s a cemen t
,

betwee n the s a n d -grai ns or a s a filli ng for crack s i n

n eighbouri n g rock -ma s s e s F urther the i nte ns e hard nes s
.
,

of the quartz a nd other grai ns whe n co mpared with ,

that of cal cite or a rago n ite cau se s a ppl icatio n of eve n

,

ge n tle p re s s ure to des troy the fi ner d eta il s of s hell s .

The re s ulti n g pi t ted s u rface ofte n with di fficulty d is ,

e n gaged from the pe netrati ng s a n d bear s l ittle or no ,

res embla n ce t o that of the origi nal s hell eve n i f ,

the fo ss il h as e s caped com m i nutio n in i ts u n co nge n ial

s u rrou n d i n g s .

Cl ay bei n g made of mi n ute flake s of s oft m ic a ceou s

, ,

m aterial a n d bei n g practically im perviou s to wa ter h a s

, ,

n ei t her of the d i s adva n tage s of s a n d as a ma t ri x fo r

'

pres ervatio n of fo s s il s I n s uch depo s it s a s the Gaul t

.

or the Barto n C lay fo s s il s ofte n retai n their origi n al

,

“ ”
m othe r O f pearl s hee n a nd pre s e n t all the detail s
- -
,

of their or name nt as clearly a s rece n t s hell s E ve n .

chiti nou s or l igam e ntou s ti s s ues m ay b e pre s erved

al m o s t u n cha n ged But i n s pite of thes e good poi nt s
.
,

argill aceou s rock s are l iable to develop two qualitie s tha t

have a ba neful i n flue n ce o n their co n tai n ed organ ic
rem ai ns The flaky a nd waterlogged co n ditio n of the
.

“ ”
m aterial s that give the clay charac t er ca n pers i s t
o nly i n u n compre s s ed s ilt Prol o n ged depo s itio n bri ng s
.

the e a rli er f orme d parts u nder the weight -pre ss ure of

overlyi n g s ed ime n t with the res ul t that the s m all flakes
,

are rearra nged i n parallel l ayers a nd i ncl u d ed water i s ,

s queezed out The sh ale thu s form ed s hows great

.

reductio n i n thick n e ss from i ts previou s b ul k a nd co m ,

 MATE R I ALS 19

pre ss io n i s bou nd to i nvolve al l but the m o s t m a s s ive of

“
H e n ce fo s s il s pre s erved i n
”
i t s ex t ra neou s co n te n t s .

s hale s are n orm ally fl atte ned the s m all qua n tity of clay
,

that filled the i r cavit i es bei n g red uced to a n al m o s t

n egl igible fil m ( s e e Pl iv fi g . N ot o nly doe s thi s
. .

proces s re s ult i n lo s s o f form but it freque ntly cau ses,

the two s ide s of the fo s s il to be tele scoped i nto o ne

a nother with very co n fu s i n g e ff ect The d ifli culty o f
, .

re cd ns t r uct i o n of exti n ct t pe s who s e detail ed s tructure

y
c a n n ot be ded uced fro m s tudy of l ivi n g relative s i s
greatly i n crea s ed whe n the character s of both s urface s
are th i s combi ned Pai n ful exam ple s of thi s u m
.

fortu nate type O f pre s ervatio n are afforded by the S e a

U rchi ns a nd S tarfi s h fou n d i n argillaceou s fla gs t one s of
the Lower L udlow s erie s S eco ndly the impermeable
.
,

n ature of a clay m atrix while e ffectively excl udi n g

, ,

de s t ructive actio n from without no le ss e fficie ntly ,

‘

e ncourage s c hem ical cha nge s withi n the rock E ve n .

the purely s oft ti ss ue s of orga n i s m s m ay b e buried i n

t he s ilt a nd by their s low decay give ri s e t o variou s
,

volatile s ub sta nce s s u ch a s s ulphuretted hydroge n tha t

, ,

s tro ngly affect m i n eral s doomed to a ss ociatio n with

them The productio n of Marca s ite ( s e e Pl v fi g 2 )
. . . .

( a n u n s table form o f I ro n s ulphide) a nd S ele n ite

( S ulphate of l ime ) are two freque n t re s ult s o f s uch
reactio n s an d ofte n thes e two wholly cry s talli n e m i neral s
,

afford the s ole evide nce of previou s exi s te n ce of fos s il s

i n cl ay . N everthele s s i n s pite o f the s e two s eriou s
,

di sa dva nt ages Clay is the be s t med iu m for the pres erva

,
'

tio n of fo ss il s.

A calcareou s m atrix i s n o less porou s tha n a n

are naceou s o ne but the S imilarity betwee n it s phy s ical
,

propertie s a n d tho s e of the co n tai ned orga n ic remai ns

( s uppo s i ng the se to be m ade of calcite) m itigate s the
da nger s to whi ch the la tt er are s ubj ect Although the .
20 I NV E R T E B R AT E PALAE ON T OLO GY

bulk of a calcareou s rock i s orga n ic i n o rigi n a nd thu s ,

i n s om e me as ure cry s tall i ne a l arge propo rti on i s s o

,

fi nely com m i n ute d that it react s alm o s t as if it were

“
a morphou s H e nce the m atrix of a l im es to ne u s ually
.

prove s m ore read ily s ol uble i n atm os pheric water tha n

the s hell s t h at it i nclud e s f a nd th ese are i n co ns eque nce
le s s l ikely t o be s eriou s ly harmed ( s e e Pl i fig ( I f . . .

t he fo s s il is m ade of Arago n ite O pali n e s ilica or Chiti n

, ,

s ol utio n ,
decay or pe trifact io n is al m o s t i n evitable ) .

I t i s u s ual t o fi n d a s eco n dary i n fi lling of calcite ,

derived from percolati ng water occupyi n g all the s m aller

,

cavitie s i n the s hell ti s s ue but thi s n eed not ob s cure

-
,

the origi nal s t ructure Agai n the m at rix co ns i s t s of

.
,

m a t erial which is theoretical ly of th e s am e har d ne ss a s

the e n clo s ed fo s s il s a n d which i n practice is s ofter owi ng
,

t o i t s powdered co n ditio n . H e nce the pitti n g that s o

ofte n defaces s hel l s i n s a nd s to n es i s u nl ikely to b e
im pre s s ed on tho s e i n a l ime s to n e .

E xcept i n m i ne s a nd bori ng s fo s s il s ca n be c ollec t ed

,

o nly fro m rock s e xpos ed a t the E arth s s urface S uch ’

.

roc k s are al way s u n derg oi ng de s t ructio n b y weatheri n g

age ncie s a nd the fo ss il s they co nt ai n m u s t s ubm i t to

,
“ ”
the s am e fate however extra neou s thei r origi n Thu s
, .

t h e m o s t perfect a nd com ple t e pre s e rvatio n is bu t

temporary in relatio n to geol ogical time I n ma ny .

d i s trict s s edi me n ta ry rock s t hou s an d s of feet i n thick ne s s

have bee n rem oved by de n udatio n at variou s geological
period s a n d no ve s tige s of their fau na s are pres erved
, ,
"

s ave for occ as io n al water - wor n relic s embed d ed i n l ater

d epo s it s or cha n ce occurr e n ce s i n the pebbles of
,

co nglom erate s But the tool of ero si o n i s d ouble—ed ged

.
,

a n d bri n gs t o l ight fo s s il iferou s rock s that wo uld other

wi se rem ai n buried a nd out O f reach O ther a nd m ore .

merci less a ge nt s m ay wo r k hav oc on roc k s a nd fo ss il s

‘
,

wi t hou t a ffo rd i ng a ch an ce for rescue by collec ti o n .

 MATE R I ALS

H ypoge ne force s which prod uce met a s om ati s m i n

,

rock -ma s se s buried to a great depth or i nvolved in cru s ta l

moveme n t s m ay cau s e di s tortio n rec rys tallization or
, , ,

eve n ma gma ti z a t i on of m aterial s affected A n e s s e n tial .

fe ature of s uch actio n i s the re -groupi n g of mole culeS }

either i nto ne w c ry s tal s O f the origi nal m i n eral s or ,

i nto e ntirely fres h m i n eral t ypes A q uartzite m ay .

occa s io nally retai n rough i n dicatio ns of the fo ss il s that

were o nce embedded i n it ;the refractory nature of
quart z te nd s to red uce it s reactio n to metam orphic
force s. But a clay -S late i n which the a n n eal ed
,

m icaceou s flak es have s lewed rou n d to po s itio n s di fferen t

from tho s e due to sedime ntatio n will i nvolve i ts con ,

t a i ned fo s s il s i n the di s tortio n a n d cleavage to which it

h as bee n s ubj ected Phyl lite s a nd s chi s t s have u n der
.

go n e s u ch com plete recry s tallizatio n that n either the

origi n al rock no r it s co n te nt s are clearly recogn izable
‘

S imilarly a m arble re s ult s from complete rearra ngeme n t

,

of the calcite m olecule s of a lime s to n e the s accharoidal

com ple x of cry s tal s retai ns n o biologi c al i n dicatio n of
orga n ic origi n F i nally whe n heat a nd pre s s ure are
.
,

applied i n full force roc k -m atter u ndergoe s that m ag

,

“ ”
m atic fu s io n o r i nte rs olut ion tha t re s ul t s later i n
“ ”
fo rm atio n of I g neo u s rock s I n thes e no s u s picio n
.

of orga nic s tructu re or i n flue nce po s s ibly excepti n g ,

diamo n d s ca n be ex pected to occur The lo n ger a

, .

rock h a s e xi s ted the greater ha s bee n i ts da nger from

,

hypoge ne force s H e nce the record of the fa una s o f

.

’
a n cie n t per iod s of the E a rth s hi s tory te nd s to become 1

m ore im pe rfect a s it become s more rem ote E xtra .

ord i n ary exceptio ns s uch a s tho s e afforded by the wo n der

ful Cambria n fo s s il s of Briti s h Col u mbia o r the al m
,

os t ,

Tertiary quality of the O rdovicia n cl ay s of N orthe rn

R u s s ia s erve merely to em pha s ize the tr a gic ded uctio ns
,

late nt in thi s gen eraliza t io n .

22 I N VE R TE B R AT E PALA EON T OL OGY

( )
I I I I M PE R F E CT I O N OF PA L A EO N TO L OGI CA L
E V I D E N CE

The accou nt of the co n ditio ns n ece s s ary for fo ss il iza

t io n give n i n the p recedi n g paragraph s s how s clearly how
rare m u s t be fulfi l me n t of tho s e co nditio ns Bu t the .

three e s se n tial s there i n dicated ( which m ay be s u m

ma r iz e d a s poss e s s io n o f hard -part s a m ode of death not
,

i nvolvi n g d e s tructio n of the s h ell a n d a plac e a nd ,

med iu m of burial whe nce age n t s of decay are excl uded )

by n o m ea ns i n clude al l that i s requ ired for fo s s il s to
“ ”
becom e palaeo n tological evide nce F or s u ch a con
.

s u m m atio n the rock i n which the orga n i s m h as bee n

,

e n t omb e d m u s t remai n o n o r be elevated i n to d ry la n d ;

, ,

it m u s t be e x po s ed by n atural or artificial e xcavatio ns

i t m u s t be vi s ited by a “ fo s s il -co llector
”
a nd at the
_

preci s e ti m e whe n the s tratu m h a s bee n l aid bare E ve n .

the n there is always the po s s ibil ity that the fo s s il m ay

be overlooked or broke n beyo n d repair d uri n g extrac
,

tio n it m ay col lap s e duri n g preparatio n d i s i n tegrate ,

d uri ng s torage or with tragic batho s be lo s t

, , , .

I t ha s bee n s hown that the chief fo s s il -beari n g rock s

are tho s e formed on the s e a floor E levatio n of the-
.

s ed ime n t s i s thu s u s ually i n evitable if their fo s s il -co n te n t s

are to be di s played I t i s po s s ible perhap s eve n

.
,

probable that i s o s tatic cru s tal m oveme n t s will n ormally

,

rai s e the detrital zo n e s alo n g the s e a m argi n s oo ner o r

-
,

later Bu t whatever views ma y be hel d a s to perm a

.
,

n e n ce of oc ea n ba s i n s it i s O bviou s that depo s it s form e d

,

o n the floor of deeper part s of the se a are likely to

reappear above it s s urface o n ly very l ocally ( i n volca n ic

regio n s) or after l o n g delay Thu s while the coa s tal
.
,

fau na i s favourably s ituated in thi s re s pect the less ,

profu s e but n o le s s i ntere s ti ng life of the o pe n ocea n

, ,

will rarely be acces s ibl e i n the fo s s il s t ate D eep se a .

-
24 IN VE R T EB R AT E PALA EON T OL OGY

M i ne s other tha n coal m i ne s are rarely d rive n i n fo s s ili

,
-
,

ferou s rock s while bo ri ngs are u s ually s o s m al l in

,

diameter that the core s ( if a ny are rai s ed) ca n no t i ncl ude

m uch fo s s il evide nce U nl ike m a ny m etallic ores fo s s il s
.
,

d o n ot t e n d t o i ncrea s e i n abu nda nce n ear the s urfa ce so ,

th a t it m ay be a s s u med tha t the va s t m aj ority of fo ss il s

are a s yet not only u ncollected bu t i n acce s s ible ,
.

The ruthl es s tool s of d e n udatio n which m ay carve out ,

fo ss il s from their ma t rix are n ot co n te n t with that,

b e ne fi ce nt work ;t hey will n ot res t Un til the orga n ic

rem ai ns a s well a s the rock co n tai n i ng them ar e
, ,

s hatte re d or d i s s olved M uch of this de s t ructio n i s

.

carried o ut be n eath the m a n tle of d rift s o that fo ss il ,

evid e n ce i s des troyed j u s t whe n i t s a ppeara nce s eem ed

i m m i ne n t Weather beat s co nti nual ly a gai n s t expo s ed
.

rock s urface s ;blow s from geological ham m er s a re few

-

a nd i n term i tt e nt F o r one fo ss il that i s fou n d

.

“ ”
weathered ou t s core s have bee n weathered awa y
-
, .

The s e are d i s a s ters be yo n d hu m a n co n trol bu t the ca s e

i s s carcely be t ter whe n artific ia l proce s s e s excav a t e the
rock s The average quarrym a n or m i ner u s ually pick s
.

out a n d po ss ibly cheri s hes large or otherwi s e s triki ng

, ,

s peci me n s ;but he h a s n either the tim e n or trai n i n g to

s ea rch for le ss O bviou s fo s s il s n or u s u ally t o keep s uffi

, , ,

cie n t record of the horizo n s from which h i s picked

s ample s h ave com e Mo s t quarried m aterial i s t ra ns
.

ferred direct to l im e -k il ns or s to ne cru s her s a nd i t s -

,

i n cl uded fo s s il s are a s lo s t to Pal aeo n tology a s if they

had n ever ex i s t ed O nly tho s e s pecime ns t hat happe n
.

t o be expo s ed o n the occa s io n s of rare a nd hu rried vi s it s

by fo s s il -hu nters are likely to be s aved ;m uch of the
evide nce that the se ca n s upply m a be lo s t i f the
collector s are am on g t he u n i n itiated A n at t emp t is .

m ade i n Ch a pter I I I to i ndicate the m ethod s t o be

.

em pl oyed i n collec t i ng fo s s il s s o t ha t their ful l val ue ca n

 I PE AT E I
‘

F IG .
~
x . BlOck of
'

l"
1 Ca b on ife
-
r ro us
'

Lm i es t o ne
i r r egu la r e . Po rt St . M a ry, I sle of Ma n .
" h as“ )

5 F IG
. 2 . B lock :
of
»
'
Ca rb dhi fe ro us _
" Lm i es t one co mtiOS e d
'

of colu mn a fs l Of
'

? A ct i nocr i n u s Cas tle ton, D e r b ysh i rex

‘

3 Block
2 1 '
of Ba t h o n i a n Lm
i es t on e co mpose d
La nsd owne ,
n ear Ba th .
 L
P A TE I I
 L
P AT E I I

FI G . 1

FI G 9 .

F IG . 3
26 I N VE R T E B R AT E PALA EON T OL OGY

( am o n g other d eb ri s ) a nd e n clo s e the m o n c rys t a lli z a

tio n S uch a proce ss u s ually re s ul t s i n utter de s tructio n

.

of orga n ic matter ;bu t s om etim e s the l ava co ns olidate s

before the whol e fabric i s co n s u m ed thu s per p e tuati n g
,

a n impres s io n of i t s form E ve n ear s o f cor n have bee n

.

“ ”
fou n d fo ss ilized i n rece n t lava s tream s - H oweve r
.
,

occurre n ce s of thi s n ature are s o rare that m agm atic

roc k s m ay b e di s m i s s ed a s practical ly u n fo s s iliferou s .

The prod uct s of vol ca n ic explo s io n fall i n to a d iffere n t

category While coar s e bl ock s of agglomeratic m aterial
.

a re likely to cru s h orga n i s m s beyo n d r e cog n itio n the ,

.

“
fi n er a sh or tu ff s ettle s s o ge n tly ( e s pecially u n de r
water) that it is no m ore de s tructive tha n cl ay o r ooze .

S i nce m o s t tuffs that are geologically pers i s te n t are tho se

erupted u n der or carried i n to the sea they re s e mble
, , ,

s edim e ntary rock s i n accu m ulatio n a n d he n ce m ay

,

i n cl ude fo s s il s A lthough the im med iate vici nity of a

.

volca n ic ve nt would s oo n be co me de s titute of l ife ,

regio n s far rem oved from the da n ger -zo n e receive great
qua ntitie s of fi ne a sh tra n s ported peaceably by curren t s
, .

Thu s while tu ffs a ss ociated with agglom era te s a nd lava

,

flow s are u s u al ly barre n tho s e i n terbed ded wi th ord i n a ry

,

s ed im e n t s are ofte n fo s s iliferou s .

Metam orphic rock s res ul t from rec rys tallizatio n of

o t her types u nd er the i n flue nce of heat or cru s tal
m ovem e n t E ve n tho s e s ch i s to s e varietie s who se s edi
.

m e n tary origi n i s probable have u n dergo n e m uch

m olecular readj u s tme nt s o that the origi nal character
,

of the rock a nd it s co nte n t s i s altered B ut m et a

.

m o rphi s m varie s i n d egree a n d s uch roc k s as clay

,

s late s i n s pi t e of cleavage a n d i n cipie n t cry s t allizatio n

, ,

O fte n c o n tai n recog n izabl e trace s O f orga n ic rem ai ns .

Thes e are u su ally m u ch d i s torted ( s ee Pl iii fi g . . .

i nd eed the s tate of the fo s s il s wa s o ne O f the phe nom e n a

,

that proved the tecto n ic origi n of s laty cleavage A s a .

 M AT E R I AL S 27

ge neral rule metamorphic rock s are u n fo s s iliferou s

, .

Thi s is d oubtles s one of the chief rea s o n s for our s par s e

k n owledge Of a n cie nt fau n a s s i nce the O lder rock s have
,

bee n i nvolved mos t freque n tly i n m etamorphic s tre s s .

There remai n the s edime n ta ry rock s i n which alo n e ,

orga n ic remai ns ca n be rea s o n ably expected to occu r .

A s h as bee n s how n i n a previou s s ectio n the three type s , ,

a re n aceou s argillaceou s a n d
,
orga n ic vary in qual ity
,

a s matrice s fo r pre s e rvatio n of fo s s il s S a n d s may be

.

terre s trial i n origi n ( like s ome of the O ld R ed a nd m a ny

Tria ss ic s a nd s to ne s) a n d will the n prove s carcely fo ss ili
,

ferou s owi n g pa rtly to the paucity of life i n de s ert s a nd

—
,

partly to the imperfect burial afforded Coar s e breccia s .

o r co n glomerate s are rarely fo s s iliferou s ;s uc h orga n ic

'

remai ns a s they e ncl o s e are u s u ally i n poor re p a Ir .

F i e grai ned s a n d s to ne s are ofte n replete with fo s s il s

n -

( g
e
. . the C a r a d o ci a n O f S hrop s hire or the Tha n e t S a n d s ) ,

but very l ittle expo s ure to weatheri n g reduce s the remai ns

to the s tate of ca s t s .

Cl ays a nd s hale s which a fford the bes t m atrices for

,

pre s ervatio n a re by n o mea ns alway s rICh ly fo ss il iferou s

,
.

The We n lock S hale s , Lia s s ic cl ay s Gault a n d Barto n ,

Bed s m ay be cited as argillaceou s depo s it s i n which ,

”
locally at lea s t rich haul s may be expected
,
“
But .

agai n s t thes e m u s t be pl aced the O ld R ed Marl s of

’

H ereford s hire the K eu per Marl s mu ch of the Weal d

, ,

Clay a n d the R e adi n g Bed s The s carcity of i n verte

.

brate fo s s il s i n the s e ro c k -s y s tem s i s probably due i n

large m ea s ure to the terre s trial l acu s tri n e or deltaic
,

co nditio n s of their formatio n The m oral to be d raw n

.

from the compari s o n i s that mottled clay s are rarely rich

i n fo s s il s
. Thi s i s well illu s trated i n the Wealde n of the
I s le of Wight where grey -lam i n ated clays are crowded
,

with Cyp r i d ea a n d Cy r ena while a s s ociated mottled bed s

,

are p ractically barre n Ar gillaceou s rock s are S pecially

.
 IN VE R T E B R AT E PALAE ON TOLOG Y
“

28

l iable to me t a morp h is m b ut the co nt or t io n a nd cl e avage

,

tha t have co n fu sed s uch s y s tem s as the L ongmynd i a n

or Ma n x S late s s eem i n s u fficie n t rea s o n s for their un ~

fo s s iliferou s n ature .O ther greywacke s i n m uch the ,

s ame co n ditio n co n tai n traces of orga n i s m s ob s cure

, ,

th ough they m ay be The barre nne s s of the va s t pile s

.

of argillaceou s Pre Ca mb rIa n s trata dema n d s a fu rther

-

expla natio n .

Typical l ime s to ne s are largely built of orga n ic remai ns ,

but thi s doe s n ot p reve nt them from ofte n di s appoi n ti n g

col lectors of m acro s copic fo ss il s N o t o nly does the
.

s im ilarity i n m i n eral com po s itio n betwee n matrix a nd

s hel l m ake their s eparatio n d i ffi cul t a nd fortuitou s but ,

O fte n l arge fo s s il s are wholly wa n ti n g Unles s the l ime

.

s to n e h a s u n dergo n e m arm aro s i s or d ol om itizatio n a ,

micrO S CO p i c s ectio n will u s ual ly reveal foram i n ifera or

'

s hel l - fr a gme n t s but eve n s uch appare ntly prom i s i n g

,

s trata a s the Coralli ne O olite o r U ppe r Chal k m ay

prove de s titu te of s eparabl e fo s s il s through a wea ry
s ucce ss io n of expo s ure s Lim e s to ne s of relatively lit
.

toral origi n are u s ually m ore prol ific tha n ooze s s o that ,

the b e s t fo s s il hu n ti n g i s to be fou nd i n s uch depo s it s

-

a s the S il uria n C arbo n i ferou s a nd J ura s s ic Lime s to n es

,
.

A m o n g s iliceou s rock s rad iolaria n chert rarely i n cl ude s

,

m a ny fo ss il s apart from the R ad iolaria the m s elves while ,

chert derived from d i s s olutio n of s p o n ge s picules m ay -

co ntai n abu nd a n t rem ai n s of larger orga n i s m s u s ually ,

a s ca s t s or s il iceou s petrifa ctio n s Carbo n iferou s a nd

.

Po rtla n d ia n chert s are familiar exam ple s of s uc h rock s .

Peat is ofte n crowd ed with s hel l s of fre s h wa fer -

m ol lu s ca bu t thes e rarely remai n in carbo n aceou s rpck s

,

approachi ng co a l i n co n s titutio n The orga n ic a cid s

.

pr oduced by pla nt decay s peed ily rem ove calcareou s

-

s tructures . Altho ugh mollu s c s a nd other I nvertebrates

ar e occa s io n al ly fou nd i n c a n nel they are al mo s t alway s
,
 M AT E R I ALS 29

repres en te d by ob s cure impre s s io ns I t is o nly in .

a ss oc iated s hale s or i n m i neral co n cretio ns tha t the

, ,

I n vertebrate fau na of t he Coal Mea s ure s is adequatel y

pre served .

( I I) F OS S I L S A S R O C K - F O R M E R S ( Pls i . . a nd

Or ga n ic rock s a s their n am e implie s co ns i s t o f

, ,

m atter t hat owe s i ts s olidi ty a nd form to the i nflue n ce

of livi ng bei ngs The mi ne fa l s ub sta nce s m o s t abu n
.

d a u tly secreted by I nvertebrate s are Calcium carbo n ate

a n d pho s phate a n d colloid S ilica
, S i n ce the s e m i neral s
.

are readily s oluble i n cru s tal water co ns iderable m ole ,

cula r rearra n geme n t i s bou n d to s upe rve n e i n rock s bu i lt

of them . H e n ce the orga ni c origi n of a rock doe s not

automatically imply per s i s te n ce of recog nizable orga n ic
remai s n . S ol utio n a n d r e depo s itio n of Calcite and
-
,

co nver s io n of O pal i nto Chalced o ny will e ntirely ,

obliterate s m all s hell s a n d de s troy the fi ner fea t ure s of

,

larger o ne s The m os t exte ns ive a n d m a ss ive depo s i t s

.

of calcareou s a nd s iliceou s rock are the ooze s who s e ,

chief co ns titue nt s are s m all Protozoa I n s uch depo s it s .

a s Chal k or Barbado s E arth F oram i n ifera a n d R adio ,

la ria however mi nu te are in pre servatio n equ al to that

, ,

of m oder n ooze -former s B ut i n l im e s to ne s of the

.

texture of the Carbo n iferou s Lime s to ne or chert s l ike ,

tho s e of the S cotti s h O rdovicia n a n d D evo n shire Culm ,

Pro tozo a are u s ually s ca rce a n d i n di s ti nct I t is n ote .

worthy that the few F orami n ifera k now n from Palaeozoic

rock s ar e o n the whole larger t ha n tho s e cha racteri s tic
, ,

o f more rece n t depo s it s ;while the s urvivi ng R adiolaria

in Cul m Mea s u re cher t s are ofte n di s ti ngui s hable with

-

the n aked eye Thi s s ugge st s that the s maller orga n ic

.

rem a i ns of a ncien t d epo s it s have di s app e ared duri ng

molecular readj u s tme nt a nd that Proto a n oozes m ay
, _,

actu ally have bee n as a bu n d ant in Pal aeoz oic t im e s a s

3 0 I N VE R TE B R ATE PALA EON T OL OGY

they are t o d ay Apart from the Chal k there are few

-
.
,

s uch rock s i n the Briti s h s eque n ce although S acca mmi n a ,

i n the Carbo n iferou s a n d N u mmuli na in the E oce n e

, ,

ofte n give character to the bed s i n which t hey occur .

The m aj ority of S iliceou s s po n ge s i n s pite of their ,

large s ize collap s e after death i n to a welter of s le nd er

,

s picule s who s e fate i s n orm al ly l ike t hat of R adiolaria

, .

S ome Lower Cretaceou s chert s co n tai n vi s ible evide n ce

of the Porifera n origi n of their s ilica while the flints i n ,

Ch alk m u s t have bee n derived l argely from d i s s olutio n

of s po ge s picu le s ( Pl ii fi g
n - C alcareou s s po n ges a re
. . .

m ore m a s s ive a nd d urable ;s ome ba n d s i n the I n ferior

O ol ite a n d the A ptia n of F ari n gdo n are l argely bu ilt
m
, ,

of their re ai ns .

The reef buildi ng capacities of Coral s are wel l k n ow n

-
,

a nd Madreporaria ( o r allied Coelen terate s of s im ilar

habit ) h ave prod uced lim est o n e s at s everal s tage s i n
B riti s h geological hi s tory I t m u s t however be .
, ,

re membered that a coral reef i s r arely built s olely of -

coral s while a l arge proportio n of tho s e a n i mal s are n ot

,

reef-m aker s The col o n ial mas s es a n d their brok e n

.
,

fragm e nt s co n s titute m u ch of a typical reef but

, ,

member s of other I n vertebrate s that s heltere d in the

wal l or becam e i n volved i n i ts rui n s form a co ns iderable
, ,

bul k Calcareou s algae have n o s mal l s hare i n reef

.

co n s tructio n The Chief l i mes to ne s i n Britai n that co n

.

tai n reef-buil d i n g coral s are the We n lock D evo nia n a nd ,

C a rbo n i ferou s Lime s to n e s ( Pl i fig 1 ) i n th e Palae ozoic . . .

,

a n d local bed s i n the Lower O olite s a n d Cor a llia n i n

the Me s ozoic .

A m o n g E chi n oderm a t a the Cri n oidea s ecrete the ,

mo s t m as s ive a nd exte ns ive s keleto ns ;group s prevale n t

i n Palaeozoic t im e s were e s pecially rem arkable for thi s
qua lity Part s O f the We n lock a nd Ludlow Lime s to nes
.
,
”
patches of D evo n ia n m arble s a nd im po rta nt m a ss es ,
 MATE R I ALS 3 1

of the Carbo n iferou s Lime s to ne ( Pl i fig are . . .

replete with Cri noid column a ls al m o s t to the excl u s io n

of other m aterial I n Me s ozoi c rock s rem ai ns of Cri n oid s
.

are ofte n abu nd a n t but they are rarely s o profu se a s

,

to m ake up m uch bul k of the strata in which they occur .

Thi n s heet s of s parry l ime s to ne i n the Lower Lia s ,

'

repre s e nti ng i nd ivid uals or clu s ter s o f P enta cr z nus a nd ,

bed s of i n th e Batho nia n ( Pl i fig a fford . . .

exceptio ns to thi s ge neral rule E chi noid s though ofte n .

,

com mo n fo s s il s n eve r attai n the ra n k of rock former s

,
-

i n thi s cou n try .

T he fi i msy colo n ie s of calcareou s Polyzoa are ofte n

‘

abu nda n t but it is rare to fi n d them i n the r ole of

,

rock -former s F enes tella s ometime s crowd s l ayers of

.

Carbo n iferou s a n d Permia n Lime s to n es ;while m ore

“
m a s s ive Coralli nes are importa n t co n s titue nt s of the
F ari ngdo n Gree n s a n d a nd Plioce ne Crag s .

B rach iopod a flouri s hed to s uch a degree i n the

s hallower Palaeozoic s ea s that they ra n k as the chief
”
s hell s of that epo ch I n co n s eque n ce the equ ivale nt s
.
,

of modern s hell -beaches were m ade of their remai ns .

V ariou s m ember s of the Or th a cea co nvert s ome l ayer s of

O rdovicia n s a nd s to n e i nto local lime s to n e s P enmmer us ,

doe s the s ame i n the Ll a n dovery L ep ta en a a nd A i m ,

/ m
p
s imilarly co n s titute part s of the We n lock while Con ,

cfi zd z u m i s largely re s po n s ible for the exi s te n ce of the

’ '

Ludlow Lime s to ne ( Pl ii fi g Ma ny m a s s e s of
. . .

D evo n ia n Lime s to ne are replete with Brachiopod s of

variou s ge n era while the broke n s hel l s a n d s pi n e s of
,

P r od uctus give character l ocally to the Carbo n i fe rou s

Lime s to ne I n later period s Brachiopod s were on the
.
,

whole l e s s n umerou s though certai n ge nera of Rh y n

, ,

ch o ne lla ce a a nd T e re b ra t u lace a give a di s ti nctive a

p
p e a r a nce t o s ome J ura s s ic Lime s to n e s a n d co ns titu t e no
,

s m a ll propor t io n of their bulk .

32 IN VE R TE B R AT E PALAEON T OL OG Y

A fter t he Palaeozoic epoch M o l l u sca as s u m e d the ,

m a nt le of the Brachiopoda I n p articular t he O s tr ei d ae

‘

,
.
,

which eve n to d ay occur i n co nges t e d clust ers rou n d

- r

our s hores are re s pons ibl e for m uch of the M iddle Lia s
,

of York s hire a nd the F ore s t Marble o f the S outher n

Cou nt ies A m mo n ite s are rarely rock former s bu t a t
.
-
,

s o m e horizo n s in the I n fe rior O ol ite o f D or s et s hire ( e g .

a t Bu rt o n Brad s tock a nd Bradford Abba s ) th ey are s o

abu nd a n t that the s ca n ty m atrix s carcely s ep ara tes the

s he ll s I n Tertiary tim e s Mollu s ca n life 1n the Briti s h
. ,

area was rarely of s uch exubera nce a s t o m a s k t he

,

char a cte r of the s ed ime nt s bu t the well -k nown Oyst e r

‘
’

bed at the ba s e of the R ead i n g s erie s t he ba seme nt

“
,

b e d of the Lo n don Cl ay l ayer s i n the B ra ck les h a m an d

,

Ba rto n s e rie s the O ligoce n e ( Pl ii fig

, a n d part s o f . . .

the Cr a gs owe m uch o f thei r bul k to the a bunda n t

,
_

Pelecypoda a n d G astropoda
‘

Trilobite s are ofte n di s s em i n a t ed ( in a m ore or les s

broke n s tate ) in great profu s io n i n Lower Pal aeozoic
rock s a nd eve n i n t he Carbo n iferou s Limes to n e but
, ,

they ca n hardly b e cl as s ed a s rock fo r mers The m o st .

e fficie nt Arthropo da i n this re s pect a re the Ost rac od a .

B ey r zc/zz a 1 n W ov i ci a n a nd S iluria n a n d C m
'
’
yp r za ,

i n the Wealde n ofte n occu r i n s warm s to the virtual

,

excl u s io n of m atrix ;bu t their s mall s ize a nd im per s is te nt

depo s itio n re s trict their rock form i n g capacit ie s to thi n
-

be d s E xam i atio of prep are

. n n d “
fl i nt meal
- s how s

a bunda nt O s tr ac od a in the Chal k ooze ;their remai ns

are u s ual ly s u bordi n a t e t o F or am i nifera a nd S po nge
s pi cule s .

( )
I I I F O S S I L -B A N D S ( PI vi ) . .

B rief ex pe r ie ce
n o f fo il hu n ti ng s u ffi ces to sh ow th a t
ss -

o rga n ic rem ai ns are ra rely s ca t tered eve nly t h ro ugh a

d epos it bu t t e n d to occ ur i n m o re or le s s d efi ned ba nd s
, .
34 IN VE RT E B R AT E PALA E ON T O LOGY

is po s s ible to d i crim i ate betwee f il ba nd s due to

s n n os s -

the m ode of life ( or ma n ner of death ) of the orga n i s m s

c o n cer n ed a n d tho s e produced by phy s iographical age n t s
,

work i n g o n d i s carded s hel l s A s a rule the form er .

,

type s co ns i s t o f well pre s erved s peci me n s of a s i ngle

-

s pecie s ,
while the others i n cl ude a variety o f m ore or
le s s d am aged fo s s il s .

The local a nd co n ge s ted habit s of life of m a ny

s ede n tary I n vertebrate s are illu s trated by the fam il i ar

m u s s el a n d oy s ter bed s o f Briti s h s hores D redgi n g in .

deeper water prove s that large area s of the s e a -floor

below low -tide m ark m ay be covered with P ecten or
E ck z nus to the virtual ex clu s io n of other type s More
’

over i t is by n o m ea ns u n com m o n to fi nd that the

‘

dredge bri n g s u p n othi n g o r dead s hell s o nly from a

, ,

place that yielded a rich harve s t of l ivi n g s pecime n s at

a previou s haul Tho s e I nve rtebrate s that lead m ore or
.

le ss s ede nta ry live s whe n ad ul t bu t have free -s wim m i ng ,

larval s tage s are apt to co n gregate i nto cl u ster s who s e

,

s ituatio n cha nge s peri od ically perh ap s with each ge n era,

tio n The evide n ce available i ndicate s that E chi n oderm s

.
,

Brachiopod s a n d Pel ecypod s ofte n s how the s e habit s ,

i n coa s tal or deep water I n a regio n where roc k

.

depo s itio n i s i n progre ss it i s ea s y to accou n t in thi s

,

way for i n cl u s io n of layer s compo s ed of on e or two

s pecie s of s hell s i n s e d im e n t which above a nd below , ,

co n tai n s fe w of t heir rem ai n s .

A gai n if s om e rapid or catacly s m ic cha n ge occur s in

,

the s urrou n di n gs of a fau n al as s em blage s udde n death ,

will overtake al l tho s e i n capable of s peedy e s ca p e S uch .

d i s a s ter s are not l ikely to com e freque n tly but they m ay ,

occa s io nally overwhel m i n habita n t s of coa s tal regi o ns .

S ud de n outpou ri n g of m ud from a river i n flood m ight

s mother orga n i s m s u n able to s wim a nd rapid s hifti n g ,

of a d s hoal s i n a s torm would have a like e ffect

s n -
.
 M AT E R I ALS 35

E arthquake s a nd
volca n ic eru ptio ns woul d hard ly work
s uch havoc upo n I n vertebrate s a s they oft e n prod uce

am o n g fi s he s ;but if they cau s ed u pheaval or other

geographi cal cha nge s their s udde n n ature would be
,

fatal to s ede n tary form s I t i s po s s ible that s ome of the

.

fo s s il ba nd s o f the uppermo s t S ilu ria n n otably the Bo n e

-
,

Bed a n d L z nguZ a -bed before me n tio n ed m ay be d u e to

’

sei s mi c cata s trophe s .

T he cleare s t a n d mos t freq ue n t type s of fo s s il ba n d s

“

are fou n d i n s trata of the l ittoral zo n e I n m os t of s uch .

ca se s the s he ll b ed s co ntai n a variety of form s although

-
,

o ne or two s pecie s u s ually pred omi n ate Al mo s t al l the .

fo s s il s are wor nor broke n a nd the m atrix m ay b e ful l,

of s hel l fragme n t s The s e fo s s il -ba n d s ca n be a s cribed

-
.

to tid al cur re n t s or waves s o rti n g o ut dead s he ll s of ,

s im ilar s hape a n d weight a nd d rifti n g them to the,

beach or sheltered water S hell -s a n d . is a com m o n

prod uct of thi s actio n O n s a ndy beache s it i s quite
.

u s u al to fi nd abu nd a nt s hell s after co nti n ued wi n d s from

a parti cu lar quarter while at the n ext v i s it n othi ng but
,

s an d a n d s hi n gle ca n b e s ee n I f i s o s tati c co n ditio n s

.

are s uch that perma ne n t s edi me n t atio n is i n progres s ,

s uch epi s ode s will be pe r petuated a s al ter n atio ns of fo s s ili

ferou s a nd barre n depo s its .

Las tly red u ctio n i n the rate of accum ulatio n of s edi

,

m e n t due to co nditio ns that do not s e r1o us ly a ffe ct the

,

local fau na n aturally i n crea s e s the relative n u mb e r of

,

foss il s i n a giv e n m a s s of rock D evelop me n t of orga n ic .

oozes o n the floor of t he o p e n s e a beyo n d detrital lim it s

'

, ,

gives a n extreme exa mple of thi s type of fo s s il ba n d -

.
”

Local ( u s ually temporary ) circum s ta n ce s m ay give co m

pa rable re s ult s 111 coas tal depos it s I llu s tratio n of s uch .

co nditio ns i s fou n d i n the I n ferior O olite I n D or s et .

s hire ,
where o wi ng to irregularity a nd hi ndra nc e i n
,

i s o s tatic depres s io n rock s o f t hi s s ys tem rarely exceed

,
3 6 I NVE R T E B R ATE PAL A E ONT OLOG Y

t we n ty feet i n th ick ne s s fo ss il s of m a ny ki nd s are so

,

crowded a s to be al m o s t i n co n tact I n Glouce s ters hire

.
,

where depre ss io n allowed accum ulatio n of s edimen t

r eachi n g t wo hu n dred feet i n thick n e s s fo s s il s are ,

ge nerally m ore widely s eparated There is n o rea s o n to

.

s uppo s e that the abu n da n ce of l ife wa s s eriou s ly di ff ere n t

i n the two d i s trict s at a n y tim e in the I n ferior O ol ite

s tage ; the app are n t d i ffere n ce i s d ue to the am ou n t s

o f s edim e n t i n which the fo s s il s are embedded The .

fo s s il iferou s ch a racter of the Chalk R ock a n d lower parts ,

o f the U ppe r Chalk i n t he S outh of E n gla n d is a

, ,

fu rther exam ple of thi s phe n ome no n .

N OD UL ES ( Pl ii fi g 2 ;Pl iv fig 1 )
( IV ) . . . . . .

Con cretio n a ry s tructu re s are characteri s tic of m a ny

s ed ime ntary r ock s e s pe c ially argillaceou s types
, Con .

cre t i o n s di ffer i n com po s itio n an d origi n bu t they agree ,

i n be i n g develope d after accu m ulatio n of the s edime n t .

C a lciu m ca rbo nate ferrou s car b on ate a nd opal i n e S ilica

,

a re freque n tly pre s e n t i n cl ay s a n d lim e s to n e s T hei r .

ready s olubility in s urface water e ns ures rem oval from

expo sed layer s while their relative i n s olubility i n n o n
,

a erated water en courage s precipitatio n at great er depths .

Pet rifactio n ofte n re s ult s from di ffere n t ial s olub il ity but ,

formatio n of co ncretio ns i s perhap s m o re u s ual I n .

s om e depo s it s s u ch a s t h e O ld R ed M arl s of H ereford

,

s hire ,the Lower Lia s s ic cl ay s of D or s et s hire or the ,

Upper Chal k calcareou s o r s iliceou s i ng redie n t s have

,
“
s egregat e d i n to appare n tly s edime n tary s heet s of

Cor ns to n e Cem e nt s to ne a nd F li n t I n other s s uch a s

,
-
.
,

the Coal M ea s ure s hale s th e Uppe r Lia s of York s hi re

-
,

( Pl . iv fi.g . or the Lo n do n Clay l ro n s to n e Ceme

,
n t ,

s to n e or Se pta r ia n n od ule s are developed Precipitatio n .

of di s s olved m i n eral s i s a s s i s ted by a n d us ual ly ce n tred

,

arou n d s om e pa rticle o f sol id m atter with s im il a r

,
 M A T E RI AL S 37

c ry s talli ne pro p ertie s T hu s the n ucleu s of a chalybite

.

or calcareou s n od ule i s ofte n a calcite s hell while that ,

o f a fl i nt -n o d u le i s u s u ally a siliceou s s po n ge Although .

the fo s s il cau s i ng the n odule m ay have bee n a m ere

fra gm e n t it i s u s ual to fi nd orga n ic rem ai ns i n concre
,

tio ns better pres erved tha n tho s e in the s urrou ndi n g

roc k The s upe rior hard nes s a n d im p erm eability of the
.

s ubc ry s tall i n e m aterial give pro t ectio n to i t s c o n te nt s .

Thu s i n the Lia s of Whitby Am mo n ite s i n n od ule s are

,

ofte n perfe ct i n form a nd detail ;while tho se i n the s h a le

are flatte n ed al mo s t beyo n d recogn itio n S imilarly .
,

S po nge s i n the C ha l k ( Pl ii fi g 2 ) are repr es e nted by

. . .

ob s cure im pres s io ns s ave for thos e e n ca s ed i n fli n t I n

, .

the latter ca se s picule s a nd ooze -form i n g orga ni s m s

, ,

m i gled with them c n be picked from the fli nt meal

n a ,
-

perfect a nd u ncha n ged .

M arca s ite a nd Pyrite s are ofte n depo s ited i n cry s tal

li ne m as se s arou nd fo s s il s bur ied i n clay Commo nly .

t he e n clo s ed orga n i s m s have bee n petrified by the

m i n eral prec ipitatio n co n ti n ui ng after replaceme nt was
,

c omplete . More rarely the origi n a l s hell is retai n ed

with a n i n cru s tatio n of I ro n Pyrites s howi ng a rough ‘

i ndicatio n of the form of the fo s s il ( Pl v fig S uch. . .

n od ules are n o t s ati s facto ry fro m the palaeo ntological

s ta n dpo i nt s i nce the hard n e s s o f the c ru s t

,
a nd i ts ,

l iability to d ecay with l iberatio n of s ul phuric acid m a ke ,

extractio n a nd pre s erva t io n of the co n te n t s di fficult or

impra cticable .

V
( ) D E R I V E D F O S S I L S

S edimenta ry rock s are built of the rui ns of othe r s .

The m aterial s e mployed m ay have formed p art of

igneou s m etamorphic or older se d im entary rocks T he
, .

rui ns of a s ed imentary r oc k are likely to i nclude s om e

of its e nclo s ed fo ss il s H e nce it follow s t hat fo s s il s
.
38 I NVE RT E B R AT E PALA E O NT OL OGY

repre se n ti n g a fau na l o ng p as t m ay be rei n terred am o ng

s h e lls o f a later period . I n the pal aeo n tological re cord ,
”
a s i n hu ma n ma n u s cript s ,
a n achro n i s m s may occur ;
bu t they ca n o nly be retro s pective i n cha racter a n d s o ,

would be m ore accurately termed Ca ta ch ro nis ms “

.
”

F o s s il s derived from d e n udatio n of older rock s are un

likely to be fou nd i n a n y bu t l ittoral depo s it s a nd eve n ,

there ca n be co n fide ntly a s cribed to local or1g1 n D uri n g .

degrad atio n of a s e a cl i ff m a ny fo s s il s will fall to the

-
,

beach but the s cour of wave s wilL d es t roy all that are
,

n o t s peedily buried i n s a n d O f t he i n nu merable fo s s il s

.

“
weathered out o n a la n d s urface few if a ny s u rvive
, , ,

the da n ger s of travel to a pl ace of perma ne n t s e d i me n

t a ti on. Glaciatio n m ay however tra n s port s hel l s un
, ,

harmed for great d i s ta n ce s a s i s s how n by the freque nt

,

occurre nce of Gryp /za ea a nd other m a ss ive fo s s il s i n the

Boulder Clay .

Co nglomerate s s uch a s the B udleigh S al terto n Pebble

,

bed m ay i n clude pebble s that are fo ss iliferou s but u nder

, ,

s u ch circu m s ta n ce s there is l ittl e ri s k of co nfu s io n .

H owever m a ny l ayer s i n the D or s et s hire I n ferior O olite

,

a n d E a s t A n glia n Pl ioce n e are rem a n i é beds m ad e u p

of fo s s il s s eparated from their m atrix A gai n the .

,

Lower G ree n s a n d of F ari ngdo n a n d Cam bridge s hire

co ntai ns ma ny Belem nite s a n d other durable fo s s il s that
are not A ptia n but K i mmeri d gia n The n orm ally
, .

“
broke n a nd water wor n n ature of s uch truly extra n eou s
-

fo s s il s i s n o t a reliable c riterio n of derived origi n ;wave

actio n ca n produce s uch co n ditio n s i n a few hour s .

O n ly by carefu l id e n tificatio n a n d com pa ri s o n ca n the

true fo s s il evi d e nce be sifted from the fal s e B ut de rived .

fo s s il s though de ceptive i n s ome re spect s are of great

, ,

geological i ntere s t s i nce th e y i ndicate the age of the

,

rock s locally ex po s ed to de n udatio n at the p eriod i n

who s e sed ime nt s they occur .
 L
P A TE I I I

Tn igom a w i th t h e li g men t pr eserv ed Corallian,

"

cla w lla t a
’

I a

W ym
. .

e ou th .

2 . Tr zgom a
‘
'

z
'

m u r va . I n te r na l mo uld . r
Po tla n d i a n , S wi nd on .

3 A ngeli na sedn ck z .
' ’

r
D is to ted b y clea va g e. T r e ma d oc s la t e s ,

W
.

M i nfford , N . a les.

FIG .
4 . Pli o ce n e , S uffolk . T h e ia ragoni te sh e ll i s

r
c u mb li ng .

5 . Ost r ea éa u lz s
’
’

. Pli ocen e , S uffolk . T he ca lci t e s h e ll r t

i s i n pe fec
p rsr e e va t ion .
 M AT E R I ALS

C
( ) T H E P R E S E R V AT I O N O F F O S S I L S

( )
I C O M PL ET E OR U NCH A NGE D P R E SE R V A T I O N
The s oft ti ss ue s of I n vertebrate s are O f s uch texture
that their pre s ervatio n eve n for s hort pe riod s after
,

death require s rapid a nd exceptio nal proce s s es S uch

, .

proce s s es are likely t o develop protect ive m atrices

which are them s elve s doomed to decay Wherea s .

pla nt s may grow i n s uch profu s io n that their m atted

remai ns exud e a nti s eptic fl uid s s u fficie n t to prolo ng
the exi s te n ce of cel l co nte n t s ( thereby givi n g op p or
-

t u n i ty for m ore perpetual pre s ervatio n) a n i mal s rarely ,

co n gregate either in l ife or death in m a s se s com parable

, ,

with peat E ve n i n coral reefs the polyp s are s eparated

.
-

by di s proportio nate qua ntitie s o f m i neral matter i nca p

able O f retardi ng the chem ical di s s ol utio n of it s builders .

“ ”
Fe w I n vertebrate s have tough s ki n s although the
,

l igame nt s of Pelecyp oda have m erit i n thi s re s pe ct

‘

S uch ligame n t s are occa s io n ally pre s erved ( with but

s light s hri n kage o r carbo n izatio n m ay be fou d
) a n d , n

i n ro ck s ys tem s a s far ba c k a s the J ura s s ic ( PI iii fig

-
. . .

They are m o s t O fte n met with i n clay s or roc k s with

m uch argillaceou s co n te nt a n d are perhap s lea s t rare in
,

the D ors et s hire Coral lia n a n d Lo ndo n Tert iarie s .

The m o s t perfect type of pre s ervatio n is fou nd i n

ma ss e s of am ber that e n clo s e s m al l i n s ect s The s ticky .

re s i n that m ay e n ta n gle a nd e n s hroud orga n i s m s ca n

d o s o o nly whe n fre s hly extruded a nd s o i s available ,

for a se ptic pre s ervatio n o f terre s trial or ae rial form s

’

alo n e
. S i nce amber i s bou n d to u n dergo grad ual
carbo n izatio n the beautiful a n d e ffective mode o f
,

\
fo s s ilizatio n that it le n d s to type s of I nvertebrate s
othe rwi s e rarely pres erved ca n be but temporary ;s uch
evi d e nce doe s not ex te n d back beyo n d the Tertiary s tage .
 40 IN VE R T E B R ATE PALA E ON T OLOGY

S pecim e ns of fo s s iliferou s amber probably d erived

,

from O lde r depo s it s are occa s io n ally fou nd i n the

,

E a s t A n glia n Plioce n e a n d Plei s toce ne .

The s hel ly s t r uctu re s of fo s s il I nvertebrate s are

com m o n ly fou n d i n a co nd itio n hardly di ffere n t from
tho s e of l ivi ng form s U s ually however com plete
.
, ,

r em oval of the e n ca s i n g a n d pe n etrati n g o rga n ic ti s s ue

“ ”
re n der s de s iccated s hell s very fr iable S i nce s olu .

tio n or replaceme n t are i nevitable for s hel l s expo s ed

to the actio n of wate r clay s or n od ul es s upply the o n ly
,

m atrice s i n which u n changed pres ervatio n ca n be lo n g

m ai ntai ned E ve n i n s u ch s urrou n d i n gs the n ature of
.

the s hel l -s ub s ta n ce i n flue n ce s the perma n e n ce of the

fos s il . Calcite m ay retai n i t s o ri gi n al ch a racter s s o
perfectly t hat it s pearly l u s tre ( a n d ev e n traces of
colo ratio n) m ay be fou nd i n depos it s a s ol d as the
Lia s E ve n i n the porou s s a n d s o f the Crag Os tr ea
.
,

pe r s i s t s Uncha nged But A rago n ite ca n rarely e s cape

.

r emoval o r al teratio n whateve r m ay be the character

,

o f i t s s urrou n di n g m atri x C om pari s o n O f fi gs 4 a nd

. .

5 ( Plate iii
.
) s how s the d i ffere n t co n ditio n o f s hell s

com po s ed of the two form s of C al c iu m carbo n ate i n

the s ame de p o s it S t riki ng ill u s tratio ns of the d i fferi n g
.

d urability o f C alcite a nd Arago n ite occu r i n t he C h al k ,

where la rge s hel l s built O f the latter ( e g A m m o n ite s . .

a n d G as tropod s ) are repre s e n ted by ca s t s a n d m ould s ,

t o the wall s O f which adhere e n cru s ti n g Polyzoa a nd

Sp ondylz which bei n g m ade o f Calcite wholly or i n
'

pa rt a re p re served al mo s t u n cha n ged

, .

O pa l i n e s il ica h a s con s iderable perm a n e n ce whe n

protected from percolati n g wate r but it u n dergoe s ,

d e vit ri fi ca t i on i n ti me o r u nder i n flue n ce O f pre s s ure

”
.
,

Chiti n m ay rem ai n ve ry fre s h if s ealed up in a nodule

or im pe rviou s cl ay ( Pl iv fi g. . The A rt hropod a
.

of the Co a l Me a s u res are but little ca rbo n ize d whe n

 L
P ATE IV
 L
P AT E I V

FIG . I . Or bzm lozd ea

' '

mtiaa
’
’
. Coa l M eas u res C lbroo kd ale
,
oa . Th e g lossy ,

la rg e ly ch i ti no u s, s h e ll is pe rfect ly pr ser ed in e v an i r
on s t on e

nod ule .

F IG . 2 . Fr gm a e n t of Pe le cyp od s h e ll r
f o m t he U pp r Gr e e e nsa n d , nea r
S id mo uth , r plac d fo mr
“

D e von , e e by ch a lced o ny in t he of

k
B e e i te

mr
.

lu lfl 3210 i f m
l
Fr
’
fl “1 I i
U
'
zr
W
v '
1 11
D af fy/t aver ns commu n e (“ p pe Lia s , . r , hitb p rk s . o m a

nod ule o f ce me n t s ton e Co mplet e . a n d un co mp es se d .

L
L a t era l! L11 1
‘

1 a! n il II" b ut! fl! J .

milar rs peci e sh r U ppe r

.
”

Per on orer a s a n n u la t u m a clos e ly s i i as ,

R uns wi ck Ba y, Yo k s r . F rom t he sh a le , fla tte ned and

t e le sco pe d .

FI G 5 Clz on a
'

s
p F li nt mou ld s of t h e c ypt sr p e rf r
o a ti n g t h e s h e ll of
U pp r Ch
. . .

l u acer a mu s . e a lk , S us se x .

F ro . 6 . A t ryp a
'

r et z cu la r z s
'

. L ud lo w Lm i e s ton e , W es t K no wle , C r a ve n

r
A ms , S a lop . I n te rn al mou ld ( v en tra l sid e ) , s h o wi n gm uscula r
a nd o t h e r i mpr e s s ion s .
FIG . 1 . cf.
g m
o al um . P
1 f
I nt érn al ’
mould ,

FIG
m
A mmo n it wholly
f: 711
e nc u t
1

r s
. 2 . e

13 4 1 Cla y, n ea r O xf rd
o .
 MATE R I A LS 41

e nca s ed in co ncretio n s a nd eve n Trilobite s a nd Gra pto

,

l ite s are o cca s io nally fou n d u naltered i n form a nd ,

hardly cha nged i n com p o s itio n R emark able s pecimen s

.

of E mypter us fou nd i n the S iluria n o f the I s le of Oe s el

( N orth R u s s i a) are s ometime s s o perfectly pre s erved

that the carapace s ca n be detached from the m a rl a nd

exami ned by tra ns m itted light Le ns e s of com po u nd
.

eye s O f Trilobites m ay al s o retai n their tra ns parency .

A m ode of pres e rvatio n i n which the origi n al ti ss ue

i s retai n ed i mpregn ated with fre s h mi neral -m a tter is
, ,

m ore u s ual tha n that i ndicated above a nd perhap s mo re ,

s ati s factory o n the whole . The m aj or ity of s hell s a nd ,

e s pe cially the s kelet ons o f E chi n o d erm s have a fairly ,

O p e n texture a n d precipitatio n O f n e w m i n eral de pos it s

,

i n the pore s s tre n gthe ns the fabric to a degree s ur p as s

i n g that attai ned duri n g life .Mi neral impreg n atio n i s
m o s t freque n t i n calcite fo ss il s embedded i n calcareou s
matrice s a nd i s s how n mo s t clea rly i n E chi nod erm
,

te s t s The crys talli ne co n ti n uity of the fres h m ate rial

.

with that O f the origi n al ti s s ue m ay lead to d is

adva n tage s by ob s c uri ng s tructural outl i ne s or i nduci ng
cleavage The former difficulty rs however us ual ly
.
, ,

m itigated by s l ight coloratio n o f th e s e co nd a ry in

'

fi lling while the latter may be hel pful i n the s tudy

,

of te s t s where each o s s icle i s i nd e p e nd e nti y or ie ntat e d .

O cca s io n ally the im preg nati n g m i n eral may b e di ffere n t

from that com po s i n g the tes t a s in fo s s il s that are
,

embedded m but not replaced by fli nt S uch s pecime ns

, ,
.

reveal m i n ute s tructu re with a d mirable d efi nitio n whe n

viewed i n thi n sectio ns .

The proc e ss of i n fi lling which i s s trictly com parabl e

,

with the ceme n tatio n of are naceou s or calcareous r oc k s ,

may at times b e carried to exces s so that i n cru s tatio ns

,

o f ne w c ry s talli n e mi n eral cover the s urface of the fos s i l .

Thi s c ondi t io n i s particularly com m o n i n the i nt e rio r of

4 2 I NVERT E B R A T E PAL A E ONTOL OG Y

s hell s or te s t s that were not filled by matrix s uch a s ,

m a ny Brachiopod s a n d I rregular E chi n oid s In the s e .

ca s e s the s parry m aterial u s ual ly cal cite but s ometim e s

,

quartz proj ect s i n to the cavity a nd m ay e ve n fi ll it I n

, .

T e r e b r a t ulid s from the F ari ngdo n Gree ns a n d s tage s ,

i n thi s i n cru s tatio n ca n be tra ced from a cry s talli n e ,

coati n g of the valve s a nd b ra chid ia to a com plete

i n ter n al m ould through who s e tra ns pare n t s ubs ta nce
,

“
the loop ca n b e s ee n Thi s s tyle of i n fi lling i s well
.

adapted for pre s ervatio n of the d elicate i nter n al a p

paratu s of the Telotrem ata ;a n d tha nk s to the method s ,

of preparatio n devi s ed by the R ev N orm a n Gla s s d oe s .

,

n ot preve n t i n ve s tigatio n o f i t s s tructu re . .

Le ss com mo nly s imilar i n cru s tatio n m ay develop

over the exter n al s urface of a fo s s il Thi s norm ally .

ob scure s the fi ner detail s O f or n ame n t ;b ut i n the c a s e ,

of E chi noid s it em p h a s iz es the plate s tr u cture by i t s

,
-

cry s talli n e rel atio n t o the os s icle s of the te s t I ncr us ta .

tio n i s n orm al l y pre s e n t o n the E chi n oid s of the Aptia n

of F ari n gdo n a n d s ometime s occur s o n tho s e O f the
,

C hal k I n fo s s il s from the former d epo s it a n outer

.

fi l m of iro n-ox id e coats the excre s ce nt calcite a n d both ,

l ayer s m ay ofte n be chipped o ff leavi ng the u n cha n ged ,

te s t ex po s ed .

( )
I I CA S T s A ND M O U L DS ( Pl iii fi g 2 ;Pl iv
. . . . . fi gs 5 .

a n d 6 ;Pl v fi g I
. . Pl vi )
. . .

I n p ermeable rock s it is u s ual to fi nd that fo s s il s

d i fferi n g from the m atrix i n com po s itio n have be e n
e n tirely rem oved i n s olutio n I f their rem oval h a s bee n
.

po s t p o n ed u n til the s urrou nd i n g rock attai n ed s om e

rigidity a n d if the text ure of the m atrix i s s u fficie n tly
,

fi ne the s car s repre s e n ti ng the m1s s in g fo s s il s will retai n

, ,

i n n ega tive all s uperficial detail s o f s tructure a n d or n a

,
“ ”
men t S u ch negative s are termed ca s t s whe n they
.
“
 L
P A TE V I
FI G r ba nd
’

. I . Pa t of a
~
foss i l-
Th e righ t h - a nd sid e was
h as r du d
e ce the o rg a n is ms mo uld s .
F a r th e r in
( to th e le ft of the fi g ur e ) (1 ,
i n ta ct .

a re t h e ch ie f fo rms pres n t e . r

FIG . 2 . r
Pa t of a fo ss i l b nd in C
- a a ra d oci fin ( A cton
‘
S co t t b ed s ) M a rsh
,

b rook , S a lo p . Co mple te ly d e ca lci fi ed . 071 71 111

”
ter t za i i fz a n
'

a
 44 IN VE RTE B R ATE PALAE ON TOL OGY

tem pora neou s m ould s of la rger s hell s The C ambrid ge .

Gree n a d
s n
“
Chloritic M arl
, a nd rock b a nd s of the
-
,
”

Chal k i n clude good exam ple s of thi s phe n ome no n ;

while the pyrit ous mo uld s of Lia ss ic A m m o n ites a nd
fli n t m ould s of Cretaceou s E chi n oid s a re s omewhat
a nalogou s .

( )
I I I P E T R I F A CT I O N

F i s s ures a nd other ope n s paces i n rock s te n d to b e

“ ”
come cl ogged by precipitatio n o f vei n -m i n eral s brought
i n s ol utio n by water The cavitie s s eparati n g ca s t s
.

from m ould s are thu s l iable to be fi lled by ne w m i neral

m a tte r
. Thi s will u s ually be di ffere n t from t hat
o r igi nally com po s i n g the fo s s il s a n d n orm al ly l es s
,

s ol uble. S i n ce the s eco n dary m i neral i s likely t o have

cry s tall i n e propertie s d i fferi n g from tho s e of the m i s s i n g
s h el l s ub s ta n ce it m ay fail to res trict i ts growth a b s o
-
,

lut e ly t o th e c a vi ty a nd will the n ob s cure s ome of the

,
‘

details of the fo ss il Thu s the petrifactio ns ( i n Beek ite

.

ch a l ced o ny ) of Mollu s ca i n the Uppe r G ree n sa n d s how

peculiar orbicular s urface s that we r e certai n ly n ot
pr e s e n t i n the origi n al s ( Pl iv fi g
. I t i s clear t h at
. .

the n atu re of the m i neral depo s ited i n the hollo w

depe n d s o n the quality of the a queou s s ol u tio n a vailable ,

a n d bea r s n o relatio n to the origi n al s ub s ta n ce .

The great m aj ority of vei n-mi ne ral s may occur a s

p s eudom o rphs after orga n i s m s but S ilica C a l cite a nd
, ,

Pyrite s are t ho s e u s ually em ployed S ilic a typical ly .

,

a s Ch al cedo n y ,
ofte n replace s cal cite or a ragon ite i n
ca lcareou s rock s co n tai n i n g opal i n e s il ic a i n the fo rm

of R ad iolaria or S po n ge s pic ules

- S uch pet rifactio n
.

i s ofte n mo s t sa ti s facto ry s i nce t he calcareou s m at r ix

,

ca n be r em o ved with a cid t he s ilic a rem ai ni ng un

,
 MATE R I ALS 45

t ouched I n favourable ca s es d e t ail s of form both

.
,

i nter n al a nd exter n al ca n b e expo s ed a s clearly a s if

,

the fos s il had n eve r bee n e n ca sed by rock E xqui s ite .

preparatio n s of Brach iopod b ra ch id ia ( Pl xi fig 9 ) . . .

a n d Blas toid h yd r o s p i re s ca n be m ad e i n thi s way ;

but ala s ! s il iceou s pe trifactio n of I n vertebrate s i s r arely

fou nd i n t hi s cou nt ry .

Calcite ofte n replace s Arago n ite i n lim e s to ne s a nd ,

.

m ay eve n form p s eudomorph s after opal i ne s po nge

s picul e s . I ro n Pyrite s a s M arca s ite or Pyrite i s the
, ,

u s ual petrifyi ng m i neral for argillaceou s rock s I t i s .

di ffi cul t i n thi s ca s e t o determ i n e whether the replace

me n t h a s bee n s ubs eque n t to rem oval of the origi nal
or by molecular replace m
,

e nt s i nce the opaci ty of the

,

m i neral hides any mi nute s tructure that m ay be p r e

s e rved. M arca s ite h as seriou s d i s adva n tage s as a
petrifyi n g s ubs ta nce s i n ce it is extremely u ns table
,

when expos ed a nd h a s a p ri s m atic habit that m ay

,

reveal it s el f i n n od ular outgrowth s i n de p e nde nt o f th e

‘

origi nal cavity ( Pl v . . Ma ny A m mo n ite s a nd

other ( u s ually Arago n ite) s hell s from the Lia s O xfordia n ,

a nd Gault are re re se n t e d b Marca s ite p s eud omorph s

‘

p y
that are ob s cure in them s elve s a nd di fficul t to prese rve
,

after extractio n Pyrite wh ich i s more characte ri s tic

.
,

of older argillaceou s rock s i s qu ite s ta ble but i t s p ro

, ,

n ounce d cubic cry st all izatio n m ay cau s e de s t ru ctio n

of s u rface details . H owever i n s ome i n s ta n ces it
, ,

ad m irable re placi n g mi neral s i nce i ts hard ,

ua lly greater tha n that of the matrix The .

S tarfi s h from Bu n de nbach which ca n be ,

their s laty coveri ng wi th a bra s s wire bru s h

-
,

m o s t s ati sfactory rOle a s a petrifyi ng

where the s ub s ta nce of the fo s s il s i s

material petrifactio ns m ay b e ,
46 IN VE R T E B R AT E PALA E ON T OL OG Y

m ade of a grea t variety of m i neral s S ilica ( O ften a s .

Q )uar t z a n d C alcite s till predom i n ate but metallic ,

m i neral s ma y O fte n form the repli ca s Mal achite .

,

Ble nd e H em atite Lim o n ite a n d Gale n a m ay be cited

, ,

a s m i n eral s occa s io n ally fou n d repl aci n g fo s s il s i n s a n d

s to n e s ( a nd eve n i n lim e s to ne s ;while Baryte s Cele s ti n e

) ,

a n d S ele n ite have bee n fou n d a s or ga n ic p s eu domorph s

”

i n greywacke s a n d clay s .

( )
b By molecula r p
re lacemen t

I t i s clear that the type of petrifa ctio n above de

s cribed ca n pre s erve n o v e s tige s of the m i n ute s tructure

of the fo s s il s repl a ced S ave for the pe culi ar character

.

O f the s hape of the cavity available the s eco ndary ,

m i neral fol low s i ts o wn cry s t all i n e m ode of accretio n

a s faith fully i n a n orga n ic ca s t a s i n a fi ss u re or geode .

But i n s om e ca s es d epo s itio n of the replaci ng m i nera l

takes place pa r i p a ss a w ith rem oval of the origi n al
'

U n der s uch circu m s ta nce s re placeme n t i s carried out

with great perfectio n s o that the fi n i s hed p s eudo m orph
,

po s s e ss es every detail of m icro s copic s tructu re of the

fo s s il S uch a ccurate repla cem e nt s are com m o n a mo n g
.

m i neral s ( e g Quartz after Crocidolite ) a nd in the

. .

s i li ci fi ca t i o n of wood y ti s s ue ;a n d they are n ot u n k n ow n

am o ng I nvertebrate fo s s il s S hell s a n d te s t s fro m th e

.

Up p e r Chalk m ay s ometi me s s how p atchy petrifactio n

by s ilica e pe ial ly whe n a s ociated with a
s
,
c s fli nt n od ule
-
,

but the repl acem e n t is rarely com plete .

( )
I V D I S T O R TI O N A ND D I S F I GUR E M EN T
( Pl iii fi g 3) . . .

Perfect s pecim e n s o f fo s s il s eve n of the com m o ne s t

,

k i n d s are relatively rare To the n u m e rou s a b nor ma li

,
.

tie s d i s eas e s a nd accid e n t s that m ay m ar a n orga ni s m

, ;
 M AT E R I AL S 47

du ri ng l ife or cau s e its death have to be ad ded in cal

, ,

l
cu a b e l ,
but alway s s eriou s ri s k
, s o f p os t -mor t e m dam age .

The m aj ority of s hell s are br oke n before they reach

the s a n ctu ary of a s ed ime ntary gr ave The s cou r O f .

wave s or i ns id iou s atta ck s O f the weather a nd the ,

u n sympathetic i nve s tigatio n s of carrio n -feeders will ,

pa ss over few of the orga n ic remai n s expo s ed to them .

S ede n tary a n im al s fi nd con ge n ial fou n datio n s o n em pty

s hell s while tho s e of retiri n g habit s welcome the s olubl e
,

fabric for excavatio n o f their crypt s .

F o s s il s embedded i n s a nd s to n es or li me s to ne s are
liable to be cru shed u nle s s their cavitie s are fil led with
s ed i me n t o r co n cretio n ary matter Brachiopod s a nd
.

I rregular E chi noid s are e s pecial ly a ffected i n thi s way ,

owi ng to the s m all s ize O f their foram i n a a nd co ns eque n t

im perfe c t i n filtratio n of s a nd or ooze But i n ge n eral .

fo ss il s i n s uch m atrice s do n ot s u ffer mecha nically

after e n tombme n t I n argillaceou s rock s o n the other
.
,

ha n d no hollow s hel l is s afe ( however well fi lled with

,

clay) u n le s s a co ncretio n h as developed withi n it The .

pre s s ure that co nve rt s clay to s hale work s equally on

the m aterial s i ns ide a nd out s ide the fo s s il s s o that the ,

latter are i nevitably flatte ned The m ore s evere s tres s es

.

a n d s trai ns that prod uce s laty cleava ge will i n volve the

fo s s il -co n te nt s i n s heari ng ;s o that the al ready flatte ned

s hell s take o n l ateral d i s tortio n At s om e horizo ns i n
.

the L i ngwla -F lags L z ngzd ella a a w s z m ay be collecte d

' '
’
'

i n great abu nda n ce but hard ly two s pecime n s out of

,

hu n dred s will have the s ame s hape E ve n whe n .

cleavage i s i n cipie nt fo ss il s are d i s torted ( s ee Pl iii

, . .

fig . while d evelopm e nt of com plete s lati n e s s u s ually

re n der s orga nic tra ce s practical ly u n recogn izable .

I n s lowly formi n g s ed im e n t s empty s hel l s become

,

e n cru s ted by orga n i s m s of s e s s ile habit F reque n tly .

Hy d ro a Polyzoa a nd Worm s grow on the s urface s

,
4 8 IN VE R T E B R ATE PALAE ON T OL OG Y

of li n i n g s hel l s i n s ym bi ot ic relatio ns hip with t hei r

“

ow ners but after death s i m ilar a n i m al s m ay m as k the

,

s upe rficial feature s exter n al a n d i n ter n al a nd pe netrate

, ,

the s ub s ta nce ; j u s t a s Bar n a cl es e ncru s t a n d s hip ,

worm s perforate a woode n hull Quite l o ng li s t s of

,
“
.

”
fo s s il s m ay s om etim e s be com piled by e n umeratio n
of the s pecie s o f H yd rozoa M ad repora ri a Polyzoa, , ,

Brachiopod a S erpul id s a nd Pelecypod a a dhe ri n g t o

,

Chal k E chi n oid s while the te s t it s el f m ay b e s o

,
.

s m othered u n der the s e growth s tha t eve n ge neric deter

mi natio n i s doubtful .

( )
V I M PR ES S I ON S A ND T R A CK S
The bod ie s of d ead a n im al s lyi n g on s oft s ilt or tho s e ,

of l ivi n g o n e s m ovi n g acro s s it l eave i mpre s s io n s m o re

,

or l es s corre s po n di n g with their s hape a nd s tru ct u re .

Thu s the s urface o f a m od er n bea ch i s b roke n by worm

cas ti ng s a n d track s burrow s of H ea rt Urchi n s R azor
, ,

s he ll s a n d [ -fle a s foot pri n t s of Crab s an d s tra nd ed

S a n d , ,

form s o f com ato s e j elly-fi s h I n m o s t cas e s the s e traces

.

of a bu n da n t life are obliterated at th e n ext tide ;but

s i n ce l ike ripple -m ark s they exte n d i nto s h a llow water

, ,

below tid al lim it s t hey m ay be co vere d by ne w l ayers

,

of s a nd o r clay without m uch d i s turba n ce i f the weather

i s Cal m a n d d epo s itio n i s i n progre s s J us t as ripple.

m ark s a n d other i n orga n ic irregularities are com m o n ly

, ,

fou n d i n littoral depo sit s of all age s so orga n ic ve s ti ges

,

a re Ofte n perpetuated The perfectio n o f s uch m a rk i n gs

.

d epe n d s l argely on the textu re of the s ilt while thei r ,

i n terpre ta tio n is bou n d to be u n c ertai n if their maker s

be lo n ged to exti n ct group s The s i n uou s gr oo ve s a n d
.

coiled ca s ts of lug worm s a nd the bl u rr ed track s of

“ -
,

N ereids look much the s am e whether m ade i n Lower

,

Pal a eozoic times or y es terd ay The co n fuse d but re .

,
 MA T E R I AL S 49

s olvable im pre s s io ns of collap sed j elly -fi s h or partly

,

decayed H olothuri a ns are n o ne the le s s di s ti nctive ,

whether the period of their prod uctio n wa s H oloce n e ,

J ura s s ic or C a mbri a n . A lthough s uch trace s rarely

provide O pportu n ity for appreciatio n of the a n atom ical
qual ities of the a nimal s that m ade them they at lea s t,

prov e the exi s te nce of variou s type s ( ofte n i ncapable of

d irect fo s s ilizatio n) i n rem ote period s I t i s i ntere s ti n g
.

to fi nd that worm -track s occur well d ow n i n the Pre

Cambria n o f the L o n gmynd a n d el s ewhere i n s late s ,

that co ntai n no other recog nizable fo s s il s Thes e rough

.

m arki n g s thu s carry back i ndicatio n s of life ( a n d that

of a fairly high order ) far beyo n d the l im it s at which

ordi na ry palaeo ntological evide n ce fail s F i n e Cambria n

.

s ilt i n Briti s h Columbia h a s yielded wo n derfully c lear

i m pre s s io n s of Medu s ae a nd H olothuria n s while the ,

U pper J ura ss ic Lithographic s to ne of S ole nho fe n i n clud e s

s im ila r relic s

Clos ely aki n to i m

.

pre ss io ns of pas t orga ni s m s are the

tubular aggregate s of fi s h s cale s fou n d i n the Chal k a nd
-
,

a scribed to the activitie s of m ari ne A nnelid s Thes e

.

“ ”
protective tube s re call the familiar cad di s ca s es of -

m oder n river s T hey a fford s om e evide n ce a s to the

.

fi s he s of the period a nd s ugge s t the exi s te n ce o f worm s

,

but they are hardly s ati s factory a s fo ss il s of either group

of orga ni s m s.
 C H APTE R III

T E C H NI Q U E

( )
A C O LL E CT I N G
1
( ) P UR POS E IN V I EW

LT H OUGH the im perfectio n of the p a la e o n t o

logical record is s uch that n o fragm e n t of fo s s il
eviden ce s hould b e l ightly throw n a s ide collectio n
,

of eve ry S pe cime n s ee n i s u s ually i m po s s ibl e . F ew a s

t he fo s sil s are i n com pari so n with the orga n i s m s the y
repre se n t they are O fte n exceedi ngly abu nda n t T o
,
.

c o llect al l acce ss ible fos s il s i n so m e dep os it s woul d

i n vol ve removal a nd s torage of whole s trat a I t is .

n e ce s s ary ,
therefore to pick a n d choo se amo ng the
,

l avi s h d i s play of O rgan ic rem ai ns s om etimes s pread

before the collector . U nle s s h i s pu rpo s e i s the id le
“ ”
o n e of picki n g u p pretty a n d s triki n g fo s s il s for u s e
a s or n ame nt s or curio s ( i n which ca s e h i s n eed s will n ot .

be co n s idered here ) the average fos s il -h u nter will have

,

s peci al i n tere s t i n s ome a s pect of Palaeo n tology a nd ,

will dire c t h is col lecti n g toward s the s ol utio n of par

t i cu la r problem s . I f hi s aim i s s t ratigra phical he will ,

s e c ure s pecime n s of types k n ow n to have re li able

“ ”
dati n g qu al itie s ; if it i s palaeo n tological he will ,

eagerly c ol lect fo s s il s of h i s cho s e n group pa ss i n g by

,

others that m ight appeal equ al ly to a colleague I n .

”
either ca s e it i s good for collectors to hu n t i n pack s
,
“
.

I t is u nlikely that a n i nd ivi d ual will s ee all the fos s il s

so
52 I NVE R TE B R AT E PA LA E ON T OL OGY

m ake m is take s The se ca n be rectified if the s pe cime ns

.

are available but m u s t alway s rem ai n as s ou rce s of

,

further error a nd co n fu s io n if t he pri n ted n am e h a s no

s olid backi n g .

( )
I I R EGA R D F OR S T R A T I GR A PH I CA L P OS I T I O N
The p reci s e horizo n from which a fo s s il wa s col lected
i s o ne o f i t s m o s t val uable qualitie s whether it i s de s ti ned ,

for geological or biological treatme n t Thi s po i nt ca n .

ea s ily be determ i ned at the tim e of collectio n ( u n les s the

s pecime n lay o n a s poil heap whe n due n ote of the fact ,

s hould be take n a n d attem pt s made on the s pot to trace

,

it s actual prove n a n ce) but it ca n n ot be s ub s eque n tly ,

ded uced with certai nty Before u n dertaki n g a n exped i .

tio n it i s well to s tudy a vailable accou n t s of the di s trict

,

a n d s trata to be vi s ited s o that d eterm i n ed s t ra ti gra p h i

,

cal divi s io ns ca n be recog nized But i n a ny ca s e the .

,

fi r s t duty for a Palaeo ntologi s t o n his a rrival at a n ,

expo s ure i s to n ote a n d if po ss ible t o m ea s ure the

, , ,

more s triki n g feature s of l ithol ogy a nd beddi n g s how n

i n the rock s N ot u ntil thi s i s do n e s hould a ny collect
.

i n g begi n S i n ce the fo ss il -evide n ce m ay O fte n s upply

‘

ot h e r a nd
,
l
better i n dica t i o ns of zo n al horizo ns tha n
,

m i n eral qu alitie s the fi rs t s ketch of the ex po s ed rock s

,

m ay n eed revi s io n I t i s u s ually be s t t o fi x on s om e

.

prom i ne nt a n d per i te n t featu re ( s uch a s a hale ba n d

s s s -

in lime s to n e a calcareou
,
s l aye r i n s hale or a fli n t -
bed ,

i n chal k ) a s a d atu m from which vertical mea s u reme n t s

'

ca n be made ;the d i s ta n ce above or below s uch a l i ne

at which a ny fo s s il or fo s il ba n d occur s s hou ld where

s -
,

po s s ible be recorded I t i s n ot e nough to gather all the

, .

fo s s il s from o ne qua rry i n to a s eparate group n or eve n ,

t o d i s ti ngui s h betwee n t ho s e from uppe r a nd lowe r pa rt s

o f th e s ectio n T oo much p reci s io n i n t hi s m a tt er is
.
 T EC H NI Q U E 53
‘

i m po s s ible ;too littl e will s oo ner or later lead to vai n

regret s S tratigraphical d ivi s io ns that seem adequate at
.

o ne time may prove far too v a gue i n the light of fur ther

re s earch - zo n al s tages u nd reamed of by o ne ge neratio n

becom e the domi n a n t feature s of s tratigr a phy for the
n ext .

I t m u s t be ad m itted that m eticulou s care i n fi x i ng

'
‘

the horizo n s from which fo s s il s have come i s tediou s ,

a n d dema n d s patie n ce a n d s te m s elf-co n trol o n the

“
part of a collector eager to try h i s l uck But .

without s uch preci s io n fo s s il col lecti ng cea s e s to have

,
-

a truly scie ntific value dege nerati ng i n to a n idle al m o s t

, ,

vi ciou s hobby N o am ou nt of s ub s eque nt repe nta n ce

,
.

ca n r e -e n dow a fo s s il with the val ue that h a s bee n

throw n away by carele s s or ca s u al collecti ng .

( )
I I I L A R G E A ND M I N U T E F OS S I L S
In ge neral the tool s required by a fi e ld -p a la e on
t olo gis t are e s s e n tially s i milar to tho s e n ece s s ary for
an geological work A light pick axe ham m r s a
y .
e , ,

cold chi s el a pair o f pl iers a bricklayers trowel and

-
, ,
’

ample s upplie s of s m al l boxe s a n d O ld n ew s paper s co n

s t i t ut e the apparat u s m o s t ofte n u s eful I t is alway s .

s ati s facto ry to have at lea s t two ham me rs both with

(
s quare head s a n d chi s el edge s weighi n g about
) o n e ,

a pou nd for quarryi ng o r hitti n g the chi s el a nd o ne ,

quite s mall for tri m mi n g s pecime ns I t is of cour s e . , ,

n eedle s s to tak e all type s of apparat us o n m o s t expedi

tio ns ;the s electio n will be determ i n ed by the n atu re

of the rock s likely to be e ncou ntered H am mers a nd .

Chi s el s are of little u s e o n s oft cl ay though the former

(
s hould alway s be take n I n V 1e w of po s s ible nod ule s ) ,

while a trowel i s i neffective o n hard rock s .

Whe n the obj ect s of search are ma cros co p ic fo s s i ls / ,

54 I NVE R T E B RAT E PALA E ON TOLOGY

fi r s t at t e ntio n s houl d be paid to we a thered rock -s u rfaces

t hat a re u s ually to be fou n d o n n atural o r art ificial
'

s ectio ns . N ot u ntil thi s exa m i n atio n i s com plete is

actual quar ryi ng advi s able I n m os t ca s es if fo s s il s
.
,

do not appear o n decayi n g s urface s they are le s s l ikely

,

to be s ee n in n ewly fractured m atrix I f a portio n o f .

a fos s il i s s ee n protrudi ng from hard roc k it s s urrou n d ,

i ngs s hou ld be carefully s tud ied with a view to fi ndi ng

crack s or other li ne s of weak ne s s alo n g which fracture s
ca n be m ade I n al l ca s e s where ham meri n g i s requ ired
.
,

care s hould be take n to let the blows fal l a s far from

the s pecime n a s i s co ns i s te n t w ith i ts ultim ate extractio n .

N ever attem pt to break o ut th e fo s s il bu t aim to collect

,

the pi e ce of rock i n which it i s em be dded W he n .

po s s ible it i s a valuable sa feguard to keep o ne ha n d

,

ove r the s pecime n while u s i n g the ham me r s i nce it m ay ,

fly off u nexpected ly a nd be lo s t to s ight .

I t i s ofte n ea s ier to pick up fo s s il s from the talu s

o r quarry -heap s tha n t o d ig them out from a rock -face ,

a n d s till le s s exhau s ti n g t o purcha s e them from the

quarrym en .S uch aid s t o col lecti ng s hould n o t be

ig nored ;bu t it i s e s se ntial to remember that s pecim e ns
thu s acqu ired have not the s am e certai nty of s trati
gr a phical ho rizo n H owever i t i s ofte n po s s ibl e to
.
,

t ra ce the bed from whic h s uch fo s s il s have come u s ual ly ,

by c om pari s o n of thei r m a t rix with that o f t he m ai n

s ectio n.

I f m i n ute fo ss il s are required ( eg F oram i nifera .

,

Os tra coda or s m all s hell s ) collectio n of i n di vidual

, ,

s pecime n s i s clearly i n fe a si ble . A ny friable or rotted

l ayer m ay co n tai n s uch orga n i s m s a nd it is bes t to
,

t ra ns fe r the l oo s e m aterial t o ti ns i n bul k taki n g the ,

r i s k as t o whether later exam i n atio n will prove it p ro

li fi c or ba rren I n s om e c a s e s it i s po s s ib le to gai n
. .

s ome i nd icatio n of the quality o f s uch m aterial be fore

 T EC H NI Q UE 55

a dd i ng it to the d ay s load A s mall qua ntity ma y b e

’
.

put i nto a gauze bag (s a y a s mall b ut t e r fly-ne t) O r a

,

fi ne s ieve . I f the rubble is d ry a S hort S pe ll of vib r a ,

tio n will drive the fi n e s t p owder through th e me s h ,

an d the gritty re s idue ca n be i nspected with a ha n d

le n s I f it is wet the s am e re s ult m ay be O btai ned by

.
,

vigorou sly s haki n g the co ntai n er u n der water I t i s .

u s ual ly po s s ible to detect s m all fo ss il s i n the re s id ue ,

a n d to form a n e s tim ate of thei r rel ative abu n d a nce .

But failure to fi n d them by thi s rough method i s no

proof of their total ab s e nce ;it S hould m e rely i n flue nce
the collector i n determi ni ng the bu l k of material worth
carryi ng S tude nt s o f Microzoa are well advi sed to
.

bri n g a way s am ple s of all s edi me n tary rock s expo sed

i n each s ectio n Ma ny com p act m a s s e s may be res olved
.

i n to powder b y s uitable m ea n s while the other s ca n ,

be S liced for m icro scopic exam i n atio n E xperie n ce .

will s oo n s how th e type s of m atrix mo s t l ikely to yiel d

s mall fo s s il s .

( V)
I F I E L D P R EPA R A T ION
The l im itatio ns of h uma n s tre n gth dem a n d th a t t he
“ ”
item s of a day s bag s hall have the s m alles t s i z e
’

c o mpatib le with their s afety H e nce whe n a fos s il is .

,

fi rm ly em bedded i n m atrix it is expedie n t to reduce ,

t he bulk O f the bloc k B u t it i s n ever s afe to aim at

.

com plete extractio n of the fo s s il at thi s s tage N ot .

o n ly doe s adhere nt matrix act a s a protectio n to the

s pecime n dur i n g carriage bu t it ofte n a ff ord s u s eful
,

proof of the layer from whi ch it wa s collected Whe n .

po ss ible use of a ham m er i n the rough trim m i n g of

,

s pecime n s s houl d be avoided The vibratio n cau s ed .

by blows often s hatters the fo s s il eve n though care is ,

ta ke n to s tr ike di s ta n t p a rts o f t he blo ck ; while t he

’

u ncertai nty of the d irectio n of fracture i nvolves ris ks

56 IN VE R TE B R AT E PALA E ON T OLO GY

that s hould n o t be l ightly tak e n A pair of blu nt .

pl ier s are the s afe s t m ea ns for red uci n g the m atrix ,

although they are o n ly effective on thi n ed ge s If .

ham meri n g i s n ece s s ary it s hould be applied l ightly

, .

A s m al l ham mer with which a rapid s ucce ss io n of

,

tap s is give n prove s le s s da n gerou s tha n a heavy o ne

, .

The block s hould whe n po s s ible be hel d i n the hand

, ,

d uri n g trim m i n g s o that vibratio n s m ay be ab s orbed

,
.

I f it is too large to hold a tuft of gra ss or m o s s give s

,

a s afer fou n d atio n tha n hard roc k N ever t ry to chip .

o ff large piece s at a tim e .

I f a crack appear s i n the fo s s il in s pite of precautio ns

, ,

three cour s e s are available U n les s t he block i s s till

.

too large to carry s top I t m ay be po s s ible to conti nue

, .

i t s re d uctio n s afely at home or i n the laborato ry If .

the m a s s i s large it is be s t to give a s ha rp blow t hat

,

s plit s the s pecime n i n hal f E ach s egm e n t ca n the n

.

be tri m med s eparately ( care bei n g take n to avoid

alteratio n of the accide n tal fracture) a nd gl ue will ,

s ub s eque n tly m i ni m ize th e e ffect s of the di s a s ter .

T hirdly throw away the s pecime n a n d l ook for a n other

, .

Thi s co n fe ss io n o f fail ure s hould be m ade o n ly if the

s pecime n h a s crum bled or if there are ple n ty more of
,

the s am e k i nd i n the sa me layer S O l o ng a s a fo ss il .

i s recog n izable it retai ns s om e value eve n i n rui n

, .

( )
V F I EL D L A B E LL I N G A ND PA CK I N G
I t c a nn ot be too s tro n gly urged that a fo il col lector ss -
,

i n com m o n with other s cie n tific worker s s hou l d proceed ,

at every s tage a s though the s word of D amocle s were

s u s pe n ded above him Thi s d oe s n ot imply that the re
.

i s a nythi n g i n tri ns ically d a n gerou s or fatal i n the work .

But eve n though bodily di s s ol utio n m ay be far d i s ta nt ,

m e n tal fail ure i s a n ever -pre s e nt da n ger N ever t r u s t .

 T E C H NI Q U E 57

to m emory I m pre s s ion s gai n ed while collecti n g a re

.

v ivid e n ough at the t ime but they rarely l a s t u n til the ,

”
fi nd s have bee n s o rted a nd labelled at home I f a .

s ucce s s io n o f expo s u re s i s vi s ited i n o ne day detail s ,

of the variou s experie nce s become co nfu s ed F urther .

,

eve n though mem ory m ay s u ffi c e to recal l each epi s ode

for a s hort time the u ncertai n ty of hu ma n a ffairs may
,

dem a n d a il ong i nterval betwee n collectio n a nd exam i n a

tio n o f m aterial The d ay s bag m ay not be u n packed
.
’

for s everal week s a n d the ha n d that u nload s it m ay

,

n ot be the o ne that filled i t U n le s s the d etail s n oted .

d u ri n g the exped itio n are e n clo s ed With the s pecime ns

i n wr i ti ng m uch e s s e n tial i n form atio n will be per
,

ma ne nt ly lo s t I t i s no t e nough to e n ter detail s i n a

"

n otebo ok although s uch record s s hould alway s be

,

made A l a bel m u s t be e nclo sed with a nd i n deed

.
,

fa s te ned to e a ch s peci me n before the quarry i s left

A s upply o f s tamp -edgi n g o r other gu m m
.
,

ed paper , ,

d oe s n ot add m aterially to the weight o f apparatu s

carried ; bu t it m ay prove the mo s t valuable item

The label for each fo s s il s houl d r e cord two fact s

fir s t the preci s e horizo n with refere nce to a fixed a n d
, ,

p er m a ne n t feature o f the expo s ure ; a n d s e c o n d the ,

locality The latter s hould be i ndicated i n s uch a way

.

that a nyo n e with the hel p of a ma p could locate the

, ,

s ectio n whether o n the s pot o r i n the s tudy La b el s

, .

are be s t writte n i n i n k or i n del ible pe n cil ;time or a ,

dre n chi ng shower m ay oblit e rate word s writte n i n ,

bl a cklead .

F or pur p o se s o f tra ns po rt e ve ry s pe cime n m u s t be ,

wrap p ed in p aper or s im ilar mate rial I f se veral fo ss il s .

a re e nclo s ed i n o ne s heet car e m u s t b e t ake n that at ,

lea s t o ne thick ne s s of paper s eparates each from i ts ‘

n eighbo ur Without thi s precautio n, s om e or all of

.
58 I N VE RT E B R ATE PALAEON T OL OGY

the s peci me ns will be rui ne d by attritio n S m al l fo ss il s .

are be s t packed i n boxe s or ti n s s u rplu s s pace bei n g,

filled with m o s s or gra s s E ve n s o they m u s t be

.

wrapped i n paper le s t gr e e n s tai ns s oak i nto a nd

,

d i s fi gure them A n excelle n t pl a n for s afe rem oval of

.

friabl e s hell s extracted from clay i s to ge ntly bury them

i n s om e of their o wn m atrix e n clo s ed i n a t in They .

wil l be s ecure agai ns t ord i n ary vibratio n a nd ca n be ,

wa s hed o ut qu ite s afely i f u n pack i ng is no t u n duly

~

dela yed .

( )
B P R E PA R AT I O N

( )
I N E E D F OR P A T I E NCE
E xtra c tio n of fo s s il s from m atrix a n d fur the r ,

proce s s e s preparatory to s tudy m ay be com pared with

,

di ss ectio n of biol ogical material I n both ca s es practice

.

a fford s the o n ly mea ns of acquiri n g s kill a nd early ,

attem pt s m u s t ofte n lead to the s acrifice of cheri shed

s pecime n s . I t i s erro neou s to i m agi ne that clea n i ng
fo s s il s i s a rough a nd -ready bu s i ne s s ;the d elicacy of
-
‘

t ouch a n d d el iberatio n required are i n n o way i n fe r 1or

to tho s e n ece s s ary for a n atom ical work I ndeed i n .
,

s om e re sp ect s the greater re s trai n t i s im po s ed upo n

the Pal aeo n tologi s t A n a natom i s t u s ually h a s reliable

.

k nowledge of the res i s ta nce that ti s s ue s will o ffe r to h is

s calpel ;but n o co ns ta n cy of texture ca n be a s s u med

fo r a m atrix . F reque nt need for applicatio n of con

s id e r a b le force dem a n d s co n s ta n t watchful n e s s le s t th e ,

tool s hould s l ip i nto s oft material a n d reach the

s pecime n .

O n e adva n tage over the Zoologi s t lie s i n the d ura

b i lity of p a la e o n tolog ical m aterial There is no need t o
.

h as t en di ss ectio n l es t the O bject s hould decay ;i n m a ny

ca se s the refractory na t ure of m atrix i s mollifi ed by
60 I NVER T E B R AT E PA L A E ONT OLOG Y

a ccou nt s of m a ny proce s s e s are publi s hed i n gu ide s

of the B riti s h Mu s eu m a n d S m ith s o nia n I ns titutio n a n d ,

i n a paper by Dr F A B ath er i n vol viii o f the

. . . . .

M u s eu m: j ou r na l I n a ge n eral i n trod uctio n to

the tech n ique of fo s s il -preparatio n it i s m ore appropriate
—

t o co n vey a s erie s of hi n t s tha n to de s cribe elaborate

s chem e s rarely adequate fo r the full s olutio n of particu lar

problem s .

The rem oval of m atrix a nd the m etho d s em ployed

,

therefor wil l depe n d chiefly on two co nd itio ns M atrice s

, .

di ffer i n texture from s oft clay to c o n cretio nary m atter

with the hard ne s s of s teel ; while variatio n i n the
d u rabil ity of fo s s il s i s hardly le s s wid e I n the relatio n .

betwee n the s e t wo qual itie s lie s the problem b e fore the

worker who s e a i m i s to des troy the o ne a nd pre s erve
, , ,

or eve n rei n force the other F o s s il s are u s ually either

, .

harder or s ofter tha n their co ntai n i n g m atrix Tho s e i n .

s oft r ock s a re u s ual ly friable but the co nver s e is not

,

al way s true .

I f the fo s s il is ha rder tha n the m atrix ( whether by

origi n al character or petrifactio n) , a n d is tolerably
coh ere n t a bru s h is the s afe s t ki n d of tool F or calcite
, .

s hell s i n loo s e lime s to n e or Cl ay n o more elaborate ,

apparatu s tha n a n ol d tooth -bru s h is required The s e .

m ay be p rocured i n varyi n g degree s o f s ti ffn e s s or can ,

be graded by cutti ng the bri s tle s R otary bru s hes m ay .

be u s ed I n a de ntal e n gi n e but ha nd work is le s s likely

,
-

to carry the proce s s too far I n m atrice s o f the n ature

.

of c halk bru s hi n g is be s t carried o ut i n the d ry ;though

,

care s houl d be take n to avoid i n hali ng the d u s t which ,

ma y cau se a k i n d of I n clays if the ,

fo s s il s are cohere n t the bru s h ma y b e m oi s te ned but

, ,

the s pecime n s hou ld n ever be s atura t ed I t ca n be .

occa s io n ally cleared of m ud by ri n s i n g for a few s eco nd s

u n der a ge n tle s tream F o s s il s t o o s m al l to be ha n dled
.
 TE CH NI Q UE 61

i ndividually ca n be s orted a nd ofte n s u fficie n tly clea n ed

, ,

by placi n g them at the top of a pile of s ieves ( be com i n g '

fi ner d ow n ward s ) a n d directi n g a s teady j et of wate r

,

through the whole .

I t ofte n happe n s that s hell s i n clay are so friable that

they ca n n ot with s ta nd bru s hi n g however ge ntle I n , .

s uch ca s e s ,both fos s il a n d m atrix m u s t be rei nforced

with glue o r S hellac The whole s pe cime n S hould be
.
'

s oaked i n dilute s olutio ns of o ne or the other reage n t ,

allowed to s e t hard The m atrix ca n the n be

'

a nd .

pai nted local ly with water or alcohol a nd gradually ,

bru s hed away .

Whe n the m atrix is too hard to yield to a bru s h ,

it may either be s oftened chem ically or chipped O ff

mecha n ically Both method s are fraught with da n ger
.
,

s i n ce i n ma n y cas e s rock a nd fo ss il s are of s imila r

chemical com po s itio n while eve n petrifyi n gs ub s ta nce s
,

are rarely harder tha n s teel .

I n the ca s e of fo s s il s pres erved i n pa rtly a morphou s

lime s to ne weatheri n g ca n be im itated by leavi n g the
,

l u mp for s om e week s i n water through which carbo n

dioxide is s lowly bubbled Thi s is preferable to .

expo ure t o rai water s i nce the l atter ofte n lead s

s n -
,

to the growth of gree n s tai n s while i ts i n term itt en t

‘

a pplicatio n e n cou rage s precipitatio n of ha rd calcareou s

cru st s O cca s io nal bout s of bru shi n g will s lightly

.

a ccelerate thi s tediou s proce ss S oaki ng i n s tro n g s ugar

.

s ol utio n h a s a s im ilar e ffect e s pecially if the liquid is ,

hot ;but great care i s nece s s a ry to rem ove all trace s of

s ugar before d ryi n g le s t i ts cry s tallizatio n s hould
,

cru mble the s urface O f th e fo s s il H ydrochloric acid .

( or
,
preferably acetic ,
acid ) m ay b e pai n ted o n t o the
m a trix remote from the fo ss il ;but u nle s s the latte r is ,

a s iliceou s petrifactio n it mu s t neve r come in co ntact

,

with the s hell I n the ca s e of c al careou s fos s il s in a

.
62 IN VE R TE B R ATE PALA E ON T OL O GY

n o n -
calcare o u s m atrix s tro n g cau s ti c potas h m ay b e
,

u s ed but it is n ot i nvariably effective a nd need s

, ,

com plete rem oval before the S pecime n i s d ried .

M echa n ical m ea ns fo r rem oval of m atrix are more

ge nerally appl ied tha n chem ical o ne s a nd give m ore ,

s cope fo r i n d ividual s kill I t i s a good pla n to leave a

.

s m all portio n of rock o n the s pecime n if thi s i s po s s ible ,

without hidi ng s ome e s s e ntial s tructure The m atrix .

n o t o n ly a ff ord s a c lue t o the horizo n from which the

fo s s il cam e ( i n val uable i n eve nt of l o s s or di s pl aceme n t

of the l abel ) but m ay s erve to s ugges t the co nditio ns
u n der which the orga n i s m l ived I n at ta ck i n g the .

m atrix n ear the fo s s il pre s s ure i s better tha n blow s

,
.

The d irectio n of pre ss ure s houl d alway s be perpe n dicula r

to th e s hell s urface N ever s cra pe or s cratch the
- .

m aterial ;try to ea s e it O ff i n chip s or gra in s I f the .

rock give s way s udde nly a n d the tool tou che s the ,

s pe c im e n o n ly a prick wil l re s ul t if the s e preca utio ns ‘

are take n ;otherwi s e a s l a s h or groove will d is fi gure it ”

S im ilarly a p oi n ted tool i s better tha n a k n ife T he .

a

m o s t e ffi cie n t i m pleme n t i s a thick n eedle ( s e t i n a

ha n dle) grou n d to a n a ngular blu n t pyramid at the
, ,

tip I n m a ny c a s e s it i s hel pful to keep the s pecime n

.

u nd er water a nd to watch the removal of each grai n

,
“

u n d er a bi noc ular di s s ecti n g m icros cope .

( I I I) D I S S ECTI O N OP F OS S I L S
With the exceptio n of E chi noderm te s t s I n ve rtebrate ,

hard -parts are n ormally exter n al s tructures They O fte n .

exhibit s uperficial excre s ce nce s a nd other or n ame n t that

have but s light rel atio n to a c tual a n atomy us eful though ,

they m ay be for s pecific ide n tifi catio n The a n omalou s .

po s itio n is O fte n reached where the s pecies to which a

fo s s il belo n g s ca n b e rea d ily determi n ed while i t s ge ne r ic,
64 I N VE R T E B R AT E PA LAE ON T OL OGY

de s troyed i n m aki ng the n ext in s eque nce The s pecimen .

s houl d be rigidly fa s te n ed to a flat pl ate a n d grou n d ,

away u ntil s ig n s of the required s tructure s appe a r T he

,
.

s urface s hould the n be clea n ed ( preferably by a w as h

'

of dilu t e acid ) a n d photographed o r carefully d r aw n

,
.

Gri ndi ng m ay the n be re s umed u n til a m ea s u red thick

n e s s h a s bee n removed a n d the del i n eatio n repeated
, .

I f the amou n t de s troyed at each epi s ode is record ed ,

a n d the repre s e n tatio n s of each s urfa ce are to t he

s am e s cale m odel s i n wax or card ca n be co n s tructed
, .

S pecial apparatu s h a s bee n devi s ed for thi s proce ss bu t ,

it ca n be carried out with fair accu racy by ha n d aft er

s om e practice The method i s well ad apted for s tudy
.

of Brachiopo d o r Pelecypod hi n ge s although it h as ,

be e n em ployed m o s t freque n tly o n Vertebrate s ku lls .

A s imple a n d val uable m odificatio n o f gri n di n g wa s

p erfected by the R ev N orma n . Gla ss for di s play o f

Brachiopod b ra chid i a the re s ul t s of h is proce s s ca n

be appreciated i n the illu s tratio ns to D avid s o n s m o n o ’

graph The tightly cl o sed valve s of Brachiopod a ofte n

.

failed to ad m it fragm e n tal d ebri s s o that their cavity ,

became filled with cry s tal l i ne calcite i n trod uced i n

s ol utio n . T h e s oft n e s s a n d t ra ns pare ncy of thi s i n fi ll
i n g depo s ited s o ge n tly that n o i n t e r na l s tructu re s are
,

dam aged m ak e s po s s ible d i s pl ay of b ra ch id ia by s crap

,

i ng away the s hell a n d s urplu s calcite poli s hi n g the ,

re s id ue ( whe n the O paque s hel l ribbo ns are reached ) by

-

dilute acid .

N atu ral m oul d s of Co nchifera ofte n retai n im pre s s io ns

of m u s cle s cars a nd pallial li ne s Whe n s uch m aterial i s
- .

n ot available artificial m ould s may be prepared pro vided

, ,

the i nfi llingm aterial i s compact The mo s t e ffi cie n t method

.

i s that o f bur ni n g I n s om e c as e s e s pecially whe n the

.
,

m atrix is o n ly pa rtly calcareou s the s hell s ca n b e ,

actual ly calci ned a nd removed by bru shi ng I n oth e r s .

,
 T EC H N I Q UE 65

the s pe cime n m ay be heated ( i n s u fficie ntly t o i nd uce ,

chemical cha nge) a nd s ud d e n ly plu n ged i n to col d water

'

The lam i n ate fabric o f the S hel l te n d s to flake O ff a n d ,

careful re p etitio n o f the proce s s will expo s e a clea n

mould R ece n t applicatio n of thi s m ethod to J uras s ic
.

Brachiopoda h a s al mo s t revolutio n ized the cla ss ificatio n

of s om e group s .

I
( )V S T A I N I N G

I n t ho s e fo s s il s tha t have s utur al li ne s or fi ne or n a

m e n t a n d pore s it is ofte n di ffi cult to detect detail s
,

owi ng t o the s i milarity i n colou r betwee n s hell a n d

matrix T hi s applie s particul arly to fo s s il s from the
.

Chal k I n t ho s e the poro s ity of the rock a n d relat ive

.
,

c ompa ct n e ss of the fos s il i nvite applicatio n of a s olu b le

s tai n
. Thi s i s abs orbed by the Chalk a n d m ay be ,

wa s hed O ff from the calcite By s uch mea n s ambulacral .

,

po re s or Po lyzoa n zooe cia are clearly brought out T he .

traci ng of s utu r e s is i mporta n t i n the s tudy of E chi n o

der mata Cephalo p oda a nd Trilo b ita I n Cri noi d s a nd
,
.

E ch i n oid s divi s io n s betwee n the plates ca n be det e cted ,

,

or made more a p pare n t by s uitable treatm e nt O n wel l

,
.

pres erved s urfa ce s s utu re li n e s occur a s very fi ne groo ve s

’ »
it
.
,

T he se ca n be i n te ns ified by appl icatio n of a s tai n givi ng

a gr an ular precipi tate O ne of the be s t reage nts i s
.

I n dia n I n k fre s hly diluted with water A few mome n t s .

after the fluid h a s bee n pai nted on to the moi s te ned

s urface before it h a s begu n to d r y the whole s pecim e n
, ,

s hould b e ligh t ly rubbe d with the fi nger or a c l o th T h e .

stai n on s moo th part s i s removed wh ile tha t i n d e p re s

,

s io ns i s left A bru s h m u s t n ot be u s ed s i n ce the bri s tle s

. ,

s cour out the groove s a n d give di s co n certi ng re s ult s .

A n ex celle n t method for di s pl ayi ng s uture lI n e s o n

pyri t ou s m ould s of Am mo nite s c o n s is ts i n s oa k i n g the
s pecime n in d il ute s ize a nd rubbi ng cha lk or prefer
, ,

5
66 IN VE R T E B RATE PALA EON T OL OG Y

ably a heavier white powder over the s u rface The ,

.

powder work s i t s way i n to the n arrow s eptal s pace s ,

while co n ti n ued rubbi ng preve nt s i t s adhe s io n t o the —

cham ber-wall s .

( )
V S T O R A G E A ND P R E S E RV A T I O N
Whether a col lectio n of fo ss il s i s di s played in ca s e s or
cabi net s or s tored i n boxe s for occa s io n al refere nce the
, ,

o n e thi n g n eedful for i t s s afety i s cotto n wool The -

.

utm o s t care S houl d be take n to keep s peci me ns from

touchi n g o ne a n other The practice of fa s te n i n g fo s s il s
.

with glue to wood or card i s to be deprecated S peci .

me ns thu s s tuck dow n ca n n ot be properly ha ndled for

s tudy ,
while attem pt s to relea s e them ofte n lead to
di s a st er M i n ute fo s s il s which m ay get lo s t i n wool ca n
.
, ,

b e st ore d i n gla s s tu be s i n the bottom s of w h i ch felt h as

‘

be e n fas te n ed .

Mo s t fo ss il s i f protected from s hock or attriti on m ay

, ,

be tru s ted to rem ai n i ntact fo r i nd e fi n ite p e riod s B ut fi

.

s om e e s pecially petrifactio ns i n marca s ite s oo n d i s int e

, ,

grate u n le s s precautio ns are take n The pre s ervatio n of .

pyritou s fo s s il s i s o ne of the m o s t di fficult problem s

a s s ociated with practical palaeo n tology Whe n expo s ed .

to moi s ture M ar ca s ite rapidly d e cay s i n to S ulphu rou s

,

acid a n d F errou s s ul phate ; d i s i n tegratio n is ofte n

i n dicated by e fflore sce n ce o f t he la tt e r The cha n ge .

d epe nd s o n the pre s e n ce of d am p air s o that i n cl u s io n ,

o f a l i ttle C alciu m chloride or wa s hi n g s od a with the

s pe cime n will retard decay th ou gh it ca n n ot preve n t it

,

e n tirely S m all s pecimen s ca n be pre s er ved i n p e trol or

.

s im il ar oil s but thi s i s u s ually i n co nve n ie n t

,
Marca s ite .

fos s il s ca n o n ly be m ai n tai ned i n good co nd itio n by

fr eque n t e x ami n atio n A t the fir s t S ig n of decay the
.
“

s pe cime n s hould b e s oak ed i n a s ol utio n o f cau s tic

68 IN VE R T E B R AT E PALA E ON TOLOGY

fewer di fficultie s appear whe n clas s ifi cati o n i s attem pted ‘

o n a broader ba s i s The res trictio n of evide nce t o

.

d urabl e s tructu re s whos e m odifi catio ns ofte n have little

,
“ ”
co nn exio n with tho s e of m ore vital orga n s ca nnot ,

but i m pair the quality of s y s tematic groupi n g M ore .

o ver it com p el s a d o ptio n of d i ffere n t pri nciples from

,

tho s e employed i n Biology S ec tio ns of the E ch i noidea

.

ba s ed o n c haracter s of ped icellariae of Gas tropoda on ,

the di s tributio n of radular t eeth or of Cephalopoda on ,

“
the n u m ber of bra n chiae ca n n ever be recogn ized by
,

Palaeo ntologi s t s by direct ob servatio n a lthough they ,

m ay be far m ore s ou n d tha n the m ake s hift s cheme s

that have to be erected Whe n rece n t re p re se nt a t i ve s
'

of fo s s il group s exi s t it i s ofte n p o s s ible to apply

,

zoo logical cla s s ificatio n provis io nally b ut t hc i n cl u s i on ,

'

of ma ny form s i n m al acological d ivi s io ns mu s t alway s

be te ntative The n orm al tre nd o f z oological re s earch
.

s eem s t o lead i t s d evotees to overlook the s hel ls or

s keleto n s of I n ve rt ebrate s i n their eage n e ss to s tudy the
r
s o ft ti s sue s . Thi s regret ta ble te nde ncy i s le s s m arked
in the c as e of V e r tebrate s but fo r ma ny I nvertebra te
,

gr oup s a n al ys i s of the ha rd -p art s of l ivi ng form s h a s

bee n left to Pa la e onto lOgi s ts who so ught i n vai n for
'

the i n formatio n they n e e ded i n the record s of biolo gical

l iterature .

There i s a co ns ta n t tem ptatio n by n o m ea ns res tricted,

to Palaeo n tologi s t s t o rega rd the applicatio n of a n ame

,

to eve ry s pecim e n a s a s u ffi cie nt e n d after wh ose a t t a in

“

,
/

“
m e n t the fo s s il i s do ne wi th excep t for s to rage or
d i s play . N ece ss ar y though n am es u s t b e they are m ,

n ot attribute s P f the orga n i s m s b u t merely co n ve n ti on a l

,

te rm s de s ign ed to facilitate refe re n ce A fo s s i l con .

t i n ue s t o d is p la y i t s m orpholog ical pecul iaritie s whether

nam ed rightly wro ngly or n ot at all ;appreciat io n of
, ,

i t s n atu ral qual it ie s ab s olute a n d rel ative i s fa r m o re

, ,
 TE C H N I QU E 69

importa n t tha n a n xiety over i ts place in artifi cial

taxon omy .F urther if the former i n ve s tigatio n i s
,

a dequate t he latter detail s hould autom ati cal ly follow

, .

I deally a n d it m ay be hoped u s ua lly n a me s are give n

, ,

to form s a s i n dicatio ns of their po s s es s io n of c e rtai n

ob s e rved qualitie s di s ti n gui s hi n g them from or a ffi li ,

a ti ng them to other form s ;add itio nal s pecime ns ca n

,

b e ide n tifi ed with tho s e previou s ly desc ribed o n ly after

morphological s tudy comparable with that m ade by the
author s who gave the n ame s At lea s t it is e ss e ntial to
.
,

m ake s ure of the preci s e s ig n ifica nce attached to n ames

before ra s hly a pplyi n g them to n e w m aterial .

Whe never po ss ible refer en ce s hould be made to the

,

pape r in which a n ame wa s origi n ally propo s ed Better .

s till the S pecime n or s pecim e n s which cau s ed the i n tro

,

ductio n of the n ame s houl d be exam i n ed a nd compared ,

with the exam ple who s e ide ntifi catio n is s ou ght F ail .

i n g either of the s e im po rta n t aid s to ac cur acy use m ay ,

be made of l ater m o n ograph s i n which the ge n e ra a nd

s pecie s were rede s cri b e d by authoritie s I f the latter
.

gu ida nce i s n e ce s s ary d ue n ote s hould be made of the

,

fa ct s i n ce ca s es are by n o mea n s lack i ng where s pec ial i s t s

,

have attached name s to form s differe nt from tho s e to

which they rightly p ertai n E s pe cially i n the publicatio n
.

of li s t s of fo s s il s for s tratigraphi cal u s e care s hould be

,

take n to give refere n ce s t o the d e s criptio ns a nd figure s

u s ed i n maki n g the ide n ti fi catio ns .The u s ual a nd ,

proper cour s e of pl aci n g im mediately after a s pec ific

,

term the n ame of i t s origi nal propo ser may prove mi s ,

lead i ng u n les s h is actual s pecime n s o r de s criptio ns have

been s tudie d A n exam ple of thi s d a n ger i s a fford e d
.

by one of th e b e s t -k n ow n chara cteri s tic fo ss il s of the

Lowe r O olites I n 1 8 29 Phillip s propo s ed the n a me
.
~
“

Clyp m: semzs ulca tus for a n E chi n oid fou n d i n the

'

Coralli ne O olite o f Mal to n the s pe cie s wa s l ater made

70 IN VER T EB R AT E PALA E ON T OL OGY

the ge notype of Pyga s ter by Aga ss iz B y a s eries of .

error s a nd m i s co nceptio ns the s pe cifi c n am e came to be

,

applied to a n allied but qu ite di sti n c t form com mo n

, ,

i n the I n ferior O ol ite of Glouce s ter s hire F rom 1 8 57 .

on ward s thi s Lower O ol itic E chi n oid wa s alway s called

.

Pj ga s ter s emzlmlca tu s the m i s appl icatio n of the s pecific

,

’
term bei n g s tereotyped i n Wright s m o nograph Late r .

writer s habitually referred to Pygmter s amb a /ca lm

whe n deali n g with the I n ferior O olite
form whe n a s s ub s eque n t re s earch h a s proved the
, , ,

specie s mea n t was “

P s emzs ulca tus Wright ( m m
’

.
,

Phil lip s) H a d their re fere n ce s be e n m ad e i n the

.

latter way m uch co n fu s io n wou ld have bee n pre

,

ve n ted s i n ce P s emir ulca tus ( Phillip s flan Wright)

, .
,

i s an e n tirely d i ffere n t s pecie s a n d occu rs i n the ,

Co rall ia n o n ly .

S i n ce co n fide n t ide n ti ficatio n of s p e cie s d em an d s lo ng

s udy a nd ofte n s pecial k n owledge of the group s co n
t ,

c er ned it i s practi cally im po s s ible for a s tratigrapher t o

,

defi n itely a n d corre ctly n ame m a ny o f the var ied form s

“
that he collect s U n les s the help of s pecial i s t s i n t he
.

di ffere n t group s ca n be obtai ne d it i s s afer a nd le ss

’
‘

, ,

m i s leadi n g to com pare fo s s il s with de s cribed type s

,

rather tha n to ide n tify them E s pe cially s i n ce true

.

ide tity betwee n ny two pecime ns i well n igh i n

n a s s -

co nceiva ble a l i s t of fo s s il s which i ncl ude s abs ol ute

,

s pecific determ i n atio n s i s le s s co n vi n ci n g tha n on e i n

which the fo s s il s are s aid to be comp a r a b le with s pe cies

previou s l y d iag no s ed Greater experien ce a nd m ore
.
,

m ate rial alway s te n d to le s s e n the co n fide nce with

,

which s pec ifi c n ame s are applied ;it i s n o t furtheri n g

the cau s e of S cie n c e if S tratigrapher s ru s h i n where
P a laeo n tologi s t s fear to tread !
The foregoi ng com m e nt s apply to ori gi nal work ;
the ir rig id adoptio n would m ake pedagogic p a la e on
7 2 I NVE R T E B R ATE PALAE ONT OLO G Y

a nd m a ny fam ilie s S i m il arly the n ame R hy m/zonella

.

i s re s tri cted to a few p e c ul iar s pecies ; the origi n al

gen u s i n cl uded al mo s t al l the m ember s of the fam ily
”

R hy n cho nellid ae of m oder n cla s s ificatio n The s t r uc .

tural character s who s e d i s cove ry h a s led to di s member

”
me n t of s uch com prehe n s ive ge n era a re ofte n o f
real i m porta n ce but n o l e s s freque n tly difficult to a p
,

p r e c i a t e without detail ed s tudy A s tude n t o f Pa la e o n .

t o lo gy for i t s o wn s ake ca n a nd s hould recog n ize , ,

rece n t ge neric d ivi s io n s a nd try to check their validity ;

,

but a s tude nt of Geology who s e pu rpo s e i n deal i n g ,

with fo s s il s i s to d i s cover i nd i ce s of S tratigraphi cal

‘

h o ri z o n s ca nn o t ho p e to p rove for h im s el f that a ny

, _

g e n e r ic di ffere n ce s s eparate s u ch well -k n ow n form s a s

Ter ebr a tula sp /za er oéa a lis a nd T s i mp lex I t i s of

'
’
. .
,

cours e po s ibl e for hi m to r m ember parrot wi s e that

,
s e -
, ,

the o ne i s a Sp ka emza e t/zy r i s while the other is a ’

P s eu d ogloss ot/zy r zs but i n form a tio n gai n ed at s eco nd

'

h a n d is n o t k n owledge a n d i n s uch a ca s e is o f very , , ,

d oubtful val ue F or al l bu t s peciali s t s mo s t o f the

.
,

Terebratulidae m u s t be cla s sed u nder the n ame Ter e

b r a tula (s ens la t ) ;all A m m o n ite s m u s t b e A mmoni tes
.

s t ill, though n o s i ngle S pecie s ca n rightly cl ai m s uch

a ge neric prefi x .
 C H A PT E R I V

G EOL OG I CAL PALA EON T OL OGY

( )
A S T R AT I G R A P H I C A L

( )
I Z O N E S A ND H E M ER A E

H RO U GH O U T the la s t hu n d red year s Geol ogi s t s ,

have fou nd i n fo s s il s e ffi ci e n t a n d reliabl e evide n ce

for the chro nologi c al cl a ss ifi catio n of s edi me n tary
“
rock s The claim m ade by William S mith that S tra ta
.
,

[
”
c a n be ] ide n tified by orga n ized fo s s il s h
, a s bee n s u b s t a n

t i a t e d i n the mai n a n d the pri nciple h a s bee n appl ied to

‘

refi neme n t s u ndream ed Of by that pio neer S ubd ivi s io n .

O f the ge ol ogical record i n to the great era s Palaeozoic

’

Mes ozoic a n d Cai nozoi c i s ba s ed o n palaeo n tological

evide nce a s the n ame s imply ; relatively brief period s
,

of time s uch a s h e me r a e are recog n ized s olely b y

, ,

d etailed s tudy O f fo ss il s .

The cla s s i ficatio n of ro ck -sy s tem s wa s a fairly sim ple -

problem for early s tratigrapher s who s tarti n g from

, ,

preco nceived idea s believed i n the ca ta s trophic do ctri n e

,

that the fau n a s O f s ucce s s ive p eriod s were di s ti nct i n ,

member s a nd origi n Lo n g year s O f pal aeo n tologic al

.

re s earch were nece s s ary to prove that t he d ati n g

evide nce a fforded by fo ssil s i s not s o sharply defi ned ,

while growth of realizatio n O f the proce s s of evo lutio n

was fo s tered by a n d co n tributed to the k nowledge th a t
, ,

each fau n al a s s emblage i s i ntimately co nnected with

i t s a n tecede nt s a nd s u cce s s or s . Whe n s pecie s ce as ed
73
74 I NVE R TE B R AT E PAL AEON T OLOG Y
‘

to be regarded a s i m m utable s harp d ivi s ions b e t wee n

,

s tratigraphical s tage s di s appeared Biological a nd

.

geol ogical sy s tem s O f cla s s ificatio n ca m e to be m oulded

i n to differe n t a n d m ore pla st ic form s who se lack of
, , ,

the O ld tim e preci s io n proved i nco nve n ie n t but n atural

-
,
.

I t is perhap s fo rt u nate that s ubd ivi s i on O f geological

, ,

time i n to i t s mai n s ectio ns wa s carried out by believer s

i n cata s trophe s a n d s pecial creatio ns s i n ce it s upplie s ,

a n ece s s ary ba s i s for arbitrary cl a s s ificatio n a nd woul d ,

be diffi cul t to e s tabl i s h with the further k nowledge

n o w available J u s t a s the ge nera a n d s pecie s diag n o s ed
.

by Li n n aeu s a n d h is follower s are s till recog nized ,

though the co nceptio n of their value h a s altered ;so

the geological pe riod s n amed by S m ith M urchi s o n , ,

S edgwick a n d Lyell s e rve a s co n ve n ie n t excerpt s from

eter n ity I t i s a s triki n g te s tim o ny to the accura cy
.

a nd i ns ight of the fou nder s of S tratig raphy th at thei r

s chem es O f C la ss ificatio n ba s ed on l im ited experie nce

,

a n d philo s ophical m i s co n ceptio n ca n s till be m ai n tai n ed

, ,

wi th but trifli n g m odificatio n after a ce n tury of d is

,

cov e r ies m ad e po ss ible b

y their e ffort s .

The s tude nt O f H i s torical Geol ogy h a s to recko n

with two factor s s trati graphical a nd chro n ological re
,

s e c t iv e l
p y . I t i s n ece s s ary to bear i n m i n d that the
a mou n t of work d o n e i s depe nd e n t o n but i n n o se ns e ,

com me n s urate with the ti me s pe n t u po n it Thu s

,
.

on e a n d the s am e i n terval O f time m ay be rep re s e n ted

by the accu m ulatio n O f a thou s a n d feet of s edi me nt '

i n o n e place te n feet i n a n other a n d the rem oval of

, ,

m ater ia l by de n udatio n i n a third While s tratigraphy .

( lite rally i n terpr e ted ) c a n record a n d correlate eve n t s

that have happe ned i n cour s e of t i m e it ca n s upply no ,

reli abl e evide n ce O f t he d ura t i on O f time i ts elf Phy s io .

g raphica l co n d itio n s
( o n which de p o s itio n o f s trata

d epen d s ) are too v a riable i n tim e a nd s pa ce to give

, ,
76 I NVE RT EB R A T E PAL AE ON T OL OGY

t heir highe s t devel opm e n t s ucce s s ively The time d ur .

i n g which s uch a l i n eage m ai n tai n s i t s a cma ic qual itie s

biological a n d nu merical is calle d the hem era O f
,
“ ”

that s er ie s I n s t ri ct t erm i n ology a ny de p o s it s th at

.
,

may accum ulate duri ng the hemer a a r e the z o ne “ ”

cha racte rized by the grou p O f orga n i s m s I f thi s d i s .

ti nctio n is bor ne i n m i n d it b ecome s m a n ife s t that zo n e s

,

m ay or m ay not be repr e sen ted i n variou s place s their

, , ,

exi te nce d epe n d i g

s n o n O pportu i tie s fo r rock d epo s itio n
n -

d u ri n g their h e me r a e A hemera i s a s ectio n of time

’

s ;while a zo n e i s the ta n g ble

( a s the n ame s ugge s t ) i
re s ult of l ocal epi s od e s tra n s acted i n t h e s pecified
i n terval Thu s whil e it is pe rfectly j u s tifiable to record
.
,

that a zo ne doe s n o t occu r i n a certai n l oca lity it would ,

be ab s urd t o s u gge s t that a hemera wa s ab s e n t although ,

it m ay be u n repre s e nt ed by actual mate rial s .

T he lea s t s ati s facto ry character of h e me r a e is their

d eterm i n atio n through evide n ce s uppl ied by particular
fo s s il s A lthough m o s t s pecie s pr ove to have wide
.

ra nge s at their tim e s O f acme they rarely become ,

world wide— the m o s t co s m op ol ita n form s are u s ually

-

tho s e of lo n g d u ratio n who s e exte n ded acm e s s how l ittle

prom i ne n ce But a race of orga n i s m s who s e vigou r is
.

s uch that i t s m ember s would be cho s e n a s hemera ]

i nd ice s wil l n orm ally have repre s e n tative s i n al l part s

of the world where co n ditio n s are s uitable ;a nd the
m arked te nd e n cy to s y n chro nou s paralleli s m i n the
evolutio n of hom oge netic s tock s wil l u s uall y p rod uce
com pleme n ta ry s p ecie s that replace the m i s s i n g guide s
a nd ful fil their r ole
. I n co n s eque n ce it i s ofte n mo s t
.
'
,

s at isfacto ry to co ns id e r a hem er a t o be i n dicated by a

”
m o rpho ge netic m od e or pha s e of s p e ci a liz a tio n tha n
, ,

by a ny s pe cial s pe cifi c type s .

~
A zo n e m ay be recog n ized by the pr ese nce O f t he
hem era] i n d ex i n a n ac ma ic s tage O f evolutio n but is ,
 G E OL OG ICAL PALA E ON TOLOG Y 77

mo re s a tis fa ctori ly d e t e r mine d by co n s ideratio n o f i t s

'

e nt ire fau na T he i n dex -fo s s il m ay ha ve bee n o f s uch

.

a n a t ure that it could n ot exi s t

~
or at lea s t flouri s h
, ,

u nde r certai n physi ographic al co nditio n s but it i s ,

probable that s om e O f i t s a s s ociate s were m ore cat holic .

I t n ot i n freque n tly happe ns that two or more organ i s m s ,

belo ngi n g to e ntirely d i ffere n t group s reach a c ma ic ,

s t a ges at abo ut the s am e tim e Th us i n the Upper

.

Chalk the Belem n i te A cti noea max qu a d r a tus a n d the

,

E chi n oid Of a s ter pi lu la bo th charac t erize the s a m e

zo ne which 15 n amed afte r the former type But while
,
.

the di s tributio n of the Belem n ite I S s omewhat s por ad i c ,

that O f the E chi noid i s m ore u n iform s o that the ,

u a d r a tu s -z o n e ca n O fte n be recog n ized withou t d i

q s
"
cove ry O f a s i n gle s pecime n of the n ame fos s il
- .

S i nce recog n itio n O f h e me ra e date s back n o fu rt her

tha n the ea rly year s of thi s ce n tu ry while zo nes were ,

propo s ed fi fty years b e fore it i s n atural that the c on

,

ce p tio n of a zo n e a s t he m aterial depo s ited d uri n g a

hemera i s a m od er n refi n eme n t H e nce ma ny zo ne s

.
“

that s till do good s tratigraphical s ervice have no real

“
chro n ological preci s io n but are of th e n ature of s ub
,

period s . Like l arger divi s io ns O f s ed i me n tary rock s ,

t hey are ide ntified by the s pecific a nd propo rt io nal

qualities O f their fo s s il -co n te n t s While s uch zo ne s ca n
.

be ea s ily recog n i z ed i n their middl e portio n s they te n d ,

t o have ill -d e fi ne bou n darie s ;i n m a ny ca se s they prove

d
‘

on further p laeo n tological s tudy to s prea d over s everal

a ,

hemerae .

It may b e a n ticipated tha t t he s tratigraphic a l

c la s s ificatio n of the future wil l be graded i n accord an ce
with the a cma ic s tages of s pe cie s or s pec ie s group s
( h e m e r a e ) ge nera ( s tages ) fam ilie s ( period s ) a nd larger
, ,

group s ( era s ) I n a rough way th i s g roup i ng i s alr eady

.
,

i n us e . The Lower P a laeozoic ( or Proterozoic ) era i s

78 I N VE R TE B R A T E PALA E ON T OL OG Y

m ark e d by the pre se n ce O f Graptolite s Trilobite s a nd , ,

certai n O rder s a n d s uborder s of Brachiopod s that a re

eithe r re s tricted withi n or rare beyo n d it s l im it s
, , , .

S im ilarly the U p pe r Pal aeozoic ( D e u te roz oic) co n ta i ns

rem ai ns O f pla n t s F i s he s Moll u s c s a n d group s of
,

Brachiopod s that are o rd ma lly d i s ti n ct from t ho s e

com mo n i n the precedi n g or s ucceed i n g era s The .

M es ozoic i s O fte n called the age O f R eptiles bu t s ome “

,
”

order s o f E ch inO id s Brachiopod s a nd Mollu s c s ( e s pecially

,

Cephalopod s ) are equally characteri s tic The C ai nozoic .

”
i s the “
age O f M am m al s H owever neithe r thes e
.
,

great d ivi s io ns no r m i nor o n e s ca n be defi n ed with

, ,

accuracy u n til Pal aeon tology h a s i ndicated the lim it s

O f h e me ra e the u n it s O f geological tim e
,
.

( )
I I C H A R A CT E R I S T I C F O S S I L S

Ac q uai nta n ce with fo ss il s chara cteri s t ic of the variou s

period s is u s ually re q uired of s tud e n t s of Geology a nd ,

i s e s s en tial for th os e propo s i ng to i nve s tigate stratigra phy

i n the fie ld The forme r s u ffe re rs d e s e rve c o ns id e rabl e
‘ ‘

a
.

s ym pathy s i n ce the exige n cie s of exami n atio n cou rs e s

,

rarely leave tim e for adequate i nquiry i nto the biologi cal
chara c ter s of the fo s s ils who s e appeara n ce n ame s a n d ,

ra n ge they are expected to lear n wh ile b oth cla s s e s O f e

workers are ofte n t roubled by the varied opi n io ns of

authoritie s a s to the particul ar s pe cie s that are
“ ”
characteri s tic Moreover N ature ofte n ad d s to the
.
,

co n fu sio n by local or s poradic di s tributio n of com mo n

form s I t i s u sual a nd correct to de s cri b e H emzez a d r i s '
' '

.
, ,

i n ter med z a a s a chara cteri s tic fo s s il of the C orallia n ;bu t

'

whil e a wo rker I n W ilt s hire m ight perhap s col lect

s pecim e n s of that s pecie s by the bag -full a nother i n ,

B erk s hire sear chi n g at t he s am e horizo n a nd i n litho

,

l ogically s im i l a r bed s would n ever fi nd a trace

,
.
80 IN VE R TE B R AT E PALA EON T OL OG Y

Cambria n the le ss bra n ched T e t ra gra p ti d s a nd D id ymo

,
-

g pr a t i d s o f the Lo w er a n d M idd le O rdovicia n the ,

D i p logr a p t i d s O f the M iddl e a n d Upp er O rd o vicia n a n d ,

the R a s tri t id s a nd M o no gra p t i d s of the S iluria n A .

m ere l i st O f the Graptol ite s pecies s elected a s zo n al i n

d ice s i s s uffi cie nt ly formidabl e ;but whe n attempt s to
recog nize a n d d i s ti n gui s h s pecim e ns or eve n drawi n gs
, ,

O f the variou s s pecie s are m ade by s tude n t s u n trai ne d

i n m ethod s O f pal aeo ntological r e s earch the fou nd a ,

tio ns of bel ief are s hake n a n d al l i ntere s t fou n de rs in a

,

whirl of bewi ld erm e n t A gitati n g problem s a s to the

.

di ffere n ce betwee n Ump lza lotr od zus d is c07 s a nd 0 r ugos ur

- '
.

( s pecie s beloved O f the exam i n er ) have littl e real value ,

eve n if s olved ;s i n ce both form s occur together i n the

We nloc k Lim e s to n e ;while eve n the family d i s ti n ctio n
bet w ee n A ct i n OCri 11 id ae a n d A mp h o ra crinid a e h a s no
‘

greater s tratigraphical s ig n ifica n ce I s it not m or e

.

u s efu l a nd le ss provocative of cram mi n g to s how

,
“
,
”

that the wri n kled T urbin id ae abou n d i n the S il uri a n ,

a n d the s m ooth E uo mp h a lid a e i n the Carbo n ifero us

while the A ct ino cri noid a are wholly cha racteri s tic of
'

the l at t er period ? The m o s t eleme n ta ry s tude n t ought

to k now that A m m o n ite s are not fou n d i n Cai n o z oic
roc k s ;but he ought not to k n ow the di ffere n ce betwee n
D ae ty lz oeems commu ne a n d P er onoeems a nnu la tu m
'

If .

the latter detail h a s bee n forced i n to hi s u nprepared

m i n d s o me poi n t of m ore ge neral applic atio n and
,

greater i mporta n ce m u s t have bee n om itted or ej ected .

( )
B P H Y S I O G R AP H I C A L ‘

I n d icat ion s a fforded by fo s s il s as to the co nd itio ns p re

val e n t at the time a n d place of their life are hard ly le s s
i m porta n t tha n their chro n ological evide nce Although .

it is u ns afe to a s s u me th a t p as t m em be rs of a group of
 G EOLOG I CAL PALAEON T OL O G Y

an im als n ece s s arily s elected the s am e ki nd of s urrou nd

i ng s as tho s e favou red b ytheir des ce nda nt s relia nce can
,

u s ua lly be pl aced o n ad aptive m orphological feature s .

The l atter ba s is for argume nt h as a further ad va n tage ,

s i n ce it c a n b e recog n ized i n e n tirely exti nct form s .

Ther e i s s till roo m for doubt a s to the preci s e s y s tematic

“ ”
po s itio n O f the Tabula te Coral s ;but the corre s p ond
e nce O f their ge neral s tructure with that o f livi n g
M ad reporaria make s sim ilarity O f habitat m ore tha n
probable Agai n the true Coele nterate affi n ities of the
.
,

Graptolite s a re n ot s ecurely es tabli s hed but there ca n

,

be no rea s o n able he s itatio n i n a s cribi n g to the se exti n ct

a n imal s habit s O f life like t ho se adopted by rece n t
H yd rozoa a n d Polyzoa po s s e s s i n g comparable m orph o
O gica l feature s .

J u s t a s s ome livi n g I nve rtebrate types are k now n to

be cap a ble of mai ntai n i ng vitality u n der a variety Of
co n ditio ns or i n s pite O f relatively s udde n cha nges of e n
,

vi r o n me n t s o s om e ki n d s of fo s s il s m ay n o t imply defi n ite

,

ph y s iographical qu al itie s for the s tratu m i n whi ch they ,

occur A s a ge n eral rule the m ore highly s pecial ized

.
,

gr oup s are se ns it iv e t o cha n ge ;s im pler type s are m ore

a d a ptable The d i s ti nctio n appl ie s equally i n h um a n
.

a ffair s : a ge n era l educatio n e n able s i t s s urvivor to l ay

h is ha nd to al mo s t a ny avail able work the s peciali s t is
,

i n capable of s ucce s s be yo nd h is part icular s phere .

Las tly it m u s t be remembered that the rel ic s O f

,

orga ni s m s may ofte n be d rifted in to very di ffere n t

phy s iographi cal provi n ces from tho s e e ss e ntial fo r the
‘

life O f their o wner s a nd m ay become emb e d ded i n i n

'

appropriate m atrice s am o n g s t ra n ge compan io ns The .

freque n t occurre n ce o f fo s s il wood i n the F ore s t Marbl e

-

a n d L ower Gree ns a n d could n ever be u s ed as evide n ce

that Co n ifer s o nce grew i n O y s ter -bed s ; s im ilarly the

pres en ce of fres h water s hell s m i n gled with tho s e of
-

6
82 I NV E R T E BR AT E PALA EON TOL OGY

m ari n e type s m u s t be as cribed to co n fu s io n Of fau na s by

tra ns porti n g age n t s F u rther d i s covery of Graptolites
.
,

i n black s hale s who s e aby s m al origi n s eem s probable i s

no proof O f the be n thic ha bitat O f tho s e orga n i s m s .

Their occurre n ce i n s uch depo s it s 13 probably d ue to the

i
s i n k n g O f pla n kto n ic s urface dwellers whe n buoya n cy
-

wa s l o s t .

( )
I T ER R E S T R IA L F A U N A S

The rarity of fo s s il s i n terre s trial depo s it s i s par

t icula rly marked i n re s pect of I n ve rtebrate s O n ly two .

p hyla the
, Mollu s ca a nd Arthropod a are ge n erally avail
,

able for fo s s ilizatio n o n la n d a n d thei r repre s e n tatives

,

are co n fi n ed to relatively few type s e n d owed with s om e

what delicate s hell s The d omi n a n t terre s trial m embe rs
.

of the two phyl a are the Ga s tropoda a nd I n secta i n the

ca se o f the former a n exiguou s proport io n o n ly are
f

“
Pul m o nate T he average s n ail h a s a le ss m a s s ive
.

“
s hel l tha n a m ari ne Ga s tropod a nd s lug s s ec rete s o

l ittle cal careou s m atter that pres ervati o n a nd di s covery

of their rem ai n s m us t alway s be doubtful E xcept i n .

the upperm os t layers O f t he s oil dead s n ail -shell s are

,

rarely fou n d owi n g to the rea d y acce s s O f bacterial a nd

,

aqueou s age n t s of de s tructio n They may Occu r i n con

.

s id e ra b le n um ber s i n fre h water depo s it s whither they

s -

have bee n d rifted by s tream s ;but i n s uch ca s es they

have pa ss ed out O f their true provi n ce a n d s erve o n ly ,

to i n d icate the pr o xim ity O f l a n d to the pl ace O f their

bu rial T O their extreme rarity i n s ub -a erial depo s it s a
.

further co n trib utory factor exi s t s i n the u s ually de s ert

origi n O f geol ogical ly perm a ne n t ter re s trial accu mula
tio n s Moreover i n rock s of co ns iderable a n tiquity i t i s
.
, ,

di fficul t to be certai n O f the p ul m o n ate character of a

Ga s tropo d s i n ce the ou twar d form O f the s hel l give s
,

l ittle guida nce toward s determ i n atio n of s o ft ”a na t omy , .

84 I N VE R T E B R AT E PALA E ON T OL OGY

agai ns t perma ne nce of pres ervatio n The proc e s s of .

filtr a tio n that co n vert s alluviu m i n to gravel i nvolves e x

os u re O f s hell s to the d eva s tati n g a ctio n O f percolati n g
p
water ( by which eve n Mam m oth s tu s k s a re d es troy e d )
’

while a n ordi n ary gravel terrace a s s um e s i n tim e a

po s itio n s o expo s ed that i t s rem oval by de n ud atio n is
wel l n igh certai n The re m ai ns of fluvia tile orga n i s m s
.

ca n o n ly attai n la s ti n g pre s ervatio n whe n drifted i n to

a l ake or s e a a n d the vici s s itude s of their j our ney are

,

co n s i s te n tly d es tructive .

S wam p or lake depo s it s u s ually po s s e s s s ome vi rtue Of

-

co n s ta n cy a nd n orm ally accu m ulate u n de r very tra n quil

,

co nd itio ns They are O fte n argillaceou s a n d s ometim es

'

. , ,

peaty affordi n g fairly s afe burial to calcareou s s hell s

, .

I n the ca s e O f aquatic M ollu s c s O f high a n tiquity the ,

s a m e u ncertai n ty a s to adaptive s tru ct u r e exi s t s a s i n

terre s trial form s ; it i s m ore from evide n ce of thei r

a s s ociates tha n from a ny i n tri ns ic qualitie s that
A r eka nod on from the O ld R e d S a nd s to n e or Ca r éan zeola
'

from the C oal Mea s ure s are regarded a s fres hlwa t e r

m u s s el s The U n i oni dae O f the York s hire J u ra ss ic the
.
,

Cyre n id a e O f the Wealde n a n d the Li mn a e i d a e o f the

,

O l igoce ne give more ce rt ai n i n dic a tio n s of the lacu s tri n e

o r deltaic co n d itio n s o f their s urrou n di ng s s i n ce they ,

cl ea rly repre s e n t group s who s e pre s e n t di s tributio n i s O f

that n ature S hell m arl s O fte n i n terbedd ed with peat
.
,

are freque nt m the l ater Tertia ry depo s it s a nd m mo s t ,

c a s e s m ark l acu s tri n e phas e s of s edime n tatio n .

The A rthrop od s i n habiti n g fre s h -water are u s ually

s ma l l a n d belo n g
,
for the m o s t part to grou p s equally
, , ,

or better repre s e n ted i n m ari n e fau n a s

,
There i s little
.

d i ffere nce in the s hell s or carapace s O f me mb e rs O f ~

allied gr oup s i nhabiti n g the two provi n ces A mo n g .

Os tra cod a the Cyp r i d a e u s u al ly p re fer fre s h water their

,

re m a i ns are particularly abu nd a n t i n the delt a ic pa r ts of

 G E OL O G I CAL PALA E ON T OLOGY

the Wealden Perhaps the s warm s of B ey r zc/u a that

’ '

characterize the uppermo s t parts O f the S il uria n m ay

i nd i cate the o n com i n g Of O ld R e d S a nd s to ne l ake

co n ditio n s ; but clo s ely allied form s abou n d i n the
Uppe r O rdovi cia n where no s uch pro s pe ct s eem s
,

pro b able I n a ny c a s e d etailed a n d s pe cial k n owledge

.
,

“ ”
o f the s e s m all bivalves is required if m ari ne form s
( whi c h are comm o n i n the Chal k ) are to be d i s ti n gu i s hed
from tho se O f fre s h water I n like ma n ner it would be .
,

di ffi c ult from s tudy O f the c arapace alo n e to a s cribe to

, ,

a crayfi s h fluvia t i le a nd t o a lob s ter mari ne habit s of

, ,

l ife
.

Perhap s the m o s t s ati s factory palaeo ntological i ndica

tio ns O f eithe r terre s trial or fre s h water origi n for a -

depo s it are of a n egative type I f fo s s il s occu r i n a .

s tratu m a n d n o n e O f them belo n g s t o exclu s ively mari n e

,

group s the prevale n ce of o ne or the other of the no n

,

m ari ne co n ditio ns m ay be pres umed While m o s t .

fluvi a t ile I nve rtebrate s have n ear all ie s livi n g i n the s e a ,

m a ny phyl a are abs olutely re s tricted to the la t ter regio n .

A gai n while it i s po s s ible a nd by n o mea n s u n u s ual for

, , ,

i nla nd orga ni s m s to be d rifted i n to the se a t he re ver s e ,

proces s is so u n likely a s to be practically negligible

A very i n s tru ctive c as e O f fa unal m od ificatio n d ue to
a s in the U pper S iluria n
h g p f l ch ge ccur
'

p y s i o r a n ca n o

rock s of S hrops hi re ;a s imil a r phe nome no n ( at al mo s t

the s am e horizo n ) h as rece ntly bee n recognized i n ,

America I n the grey flag s of the Upper Ludlow s erie s

. .

. ,

Pro t re ma t ous Brachiopod s s uch a s Or tfu s lu na ta a n d

'

C/zonetes s tr za tella are pre s e n t i n extraordi n ary pro

'

fu s io n ( Pl vi fi g. . They are accom pa n ied by a s ub

.

'

ordi n ate Mollu s ca n fau n a i n which s pecies O f P ter z nea , ,

M od i olop s zs a nd Or t/zoeer a s are the mo s t abu n da n t types

’

A s the flag s pa s s up t oward s the Downto n ia n s a nd s to n es ,

the propo rtio ns of the group s u n dergo revers al Al mo s t .

86 IN VE RT E B R A T E PALA E ON T OL OGY

the o nly Brachiopod s are L z ngula e Or r/wa r d s d i sappear s

’

a n d the bulk O f the fau n a i s com po s ed of Pelecypo d s ,

with occa s io nal Tubicolou s A n n elid s a nd E u rypteri d

“ ”
A rthropod s The Ludlow Bo ne Bed give s s triki n g
.

"

te s timo n y to al m o s t cata s troph ic cha nges at thi s s tage ,

a nd i n the true D o wn t o n i a n that s uperve ne s eve n

‘

Pelecypod s have go n e ;L z ngu la alo ne s u rvive s re p re

'

s e n ted by dwarfed form s s uch a s L mi m ma The

'

. .

micaceou s s trata O f the Pa s s age Bed s e ncl o s e L comm .

,

a n d the n al mo s t all palaeo n tol ogical evid ence i s l o s t i n

the m arl s O f Lake Mo n m outh T he grad ual tra n s itio n

.

from m ari n e through b rack i s h to fre s h -water co n ditio ns

, ,

i s very clearly s how n i n thi s fau n al s eque n ce The .

higher Brachiopod s are excl u s ively m ari ne a n d d is ,

“ ”
appear a t the fi r s t s u s p icio n O f e s tuari n e co ntam i n a
tio n Ma ny Pelecypod s ( e s pecially the Mytilid ae a
.
,

group n early rel ated to M od i olop s zs ) are i nd i ffere n t to

'

d ilutio n of s e a wate r al though they ca nnot s urvive whe n

-
,

s weet n e s s h a s bee n quite attai n ed The Ceph alopod s

.

are a s defi n itely m ari n e a s the Brachiopod s ; bu t

L i ngu la o n e of the s im ple s t member s O f the latter
,

phyl u m m ay well have po s s e s s ed the catholicity o f

,

ta s te com mo n to m o s t l owly form s A s omewhat .

s im ilar s eque n ce carried o u t with other ge n era a n d i n

,

the revers e d irectio n ca n be traced through the R haetic

,

fau n a s from the Tea Gree n M arl s to the Lower Lias .

( )
I I I M A R I NE F A U NA S
The l ife o f the s e a ca n be grouped broadly i nto
two zo n e s e a ch c a pable of further s ubd ivi s io n T he
two m ai n provi nce s c o rre s po n d fairly clo s ely w
.
,

ith t he
d etrital a n d pelagic z o ne s of s edim e ntatio n re s pe ctively .

F orm s i nhabiti n g the l ittoral be lt ca n be s ep a rated i nto

tho s e dem a n di n g a clea n e nv
“
iro n me nt a n d thos e
”
,
88 I NVE R T E B R AT E PALA E ON T OLO G Y

co n d itio ns I t i s u s ually n ece s s ary to d eterm i ne the

.

bathyme t ric ra n ge of the fau n a of ea ch pe riod by variou s

m ea ns ( largely i nd uctive ) a n d to apply that k n owledge
,

to the fo ss il s O f the period co n cer ne d I n thi s way .

i n tere s ti n g co n cl u s io n s ca n be reached a s to variatio n s

i n the depth O f the Chal k s e a A m m o no id s a n d .

G a s trop od s are fairly abu n da n t i n the Lower Chal k ,

be com e rare I n the Middl e Chal k reappe ar i n great ,

n u m be rs i n the Chalk R oc k a n d practi c ally di s appear

-
, .

from t he U pper C hal k Their pre s e n ce corre s po n d s

.

with the prevalen ce o f rel atively s h a ll ow water ;a n d for ,

the Cretaceou s period at lea s t they may be u s ed as ,

evide nce for s uch co n d itio n s S imil arly D i scard ed a n

.
, ,

E chi n oid a n ce s tral to m oder n

“
Cake U rchi n s a nd
- ”

“ ”
S a n d D ollar s ( who s e d i s tributio n i s m ai n ly lit toral ) ,

occur s i n the Lower C hal k a nd i s repre s e n ted by

'

mi n ute s pecies i n the M iddle Chalk ;but it is e n tirely

ab se n t from higher zo n e s in thi s cou n try I n o t her .

di s trict s however s uch a s S Ou th er n F ra n ce a n d N orther n

, ,

A frica the ge n u s a nd i t s all ie s are abu n da n t throughou t

,

the S e n o n ia n i n d epo si t s O f le s s pelagic cha racters tha n

,

th e Briti s h Uppe r Chal k .

( )
I V F O SS I L S AS E VI D E N CE OF
'

C L I M A TE
U ncertai nty as to the habit s O f exti n ct ty p e s o f
I nvertebrate s a n d virtual res tri ctio n O f their remai ns to
,

“
m a ri n e type s m ake attempt s at clim atic re co ns t r uc
,
”
tio n of d oubtful val ue whe n ba s ed o n s uch evide nce .

Althou gh reef-buil d i n g Coral s flouri s h in warm seas a t

pre s e n t there i s n o more rea s o n to S uppo s e that their
,

pas t d i s tributio n was s imil arly determi ned tha n to

r egard Probo s cidea a s ex c l u s ively tropical form s T he .

M am m oth e n d ure s a s a la s ti n g war n i n g agai ns t a ss ump

tio ns of thi s k i n d I n the ca s e o f relatively m od er n
.
 G E OL OG I CAL PALA EON TOLOGY

fau n a s s ome Qonfi d e nce ca n be attach e d t o deductio ns

,

with regard to cl im ate e pecially whe n hallow water

,
s s -

form s are co n cer n ed T h u s the Lower Tertia ry Mollu s c s

.

of E n gla nd s how a defi n itely s ubtropical facie s whe n ,

com pared with livi n g type s ;tho s e of the O ligoc en e

s ugge s t temper ate co n ditio n s while tho s e of the Pl io ce n e
,

a nd Lower Plei s toce ne are d i s ti n ctly arctic I n al l

.

cas e s however terre s trial orga n i s m s ( e s pecially pla n t s )

, ,

are m ore expo s ed to climatic i n flue n ce tha n tho s e i n

habiti ng the s e a a nd therefore give m ore delicate a nd
,

reliable i ndicatio ns of cha nge .

I t is chara cteri s tic of partly e ncl o se d gulfs t o develop

s our qualiti e s in tropical climate s owi n g to ex ces s ive
,

loc al e vapo ratio n Co n ce n tratio n of th e se a -water ( s uch

.

as that com m o n ly f u n d o i n coral l agoo n s ) adv e

- rs ely
a ffec t s the fa un a giv i ng s tu nted race s pro n e to d i s ea s e
,
'

, .

Perhap s the s m all s ize o f mo s t fo s s il s in the Mag n e s ia n

/

Lime s to ne ( a n d i n m a ny Tria s s ic d olom ite s on the

Co n ti ne nt ) m ay be a s cribed to s uch circum s ta n ce s a nd ,

s o giv e i n direct evide n ce of the c l imatic co n ditio n s

prevale n t i n the two period s .

 CH A PTER V

B I OL OG I CAL PALA EON TOL OGY

N the s m all s pace av a ilable it i s impo ss ible t o d i s cu ss ,

or eve n to catalogue the variou s biological problem s

,

toward s who s e s olutio n Pal aeo ntology ca n co ntribute .

S i n ce detailed refere n ce to the m a ny bra n che s of s uch

re s earch i s im practicabl e a n attem pt h a s bee n m ade to
,

give a ge neral accou nt of the broad outli n e s of phylo

ge netic evol utio n . Lo n g a n d ted iou s s tudy will be
n ece s s ary before ma ny of the propo s itio n s h e re m ade

ca n be proved or di s proved ;a ny s timulu s to s uch e ffort ,

wi t h either i nte n t that m ay percha n ce proceed from

,

thi s chapter would am ply j u s tify i n clu s io n of theoretical

,

m atter i n a n i n trodu cto ry e s s ay Th e whole chap ter i s

.

i nte nded to co nvey hi nt s a s to l i ne s of i n ve s tigatio n

th at ca n not fail t o p rov e profitable ; i t i s i n n o s e ns e
a n epitom e of all that h a s b ee n achieved towar d s e luci

datio n of evol utio n al problem s .

( )
I P H Y L OG E N Y AN D MO R P H O G E N Y
The s tratigraphical record who s e i nterp retatio n is
,

ultima t ely ba s ed upo n the i n co ntrover ti b le e vide nce of

“

s uper po s itio n repay

, s i n ful l the debt it owe s to p a la e on

t o logica l pri n ciple s si n ce it s uppl ies a hi s t orical se que nce

,

for the fau na s of s ucce s s ive period s I t i s p o ss ible to

“

t r ace the directio n a n d progre s s of evol utio n i n i ts O pe ra s

t io n through va s t age s of tim e with co nfide nt k n owledge

tha t the order of eve nt s i s e s tabli s hed I n s pi t e of the
.

9°
 92 IN VE R T E B R ATE PA L A EONT OLOG Y

Be s ide s com pl icatio n p roduced by geol ogical im per

fe ct io n s i n tri ns ic d i fficultie s ob s cu re the tre nd of e volu
,

tio n Like a river -s y s tem flowi n g over a broa d plai n

.
,

the s tream of orga n ic progre s s is no t con fi ned with i n

o n e cha n n el n o r d oe s it follow a d irect cour s e
,
I t is no
.

ea s ier to ide nt ify the m a in cu rre n t ( if s uch ca n be s aid

to exi s t ) i n a delta tha n to d i s ti ngui s h betwee n the
,

d iverge ncie s a n d co n verge n cie s i n ev o l utio nal h i s to ry .

A ge n eral d irectio n of progre s s ca n be a s s u med in the

one ca s e n o le s s tha n i n the other ;bu t the bour n e to

which evol utio n i s ten d i n g h a s n ot yet bee n attai n ed ,

i t s s ource i s s omewhe r e i n the ter m z mogmta of the

' '

rem ote pa s t while the com plexitie s reveal ed by eve ry

,

gl im p s e available dema n d clo s e a n d d etailed s tudy b efore

their relatio n to the m ai n s y s tem ca n b e appreciated .

The gu idi ng force that d etermi n e s the flow of a river is

t hat of gravitatio n to that pr i n ciple al l ob s erved feature s
ca n be referred a n d by it u n s e en qualitie s ca n be con
,

j e c t u r e.d But what i s the co n troll i n g age n t that s teer s

the cour s e of orga n ic evolutio n ? Th at one exi s t s i s

beyo nd d oubt ;it s n ature a nd operatio n are a s yet but , ,

d im ly vi s ible .

J u s t a s a group of orga n i s m s i s the s u m o f i t s i n di

v i d u a ls s o a n
,
“
i nd ivid ual i s the s um of it s organ s
”

A t a ny on e ti m e a group m ay i n cl ude precocio us a n d

,

backward mem ber s ;eve ry i n dividual po s s e s s e s s tr uc

tures i n va ry i n g s tat e s o f e ffi cie ncy . T h e qua lit y or
’

s tage o f evol utio n attai n ed by a stock m u s t be cal e u

‘

~
,

l ated a s the average for it s com po n e n t s ab o ve a nd below

,

which the relatively s carce progre s s ive a nd r egre s s ive

member s flouri s h or la ngui s h . A group i s in the
a s ce n d an t if the progre s s ive member s out n um be r t he

regre s si ve ;it i s dec ade n t if the proportio n s are rever se d ,

for the average m u s t te nd to m ove toward s the p re

d o m i na n t extrem e The d egre e of s ucce s s reached by
. .
 B I OL OG I CAL PALAE ON T OL OGY

an i ndivid ual m ay b e mea su red by the relative harmo ny

with wh ich it s com po ne n t s tructure s collaborate H e n ce .

a high s tage of elaboratio n i n o ne particular characte r

m ay ofte n prove i n co nve n ie n t or fatal to a n orga n i s m if ,

it fail s to co n form to the fu nc tio n al requi re me nt s o f le s s

prog re s s ive feature s The n ame i nd ividual thu s prove s
.
“

m i s leadi n g whe n appl ied to a n orga ni s m s i n ce althou gh , ,

the pr es e n ce o f all it s s tru cture s m ay be n e ce s s a ry for

l ife ( a n i nfreque nt a n d da ngerou s co n ditio n) each one of ,

i t s orga n s ca n have s om e i n depe nde n c e of developme nt

“ ”
duri n g i n dividual growth a n d evolutio n al hi s to ry .

F rom the s ta n dpoi n t of evol utio n it i s e s s e n tial t o,

”
regard a n i n d ivid ual a s d ivi s ible i nto i t s s everal
“

s tructu res .

The orga n i s m is t he unit of the group a n d the orga n

'

“ ”
the u n it o f the i nd ividu al ;the rel atio n of the two
u nit s to their re s p ective com pou nd s i s i n s ome mea s u re
d i s s imilar. Thu s it follow s that s tudy of race -hi s tory
a nd that of s tructural evolutio n m u s t be i n s ome d egree

i nd epe nde n t H e n ce the two word s i n the title of t hi s

.

s ectio n . Phyloge ny i s ra cial evolutio n m orphoge ny that ,

o f s tructure .The two s tudie s are as i n separa ble a nd

.
,

a s d i s ti n ct a s orga n i s m s a n d their compo n e n t or ga ns

,

R ace -hi s tory ca n be traced o n ly through the s e que n ce

of i n divid ual s ;progre s s or rev er s io n i n the l atter is i n di
ca t e d by the qual itie s of their m orphological characte rs .

I t follows that morphoge ny m u s t s upply the bas is on

'

which ph yl oge ny ca n be reared S imilar relatio ns

.

exi s t betwee n Morphology a nd Taxo n omy S y s tem .

atic clas s ifi catio n mu s t b e fou n d ed o n morphological

di s covery ;phyl oge n etic s erie s ca n n ot be traced u ntil
morp hog
’

e n etic tre n d s have bee n recog n iz ed But it i s .

i nex p edie n t i n either ca s e to await completio n of t h e

, ,

fou ndatio ns be fore erecti n g the s upers tructure Pro .

vi s ion al cl ass ificatio n s ubj ect to co n ti n ual rev is io n i s

, ,
94 IN VE RT EB R A TE PALA E ON T OLO GY

n eces s ary if morphol ogical re s earch i s to be co o rdi nated

a n d m ade i n telligible ;te n tative s chem e s of phyloge ny

s erve a s co n ve n i e n t if i mperfe ct corr e latio ns of e s t a b

, ,

lis h e d data i n m orphoge ny .

O Pl vii fi — ttempt i m ad e to i dicate

n .
g s
. 2. 1 3 a n a s n

the n ature o f a m orphoge netic s eque nce a nd to s how ,

i t s relatio n to phyloge ny The apical sy s tem s of

.

I rregul ar E chi noid s u n dergo di s orga n izatio n ( i n o nto

g e n y a n d morphoge n y ) by retrogre s s io n o f the periproct .

The po s terior ge n ital plate i s re s orbed ; the gap i n

the cycle a n d the ce n tral area o n ce occupied by the
,

periproct have to be fi lled F igs 2 a n d 3 s how two

,
. .

D i a d e moi d ( R egu lar ) type s of apical s y s tem ;i n both

the backward te nde ncy of periproct -m ovem e n t i s a p
c i a b le b ut d oe s n o t d i s turb the ge n ital a n d o cular
p r e ,

ri ng s ( The alter native arrows co nn ecti n g the s e figure s i n

.

d ica t e u n certai n ty a s to the m orphoge netic relatio n of

D i a d e mi ne a n d C alyci n e s ys tem s ) I n P les z ecfi i rzu s ( fi g
'

. .

the m orphoge netic ( a nd appare ntly phyloge netic )

a nce s tor of I rregula r E chi n oid s th e peripr oct occupie s ,

the po s te ri or part of the s y s tem breachi n g the cycle ,

a nd d i s placi n g the s urvivi n g pl ate s F rom thi s con .

ditio n two mai n tre n d s o f recove ry ca n be traced I n .

that s how n by the s eque n ce of figure s 9 —1 I n o attem pt ,

i s m ade to re s tore the lo s t ge n ital ;i ts place a n d the ,

ce n tre o f the s y s tem a re fill ed by exte ns io n o f th e

,

m ad reporic plate A s l ight m odificatio n of thi s m ethod

.

fi
( g .s 1 2 a n d I 3 ) clo s e s up the s y s tem after the pe r iproct

h a s pa s s ed to the rear prod uci n g i n i ts e xtrem e pha s e

,

a n el o n gate apic al s y s tem with alter n ati ng pl ates The .

s ec o n d tre n d o f recove r y is s how n i n fi gs 5- 8 H er e . .

a ne w pl ate i s formed i n the w a ke o f the r ecedi n g

periproct i n cou rs e o f time it a ss u me s the proportio ns,

a nd fu n cti on o f the m i s s i n g ge n ital The cen t re of .

the s y s tem is filled by i nwa rd exte ns io n of the ge ni t a l s ,

9 6 INVE R TE B R AT E PALA E ON TOL OG Y

s t ruct ure s , a nd al mos t a ll are m i n ute that p re se rv a ti on

so

a nd i s olatio n are u n likely F urther i n the rare ca s es

.
,

whe n early devel opm e ntal form s a re d i s covered it i s ,

u s ually i mpo s s ible to d eterm i ne the relatio ns hip b etwee n

larva a nd ad ult Si n ce it i s too l a te to watch the g ro wth
,

of o ne i nto the other N everthele ss Palaeo n tology ca n

.
,

m ak e us e of certai n o n toge n etic qualitie s i n a d ult or

a dole s ce nt organ i s m s that withi n re s tricted l imits give
, ,

a ki nd of re capitulatio n .

There are two law s of s tage s of developme n t reco g

n iz e d
, but they d iffer chiefly i n applicatio n a nd m od e
“ ”
of expres s io n O ne the law of s tages i n o n toge ny
.
,

( H yatt , ) i s merely a n exte n s io n of that of recapitulatio n .

I t s tate s that i n you n g form s s tage s Occur which cor ,

r e s po n d with the ad ult s tage s of a n ce s tral type s ;while

i n s e n ile i n d ivid ual s s t age s occur which m ay repeat

, ,

i n revers e order tho s e fou n d i n the you n g a n d are i n

, ,

s om e mea s ure pr ophetic of comi n g cha n ge s i n s ub s e

quen t evolutio n o f the group co n cern ed E xpre s s ed

.

i n a s im pler form the law s tate s that recapitulatio n

,

co n ti n ues till m atur ity a nd i s followed by a n ticipatio n

,

o f future ch a n ge s i n the s tock H e nce it foll ows that

.

s tudy of the com plete l ife hi s tory of a n i nd ivid ual

-

s upplie s a n abrid ged e pitom e of th e hi s to ry of th e

group t o which it belo ngs The other law that of
.
,
“ ”
s tages i n m orphoge ny ( u s ually a nd o r igi n al ly called
the la w of l oca l ized s tages i n develop me n t J ack s o n )
“
,
”
,

s tate s that throughout the life of a n i ndivid ual there ,

occur i n local s tructure s s ta ge s com parable with tho s e

fou n d i n youth o r s e n il ity s upplyi ng evide n ce on raci al

,

evol utio n s im ilar to that a fforded by o n to g en etic s tage s .

I n other word s t he growth o f s eparate s tr ucture s or

,

orga n s follow s recapitulatory o r a nticipato ry cour ses in

s ome r e s p ects i n dep e n de n t of the age of the i nd ivid u a l .

Both o f th ese hypoth es e s a re c a pable of appreciatio n ,

 B I OL OGI CA L PALA EON T OLOGY 97

a nd ve rificatio n in pa l aeo n tological m aterial Although

, .

ful l s eri es of growth - s tage s of s pecie s ca n rarely be

collected a l arge proportio n of I n vertebrate s retai n
,

the p roduct s of their early growth throu ghout life s o ,

that a s i ngle s pecim e n ca n by d i s s ectio n or a naly s i s

, ,

b e m ade t o yiel d evide n ce of i t s o n toge n y Thu s .

A m mo n ite s added s ucce ss iv e whorl s duri n g l ife b ut ,

did not des troy the fi rs t -formed part s of the s hell By .

”
u n wrappi n g a n Am m o n ite ( removi n g o ne chamber
at a time) the whole po s t -l arval hi s tory of s hell growth -

i s gradual ly revealed i n i n ver s e order u n til the proto

,

co n ch i s reached S im ilarly Ga s tropod s d i s pl ay their

.
,

s ucce s s ive on toge n etic character s although the expo s ed

,

po s itio n i n whi ch the early wh orl s are u s ually left ( at

the apex of the s pire ) ofte n re s ult s i n dam age or removal .

Pele cypod s a n d Brachiopod s s how i n the u m bo n al

regio n s products of early s he l l formatio n a nd m ay
-
,

oc cas io nally retai n their p r o d i ss oco n ch s or p r ot e gu la .

The ba s al part of the theca of a coral h a s the fir s t

s im ple arra n geme n t o f s epta a nd di s s epime n t s while ,

more adva nced s tages of complicatio n ca n be traced

upward s toward s the calyx .

Po s t -l arval o n toge n y i s reple t e with d i fficultie s l ike

tho s e that compl icate embryology The s h ort time
.

avai lable i i ndividual l fe f o

n i r
“
revi io
s n -
”
of age lo n g
evol utio nal hi s tory re s ult s i n compre s s io n om i s s io n a nd
, ,

eve n d i s turba nce o f s eque n ce of s tage s pa s s ed thr ough

, .

D evelopme n t s of late i ntrod uctio n m ay be im pre s s ed

upo n s tructure s of prim itive s impl i city Thu s while
.
,

the u s ual typ e of Ga s tropod protoco nch i s a s mooth

, ,

s li ghtly coiled s hell feature s of or n ame n t characteri s tic

,

of more adva n ced s pecializatio n may occur upo n it

‘

as h a s bee n well s how n by Graba u i n the F us id a e .

‘
But i n s pite of com plexitie s the pri n ciple of o ntoge n etic
,

s tage s gives ge neral ly s ati s facto ry i nd ica t io n of phyletic

9 8 I NVE R TE B R AT E PALAEON TO LOG Y

evoluti o n Beecher pr oved i ts val ue i n grea t nu m ber s

.

of Brachiopod s a n d Trilobite s while m a ny author s have ,

a pplied it to the s tudy of A m mo n ite s Ga s trop od s a n d ,

Pelecypod s F au nal s ucces sio n i s available to co nfirm

.

or di s e n ta n gle the record s of po s t l arval o nt oge n y

-
.

I n s everal I n vertebra t e grou p s n otably the E chi no ,

derm ata ne w s tru cture s which are practically repetitio ns

,

of tho s e already pres e n t are developed throughout

life I n Cri noid s the s tem i s le ngthe ned by ad d itio n
.
,

of ne w co lu mn a ls ;i n E chi n oid s the coro n a i ncrea s e s ,

i n s ize pa rtly by i n trod uctio n of fre s h plate s arou n d

th e m argi n of the apical Sys tem O rga n i s m s with s uch .

m ode s of growth will alway s po ss e s s s o me you ng “

“ ”
s tructure s however
,
old the i nd ivid ua ls m ay be .

F or exam pl e the co m m o n H ol ectypoid P les zec/z i nus

'

, ,

am a tus ( Pl v u fi g . 1 ) s how s certai n coro n al plates a t

. .

t he apex that are s mall s im pl e a nd i n cour s e of develop

, ,

m en t . Thes e m o rphoge n etically n ea n ic o s s icles a re ,

i n m o s t re s pect s s imil ar t o the corre s po ndi ng s tructure s

,

produced i n early s tage s of i n d ivid ual o ntoge ny .

_

the ti me o f their i n troductio n o n ward s such pla t e s ,

progre s s i n growth a nd elaboratio n u ntil t hey atta i n

a degree o f s pecializatio n harm o n izi n g with the o n t o

ge netic s tage of their ow ner S tudy of the i n t er .

a mbul acrum of a n adul t P les i ec/zz n us s how s that the

'

plate s n ea r the p eri s tome which date from th e early ,

youth of the s pe cime n are s m all a nd s uppor t few , ,

tubercle s They repre se n t the s ligh t adva nce of mor

.

h o ge n e s is corre s p o n d i n g with the s hor t progre s s m ade

p t

i n o n toge ny at the tim e whe n t hey co n s tituted th e .

whol e a re a A s the colu m ns are traced away from

.

the p e ris t o me th e s ize a nd tubercular or na me nta tio n

, ,

of the plate s s how s teady i n crea s e u n til at the m id , ,

o
z n e ( hal f-way u
p the coro n a Pl ate they exhibit ,
‘

the highe s t pha s e of m orphological com plexi t y p rope r

I OO I N V E RTEB R ATE PALA E ON T OL OG Y

i ts ac m e O n the o t her ha n d if the m id -zo nal plate s a re

.
,

le s s elaborate tha n s om e n earer the peri s tom e the s pecie s ,

i s regre s s ive i n th at the cul m i n atio n of m orphoge netic

,

develop me nt i s reached a n d pa s s ed before the a n i m al is

, ,

fully grow n Great care i s n eed ed i n i nterpretatio n of

.

r ecord s o f thi s type s i nce

, s eco n dary s pecializatio n

due to l ocal requireme n t s ofte n ob s cure s the origi nal

s t ructure B ut with d ue p recautio n s tudy of m or
.
,

p h og e n e t i c s tage s of developm e n t m ay ofte n lead

to d ed uctio ns that ca n be verified by i ndepe n de n t
evide n ce .

( )
I I I S P E C I A L I Z AT I O N A ND R E G R E S S I O N

A rgui n g from the ob s erved feature s of o ntoge ny it i s ,

po s s ible to co n ceive tha t a group of a n imal s at i ts i n ,

ce p t ion m ay be capable of im proveme n t the s tage of

, ,

perfectio n bei n g attai ned whe n ge neral e fficie ncy i s

acqu ired But it i s im po s s ible t o mai ntai n s eparatio n
.

betwee n the orga n i s m s a n d their e nviro n me nt O rga ns .

tha t would do good s ervice i n r ocky s urrou nd i n gs woul d

probably prove worthles s o n s a n d or ooze ;a nd s i n ce ,

group s are liable to cha n ge s of habitat either of thei r,

o wn accord or owi n g to phy s iographical i n flue n ce the ,

efl i cie n cy of on e period m ay becom e the imperfectio n of

a n other . The E chi n oidea m ay be cited i n ill u s tratio n
o f thi s ph e n ome n o n A l arge proportio n of the Pal ae
.

“
oz oi c S e a -Urchi n s had flexible te s t s appare n tly ,

s u ited for l ife a m id yieldi n g materia l s ;s om e m oder n

m ember s of the group with s im ilar habitat show

, ,

corre s po ndi n g characters D uri n g Tria s s ic tim e s the

.
,

E ch in oid s s eem to have cha n ged their m ode of life ,

cho os i n g reefs a nd rock s a s their e nviro n me n t E xpo s u r e .

t o wave s d em a n ded s t re ngth a n d ri gidity of fabric a n d ,

Pcris choe chi noi d flexibili ty was a ba n do ned Thr ough

”
.
 B I OLOG I CAL fp hizs noisif otb é ? 101

ou t the Mes ozoic a nd Cai nozoic era s s teady im prov e ,

ment s al o n g vario us l i n e s were i ntrod uced u ntil at th e

, , ,

pre sen t day s uch form s a s the E ch i n ome t r id ae m ay be

co idered thoroughly efficie n t for rock dwelli n g habit s
n s - .

B ut s oo n after thi s cou r s e of evol utio n had bee n e ntered

upo n s ome group s of E chi n oid s reverted to the s a n d
,

habitat of their a nce s tor s The s pecializatio ns for r eef

.

life i n itiated a nd elaborated by the on e s eri es would have

bee n u s ele s s a n d eve n da n gerou s for the other s o that
, , ,

a grad ual retur n to qualitie s like tho s e of Palaeozoic

type s wa s begu n I f racial progres s is to be m ea s ured
.

by the degree of s ucce s s attai ned i n adaptatio n of a n

orga n i s m to it s e nviro n me n t Colobocen tr otus of the ,

s outher n coral -reefs a n d E ch z noca r d z u m of the Briti s h

' ’

sa nd fl - a t s are equally progre s s ive

,
B u t i n s tructure the
.

E ch in ome t r i d a e are far m ore elabo rate tha n Palaeozoic

E chi n oid s while the S p a ta n gi d a e with all their com
, ,

“
l
”
p e x i t y s,
how m a n y Pe r i s ch oe ch i n o i d featu res Morpho .

ge netically the former fam ily i s progres s ive ( a n age n etic) ,

th e latter r e gre s s ive ( catage netic ) i n e fficie n cy both are ,

equal The s olutio n of s uch a n a n om aly is probably to

.

b e fou n d i n recog n itio n that choice of e nviro n me n t i s a

character i n dicative of evol utio n al activity n o le ss tha n ,

s tructu ral s pecializatio n The prefere nce s how n by

.

Lia s s ic I rregul ar E chi n oid s for m uddy or s a ndy

s urrou n di n g s wa s a n expre ss io n o f their catage netic
te nde ncy ;m orphoge netic regre s s io n followed i nevitably
after that backward s tep .

I n the di s tributi o n of l ivi n g orga n i s m s there is a

pro n ou n ced te nde n cy for moribu nd or a n tiq uated type s
t o avoid po s itio n s i n volvi n g expo s ure to d a n ger or com
petitio n Thu s the s tal ked Cri noid s ma ny Brachiopo d s
.
, ,

a n d Pleu r otoma n a n ow exi s t i n the s heltered s eclu s io n

'

of deep-water although their m ore vigorou s a nce s tor s

,

j oi n ed s ucce s s fully i n the s trug gle for life i n littoral

1 02 ~
'

I NV Q
ERTEBRATE PA LA EONT OL OGY
zo n es S im ilarly Co n i fer s a nd Lycopod s for the mo s t
.
,

part have aba ndo ned com petitio n with A n gio s perm s
, ,

a n d prefer to brave the eleme n t s o n m ou n tai n t op s or i n

high latitudes With t hi s evide nce an d by a n alogy

.

with i nd ividual l ife i t s eem s rea s o n able to s uppo s e that

,

race s of orga n i s m s dem a n d s om e m ea s ure of s helter

d uri ng early a nd late s tage s of their developm e nt a nd ,

j oi n i n the tu rm oil of robu s t activity o n ly whe n racial

vitality is at it s height Choice of a s hel tered e n viro n
.

m e n t by a gr oup who s e a nce s tor s flouri shed am id

com petitio n is equivale n t to co n fe s s io n of failure ;
morphoge n etic cha nge s thu s i n duced are rightly con
s id e re d r egr e ss ive however they m ay ha r m o n ize with
,

the ne w s u rrou nd i ngs D evelop me n t of ad ipo s i ty or

.

arthriti s is em i ne ntly app r opria t e for i nd ivid ual s who s e

chief des i re i s for re s t ;the di s ea s e s a nd the de s ire are
bo t h sym ptom atic o f old age a nd com i n g d i ss ol utio n .

S pecializati on ca n therefore be defi n ed a s mod i fi ca

, ,

ti o n of s tructure co nducive t o s ucce ss of a n orga ni s m i n

the m o s t t re nuou s s ectors of the battle grou n d of life ;
s -

regr es s io n a s that co n n ected with retireme n t to s hel t ered

s u rrou n di n g s Both tre nd s of evol utio n m ay be la rgely

.

adaptive although both are probably i n flue nced by

,

i ntri ns ic qual itie s of racial ri s e a nd decay The d e fi ni .

tio n i s n either com plete n or exclu s ive but i t s erve s a s ,

a u s eful d atu m whereby evol utio n al tre nd s m ay be

pr o vi s io nally correl ated .

The A m mo n ite s have l o n g bee n recogn ized a s a

group a ffordi n g a n u nu s ually complete a n d obviou s /

exam ple of racial life h i s to ry Al though there is room

-
.

for di ffere n ce of opi n io n a s to the i ntri ns ic or ad aptive

qual itie s of the var iou s m od ificatio ns fo und i n the group
the ge neral s eque nce of s tructural cha nge i s s o regular
a nd s o o ft -repeated that i t m u s t be a s cribed to s om e
,

(
n
fu dame tal n te n de n cy The s t r aight
..s mooth m a n
,y ,
m4 i NVER TEBR ATE PA LA E ON TOLOG Y
t io ns bu t failed to a chieve or m ai ntai n e nrolm e n t
, , ,
.

Thi s type of revers io n cul mi n ated i n the Cretaceou s

B aculi tes , which , apart fro m i n tern al detail s wa s ,

e s s e ntially a s eco n dary B actritico n e O ther grou p s ex .

,

e mp li fi e d by the Pulch e llii d ae of the Cretaceou s peri od ,

“ ”
retai n ed the A m m o n itico n e i nvolutio n bu t allowed ,

their s epta to revert to a d egree of complicatio n hard ly

”
greater tha n that of t he Go niatite s F or exam ple the .
,
'

s uture-l i n e of Tis s otzcz from the Ce n om a n ia n i s hard ly

, ,

to be d is ti ngu i s hed from that o f Cer a tz zes from the

'
‘
,

M u s chelkal k A lthough either of the s e regre ss ive de

.

ve lop me nt s m ay have bee n a s s ociated with e n viro n m e n tal

i n fl ue n ce thei r appeara n ce s eem s co ns ta n tly to have

,

preceded exti n ctio n of the group s a ffected Whether .

s uch group s peri s hed i n the Tria s or per s i s ted u n til th e ,

fi nal collap s e of the cla s s in the Cretaceou s period they ,

u s ual ly di s played i n thei r l a te s t m embers revers io n

, ,

toward s a nce s tral feature s The co nd itio n h a s bee n aptl y

’

”
de s cribed a s s eco n d -childhood I t m u s t be ad m itted
.
,

however that certai n appare n t exceptio ns to thi s rule

,

are k n ow n F or exam ple I nd orer rzs a n U pper Cre

.
, ,

t a ce o us ge n u s from I n d ia i s n o t merely a n
,
A m mo n ite
i n s hape bu t h a s s epta which i n s om e pa r ti cu lar s rival
,

tho s e of the m o s t elaborate Tria s s ic types The s ign i fi .

ca n ce of thi s a n d s imil ar ca se s is ope n to doubt ;but ,

i n s pite of exceptio ns regre s s ion duri n g d ege neracy

,

se em s a defi n ite pri n ciple o f evol utio n .

Two com mo n feature s o f s pecializatio n i n crea se i n ,

s ize a n d additio n o f s hel ly m atte r are readily t raceable ,

i n fo s s il m aterial The form er qu ality i s n ot s o m arked

.

a mo n g I nvertebrate s a s am o n g R eptile s a nd Mam m al s

‘

e s pecial ly i n the c atage netic s tage s of i t s over d evelop

-

m e nt the latter i s very clearly s hown i n a great variety

of group s Both te n de ncie s s eem particularly liable
.

t o per s i s t beyo n d the s tage i n wh ich they m a ke for

 L
P ATE V I I

15 3 32
 PL A T E V II I
PH Y L o G EIfloN faC C H A RA CT E RS
’

Pa r ézm ow ( much rs d goszd l 111133t

[d ow e/e w es
l Burm a 1

3g, Bra d s t Q Ck D o rs e t The d n ne r ( e arly) whorlsf a re s trong

, . . ly

a u
r i b b e d fi e ro i . l
gt g aa yig
e a t i op ob li t e ra t eg3 gp r a m e n t i n t h e

m rer whq s he -
a nd ; 1a p u .

; m ] 1 1 1 1 5 111 1 1
3
1

Claw
H/d léfi '
a va “1
13511011 , H a fifi d n f
l he
‘

p r
otocon ch

i s fo llo we d b y on e wh o r l ’
s h Ow i ng
‘i
F fi si tf
’W‘
ifross o rn anw n t .

1b m d d fi Thf3 tW0
r
1
n e xt whorls an
v e s

ri ly
moo thefl b y gfi
w n ic
fi b e vi
ga nonn thc
fi
4 m 11 1
.

{
est
beco me . seco n d a ,
sp i nous
w b y t exces s i ve
q ca lci ca ti on

( bag mi
’
. ed 01 ;Gxab a u s fi gurel m )
,
4 ,

61 1 1 43 51 mi la r to

W é i aefi rgkt i on .

Q hr
g
o ngf
re r ou s
j
i me L
s to n e . S h o wmg . g e r ont c
i . fea tu e s r ,
np fi g
i n’ fl r li c t i on o f t h e

g l be ll
a a , mu lt i p le pygi d mm , a nd
p u s t u la r o r n a me n t .
1 06 I N VE R TE BR ATE PALA EON T OLOGY

it s acm e a nd the n de sce nd s alo n g a cour s e rem i n i sce n t

,

of that by which it r o s e dyi n g of old age after a pha s e

,

of eco d childh ood whe n i ts vitality is exhau s ted I n

s n -
, .

t he othe ca s e over s pecial izatio n co nti n ue s to p rod u ce

r -
, ,

e ffect s which after t he acm e h a s bee n pa s s ed becom e

, ,

po s itively d i sadva n ta geou s i n hypertrophy howeve r ,

profitable they m ay have bee n in re s trai ned develop

“ ”
me nt B ut thi s s u icidal te nde ncy i s no n e t he l es s
.

com parable with phe n ome n a ofte n s how n i n i ndividual

s e n e s ce n ce Morb id growth s either i n bulk or s clero s i s
.
,
,

freque n tly precede a nd i nd uce death ; s o t hat t he

, ,

a n alogy be t wee n racial a nd i n d ividual life is s till

t e n abl e.

( V)
I A C C EL E R A T I ON A ND R E T A R D AT I O N
The rate at which s pecializatio n proceed s varie s t o a
m arke d d egree i n d i ffere n t group s o f a n im al s a nd is n ot
‘

c on s t a nt duri n g the evol utio n of a s i n gle s erie s ;it m ay

d i ffe r i n the s everal part s of a n i n d ivid ual orga n i s m .

S om e s tock s s eem bor n t o s tag natio n others to progre s s , .

The r acial hi s t ory of the former type s te nd s to b e lo ng

a nd u neve n t ful that of the latter s hort a nd crowded
,

with m orphoge netic epi sod e The co nt ra s t coul d hardly

.

be d i s played m ore clearly tha n i n the ca s e o f the

Tetrabra n chiate Cephalopod s al ready d i s cu s s ed The .

N autiloid s l ike the A m m o nite s origi nated a s fo r m s

, ,

with s traight chambered s hel l s which have s how n

,

a s teady t end e n cy toward s e n rol m e nt fr om Lower

Palaeozoic t i me s o n ward s B ut wherea s t he A m mo noi d s
. .

attai n ed the acm e of their s pecializatio n i n the T ria s ,

a nd ha s te ned to de s tructio n i n the Cretaceou s pe riod ,

the N autiloid s have n ot yet r eached a degr ee of el abora

tio n comparabl e with tha t of a n A m m o n ite a nd s how no
“

s ig n s of a s pira t io n t o s uch a n ide a l Although s om e .

 BI O L OG I A L C PA LA E ON TOL OG Y

amo unt o f en rol me nt wa s developed i n tha t co ns erv ative

group as early a s the C arbo n i ferou s ( while s tro ng i nvolu
tio n h as cha ra cter ized N a u tilus s i n ce J ura s s ic time s) the ,

s epta u s ual ly retai n great s impl icity N ot u n til the .

C ai n ozoic d o co rrugatio n s com parable with tho s e of

D ev on ia n “ ”
Go n iatite s affect the N autiloid s eptum ,

a nd eve n thi s s m al l deg ree of compl icatio n h a s be e n

aba n do ned by l ivi n g form s The d i ffere n t s peed of
.

evolu tio n i n the s e pre s u mably homoge netic cla ss e s i s

as s ociated with i mporta n t res ul t s from a s tratigraphic al
poi n t of view The A m mo n oid s appear i n the D evo nia n
. ,

a n d di s appear i n the Cretaceou s attai n i n g i n their r ela

, ,

t ive ly brief career a cma ic prom i ne n ce of a n extrem e

,

t ype The N autiloid s aro s e i n the Cambria n ( or before)

.
,

a n d are s t ill livi n g but they ca n n ot compare with their

,

m o re progre s s ive relatives i n s pecific va riety or n u merical

a bu nd a n ce .Their delibe rate a nd u n a s piri n g progre ss
may be correlated with their early i ntroductio n I t i s .
'

us ual for the le ss -s pecialized a nce s tral s tock s of group s

,

that have produced progre ss ive bra n che s t o pers i s t lo ng

after their preco ciou s offs pri ng have falle n from the
a cma i c heights The Atrem atou s Brachiopod s are
.

repre s e nted to day by s pecies of L zngu lcz rem arkably

’

like m a ny of tho s e prevale n t i n O rdovicia n s ea s ;but

the O rt h a ce a St rophome nacea a n d S piriferacea group s
, ,

whic h s eem certa i nly to have had a n Atrem atou s

a nce st ry have had their day a n d qui te or al mo s t
, , , ,

cea s ed to be The E chi n oid family of the Cidaridae

.

i s repres e n ted by clo s ely s i milar form s i n the Lia s a n d

the H oloce ne but m a ny of the J ura s s ic a nd Cretac eou s
,

types devel oped from the Cidarid s tock a nd of far

,

greater mo m e n t i n their time have bee n lo n g exti nct

,
.

Al mo s t every phylu m ca n s how i t s co nservative s a n d

progre s s ive s ;while the practically s tatic qual ities o f
tho se Protozoa who se palaeo n tological hi s t ory ca n b e
1 08 I N VE R T E B R ATE PALA E ON T OL OG Y

traced s eem s to i n d icat e t hat t he pri n ciple a pplie s

equally to l arge a n d s m al l grou p s of orga n i s m s .

Te n de ncy to s pecializatio n appea rs t o vary i n i nte n

s ity a nd s cope at di ffere n t s tage s of racial evol utio n
'
.

There are three s tage s d uri n g which rapid m orpho

ge netic acceleratio n typically occur s coi ncid i ng fairly ,

cl o s ely with the youthful m ature a nd s e nes ce n t pha s e s

,

of race -life .O f the s e the fir s t i s the m o s t s tro n gly

,

m arked a n d i s u s ually productive of the m o s t far

,

reachi n g i nn ovatio ns Whe n the geological ra nge s of

.

the s everal cla s s e s of a phylu m are com pared i t is ,

com m only fou n d that wh ile the period s of acme or

.

exti nctio n m ay be very d iver s e the tim es of i ncep t io n

,

follow very s hortly after di ffere ntiatio n o f the whole

grou p I n the ca s e o f m o s t I n vertebrate phyla thi s
.
‘

quality is ob s cured b y i t s pre -Cambria n remote n e s s ,

but am o n g V ertebrate s i t i s very appare n t The rich .

variety of the Ca mbria n fau n a ca n d oubtle s s be a s c ribed

to i ts operatio n That the s ame te n de n cy work s th rough
.

s m aller d ivi s io ns i s abu n da n tly m a n ife s t I ndeed t he .

,

rarity of ca s e s where grad ual tra n s itio n from on e fam ily

to a nother ca n be d em o ns trated u s ed to s upply s ome
of t he mo s t powerful argu m e n t s of Oppo n e nt s of the
evol utio n -theory H owever a n alogy wi t h i n dividu al l ife
.
,

would m ake s uch nea n ic acceleratio n po s tulated we re it ,

n ot actually ob s erved The early on toge ne t i c p h a se s o f

‘

a n orga n i s m are pl a s tic a n d protea n a s on e s tructure ,

after a n other i s prod uced a nd developed ;relati ve s tability

characterize s maturity .

A m o n g the m a ny i n s ta n ce s of rapidity o f s p eci a li z a

tio n i n you n g group s th at s how n by the E chi noidea i s
,

well m arked A fter a s erie s of experime n t s in Palaeozoic

.

tim e s the cla s s acquired a certai n fi x ity o f character

,

i n the Tria ss ic Cidaridae ( The P i h hi i d i

‘

e r s c. oe c n o e a n ,

their ma n ifol d a nd extraordi nary varietie s ca n perh a p s ,

I Io IN VE R T E B R AT E PAL A E ON T OLOGY

progres s a t tai ni n g i ts prime after a n i nt erval l o nger or

,

s horter tha n that O f i t s fellow s ;but there i s com mo n ly

s om e mea s ure O f s im ilarity i n the quality O f s tructural

evol utio n in a ny homoge n etic s erie s S uch corre s po n d

.

e nce h a s bee n i n dicated above i n th e ca se of the Tetra

bra nchiate Cephalopod s The N autiloid s i n s pite O f the
.
,

extrem e deliberatio n of their progre ss pur s ued li nes of

,

s pecializatio n which ca n be correl ated with tho s e p re cip i

t a t e ly followed by the A m m o n oid s . The pri nciple o f

paralleli s m of developme n t i n hom oge netic form s ( ortho
ge ne s i s) s eem s applicable to al l grade s of cla s s ificatio n .

Ca s e s are not wa nti ng where thi s m orphoge netic corre

s p ond e n ce i s empha s ized by i ts co n tempora n eou s occur

ren ce i n d iverge n t s tock s A m o n g the E chi n oidea s everal

.

s uch phe nome n a occur . The H olectypoid ge n u s F i lms

a nd the D i a d e moid fam ily of D i p lop od i i d a e — type s ulti

m ately hom oge n etic but s triki ngly d i s s i m ilar in s truct ure
,

a n d probably i n habitat — both s how the u n u s ual feature

o bi erial pore arra n geme n t a d a p i ca lly a n d both aro s e i n
f s -
,

the Upper Ju ra ss ic Agai n the cu riou s form L a mer za a n

' ’

.
, ,

H olectypoi d with m a ny a ffi n itie s to H olec s acquired ,

i n Upper Cretaceou s time s a type of am bulacral s tructure

ide n tical with that of the co ntem pora n eou s Con ulus wi th ,

which it h a s few other feature s i n com m o n Thi s ca s e .

i s m ade m ore s trik i n g by the occ urre nce i n the Lower

Cretaceou s o f the L a n ieriid ge n u s D is c/zolea jypu s with ,

D zs ck olectypus ca n hardly
’

s imilar a mbulacral plati n g .

be a nce s tral to L a ni er i d but i ts develop me n t of Comd us

,

like s t ructure coi ncide s i n time with the origi n of the

Py n n s tock from which Conulus i s cle arly d e s ce nded
-
’

a ,
.

The N ucle olitoid ge n u s Tr ema topygm who s e phyletic ,

co nnexio n with the H ole ctyp oid a m u s t be very remote ,

prod uced al mo s t ex actly the s am e type O f s tructure i n

Lower Cretaceou s t ime s .

The seco n d a ccelera t io n of evolu tion occur s wh en a

 B I OLOG I CAL PA LA E ONTOLO GY

roup of orga n i s m s i s appr o achi n g i ts acm e Thi s out

g .

b rea k O f m orphoge netic activity rarely if ever equal s , ,

the ear lier pha s e i n e nergy or s cope D i ffere n tiatio n of

.

ge nera a n d s pecie s rather tha n familie s or order s m ark s

, ,

i ts i n cide n ce . S i n ce the a cma i c s tage i s u s ually on e i n

whi ch the variou s m ember s O f a group excel i n n umerical
wealth it may wel l be that the s eparatio n o f relatively
,

s m a ll s ectio ns i s a n ex pre s s io n of the abu n da n ce of

i ndivi d ual variatio n available for s electio n Mo s t ge nera .

or s pecie s evolved at the acme of a group are s ho rt

lived though i n their brief career s they m ay attai n
,

remarkable s ucce ss The type s that s urvive to follow

.

the de s ce n t of the group from i t s acme are u sually

t hos e t hat took th e le as t promi n e nt par t i n acma ic
s pecial izatio n .

The third a nd fi nal outbur s t of evol utio nal activity

, ,
,

occur s in the gero n tic pha s e of a s tock D i ffere nt iatio n .

of ord i nal q ual ity is rarely d eveloped at thi s s tage but ,

ne w often s u rpr is i n gly aberra n t fam ilie s m ay appear

, , .

Modificatio ns of s uch import a nce th at a rej uve n ated

stoc k i s produced m ay occa s io n ally be attai ned An .

exam ple of thi s phe n ome n o n is probably affo rded ,

amo ng V ertebrata by the O phidia which s eem t o ra n ge

, ,

back no fur ther tha n the Cretaceou s period whe n ,

R eptile s a s a cla ss were on the wa n e Perhap s t he .

S e p ioid s are s imilarly re lated to the B e le mn oi d s a nd the ,

Clyp e a s troid s to the H ole ctypoid s U s ually however

.
, ,

the fam ili es tha t bra n ch from a gero ntic s tock s how
ma ny feature s of racial O ld age eve n at their i n ceptio n
in t o oblivio n together with t hei r m
,

and s ink ore

o rthodox a s sociate s .

The Brachiopoda m ay b e c ited afre s h i n illu s tra t i on

o f t he t hree s tage s of rapid s pecializatio n The Telo .

t re ma tous s upe rfamily S piriferacea who s e relative ly

,

brief ca reer la st ed from O rd ovicia n t o Jura ss ic time s ,

1 12 I N VE R TE B R AT E PAL AEON T OL OG Y

reached a prom i n e nt acme i n the D evo n ia n O f t he .

n i n e familie s n o le s s tha n s eve n were d i ffere n tiated i n the

,

O rdovicia n a n d S iluria n period s B.ut the pre -D evo n ia n

ge ne ra n u mber On ly twe n ty -four wherea s twe n ty -eigh t
,

origi n ated i n the D evo n ia n period alo ne ;s pecific a naly s i s

would bri n g out yet m ore s tro n gly the a cma i c q u al ity o f
m i nor d i ffere ntiatio n at that tim e O nly twelve ge n er a
.

aro s e i n the Carbo niferou s a nd five in the Perm ia n ;but

, ,

i n the Tria s whe n the s uperfam ily wa s approachi n g ex

,

ti n ctio n no le s s tha n twe n ty -o ne ne w ge nera app eared

, .

I n s pite o f thi s gero ntic effort i n s pecializatio n the whole ,

group d i s appeared before the e nd of the Lia s s ic s t a ge .

I n a d d itio n to the di ffere n tial rate s of evol utio n i nd i

ca t e d above ( wh ich are com m o n t o all o r a l m o s t all , ,

s erie s) there i s a type of irregularity which while a s ye t ,

u nrecog n ized i n ma ny grou ps s eem s t o mod ify the early

,

pha se s o f racial -life i n s everal ca s e s I t m ay be de .

“
s cribe d a s
”
deferred s pecializatio n The o uts ta n di ng
.

exam ple of thi s ea rly retardatio n i s t o be f ou n d i n the

M a m m al ia Al though the Mam m alia n s t ock aro s e i n
.

R haetic or perhap s Ferm o -T rias s ic time s its m em be rs

, , ,

s howed littl e progre s s throughout the M e s ozoic e ra At .

t he com me nceme n t o f the Cai n ozoic extraord i nary ,

”
acceleratio n typically nea n ic i n character a nd p ro
,
“

d uct s rapidly brought the cla ss i n to a po s itio n of fau nal

,

pred om i na nce I t i s po s s ible t o argue i n thi s ca se tha t

.
, ,

t he i mp orta nce Of M e s oz oic R eptil ia was s uch that the re

‘

wa s n o room for a s econ d powerful gro up of V ertebrate s

a n d it i s true that the ri s e of Mam mal ia wa s deferr e d

u n til the R ep tilia were red uced to i ns ignifica n t propor

tio n s in the fau n a But s uch a n argu me n t would in no
.

way accou n t for t he s low s pecial izatio n O f the Ma mma lia .

I t m ight have bee n expected that the two great cla ss e s

‘

woul d have competed fo r m a s tery the Mam mal s di ss i ,

p ati n g thei r racial e n ergy i n vai n e nd eavour s t o m ak e

1 14 I N VERT E B R A TE PAL AEONT OL OGY

O lder realizatio n O f the u nderlyi n g u n iform ity t hat a pper

tai ns to all l ivi ng thi n gs was i n s ome ways o f eve n
, ,

greater a s s i sta n ce toward s u n der s ta nd i n g of vital phe no

m e n a But there i s n o t yet a ny real k n owledge O f the
.

force s that drive or the law s that guide the s tream of

, ,

orga n ic progre s s ;a ge neral view of fau n as a n d fi ora s ,

rece n t a n d fo s s il give s a n impre s s io n of a m ob local ly

,

s egregated i n to cl ique s rather tha n of a proce s s io n with

defi n ite order a nd aim I t m ay be take n a s certai n that

.

the d i s order i s m erely appare t the prod uct of i m

n —

perfect a nd re s tricted k n owled ge Palaeo n tol ogy gives

.

s cope for lo n ger a n d wider view s tha n the S cie n ce s that

deal with exi s ti n g form s alo n e ;i n ful ler k n owledge of

fo s s il s l ie s the hope for d i s co ve ry of the fu nd ame ntal
l aw s o f l ife .

I n the earl ier part s of thi s chapter a n alogie s have ,

bee n m ade betwee n i n divid ual a n d phyletic career s ;

“
com pari s o n of the two k i n d s of l i fe -hi s tory ca n be
a mplified with p ro fit .I t m ay be a ss u med that the l aws
that gover n i nd ivid ual life are O perative throughout the
orga n ic world . S uch a n a ss um ptio n s uppo rted by ,

k n owledge of the worki n g o f com parable law s lead s ,

logically to the s ugge s tio n that a cl ue to the n ature of

vital pri n ciple s m ay be fou n d by s tudy of the l ife -hi s tory
o f a ny s i n gle orga n i s m Whether th e expres s io n of the
.

law s m a n ife s t e d i n the brief s pa n of i n divi d ual life is

s im pl ified ,
or co n ce n tr ated ( with re s ulti n g com pl exi ty ) ,

ca n be determ i n ed o n ly by compari s o n with i t s exte n ded

record i n phyloge n y B ut s i n ce every orga n i s m gives

.
,

s o me a mou n t of phyletic recapitulatio n i n o n toge ny of

the ma tter of which it i s built it i s rea s o n able to expect

,

that the ma n ner a nd p roce s s of i ts livi ng m ay bear com

parable relatio n to the l aw s o f evol utio n “
K n ow .

”
thy s el f m ay s till be the oracul a r reply to a s eeke r a fter
biol ogical k n owl ed ge .
 BI OLO G I CAL PALA E ON T OL OG Y

There are fo ur cri se s in the l ife Of mo s t a n imal s ,

particularl y o f the Metazoa The fi r s t that of fertil iza

.
,

“
tio n of the ovu m m ay be cl aimed as al m o s t creative
,

i n e ffect ;but i t s s ig n ifica n ce i s fa r from clear a nd the ,

phe n om e n a of parthe n oge n e s i s s how that it i s not

u n iver s ally n ecess a ry The s eco n d cri s i s occur s after a
.

pe ri od of l arval or emb ryo ni c developme n t whe n meta ,

mo rpho s i s or bi rth i s reached Thi s n orm ally take s

, , .

pla ce whe n o n toge ny h a s produced s om e approxim atio n

to ad ul t s tructure The third cri s i s m ark s ar rival at
.

m aturi ty the fourth i s death I n compari s o n with race

,
.

life i nd ividual birth m ay be take n to repres e n t the

,

i n ceptio n O f a ne w s tock maturity corre s po nd s with the

,

acme a nd death s ugge s t s exti n ctio n By a n alogy it is

,
.
,

pa rdo n able to s peculate that fertilizatio n is a recapi t ul a

t io n of the or igi n of life .

The cri s es are of rel atively s ho rt du ratio n ;bu t t hey -

are clearly recog n izable i n phyloge ny i n all ca se s where

reas o n ably complete k n owledge i s available They s erve .

a s co n ve n ie n t divi s io n s i n developme n tal s eque n ce O f

i ndividual o group life

r - The i nte rval s betwee n the
.

cri s e s vary m uch i n le ngth i n differe nt type s of Metazoa ;

their d uratio n is n ot neces s arily com me ns urate with the
amou n t of o n toge neti c develop me nt attai n ed There .

is however a t e n de ncy for m ore highly s pecialized

, ,

orga n i s m s to require longer period s for growth ;though

“ ”
the phyletic age O f the s tock seem s to have s om e
i n flue nce .

The larval ( or embryo n ic ) s tage of developm e nt i s so

.

clearly co n nected with phyletic evolutio n that it need s

no further em pha s i s i n thi s place I t i s i n the later
.

pha s es of i n d ivid ual growth that compari s o ns with

phyloge ny are m ore fea s ible i n V iew of the n ature of
,

palaeo ntological evide nce Betwee n birt h a nd m atu rity

.
,

t hree s tage s i n a n age ne s i s ca n be arbitra rily s eparated

1 16 l NVE R TE B RA T E PALAE ON T OL OGY

co rre s po n di n g wi th i n fa ncy childhood an d adole s ce n ce

, .

The epi s ode s O f the s e s tage s m ark the progre s s of a n

orga ni s m toward s the degree of developm e n t attai ned
by the group of which it i s a m ember I n com mo n .

phra s eo logy a new b or n i n fa n t may be s aid to s tart

,
“

” “
late s o that it need s m u s t hurry to catch up the mai n
,

group of its fellows B ut s peedy a s early progre ss m ay

.

be it lead s alo n g the s am e cour s e that was take n

,

d ur1 ng phyle t ic a n age n e s i s of the s tock The dom i n a n t .

characteri s tic s of youth ca n be a s cribed to exubera n t

vital i ty The child h as t o fi nd its pl ace in race -l ife a nd
.
,

to fit it s el f for the activitie s of m aturity S tructu rally .

it n ot o nly grow s but develop s ; p sychologically it

,

lear ns chiefly by experim e n t the o n ly s ou n d me t hod

, , .

H abit s the s ubco n s ciou s s equel s to experie nce are

, ,

a cqu ired ;s om e m ake for s ucce s s other s m ay be ha r m ful , .

I f m aturity i s to bri n g full fruitio n g ood habit s m u st be ,

co n fir med a nd exte n ded bad o ne s d ropped The m o s t

, .

e ndu ri ng im pre s s io ns are tho s e gai ned in i n fa n cy ;they

are co ncer ned with fu ndame n tal s a n d their n atu re m ay ,

i n flue nce the cou r s e of al l l ater develop me n t of characte r .

A s childhood pa s s e s to adole s ce n ce detail s are added ,

t o the early outl i n e ; the rate of their i nt roductio n

become s s l ower u ntil m atu rity i s reached .

I t i s clear that thi s s ketch O f early i n divid ual life

could be appl ie d al mo s t verbati m to phyletic a n age nes i s
, , ,

r eplaci ng the word s i n fa n cy chil d hood a n d adole s ce n ce

,

by their phyletic equivale n t s n ea n ic n epio n ic a nd ,

epacm aic s tage s The rate a n d quality Of d i ffere ntia

.

tio n i n a s tock are greate s t i n the n ea nic s tage whe n ,

m o s t ordi n al ( a nd eve n fam ily) s erie s are evolved I t i s .

,

m oreover particularly characteri s tic of t hi s s ta ge t h a t

,
'

great n um ber s of s hort -lived ofte n ve ry aber ra n t , ,

”
of
“
bra n che s ari s e the
— u n s ucces s ful experime nt s
i nfan t life A fte r ne a ni c accele ra t io n follow s rel a t ivel y
.
1 18 IN VE R TE B R ATE PALA E ON TOL OG Y

perhap s s om e Prim ate s) s u ffer lo s s O f d e ntitio n Ma ny .

s how phyle t ic rever s io n n o n e the le s s a nalogou s with

,

s eco n d -childhood becau s e i n com plete The S piriferacea .

have already bee n cited i n il lu s tratio n of the outbur s t

of s pecial i z atio n ( of ge neric a nd eve n fam ily quality)
t hat precede s exti nctio n ; m a ny other group s m ight
have bee n u s ed for the purpo se .

I t i s po s s ible therefore to s u m marize the matter

, ,

i nclu ded I n thi s chapter by the s ugge s tio n that i nd ividual

l ife d oes pre s e n t i n epitome the s alie n t characters O f
, ,

phyloge ny The i n te ns ity of developme nt varie s i n

.

i ndivid ual s n o le s s tha n i n group s There are roughly .

, ,

two ex t reme type s of i n d ividual l ife ;that ofte n re com

me n ded a s s hort a nd merry a n d that hu mdru m
“ “
,
” ”

exi s te nce that h a s u s ually bee n the lot of a n olde s t “

i n habita nt . M uch m ay be accom pl i s hed d uri ng the

brief cours e of the former type ;little achieveme n t l o n g ,

draw n o ut charact erize s the latter; S o i n phyloge ny

,

the s tock s tha t ri s e rapidly t o promi n e n t acme s fall

precipitately ; they have s hort but effe ctive c areer s , ,
.

Th os e gro up s that cli mb s lowl y a n d without di s ti n ct io n

to i n gloriou s m aturity l i nger through a n u n eve n tful a nd
protracted p eriod of s e n es ce n ce Accid e n t m ay s horte n
.
,

o r s helter prolo n g the live s of i n dividual s ;evide n ce

,

i s n ot wa n ti n g that comparabl e circu m s ta nce s have

i n flue n ced phyletic evol utio n .

Life whether i n divid ual or racial se em s n orm ally to

, ,

fo llo w a parabol ic cour s e A n orga n i s m ri s es fro m birth

.

to m aturity a nd the n with l ike s peed s i n k s back to the

, , ,

u n k n ow n from whe nce i t s pra ng A phyle t ic s tock .

pur s ues a s imilar cour s e a n age netic a n d catage netic

, ,

from i nceptio n to exti nctio n The a n alogy i s too pe rfect

.

to be the outcome o f m ere coi ncide nce or s ophi s try it

i s a n ex p re s s io n O f the s implicity a n d order that are
fu nd a m en tal attr ibutes o f th e U n ive rs al C o s m o s .
 PA RT I I .
- S
H I T O RI CA L B I O L O GY

C H AP T E R I
PRE -CAMB R I A N F A U NA S

HE rock s that u n derlie the lowe s t bed s of the

Cambria n sys tem cover a co n s iderable part O f the
E arth s s urface a n d attai n a n e n ormou s thick n e s s
’
.

They ca n be divided i nto two s ectio n s : a lower s erie s

of cry s tall i ne g nei s s a nd s ch is t a nd a n upper of le s s
,

al tered s edime ntary a nd ign eou s m aterial M eta .

m o rphis m h a s de s troyed al l origi n al character s i n the

fo rm er s erie s so that there i s little po s s ibility of a ny
,

palaeo ntological evide n ce rem ai ni n g es pecially s i nce a ,

large proport io n s eem s to have bee n pluto n i c i n chara cter .

E 03 00” o nce regarded a s a reef buildi n g F o ram i n ifera n

,
-

of Archaea n time s h a s bee n defi n itely reclaimed from

,

the Biol ogical s phere a nd proved to be a pecul iar i n ter

,

growth of mi neral s i n the cry s tal li n e com plex of

O phicalcite . B ut while no determ i nabl e fo ss il s ca n be
“ ”
expected i n the Lau ren tia n s erie s of fu n dame ntal
rock s m i n eral matter s ugge s tive of orga n ic activity i s
,

n ot wa n ti n g Ma s s e s of m arble ( c ry s talli n e calcite) a n d

.

graphi tic s chi st s s ugges t the exi s te n ce of l ime s to n e s a n d

c a rbo n aceou s s hale s amo n g primeval depo s it s H ow .

ever k n owledge Of the co n ditio n s preval e n t i n Archaea n

,

time s is so imperfect that it is at lea s t a s rea s o n abl e t o

ascribe th es e s ub s ta n ce s to i norga n ic precipitatio n o r
1 19
1 20 IN VE R TE B R ATE PALA EON T OL OG Y

d i stilla t io n a s to i magi ne t hat they are the prod uc t s o f

l ife The extreme s carcity O f ooze -build i ng a n im a l s a nd
.

pea t -m aki n g pla n t s i n the Ol der fo s s il iferou s st ra ta

les s e ns the p robability of their occurre nce i n yet ea rl ie r
period s .

S eparated from the fu n dame nt al A rchaea n compl ex

by a com plete a nd i nt e ns e u n co nformity the p re ,

Cambria n rock s pre s e rve i n the m ai n tho s e l ithological

, ,

charac t er s with which they were origi nal ly e n dowed .

G reywacke s cleaved b ut hardly recrys tallized co ns ti t u t e

, ,

the bu lk o f the n on -volca n ic part s ;although the upper

portio n s s eem to co ns i s t of coars e arko s e a n d con
glom erate i n m o s t regio ns A m o n g B riti s h r e p re
.

s e nt a t ive s of thi s s y s tem the U rico n ia n ( a nd comparable

,

s eries ) a nd Torrido n ia n a re u ns uited for i nclu s io n of

palaeo n tological evide nce the former be i n g m ai nly
,

volca n ic a nd the latte r pro bably terre s t rial i n origi n

, , .

But the s late s of the L on gmyn d i a n s e ries i nterbedde d ,

with are n aceou s and a s hy bed s are not ve ry di ffere nt

m
,

i n l ithology fro m uch of the Lower Palaeo z oic grey

wacke a nd s eem to have bee n depo s ited u nder com
,

parable co n ditio ns Worm track s a n d burrow s which

.
,

“
occur i n them prove t hat the de s ig n atio n Azoic i s
,

i n appropriate ; but they have yielde d no m ore s a ti s

facto ry fo s s il s I n A m erica a certai n n u mber of s ome
.
,

what O b s cure form s have be e n di s covered i n the

Algo n k ia n ( a form atio n n ot u n like t he L ongmynd ia n) ,

but the i n tere s t of thi s di s covery i s a l ittle d i s cou nt ed

by the appare n tly Lower Cambria n facies of the fau n a .

I n s pite of the al mo s t com plete ab s e nce o f di rect

evide nce it m ay be take n a s certai n that orga ni s m s
,

i nhabited the pre-Cambria n s ea s The wealth a nd

.

variety of the Cam br ia n fau n a dem a nd precedi ng '

epi s ode s o f evolutio n ;a n d ye t with i n it s el f the fau na

'

.
, ,

s eem s t o i ndicate th a t i ts precur s or s w ould not hav e

1 22 IN VE R TE B R ATE PALA E ON T OL OG Y

d esce nda n t s . The rarity of fo ss il s i n Cambri a n rock s

m ay wel l be d ue to the i n ad equacy of co n temporary
s keletal ti ss ue for pre s ervatio n— i t ca nn ot be a s cri b ed

wholly t o u n s ati s factory l ithological character s I t is .

therefore rea s o n able to s uppo s e th at the pa rtial failure

of pal aeo n tol ogical evide n ce i n the Cam bria n p eriod
give s ( i nver s e) pre mo n itio n O f it s total ab s e nce fro m
earl ier record s N ot u nt il the ope n i ng of the Palaeozoic
.

era had evol utio n produced orga ni s m s capable of

“ ”
s ecreti n g hard -part s J u s t a s i n o n toge ny early
.
,

d evelopme n tal s tage s are characterized by s oft n e s s or

d elicate m embra n ou s coveri ngs ;s o i n phyloge ny the ,

a n ce s tral form s of a s tock m ight be expected to lack

s tructure s capable of n orm al pres e r vatio n .

The o nly I nvertebrate phyl u m at prese n t u ndi s covered

i n C ambria n s trata i s that o f the Polyzo a I n all of
.

the other s the type s d i s covered are u ndoubtedly

,

prim itive i n s tructu re although no i n d ication of form s

,

tr a ns itio n al betwee n the phyl a appear s The bou ndaries

.

betwee n orde rs are O fte n O b s cure but the es s e ntial

,

characteri s tic s of the great d ivi s io n s had already bee n

di ffere n tiated The palaeo n tological record fail s j u s t
.

where the m o s t i mporta n t eve n t s i n evol utio n wou ld be

expected to appear .

I t h as O fte n bee n argued that the occurre nce of s uch

d iver s ity i n the fau n a of the Cambria n period require s
i nvocatio n of a proportio n ately lo n g s eque nce of previou s
-

fau n a s i n pre -Ca mb ri a n tim es B ut the method s O f e volu

.

tio n that ca n be traced i n later period s an d i n s m aller,

group s m u s t s urely have operated co n s i s te ntly I t h a s

,
.

bee n s how n that the mo s t s triki n g characteri s tic of nea n ic

pha s e s of evol utio n i s the rapid ity with which im porta n t
i n n ovatio ns ( ofte n of ordi n al qual ity) are attai ned .

H e n ce the e la b Ora t i on of the fi r s t Metazoa n would

,

robably b follo wed i n a relativel

y s hor t t ime b y
p e , ,
 PRE -CAM B R I A N F AU N A S

d i ffer e nt ia t io n of a ll
the m ai n phyletic s t oc k s The .

s im ple qualitie s s how n by m o s t Lower Cambria n

orga n i s m s imply that the phyla were but n ewly
s pecialized at that time Perhap s s ome a n ces tral
.

Metaz oa n s tock had bra n ched al m o s t at a s i n gle n ode

, ,

i n to the variou s phyla at a s tage no t m uch more rem ote

,

tha n the upper pre -Cambria n Prior to t hi s epi s od e .

,

comparable acceleratio n may have d i ffere n tiated the

three great s ectio ns of the a n im al ki ngdom withi n a
relatively brief period .

I t m ay be objected th at the foregoi n g hypothe s i s i s

reactio na ry i n that it revive s co n ceptio ns which i n
, ,

geologi cal di scu s s io ns have bee n d i s carded a s u nsou n d

, .

E arly s chool s of Geologi s t s i n troduced idea s of the

,

acceleratio n of phy s ical proce s s e s i n rem ote period s ,

largely u nder the ( s ubco n s ciou s) pre s s ure of theological

prej udice Accordi ng to their beliefs rai n fell more
.
,

heavily river s flowed m ore s wiftly a nd the s e a ( perhap s

, ,
“
boili ng hot ) wa s m ore rapaciou s the n tha n n ow s o
-
”
,

that va s t qua n titie s o f rock -m atter were eroded a n d

depo s ited d uri n g s ho rt i n terval s o f time All of which .

i s n ot prove n a nd d i fl i c ult of belief But argu me nt s fo r

.

orga n ic acceleratio n fall i nto a di ffere n t category Al l .

e evid ence poi n t s to the vigour a n d variability

whether i n dividual or racial I t i s n ot m ere .

ct i on o f a hypothe s i s ba s e d o n

to s ugge s t that differe n tiatio n a nd

of orga n i s m s m ay have proceeded rapidly
“
by l ater s ta ndard s ) whe n the worl d wa s
 C H A PT E R I I

L O WE R PALA EOZ OI C F A U N AS ( Pm . 1 x. A ND x .
)
A
( ) GEN ERAL A CC O U NT
H E s trata of the C ambria n , O rdovicia n a n d
S il uria n period s exh ibit , i n Britai n , a certai n
m o noto ny of l ithol ogi cal qual ity relieved l ocally , ,

a nd a t variou s horizo n s by develop me n t of volca n ic

,
‘

rock s The s e i n terlude s were m o s t widely d i s tributed

.
,

a n d prod uced the greate s t bul k of pyrocl a s tic a n d other

effu s ive m aterial s i n the O rdovicia n period ; but i n

,

s pite of t h e s carcely fo s s iliferou s n atu re o f the A re n ig

s erie s of Wale s a n d the Borrowd ale s erie s o f the Lake

D i s trict a practical ly co n ti n u ou s fau n al record ca n be

,

d i s cer n ed i n o ne d i s trict or a n other throughout the

, ,

Lower Pal aeozoic era The domi n a n t type o f s ed ime n t

.

“
i s be s t ex pre s s ed by the term Greywacke — de t rital
m atter with a n average com po s itio n of s a n dy s late
co n tai n i n g s mall qua ntitie s of drifted volca n ic a s h .

A ctual ly al l o rd 1n a ry grade s from co n glomerate to

pu re cl ay -s late occur bu t a certai n argillaceou s qua li ty
,

pervade s the whol e s erie s except i n the n orth -we s t

,

H ighla n d s of S cotla n d With the s a m e exceptio n

.
,

the few relatively thi n a nd i m per s i s te nt l ime s to ne s of

, ,

the s erie s a re u s ually earthy eve n the c oral reefs bei ng

,

i m pregn ated with grey s hal e The s e lim es to nes are

.

either coral li ne or s hell y ;i n n o ca s e s do they a ppear I

t o repres e n t depo s it s of the ope n s ea I n ma ny .

d i s trict s n ot a bly the Wel s h border a nd A ngle s ea t he

, ,
1 26 I NV E R T E B R AT E PALAEON T OL OGY

The form s o f l ife pred om i n a n t duri n g the e ra were

the G raptolite s Bra chiopod s a nd Trilobites The fi rs t
, .

n am ed group aro s e i n the later s tage s o f the Ca mbria n

_ ,

a nd becam e practically exti n ct

be fore the e nd of th e

S ilu ria n
. D uri n g i t s s hort career it attai n ed a ve ry
promi ne n t acme ; a n d by i t s rapid a n d abu n da n t
variatio n provided s at i s factory zo n al i nd ice s Brief .

s earch i n a ny O rdovicia n or S iluria n s hales u s ually

reveal s i n n u merabl e re mai n s O f G raptolite s The .

Brachiopod s belo n ged chiefly to the three order s

Atrem ata N eotrem ata a nd Protrem ata O f the s e the
,
.
,

t wo fi r s t were m ore abu n da n t a n d varied d uri n g the

Lower Pal aeozoic tha n at a ny l ater s t age ; but they
were over s hadowed by the a cma ic exu bera nce of the
O rt h a ce a S trophome n idae a n d Pe nt a mer a ce a of the
,

Protrem ata I t i s rarely po s s ible to vi s it a fo ss iliferou s

.

horizo n i n Proterozoic s trata withou t di s covery of m an y

s pecie s that would have bee n called Or tlzzs b y early
'

Pal aeo ntologi s t s ;while Pe nta me r id s are im porta nt roc k

former s i n the S il uria n The Trilobite s are the m o s t
.

s triki n g a nd n ot the lea s t characteri s tic fo s s il s of the

, ,

Lower Palaeozoic H ypoparia a nd O pi s thoparia abo u nd

.

i n the Cam bria n while the Propa ria give them full
,

s upport i n the O rd ovicia n A lthough Trilobites ra nge

.

u p to the Permia n po t S iluria n type s are u su a lly

s -
,

s carce i n occurre n ce a n d gero n tic i n character The .

O rdovicia n i s the period of their acm e twe n ty -t w

, o
fam ilies ( ou t of a total of twe nty -eight ) bei n g there
repr es e n ted Trilobite s are u s ed a s zo n al i ndices i n
.

th e Ca mbria n ( i n the ab s e n ce of Graptoli t es ) b ut t he

‘

worl d -wide ra nge o f m a ny O rdovicia n a nd S il uria n

ge nera give s tho s e later types great s tratigraphica l
val ue s lightly di s cou nte d by te n den cies to s pecific
,

l ongevity .
 LOW E R PAL AE OZOI C F AU N A S

( )
B P R OT OZ O A
The relative s im pl i city O f the Protozoa would s u gge s t
that they were abu nd a n t if not predom i n a n t mem ber s
, ,

of early faun a s Although s uch a co n ditio n m ay have

.

bee n realized the fact rem a i n s that rel ic s of the two

,

order s capable of fo s s ilizatio n are le s s abu n da n t i n the

Lower Palaeozoic tha n at later period s R adiol aria .

occur s pari ngly i n a ll three s tage s ( a n d i n deed i n the

, ,

pre -Cambria n) but F oram i n ifera are u n k n ow n i n the

,

Cambria n barely record ed from the O rdovicia n a nd

, ,

ex ceed i ngly rar e i n the S il uria n . Two expla n atio n s

of thi s a n om aly are available The i n durated qual ity
.

of m o s t Lower Palaeozoic rock s make s s eparatio n of

m i n ute fo s s il s d i ffi cult ; fu rther it h a s r e s ulted fro m
,

i ncipie n t meta morphic cha n ges that would n ormal ly

O b s cu re s m al l obj ect s by m ole cu lar readj u s tm e n t The .

pe r s i s te n ce of s iliceou s R adiolaria s ugge s t s the opera

tio n of thi s proce ss s i n ce their s ub s ta nce woul d O ffer
,

re s i s ta n ce to altera tio n
. S eco n dly the retarded nature
,

of Protozoa n evoluti o n m ay have fou n d expre s s io n

i n delay i n the s ecretio n O f hard s tructure s The
.

alm o s t complete ab s e n ce of F oram i n ifera from O rd o

vicia mdep o s its s eem s expl icable o nly o n the a s s u m ptio n

that fe w mem ber s of the order po s s e ss ed durable te s t s
at that time R adiolaria are l ocally pre s e n t i n rock
.

form i n g qua ntitie s i n the O rdovicia n of S cotla nd .

rt for thi s bel ief is fou n d i n the n ature

S iluria n F oram i n ifera S a cca mmz na
'

“ ”
are bo th A gglutina nt ia buildi ng
,

tests i n which adh eren t foreign particle s are bou n d

together by a n i ns ig n ifican t amou nt of s ecreted m atte r .

The few vitro calcareou s ge n era of the period are u s ually

fou nd a s glau co n itic m ould s ;while thi s character is
sh ow n eve n by H oloce ne type s i t is s ugge s tive of
,
1 28 IN VE R TE B R ATE PA L A EON TOLOGY

s pee d y d i ss olu tio n of very del icate te s t s There are .

ple n ty of S il uria n l i me s to ne s which migh t be expected

to a flord com plete pre s ervatio n to calcareou s F ora min
i fe ra if they had bee n available .

A m o n g R adiolaria two s uborders o n ly S p ume lla ria ,

a nd N a s s e la ria are clearly recog n ized in the fo ss il

,

s tate O f the s e the form er m ay with propriety be

.
,

co ns idered the m ore prim itive s i nce i ts member s are

,
~

roughly s pherical a nd h ave u n iform ly perfora t ed

skeleto ns The Na s se la ri a with divers e form a n d
W
.
,

W
It i s in
accord with expectatio n t o fi n d t hat Lower
P alaeozoic ( a nd pre - Cambria n) R ad iolaria are all
S p ume lla r i a n s T hey a re rarely wel l p res erv e d ( thei r
.
-

s ilica bei n g u s ual ly d e vi t ri fi e d i nt o chert d

) a n ca n be ,

recogn ized i n thi n rock -s ectio ns ra t her tha n collec t ed

i nd ividually .

( )
C POR I FE RA
X Two group s of S po nge s s ecrete s keletal mat t er O i
d urability s u ffi cie n t for fos s ilizatio n O f t he s e the
.
,

S i lici s pgn gjae t e n d to prod uce loo sely a s soci ated s picule s
l
‘

which collap s e after dea th ; L it h is t id s a nd D ictyonine

H exacti nellid s alo n e have well -k n it s keleto n s The .

form atio ns from the Cambria n o nw ard s .

De t ached s picule s of M o n acti nelli d s po nge s appear

fi rs t i n the S iluria n but are rarely s uited for preci s e
,

ge neric d iag n osi s Lithi s t i d s repres e nt i ng al l five s ub

.

o rd e rs occur i n the Ordovici a n a nd S iluria n ; Tetr a

1 30 I N VE R T E B R ATE PALAE ON T OL OGY

O lder s trata They i m med iately a ss u me d biological

.
)

a n d ge ological i m porta n ce Al cyo n aria ( or O ctocorall a )

.

have a l ike ra n ge bu t are le ss s ati s factorily k n ow n a s

,

fo s s il s Lower Palaeo z oic Zoa n tharia belo nged chiefly

.

”
to two k i n d s Tetracoral la ,
R ugo s e Coral s ) a n d
Tabulata Tru e Coral s ( i n the re ce n t s e ns e) were wa nt
.

i ng u n le s s t he Archaeocyathidae belo n g to that group

,
.

The pr eci s e a ffi n itie s of both s ub -cla s se s are doubtful ,

a n d recog n itio n of the forme r i s n o t u n iver s al While .

T e t ra cora ls have m uch i n com mo n with the later H exa

corall a ( a n d m ay perhap s b ea n early a nd peculiar m a n i
fe s ta ti on of that group ) Tabul ate s with ru dime n tary or
»
, ,

ab s e nt s epta s eem at fi r s t s igh t m ore aki n toAl cyo naria

,

or H ydrozoa Both group s were im po rta n t rock builder s

.
-

i n Lower Pal aeozoic ti me s co n ti n u i n g through the Up per ,

Pal aeozoic a nd the n becom i n g virtually exti n ct .

R el atively few T e t ra co ra ls are k n ow n from the

O rdovicia n ;s m all s olitary form s calle d P etm za are
'

the m o st abu nda n t A mp lexus a nd Cya thop hyllu m al s o

.

o ccu r u s ual ly a s s m al l a nd s im ple s pecie s I n the

, .

S iluria n a n extrem ely rich s erie s appeared Petr a za i s

'

s till fo un d i n the Ll a n dovery while i n the We n lo c k Palaeo

,

cycli d a e Cyathophyllid ae a n d Cys ti ph yllid a e formed

,

n o i n co n s iderable bul k of the co n tem po rary coral -reefs .

S ol itary type s ( e g Pa la eocyclus Cy a tk op fzyl/u m Omp IzJ/ma

. .
, , ,

Gon top kyllzt m) were accom pa n ied by m a s s ive colo n ies

( C h fi lt m S mb ( é w l )
’

e.
g y.a t op y u t r o oa es Pl ix,fig 3 A c r a r
’
z a . . . .

Tabulate s were u n doubtedly the predom i n a nt group

o f Zoa n tharia i n t he L oiver Palaeozoic F a vos it id s with .

m a s sive fu ngiform colo n ies ( e g F a vor i tes A lveolz tes )

'

, . .
,

are n ot u ncom mo n i n the O rdovicia n a n d abou n d i n the ,

S il uria n ;A ulop or id a e S yr i ngop or id a e a n d H a lys itid ae

,

wi th re cumbe n t or fa s ciola t e habit are s carce at the lower

h o r izo n but ( e s pecially the la s t -n amed ) profu s e i n the
,

S iluria n The fibrou s Cha e t etida e a r e als o k n ow n from

.
 L OW E R PALA E OZ OI C F A U N A S 1 3 1

t he se period s but Cita etetes wa s m ore fully developed i n

,

the U pper Palaeozoic F a vor i tes ( Pl ix fig 4 ) a n d H a ly

. . . .

s i tes ( Pl ix fig 5) were the chief reef builder s of the era

. . .
-
.

Alcyon aria were appare n tly repre s e n ted by the

abu nd a n t a nd s hort -lived fam ily H el iolitidae H eli oli tes .

pers i s ted i n to the D evo n ia n ;the m aj ority o f ge nera

were re s tricted to the Lower Pal aeozoic attai n i n g chief ,

promi ne nce in the S iluria n .

There are two s erie s of Pal aeozoic orga n i s m s referred

to the H ydrozoa O f the s e the S tromatoporoidea have
.
,

had a chequered s y s tematic hi s tory havi ng bee n cla ss ed ,

at variou s ti me s a s Calcareou s Algae F oram i n ifera , ,

“ ”
Porifera Zoa n tharia a n d Polyzoa M a s s ive s po n gy
, .
, ,

e n cru s ti ng s heet s of Cla tft r oa ictyon a n d S tr oma top or a

’

play a n i mporta n t part i n the co ns tructio n of S ilu ria n

reefs but the group wa s le s s abu nda n t i n the Lower
,

Palaeozoic tha n i n the s ucceed i n g era .

The H ydrozoa n a ffi n itie s of the Graptol i t oidea are

tolerably e s tabli s hed but they have b ee n regarded
,

as Pl a n t s Porifera Polyzoa a nd eve n Cephalopoda !

, ,

Po ss e s s i ng whol ly chiti n ou s e nvelopes Graptolite s are ,

rarely well -pre s erved but thi s faili n g i n no way detract s

,

from their abu n da n ce a nd s tratigraphical val ue The .

t wo order s D e n d roi d a a n d Gra p t o lo i d a s eem to ill us trate

, ,

evol utio n al pri nciple s i n a co nvi n ci n g m a nner The .

fo rmer , with bu s h -l ike colo nie s s upporti n g ve ry s mall ,

s imple thecae aro s e i n the U pper Cambria n D i t ty

, ( /an gi n a
)
per s i sted to the Carbo n iferou s rarely attai n i n g ,

da n ce a nd s howi n g little m od ificat io n The latter .

,

le s s complex habit but far m ore elaborate thec ae ,

i n the O rdovicia n a nd with the exceptio n of

’

, ,

rare type becam e exti nct i n the S iluria n

, .

their s hort a n d varied career a nd prom i ne n t ,

ida are o f s upreme i mporta n ce a s zo n al

mo period s o f their exi stence Mos t .
 I 3 2 IN VE R TE B R ATE PALA E ON T OL OG Y

O rdovicia n type s bel o n g t o t he m ore pri mi tive s e ct io n

Axo n olipa ( i n which there i s no co nti n uou s a x 15 I n
the rhabd o s ome ) ; the m o re s pecialized Axo n ophora
pers i s ted i nto the S il uria n Co ncurre ntly with thi s
.

m orphological trait occurs a habit that i s readily

recogniz abl e O rdovicia n types of both s ectio ns were
.

either bra n ched ( all A xo n olipa ( Pl ix fig . chiefly

. .

Are nig a n d Ll a n deil o ) or D iprio n idi a n ( e g D ip/o

“
. .

g p
r a t a s, G los s og p
r a t a s ) While
. both of the s e m ode s o f
“
growth were retai n ed in the Lo wer S il uria n Mo n o ,

p r i o n i d i a n
”
form s ( g
e .M o n o.r a tus
g p ( Pl ix fi g . . .

R a s tr i tes) are e s pecially Characteri s tic of that pe r iod .

( )
E E C H I N O D E R M ATA
The E chi n oderm ata al o ne of Metazoa n I nve rt ebrates
, ,

s ecrete m e s oderm al s keleto ns which like th e a n alogou s

,

bo ne s of V er t ebrate s are i n ti m ately co nnected with the

, .

peri s hable Orga n s O f the b od y H e n ce the phyl um i s

.

p eculiarly well -ad apted for palaeo n tological s tu dy T h e .

characteri s tic reticulate s tructu re of the s te reom is a s

di s ti n ctive a s that of bo ny ti s s ue a n d m ake s eve n ,

ob s cure fragme n t s of E chi n oderm s recogn izable a s s uch .

O ne cl as s o n ly the H olothurioidea i s deficie n t i n

, ,

s keletal t i s s ue ;s tra n gely e n ough rem arkably well -pre

s erved impre s s io n s o f the s e form s are k n ow n from the

C a mbria n S tarfi h
. s a n d B ittle S tar s te nd t o have
r -

l oosely co n st ru cted s keleto n s So that their remai ns

, ,

though far from rare,are u s uall yfragme ntary But the .

m a s s ive a nd cohere nt te s t s of m o s t E ch ihoid s the bul ky ,

crow ns a n d exte ns ive colu m n s of m a ny C ri n oid s a nd ,

the com pact thecae of Bl a s toi d s a nd adva nced Cy s tid s ,

give am ple opportu nitie s for fo s s ilizatio n .

[
M orphoge ny a n d on toge ny alike i n dicate that the
u l tim a t e a nce s tor s O f the phylu m were bilate ra lly
1 34 I N VE R TE B R ATE PALA E ON T OL OGY

wa s al mo s t re s tricted to t hat s tage di s appeari ng i n ,

the Lower D evo nia n Both group s were repre sen ted
.

i n the Lower Cam bria n but the e arly Cy s tid s were ,

obv iou s ly the retarded s urvivor s O f a far m ore primitive

s tock tha n the Cri n oid s The s im ple s t Cys tid order
.

( A m p h o r id e a ) m ay wel l be reg arded a s m orphoge n etic

al ly repre s e ntative o f the fi r s t s tage i n E chi no d erm

fixati o n . Mo s t A mp h ori d e a have il l - d e fi n e d radial
s ym metry ,
bei n g irregularly s a c - l ike form s who se
E chi n oderm al n ature i s appare n t chiefly i n s tereom
m e s h a n d po s ses io n of a water va sc ular s ys tem A r i s to
s - -
.

cys t i s,
a n O rd ov1c1 a n ge n u s i s the lea s t e labo rate of ,

k n ow n t ype s ;E ocys tis a Cambria n form fou n d u s ually

,

a s d i s s ociated pl ate s m ay s how a n early s tage i n the

,

developm e n t of a cohere n t theca The O r d ovicia n .

D ea a r ocys ti s ( Pl ix fi g 6 ) m ark s a great adva n ce on

’ ‘

. . .

the prim itive co n d i tio n s i nce it po s s e s s es a defi n ite

, ,

partly tubular s tem a nd a n arm -l ike outgrowth from

,

the oppo s ite pole I t h a s been fou n d i n the Ca r ad oci a n

.

of Girva n ; but the m o s t fam il iar Briti sh member of

the order i s P la t ocys ti s from the S iluria n Thi s i s a
, .
f

curiou s ly sp ecialized l ate for m laterall y compres s ed ,

“

a n d s howi n g divers e plati n g character s o n the two side s - .

A well marked s tem i s pre s e nt a n d two s le nd er s pi ne s

-
”

s eem to take the pl ace o f arm s The, ex t raordi n ary .

Cotft um ocy s ti s rece n tly de s cribed fro m the O rdovicia n

,

of S cotla n d s hare s m a ny of the featu re s of Pla t ocys tis

, ,

“
but i s u n ique i n the phyl u m i n havi n g s everal oral
”
apert ure s . The s tra n ge s pecializatio n o f thes e flatte n ed
form s i s a s cri be d t o re cu mbe n t h abit s of l i fe
‘

The order R h o mb i fe ra i ncl ude s the m o s t

type s O f Cy s tid s I t s how s a n adva nce in s
.

fro m the A rn p h o ri d ea n patter n a nd i n later , ,

“ ”
p o s s e s s al m o s t c ri noid a l regularity of p
peculiar a nd u n iver s al fe a t ure i n the order is
“

.
 L OW E R PALA E OZ OI C FA U N A S 1 35

of s tereom -fold s matchi ng on co n ti g uou s plates an d ,

becom i n g localized i n to p ect in i rho mb s i n s pe cialized

“ ”

form s M acr ocys tella a cri n oid -like form with a lo n g

.
,

s tem a nd fi ve bifurcati n g arm s i s not u n com m o n ,

( a s di s s ociated pla t e s) i n the S h ine ton S hales Th e .

O rdovicia n Ca ryocr i n id a e s how m uch s uperficial re s em

bla n ce to Cri n oid s a nd Bl as toid s E c/i i nosp /za er a a n .
,

r h o mb i fe r o u s
“ ”
al mo s t globular form with n u m erou s

plate s is fairly abu nd a n t in the O rdovicia n but is

, ,

rarely fou nd e n tire i n thi s cou ntry S iluria n R hombi .

fera s how ob s cure paralleli s m with co n tempora n eou s

A mp ho rid e a i n s hape S c/ziz ocys tis a nd L ep a d ocr i tz us
.
,
“
with re cu mbe n t arm s s ugge s tive O f Bla s toid am bulacr a ,

are am o ng the few Cys tid ge n era fou n d i n good pre s erva
tio n in Britai n They occur i n the We n lock Limes to n e
. .

A s m all a n d i n co ns picu o u s order the A p orit a occur s , ,

i n the O rdovicia n a n d S ilu ria n Cryp tocr i n us i t s typical

.
,

repre s e n tative h a s a theca comparable with a cri nOi d

,

calyx but phyletic co nn exio n betwee n the two group s is

,

ve ry improbable .

The D ip lop orita i n whi ch paired pore s perforate the

,

media n s tereom layer are perhap s the m o s t i n tere s ti n g

-
,

order of Cys tid s s i n ce Bl a s toid s were certai nly derived

,

through them a n d there i s rea s o n to co ns ider them

,

a nce s tral to Cri noid s a s well The order i s e s s e n ti al ly

.

O rdovicia n i n date a n d s eem s clearly des ce n ded from

,

t h e A mp h or i d e a The theca is u s ually gl obular or p e n

.

t a me ro us i n sym metry with irregul ar plati n g ( be com i n g

,

m ore defi n itely arra nged in later types ) a n d food groove s -

,

termi n ati ng i n b ra ch ioles p a s s mg with i n crea s i n g pe n

,

t a me ro us regul arity over the s urface s O f the plate s .

Sp /ta er oni s h a s food groove s ex t e n di n g s carcely beyo n d

-

the m outh ; Giyp tosp k a er a h a s five bra n chi n g groove s

pa s s i n g for s ome d i s t an ce over the theca M es ocys ti s .

a nd P r oter obi a s tus s how a s tro n g te nden cy to s tereo

1 36 IN VE R T E B R ATE PALA E ON T OL OGY

typi n g O f sym m et ry on the pe nt amerou s pl a n a n d l ocal ,

s pecializatio n of the pl ate s traver s ed by the s traight

food groove s
-
. Both of the s e ge nera are al mo s t a s
m uch Bla s toid s a s Cy s tid s ;P r otocr i na s bear s n o s m al l
re s e mbla n ce to s om e E drioa s tero id s .

The l a s t -n am ed cl as s ( regarded by m a ny a s a n order

o f Cy s tidea ) s eem s to have be e n either ex t remely s e s s ile

( without s tem s ) or i mperfectly

,
e le u t h e r o z o i c ( after the
m a n ner of limpet s ) E drioa s teroid ea ra n ge from the
.

C a m bria n to the Carbo niferou s a n d have bee n claimed ,

a s tra ns itio n al betwee n Pel m atozoa a n d E leutherozoa .

That they have s tro n g Cy s ti d ( eve n A mp h or id e a n)

a ffi n itie s s eem s certai n bu t i n a mbul acral s tructure
,

they a re d efi nitely u nl ike the Pel m atozoa a nd s how a

, ,

m arked r es embla n ce to e le ut he roz o ic type s S troma ta .

cy s t i s i s a C ambria n ge n u s ; E d r i oas t e r typical ly O rdo

vi ci ah H emicys tis i n the S iluria n foreshadow s the
’
‘

.
, ,

s caly type s characteri s tic O f the U pper Pa la z oi c .

The cla ss of s mal l Pel m a tozoa n ame d Bla s toide a

“ ”
com pri s e s form s tha t developed al o ng cri noid al li nes ,

except i n ambulacral character s I ns tead o f bei n g

.

rai s ed o n flexibl e arm s the food grooves are recu m be n t

,
-
,

a n d pa s s with ex t reme re g ul arity dow n five radii from -

the oral pole s om eti me s al m o s t reachi ng the oppo s ite

,

ex trem ity ( whe n ce pr oj ect s a s le n der s tem ) The s e .

“ ”
a mb ulacra s how s tra n ge s u perficial re s em bla nce to
t he a rea s s im ilarly n am ed i n E chi noidea ;the com
paris o n i s ampl ified by the developme n t O f s im ple -

“ ”
a nd petaloid s tructure s Prim itive Bl as toid s ( Proto
.

b la s t oid a ) are re s tricted to the O rd ovic ia n a n d po ss e s s ,

s om e d ip lop o ro u s plate s thu s demo n s trati n g their a ffi n ity

,

with Cy s tid D ip lop or i t a Tru e Blas toid s appeared fi rs t

.
‘

i n the S il u ria n wh e re Coa a s ter s howed a n early a nd

’
,

i nco mplete phas e of the developme nt of i nt er n al s tereom

fold s ( hyd r os p ire s ) cha racteri s tic of typical fam ilie s .
 L
P AT E I X
 L
P AT E I X

L O WE R PA A E O ZO I C O SS I S L F L
X S TA I I
L
‘
L

FIG . 1 . D i ay mogr ap t u s
’
‘
mu r t lz zsom lan d e i lo , A
'

. br e e id d y .

F IG .

FIG 4 a si e t sflgnflz lan a’ ca

i L I me
j
. .

st e P es t h o S lop
p r pe a“ t
Lm
.
,
n ,I ,

W
i " h
F IG 5 H a ly s i tes ca tc mla r z a e n lo ck i es t o n e , M u ch We n lo ckf
z
. . .

W
FIG 6 D en d r ocy st z s st a t
'

. .

j n
W
”v 4 1
i
r
i
L I In e st one , Much f
K
1
it /z ot r i n t s s p e n lOck
i

e n ock
‘
FIG
’ ‘ ‘

. c . .

mb
’
8: i l T r e matlo c
'
" ‘ r

FI G n u lel a dd éz si
f
. Ca n an , .
( Pa rt ly b a sed
‘

on

D a v i d s on
’
s fig u re )
L
.

FIG 9 . Or éz cu lozd ea ru
ga ta m .
U p pe r u dlo w, O n i b u y, 8 3 1013r 1
-x

FFG .
‘
10 .
( PD a lma m lla ) ’
tes t u d m a ma .
,
.
_
Car ad oc ia n, Ma rsbi o

b ro ok , S a lop . R e st o re d gfr o m ca s t s Pl. v1.

ex p a n va t . Ca ra d OCI an, S o u d l y, eZ p
Sa lO . Res t o d re
( pa rt ly b as ed
’
1d of ve ntra l v a lve on D a vi d so n s

F IG . 12 . C/z on et es st r i a t etta . U pp r L e u d lo w, O ni b u ry, S a lop .

 L
P ATE X

L O W ER Pi
fL a EOz o
ie F O SS I
L S

m
‘ “

FIG . L
u d lo wL i es ton e , We o Ed ge;S alop .
f

ort fi oizh Whett leton, S alop

z
FIG ta rigi d a )
'
a
. . .

L
O

9 7 1/1 9 c bu lla t um . Upper ud low, O ni bury , S a lop .

Pa r a boti nella t r i a r t lz r a . S h ine ton S h a le s, Be lswa rdi ne , B I Ok ,

O
S a lop .
( S li gh t ly r e stored ) .

Ogygz ot a r i s b u r/i i . Lland ei lo , : B ui lth a an.

»
0

Tr i n m leu s t on mn t n ms » Onu y
'

Cryp toli t lz d s Ca rad ocia n, »1

' ’

F 1G
,
. Cer a t i oca r z s
’

N ep toa acty lu s
’
. L o we r L ud lo w ,
Mg
r ti n s
’
S h e ll ,
 LOW E R PA LA E OZ OI C F A U N AS ,

1 37

Cri no idea the o n ly e n d uri n g cla s s of Pel matozoa

, ,

occu rred i n the Cam bria n but the O ldes t type s are not ,

wel l k n ow n I t is i n tere s ti n g to fi n d that one O rdovicia n

.

ge n u s Hybocr i nus had defi n ite though eleme nt ary

, , , ,

hyd ros p i re s s ugge s ti n g deriva t io n of the cla s s through

, ,

o r parallel with early Bl as toid type s E xtreme regu

,
.

la ri ty i n plati n g a nd prolo n gatio n O f the food groove s

,
-

”
on to
“
arm s are the dom i n a nt chara cteri s tic s O f the
,

clas s The calyx ( pati na ) m ay be b uil t O f two or three

.

cycle s O f plate s ;it i s d i fficult to b el ieve that the two

chara cter s ( which im ply co n tra s ted arra ngeme nt of
ne rvou s a n d other s tructures ) ca n have be en developed

m ore tha n o n ce in Cri noid hi story The two s ub .

cla ss es thu s recog n ized Mo nocyclica a nd D icycl ica , ,

followed n early parallel a nd sy n chro nou s cour s es of

evolutio n but Mo n ocyclic form s are al m o s t exti n ct
,

t o -day while m odified D icyclic familie s are profu s ely

,

r epres en ted i n m oder n fau n as .

Mo nocyclica are divid ed i n to three orders I nad una ta , ,

Adu nata a nd Camerata The I n a d unat e s with calyce s .

,

built o f ba sa is a nd radial s ( with or without i nter calated

a nal plate s) are k now n from the Cambria n a nd s till
, ,

s u rvive They are ge nerally s mall form s a nd rarely

.
,

prove abu n da n t Hy éocr i nus from the O rd ovicia n h a s

.
, ,

bee n me n tio n ed abo ve ;the S iluria n P i socr i nus i s the

o n ly ge n u s well k n ow n from Briti s h L ower Palaeozoi c
horizo ns Ad u n ata with calyce s like tho s e O f the I na
.
,

d una ta but s upporti n g m a ss ive t e gme n s were mai nly

“

, ,

Of U pper Palaeozoic d ate ;but H ap a locr i n us a nd M a r s i

p or r i flus from t he We n lock Lime s to n e prepared t he
, ,

way for the abu ndan t Pla tycri n i d a e O f the Carbo niferou s
Camerata i n which the calyx co n tai n s ma ny i ncorporated
,
-

arm plate s were s imilar i n s tratigraphical ra n ge to the

-
,

Adu na ta ;P er i ec/zot r i n us is a fam iliar S iluria n fore ru n ner

O f the la ter Acti n ocri n idae .
1 38 IN VE R TE B R AT E PALA EON TOL OGY

D icyclic Cri noid s are d ivide d i nto I na d unat a F le x ib ilia

‘

a n d Ca merata . The fir s t a nd la s t order s are parallel ,

a nd practically co n tem pora n eou s with th e s im ilarly ,

“ ”
n amed divi s io ns of Mo n ocyclica ;a n Adu n ate group
m ight well be e s tabl i s hed for the D icycl ic Crotalo
cr i n i d a e
. A part from the S il uria n Cr ota /ocr i mt s ( one o f
the co mm o ne s t O f We n lock Cri noid s ) Cya t/zocr i n a s ( Pl , .

ix fi g Gi ss ocr i n us P eta tocr i n us a n d D end r bcr i nus are

'

. .
,

the m o s t fam iliar I n a d un a te D icycli ca of the Lowe r

Palaeozoic R eteocr i n a s wa s a n O rdovicia n precur s or
.

of the s m all ,
mo s tly Carbo ni ferou s Camerate group , .

F le x ib ili a are chiefly Me s ozoic a nd R ece n t form s bu t ,

Ta xocr i n us i s not u n com mo n i n t he We n loc k Li m es to n e .

Lower Palaeozoic E leutherozoa are rarely abu n da n t ,

a n d u s ually O b s cure W o n derfully perfect im pres s io ns

.

( al m o s t amou n ti n g to petrifactio n s ) O f appare n tly p elagic

H olothu ria ns di s covered i n t he M iddle Ca mbria n of
,

Briti s h Colu mbia carry back the hi s tory O f that debat

,

able cla ss to ve ry early t ime s The two m ai n s ectio ns of

.

S t e lle r oi d e a ( A s teroid s a nd O phiuroid s ) are both re p re

s e n ted i n the Lower Palaeozoic ( the form er i n the
‘

C ambri a n) b ut their collap s ible n ature m ake s i nter

,

p r e t a t i o n O f their s tructure s a n d a f fi n itie s d i fficul t The .

chief Briti s h Proterozoic depo s it s i n which S t e lle roid s

are a t al l n umerou s are thi n l ayers i n the Ca ra d ocia n
of Girva n a n d the Lower Ludlow O f Lei ntwardi ne
( H ereford s hi re ) . L ap wor tk u r a a n
“
, O phiuroid for”
m ,
”
“
i s extraordi n arily abu n d a n t i n the latter S tarfi s h bed .

I s olated occurre n ce s i n O rdovicia n a n d S il uria n rock s

of other Briti s h l ocalities are n ot i n freque n t but the ,

u s ual ob s curity o f the fo ss il s m ake s them u ns uited for

d i s cu s s io n i n ge neral term s .

'
The earlies t recorded E chi noid i s B ot/t r i oci a a r i s a ,

s mall ; s ubglobul ar form from the O rdov icia n . I t d iffers

from all o t her members o f the clas s i n havi ng i nter
1 40 IN VE R TE B R AT E PALAE ON T OLOG Y
t hread -l ik e colo n ie s of the Cte n os t om a t a ( e g Vi nt i la ). .

are m ore abu n da n t i n the Lower Palaeozoic tha n at

la t er horizo n s but they n ever attai n promi n e n ce Cyclo
, .

s t omata s eem to be repres e n ted i n the O rdovicia n

a nd S iluria n by the s uborder Ce ra mop oro i d e a ( e g . .

Cer a mop or a a nd F i stu lzjaor a ) ;thes e s omewhat problem

a t ical form s co n s tit uted a l arge proportio n O f the
Po lyz p a n fau n a of tho s e period s The abu n da n t s erie s
.

of form s k n ow n a s T repo s tom ata are part icularly char

a cter is t i c of the Lo wer Pal aeozoi c ( though e n d uri n g to

the Perm ia n) U n der the n a me of M o nticu li po roid s

.

their m a ss ive colo n ies were lo n g regarded a s thos e

of C o ele n terate s of Alcyo n aria n a ffi n itie s but th eir ,

Polyzoa n pos itio n se em s e stabli s hed C ryptos tomata .

,

a n other excl u s ively Palaeozoic order were le ss abu n da n t

,

i n the Lower tha n i n the Upper pe riod s but P ti loa ictya

’
,

s ometime s occur s i n profu si o n i n the Lud low Lime s to n e .

Thi s order wa s m o s t probabl y a n ce s tral to the Cheilo

s tomata t h e predom i n a n t group O f Tertia ry a n d R ece nt
,

Polyzoa which are n ot k now n from the Palaeozoic

, .

( )
G B R AC H I O P O D A
In clear ne s s a nd com plete n e s s the palaeontologi c al
record of the Brachiopod a i s s u perior to that available
for a ny other p hylu m N ot o n ly do al l member s of
.

the group s ecrete s hel l s s uited for pre s ervatio n while ,

“ ”
retai n i n g s om e evide n ce Of s oft a natom y ;but there
i s rea s o n to b e l ieve that the e arl ie s t Brachiopod a k n o wn
a r e ve r y nearly the s i mple s t a n d m o s t a rch aic O f the r
i
ki n d I n t he Ca mbria n R us te lla ce a there i s perpetuated
. .

the pri m itive s tage o f evol utio n whe n ad ult s were

e nca s ed in s hell s hardly d i ffere n t ( s ave i n s ize ) from the
“ ”
fi r s t po s t larval p rote gula O f later type s A l arge
- .

proportio n of k now n B ra chiopod a are no t o n ly fo ss il but

 L OW E R PALAE OZ OI C F A U N A S 14 1

Palaeozoic ;duri n g tha t era they rivalled in nu m ber s ,

a n d di ffere n tiatio n the later achieveme n t s of M ollu s ca

, .

The simple i n articul ate largely cor ne ou s order

, ,

Atrem ata e nj oyed i n the Lower Palaeozoic s u ch

repre s s ed prom i n e nce a s it h a s attai ned A fter the .

cl os e of the S ilu ria n peri od but o ne fam ily li n gered to

,

pe rpe tuate the s toc k i n th e exi s ti n g fau n a The .

exceed i n gly rapid a nd relatively profu se s pecializatio n

, ,

of fam ilie s i n C ambria n a nd O rdovicia n time s i s

s ugge s tive Of n ea n ic acceleratio n ; a nd s i n ce the ,

Atrem ata are u nd oubtedly a n ce s tral to all other orders

of Brachiopo d i dicate a l ate pre Cambria n st age
s n , s -

as the tim e of i n ceptio n of the phylu m The Lower .

C a m b ria n K a s tella ( u n k n ow n from Britai n) ca nn ot be

far rem oved i n m orphology from the fi rs t d efi n ite
Bra chiopod I t s co n tem porary K u torgi na s h ow s m odi
.

fi ca ti on s poi n ti n g toward s the Protremata Th e mo s t .

abu n da nt fam ily o f Cambria n Brachiopod s the O b o li d a e , ,

i s repre s e n ted by L i ngu lella ( Pl ix fi g 8 ) i n the m iddle

. . .

a n d upper Cambria n O f thi s cou n try a n d wa s co n ti nued ,

i nto the S il uria n by the m a s s ive phylogero ntic s tock of ,

the T ri me relli d s S ome early O b olid s s eem to have

.

give n ri s e t o the Neot re ma t ou s group O b ole llid ae I t .

i s probable though le ss certai n that the s am e s erie s O f

, ,

Atrem ata are on the d irect l i ne of de s ce n t O f the

Tel otrem at a The L i n gu lace a de s ti ned to s urvive with
.
,

s carcely a n importa n t m odificatio n u n til the pre e n t

y s

day s eem to be co n ti n uou s with the O b olid group

,

repre se n ted by L i ngulella appeari ng fi r s t a nd i n

, ,

greate s t profu s io n i n the O rd ovi cia n L i ngula it sel f

,
.

wa s ofte n exceed i ngly abu nd a n t i n s hallow S il u ria n s ea s ,

a nd attai n ed m uch s pe cific di ffere n tiatio n d u ri n g t h a t

period I n pa rticular the bed s O f L i ngula cor nea a n d L

. , .

mi ni ma i n the U ppermo s t S il uria n are de s erv i n g O f noti ce .

The s m all c uriou s ly s pecialized m ember s of the

,
 142 I N VE R TE B R AT E PALA E ON T OLOGY

N eo t rema t a ca n b e cl a ss e d i nto four gro up s t wo of ,

which were res tricted to the Lower Pal a e ozoic The .

other t wo ( D is cin a ce a a n d Cr a n i a ce a) did n o t app e ar

u ntil the O rd ovicia n a n d l ike the co ntem pora neou s , ,

L i n gulid s ha ve proved per s i s te n t though s howi n g th eir

, ,

greate s t d i ffere ntiatio n i n pre-D evo n ia n time s A t r otr eta .

i s the bes t k n ow n ge n u s O f N eotrem ata occurri n g i n the

Briti sh Cam bria n a nd O rd ovicia n Or b it a loi a ea ( Pl ix
‘

,
’
. .

fi g 9 ) i s equ al ly fam ilia r i n the We nl ock a nd Lu dlow

.

s tage s of the S iluria n .

V e ry few Protrem ata occu r i n the C ambria n but ,

arch a ic form s O f O r th a ce a a n d Pe nt a me ra ce a a re k n ow n .

B illi ng sella a n d E oor t/ t i s are the o n ly type s certai n ly

recogn ized i n Britai n I n the O rd ovicia n the O rthid ae .

,

aro se a nd reached their acm e Or t/zis ( Pl ix fig . . . .

Plector t/zis a nd H eber tella were abu n da n t ge n era Com .

parable a cceleratio n wa s s hown by the S t rophom e nid ae ,

S tr op fzomena ( PI ix fi g I i ) R afi nes ga i na a nd P lecta m

. . .
,

b omtes occurri n g i n great profu s io n i n s om e bed s of the

'

Upper O rdovicia n T he fi r s t O f the C/zonetes s tock .

—
,

from which the Upper Palaeozo ic Prod uct id s were ,

ev ol ved m ade i t s appeara n ce i n thi s period S m all

, .

Pe nta mera ce a s uch a s Cli ta mbon i tes a nd P ar a méom tes

’

, ,

a re l ocally abu n d a n t I n the S il uria n the O r t h a ce a .

,

S t rophome n acea co n ti n ued the a c ma ic s u cce s s

[

a nd .

a ttai ned i n t h e previou s peri od Plector tb i s D a lma nel/a , ,

a nd B i lob i tes ge n erou s ly repre s e n ti n g the former a n d ,

L ep ta ena S tr op k on ella S tr ep t i s a n d Cfi om tes ( P1 ix fi g:

, ,
. .

1 2 ) the l atter Pe n ta me r id s attai n ed their ac me i n thi s

.

period ;the s m ooth P en ta mer us form s bed s of li m es to n e

i n the Lla n dovery the partly plicate Gyp i a u la abou nd s I n
’
,

the Wen lock a n d the large eve n ly ribbed Comk ia i u m

, ,
-
’

( Pl ii .fi g 1 ) build
. s mu ch. of the Ludlo w Lim es to n e .

Telotrem ata are the lea s t importa n t order O f Lower

Pal aeozoic Brachiopod s ; a n d were it no t for the ,
 1 44 IN VE R TE B R AT E P ALA E ONT OL OGY

the s e excelle nt qual itie s which re Sult i n abu n da nce O f

,

palaeo n tol ogical m a t erial Mollu s ca are i n the m ai n

,
)

, ,

un s a t i s fa ct ory f o ss i ls The s hell s are wholly exter nal

.

s tructu re s a n d s o u s ually fail t o receive m uch im pre s

,

s io n from peri s habl e m orphological character s S tudy .

of rece n t type s s how s that the featu re s O f chief import

a n ce in taxo n omy are s e n s o ry re s pirato r y reproductive

, ,

a n d m a s ticatory orga n s Al l e vide n ce of the s e q ualities

.

i s u s ually i n acce s s ible i n fo s s il m aterial s o that th e ,

cl as s ificatio n O f exti nct Mol l u s c s i s i nevitably s om ewhat

empirical S ter n nece ss ity a nd fortu n at e coi n c ide n ce
.
, ,

have m ade the Cephal opod a reveal Character s O f tax o

n om ic value ;i t i s probable that fu rth er re s earches on

Pelecypod s a nd Ga s tropod s may m ak e palaeo ntol ogical

cla s s ificatio n po s s ible Moderately su cc es s ful attempt s
.

have bee n m ade i n the ca se O f Bivalves but U n ivalve s ,

are s t il l group ed i nt o fam ilie s by a n alo gy rather tha n

m o rphology .

O f the five cla ss e s O f Moll us c s s ui t able for fo s s iliza

tio n Ga s tropod s alo n e a re defi n itely recogn ized i n the
,

Cambria n I t i s po s s ible that Pelecyp od s a nd Cep ha

.

lop od s m ay occur but m uch u ncerta i n ty s u rro un d s

,

the O b s cure a nd p ri m itive type s r e corded Ca mbria n .

“ ”
B ivalves have o n s everal occa s io ns proved to be
Bra nchiopod Cru s tacea (o r eve n d i s torted Brachiopod s) ;
while pre s u m ed Or thocer a s form s from that horizo n -

migh t be a s c ribed to utterly di ffere n t org an i s ms

equal probability Both Pelecypod s a nd Cephalo
.

were repres e nted i n the O rdovicia n where they were ,

accom pa n ied by the earl ie s t A m phi n eura S caphop od s .

po s s ibly occ ur i n the S il urian but thei r s imple tubul ar,

s hell s are d i fficult to d i s ti n gui s h from tho s e of Pteropod s

or eve n Tubicolou s A n nel id s .

The Pel ecypod a are d ivided i nto three s ubcla ss es ,

P rio no d e s m a cea A nom alod es ma cea a nd Teleodes mac ea

,
.
 L O W E R PALAEOZ OI C F A U N A S 145

The t wo la st -n am e d s eries m ay po s s ibly occur i n the

O rdovicia n but L ower Pal aeozoic form s are p rob le
,

ma t ica l a nd rare Two ge nera Or tk od on tzseus a n d

.
,

A lloa esma are s ometim es clas s ed with the T e le o

’
,

d es ma ce a They were probably S ch iz od ont Pri ono

.

d e s mace a ha vi n g po ss ibly a n ces t ra l relatio n s wi t h th e

more elaborate group Prio n ode s m acea were the chief
.

bivalves of the Palaeozoic ;their promi nen ce i n early

fau na s wa s du e rather to the feeble developme nt of the
'

othe r t wo s ubcl as s e s tha n to i n tri ns ic abu n da n ce I n .

a ctual n u mbe r s M e soz oiE form s exceeded ma ny -fold

thos e O f the earl ier era A n O b s cure s erie s O f Priono
.

d e s mace a ns with u n hi nged valve s ( Palaeoco n cha ) ,

ra n gi n g from the S iluria n to the H ol oce ne is r ep re ,

s e nted i n the form er per i od by a few form s O f which ,

Ca r d i ola i s a familiar fo ss il o f the We nlock a nd Ludlow

s tage s The T a x o d o nt s which i n clude form s who s e
.
,

primitive qualities ca n be a s s e s s ed by d is sectio n ,

appeared i n the O rdovicia n The two ge nera O f chief

.

i n tere s t are N uca ia a nd L ea a both of which have ’

,

pe r si s ted from th e S iluria n to the pre s e n t day with

remarkably little cha nge They clearly repre se n t the
.

“
L i ngula type O f evolutio n wherei n early s tock s are
perpe t uated by almo s t s tat ic m orphogen y The m o s t .

im porta nt S chi z od ont s ( a s eries that occurred i n extrem e

profu s io n i n the Mes ozoic) were the radical s of the
Pteriidae P ter i nea is com m o n i n the U pper S iluria n
.
,

a n d ca n be fo un d i n the O rdovicia n ;while R /i omb opter i a ,

claim e d a s the true m orphoge netic a nce s tor of the

fam ily wa s of S iluria n date I s odo nt s are repres e n ted
, .

by early form s Of P ter i neop ecten i n the S il uria n but are ,

rare The D ys od onts s o familiar u n der the n a me of

.
,
“ ”
M u ss el s a t the prese n t day were perhap s t he ,

co m mo n e s t s erie s of Lower Palaeozoic Pelecypoda .

M od iolops zs is not u n com m o n I n t he O rdovicia n a nd ,

 1 46 IN VE RTE B R AT E PALA EON T OLOG Y

with Or tiwnota ( P a Taxodo nt ) abou n d s i n the Ludlow

s erie s O f the Upper S iluria n ( Pl x fig . . .

Ga s tropoda are s eparated i n to two m ai n s ectio ns

( S trepto n e ura a n d E uthy n eu ra ) by the di s tributio n O f
their chief n erve -tru n k s ; cla s s ificatio n O f the n ext
g rade i s ba s ed o n qualiti es O f the re s piratory orga ns .

M o s t s mal ler Sub d ivi s io ns are recog n ized by the n ature

of the radu lar teeth I t i s clear therefore tha t i n cl u s io n
.
, ,

O f ma n y fo s s il U n ivalves ( e s pecially early form s ) i n s uch

or d er s a n d fam ilie s m u s t b e te n tative A s far a s ca n be .

a s certai ned bo th S trepto n eura a n d E uthy n eura form ed

,

part O f the Cambria n fau n a Both s ectio n s Of the

m
.

for er group ( A s pidobra n chiata a n d Cte nob ra nch i a t a )

s eem to have exi s ted i n tho s e early time s but Pteropoda ,

are the o n ly E ut hyne ura n type s k n ow n The s e form s .

s ea -m oth s are abu nd a nt i n rece n t pla n kto n but are ,

m os tly ill —s uited for pre s ervatio n a s fo s s il s I t i s po s s ible .

that the Lower Palaeo zoic Pteropod s m ay repre s e nt a

truly primitive s tock fro m which n ot o n ly Ga s tropod s
, ,

b ut eve n Cephalopo d s m ay have s pru n g while m oder n

, ,

s e a -moth s per p etuate the early form s with m uch r e

ductio n of s hel l -s ub s tan ce Hyoi i t/i es which ra nged

.
,

throughout the Palaeozoic wa s s u fficie ntly abu n d a nt

,

to be al m o s t a rock former in part s O f the C ambria n

- .

There i s l ittle di s ti n ctive i n it s t ubular s he ll b ut s peci me ns ,

have bee n fou n d i n M iddle Cambria n s hale s O f Briti s h

Col u mbia which by pre servatio n of the s oft ti ss ues
,
-
,

co n fi rm bel ief i n the Pteropod a ffi n itie s of the ge n u s .

A n other group of po ssible Pteropod s was that il lu s tra t ed

by Tefi tat ut i tes ( extraordi narily abu nd a nt i n the O rdo
'

v i ci a n ( Pl x fig
. . .a n d s u rvivi n g to t he D evo n ia n) a nd

Cami /a r i a ( O rd ovicia n to J u ra ss ic lea s t rare i n the ,

S il uria n). T he latter t ype had a partly s ept ate s hell _

with rou ghly qu ad ra n gular s ectio n .

A s pidobra nc h ia t e S trep t o neu r a were es se nt i a llyPalaeo

1 48 IN VE R TE B R AT E PALA E ON TOLO GY

Cephalo p od s which appeared in the O rd ovicia n reach ed ,

i t s acm e i n the Palaeozoic a nd h a s l i ngered with u n ,

di s ti n gui s hed pers i s te nce s i nce the Perm ia n I t is .

i ntere s ti n g to n ote that thi s grou p O f T e tra b ra n chs the ,

earlie s t to ari s e i s s till l ivi ng ;while the A m m o noid s

,
, ,

whos e d iffere ntiation wa s d eferred u n til the D evo n ia n ,

a r e utterly exti nct N o type s referable to D ibra nchiata

.

( the o n ly s ucce s s ful m oder n Cephalopod s

) are k n ow n

from the Palaeoz o ic The m os t abu nd a n t N autilo id s

.

“
o f the O r d ovicia n a nd S ilu ria n are
”
Or t hoce racon e s
- s traight tubular s hell s with s aucer -s haped s epta
, .

O r t/wcer a s ( Pl x fi g v ) i s rare i n the O rdovicia n a s

. . . .
, ,

i nd eed are m o s t other ge n era ;but with i t s or n ame nted

, ,

r el ative D a ws ou oeer a s it i s p rofu s ely di s tributed i n th e

,

S ilu ria n . I n the Ludlow s erie s rem arkable probably ,

phylogero n tic form s allied to Gomp /zocer a s are not un

,
“
com mo n Cyrt oce ra co ne s re s embli n g ben t Or tlweer a s
.
,

form s were repre s e n ted by the S iluria n A scocer a s while

, ,

Op /t i a i ocer a s a n d L i tu i tes s howed s ome degree O f e n

’

r olme nt
“ ”
preparatory to the N autilico ne co nd itio n .

D efi n itely “
N a t
u i lus -like form s s eem no t t o h a ve
d eveloped before the D evo n ia n .

K
( ) A R T H R O P OD A
W ith t he p robable exceptio n of Pro t ozoa Arth ,

a re by fa r the m o s t abu n d a n t I n vertebrates

pres e n t period M ari n e a nd lacu s t ri ne p la nk to

.

with s m all C ru s t a cea whil e la rger aq uatic form s abo u n d

,

i n deep a n d coa s t al water s O n la n d a n d i n the air

.

i ns ect s a nd arach n id s for al l their norm al ly s m al l s ii e

, ,

can m ake other life m i s erable a n d precariou s by t heir

i n s i s te n t h a bit s although,
they excite wo n der by the i n

t r i ca cy a nd bea uty O f their s truc t u re s Mo s t A rth ropods .

“ ”
s ecrete s hell s O f chi t i n but i n mari ne form s a fa i r
,
 L O W E R PALA E OZ O I C F AU N A S I 49

propo rtio n O f l ime s alt s i s add ed Although chiti n i s

-
.

a remarkably re s i s ta n t s ub sta n ce the u s ual delicacy O f

,

Arthrop od coveri n g s a nd the loo s ely s egme n ted quality

O f m a n y type s m ake their s ati s facto ry pre servatio n rare
, .

The n orm al mode of growth by ecdy s i s while provid i ng ,

a s ucces s io n O f e nvelopes from each i n divid ual res ul ts ,

i n prod uctio n of m uch fragme n tary material A more .

or le s s argillaceou s tomb i s n e ce s s ary for prolo n ged

pre servatio n ;but with s uch a m atrix there i s no l imit

to the d uratio n O f chiti n .

F rom evide n ce available there s eem s j u s tificatio n for

,

belief that A rthropod s have been relatively a s abu nda n t

in all recorded period s O f the pa s t as they are to d ay .

I n the Pa la e o iC era Tril obite s s eem to have m ore tha n

filled the p os itio n occupied by Crabs a nd Lob s ter s at
the pre s e n t tim e ;while s mall O s tracod a are s cattered
through s uitable m atrices with equal profu s io n i n the
O rdovicia n a n d Cretaceou s I ns ect s are ill adap t ed for
.
-

fo s s ili zatio n the co n s iderable n umber s k n own eve n from ,

the Carbo n iferou s are s ugge s tive O f their exi s te nce i n

,

va s t s warm s I t i s a s triki ng fact that the n u mber a nd

.

v a riety O f large A rthropod s k now n from the Palaeozoic

‘

are m uch i n exce s s O f tho se of later fau na s With few .

exceptio n s Cambria n form s were Crhs t ace a ;all Lowe r

Palaeozoic type s were Bra nchiata Myriopo d s a nd
.

I nsect s are u nk n ow n before the D evo n ia n .

T rilobites are am o n g the mo s t fam iliar fo ss il s of the

Palaeozoic I n s p ite of their app a re nt com plexity th e s e
.
,

exti nct Cru s tacea ns were amo ng the s imple s t form s of

the phylum The lo ng s ucce s s io n of s eparate s im ilarly
.
,

s p ecialized s egme n t s that characterized m o s t Cambrian

type s ca n be compa red with the s i mpl e repetitio n of

parts i n a caterpillar I n mo s t Trilobite li neage s there
.

appears a progre ss ive te nde n cy toward s i ncorporatio n O f

m o re se gme nt s i nto the pygidiu m ( se e Pl viii fi g 4 ) . . .
1 50 IN VE R T E B R ATE PAL AE ON T OL OGY

the cephalo n never s eem s t o com pri s e m ore or le s s , ,

tha n five .

O f t h e t hree order s u s ually r ecog n ized amo n g T r i lo

bite s two ( Hypoparia a n d O pi s thoparia ) were wel l
,

developed at the lowe s t horizo ns O f the Cambria n .

They reached well d e fi n e d acme s i n the O rdovicia n

-
,

whe n m a n y n e w fam ilie s origi nated b efore th e earlier

o ne s had di s appeared The Proparia aro s e i n ful l
.

di ffere n tiatio n i n that peri od D uri n g the S il u ria n .

Trilobite s co nti n ued to s how s pecifi c abu n da n ce bu t a ,

m arked dec li ne i n fam ilie s s e t i n Thi s became m o re .

m arked in the D evo n ia n after which a few gero n tic

,

ty pe s li n gered to the e n d of the Palaeozoic .

T h e H ypoparia were repre s e n ted i n the Cambria n by

two fam ilie s o nly both of which were virtually re s tricted
“

“
to that period The m i n u te al mo s t larval A gn os tus
.
, ,

s erie s with but two body -s egm e n t s a n d el u s ively s im ilar

,

cephala a n d pygidia are lo cally abu n d a n t i n the Middle

,

a n d Upper Cambria n of Wale s ;the grou p ra n ged from

the Lowe s t Ca m bria n to the Lo wer O rd ovicia n A .

s omewhat di ffere n t s erie s of H i aro e i the

yp po a r a n s s n

latter p e ri od co mp r is i ng larger a n d curio us ly s pecialized

,

type s O f which Tr i nu cleus ( P1 x fig 8) a n d A mpyx are . . .

the b e s t -k n ow n exam ple s .

O pi s thoparia were the m o s t abu n d a n t a n d varied of

Lower Palaeozoic Trilobite s N o fewer tha n eleve n
.

fam ilies occurred i n the Cambria n ;o f the s e three were

ab s olutely res tricted to that s tage a n d the rem ai nder ,

d is appeared i n the s ucc e ed i ng peri od I n the ab s e n ce .

of Graptol ites Trilobite s prove well adapted for zo nal

,

s ubd ivi s io n of the Cam bria n T he s pi n ou s M e s o na cid ae

.
,

with large cephala n u merou s thoracic s egm e nt s a nd

, ,

m i n ute pygid ia characterize the Lower Cambria n ( Olen

,

ellus s tage
) N.early allied Pa r a d o x i d a e O fte n attai n i n g ,

very great s ize , are re s tri cted to t he Mid d le Cambri an H

1 52 I N VE R TE B R ATE PALAE ON T OLOG Y

t he Ca ra d oci a n i s clo sely com parable with Tr i mer us

a nd H oma lou otus of the U pper S iluria n a nd D evo n ia n .

The Ch e ir ur id a e i n cl uded the m os t bizarre type s O f

Trilobite s .C/t ei r u r us it s el f h ad a d i s proportio n ately
wide gl abella while Sp /ta er ex oeltus a nd S ta ur oeep/t a lus
,

had that feature i n flated to a grote s qu e degree Perhap s .

the m os t extraordi na ry member of the group wa s the

S il uria n D ei/ lzou i n which the carapace wa s red uc e d to
i
,
“ ”
mere s ca ffoldi n g except for the gl abella which form ed
, ,

a bulbou s a n d warty protubera n ce The Phacopidae .

-
,

with com pou n d eye s comparable with but i n ferior i n ,

s ize to ,
tho s e of A egli ua were a fam ily hardly les s
, ,

im porta n t tha n t he Ca lyme n id ae Large relatively .

,

s m ooth form s of D a lma u i ti u a are fairly com m o n i n

the Ca ra d oci a n while P ka eop i a ella a n d D a lma ni tes are
,
’

abu nda n t i n We n lock a nd Ludlow bed s All of the s e .

fam ilies except the E n crin ur i d a e pe rs i s ted i n to the

, ,

Devo n ia n but the Phacopidae were the o n ly abu nda n t

,

P roparia of that period .

A m o n g t rue Cru s ta cea the four great group s O f ,

Bra n chiopod a O st racoda C irriped ia a nd M a l aco s traca

, ,

h ave repre s e n tative s i n the Lower Palaeo z oic The .

Bra nch iopod s s urvivi n g at the pre s e nt d ay i n the well

,

k n ow n Ap us are probably the m o s t prim itive Cru s t acea

,

d i s covered a n d are bel ieved to be a n ce s tral to all other

,

m em bers O f the cl a ss The deli cacy of their chiti nou s

.

coveri n g s m ake s them u ncom m o n a s fo s s il s b ut it i s ,

i n tere sti n g to n ote that P r otoca r is a gen u s clo s ely ,

re s embl i n g Ap us occurred i n the Lower C ambria n

,
. .

The s m al l bivalved O s tracoda are ofte n abu n da nt .

Their fir s t occurre n ce wa s i n the O rdovicia n a nd s uch ,

type s a s L ep er a i ti a B eyr ic/ ua a nd E n tomi s may be

'
’
,

fou n d thic kly s cattered i n s hale s of Car a d oc a nd S iluria n

age Cirripede s ( Bar n a cle s) O f Lower Palaeoz oic date
”

are rare a n d u ns ati s facto ry There i s m uch u n certai nty .

 L OW E R PAL AE OZ OI C FA U N A S 1 53

as t o whether ce rt ai n s m al l fo s s il s are d etached Cirriped e

plate s or p rim itive Pelecypod s hell s The curiou s fi r-co n e
.
,

like Tu r r ilep as i s k now n from the Cambria n o nward s a n d ,

s eem s to have di s appeared i n D evo n ia n time s .

Lower Palaeozoic Malaco s traca all be lo nged to the

primitive serie s Phyllocarida u s ually s howi ng s u p er
,

fi cia l res embla nce to s hri mp s Hy meuoca r is from the

.

Cam bria n a n d Cer a ti oea r i s ( Pl x fig 9 ) from the two

, . . .

s ucceedi n g period s are the be s t k now n Briti sh re p re

,

se n t a ti ve s of t he group
.
5

Ar ach n id s of the e ra belo nged to the orders S ynx i

p h o s u r
,
a E u rypterida a n d S corpio n id a T h e fi r st .

n amed group wa s re s tricted to the Lower Palaeozoic ,

a n d i n cluded Trilobite -like form s that were probably

a ce tral to the L i mulus s eries a nd E u rypterid s Tha t

n s -
.

order i s k now n from the Palaeozoi c o nly but ra nged ,

from the Ca mbria n to the Permia n with a m ark ed ,

acme i n S ilu ria n a nd D evo n ia n time s B elti ua a prob .

,

le ma t i ca l E urypterid from the p re -Cambria n i s O f ,

doubtful sy s tematic po si tio n The two m o s t fam iliar

.

members of the order are E u ryp ter us a nd P tei jygotus .

The former ge nu s li ngered to the Permia n but wa s ,

m o s t prom i ne nt i n the Upper S il uria ns The cla ss ic

locality for i ts occur re nce i s i n the I s le of Oe s el whe nce ,

S pecime n s have bee n obtai ned from the fi n e S iluria n

marl s i n s uch perfect pre s ervatio n that they ca n be

detached from their m atri x a nd m ou nted for m icro
s copic s tudy like r e ce n t m aterial P tei ygotus a giga n tic
.
,

form with peculi ar outward s im ilarity to certai n con

temporary An tiarchi ( e g P ter i clzt/zys ) appeared with
. .
,

E ui j /p ter u s i n the O rdovicia n but i s be s t k n o wn i n

'

Britai n from the O ld R e d S a n d s to ne S corpio ns are .

fou nd i n the U pper S iluria n of S cotla nd a n d el sewhere ;

they belo n g to a d i s ti nct fam ily a nd their order wa s
,

n o t fully di ffere ntiated u ntil the Carbo niferou s period .

 C H A P TE R III

U PPER PAL A E OZO I C F A U N A S ( PLs . XI . A ND

( )
A G E N E R A L AC C O U N T
HE produ ct s O f al mo s t all co n ditio ns of s ed i
m e ntatio n are to be fou n d a m o n g Brit is h Upper
Palaeozoic s trata S om e part s of the O ld R ed
.

S a n d s to ne a nd certai n s a n d s to n e s a n d m arl s of the

,

Perm ia n s eem to have accum ul ated o n la n d a s m ou n

,

t a i n s cree s or de s ert d u n es O ther local portio ns O f

.

the O ld R ed S a nd s to n e give evide n ce of tra n s ito ry

lacu s tri ne s u rrou n di n g s while m u ch of t he U pper
, ,

a n d part s O f the Lower Carbo n iferou s s erie s were de

,

po s ited i n fre s h water mea n d eri n g through d elta

,

s wam p s . Broad t idal s a n d fla t s of M id -Carbo n iferou s

-

d ate pro d uced the Mill s to ne Grit while s hale s a nd coral

,

reefs i n dicate d i ffere n t littoral co n d itio n s at lower

horizo ns .The Perm ia n Mag ne s ia n Li me s to n e wa s
formed i n a gul f probably com parable with the R ed
S ea . M uch of th e Carbo n ife rou s Li me s to n e appear s
to have developed on the floor O f a n ope n but n o t ,

deep s e a ; the R ad iola ria n chert s O f the D evo n s hire

,

Cu l m s ugge s t s depo s itio n i n aby s m al depth s .

Great a s i s the variety of m at ri x the s elect io n of ,

d ivers e fau n al a s s embl age s i s s om ewhat limited Ter .

re s t r i a l dep o s it s are o n the whol e

,
s adly u n fo s s iliferou s
, ,

a n d eve n the Coal Mea s ure s with their weal th o f pla n t

,

remai n s e nclo s e but fe w rel ics O f I n vertebrate s The

, .

M a gn e s ia n Lim e s to ne m ay h ave lo s t s om e propor t io n

1
54
1 56 I N VE R TE B R AT E PALA E ON T OLOG Y

a nd o nly occa s io n al i nroa d s O f mari ne co nditio ns reach ed

“
H owever proximity of
”
the Midlothia n l ake ba s i n -
.
,
“
the se a i n d uced clim atic cha nge s there ; the Coal
”
M eas ure co n ditio n s O f the Cal ciferou s S a n d s to n e s t age
are i n m arked co nt ra s t to the de s er t feature s s how n
by the Upper O ld R ed S a nd s to ne M e a n whil e the
.
,
”
D ev o n s hire area s eem s to have u n dergo n e varied
a n d viole n t cha n ge s of level re s ulti n g i n the complex
,

roc k s e rie s O f the C ul m Mea s ure s which co ntai n R ad io

-
,

l aria n chert s i ndicative of deep water a nd A n thracitic ,

m aterial s ugge s tive of al m o s t terre s trial co n d itio n s .

The brief s ucces s O f the Carbo ni ferou s Lim e s to ne

sea i n i nvadi ng
“
Lake Caledo ni a wa s follo wed by ”

diver s io n O f a v as t am ou n t O f s edi m e nt o n to the B ri t i s h

a rea . S a n d ba n k s a
- n d mu fla ts bega n to s pr ead
d -

s outhward s from the Cheviot s s ilti n g u p the s ea -ba s i n

,

a n d s motheri n g i s l a n d s that had d efi ed the s e a it s el f .

Grad ual ly the s e depo s it s prod uced a delt a from who s e ,

m ar sh y flat s m ari n e co nd itio n s were eve n tu al ly excl uded .

M ari ne ba nd s ofte n cro wded with fo s s il s occur i n the

, ,

Lower Coal M ea su re s p b ut s u ch i n tercal at io ns are rare

i n the m ai n part s of the coal beari n g s trata
-
.

A fres h paroxy s m of u pheaval the Pe n n i n e Uplift

, ,

bega n t o a ffect the Briti s h area before the e n d o f the

Coal Mea s ure s a n d co n ti n ued more or le s s per s i s te ntly
,

throughout the Permia n O n the we s ter n s ide Of t he

.

Pe nn i n e chai n topo graphy rem i n i s ce n t O f pa rt s O f the

,

O ld R ed S a nd s to n e l a n d area wa s prod uce d res ulti n g i n

-
,

“ ”
lacu s t ri ne marl s a nd brockram s fal le n from fault
“

s carp s or volca n ic peak s . The va s t pl uto n ic i n tru s ion s

of D evo ns hire a nd Cor n wal l belo n g to thi s epi s ode ;
the ir developme n t m utilated a n d meta morpho s ed all
ea rlier rock -s ys tem s of that d i s tri ct O n the ea s ter n .

s ide a
‘
s n n
y cl i al d epre s s io n ( com pl em e n tary to the
Pe nni ne an ticl i ne) let i nm a gul f of the s ea i n N o rt h
 U PPE R PALAE OZ OI C F AU N A S 1 57

bu t the Mag ne s ia n Li me s to ne therei n

E a s ter n E ngla n d ,
depo s ited gives but feebl e i n dicatio n O f the i mporta n t
mari ne depo s it s O f the E uropea n a nd A s iatic Permia n .

Before the e nd of the period the gul f wa s s ilted up a nd

, ,

al mo s t the whole of the Briti s h area wa s brought u nder

the des olate m o noto ny of the Tria s s ic des ert .

Although ma ny of the group s of orga n i s m s domi n a n t

i n Lower Pal aeozoic times per s i s ted through thes e l ater
period s there is a m arked d i ffere n ce i n the ge neral fau n a s
,

O f the t wo era s Graptol ite s have become exti nct ( for

.

al l practical purpo s e s) ;Brachiopod s though s till ab u n

,

da n t are repres e nted by a cma ic s tages of quite di ffere n t

,

group s from tho se s ucce s s ful i n the pre cedi ng era ;while
Trilobite s have l o s t m uch of their earlier prom i n e nce .

I f four type s are to be s elected a s pa rticularly im po rta n t

i n U pper Palaeo zoic time s the choice would un
,

doubted ly fall o n T e t racor a ls Camerate Cri n oid s

, ,

Productid a nd S piriferi d Brachiopod s a nd Go n iatite ,

Cephalop od s .Ma ny group s of Arthropod s are m ore

fully k now n from th e Ca rb o ni fe ro us tha n from a ny other
‘

ho rizo ns ;but the s pecial n ature O f their occu rre nce i n

the Coal Meas ures m ake s com pari s o n with previou s or
la t er fau n as im prac t icable .

( )
B PR O T O ZO A

The Pro t ozoa are m ore fully repre se nted i n the

Upper Pal aeozoic tha n i n O lder p eri od s F oram i n i fera.

a n d R ad iolaria both pl ay the part of rock -former s i n

the C a rbo n iferou s s erie s ;the R adiolaria n C ul m of

“ ”

D evo n s hire i s o ne of the riche s t s tore s of thes e s mall

fo s s il s at a ny horizo n i n Briti s h s trata I n the N ort h of
.

E n gla nd i mpo rta nt m a ss e s of the Carbo n iferou s Lime

,

s to n e are built O f F orami n ifera I t i s a poi nt O f i ntere s t

.

tha t i n both c as e s th e re s pec t ive ord e rs i ncl ud e ma ny

1 58 IN VE R T E B R AT E PALA EON TOLOGY

s pecie s of s u ch s ize that they ca n be readi ly s ee n with the

n aked eye a n d m ay eve n be s eparated from the m a t rix
, .

D evo n ia n F ora m i n ifera are hardly s uperior to tho s e O f

the S iluria n i n abu n da n ce or varie ty ;a n d i n view of t he
,

O b s cure co n ditio n of Briti s h s trata o f thi s type ca n be ,

ig n ored B ut i n Carbo n iferou s ti m e s the order ro s e to a

.

po s itio n of fau n al im porta n ce Ple n tiful developme nt o f

.

m ore or le s s ope n s e a li me s to ne s m ay accou nt i n s ome

-
,

mea s ure for the abu n da nce of F ora m i n ifera but

, ,

evolutio n al progre s s i n the s ecretio n of d urable te s t s

i s probably a co n tribu tory cau s e N owhere i n Britai n
.

do the Ca rbo n iferou s s tr ata s how the character s of

F orami n iferal ooze but the le s s pure part s of the
,

Mou n tai n Li m e s to n e of N orther n cou ntie s are locally

rich i n the are n aceou s for mS a cca mmi na A com
“ ”
.

parable part i s pl aye d by F us u li na ( a m em ber of the

N u mmuli te fam ily ) i n Upper Carbo n iferou s Lime s to n e s
i n E a s ter n E urope T he l o n g-l ived group of the T e x tu
.

l ariid ae i s wel l repre s e n ted e s pecially by form s with

,

“
a ggluti n ati n g habit Permia n F oram i n ifera corre
.

s p o n d very clo s ely with tho s e O f the preced i n g period ,

but they are n atural ly ve ry s carce in Britai n .

A mo ng R adiolaria the S p u me lla ria s till pred om i nated

,

i n the U pper Palaeozoic the m ore com pl ex N as s e la r i a

'

occur i n rela t ively s mall though far from negligibl e ,

nu m
,

ber s .

( )
C PO R I F E R A
I m porta n t additio ns to the Porifera were i n itiated in
the U pper Pala eozoic All th e type s found i n the O lder
.

s e r ie S con t i nue d ( R eceptac ulitidae di s appeari n g i n the

’

Carbo n iferou s ) while s il iceou s T e t ra cti nellid s a nd the

, ,

C alci s po ngiae m ade their fi r s t appe ara n ce With the

,
.

exceptio n of the H exacti n ellid Hya los teli a s po n ge s are ,

u s ual ly r ar e i n Briti s h D evo nia n a nd Carbo n ifer ous rock s

1 60 I NVE R T E B R AT E PALA EON T OLOG Y

k now n a s Pleu r od ietyu m ( occurr1ng a s c as t s i nc ludi ng a n .

appare n tly s ymbiotic worm ) is a well -k n ow n M iddle

D evo n ia n fo s s il ; the ge nu s l i ngered i nto the Lower
Carbo n iferou s i n which s tage the m a ss ive ho neycomb
, ,

l ike col o n ie s of M iefi eli u i a were the m os t effective

'

repre s e n tative s O f the fa m ily Sy r z ugop or a a form

.
,

s im ilar to the rece n t Tu b ipor a i n habi t i s l ocally ,

abu nda n t i n the Lower C arbo n ifer ou s ;while C/za etetes ,

the o nly Tabulate t o s u rvive beyo nd the Pal a eozoic ,

s ometime s build s m a ss ive s heet s of l ime s to ne .

A m o n g H ydrozoa the S troma tOp oroid e a were O f fir s t

,

i mporta ce i D evo n ia n I eef l im e s to ne s bu t s u ffered

n n - ‘
,
,

exti nctio n i n the C arbo n iferou s D e nd r oid Graptolite s

.
,

whi ch m ay either be pers i s te n t type s or rever s io n ary

m im ic s of early form s are to be fou nd though rarely
, , ,

i n t he D evon ia n a nd Carbo n iferou s ;while t he es s e n tially

S il uria n ge n u s M ou ngv ap tus h a s bee n recorded from the
D evo n ia n . S porad ic di s cove rie s O f s uch a n atu re fail to
d etract from the val ue of the Graptolites as i nd ice s of
Lower Palaeozoic horizo ns .

( )
E E C H I N O D E R M AT A
In re s pec t of E chi noderma t a the U pper Palaeo zoi c
,

coul d be des cribed a s the age of Bl as t oid s Camerate

“
,
”
Cri n oid s a nd Pe ris ch oech i n oid s The se type s all had
.

repres e n ta tives i n the S iluria n b ut no n e s u rvived the

,

Perm ia n The extrem e s carci t y of E chi n oderm s i n the

.

la s t Palaeozoic period i s s omewha t mys terio u s but it ,

help s to em pha s ize the c on t ra s t betwee n C a rbo niferou s

a n d M e s ozoic types .

L a te form s of Rh o mb i fera n Cy s tid s a nd perhap s of

,

A mph or id e a have be e n recorded from the D evon ia n

, ,

b ut t hey are i nco ns picuou s — comparable with s u rvivi ng

Grap t olites i n st ratigraphical ins igni fi ca nce Mi nu t e sca ly .
,
 U PPE R PALA E OZ O I C FAUN A S

Edrioa steroid s u s ually ceme n ted by t he aboral s urface

, ,

perpetuate d the cla s s (or order) beyo n d the D evo nia n .

Agelaer i uus is typically D evo n ia n a nd L ep za oa zseus h a s

’ '
'

been fou n d in the Briti s h Lower Carbo niferou s .

Bl as toid s were pre em i n e ntly D evo no -Carbo n iferou s

-

form s Like other recog n iz able fo s s il s they are rare i n

.
,

the Bri ti s h D evo n ia n but Or b i tr emi tes ( Pl xi fi g 3)

, . . .

t is ma m ay be fou n d i n l ocal
“
a nd P/ ta eu osc/ nests
”
in
part s Of the Carbo n iferou s Li me s to n e The Bolla n d .

di strict O f La nca s hire i s m os t prolifi c i n the s e s m all

form s The D evo n ia n N ucleocr i u us is a precurs or of
.

the form er ge n u s ; P en tr emi tes from the A me rica n ,

“
S ub C arbo n iferou s ( s howi n g petaloid ambulacra a n d
-

elaborate hyd r os p ire s ) is m ore nearly re lated to the la t ter

, .

The la s t k n ow n Bl as toid d ied i n early Permia n tim e s .

I n a d unat e Mo n ocycl ic C ri n oid s per s i s ted i n their

u na s piri n g cou r s e Sy mba taoer i uus i s not u n com mo n
.

i n th e Carbo n iferou s Lime s to n e Both Adu n ata a nd

.

Cam erata failed to s urvive the C a rbo niferou s pe riod ,

but in their s hort ca reer s attai ned grea t abu nda nce .

H exa ct i u us i s a wel l k n ow n D evo n ia n A d u nate ;P la ty

-

er i nus ( Pl xi . . fig 4 ) is o n e O f the com mo n es t Cri noid s

.

of the Carbo niferou s Lime s to ne A cr oer i uus one O f the

.
,

late s t A d u n ata i s r emark able for i n clu s io n of n u merou s

,

acce ss ory plate s bet wee n the b as al a nd radial circlet s ,

givi n g a s pu riou s Camer ate appeara n ce to the calyx .

True Mo nocy clic Camerata were the chief r ock form er s -

amo ng E chi n od erm s They attai ned their acme a n d

.
,

reached exti nctio n i n the Carbo niferou s M elocr i uus i s

,
.

a fam iliar D evo nia n type on the Co nti n e n t but from ,

th e Briti s h s ta n dpoi n t A ct i nocr i nus a n d A mp horacr i uus

( excl u s ively Carbo n iferou s ge n era ) are the m o s t i m

po rta n t Column als O f thes e form s co ns titute m uch of t he
.

C a rbo n iferou s Lime s to n e ( Pl i fi g a nd thei r m as s ive

. . .
,

t egmi na t e crow n s are loca llya b und a nt a nd well pres e rved

-
.

I I
162 I N VE R TE B R ATE PALA E ON T OL OGY

D icyclic Cri noi d s i n s pi te O f t heir l o nger ra nge a nd

,

l ater s pecializatio n s how comparable diver s ity i n t he

,

Upper Palaeozoic I na d una ta are e s pecially abu nda n t

. .

Cup r es s oer i uus with s imple curiou s ly m a s s ive arm s

, , , ,

i s one of the m o s t charac t eri s tic D evo n ia n C ri n oid s .

Cya tkocr i u us s u rv ived from the S iluria n ;while P ater i a

er i n u s
, Gr ap lz i ocr i uus a n d Wood ocr i uus repre s e n ted i n
the Carbo n iferou s the m o s t prol ifi c s uborder ( De nd ro
Cri n o i d a ) of the Me s ozoic Fle x ib ilia were n ot yet
.

abu nda nt bu t the s m al l M esp iloer i uus occur s i n the

,

Carbo niferou s Lime s to ne D icyclic Ca mera tes are few .

,

but s uch a s t hey are corres po nd with their M o n ocyclic

equivale n t s i n ra nge R /i oa oer i u us a nd Gilaer ts ot r i uus
.
’

are the m o s t fam iliar Carbo n iferou s ge nera The l atter .

i s r ema r kable i n po s s e ss i ng hol low bra n ched exte ns io ns , ,

o f u n k n ow n fu n ctio n from the tegme n be t wee n whi ch

, ,

a ri s e the s le n der pi n n ul ate arm s

, .

The o n ly n oteworthy featu re i n the s c anty H olo

th uria n fau n a O f the Upper Pal aeozoic i s the occurre nce
O f true S yn a p t id a e ( i n dicated by characteri s t ic
”
wheel
s picul e s ) i n th e Perm ia n S te lle r oid s are n ot m uch
.

m ore s ati s facto ry tha n tho s e of the earl ier era .

B u n de nbach i s a cla s s ic local ity for D evo n ia n form s ,

which occur a s petrifactio ns i n pyrite s readily cleared ,

from the s urrou nd i n g s late Oriya/ta s ter is a cu riou s .

O phiuroid from the Carbo n i ferou s .

Upper Palaeozoic E chi n oidea belo n g to the peculiar

“ ”
group s ometime s called Pe ri sch oe chi noid e a U n l ike .

all l a t er type s they have more tha n t wo col um ns O f

, .

i n ter a m bulacral plate s a nd m a ny po s s e ss extreme

,

ambul acral m u ltiplicity The t e s ts were com mo nly

.

el lip s oidal i n form ;a n d the coro n al pla t e s though O fte n ,

m a s s ive were i mbricate i n arra n geme nt givi ng flexibility

,
-

comp arable with that s how n by the abe rra n t E chi n o

t h ur iid a e of l ate r time s Two m ai n t re n d s O f m o rpho
.
 1 64 I NVE R T E B R AT E PALA E ON T OL OGY

pur s ued their quiet a nd i n co ns picuou s cour s e while , ,

Cheilo stom ata were n ot yet d i ffere ntiated .

B RA C H I OP OD A
( )
G
I n s pite of t he rapid ly i n crea s i n g el aboratio n of other
phyla ( pa rt icularly Mollu s ca) the B rachiopoda m ai n ,

t a i ne d a po s itio n of fir st i mporta n ce i n the Upper

Palaeozoic . The virtual d i s appea ra n ce o f the
Atremata a n d N eotrema t a wa s ful ly com pe ns ated
by the a cma i c exubera nce of the Prod uctidae am o n g
Protrem ata a n d the m ai n tai n ed prom i n e nce o f
,

S piriferacea with awak e n i n g s pecial izatio n of R hy n

,

ch one lla ce a amon g Telotrem a t a

, I n the Devo n ia n a n d
.

Carbo n iferou s periods Brachiopod s s till cou nt ed a s l oc al

,

rock -form er s a nd they are am o n g the m o s t abu nd a nt

,

mem ber s of the exiguou s Briti s h Perm ia n fau n a .

The o n ly group O f Atremata to s u rvive the S ilu ria n

wa s the la t e s tock of the Li n gul idae L i ngula oc cur s
.

with s ome freque n cy i n the D evo n ia n a nd is there ,

accompa n ied by a few other ge n e ra But i n the .

Carbo niferou s the family decli n ed toward s t ha t hu mble

co n d itio n i n which it s til l ex is t s .

The record of the N eotrema t a is s im ilar D is cina cea .

per s i s t ed i n s ome v a riety i n to the D evo ni a n but became ,

s o red uced i n Carbo n iferou s time s that Or b iculoi d ea

( Pl . iv. fi g 1
.
) w a s al mo s t their s ole repre s e n ta ti ve .

Cra n ia ce a are n ot u nk n ow n from the U pper Palaeozoic ,

but d es erv e n o m ore flatteri n g m e ntio n .

A m o ng Protrem ata the prolific s t oc k of the O rt h acea

,

met it s d oom i n the Permia n While typical genera .

( s uch a s D a l m a uella ) rem ai n ed i n the D evo n ia n the ,

m aj or ity of C arbo n ifero u s form s were clearly phylo

gero n tic . S clt i z op /z or i a with i t s s pi n o s e s u rface a nd
, ,

E u teletes with elaborate or n a me n t ill u s trate l ate phas es ,

i n O rthid evol u t i on . The S t rophome n acea though ,

 UPPE R PALA EOZO I C FA U N A S 16 5

s till s urvivi ng a ls O collap s e d i n Perm ia n ti me s

,
.

L ep ta eua occur s i n the Lower Carbo n iferou s with

characters hardly altered from tho s e of S ilu ria n s pecies ,

while other type s of that group ( e g D a vi d s ou i a ) as s u med . .

the Ne ot re ma tous habit of com plete s e s s ility with

“ ”

atrophy o f the ped i c le Or t/zotetes a n ally of S t rop h o

.
,

m e n a which i s n ot u ncom mo n i n the Carbo n iferou s

Lime s to ne s hows s imilar te nde n cies The fi r s t member s
,
.

of the s u rvivi n g gro up of T h e cid i i d a e made their appear

a n ce i n the C ar bo n iferou s a nd were u s ually m uch larger
,

tha n their de s ce n da nt s The fam ily Prod uctid ae far.

excel s i n importa nce all other Upper Palaeozoic Pro

tremata Ck ouetes i t s s im ple s t expre s s io n, ra n ge s from
.
,

the S iluria n to the Permia n True Prod uctid s appeared .

i n the D evo n ia n a nd are al mo s t the com m o ne s t of

,

Carbo n i ferou s m ari n e fo ss il s Profu s e developme n t of .

s pi n e s , attai n me nt of giga n tic proportio ns a nd pro ,

g r e s s i ve lo s s of ped i c le -fu n ctio n s how th at P r o d uctus

( Pl . xi fi
. g 6 ) .i s i n , s om e way s a phylogero n
,
tic s toc k
a mo n g Protremata Ceme n tatio n either by s pi n ou s
.
,

outgrowth s or actual Ne o tre ma t o us adhe s io n be cam e ,

m ore a n d m ore typical a s the group pa s s ed through the

Carbo n iferou s to the Perm ia n ; it culmi n ated i n the
extraordi ary n H i
pp u r i tes -like R ic k tlt of eu i a O f the l atter .

period . Pe n t a me r a ce a li n gered through the U pper

Palaeozoic bei n g repre s e n ted by s uch s m al l a n d
,

s pe cialized type s a s Ca ma r o / r i a i n the Carbo n iferou s

p t o

a n d Perm ia n I n that pe riod they d i s appeared

. .

Telotrem ata s howed wel l -d e fi n ed membe r s of all three

sectio n s i n the D evo n ia n T h e R h yn chone llid s had
.

i n crea s ed far beyo n d their S il uria n quality P ugu ax ,

fig Hyp otlty mtli ua a nd A llor /ij uc/z us bei n g

'

.
f

particularly abu nd a n t i n the Carbo n iferou s Lim e s tone .

T e re b r a t ula ce a with s hort loop s appeared for the fi r s t

ti m e i n the D evo n ia n Thes e T e re b r a t uloid a bra n ched
.
1 66 I NV E R T E B R ATE PALAE ON TOL OG Y

o uti nto three fam ilie s two O f which ( a nd a n im p orta nt

,

part of the third ) are re s tricted to the Up per Pal aeozoic .

Ceu tmu ella S tr i ugoeep /z a lu s M ega la u ter is

, , a nd D i ela s ma
( Pl xi fi g 8) repre s e nt the s e early s hort l ived e x p e ri
. . .
,
-

men t s O f the s erie s True Terebratul a s are u n k n own

.

before the Tria s A m o n g the S piriferacea the be s t

.
,

k n ow n a n d mo s t fully s pecial ized repre s e ntative s are of

D evo n ia n a nd Carbo n iferou s age The A t ryp i d a e .

l i n gered i n to the D evo n ia n but were overs hadowed by,

the S piriferidae Sp i r if er ( Pl xi fi g 9 ) it s el f ( in the

. . . .

broad s e n s e of the term ) i s s carcely le s s com mo n i n the

Carbo n iferou s Li me s to ne tha n P r od uctus a nd attai n ed ,

peculiar proportio ns i n s uch s ectio ns a s Sy r i ngot/tyr is .

The S ue s s i id a e which s u rvive d i n to the J u ra s s ic are

, ,

rarely abu nda n t a n d the M e ri ste lli d a e dwi n d led to

,

exti nctio n i n the D evo n ia n Two ne w fam ilie s aro s e .

i n that period The a n om alou s U n cit id a e with d or s o

.
,

lateral pouches a n d ro s trate form are typ ically D evo nia n , ,

but pers i s ted to the Perm ia n ;while the s uperfi cially

s i mple but i n wardly Com plex Athyridae are a mo ng
the m o s t characteri s tic U pper Palaeozoic Brachiopo d s .

A t/ty r is i s locally abu n da n t i n the Carbo n iferou s Lim e

s to n e ;i t s aberra n t ofte n cem e n ted d e s ce n
,
da n t s reached ,

their pha se o f gero n tic accelera tio n i n th e Tria s .

H
( ) M O LLU S C A
U pper Pa la ZO I C M oll u s ca s how a m arked i ncrea s e
i n n u mbers a n d variety u po n the exiguou s fau n a of the
precedi n g era O pportu n ities for di s covery of terre s trial
.

a n d fre s h -water ty pes have bu t s light i nflue n ce o n thi s

quality M a ny ge n era a n d fam ilie s of Pelecypod s that

.

aro s e i n the S iluria n reached their acm e s i n the Car

b o n i fe rou s while a n eve n greater n um ber of fre s h
,

type s appeared Ga stropod s mai n tai n ed thei r previou s

qualitie s but s howed s m
.

,
all te nde ncie s t o d iffere n tiatio n
 168 IN VE R TE B R AT E PALAE ON T OL OG Y

( probably brack i s h ) hor i zo n s i n the Coal Mea s ure s ,

while A r caa uod ou from the O ld R ed S a nd s to ne ( doubt

fully placed i n thi s group ) is the O lde s t k n ow n fre s h
water Pelecyp od I s od on ta bega n to s how s om e p ro
.

gre s s i n Carbo n iferou s tim e s ; both of the com m o n

s tock s ,
P eeteu a n d L i ma d ate from that period ,
But .

the m o s t importa nt m ember of the s erie s is P ter i ueo

p e ct e u ( Pl xii
. fi g . o n e of
. the com m o n e s t fo s s il s o f
m ari n e ba n d s in the Lower C oa l Mea s ure s M oa i olus .
’
,

a li vi ng ge n u s of D ys od o n ts appeared i n t he D evo n ia n , ,

while the rock bori n g L i t/i oplzagus h a s bee n recorded

-

from the C arbo n iferou s The o n ly Palaeozoic fam ily .

of A n oma lod e s macea ( Ph o la d e llid ae ) i s repre s e n t ed by

ll
A o is r m a a m u s el s haped s hel l n ot u n com mo n i n the
, s -

Carbo n iferou s but rare i n the Perm ia n R ecord s o f

, .

Teleode s m acea from the Upper Palaeozoic are doubtful ,

bu t Pleu r op /tor us if rightly pl aced i n thi s order gave

, ,

it abu nda n t repre s e ntatio n i n the Perm ia n The d e .

~

ge n erate burrowi ng type s O f Pitatus a nd Ter ea o are s ai d

,
'

to occur i n the Car b o n iferou s but quite d i fferen t a n i mal s ,

might have prod uced t he c rypt s a s cribed to them .

A s pidobra nchiate S trepto neu ra retai ned their prom i

ne n ce am o n g Upper Pal aeozoic G a s tropod s Ma ny of .

the ge nera th at occurred i n the S il uria n s uch a s B elle ,

r opltou ( Pl M u r clt zs ou ia a nd E uomp /za lus ,

'

xii fi g. . .

s urvived i n to the l ater era with equal or greater , ,

n u m ber s . The E uomp h a lid a e are particularly charac

t e ri s t i c of the Carbo n iferou s Lime s to n e E u omp /za lus ,

( Pl . xii .fi g S.e a i z os to m a S t r ap a r ollu s a n d,

P l a tys c /ti s m a

bei ng fa m iliar ge n era The s m all lim pet -l ike Fis s urel
.
,

li d a e m ade their appeara n ce at that horizo n ;N a t i eops is

wa s perhap s a precu rs or of the com m o n N eritidae of
to -d a y A m o n g Cte n ob ra nch i a t e type s few i n n ovatio ns
.

were m ade at thi s s tage Gym n oglo s s a were s till rep t e .

s e n ted by M at r ocitezlus e sp ecially by L ox ouema ;

'

a nd ,
 L
P AT E X I
 HLATE OQ
II

U PPE R 31 AL4 Eoz 0 1 0 3F Oss ILs

’

F IG . I .

F IG 12 .
‘
Ptcr i u eop eeteu papy mr eus .
L w rC
o e oa l Me asuring S ta lyhr idge, z 4
FIG .
3 . E u omp/z a lu s p en ta ugu la t u s . Ca rb oni fsfous
‘
Li ms
e
‘
tolih, Ca s le t on
t .

F 111 .
a aiizmp m
m
sp :
“ ’
Carb én rfe rsus Li mestOhé, K
‘
mdaL W
Esm
”

1afi i
FIG .
5 . Mhb s u iu . l d d le Devmii a n , Ga
’

me rs “ ? 5 JO ‘1

f t’ "
l I Gi Gi Gl ypj lli déef as‘ cr eui st r z ? “
‘

a Cér b o niferb us Lmi est one P0 11

“v u lafi i
: 9 Va ishl
’
I
'

I sle of M an . ,

( After W ood r
wa d ) .

P1 11 .
9 1 4 .,»
1 1
q aeu fi z z iia ta : o n Red sai d s tdnaw
wméet aufi e s
)
4 “
 L
P ATE X I I
1 70 IN VE R TE B RAT E PALA EON T OL OGY

the s hel l -cavi ty I t i s quite re s tric t ed to the D evo n ia n

.

( where with
,
i t s a l lie s it occur s
, i n abu n da n ce ) a n d ,

probably repre s e nt s a n u n s ucce s s ful n ea n ic expe ri me nt

i n the early evol utio n o f the order Typical Go n iati t e s
.
,

s u ch as A na r ces tes Tor uoeer a s A a u i a es a n d Gep ky r o

'
, g ,

cer a s a re n o t u n com m o n i n the D evo n ia n

, a n d o fte n ,

s ho w s t r o n g i n vol utio n a nd pro n ou n ced s eptal corruga

tio n The la s t -n am ed ge n u s is i n tere s ti ng s i nce it wa s

.
,

probably d irectly a n ce s tral to the fam iliar Cer a ti tes

s tock of the Tria s The com m o ne s t Go nia t ite s of
.

the Carbo n iferou s are P r oleea u i tes a nd Glyp /zi ocer a s

( Pl . xii fig 6.
) (
. Lower a n d U p p er Carbo n ife rou s Lime
s to n e re s pectively
) a n d G a s t r i oce r a s ( Coal Mea s u re s ) .

I n s ome pa rt s of the Briti s h Carbo n iferou s s eque nce the

'

fi ve la s t ge n era are al m o s t rock -formers Ga s tr i oeer a s

‘

bei n g pa rticularly abu nda n t i n m ari n e ba nd s o f the

Lower COa l M e as ure s a nd s upplyi n g m uch O f the cal
,
“ ”
ca reo us m atter that cau s ed developme n t O f bul lio ns
“ ”
a nd coal ball s
- i n s uch s eam s I n the Perm ia n
.
,

M edlieotti a co nt i n ued a nd ampl ifie d the qualitie s s how n

i n P r olet a u i tes ;while P op a u oeer a s repre s e nted a s im ilar
po s t -G o niatite s tage wit h c om plete i nvolutio n a nd
,
”
practically A m m o n ite s utures i n a d i ffere n t l i neage , .

N o A m mo noid s have bee n fou n d i n the Magn es ia n

Lime s t o ne ;the u nhealthy w at er s O f the e n cl o s ed gul f
i n which that depo s it wa s form ed were evide n tly re p ug
n a n t to robu s t type s N o form s referable to D ibra nchiate
.

Cephalopod s have yet bee n d i s covered i n the Palaeozoic ,

al though B e le mn oid s ( probably m od ified from s ome O rtho

ce r a t i d a n ce s tor were defi nitely e s tabli s hed i n the Tria s
) .

A RT H ROPO D A
K
( )
The t wo Arth r opo d group s m o s t characteri stic of the
Lower Palaeozoic ( Trilobita a n d E u rypterida ) pe rs i s ted
throughout the late r period s bu t s howed rapid d i minu ,
 U PPE R PALAE OZ O I C FAU N A S 17 1

tio n i n importa n ce Both di s appeared com ple t ely i n

.

the Perm ia n The failure o f the s e types wa s i n large

.

meas ure cou nterbala nced by the ri s e of X ipho s ura a n d

S corpio n id a am o ng Arach n id s a n d by the appeara n ce ,

of Myriopod s a nd I ns ect s The s wa mp -co nditio ns of

.

Coal Mea s ures gave u n u s ual opportu n ities fo r pre s erva

tio n o f the la s t two s erie s ;their ab s e nce from Lower
Palaeo z oic horizo n s m ay well be due to lack O f evide nce
rather tha n to actual n on -exi s te nce .

H yp op a r i a n Trilobite s were repre s e nt ed i n the

D evo n ia n by the s ol itary ge nu s H a rp es a form s howi n g ,

m a ny a ffi n itie s with the T ri nucle id ae of the O rdovicia n .

H a rp es com pri s e s i n a curiou s way feature s of over

S pecializatio n i n the cephalo n with revers io n a ry ( or
r etarded ) te n de n cies i n the pygid ium O pi s thoparia .

m ai ntai ned four fam ilie s i n the D evo nia n ;the s e were
reduced to three i n the Carbo n iferou s a n d o ne ( Proétid ae )
'

i n the Permia n Gola i us ( u s ually called B r ou tes) t he

’
.
,

ge n u s that failed i n the D evo n ia n s hows clearly the ,

gero ntic quality o f a large pygidiu m The extra « .

ordi n arily s pi n ou s L ich a d id a e a nd O d o nt op le ur id a e ,

which d i s appeared in the Lower Carbo n iferou s s how ,

comparabl e over -s pecializatio n i n a di ffere n t directio n ;

while Carbo n iferou s a nd Perm ia n Pro etidae combi ne the
feature s O f m ultiple pygidia with warty outgrowth s from
the carapa ce Pk i llips i a ( Pl viii fig 4 a nd PI xii
'

. . . . . .
,

fi g 7 ) a nd Gr ifi t/t i a es are two well k n ow n ge n era from

.
’
-

the Carbo n iferou s Lime s to n e Proparia di s appeared i n

.

the D evo n ia n but w ere repre s e n ted by three o f thei r

,

four familie s i n that period S pi nou s s pecie s of H oma

.

lanatus ma r k e d t h e decli n e o f the Ca lyme ni d a e Cr ota lo ,

cep /za la i s the e n d form O f the Ch e i r u r i d ae ;while a fair ly

-

exte n s ive s erie s of Phacopidae s howed gero n ti c feature s

i n s implificatio n of the glabella warty or name n tatio n , ,

a nd i n s ome ca s e s dege neratio n of the compou n d eyes

, ,
.
172 I N VE R T E B R AT E PALAE ON TOL OG Y

True M alaco s traca occu r fi rs t i n the C a rbo niferou s .

,

though ce rtai n problem atical I s opod s have bee n de

s cri bed fr o m the D evo n ia n The grou p was but feebly
.

repres e nted i n the Upper Palaeozoic .

A rach n id s of the L i mu lus type are n ot u n com m o n i n

-

the Coal Mea s ure s ( where they are ofte n beautifully

pre s erved i n iro n s to n e nod ule s) but L i mulus its elf i s n ot
,

k n own before the Tria s Cyclus B eli nur us ( Pl xii fi g 8 )

.
, . . .

a nd P r es t wick i a are well -k now n Carbo n iferou s type s ,

the la s t beari ng a rem arkable ge n eral re sembla nce to

Trilobite s E urypterid s are fam iliar fo s s il s of the O ld
.

R ed S a n d s to n e where their ofte n e n orm ou s s ize a n d

,

grote s q ue form are Clearly gero ntic characters prepara ,

to ry to their im m i ne nt ex ti n ctio n P tei jygotus S li mou i a

.
,

( Pl. xii . fi g 9)
. a n d S tylon u r u s are am o n g the m o s t
fa m ili ar B riti s h ge n era The la s t -n am ed type s ometimes
.

attai n ed a le n gth of nearly s ix feet E u ryp ter us it self i s.

rare i n the O ld R ed S a n d s to n e but occurs in the Coal

,

Mea s ure s a nd i s k n ow n by one s u rvivo r in the Perm ia n .

S corpio ns a n d S pi d er s occu r i n s om e v ariety i n the

Coal Mea s u res where they are accompa n i e d by M yri o
,

pod s That group appeared fi r s t i n the Lower Old R ed

.

S a n d s to n e of S cotla nd The m o s t prolific l ocality for

.

di s cove ry of Carbo n iferou s Arthr opod s i n E n gla nd i s at

S parth Bottom s n ear R ochdale,
.

The I ns ecta which s how s uch overwhelm i ng pre

,

po ndera n ce i n the rece n t fau n a are fou nd fi rs t i n the

,

C arbo niferou s Gre at n u m bers o f type s occu r there but

. ,

al m o s t al l belo n g to exti n c t ord er s I n s om e re s pe ct s .

these early form s were s y n th etic com bi n i n g featu re s no w

’

di tributed am o n g d i s ti nct order s I ns ect wi n gs ( ofte n

s -
.

d a n gerou s ly like fer n pi n n ule s) are n ot u ncom mo n i n

the s hale s of the Briti s h Coal Me as ure s but the m o s t ,

n oteworthy locali t y for C arbo n iferou s I n s ecta i s un

do ub te d ly t he di s trict O f Com me ntry i n F ra nce ,

.
1 74 I N VE R TE B R ATE PALA E ON T OL OGY

The early Mes ozoic hi s to ry of the Briti s h area i s

directly co nti n uou s with that of Perm ia n tim e s D uri ng .

a n i n terval that m ay well have ex t e nded through half

the era the terres trial m ai n ly de s er t co nditio ns that

, , ,

s ucceeded the Pe n n i n e uplift were m ai n tai n ed E m .

clo sed pa t che s o f water that s urvived throu gh the

Perm ia n period pas s ed through the s everal s tage s of
des iccatio n u nt il the whole area wa s covered with
s hifti n g s a nd d u n e s a n d wi n d s wept loe s s
- -
. The R haetic
s tage had prevailed over S outhern E u rope lo n g befo re

a ny m odificatio n of Tria s s ic co n d itio n s a ffected thi s area .

J u s t before the cl o s e of the R ha etic per iod mari ne ,

e ncroachm e n t a n d s teady though l ocal i so s tatic d e p re s

, ,

s io n , brough t the coa tal water of the M id europea n

s s -

“ ”
s e a over part s o f Britai n a n d let i n a gu lf from the
,
”
Atl a n tic i n the n orth -we s t er n part s of the di s t rict .

T HC M a i n R haetic coas t li ne pa s s ed from the ea s ter n

s lope s o f D artm oor n orthward s to t h e n eighbourhood

of Cardiff a n d the n took a n o rth ea s terly cour s e pa s t

,
-
,

the s ou t h of the Pe nni ne s to N orth York s hire with a ,

we s terly exte ns io n toward s the Che s hire plai n A large .

i s la nd or pe n i ns ula covered m uch o f E a s ter n a nd S outh

E a s te rn E ngla nd so t hat the s e a area wa s of the n ature
,

o f a s trait wide n i n g a t i t s two extremitie s I n no part

.

o f the Briti s h area wa s it po s s ible to get far from la n d .

The i n road of the se a over the Tria ic de e rt s urface

s s s -

res ul t ed i n adulteratio n of the water by s olutio n of i n

c ru s ta t io n s The R haetic Bo ne Bed a n d a curiou s ly
.
-

l im ited I nver t ebrate fau n a bear tes timo n y to the

un healthy q uality o f t he R haetic coa s tal belt .

The chief feature s of R haetic topography were

m ai n t ai n ed t hrough the J u ra s s ic period — all B riti s h
depo s it s of that s tage havi n g bee n form e d withi n detrital
limit s Lia s s ic s t rata are largely argillaceou s ;s hal es
.

a nd cl a y s become i n terbed d ed with s a nd s a nd lit t oral

 M ES OZ OI C FA U N A S 175

l ime s to ne s in the O ol itic s erie s Throughout the J ura s s ic

.

period co n ditio ns comparable with tho s e of the Coal

Mea s ure s ( o n a s m all s cal e) O btai n ed arou n d the
H ighla n d coa s t s ;a nd e s tuar i n e depo s it s form the bulk
of Lower O olitic s trata i n N orther n E n gla nd Toward s .

the cl o s e of the O olitic s tage delta -form atio n wa s s hi fted

t o the s outher n area a n d the huge but local m a ss of
, , ,

the Purbeck -Wealde n s erie s was formed .

The Cretac eou s period wa s m arked by t he com me n ce

me n t of grad ual a nd co n ti nuou s depre s s io n whereby ,

the amou nt O f la n d i n the area wa s s teadily dimi n i s hed .

Although i ts m ai n coa s tli n e s eem s n ot to have pas s ed

m uch farther we s tward s tha n that of the R haet ic the ,

Cretaceou s se a becam e ope n by s ubmerge nce of the

E a s ter n I s le
. Littoral facie s of U pper Creta ceou s
”

rock s rem ai n i n s uch fragme nta ry depo s it s a s tho s e of

,

H a ld on H ill i n D evo n s hire the H iber n ia n Gree ns a n d of

,

A n trim a nd outlier s in a n d near the I s le of Mull B ut

, .

calcareou s ooze O fte n al mo s t free from detrital m atter

, ,

repre s e nt s the typical product O f thi s pha s e U pheaval .

brought the s e thala ss ic co nditio n s to a n e n d bu t the ,

s tage s O f i ts O peratio n are lo s t ( i n Britai n) by m uch

de nudatio n of the Upper Chal k i n pre -E oce n e time s .

The M es ozoic era is ofte n called the age of R eptile s

“
,

a nd n o more appropriate de s ig natio n could be applied .

But there are m a ny feature s i n the I n vertebrate fau na

o f the era that are e qually a s d i s ti n ctive if n ot s o
,

s uperficially evide n t a s the reptilia n r égime

, I n Britai n
.

there i s a particul arly marked co n tra s t betwee n the

fau na s O f the U pper Pal aeozoic a nd Me s ozoic s trata .

Thi s diver s ity i s n ot e n tirely due to the lo ng Pe r mo

Trias s ic i nterval duri n g which very s ca n ty p a la e ont o
logical record s are available I n other regio ns where
.
,

fos s iliferou s Permia n i s overlai n by equally prolific

T ria s the exti n ctio n of old type s a n d i nceptio n of
,
170 I N VE R TE B R ATE PALAE ON T OLOGY

n e w on s e is m a nifes t Te t r acoralla S t roma to poroid s

.
, ,

B l as t oid s Camerate Cri n oid s Pe ri sch oe chi no id s C ryp to

, , ,

s t ome a n d T re p os t o me Polyzoa O rthid S trophome n id

, , ,

Productid a nd S piriferid Brachiopod s ma ny grou ps O f ,

Pelecypod s a nd Ga s tropod s O rt hocer a t id a nd Go n ia

,

t it ic Cephalopod s Trilobi t e s a n d E u ryp t erid s are s o me

, ,

of the more i m porta nt s eries of Upper Palaeozoic

I nve rt ebrate s t hat either failed t o s urvive the Permia n ,

o r li n gere d red uced to al m o s t negligible propo rtio ns

‘

, ,

i n to the Me s ozoic I n pl ace o f the heavy ca s ualtie s i n

.

F erm o-Trias s ic evol utio n appeared the H e x a cora ls true

, ,

E chi noid s R hyncho n e llid a n d Terebratulid Brachiopo d s

, ,

O y s ters a n d T rigo n ia s N atica s a nd Pleu rotomari a s a nd

, , ,

above all A m m o nite s a nd Belem n ite s Ma ny of thes e

, .

type s had bee n i n exi s te nce i n the Upper Pal aeozoic ,

bu t their pr om i n e nce wa s po s tpo n ed u n til the Tria s .

I n l ike fa s hio n m o s t O f the type s predom i n a n t i n the

,

Cai nozoic had their origi n in the Tria s ( the Ma m malia

bei n g n ot ewo rthy) but were co nt e nt wi t h a lowly
,

po s itio n in Me s oz oic fau n a s .

( )
B PR O T O ZO A

F oram i n ifera are u s ually abu n da nt eve n to a rock ,

form i ng degree i n Me s ozoic depo s it s but R ad iolaria are

, ,

le s s co ns picuou s The former group wa s s pari n gly

.

d eveloped i n the T ria s ( ab se nt from Briti s h N ew R ed

“

but i n J ura s s ic clay s a nd lime s to ne s i t

bega n to s how that divers ity a nd abu nda n ce tha t
,

cul m i n ated i n the Upper C retaceou s A s rega rd s the .

quality of the t es ts Me s ozoic F ora m i n ifera a re clo s e ly

,

com pa rable with thos e Of the pres e n t day —calcareou s ,

m ultiperforate types predom i nati n g over are na ceou s

forms The mo s t i m por t a nt s e ri e s of the latter type is
.

tha t of t he L ituolid a e ;s om e members of t he fam ily

 178 I N VE R T E B R AT E PALAEON T OL OG Y

s keleto ns withi n the s a n ctuary O f fli nt -n od ules co ns t ru cted

ou t of the rui n s of t heir derm al s picule s ( Pl ii fi g
‘
. . .

I n part s of the O ol ite s a n d i n the Lower Cre

,

t ace ou s Calca reou s S po n ge s ( e s pecially Ph ar e t ro n es) are

,

s u fficie ntly n u merou s to ra n k a s s ubordi n ate rock

formers I n the Lower Batho n ia n of D ors et s hir e H olco
.

sp ongi a a nd R ap /i t d ou ema for m well marked

“
S po n ge
'

bed s whi le t h e A ptia n S po n ge gravel s of F a ri n gd o n

are famed for the abu nda n ce of m a ny s pecie s of the latter
ge n u s The S yco n B a r r ois i a i s fairly com m on i n th a t
.

depo s i t .

D
( ) CO EL E NT E R A T A

The Ool itic s tage is t he o n ly part of the M es ozo ic

s eq ue nce o f Br itai n i n which coral -r eefs are fou n d ,

although s cattered coral s ( usfiua lly s ol itary form s) occur

a t al l horizo n s from the Li as to the Chal k Practically
.

a ll Me s ozoic Coral s are H exacoralla ; a nd i n the ,

ab s e nce of S tro ma t O p o roid s a nd G raptolite s they con ,

s t it ute m o s t of the Coele n terate record of the era .

M o s t k n ow n fam ilie s of H e x acor a ls appear ed in the

Tria s or Lia s a nd Me s ozoic m em ber s of th e group bea r
,

a s tro n g re s embla nce to tho s e s til l livi n g The great .

fam ily of the A s traeidae i s repre s e nt ed by M ou tli ualti a

a mo n g s olitary type s a nd by l sas tr a ea ( Pl xiii fig 1 )
, . . .

a nd Tli et os mi li a ( chiefly i n the O olite s ) amo n g reef

builders O f the o ther fam ilie s the s mall A na b aciid ae
.
,

are com m o n i n the Lower O olite s an d the m a ss ive

‘

Tfi a muas ter i a a nd S tyli ua are importa n t i n the Corallia n .

The l a s t ( p erhap s al s o the fir s t ) of th e T a b ula t /za etetes ,

l i n gered throughou t the era a n d m ay be fou nd i n s om e

,

abu nda nce i n the I n ferio r O olite o f the Me ndip area .

There a re but fe w corals i n B riti s h Cretaceo u s roc k s ;

they are typ ified by s uch s m all s ol itary form s a s,

Tr oek ocya taus a nd Par a s mili a . B ut i n the Uppe r

 M ESOZ OI C F A U N A S I 79

"
Cretaceo u s two pre s u mably H ydrozoa n ge n e ra appear
i n co ns iderable profu s io n P a r k er i a i s a com m o n fo ss il
.

o f the Cam bridge Gree ns a n d a nd P or osp /ta er a i s ,

fam il i ar ( a n d zo n ally val uable ) t o all worker s in the

U pper Chalk Both of thes e ge n era are co ns idered
.

to be n early alli e d t o Hyd r a cti ni a but they have bee n , .

referred by s om e Ob s e rver s to the Pori fera or eve n to

the F oram i nifera .

( )
E E C H I N O D E R M AT A
Although s ome for m s of C ri n oidea are locally abu n
da n t a n d fragme nts of S te lle roid s occur i n m o s t m ari n e
'

depos i t s Echi noid s are the predomi na nt Mes oz oic

,

E chi n odermata The rapidity of s pecializatio n a nd

.

wealth of va riety s how n by that cla s s d uri n g the e ra

, ,

are i n accord wi t h it s exubera nt n ea n ic elaboratio n i n

the U ppe r Palaeozoic S o n um erou s ( a n d withal beauti
.

ful ) are fo ss il S ea U rchi n s i n the O olite s a nd Chal k

-
,

that the utter a b s e nce of Cys tid s E drioa s teroid s Bl as , ,

t oi d s a nd A d u n ate a n d Camerate Cri n oid s i s i n s u fficie n t

,

to red u ce the i ntere s t a nd i mporta n ce of the phyl um .

I na d un a t e Mo n ocy clic Cri n oi d s the so le s urvivi n g ,

order of the s ubcla s s a re fully repre s e nted ;the m o s t

,

n oteworthy type i s S a ccoeoma be s t k n ow n from the ,

S ole nh ofe n S to n e but fou nd i n t he B riti s h K im m e r

,

T li ls s m all fea ther s tar wa s o n e of the very

'' “ ”
i d gi a n.
-

few Mo n ocyclic type s to a s s ume e le ut he roz o ic habit s

by atrophy o f the s tem ; the quality wa s developed ,

a n d i s m ai n tai n ed i n the m aj ority of D icyclic F le x i b i li a

, .

D e nd rOCr i n oid D icyclic I n a d u na ta are es pecial ly

’

a b u nd a nt i n Mhs oz oic depo s it s E ncr i nu s the ty pe o f

‘

.
,

the whole cla ss occurs a s a rOCk former i n the Trias s ic

,
-

Mu s chelkalk P en tacr i nus a s ide li ne of the per s i s te nt

.
-
,

l s ocr i uus -li n ea ge i s a fam iliar fo s s il of the Lower Lia s

,
 1 80 IN VE R TE B R ATE PAL AE ON T OL OG Y

i ts pe n t ago n al column a ls abou nd i n rock s of J ura s s ic

age The t wo m o s t peculiar I n a d un a te s are Ui u tacr i u us
.

a n d M a r s up i tes both re s tricted to the U pper C retaceou s

,
.

U n like m o s t free -s wim m i n g Cri n oid s the s e type s i n ,

c rea s ed rather tha n reduced the s ize o f their calyce s

, , ,

the form er by i ncorporatio n O f n u merou s s m all brachial s

( a s eco n dary Cam erate quality ) the latter by e n large ,

m e nt of the p late s of the normal cycles Bo t h type s .

had exceedi n gly s hort career s that of Ui u ta er i uus ,

i m m ed iately precedi n g t hat of M a r s up i tes They are .

u s ed a s zo n al i n d ice s i n the Upper Chalk F lex i b ilia

.
.

were equally im porta n t i n the Me s ozoic Ap i oer i u us .

“ ”
n ite ) ( Pl
( the pear -e n cri xiii fi g 2 ) g
i a wel l
. k n ow n . .
-

Batho n ia n type S peci m e ns com plete from root to

.

crow n have bee n fou n d i n the Brad ford Clay ;the d is

coid colu mn a ls are abu n d a n t o n s l ab s of F ore s t M arble .

B ou rgueti cr i u us a s m all allied ge n u s i s u niver s ally

, ,

abu n da n t i n the Upper Chal k ;i ts varie d s hape h a s

s ome zo n al s ig n ifica n ce R ai z oer i u us co n ti n ue s thi s s erie s
.

i n m oder n fau n a s F ree s wi m m i ng type s of thi s order

.
-

predom i n a t e at the pre s e n t d ay a n d were wel l d i ffere nti ,

ated i n the Me s ozoic The s mal l s ize a nd delicate .

”
co ns t ructio n of F eather -S tar s m ake them rare a n d
“

ob s cu re i n the fo s s il s t ate M i llet i er i uus ( Pl i fi g .

f
. . .

a type local ly abu n da n t i n the Batho n ia n a nd Corallia n ,

though ak i n to Ap i oer i uus s how s a n early s t a ge i n

‘

atrophy of the s tem A u tea ou (s ea s la t ) i s k n ow n from

.
’
. .

the Lias a n d m ay ofte n be collected d uri n g s earch for

,

Chalk M icro a .

H olothuria ns of e ss e n tially m oder n a s pect occur i n

”
S ole nh o fe n S lat e S t e lle ro i d s are abu nda n t through
.

o ut the era bu t e n tire s peci me n s are rare

,
Op aiod er ma .

“ ”
pr od uce s a local s t ar fi s h -bed i n the Midd le Lia s .

L arge margi n al s of Pha nerozo nate A s teroid s are

a bu n da n t i n the O olit es a n d ubiquitou s i n the Chalk , .
1 82 I N VE R TE B R ATE PALA E ON T OLOGY

ra d iole s . Both of the s e qualitie s were carried far d uri n g

the Me s ozoic but have bee n yet m ore el aborated i n
,

l ater time s S m all s om ewhat ob s cure form s u s ually

.
, , ,

a s cribed t o A cr os a leu i a a nd H emip ea i u a occu r i n the ’

,

R haetic a nd Lia s The I n ferior O olite i s rich i n s pecie s

.

of Ps eu d oa i a a ema H euz s ed i ua a n d S tomee/zi uus ;H emi

’ ’
,

e i a a r i s a n d A er os a leu i a are i m porta n t i n the Batho n ia n

’
.

The Coral lia n i s o ne of the m o s t prolific horizo n s for

D i a d e moi d a ( coral -reefs are favou rite r es ort s of m od er n
t ype s) ;H emi ei a a r i s a n d D it lop oa ia are repre se nted by
’ ’

abu n da n t a nd Characteri s t i c s pecie s The latter ge n u s .

,

with s m all form s of S a lema a n d P elta s tes predomi n ated

'

i n the Lower Cretaceou s ;P ky uz os oma ( Pl xiii fi g . . .

with greater a mbulacral complexity tha n is u s u al i n

Me s ozoic form s i s s om etime s com m o n i n the Chal k
, .

The pro b ably regre s s ive E chin o t h urii d a e repr es e n t e d ,

by the collap s ible ge nera A s tfi euos oma a nd Pitar uto

”

s oma at the pre s e n t d ay were fore s had owed by P ela u

,

et /z i u us i n the Corall ia n a n d E ek i u otlt u r i a i n the Chal k .

The R egular order s were j oi n ed by their I rregular

“ “

o ffs pri ng i n the Lia s if not before T wo d i s ti n ct order s

,
.
,

with very differe nt m orphoge netic qualitie s ,are certai n ly

k n ow n from that period H ole ct yp oi d a the lea s t
.
,

I rregular of their ki n d are rep re s e n ted by Ples i eclt i uus

, ,

which becam e abu n da n t i n the Baj ocia n ;Pyga s ter con

ti n ned thi s s to c k i n the Coral lia n H olectypu s di fferi n g .
,

from the Pyga s te rid a e i n the po s terior po s itio n of the

periproct i s k n ow n from the U pper Lia s a nd ha s
, ,

abu n da n t s pecie s i n the Lower O olite s Cor n bra s h a n d ,

Corallia n D is eoi a ea a n i m porta n t a nd a mb ulacra lly

.
’
,

elaborate d es ce n da n t of H olectyp us i s e s s en ti ally a ,

L ower Cretaceou s a n d Lower Chal k ge nus ; Conulus

ra nged thro ughou t the Chal k but i s abu nda n t o n ly ,

a t re s tr icted horizo ns .

N ucle olit oi d a which show m uch greater adva nce s

'

,
 M E S OZ OI C F A U N A S 1 83

toward s U rchi n character s tha n eve n the l ate s t

H eart -

H olectyp o id s were repre s e n ted i n the Lia s by Ga ler o

,

pygus which l ike Ples i eeai uus i s abu n da n t i n the

, , ,

I n fe rior O olite At that horizo n it wa s j oi ned by

.

N ucleoli tes Clyp eus a n d Pygu r u s

,
Giga n tic te s t s of .

”
Clypeu s s i n ua tus give the n am e to the Clyp eus -grit ,

of Batho n ia n age in the Co tes wold s ,

The s peci es .

i s hi s tori cally i n tere s ti ng havi n g bee n recorded a n d , ,

credited with s u nd ry virtue s i n the seve n tee n th ce ntury ,

.

N ucleoli tes ( Pl x iii fi g . o ne of the s im pler ge nera

. .

o f i t s order i s n ot u n com mo n i n the Cor n bra s h a n d

, ,

ofte n s warm s i n the Corallia n I t ca n boa s t of a s i ngle .

s urvivi n g s pecie s Pygu r us li n gered to the Lower

.

Cretaceou s but i s rarely com mo n i n thi s cou n try

,
.

The Collyri t id a e a peculiar group o f s m all E chi noid s

,

( probably a n aberra n t of fs hoot from the N u cle o li t o id

s tock ) ,
are repre s e nted by abu nd a nt s pecie s O f Pyga
r /zy t i s i n the I n ferior O ol ite Hy bot lyp us from the s ame .
,

s tage wa s perhap s a n i n com pletely s pecialized m ember

,

of the fam ily S pata n goida appeared at the begi nn i n g

.

O f the Cretaceou s period a n d a t o n ce attai ned great

,

prom i ne n ce The imperfectly cordiform E chi n o

.

coryt hid a e are particularly characteri s tic of the period ,

but per s i s t in deep water at the pre s e n t day H ola s ter .

s upplie s two zo nal i ndice s to the Chalk a nd com ,

pri s e s ma ny other com m o n Upper Cretaceou s s peci es .

E e/zi uoeorys a n e ss e n tially Upper Chal k ge n u s ca n be

, ,

fou n d at all horizo n s i n that s tage a nd give s valuabl e ,

s tratigraphical evide n ce by i t s protea n cha n ge s of form .

The s m al l Ofi a s ter i s characteri s tic of high zo ne s T ru e .

H eart Urchi ns ( S p at a n gid a e ) are eve n m ore ab un da n t

- .

H emia s ter i s n o t u n com mo n i n the Gault a nd G rey

Chal k but i t s few Briti s h s pecie s give l ittle i nd icatio n of
,

the overwhel mi ng vigour of differe n tiatio n s how n el s e

where by Cretaceou s member s of the s tock Ep ia s ter .
 1 84 I N VE R TE B R AT E PALAE ON T OL OGY

is rare i n low zo ne s of the Ch al k but abu nda n t a t ,

higher horizo n s M zcm s z er ( Pl x iii fi g 6 ) i s e s pecially

’
’
. . . .

“
d omi n a n t i n the Upper Chal k Two zo ne s are n am ed
.

after s pecie s of the ge n u s ;but without a ny attemp t

a t s pecifi c determ i n atio n it i s po s s ible to recogn ize the

preci s e horizo n of a ny M zcms ter by it s form a nd s urface

’

d etail s .The ge n us h a s s uppl ied s om e of the m o s t

s ati s factory m aterial a v ailable fo r traci ng the evol utio n

of co nti n uou s s erie s l ivi n g am id u niform s urrou ndi n g s .

( )
F PO LY ZO A

A part from a few repres e ntative s of the Cte no s tom ata ,

Me s ozoic Polyzoa were either Cycl o s tom ata or Cheilo

s tom ata The former order s howed a m arked re
.

awa k e n i n g o f s pecializa t io n a s com pared with i t s Upper

Pal aeozoic dorm a n cy The Ce r iop o ri n a perpetuated
.

m a n y of the characters of the Ce ra mop o roi d e a ( from

which t hey m ay have bee n derived ) a nd e ncru s ti ng ,

form s s uch a s B er emcerz a nd Ri mm ed s pread abu nd a ntly

'

, ,

over the s hell s a n d rock -s urface s of the era I t is quite .

exceptio n al to collect s pecime n s of Chal k I r regular

-

E chi noid s that are free from colo n ie s of tho s e form s .

The Cheilo s tom ata ( which are probably direct de

s ce n d a n ts of the Crypto s tom ata ) fi r s t appeared i n the

J ura s s ic but did not become abu nda n t u n til the U pper
,

C retaceou s S heet s o f M emém mfi om are com m o n ly

.

fou n d a s s o ciated with e ncru s ti n g Cyclo s tomata o n Chal k

fo s s il s a n d free growi ng ty pe s are co ns ta n t though
,
-
,
”
fragme ntary i ngredie nt s of fli n t m eal
,
.

Perhap s the mo s t im porta n t Polyzoa n fau na i n the

“
Briti sh Me s ozoic i s that of the F ari n gdo n S p o nge
Gravel s of Aptia n age I n m o s t part s of thi s local
, .

facie s of Gree ns a n d the S po nge s are s ubordi n ate to

“ ”

Cyclo s tom atou s Polyzoa in qua ntity a nd variety The .

 L
P AT E X I I I
 L
P ATE X I II
M
wws E 9 51 “
5
1 . [s ast r aea ex p la n a ta
“ . Coralham8 hefli ngford , Be rk s
, .

Ap i pm nw p ar krm(onig Bra d ford C la y, B r

a d fo rd Wi lt
, s.

;
a )” 1 38 3 . B
( 1 '
l
Pa r a a d a r i s flor zl
I"
g em a
:
m C m
o l
h a n S tan ford m th e V a le
. .

a koemgz U Cha lk , nea r :K i ngscl_ere H a n ts

p
.
p e r , .

u
( T h e recon s truction of t e a p i ca l Sys t e m is hypb lhe tic
’
a l:

FIG

aw
.

Fm . 6 fi u mM
Mk ”. U p p r Ch in;
“
E h i sw
e ell a
“
cc n , H ams :

ar N ew
‘

L Upp r Ch lk
‘
i
‘ ’l “
FIG .
7 . C I fiizy r z s fi
’

li ca i li s . e b y?
a , ne ur
"
Fm
“
h i thoiiia n téek , b
-
éif
‘

surroii B iéafE
‘

. 8 . Ter ebr at u lid , or s

Bra d fo rd vé ?
Fm .
9 . 26 2110 7 2; d zgma . v al

F Spi n f m na cf. walwtti a lL o w a i a s i msb gk

'

, Raé s t o
'
‘
1 0.
uI
y .
 m s "
M éso z oré sros s m i a “w
’

i f 4 “M i th

ml' 1: 8 m
Bth a t!
b g
dst ockt fi bls e t
1 ‘
B urtdt
1

fii
l' n

Tr zgoma
t
FIG . I . s tr z a t c
z. a o a n, ra
‘

9 18 ! v: m“t fl nh n m
Cora llian
1
fla t a
.

FI G Tr zgoma cla w a am
t
.

. 2 . .
, re

FIG .
4 .

m t?
fl

FIG N 6 Bel B ur B r d t o cl
L5 H
at a ?
to
“
on
55 1 3 gQ é s t
6 70 4 1 5
. .
” 5 .

y
”
,
1
‘

FIG . 6 . rk s
o

F-
Ig n
t
Cos mocq a S J aS
f % .

11 Q¥ 9 f 1

Si s g
E1 9 cap fi fi es p
§ U p pe }
g hall
$5 t
.

1 3 ?
'
f gfiu lx s tfi
'
'
“1 13 h
t "

U pp r Ch Wig h
l

m
I

FI G Belemm tella mu cr on a ta ; ét w ilsle of

'

.
9 . . e a lk n o r t .
 M E S OZ OI C F A U N A S 1 85

depo s it wa s at o ne time co ns idered to b e a n ou t lyi ng

“ ”
port io n of the Plioce n e Coralli n e Crag on accou nt
of i t s Polyzoa n wealth but detailed s tudy of the fau na
s howed that it m ark s comparable co n ditio n s of s ed i

me n tatio n i n Lower Cretaceou s time s .

G
( ) B R A C H I O P OD A

S ave for a few dege nerate or ove r s pecial ized -

s urvivor s of Atremata N eotrem a t a Protrem ata a nd

, ,

S piri feracea the Me s ozoic Brachiopod fau n a m ay be

,

s um marized by the broad

“
ge neric term s R ky nckonella ,

Tereb m tula Ter ebm tel/a L z ngu la , Or biculozd ea a nd

'

a nd
'

Cr a ma repres e nted the two fir s t order s with no greater

'

d i s ti n ctio n tha n duri ng the U pper Pal aeozoic S mall .

,

u s ually ceme nted form s allied to Tfzecza ea were the o n ly

'
’
-

Protremata of the Me s ozoic a s they are of the pre s e n t d ay

,
.

The Rhynch one llid s are abu nda n t throughout the

Me s o z oic a n d m ay be fou n d at al mo s t all m ari ne
,

ho rizo n s of the Lia s s ic Lower O olitic a n d Cretaceou s

,

s trata of Britai n Mo s t of the later Me s ozoic type s

.

b ad a s harp parapet arou n d the pedicle ape rtu re a n d

a
-
,

are the nce s tyled Cyclot/zy r zs ( Pl xiii fig but a va s t

’

. . .

s erie s of ge n eric term s are applied to J ura s s ic form s .

There s eem s a fairly s teady te n de ncy toward s i ncrea s i n g

delicacy of or name nt a s the group i s traced from l ower
to higher s tages S ome Lower O olitic type s s uch a s
.
,

R izy nck onellcz cy nocep /za la are almo s t P ugna x - l ike i n

,

form a n d bold ne s s o f plic a tio n ;while the Cretaceo u s

facies s hown i n Cyclot/zym la tzs s z ma or C p lzba tzlzs
' ' ' ’

( Pl . xiii fig .
7) i s .flatte n ed a n d fi n ely ribbed O n e .

s erie s illu s trated by A ca n tfi otky r zs developed a s pi n o s e

’

, ,

s urface s ugge s tive of rac ial decade n ce ;the ge n u s n a med

i
. s locally abu n d a n t i n the I n ferior O olite .

T e reb ra t u li d s were al mo s t more prom i ne n t tha n

1 86 I NVE R T E B R AT E PALA EON T OL OGY

R hyncho ne llid s i n Me s ozoic time s Glossa .

t/zyr zs a n d M us cu la r s howed the typical s m ooth s urface

'

with u ndulate margi n ( Pl xiii fi g while D zcéyot/zy r zs

’ ’

. . .

i n the Great O ol ite a n d Ter eb r a tulz na i n the Cretaceou s

’

, ,

exhibited featu re s of s u r face or n a me nt u n u s ual in the -

group Ma ny Chalk form s (e g Ter eém tu la 6 627 728 62)

. . .

s howed dege neratio n of the ped icl e without a n trace

y
o f acce s s o r y m ea n s of adhe s io n The T e re b ra te llid s .
,

which are the predom i n a n t group of rece n t Brachiopod s ,

appeared i n the Lia s with the s mal l M ega tizyr zs -s erie s

'

that s till s urvive s ;while Coenotfiy r zs which retai ned

’

m a n y T erebratulid featu re s i s k n ow n from the Tria s , .

S m ooth elo n gate type s ( e g A ula cot/zyr zs M in ot/Lyn }

'

, . .
,

a n d Zezl/ema ( Pl x iii fig
’

are n ot u n com m o n i n the

'

. . .

Lower O olite s ; a nd s mall s hort ofte n s u pe rfi cially , ,

or nate ge n era (K z ngefla Tr zgonos emus M aga s a nd Ter e

'

, ,

b r a tella ) a re m ore characteri s tic of the Cretaceou s .

The S piriferacea were repre s e n ted by two fam ilie s

which ra nged through the Tria s i nto the Lia s a nd there ,

s u ffe red exti nctio n The rel atively s i mple a nd per s i s te n t

.

S ue s s i id a e have a fam il iar Lia s s ic member i n s r ifer z na

' '

( Pl. xiii fi g. .Tria s s ic relative s of thi s beautiful

form we re ofte n ceme nted Th e l ate group Athyrid ae .

s im ilarly s u rv ived attai n i ng rem ark able complexity ( of

,

a gero n tic type ) i n the Alpi n e Tria s The d ouble s p iralia .

b losp z r ella or K ommé z na give s triki ng

'

o f s uch ty pe s a s D é
' '

te s tim o ny to the capacitie s of over -s peci al iza tio n .

( )
H M OL LU S C A
The d i s ap peara n ce or decl i n e o f the mo s t prolific [

s t ock s of Brachiopod s at the clo s e of the Palaeozoic

era l eft the Moll u sca i n a predom i n a n t po s itio n amo n g

s hell -be ari n g a n i m al s B ut the abu nd a nce of Me s ozoic
.

M oll u s c s i s not merely co mpara t ive — i t i s ab so lute .

.
1 88 IN VE R T E B R AT E PALA EON T OLOG Y

zo n e in t hat s t age A m o n g Pte rud ae Ps eud omonotzs is

'

.
,

al m o s t a rock -form er i n the R haetic while P ler za

'

, ,

abu nd a nt i n tha t period i s com mo n i n the Lia s a nd ,

Lower O olite s O y s ter s belo n gi n g to all ge n era re

.

cogn ized i n th at protea n group occurred a nd abou n ded , ,

throughou t the era Os tr ea it self i s a rock former i n

.
-

the Lower a n d M id dle Lia s Great O oli te K im meridge , ,

Cl ay a n d Purbeck bed s Gryp /za ea i s on e of the be s t .

k n ow n fo s s il s o f the Lower Lia s a n d i s com m o n in the ,

O xfordia n A lect ry oma i s e s pecially characteri s tic of

'

the O olites but occur s abu nd a n tly i n the Cretaceou s ;

,

while E x ogym i s al mo s t re s tricted to the Upper O ol ite s

a nd Lower Cretaceou s I n the Lower Gree n s a n d
‘

s pe ci me n s o f E x ogym s m
,

u a m m ore tha n a foot

'

in
le n gth are n ot u n com m o n Ca r d z n za a very com mo n
' ’

.
,

fo s s il of the Lower J ura s s ic wa s po s s ibly a de s ce nda n t ,

of the Ca r éon i cola s tock o f Carbo n iferou s date-

In .

e s tuari n e facie s of the Lower O olite s a nd agai n i n the ,

d eltaic m arl s of the Weal de n Umo hard ly to be d is

’

, ,

t i n gui s he d from the fre s h -water M u ss el s o f exi sti n g

r iver s occu rred i n s ome abu n da nce La s tly n oma
'

, .
,

( Pl iii fi pru g from S f the

’

k
’
s e e . .
gs I a n d s . n c z z oa us o

Perm i a n ( through the T ria s s ic ca n be

coll ected i n hu nd red s at m a n y J ura ss ic horizo ns a nd ,

m ai ntai ned it s abu nda n ce i n the Lower C retaceou s .

O n the whole there i s a te n d en cy for Lower J u ra s s ic

,

Tr zgon i a s t o have l i n ear or n ame n t ( e g T s z r z a m Pl x iv

'
‘
. . .
. .
,

fig . while tho s e from the Upper O olite s have p u s tular

s urface s ( e g T . cla vella m Pl
. .xiv fi g Morpho , . . .

ge n e tic pha s e s s howi n g derivatio n of the latter from the

form er type c a n be s tudied on wel l -pre s erved s hell s from ‘

high horizo ns Three fam ilies of I so d o n ta were p a r t icu

.

la rly p ro m1ne n t i n the Me s ozoic Pecti n idae abou n d .

i n the M id dle Lia s a n d i n m o s t clear -water depo s it s

o f la t er d at e I n the Upper Cre t aceou s the s ubge nera
.
 M E S OZ OI C F AU N A S 1 89

A egmjfieczen ‘
a nd C/zla mys
were es pecial ly im porta n t .

S po n dyl idae probably derived from the Pecti n idae a nd

, ,

h
s o wi n g phy logero n tic te n de n cie s toward s s e s s ility were ,

repre s e n ted by P lzca tula a n d Sp ondylus both ge nera

'

bei ng very com mo n i n the Cretaceou s Lim idae are .

very u n iform ly di s tributed through the period s Plagi a .

s toma i s com m o n i n the Lower Lia s a nd agai n i n the

Chal k L z ma tula a nd Ctenos tr eon are e s se n tially O olitic

’

. .

The D ys od ont ge n era My lzlus a nd M od zolus ( Pl x 1 v

' '

. .

fig . both s til l l ivi n g were repre s e n ted throughout the

,

Mes ozoic the latter bei ng e s pecially abu nda n t i n the

,

R haetic a nd Lower Jura ss ic H zpp op od z u m a wel l

'

.
,

k now n Lia s s ic type i s probably ak i n to the M u s sel s

, ,

a n s e em s con ti n uou s with the S iluria n M od zolo

d erie
' '

-
p r z s s s .

The s m all order A n o ma lo d e s macea was m ore ful ly

d eveloped in Mes ozoic tha n later period s Pleur omya .

a n d Gr es s b/a were two abu n d a n t ge n era of the Pleuro

myacidae a family res tricted to the era The former

,
.

ge n u s i s e s pecially com m o n i n the I n ferior O ol ite a nd

Portl a ndia n P kola a omya a nd Gon i omya s i m ila rly
’
.

repre s e nted a n other fam ily but the former h a s o ne ,

s pecie s s till livi n g I t i s es pecially abu n da n t in the

.

Lower Lia s a nd I n ferior O olite .

A mo n g Teleode s m acea all type s of hi nge-s tructure

,

appeared although the m o s t abu n da nt Me s ozoic form s

,

“ ”
had the relatively s imple D ioge no d ont d e n titio n .

Tr ap ez z u m a nd A n ti“: ( Cyp r z na ) r e pre s e n ted t h e Cyp ri

' '

cardiace a A s ta fi e Cr a ss z nella a nd Op z s are com m on

' '

A s t a rt a ce a of the O olite s ( the fi r s t n am e d bei n g m ore

-

importa n t i n th e Ca i noz oic) ;while Cy r ena ( abu n da n t

.

i n t h e We alde n) Ca m z ta a n d D icéms ( e s pecially

'
’

N eocomia n) i ntroduced their re s pective s uper -fam il ie s .

The mo s t extraordi na ry s eries of Pelecypoda k n ow n ,

the R ud i st a ce a were probably d erived from the l as t

,

nam e d type They were ma ss ive s es s ile form s re

.
,
1 90 IN VE R T E B R ATE PALA E ON T OL OG Y

s embli n g O perculate Coral s m ore tha n other Mollu s ca ;

their morphoge n etic a n alo gue s are to be fou n d i n the
'

R i ch th o fe n i id Brachiopod s of the Permia n Like them .

,

R a d iolz les a n d H ipp u r z tes were phylogero n tic form s o f

' ’

s hort d uratio n They were ab s olutely re s tricted to the

.

Cretaceo us period a nd played the part of r oc k -form er s

,

i n s ome d i s trict s Though rare i n the E n gl i s h Chal k

.
,

they are n ot u n com m o n i n lit t oral d epos it s s uch a s the

Ca mbridge Gree n s a n d Lu ci n acea were repre s e nted by .

Ta ncr ed z a ( e s pecially J ura ss ic ) L ucz mz a nd Sp /za er a

' '

n g Cyclo d on ts the Car d i id ae

( Lower Cretace o u s) A m o .

wer e well devel o ped but they a re ra rely ,

P r otoca m za i s l ocally abu n da nt i n the

’
’
com m o n .

Portla n dia n a nd Blackd own Gree n s a n d T e le od ont s .

are rare i n the Mes ozoic but repre s e ntative s of m o s t ,

fam ilies a ppea r ed i n or before the C reta ceou s period ;

while Cor b ula a n d m a n y b urrowi n g form s ill u s trated
'

the degen erate A s th e nod on t a .

S caphopod s are rarely importa n t fo s s il s but ,

L a evid enmllu m i s locally abu n d a n t i n the Lia s a n d ,

A n ta lzs i n the Gaul t

'

M e s ozoic Ga stropod s are chiefly rem ark abl e for the

decli ne of A s p id ob ra nch s a nd t he ri se of Cte nob ra nchs ,

Pleu r otoma ma ( Pl xiv fi g 4 ) c arried

’

a nd Tectibra n ch s
.

. . . .

o n the Palaeozoic traditio n a n d m ai ntai n ed i ts im port ,

a n ce throughout the era ;wh i le T roch one ma ti d a e ( e g . .

A mber leyd a nd Ci rr us ) are ab u n da nt i n the J uras s ic a n d ,

Tr ochidae Ne ri t op s id ae a nd N eritidae occur locally

,
.

'

Gym noglo s s a a re repre s e nted by B ou rget za i n the

Corallia n ; Pte n oglo s s a by a nd

S ola r i u m both abu n da n t i n the Cretaceo u s

,
Ma ny .

im po rta n t type s of Tae n ioglo s s a appear ed Ma ss ive

”

are com mo n i n the Great O olite ,

a nd L z tlor z na Capu lus a n d N a tion ( Pl xiv fi g 5) occur

’ '

. . .
,

i n the B aj ocia n a nd Bat ho n i a n The la s t s e ries is ‘

.
 192 I NVE R TE B R ATE PAL AEON T OLOG Y

In the Upper Lia s Plzyllocer as repre s e n t e d i t s family ,

H ila ocem s wa s a n early H a rp o ce ra t id a n d D a czjyli ocer a s

’

( Pl iv fi g
. .P er ouocer a s ( Pl iv fi g 4 ) a n d Coelocer a s
. . . .

i n t rod uced the im porta n t S te p h a n oce ra t id a e .

Lower O olitic ge nera were i n the mai n member s o f , ,

the l a s t two fam ilie s L i ocems ( Pl v fi g . L ud wigi a-

. . .
,

Opp elia a nd S tr igocem s are abu n d an t Baj oc i a n a nd

'

Batho n ia n H a rp oce ra tid a e a nd S tep lza u ocer as Sp /za er o

, ,

cer as M orp /wcer as a n d P a r k i ns ou i a ( Pl viii fi g 1

, ) gave . . .

s im ilar repres e n tatio n to the S te p h a noce ra t id s t ock .

M a cr ocepli a li tes wa s a ge n u s of that fam ily e s pec ial ly ,

characteri s tic of the Cor n bra s h a nd Callovia n Ly tocer a s .

wa s a lo n g lived type who s e pre s u m ed C retaceou s d e

-

s ce n d a nt s s howed m a n y poi n t s of i n tere s t A m m o nite s .

s how a m arked decrea s e i n abu n da n ce i n the Upper

J ura s s ic Ca r a i ocer a s a n d P er i sp ui ucles co nti n ued the

.
’

S tep ua uocems trad ition i n O xfordia n a n d higher ho riz ons

re s pectively ; while the Cos moce ra tid a e ( a m ai n ly
C retaceou s fam ily ) were i nt roduced by Cos mocer a s
( Pl xiv fi g 7) i the O xford Clay A p id w a ’

. n. . s ocer a s a s .

form res tricte d to the Upper O olite s ;it ofte n occur s a s

l oo s ely -fi tti ng coi l s of i nter nal m ould s of the cham b ers ,

“
givi n g a vertebrate appeara nce that j u s t ifie s the a pp li
c at io n o f the term s nak e -s to ne s t o A m m o nite s .

In the L ower Cretaceou s, gero ntic acc eleratio n ,

coupled with lap s e of m a ny pre viou s ly importa n t type s ,

b ecame a bu nd a n tly m a n ife s t Olcos teplia u us wa s one of .

the late s t of the S te p h a nocer a ti d a e H op li tes repres e n ted

«
,

the Cos moce r a t i d a e a nd D es mocer a s i ntrod uced a ne w

,

family with thicke ne d or co ns tricted apertures A my lo .

cer a s ( a Cos mo ce r a t i d ) s howed u n coili ng characteri s tic

of late evol utio n al s tage s Upper C retaceou s A m m o .

n oi d s were the l a s t p f their ki n d F orm s a s s ociated .

with the L ytoce ra tid ae s howed the i n flue nce of

-

phylogero ntic s peciali zatio n i n s hel l form H a mi tes -

.
 M E SOZ OI C F AU N A S 193

( com mo i he G aul ) prepared the way for B l

’

n n t t acu z tes

( chiefly Chalk Marl a n d Chal k R ock i n Britai n ) i n which ,

th e s tr aight s hell is s uperficially lik e that of B actr i tes ,

though retai ni ng elaborate s u t ure s Tu r r ili tes com mo n .

,

i n the Lower Chal k s howed deceptive re sembl a n ce to a

,

G a st ropod Cos moce ra tid ae were repre s e n ted by H op

.

li tes ( com mo n i n the G ault ) A ca utlzocer a s ( Ce noma nia n) ,

a n d u n rolled form s s uch a s Cr i ocer a s A mylocer as a nd

'

S t ay/l i tes ( Pl xiv fig . The la s t ge nu s is s ometime s

. .

abu nda n t i n t he Lowe r Chal k a nd Chal k R ock .

P a ola d i scus at the l as t -named horizo n brought t he

/
, ,

De s moce ra t id ae to a clo s e ;while S clzloeubac/z ia one of ,

t he com mo n es t fo ss il s of the Chal k M ar l repres e nt e d a ,

fam ily ( Pri onotrop id ae) who se m embers were s t r angely

r em i n i s ce n t of the Tria ss ic Cer a titid a e At t hi s s t a ge .

fo r m s wi t h i n t e ns e i nvolutio n bu t s ep t al s implificat io n , ,

a ppe a red I ua ocjer as a n d e s pecially Ti ss otia had s utu re s

’
.
, ,

almo s t ide ntica l with tho s e of Cer a ti tes No Am m o n ites .

s urvived the Uppe r Cretaceo u s I n the B riti s h Chal k

‘

thei r rem ai ns are rare excep t i n the l owe s t zo n e s a n d

the s hallow -wate r facie s repre s e n ted by
”
roc k -ba n d s .

D ibra n chiate Cephalopo d a m ade their fir s t appear

a nce i n th e Tria s ( A ulaaoceras ) a nd the Belem n i t i d a e ,

vie with the A m mo noid s i n abu nda nce th roughou t

Briti s h ma ri ne Mes ozoic s t ra ta I n the Lia s e s pecially .

their guard s are so th ickly co nge s ted as to co ns titute

l oc a l rock former s V ery large s pecie s oc cu r i n the
- .

O xford ia n s m all o ne s are m o re char a cteri s tic of the

,

Up pe r O olites a n d Lower Cretaceou s The well .

‘

k n ow n B elemni tella ( Pl xiv fi g 9 ) wa s one of the l a te s t . . .

Briti sh m embe rs of the family ;it i s u sed 3 a zo nal

'

i ndex in the higher part s of the Chalk D enerate .

,

form s with reduced guard s per s i sted i nt o the E oce ne ,

b u t the Belem n itid ae were e ss e n ti a lly a Mes ozoic family .

The allied Bele mnoteuthid ae had a s im il ar b ut m ore re ,

1 3
1 94 I N VE R T E B R ATE PALA EON T OL OGY

s tri cte d , ra n e
g S pe cim e ns s h owi n g t he chiti nou s h ook s
.

of t he te t cle
n a s a n,
d t he i n k -bag with its co nt en t s ,

a re not u ncom m o n i n t he Lia s a nd Oxford Clay .

O cca s io n a l rep r e s e n t a t ive s o f the S epioide a ( a

C tt l e -fi s h)
have bee n fou nd i n the Lia s a nd l a t er s t ra t a The fi rs t .

of the O c t op od a ( Ca la i s) occu rs i n t he Upper Cre t a ceou s

of Le ba n on .

K
( ) A R T H R O P OD A
The T r i ass ic period t ho ugh poor i n A rt hr op od
, .

rem ai ns m u s t have bee n a cri t ical tim e i n the evol u t io n

,

of t h e p h ylum . Pr a c t ically a ll t he or d ers ch ar act e ri s tic

o f t he Pal aeozoic failed t o s u rvive t he P ermia n a nd

m
,

tho s e of the Lias h a d a a rkedly m od e rn a spec t .

Me s ozoic Cru st ace a were m o s t ly O s t ra cod a Cirripedi a ,

a nd Malaco s t rac a The fi rs t n amed grou p i s exce s s ively

-

a bu nd a n t i n ma ny pa rt s of the Briti s h Wealde n a n d ,

occu rs i n profu s i on i n t he Chal k s ome t ime s out nu mbe r

,

i ng eve n t he F oram i n ifer a B ar nacle s of s uch fa m ilia r

.

r ece n t t ype s as Pollicip es a nd S ca lp ellum were wel l

r epre s e nt ed i n J ur a ss ic a nd par t ic ularly Cre ta ce ou s
, ,

d epo si t s . s W
I s opod ( ood l ice) a re u s ually r a re fo ss il s
-
,

bu t A r cha eouiscus occur s i n local s warm s i n the Purbeck

.

be d s e s pecially i n t he V ale of W a rd our True D ecapod s

, .

C ab d Lob s t er s ) a ppea re d fir s t i n t he T r i as The

( r s a n .

lithographic s t on e of S ole nh ofe n ( Uppe r Oo l i tic) co nt ai ns

beau t ifully pres erved s pecim e ns of E mma the pr a w n
like Glyp /za ea a nd crab Pa la eocom f s les
( Pl xiv . fig 1 0)
. .

ar e n ot un com m o n i n the S tones fi eld S la t e and

S el bo r nia n re s pec t ively .

A ra ch ni d s Myr iopod s a nd Ins ec ts are not well k now n

,
'

f r om B r i t i s h Me s ozoic s t rat a a ltho u gh “

b ee t l e ca se s a re
- ”

m
,

co m o n i n th e Li as a nd S tones fi eld S l a t e Thos e i ns ec ts .

t ha t occu r s eem re fe rable t o exi s t i ng ord ers s howi ng i n ,

t his cha ra c t er a m a rke d co nt r as t from P alae ozoic types .

1 96 IN VE R TE B R ATE PALA E ON T OL OGY

F ollowi n g t he upheaval that brought Ch a l k form a tio n

to a n e nd a l o n g i nte rval of de nudatio n s upe rve ned
,
.

A t le ngth the S ou t h -E a ster n part s of the Briti s h area

becam e dep re s sed s u fficie n t ly for accu mu la t io n of la s ti n g
sedim e n t .A t the out s et of t hi s epi sod e e s t u ari ne s an d s
a n d cl ay s s ee m t o h ave s pread wi t h practical co n t i n uity
, ,

from s uffol k t o the I s le of Wigh t ;but thi s b road a rea

s hor t ly beca m e d ivid ed i n t o two trough s s epa ra t ed by

t he Wealde n axi s of upheaval I n the Lo nd o n B a s i n

.

“ ”
i n crea s ed but localized
,
s aggi n g
, ad m itted the se a
du ri n g form a tio n of the Lo ndo n Clay ;bu t a rever s e
te n de n cy followed whereby s ed ime ntatio n ( s ave for
,
[

fluv ia t i le depo s its ) wa s preve n ted I n t he H amp s hire

.

Ba s i n 9 s im il ar s erie s of eve n t s occu rred bu t they were

.
,

ex t e nd ed over a l o n ger period of tim e a nd prod u ce d a ,

grea t er bu l k of s edim e nt N o t u ntil the Upper E oce n e

.

“
whe the Lo n do n Ba s i n wa s ful l d id the s ea gai n
( n

u n di s puted s way i n the s outher n trough The s ucceed .

i n g O ligoce n e s trata m ark a re newal of th e s t r uggle

betwee n la nd a nd s ea com pri s i ng depo s i t s that ar e
,

e s tuari ne or lacu s tri ne 1 n the m ai n With the infi lling .

o f the H am p s hire B a s i n s ed ime n tatio n ( of a d urable

,

type) cea s ed i n B ritai n the whole area m ay h ave

,

u n dergo n e co n s iderable elevatio n d uri n g the M i ocen e

“
pe ri od. I n Plioce n e tim e s the N or t h S ea covered
pa rt s of Ea s t A nglia tha t are now la n d a nd left thi n , ,
“
bu t ch a racteri s tic l ayers of crag a nd cl a y
, Thes e .

are o verlai n by d epo s it s s ugges t ive of co nditio ns l ike

t ho s e prevale n t i n the Wa sh a t the pres e nt d ay The .

fo ss il s of t he s ub gl aci al pea ts a nd m a rl s give clear

-

a nd co nvi n ci ng evi de nce of the a pproachi n g cl i mati c

rigo urs of the Lower Plei s t ocen e ; bu t ap a rt from
-

their bo real a s pec t the y are e s s entially m oder n i n fa cies .

Plei s toce n e depo s its are largely Boulder Cl ay a nd

s im ilar gl aci a l re l ic s ; the fluvia tile a nd pea ty be d s ‘
 CA IN OZO I C FA UN A S 1 97

t ha t occu r a mo ng t hem co nta i n few I nver tebr a t e

remai ns .

The Cai nozoic era is the age of Mam mal s a nd

“

”
Bird s . A fter l o n g s tagn atio n i n the Mes ozoic the
, ,

warm blo oded d e s ce nd a nt s of Perm ia n T he romorp hs

-

a n d Tria s s ic D i n o s aur s have by d eferred n ea n ic accelera

,

tio n amply a nd worthily a s s u med the soverei gnty abdi

,

ca t ed by s e ne s ce n t R eptile s i n the C re t aceou s period .

B u t i n the humbler s phere of I nve rt eb ra t e l ife a com

parable revol utio n ca n be recorded The teemi ng .

R hyn cho nellas a nd Terebra t ula s of t he Me s ozoic are

red uced to i ns ign ifica n t n umber s ;Trigo nia s a n d ma ny
o ther Prio node s m acea n Bival ve s s u ff er a like fate ;
Pleuro t o maria s s i n k al m os t to exti nctio n ;A m mo n ite s
have e nti rely go ne a nd Belem n ite s are rep re se n ted
,

by s carce u s ually s mall de s ce nda nt s that b ear little

, ,

s u perficial re s embla n ce to their predeces s or s Ma ny .

of the dom i na nt Cai n ozoic types appeared i n previou s

epoch s ( like the Ma mm al ia) but po s tpo n ed active ,

s pecializ atio n u n til the E oce n e Mos t of the T e le o.

d e s ma ce a n Pelecypod a Cte nob ra nch ia t e a nd Pul mo nate

,

Gas tropoda a n d Nu mmuli t id a nd Glob ige ri nid Fora mi ni

,

fera belo ng to thi s cla s s “ “

Cake Urchi ns a nd S a n d
- .
”

D ollar s ( Clype a s t roi d s ) are ex cl u s ively Cai nozoic while ,

”
bu rrowi n g H eart -Urchi ns ( S p a t a ngid s) though locally ,

a bu n da n t i n the Cretaceou s are e s s e ntially Tertiary

,

form s .

I t i s appropriate i n thi s place t o l ay empha s i s o n t he

fact that there is no co n tra s t betwee n exi s ti n g fau n as a nd
tho s e of precedi ng Cai n ozoic period s that ca n compare
with that di s ti ngui s hi ng E oce ne from Cretaceou s group s .

There is no Palaeo ntological w arra n t for recog nitio n

of a Quater na ry era I n deed it may be doubted if
”

.
,

Cai nozoic hi s t ory ha s ye t reached its middle phas e .

198 I N V E R T EB R ATE PALAEONT OL OG Y
( )
B P R OT O Z O A
The ou ts ta ndi n g fea t ures i n the F oram i n ifera n fa una
o f the C ai n ozoic era are the rem ark able abu nda n ce s ize ,

a nd elabo ratio n of Nu mmuli ti d s a nd M i lioli d s i n the

earlier peri od s a n d the overwhelm i ng pred omi n a n ce of

,

G b g i d s i n late r a d exi s ti g time s

l o i e r n i n n The coi like
. n -

t e s t s of N u mmuli tes ofte n attai ned diameters exceed i n g

two i che ;they were t he chief lime to e buil d ers i n the
n s s n -

M edi t err an ea n d i s t rict d uri n g t he E oce ne a nd O ligoce ne .

M ilioli d s play a corre spo n di ng p a r t i n the co ns t ru c t io n

o f the wel l k n ow n E oce n e
- “
C alcaire g ross ier V as t .
”

n umber s of s mal l N u mmulit i d s (N u mmuli ua ) a nd rela

t ive ly large M i li olid s (A lveoli ua ) occur i n s om e o f the

Upper E oce ne clay s of the I s l e of Wight b ut CFora mini ,

fera are n eve r r oc k -form er s i n B ri t i s h Tert ia ry s trata .

Glob iger i na i s the m o s t abu nd a n t co ns titue n t of fora mi ni

feral ooze i n the late Te rtiary a nd H oloce n e .

R ad iolaria are we ll repres e n ted i n m a ny d i s t ric t s ,

a nd their r em ai ns practically u n al tered are s ometime s

, ,

s u ffi cie n tly abu n da n t to have a n eco n om ic v a l ue The .

“ ”
wel l k n ow n Barbado s E a r th a nd th e eq u a lly famou s
-
,
“
Tripol i Powder are n othi n g m ore tha n de p o s its o f
,

u ncom pacted R adiol aria n oo ze They a re u sed as

.

poli shi n g powder a n d for dil utio n of ex plos ive s Bo th .

the s e d epo s i t s are o f M i oce ne age but the m aj ority of

,_

ge nera com po s i n g them are s till livi ng Na s se la ri a

.

a re co n s iderably i n exce s s of S u me lla ria a nd their

p ,

grotes que bu t exqui s ite te s ts a re fam iliar t o a ll m ic ro

, ,

s cop is t s .

( )
C P O R I F E R A
R em ai ns of S po nges are rare in Briti s h Te rtia ry
depo s its The m uddy and ofte n e s t uari n e co n di t io ns
.

pre vale nt i n sedi me nt atio n a rea s were u nco nge ni a l to

2 00 IN VE R T E B R ATE PALA EON T OLOGY

va ri ety devel oped by F ea t he r-S ta rs S t a rfi s h B ri tt le

, ,

S t ar a d S ea Urchi ns is available i n thi s cou nt ry u ntil

s n -

the p r es e nt perio d i s reach e d I n s ome mea sure thi s

.

s carcity o f fo ss il s i n the Te rt ia ry m ay be as cribed to

the reduc t io n i n m ass ively -built typ es of E chi noderm s

begun in Mes ozoic ti m es S t al ked Cri n oid s a re r are
.
,

a nd h a ve u s ually ret r ea t ed t o a by s m al d epth s Their

.

e le u th e roz oi c offs pri n g a re t oo d el ica t e for n o r mal

pre s ervatio n M oder n S t e lle roi d s a re over whel mi ngly
.

n u merous bu t mo s t of t hem s eem eve n le s s suited for

,

fo s s ilizatio n t ha n their a n ce s tor s The abu n da nce o f

.

E chi noid s which are em i ne ntly a dapted for pre s erva t io n

,

( whether m a s s iv e rock -
d w el ler s or fl i m s y burrower s ) i n

Cai n ozo ic r oc k s of other d i s t ric t s no le s s t ha n i n m ode rn

fau n as s hift s the re s p ons i b ility for the feeble ne s s of the
,

Brit is h record on t o phy s iographical co nditio ns .

Cri noid remai ns are al ways r a re i n the Tert ia ry .

I s om uus ( u s u all y r epr es e nt e d by columna ls ) occur s i n

'

th e Lo nd o n C l ay an d the ge n u s i s t he le a s t r a re o f
,

l ivi ng s t al ked types S uch S te lle roid s as a re k n ow n

.

are e s s e n tial ly m od er n i n fa cie s .

B oth R egul ar a nd I rregula r E chi n oi d s seem n e ar to

t heir acme s i n t he m oder n fa una a nd t heir te s t s occur
,

in rock form i n g profu s io n i n the Med iterr an ea n r egio n

-

a n d p a r t s o f N orth A merica . The Cidaroid a s how li t tle

cha nge i n s t r uc t ure o r n um be r s b ut they s eem to have
, ,

betake n t hem s elves to par ts o f the s e a r e mo t e from

l it t o ral bel ts ( on which they flouri s hed i n J uras s ic t im es ) .

The S a le niid s e rie s of D ia d e moid a i s com parably

retiri n g a n d h a s dwi n dl e d t o ve ry s mall fau n al pro
,

po rti o ns The A rb a ci oid s er ies ( wi th a mbulacra l

.

s t r uctu r e s imilar to that o f H emicid a r i s ) i s fully re p re

s e n ted by A r éacia a nd Coelopleu r us the latter bei n g

,

a b und a n t i n I n dia n M ioce n e depo s it s D ia d ema ( Cen

.

tr eclzi uus) is appar n t ly a r eve rs i o n ary member o f i t s

e
 L
P ATE X V
 L
P A TE X V I

CA I NO ZO I C F oss rL s

FIG . 1. Vm u s cas i u a . Co ralline Crag , S utto n , S uffolk

v .

F IG . 2 . D os i n i a qxp le t a . Red Crag, . Wa lton a

-
k

3 N er g 91 5377 224
.
i a. 5
“ O i g
l c , ea
o e n e H d on H ill, I l
s e o f1 fily hrg t

FIG .

Tu r r z tella i mb r icat a r i a ( B ta clfleSha mJQ Q w i E i-nfl

'

F IG E d
Q s p EEi
'

5 ocene

W ig h
. .
a

Ba y, I sle of t .

FIG . 6 . B a ti lla r i a ser r a t a . Eoce n e ( Ca lca i r gr

e oss ie r) .

FI(
2 1 41 11 1 11 1 J I
FIG . 8 . S u r cu la Pleu r otoma
’
fei et r z u m . E oce n e , B a to n , H an t s r .

FIG . IO . Pla u or éi s euo mp /ra lu s . O li g oce n e , Be mb i d r g e, I sle of Wigh t .

FIG . 1 1 . X a n t /z op s i s b i sp i n as a . E o ce n e , S h e ppe y .
 L
P AT E X V I
2 02 I N VE R TE B R ATE PAL AE ON T OLOGY

sp icuous t h a n the m a s s ive s urface -d weller s S c/ziz as ter

.

( which o ccu rs i n local cl u s t er s i n t h e L o ndo n Clay )

com petes with the Cre t aceou s H emia s ter i n s pecific
d i ffe re n t i a t io n ( Pl xv fig . . while B r is sa s B r i ssop s is
.
, ,

E clzi uoca m i u m M oi r a a nd Sp a ta ngus help to prove t he

’
,

co nt i nu ity of C a i nozoic wi t h rece n t fau nas .

( )
F PO L Y ZO A

The m os t n o t ewor t hy a s s embl a ge of Polyzoa i n

Briti s h Te rtiary s tra t a i s t ha t of t he Coralli ne Crag o f
the E a s t A n glia n Plioce ne Cyclo s t om a t a a re there
.

rep re s e nt ed by m a ss ive colo n ies of Tlzeouoa A lum/a r i a ,

a n d H or ner a but they are i n fe rior i n nu m ber s to t he

,

Cheilo s t om at a of which R etep or a i s the mo s t fam il iar

,

ge n u s Thi s proportio n of t he t wo im porta nt orde rs

.
‘

of Gy mn olaem a t a i s tha t n o rm a l t o Ter t iary fau na s .

Ap a r t from t he C ra gs Polyzo a a re ra re i n the

,

a rgill a ce ou s or s a n d y depo s it s of t he Briti s h Tertia ry ,

bu t at th e pres e nt d ay M emb r a uzpor a a n d F lus tr a

a b ou nd i n sh a llow wa t er arou n d t he coa s ts .

( G) B R A C H I O P OD A
The o n ly s eries of Brachiop od a t o s how any phyletic .

vigour i n Cai nozoic time s i s that of the T e reb ra tula cea ,

a n d a m on g them the T e reb ra t e lli d s ( the l a t e s t group

t o a ri s e) a lo n e a ttai n fau nal promi ne n ce I t m ay be

.

co n fi d e n tly a s s u m ed th at the a bu nda nce of recen t

ge ne r a re c og nized i n thi s s ectio n is the ou t come of more
co m plete m orphol ogic a l k now led ge a nd wil l s i n k i n t o
,

r elati ve i ns i gni fi ca n ce as co nt i n u e d s tudy i s a pplie d t o

fo s s il type s .

The Lingulid s T heci d ud ae a nd Cra nud ae co nti nue

,

their u n as piri n g progress throughou t t he e ra a nd th e ,

R hynchone llid s a re ha rd ly m ore imp ort a n t Ter ebr a tula

.
 CA IN OZ OI C F AU N A S 2 03

is rep re se nt ed i n the Bri t i s h E oce ne a nd a gai n by s uch

,

l arge s pe cie s as T p erf or a ta in the E a s t A n gli a n

.

Plioce ne ;but throughout the Cai n ozoic the T e re b rat

e lloi d s s eem t o have favou red the clea r a nd au s tr a l
water s that t hey n ow i nhabit O f Cai nozoic Brachio pod a
.

i t m u s t he c onfe s s ed that r a rity m ake s them s t ra ti

graphically ne gligible a nd s im ilarity t o rece nt type s
,

red uce s their palaeo ntol ogical i ntere s t R eflecti ng u pon

.

the ove rwhel mi ng i mpo rt a nce of the phyl u m i n

Pa laeozoic time s a nd the abu nda nce of s ome groups in
,

the Me s ozoic it is i m possi bl e to repres s the s e ntime n t

,

co ntai n ed i n the hac k neyed q uo t atio n S ia tr a ns i t glor ia

'

H
( ) M O LLU S C A
Te rtiary Mollu sc a h a ve a defi n itely m oder n facies .

N o t o nly a re the propor t io n a t e val ue s of the seve ral

orders m ai nt ai ne d without cha n ge from the E oce ne t o
the pre se nt d ay but a great n u mbe r of gen era have
,

pe rs i s t ed throughou t t he e ra R ed uctio n a n d de
g e ne r a cy i n the Cephalopo d a are more tha n cou n t er
bal an ced by extreme d ivers ity i n Pelecypod a a nd
Ga s tropod a The extraordi na ry ab u nda nce of terr est rial
.

form s of the la t te r order i n m oder n fau n a s i s probably

deceptive— thew choice of habitat de nie s them ordi n ary
cha nce s of fo s s ilizatio n B ut Teleo d e s macea n B i valves
.

a nd Cte nob r a nchia t e U n ivalve s s how a real i nc reas e

( fore s hadowed i n t he Cretaceou s period ) a nd their ,

u s ual pre fere nce for m uddy coa s tal water h as ena bled
t hem t o lea ve abu nda nt r em ai ns in B riti s h C a i nozoic
depos i ts.

Al though Teleode s m acea a re the predom i na nt Pele

cypo d a o f to -day the old -e s tabli s hed Prio n od es macea
,

are s till import a nt a nd m ay eve n o u s t their mo re

,

progres s ive relatives locally T a x od ont s are repr es e nte d

.
2 04 I N VE R TE B R AT E PALAE ON T OLOG Y

by N ucula ( which is oft e n c om m o n i n the Lowe r

Tertiaries of t he H amp s hire B a s i n Pl xv fi g ,
. . .

Cuculla ea ( chiefly i n the Woolwich bed s) a n d Glyci mer is

( Pl .i ii fi g .
4 a n d P I
. xv fig The . l as t ge n u s
. s o .
,

a bu n da n t on rece n t beache s i s fou n d i n the E oce n e , ,

but i s e s pecially cha r ac t eri s tic of the Plioce ne A mo n g .

S ch iz od ont s grea t reduc t io n on Me s ozoic n u mber s h a s

,

t o be n oted P i uua i s o ne o f the few s u rvivi ng t yp es

.

o f t he Pt e ri a ce a ;it i s n ot u n com m o n i n the Lo n do n

Clay The I nocer a mus -s eries h a s d i s appe a red but Per ua

’

.
,

s till li nger s a n d eve n Ger uilli a h a s bee n fou n d i n t he

,

E oce ne The O s treid a e are th e o n ly S ch i z od on t fam ily

.

t o m ai ntai n s uprem acy a n d Os tr ea it s el f ( Pl iii fi g

, . . .

com m o n i n all bracki s h -wa t er facie s of t he Ter t iary ,

Umo
'

s how s few s ig ns of decli n e a t t he pre s e n t d ay .

oc cu rs i n s ome pr ofu s io n i n O ligoce ne clay s ; but

Tr igou ia i s al way s s carce i n t he C ai n ozoic a nd is ,

r epre s e nted by few 11v1 ng S pecie s The I s od o nt s r e t ai n . ,

s ome mea s ure o f their Me s ozoic im po rta nce Pecten

‘

( Pl . xv fi g . S p on dy.lu s a n d A nom i a are the p re

d om i n a nt type s The fi r s t is es pecially a bu n d a n t i n

.

the Plioce ne Ma ny rece n t s pecie s of Sp ondylus s how

.

com ple t e fi xati o n i n Oy s ter -fa s hio n— a ppa re nt ly i n d i

ca t ive of phylogero ntic s pecializatio n ak i n t o t hat
p revale n t i n m a ny Brachiopod li neage s M oa iolus a nd .
’

M y ti lus repres e n t the D ys od onts bu t the l atte r i s le s s

.
,

a bu n da n t i n the fo s s il s ta t e tha n i t s m oder n ex ube ra n ce

wou ld s ugge s t .

A n oma lod e s ma ce a a re di s t inctly le ss pr o m i n e nt tha n

'

i n the Mes ozoic P aola d omya s till pe rs i s t s a nd i s

‘

.
,

com m o n i n the Lo ndo n Clay Tur aci a a ge nu s tha t .

,

m a de i ts appe ara nce i n the Tria s i s bes t k n own in

.

Britai n fro m the Plioce n e .

Teleo d e s m acea are d evel oped i n overwhel m i ng 0

“

n u m ber s The D io ge nod ont s which were the o nly

.
,
206 IN VE R TE B R ATE PALAE ON T OLOGY

Pli oce ne fo ss il s ;bu t Pleu r otoma r ia ca n s how o n ly fou r

livi n g s pecie s ( a n d few po t Cre t aceou s o n e s ) o f great
s -

ra ri ty The s m al l Trochidae a nd N eri t idae are the m o s t

.

s ucces s ful s e r ie s of rece n t A s p id ob r a nch s Tr oe/ms i s .

a bu nd a nt i n t he Plioce ne ; N er i ta ( Pl xvi fi g 3) a nd . . .

N er i ti na s w a rm i n t he O l igoce n e t heir s hell s ofte n ,

re t a i ni ng t ra ces of col or a t io n A mo n g Cte nob ra nch ia t a

.
,

Gym no gl oss a are repre s e nt ed by s m a l l form s a ki n t o

Tur bomlla bu t t hey a re n ever com mo n S cala r ia is
'

.
,

a n abu nd a nt ge n u s o f P t e n ogl os s a Tae n ioglo s s a a re .

very fully d evel oped L i ttor i na which co n grega t e s

.
,

i n s w ar m s on m o d er n co a s ts is ha rdly le ss a bu n d a nt ,

i n t he Pl i oce n e Cap ulus em ula t es Pa tella i n pers i s t e nce

.

as i n fo rm ;Ca ly p t r a ea a n d X e uop lwr a a r e e ss e n tially ,

C a i nozoic ge ner a N a t icid ae wi t h grea t ly i n flated

.

bod y -wh orl s (A mp ulli ua Pl xvi fig 4 ) are a mo n g the , . . .

c o m m one s t fo ss il s of the E oce ne a n d abou n d in t he ,

Cr a g F res h wa ter V ivip a r id a e s t riki ngly like Vi vipa r a

.
-
,

of m od er n river s a re com mo n i n the O l igoce n e

, Ta r .

r i i ella ( Pl xvi fig 5) i s a t i ts be s t i n t he E oce n e s om e

. . .
,

s pecie s a tt a i n i n g ver y grea t s ize M ela n i a Cer i tlzi u m .

,

( Pl . xvi fi g
. 6 ) a n d. P o ta m i a es are com mo n
’
throughout
the Te rt iary the t wo fir s t oc curri ng i n m ari ne depo s it s
, ,

t he la s t in br a cki s h l ayer s of the O lig oce n e Ap or r /za is .

i s a wel l -kn ow n Pl i oce n e fo ss il ;while t he s m all R i mella

a nd giga n tic H ipp os/tr ams abou n d i n the E oc e ne .

Cow rie s are es pecial ly Cai nozoic form s ;Tr i vi a oc cur s

profu s ely i n the Plioce n e a n d Plei s toce n e
‘
Large .

s pe cies of Py m la are com mo n i n the Lo n d o n Clay .

R a chiglos s a a n d Toxoglo s s a a re pre - em i ne nt ly

Te rt i a ry group s The form er a re r epres e n ted by
.

”
a bu n d a nt B ucci n i d a e whelk s ) t hroughout the e ra ,

B ucci uu m, Pi sa ui a , N as sa , S tr eps i d u r a , Al a b a m a a nd

N ep tuuea be ing th e m o s t i m po rta n t ge nera The l as t .

na med is pa rt ic ula rl y i nt eres ti ng , in that it s m os t

 CA IN OZ O I C FAU N A S 207

cha rac t e ri s ti c Pl i oce ne s pec ies wa s s i ni s t ral while ,

m ode rn types are n or m ally d ext ral S p i n ou s Mu ricidae .

a re hard ly l es s abu n da n t ; Typ fi i s with s m a l l s hel l s ,

o rn ame n t ed by hollow excre s ce n ces bei ng es pecial ly ,

com mo n i n t he Barto n Clay Pu ra

( T li a i
. s ) i s m
a bu n d a n t i n the Plioce n e a s at pre s e n t The F usid a e '

.
,

ar e one of the m o s t fam iliar Cai n ozoic fa milies F us us .

i s com mo n i n the E oce ne a n d i s there a s s ocia t ed with

,

gero ntic s ide-li ne s s uch a s Sycum a nd Cla vella The .

B a rto n C lay is p a rticularly fam ed for it s a bu nd a nt

V oluti d ae s co res o f exqui s ite form s of Volu ti li tlzes
( Pl xvi
. fig.
7 ) occur
. togethe r with
, t he le ss o r n ate
s pecie s of Lyr ia M a culopeplum is a well -k n ow n
.

Pl ioce ne ge n u s The chief fam il ie s of Toxoglo s sa a re

.

the T urrit i d ae a n d Co nidae Ta r r i s ( Pleur otoma Pl xvi

.
, . .

fig . divi ded i n t o n u m erou s s ub -ge nera i s com m o n ,

throughout the E oce ne ;Couus while enj oyi ng i ts acme ,

a t the pre s e n t day i s well repre s e n t e d by less i nv ol ute

,

form s ( Couor b is ) i n the E oce n e .

C uriou s ly e n rolled types s uch as B ulla r ia a nd ,

S er ap lzs are familiar Te rtia ry m e mber s of t he Tec t i

,

bra n ch E u t hy neura but it is i n the Pul mo nate s ec tio n

,

t hat the s ubcl as s fi nd s i ts fulle s t C a i nozoic expre ss io n .

L i mnaea ( Pl xvi fig 9 ) a n d Pla nor bi s ( Pl x vi fig

. . . . . .

rep res e n ted by s pecie s clo s ely re s embl i n g moder n typ e s ,

a re l a rgely re s p o ns ible for the B embridge Lime s to n e a n d ,

occur abu n da ntly in o t her fre s h -water O ligoce ne s trata ,

e s pec ial ly the H eado n Bed s S nail s are u s ually s ca rce

.

a s fo ss il s bu t H eli x (sens la t ) a nd A m i m ’
, p./z a r o . u s are
not u n co m mo n I n t he Bembridge Bed s The dege nerate .

s hell s of S lugs a re o cca s io nal ly fou nd i n t he s a m e

depo s i t .

Cai nozoic Cephalopod a cal l for l ittle pal aeo n t ol ogical

com me n t N autiloid s are al mo s t res tricted to the fam ily
.

N autilidae one s pecies o f N a u ti lus s till s u rvivi ng Large

,
.
 208 I NVE R TE B R AT E PALA EON TOLOG Y

s pecie s of tha t ge n u s a ccom p a nied by the e nd -form s of

,

the Clyd onautilid a e (A tur i a ) occur i n the Lo ndo n Clay , .

A m m o n oid s are al togethe r wa n ti n g but d ege nerate ,

B e le mno i d s ( e g B elop ter a ) wi t h i ns ig nifica nt g uard s

. .
, ,

l i ngered i nto the E oce ne The rece n t Sp i r ula with a.

,
”
c oil ed, al m os t f Gyrocera co ne s hel l s eem s t o s how
‘
, ,
'

develop me nt of lo ng d eferred e nrol m e nt i n thi s u s ually

-

“
O r t hoce raco n e group Cu t tle fi sh a nd O ctopod s
- .

occu r s pari n gly The m aj ority of the l atte r a re devoid

.

of hard par ts a nd t heir freque ntly giga ntic propo rt io ns

-
,

s ugg e t phylogero ntic over developme nt comparable

s 1 -

wi t h that of the E lepha n ts .

K
( ) A R T H R O P OD A

Little co mme nt is n ee d ed on Ca inoz mc Ar t hrop ods .

The prevale n t order s are tho s e d om i na n t at the pre s e nt

d ay ;a l a rge p rop ort 10n of ge n era have pers i s t ed from
“
the Eo ce n e The com m o
. n aco r n bar nacle ( B a la uus )
-

fi r s t appeared i n the E oce ne ;it is abu nda nt i n t he

Pl ioce n e C ra gs Crab s a nd Lob s t er s are com m o n i n
.

t he Lo ndo n C lay es pecial ly i n the I s le o f S heppey

, .

H op lop a r ia a na u tlzo
p s i s ( Pl xvi fi g 1 1 ) are wel l . . .

k now n r epres e nt ative s of the two s eries More delicate .

A rthropod a s uch as My riop od a a nd I ns ec ta are bes t

, ,

k now n by form s e nclo s ed i n a mber ( u s ually of O ligoce n e

d ate) ; but a n abu n da n t a n d wo n derfully pre s e rv ed
-

- h
i ns ect fau n a a bee d i covered
s n s i n fre sh wa t er m arl s
-

at F l ori s s a n t Colorado
, L epidoptera Diptera a n d
.
,

H yme n opter a of es s e n t ially moder n a s pe ct oft e n re ta i n ,

i n g m uch of their c olor a t i on h a ve bee n fou n d i n t h at ,

remarkable depo s it S pecime ns e nclo se d i n a m ber a re

.

n o rm ally pe rfect a nd u n compres s ed K n owl ed ge of .

s mal l form s s uch a s the D ip t er a i s m ade a cces s ible by

s uc h a m ode of p re s erv a ti on .
 210 I NVE RT E BR AT E PALA E ON T OLO G Y

em pha s ize the d a n ger of formi ng opi n io n s on s uch a

m atter from the evide nce provided withi n a s mall area .

I n B ritai n the arbitrary d ivi s io ns betwee n the Lower

,

and Upper Palaeozoic ( Proterozoic a nd D e ute roz oic) .

Pal aeozoic a n d M e s ozoic a nd Me s ozoic a nd Cai nozoic

, ,

co n form s o clos ely to o b served fac t s i n S tratigraphy

a nd Palaeo ntol ogy that they appear to coi ncide with
n atu ral break s N ot o n ly were the U p pe r S iluria n
.
,

Pe r mo Carbo n iferou s
-
and Late Cretaceou s period s
m arked by s eriou s ea rth moveme n t ( i nvolvi n g importa n t
-

phy s iographical cha nge s) b ut they were followed by ,

epi s ode s which whether i n s edime n ta tio n or den udatio n

.
, ,

have left great lacu n ae i n palaeo n tological eviden ce .

I n co n s eque n ce the fau n a s s eparated by thes e gap s i n

,

the record prove to b e exceedi n gly di ffere n t i n character .

The fo ss il s of the M ag ne s ia n Lime s to n e s eem to have

l i t tle co nn exio n with tho s e of the R haetic a nd Lia s ( the
n ext truly m ari ne s erie s a cce s s ible i n t hi s cou nt ry ) .

Betw ee n the orga n i s m s of the Chal k s e a a n d tho s e o f

th e E oce ne e s tuarie s there i s a co n tra s t that ca nn o t
wholly be a s cribed to d iver s e phy s iographical co n ditio ns .

I t i s d i fficul t t o avoi d the co nceptio n that s ome great

accelera t io n m u s t have i n flue n ced evol utio n d uri n g the
barre n i nterval s Where po s itive a nd n egative evide n ce
.

i s i nvolved a ki n d of me ntal halatio n te nd s to mag n ify

,

t he impre ss io n of the former at the expe ns e of the

.

la t ter .

I nd icatio n of the u ns ou nd ne s s of s uch rea s o ni n g ca n

be fou nd i n purely palaeo n tological featur e s eve n i n
, ,

thi s cou n try The s ca n ty a nd ill -pre s erved fau na of the

.

D ev o n ia n co n t ai n s a ble nd of type s ( particularly amo ng

Coele ntera t e s Brachiopo d s a n d T r i lob ite s ) §which have
,

a s m uch i n com m o n with their S il uria n predece ss or s

a s with their Carbo n iferou s de sce nda nt s O n s trati

.

-
raph ical grou n d s the va s t accu m ul a tio n of Ferm o
g ,
 C R I S ES I N EVOL U T I ON 21 1

Tria ss ic N ew R ed S a nd s to n e a nd the proofs of

“
,

e n ormou s de n udatio n prior to the com me nceme nt of

Te rtiary s e dime ntatio n m u s t be accep t ed a s rep re s e nt
,

i ng lo n g i nt e rval s of time duri n g which orga n ic e volu

,

tio n wil l have m a de great progre ss eve n if proc eed i ng ,

at i ts n ormal ra t e .

Whe n the record s of the s e critical times are s tud ied in

di s ta nt regio ns th e bou n darie s betwee n era s ofte n prove
,

n o le s s ob s cure tha n tho s e betwee n their s everal period s ,

both i n S tratigraphy a nd Palaeo ntology The fam iliar .

term Ferm o -T ria s expres se s t he u ncertai n ty s urrou nd

i ng s eparatio n of the Palaeozoic a n d Me s ozoic while the ,

d i fficulty of de t erm i n atio n of the upper lim its of the

Carbo niferou s a nd diver s ity of opi n io n a s to the val ue
of the R haetic s how that the d ivi s io n betwee n the t wo
era s ca nn ot be s hifted i n eithe r d irectio n with be tter
hope s of precis io n The A m mo n ite s which appear s o
.
,

s udde nly i n ou r Lia s s ic record were the de s ce nda n t s of

,

form s tha t flouri s hed el s ewhere i n Perm ia n a s well a s i n

Tria ss ic tim e s The fir s t Belem n ite certai nly s ee m s to
.

have bee n Tria ss ic but s o wa s the la s t Or tuocer a s

, .

I n ma ny pa rt s of E urope there i s comparable reas o n

fo r hes ita t io n in drawi ng a li ne be t wee n the Cre t aceou s
a nd E oce n e .The fau n a s of the M aes t richt ia n a nd
D a nia n s tage s ( u n repre s e n ted in Brita i n) s how s teadily
“ ”
i ncreas i ng Tertiary qual ities a s they are traced
upward s M o s t Cai nozoi c I nve rtebrate group s ra n ge
.

back to the Cretaceou s if not farther ;while Belem n ite s

, ,

at lea s t li n gered after Me s ozoic time s T e le od ont s a nd

,
.

Cte nob ra nch s had climbed far toward s their acme s before
A mmo n oid s di s appeared .

I n fi n e the world evide nce at pre se nt available s eem s

,
-

t o p oi n t toward s a n egative a ns wer to the two fi rs t .

que s tio n s as ked i n the ope ni n g paragraph of thi s

chap t er I t i s pe rhap s perilou s ly hypothetical but a t
.
,
212 I NVE R T E B R ATE PALA E ON T OL OG Y

lea s t in accord wi t h pre s e n t k nowledge to s ugge s t that

,

t he cours e of evol utio n ca n be like ned to a flowi ng tide .

O n e wave afte r a n other rears i t s cre s t a nd fall s i n rui n

each one ru s he s u p the beach only to s l ip back a nd
, ,

yiel d place to a n other which m ay s weep farther or fall

s ho rt
. A m id the welte r of adva nce a nd retreat — t he
r oar of the o nc om i ng b reaker a nd t h e hi s s o f its
s h at t ered recoil — i t i s hard to see tha t a ny la s tin g
progres s is achieve d But behi n d the seem i ng futility
.

o f each fored oom ed e ffort s tead ily a nd i nvi ncibly the

,

tide is com i n g i n
.
2 14 IN VE R TE B R AT E PALA E ON T OL OG Y
A rachni d s 1 53 1 7 2 , , B a ctr i tes 1 6 9 1 9 3
, ,

A ragoni t e 1 4 20 , , a r
B ct i t i cone 1 0 3 ,

d ura b i li t y 4 0 , B acu li te s 1 04 1 9 3 , ,

A r b a ci a , 2 00 B a k ewelli a , 1 6 7
r
A ch a e n 1 1 9 a , B a la u ophylli a , 1 99
A r ch a eoci d a r i s 1 6 3 , B a la n u s 2 0 8 ,

r
A ch aeocy hi d e 1 2 9 at a , B arba d os e h 29 1 98 art , ,

A w h a eon i s cu s 1 9 4 , ar
B n a cle s , 4 8 1 52 ,

A r cha n od on , 8 4 , 1 6 8 B a r r oi s i a 1 7 8 ,

r
A ci d a e , 1 8 7 B art on cla y, 1 8 2 7 , 3 2 ,

A r cti ca , 1 8 9 , 2 0 5 ar t
B y es 4 6
a r F
,

r a
A e n ce ou s oc s , 1 7 , 2 7 r k B the ,
A 60 .

r
A e n i g, 1 2 4 a a
B t h on i n 3 1 ,

r a
A gi ll ce ou s oc s , 1 7 , 2 7 r k r
B e e ch e 9 8 ,

A r i s tocys ti s , 1 3 4 kt
B ee i e 4 4 ,

r r
A t h op od , 3 2 a B e e t le -cas e s , 8 3
ca i n o oi c, 2 0 8 z B ele mn i tella , 1 9 3
a
co l-mea s u e s , 4 0 r B e le mn i t e s 3 8 ,

d e u t e r oz oi c 1 7 0 , B e le mn oi d e a 1 I 1 , 1 9 3 ,

flu v i a ti le 8 3 , B e le mn ot e u t h i d a e , 1 9 3
h ab ta
i t, 87 B eli n u r u s 1 7 2 ,

me s o oi c 1 9 4
z ,
B eller oph on 1 4 7 , 1 6 8 ,

r t r z
p o e o oi c 1 4 8 , B e lop te r a 2 0 8 ,

e t rr tr a
es i l 8 2 ,
B elti u a I 53,

A s a phu s 1 51 ,
t
B en h i c f un 82 87 a a , ,

a t t k
A sce n d n s oc s 9 2 , B er en i cea , 1 8 4
A s cocera s 1 4 8 , B eyr i chi a 3 2 8 5 1 52 , , ,

A S p i d ob r a n chi a t a B i lli n gs ella 1 4 2 ,

ca m oz oi c 2 0 5 , B i lob i tes 1 4 2 ,

d e u t e r oz oi c 1 6 8 , B i ology 1 9 , ,

r
B i th 1 1 5
,

p rote roz oi c ,
1 46 t
B las oi d e a
4 5, 1 36 1 6 1 , ,

A s pi d ocer a s , 1 9 2 B le n d e 4 6 ,

A s ta r te 1 8 9 2 0 5
, , B olla n d , 1 6 1
A s ter ocem s , 1 9 1 r g
B o in s 2 0 2 3 , ,

tr
A s e oi d e , 4 5 a rr
B o ow d a le se i e s 1 2 4 r ,

A s t h e n od on ta 20 5, 1 9 0 ,
B othr i oci d a r i s , 1 3 8
A s then os oma , 1 8 2 r a
B ou ld e cl y 3 8 ,

A s tr opecten I 8 1 , B ou rgeti a 1 90 ,

A ilzyri s I 6 6
-
, B ou r gu eti cr i n u s 1 8 0 ,

A t e mar 1 07 ta ,
ra
B ch i op od a 1 5 3 1 , 34 , , ,

d e u t e r oz oi c, 1 6 4
p o t r
e o o i c,
1r z
4 1 b r a ch i d i a , 6 4
A tr ypa , 3 1 , 1 4 3 1 6 6 ,
z
ca i n o oi c, 2 0 2
A tu r i a , 2 0 8 d e u t e r oz oi c, 1 6 4
A u la cocer a s 1 9 3 ,
v t
e olu i on 1 I 1 ,

A u la coflzyr i s , 1 8 6 h ab t t
i a 87 ,

lop or i d a e 1 30 g
hi n e 4 3
g on oh a r 3 z
,

z
me so oi c 1 8 5
,

g
, ,

A x on Op o ra I 3 2 , mo p h o en y, 9 7
r g
z
A oi c, 2 , 1 2 0 p r o t e ro z oi c , 1 4 o
r a n e , 1 09
B a ct e i a , r 1 5 r k
B a c es h a m e d s , 3 2 b
 IN DE X 21 5

B ra rd fo d A s , 32 bba Ca r d i omorph a , 1 6 7
r
B a n chi opod a , 1 52 C a r d i ta 3 3 1 8 9 2 0 5
, , ,

r a
B e cci ,27 Ca r d i u m 2 0 5 ,

B r i ss op s i s , 2 0 2 Car yocri n i d a e 1 3 5 ,

B r i s s a s , 2 02 Ca ss i d u loi d a 2 0 1 ,

r
B i t i sh Colu m i , 2 1 , 4 9 ba ,
121 Cas t s 1 7 2 7 4 2
, , ,

r r
B ock a m 1 56 , Ca t a ch r on i s ms , 3 8
B r ongn i ar tella 1 51 ,
g
Ca t a e n e s i s 1 0 1 , 1 1 7 ,

B r ontes , 1 7 1 C ata r s t op h es 3 4 ,

B u cci n u m, 2 06 Ca u s t i c p ot sh , 6 2 a
g
B u d lei h S a l e t on , 3 8 t r t
Cele s i n e , 4 6
k
B u l , 1 04 1 1 7 ,
Ce me nt -s on e , 3 6 11 t
B u lla r i a , 2 0 7 tr
Ce n e ch i n oi d a , 1 0 9
B ulli on s 1 7 0 , Cen tr echi n u s 2 00 ,

B un d en ba
ch 4 5, 1 6 2 , Ce ntr on ella 1 6 6 ,

r
B u i a l, I 5 Ce p h a lop od a
r
B u ni n , 64
g a
c i n o oi c 2 0 7 z ,

B urr ow mg 4 8 , d e u t e r oz oi c 1 6 9 ,

rt
B u on B d s oc , 3 2 ra t k me s o oi c 1 9 1 z ,

r
p ot e o oi c 1 4 7 r z ,

Cad d i e-ca se s 4 9 , Cer a mopor a , 1 4 0

Cai n oz oi c 7 8 1 9 5 Cer a ti ooa r i s , 1 53
k U
, ,

Ca e - rchi n s 8 8 , Cer a ti tes , 1 04 1 70 , 1 9 3 ,

Ca la i s , 1 9 4 Cer a tocepha la 1 5 1 ,

Calcai re gr os s i e r 1 9 8 , Cer i thi u m 1 9 1 , 2 06 ,

Calca re ous algae 30 , Ch a etetes 1 3 1 1 60 1 7 8 , , ,

rock s ,
1 7 Ch a lce d on y, 2 9 4 4 ,

C a lceola 1 59 Ch a lk , 2 9 4 0 8 8 9 1
R
, , , ,

r
Ca lci fe ou s s n d s on e 1 56 a t , Ch a lk ock 36 ,

a g
C lci s p on i a e ;1 2 8 , 1 7 7 Ch a ly i t e 3 7 b ,

a
C lci t e 1 4 1 9 2 9 , 4 4
, , , Ch a ma 2 0 5 ,

r b t
d u a ili y 4 0 ,
r t r
Ch a a c e i st i c f oss i ls 7 8 ,

a ar b at
C lci u m c on e , 36 Ch e i los t oma t a 1 4 0 1 8 4 , ,

h OS p h a t e 2 9 , Chei r a r a s 1 52 ,

Ca e d oni a n u p li ft , 1 2 5 r
Ch e t 2 8 2 9 , ,

Ca lyci n a , 9 4 Child h oo d , 1 I 6
Ca lyme ne, 1 51 Chi t i n 1 4 , ,

Ca lyptr aea , 2 0 6 r b
d u a i li t y, 4 0
Ca ma r ophor i a 1 6 5 , Chla mys 1 8 9 ,

Ca ma r otoechi a , 1 4 3 r
Ch lo i t i c m l 44 ar ,

br
Ca m i a n 2 1 1 2 4 , , Chon ezes 3 3 8 5, 1 4 2 1 6 5 , , ,

a b r g gr
C m id e ee ns a n d 3 8 , 44 , r
Ch on olo y 7 5 g ,

ra
Ca me t a 1 3 7 ,
r a
Ci d a i d e , 1 0 7
Ca nn el, 2 8 Ci d a r i s , 1 8 1
Ca pu lu s , 1 4 7 , 1 6 9 1 9 0 , 2 06 ,
rr
Ci i p e d i , 1 52 , 1 9 4 a
Ca r a d oci a n , 2 7 Ci r r u s 1 9 0 ,

rb
Ca ona ce ou s oc s , 2 8 r k fi t
Clas si ca i on , 6 8
C a r b on i cola , 8 4 1 6 7 , 1 8 8 , Cla thr od i ctyon 1 3 1 ,

C ar b r
on if e ou s , 2 8 3 1 , 1 5 5 , Cla vella 1 0 5 2 0 7 , ,

Ca r d i n i a 1 8 8 ,
Cla y, I 7 2 7 ,

Ca r d i ni i d a e 1 6 7 ,
Cla y-sla t e , 2 1 , 2 6
Ca r d i ocer a s 1 9 2 , Clea v a e 2 1 , 4 7g ,

Ca r d i ola , 1 4 5 Cli ma e , 8 8 t .
2 16 IN VE R T EB R AT E PALAEON T OL OG Y
Cli ona 1 7 , Cr ani acea , 1 4 2
Cli ta mb om tes , 1 4 2 Cr a ss a telli tes , 20 5
Clymen i a 1 6 9 , Cr a s s i n ella 1 8 9 ,

Clypea s ter 2 0 1 , Crayfi sh , 8 5

r
Clyp ea s t oi d , 9 5, 1 1 1 , 2 0 1 a Cret ace ous 30 1 7 5 ,

Clypeu s , 6 9 , 1 8 3 Cri n oi d e a 30 9 8 ,

a ba
Co l- lls 1 70
a a r
Co l-M e s u e s , 2 9 , 36 , 4 0 , 1 56
Cochlocer a s , 1 9 1
C ad a s ter , 1 3 6
Coele nt e rata
a z
c i n o oi c 1 99
d e u t eroz oi c I 59
mes o oi c, 1 7 8
z Cr i s tella r i a , 1 7 7
p ort e o o ir z
c, 1 2 9
Coe locer a s 1 9 2
Coelon a u ti lu s , 1 6 9
Coelopleu r u s 2 00
C oen othyr i s 1 8 6
Collyr i ti d ae 1 8 3
Colob oce n tr otu s ;1 0 1
Colou , 4 0r
Comme n y 1 7 2 tr
Con chi d i u m, 3 1 , 7 9 , 1 4 2
rt
Co n c e i on s 3 6
, ca i n oz om, 2 00
d e u t e r oz oi c, 1 6 1
h a b i tat , 8 7
mes oz oi c , 1 79
p r ote r o z oi c , 1 37
, Cr i ocems , 1 9 3
, r
C i s es, 1 1 5, 2 09
r
C oci d oli t e , 4 6
, Cr ota locepha la , 1 7 1
Crota locr i n u s , 1 3 8
, r a a
C u s t ce , 1 4 , 1 6 , 8 3 , 1 4 9
, Cryptocr i n us , 1 3 5
, r
C yp t os t om , 1 40 ata
Ct e nob ra n chi a t a, 1 4 6 , 1 6 8
Ct e nos t om t , 1 4 0 aa
, Cten os tr e on , 1 8 9
Cu ou lla ea , 1 8 7 , 2 04
, Cu lm-me s u e s , 2 9 1 56
a r ,

g
Con lome rat e , 2 0, 2 7 Cu pr ess ocr i n u s 1 6 2 ,

r
Con ife s 8 1 , 1 02
, t
Cu t le -fi sh , 2 0 8
C on oca r d i u m 1 6 7 , Cya th ocr i n u s , 1 3 8 , 1 6 2
Con or b i s 2 0 7 , Cya thophyllu m 1 30 1 59 , ,

r a v
Con se v ti e s , 1 0 7 Cyclod on t a 1 90 , 2 0 5 ,

C on u la r i a 1 4 6 , Cyclos t om t , 1 4 0 aa
C on u lus 7 1 7 9 9 5, 1 1 0 1 8 2
’

, , , , Cyclothyr z s , 1 8 5
C on u s 2 0 7
, Cyclu s , 1 7 2
r
Cop oli t e , 4 3 r
Cyli n d i t e 5 1 9 1 ,

r
Co al-li me s t one s , 30 Cyp ri d a e , 8 4
re ef s , 30 3 9 , Cyp ri d e a , 2 7 , 3 2
ra a
Co lli n 30 3 9 , , Cypr i n a , 1 8 9
ra
Co lli ne c g, 3 1 ra Cyr en a , 2 7 , 1 8 9
ooli t e 2 8 , Cyr eni d a e , 8 4
a
Cor lli n e s , 3 1
’

Cyr tz a , 1 4 3
r
Co als 8 1 6 6 3 , 8 8
, , , m ‘

Gy n a , 1 4 3
Cyr tocems , 1 6 9
morp h og e ny . 97 t a
Cys i d e , 1 3 3 , 1 60
C or b i cu la , 2 0 5
Cor b i s , 2 0 5 D actyli ocer as , 8 0 , 1 9 2
Cor b u la , 1 9 0 , 2 0 5 D a lmcmella , 1 4 2 , 1 6 4
r
Co nst on e , 3 6 D a lma m tes , 1 52
’

C or on i cer a s , 1 9 1 D a lma m li n a , 1 52
'

C os gi n op or a , 1 7 7 a
D mocle s , 56
C os mocer a s 1 9 2 , r
D a wi n , 9
Cothu m ocys ti s 1 3 4 D avi d s om a , 1 6 5
‘

t
Co t on -w ool, 6 6 D a ws on ocem s , 1 4 8
Cr a n i a 1 8 5
, De at h , 1 1 5
21 8 IN VE RT E B RA T E PALA E ON T OL OGY
E ut h yne u , 1 4 6 ra G ast rop od a morph ogeny , , 97
v t
E olu i on , 5, 7 9 , 9 1 , 9 2 , , p r ot e r o z oi c 1 4 6 ,

G a u lt 1 8 , 2 7 4 5
, ,

E xogyr a , 1 8 8 Ge n ea lo i ca l ee s , 1 09g tr
ra
E x t n e ous fos si ls , 10 Geo d e s 1 0 ,

G e ology, 2
Fari gd 3 3
0

n on , 0, 1 , 38 , 4 2 , Gephyr ocem s , 1 7 0
84 1 r
G e on t i c s t e, 1 1 1 ag
Fa al li t 5
un s s, 1 Gew i lli a , 1 8 7 , 2 0 4
Gi lb er ts ocmn u s , 1 6 2
’

F
’

t
a vos z es , 1
3 0
F ath r tar 8
e e -
s s, 1 0 r a
G i v n , 1 34 , 1 3 8
F t ll Gi s s ocmn u s , 1 3 8
’

ene s e 3 63 1, 1
F rr
a,
e ar b t
ou s c on a e , 36 a
Gl ss N , 4 2 6 4
F rt ili zati
.
, ,

5 on , 1 1 a
Gl ucon i t e , 4 3 , 1 2 7
Fi r lli d
e
ss u e 68 a e, 1 Glob i ger i n a 1 9 8 ,

F i s t u li p or i d a e 1 4 0 , 1 6 3 , Glos s ogm ptu s , 1 3 2

F la b ellu m 1 9 9
'

, Glos s othyr z s , 1 8 6
F le x i b i li a , 1 3 8 , 1 6 2 Glou ce st e sh i e 3 6 r r
F
,

Glyci mer z s , 2 04
‘

li nt 30 , 3 6
F
,

li nt -me al , 3 2 , 3 7 Gly fi h a ea , 1 9 4
mou ld s 4 4 Glyphz ocems , 1 7 0
’

F
,

r a t
lo i ss n , 2 0 8 Glyptosph a em , 1 3 5
F lu s tr a 2 02 Gn e i ss , 1 1 9
F
,

a
lu v i t ile f un s 8 3 a a Gold i a s 1 7 1
F
-
, ,

o mi ni fe , 2 8 , 4 3
ra ra Gom b ocer a s , 1 4 8
a
c i n o oi c, 1 9 8 z tt
G oni a i e s , 1 04 , 1 0 7 1 6 9 ,

Gom omya , 1 8 9
’

d eu t e r oz oi c, 1 58
me s o oi c 1 7 6
z ,
Gon i ophyllu m, 1 3 0
p r tr z
e i c 1 27 r b
G a au 9 7
F
o o o , ,

r tb
o es - e d s 8 3 Gmpk i ocr i n u s , 1 6 2
F
,

r
o est M a rb le 3 2 r t
G a ph i e , 1 1
F
,

oss i l-collec i n , 5, 2 5, 50 t g r t t
G a p oli e s , 4 1, 1 26 ,
F os sili a t i on z 31 1
F
19 ,

oss i ls 1 0 , 1 1 , 3 9
, Gr es s lya , 1 8 9
ba nd s 32 ,
Gr e a k
c s , 2 8, 1 20
rv
d e i ed , 3 7
’

Gr z thi d as , 1 7 1
d i sse ct i on , 6 2 Grypha ea , 3 8 , 1 8 8
occu rre n ce 2 5 , G ymn o los s 1 47 1 6 8, a , ,
1 90 ,
r ock f ormers 2 9
-
G ymn o a e ma t a 1 39
F
, ,

os s i l-w ood Gypz d u la , 1 4 2

‘

, 81
F r on d z cu la ma , Gy m
' ’

1 77 d es , 1 9 1
F u si d a e , 9 7
F u s u li n a , 1 58 H ab i t at 1 5 , , 34
F usu s, 2 0 7 H ab i t s 1 1 6 ,

H a ld on H ill 1 7 5 ,

a
G le n , 4 6 a H a lys i tes 1 3 1 ,

Ga ler i tas , 7 1 H a mi tes , 1 0 3 , 1 9 2

Ga leropygu s , 1 8 3 a
H mme s , 54 r
Ga str i ocer d s , 1 7 0 H a mps hi e , 1 9 6 r
tr
Ga s opoda 1 3 , 3 2 , 4 0 , 8 2
-
H a pa locr i n u s , 1 3 7
ca i n oz mc 205 H a r pe s , 1 7 1
rt U r
,

d e u t e r oz om ,
168 He a - ch i n s , 8 7 , 1 8 3, 2 01

H eb er tella , 1 4 2
 IN DE X 2 19

H eli oli tes , 1 3 1 , 1 59 I n d ocems , 1 04 , 1 9 3

H eli x, 2 0 7
H emati t e 4 6
H e mer a e 7 3 7 5
H emi a s ter , 1 8 3
I nf a n c y, 1 1 6
, r r
I nf e i o O oli t e , 30 , 3 2 , 3 5, 3 8
, , I n ooem mu s , 1 0 5, 1 8 7
I n se ct a 1 6 , 8 2 , 1 7 2 , 1 9 4
,

H emi oi d a ms , 7 8 , 1 8 2
‘

I s a s tr uga , 1 7 8
H emi cys tz s 1 3 6
’

I s ch ad z tes 1 2 9
’

Wg
, ,

H em/iped i n a , 1 8 2 '
I sle of i h , 2 7, 33 t
H er cyn ella 1 6 9
’

, I s ocr z n u s 1 7 9 , 2 00 ,

H e ref or d shi re , 2 7 36 , I s od on t a , 1 4 5, 1 6 8 , 1 8 8 , 2 04
H eter ocentr otu s , 2 0 1 I s op od a , 1 7 2
H exacr i n u s , 1 6 1 t t
I s os a i c mo e me n , 2 2 v t
a t
H e x c i n elli d , 1 2 8 a
br
H i e ni a n G ee n s a n d , r 1 75 Ja ck s on 9 6 ,

H i ld ocem s , 1 9 2 J y
e ll -fi sh
4 8 49 , )

H i ppochr en es , 2 06 J erea 1 7 7 ,

H i ppopod z u m, 1 8 9 Ju rassi c 8 2 8 1 7 4
’

, , ,

H i ppw z tes , 1 0 5, 1 90
’

H old s ter , 1 8 3
'
K pr 7
eu e ,
2
H olcos pongz a , 1 7 8 Ki g 86
Ki rk by M r Flag
n an a , 1
H ole ct yp oi d a , 9 5, 1 1 1 , 1 82 oo s, 1 4 7
H olectypu s , 1 1 0 , 1 8 2 K i t ch e n -mi d d e ns , 1 1
K omn ck i n a 1 86
’

H olopella , 1 4 7 ,

t r
H olo h u i oi d ea , 1 4 , 4 9 , 1 38 , 1 62, K u torgz n a 1 4 1
'

Lab lli g 5
1 80
H oma lon otu s , 1 52 , 1 7 1
La tri fa a
e n ,
6
H oplz tes , 1 9 2 1 9 3
’

, cu s ne un s , 83
h m
Lg
’

H op lopa r i a 2 0 8
’

,
L a evz d en ta u , 1 9 0
H ord e rle y S a n d s tone , 3 3
La l a 5
a oon s , 8 9
H ori on 52
z ,
k e Ca e d on i 1 6 ,

Ho mem ,
2 02 d i st i ct , 1 2 4 r
M on mou t h , 8 6
Lak
’

H ya los telz a , 1 2 9 , 1 58
at
L am r k
Hy t 9 6 , es , 8 4
H yb oclypus , 1 8 3 a c , 9
H yb ocr z n u s , 1 3 7
'

L a nfer i a , I 1 0
H yd r a ctz m a , 1 7 9 , 1 9 9 L a pwor thu m , 1 3 8
Lar a tag
’ ’

r r
La r a
H yd ochlo i c aci d , 6 1 v ls e s , 1 4 , 9 5, 1 1 5

r z a
H yd o o , 4 7 , 1 2 9 u e nt i n, 1 1 9
H ymen occm s , 1 53
L ar
’

L ed a , 1 4 5 1 6 7 , 1 8 7 ,

H yoh thes , 1 2 1 , 1 4 6 e in tw
'

d ine , 1 3 8
H yp oge n e fo ce s , 2 1 r L ep a d ocr z n u s , 1 3 5
’

H yp ol ar a
i , 1 50 , 1 7 1 L eper d z tz a , 1 52
’ ’

z
H ypot yr i d z n a , 1 6 5
'

L epi d es thes , 1 6 3
L epz d oce n tm s , 1 6 3
’

fi t
I d e n ti ca i on , 6 9 L epi d od z s cu s , 1 6 1
'

I d mon ea 1 8 4
L
,
L epta en a , 3 1 , 1 4 2 , 1 6 5
r k
I gn e ou s oc s , 2 1 , 2 5 i a s s i c 2 7 , 36 , 4 5, 1 7 4
r g t
I mp e n a i on , 1 2
L
,
2 0, 41 L i ch a d i d a e , 1 5 1 , 1 7 1
r
I mp e ssi on s , 4 8 r
Lg
i fe -cu v e , 1 1 3
I n a d un a t a , 1 3 7 i a me n s 1 8 , 39 t ,

r a
I nc ust t i on 4 1 ,
L i ma , 1 6 8
a
I n d i n i nk , 6 5 L i ma tula , 1 8 9
I nd i vi d uals , L i me -k i lns , 24
 2 20 I N VE R TE B R AT E PALAE ON TOL OG Y
Li mest one M a ga s , 1 8 6
Li mi d a , 2 0, 2 8
e, 1 89 M agma , 2 1
Li m n e si a n Li mest one
Li m i d a fig
n aea , 20 7 89
ghi t
,

n ae e, 84 1 c e 46
Malacos traca , 1 53 ,
L i mo si d ae , 1 87 M alt on , 6 9
L i ma us , 1 7 2 M alve rn axi s , 1 2 5
L i n gu la , 3 3 , 8 6 , 1 07 , 121 ,
141 M ammali a, 1 04 1 1 2 ,

M a mmot h , 8 4 8 8
Li g la flag
n u s, 47 M an x S lat e s 2 8 ,
,

M ar ble 1
Li a 7 4
L i g l ll
n u e 7 a, 4 ,
14 1 ,

nn e us, M arcasi t e 1 9 37 4 5 6 6 , , , ,

M ari ne f a u nas 8 6
L
L i ocer a s , 1 9 2 ,

f l
i s t s of os si s , 8 M a r marosi s 2 8 ,

M a r s i pocr i n u s 1 3 7
Li thi ti d a
L i t a r aea , 1 9 9
h ,

s ,
128 M a r s u pz tes , 1 8 0
M atri x 1 7
o

Li thopha gu s , 1 6 8 ,

M at u ri t y 1 1 5
Lt
L i thos tr oti on , 1 59 ,

ra b t
i t o l el , 1 6 M ed h cottz a 1 7 0
a a
f un , 8 6 M ega la n ten s 1 66
M ega thyn s 1 8 6
L
’

L i ttor i n a , 1 9 0 , 2 06 ,

r
i t t o i ni d e 1 4 7 a ,
M ela n i a 2 06 ,

M elocr z nu s , 1 6 1
’

L i tu i tes , 1 4 8

La
L i t u oli d a e , 1 7 6 M elon echi n us 1 6 3 ,

v r b
l nd o e y e d s , 3 1 M elongen a 2 0 6
M e mb r a mpora 1 8 4 2 02
L a
L ob ltes , 1 9 1 ,

z a M e n d i p a x i s 1 55
L
oc li ed st ge s 9 6 , ,

on d on ba
si n 1 9 6 , M er etr z x 2 0 5
a
cl y. 3 2 . 3 6 M e r i s t e lli d ae 1 6 6 ,

L on gm d i n, 2 8 , 4 9 1 2 0a , M er i s tz n a 1 4 3
'

L
L on s d a ei a , 1 59 M es ocys tz s , 1 3 5
'

ow e r r
G ee n s n d , 3 8 a M e s on a ci d ae 1 50 ,

O oli te s , 30 M e s oz oi c 7 8 1 7 3 , ,

L oxone ma 1 4 7 , 1 6 8 ,
M es pi locr i n u s 1 6 2 ,

M e ta morp hi s m 2 6
L
L u ci n a , 1 9 0 2 0 5 , ,

u d low B on e b e d 3 3 3 5 8 6 , , , M e ta morp h osi s 1 1 5 ,

t
li me s one , 30, 3 1 M e t as omati s m 2 1 ,

se ri e s , 19 M e ta z oa 1 1 5 ,

L u d wi gla , 1 9 2 M etopa s ter , 1 8 1

M i cheli ma , 1 60
'

L yco od s , 1 0 2
L ye 74 M i cr as ter , 7 9 1 8 4 ,

L yr i a , 2 0 7 M i cr os cop i c f os si ls 54 6 7 , ,

L ys s a ci n a , 1 2 9 M i cr othyr i s , 1 8 6
L ytocer a s , 1 9 2 M i croz oa 55 ,

Mi d -z on e 9 9 ,

M acoma 2 0 5 , M ili oli d a e 1 9 8 ,

M a cr oceph ali tas M i ller z cn n u s 3 1 1 8 0

’ ’

,
1 92 , ,

M acr och ei lus 47 168 ,

1 , M i nerals 1 0 ,

M acr ocys tella , 1 3 5 M ine s 2 0 2 4

, ,

Ma r
c oscop i c f os sils M i ocz d a r i s 1 6 3
’

53 , ,

M a ct; a, 20 5 M od e 7 6
,

M a cu lopeplu m, 2 0 7 M od e ls 6 4 ,

Ma r i a , 30rar M od i olops z s , 8 5, 1 4 5
'

d e po
222 I N VE R T E B R AT E PALA E ON T OL OGY
Or th ocer a s , 8 5,
Or thod on ti s cu s , 1 4 5
r
O t h oge ne si s , 1 1 0
O r tk on ota , 1 4 6
Or thotetes , 1 6 5
ra a
Os t cod , 3 2 , 8 3
mes oz oi c
, 1 94
p r ot e r o z oi c ,
Os tr ea 4 0 ,
, P em a 1 8 7 , 2 04
r
Ost e i d e a , 1
r a
Oxf o d i n 4 5 , P er on ocer a s , 8 0 , 1 9 2
Oxyn oti cer a s , 1 9 1
P a chyd i s cus , 1 0 5, 1 9 3
P a chypor a , 1 59
a k g
P c in , 57
P a la eechi nu s , 1 3 9 , 1 6 3
aa
P l e ocon ch , 1 4 5, 1 6 7 , a
P a la eocor ys tes , 1 9 4
P a la eocyclus , 1 30
a a eod i s cu s , 1 59
af
g a e on t o gr 8 ? y 4 6 Ph i lli psi a , 1 7 1
P a la d i na , 1 9 3 Ph ola d e lli d a e , 1 6 8
P a n op e, 2 0 5
P a phi a , 20 5
P a r ab oli n ella , 1 51
P a r a d oxi d es , 1 5 1
P a r a llelod on , 1 6 7
P a r a s a len i a , 2 0 1
P a r a s mi li a , 1 7 8
P a r k er i a , 1 7 9
P a r k i n son i a , 1 0 5, 1 9 2
ar
P t h e n oge ne s i s , 1 1 5
P a tella , 1 4 7 , 2 0 5
a
Pe t , 2 8 , 3 9
P ecten , 3 4 , 1 6 8 , 2 0 4
Pe ct i ni r h omb s 1 3 5
P ed a li on , 1 8 7
a z
Pe l gi c one , 8 6
Pe la n echi n u s , 1 8 2
Pe le s yp od a 3 2 . 34 , 3 9 , 4 3 . 6 4
a z
c i no oi c, 2 0 3
d e u t er oz oi c, 1 6 7
mes o oi c, 1 8 7
z P latyoer a s , 1 4 7
mo p h o en y . 9 7
r g
r ot e r oz oi c, 1 44
Pe ma t oz oa , 1 33
P elta s tes , 1 8 2
Pe nni n e p U
lif ,
1 56
P en tacr i nu s ,
P en ta mem s , 3 1 ,
Pen tr emi tes , 1 6 1
r rata
Pe fo , 1 59
P er i eck ocr i n u s , 1 37
r
Pe i od s , 7 7
P er i s ck od omu s , 1
1 62
A
Per i sphi n ctes , 1 9 2
1 52 Per mi an 3 1 , 1 56
,
,
67 Pe rn i d ae , 1 0 5, 1 6 7
P eta locr i n u s , 1 3 8
P etr a i a , 1 30
tr t
Pe i fa c i on , 1 2 , 1 7 , 44
-
r r
Pe t i fyi ng S p i ngs , 1 2
a a
Ph cop i d e , 1 52 , 1 7 1
P h a copi d ella , 1 52
1 87 P ha en os chi s ma , 1 6 1
Ph ar e t r one s , 1 7 8
Phi lli s , 6 9
P hi lli ps a s tr a ea , 1 59
P h ola d omya , 1 8 9 , 204
P hola s , 8 7 , 1 6 8
P hor mos oma , 1 8 2
Ph yla ct ol e m a , ata
1 39
Ph ylli te , 2 1
ar a
Ph ylloc i d , 1 53
P hyllooer a s , 1 9 2
Ph yloge n y, 1 5. 90 . I I 4
P hymos oma , 1 8 2
ra
Ph ysi og p h y 8 0 ,
Pi leu s , 1 1 0
P i n a cocer a s , 1 9 1
P i nn a 2 04
Pi s a m a , 2 0 6
, Pi s ocr i n u s , 1 37
g
Pi t t i n , 1 8
Pla cocys ti s , 1 3 4
P la cops i li n a , 1 2 7
P la gi os toma , 1 8 9
a k
Pl n t on , 8 7
P la n orb i s , 2 0 7
P la tycr i n u s , 1 6 1
P la tyschi s ma , 1 4 7 , 1 6 8
’
P lecta mb on i tes , 1 4 2
P lector thi s , 1 4 2
t P les i eohi n u s , 9 4 , 9 8 , 1 8 2
P leu r od i ctyu m 1 60 ,
P leu r omya , 1 8 9
Pleur ophoms , 1 6 8
 I NDEX 223

Pleur otoma 2 0 7 , r t z a r
P o o o , p ot e o oi c, 1 2 7 r z
P leur otomar i a , 1 0 1 , 1 4 7 , 1 90 , P o em
r tr ata , 1 4 2 , 1 64
2 06 P s a mmob i a , 2 0 5
P li oa tu la 1 8 9
, P s eu d od i a d ema , 1 8 2
r
Pli e s , 56 P s eu d oglos s othyr i s 7 2 ,

Pli oce n e 3 1 , 3 2 , 3 8
, Ps eu d omon oti s , 1 6 7 , 1 8 8
Plut oni c oc s , 2 5r k P s i locer a s , 1 9 1
P olli ci pes , 1 9 4 g
Pt e n o loss a , 1 4 7 , 1 90 , 2 0 6
z a
Poly o , 1 , 4 0 , 4 7 , 1 2 2
3 P ter i a 1 6 7 , 1 8 8
,

ca moz 01c , 202 P ter i chthys , 1 53

d e u t er oz oi c, 1 6 3 Pter i n ea 8 5, 1 4 5 ,

hab a it t 87 , Pter i n eop ecten , 1 4 5, 1 6 8

z
me so oi c, 1 8 4 r a
Pt e od ct yle s 8 3 ,

pr r z
o t e o oi c,
1 39 r a
Pt e op od , 1 2 1 , 1 4 6 , 1 6 9
P opa n ocer a s , 1 70 P terygotu s 1 53 , 1 7 2 ,

P or a mb on i tes , 1 4 2 Pti lod i ctya 1 4 0 ,

r ra
Po i fe Pu n a x 1 6 5
f
,

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2 24 IN VE R TE B R AT E PALAE ON T OL OG Y
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2 26 IN VE R TE B R AT E PALAE ON TOL OGY
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,
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1 59
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. .

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143
 TH E
S N A KES OF E U RO PE

G . A . B O U L E N GE R
LL D . .
, B se. .
, Ph D. .
,

I/Vi tb 14 Pla tes a nd 4 2 F igu r es i n tiw Tex t .

S econ d Ed i ti on . Cr own 8 00 .
73 . 6d . net

H I S work co n a ins t d escr ip t ions of l

al

E u rop ea n sp e cie s o f s na k es , t he cha ra c e rs t

of which a re illu st ra t e d by nu me rous t e x t

fi gu re s a nd l t wit h t of
‘

1 4 an accou n
p a e s ,

t he i r h a b i t s a nd d is t rib u t ion . In t he in t r o

d u ct ion t he au t hor d ea ls wi t h t he mo rp hology ,

e thology, class i fi ca t ion , a nd d is t r ib u t ion of

s na k e s n e ra lly
ge .

METHUEN CO LTD , 36 ESS EX STREET,

. . WC 2 . .
 TH E

FR ES H WA T E R FI S H ES

OF TH E B RI T I S H I S L ES

C TA T E R E GA N , M A
. .

27 Figur es a nd 3 7 Pla tes by Me A utfi or .

Cr own 8 210 .
7 S . 6d . net

PO PU LA R accou n t of t he fi s he s of

our la k e s a nd ri v e rs . T he b oo k is u
p

to d a te a nd t he r es ul s t of t he mos t re ce n t

re se a rche s a re inclu d e d .

METHUEN CO; LTD 36 ESS EX STREET, WC 2 . .

 TH E O X
A N D IT S KI N D R E D

R I CH A R D LY D E K K E R
B A
. .
, F G S
. . .

55 I llustr a ti ons . Cr own 8 00 .

7 3 . 6d . net

H I S work d e a ls wi t h t he ge ne ra l s t ru ct u r e

o f t h e s p ecie s , t he a n cie n t E u rop e a n wild

ox a nd it s t
ex er min a t ion ,
B ri t is h whit e pa rk

ca ttle d ome s t ica t e d b re e d s Cont i ne n ta l b re e d s

, , ,

a nd so on t o t he hu mped ca tt le of A s ia a nd

A fri ca .

METHUEN CO LTD
. 36 ESSEX STREET. WC 2
. .
R E PT I L ES , A M PH I B IA ,
FI S H ES , A N D
L O WER C H O R D A T A

J T CU N N I N G H A M, M A
. . . .
, F ZS . . .

R . LY D E KK E R , B A . .
, F GS . . .

G A B O U L E N GE R , D S c , F R S
. . . . . . .

J A R T H U R T H O M S ON, M A
. . .

E dited b y J . T . C U NN I N GH A M

Pla tes i n Colour , 3 3 i n M onotone, a M ap ,

a nd 3 2 Tex t Figur es D emy 8 00
. 1 28 6d net . . .

H IS volu me t ra ce s in b roa d ou t lin e s t he

e v olu t ion a ry h is t o r y of t he R e p t ile s,

A mphi b ia , F is he s , a nd t he le s s fa milia r, b ut as

t hes e p a ge s will s ho w ,
no le s s i nt e re s t ing typ e s

co mmonly d escrib ed u n d er t he t e rm “
lowe r

v e r e b ra e s .
t t

METHUEN ! CO LTD 36 ES S EX STREET,

‘

.
‘

WC 2 . .
 TH E L I F E

OF TH E M O LL U S C A

B . B . W OO DW A R D ,
E L S
. . .

lVi tlz 3 2 I llustr a ti ons a nd

/
a M ap .

Cr own 8 00 7 8 6d
. . . net

H I S b ook gi v e s a o
p p ula r b ut a ccu ra e t

su mma ry of wha t is k nown conce rn in

g t h e

wa ys , ha b i t s , a nd mod e of life of this b ra nch of

t he a ni ma l k ingd om , of which t he s na il, t he

oys e r, a n d t he cut tle fi s h a r e

t fa milia r e xa mple s .

METHUEN 5. co LTD . 3 6 ES SEX STREET, wc 2

. .
G L O SS AR Y A ND N O T ES
ON V E R T E B RA T E
PA L A EO NT O L O G Y

S . A . PE L LY , M A . .

Ft ap 800 . .
53 . net

HE ai m of t his work is t o a s s is t p e rs on s

who wis h to ob a i nt a k nowle d ge of t h e

ma ny i n t e re s t i ng fos s il cre a ure s t see n in

Na t ional M us eu ms . T he mea ning of the na me ,

a nd the chie f p o i n s t i n th e t
s r u c ur e t of t h e s e

a ni mals is see n at a
gla n ce , whe the r it b e
. a

ma mma l , or b ird , or fi s h, o r re tile

p j a nd for th e

sa ke of e a s y a nd q u i ck r e fe r e n ce t he crea u r e s t

a re
p la ced i n al
p ha b e t i ca l o rd e r .

METHUEN CO LTD. 3 6 ESS EX S TREET, WC 2 . .

 A S EL E CT I O N OF

MES S RS . MET H U EN ’

S
PU B L I CA T I O N S

M ETH E N N
IN t his i s accor d in g t o a ut h or s t h e or d e r
M
.

us ar e p u b lis he d of a ll ess r s U §S ovels is s u e d .

a t a p r i ce a b ov e 4s ne t , a nd s i mila r e d i t i ons are p u b lis h e d of s ome w or k s of

L
.

Ge nera l it era t ure Colon i al E d i t ion s ar e only for c i rcu la t i on i n t h e B r i t ish

.

Colon ies a nd I nd ia

B
.

All b ooks mar k ed net a fe n ot s u bj ect t o d iscoun t , a nd ca n not b e b ough t

a t le ss t ha n t h e p u b lis h ed p r ice ook s not mar k ed ne t ar e s ubj ec t t o t he . .

d iscou n t wh ich t he b ook se lle r a llows .

T h e p r i ces i n t h i s Ca ta logu e ar e lia b le to a lt e r a ti on wi th ou t p re vi ous n ot ice

M M ETH E N
U S b ook s a re k e p t in s t ock b y a ll good b ook se ller s I f
.

’
e s s rs
M M
. .

fi cu lty i n se e ing cop ies ,

t h er e i s a n y d i f ess rs e t h u e n wi ll b e v e ry gla d t o .

ha v e ea r ly i n for ma t i on , a n d sp e ci me n copi e s of a ny b ook s will b e se nt on

r e ce i p t of t h e p u b li sh ed p ri ce p lu s p os ta ge for ne t b ook s , a nd Of t h e p ub li sh ed

p r ice for ord i n a ry b ook s .

T h i s Ca ta logu e con t a i n s only a s e lec t i on of t h e mor e i m

p ub li sh e d b y essrs et hu e n M . M
A c omp le t e ca t a logu e of t he i r p u b lica t i o ns .

ma y b e ob ta i ned on a p p lica t ion .

An d re we s ( a ncelot) L PR E CE S PR I A R U
D A G H T O F TH E B LU E

R IG HT M A N C
. .

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F
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.
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.

ne t .
.

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sr .

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HN
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5s .
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T E CT U E . I llus tr a t e d . F ou r t h E a i t i on
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. .

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R
.

R U S S I A N P E O PL E
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I N T H E G E AT GO D S H A I R
’
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. . . . E d i t i on . D emy 80 0 . ne t .
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A YEAR I N R U SS I A Cr 800 73 6d Bennett Ar nold ) . TH E T TH A BO RU UT
R
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L L N R
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N
.

tr a t e d . S e0e n th E d i t an R oy a l i ss . .
B I B I CA I T O D U CT I O Wi t h a .

co ncise B ib li ogra p hy

R N WA LL I ll
S ix t h E d i t ion C7 . . .

800 8s 6 d n et A ls o i n T w o V olu mes

A K
B OO O F CO . us t ra t ed .

Cr 800
.

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. . . .

A B OO K O F D A R T MO O R B
. .

T hi r d E d i ti on
8 Cr 6d
. .

7 . . 00. s. . ne t .

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Thi d E d i t i I llus t ra t ed S econ d E d i t i on Cr

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1 2 s 6d .n et . .

d Thi r d
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M OT O R I N G . I llu s t ra t ed . D e iny 8 00 .

B
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S ongs wi th t he ir T r a d i
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. . . . .

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.

t h e mus ica l e d it ors h i p of CE cu J S P . . HA R . AN E SS V S T I CA L T OL O HE .

S econd E d z t z on L a rge I mt m a l 8 0 0 . S event h E d i t i on . Cr 800 5 s n et

. . . .

7 s 6d n e t
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R E E K P O L I T I CA L
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.

O . F ou r t h E d i t i on . Cr . 8 00 .

5s ne t
R
N N M
. .

Bu ta b le ( c. E ) . T H E CO M E CE O F
ATI O S E igh t h E d i ti on Cr 8 00 5s
TH E E N GL I S H CH U R CH A ND RE
. . . .

UN IO N C 8 r. 00 5s ne t

B
. . . .

net .

ra b a n t ( F R AMB L E S I N S U S S EX
H P AG E T
T H O G H TS U .

ON
.

Backrord I llus tr a t e d Cr 800 6d ne t .

H U T I G Ed it e d b y J OT O
N N 7s

B
. . . . . .

. . .

I llu s t ra t e d Th i r d E d i t i on.D emy 80 0 .

ra i d ( J a mes) A D V A CE D
. GO F N L .

7 s 6d ne t I llus tr a t e d E ight h E d i t i on D emy 8 00

B
. . .
. . .

PA R I S 1 2s 6d net

B
. . .

elloc ( IL ) . . I llu s tra t ed .

E d i ti on . Cr . 80 0 .
7s . 6d ne t . .
ulle y (H A CI E T A N D N N M ED I
LL L ART
.

HI S A N D TH E S EA . N i nt h E d i t i on .
EVA . Illus t r a t ed Cr 800 . . .
7s . 6d .

N
F eat 80 0 és ne t . . . .
n et .

ON OTH I G A N D I N K N D RED SUB Ca r lyle (Th omas ) TH E E FR NCH

R L R L
E T C H ER
.

J E CTS F ou r th E d i ti on
ne t .
. .
EVO U TI O N . E d it e d by C . . .

FL Th r ee Volu mes Cr 80 0 1 8s
RYT H I N G F
. . . . .

O N EV E . ou r t h E d i ti on F ed t . . ne t .

8 6
00. t s . ne .

TH E L
E TT E S A N D S E E CH E S O F R P
ON S OM E T H I N G Th i rd E d i t i on F e at O I VE L C O WE R R M
Wi th a n I n LL
IR TH
. . .

H F
.

8 é
00. t s . ne .

0
t r od u ct i on b y C , a nd o t es . . N
F I RS T A N D LA S T . S econ d E d i t ion .
a nd A p p e n d i ces b y S
Volu mes B er ry 80 49
C LO M A s
1 83 n e t
T h r ee
. .
. .

. .
.

T H I S A N D T H A T A N D T H E OT H E R Ch a mb e r s ( Mr s a mb er t) L LAWN
L
. .

NN
.

S econd E d i ti on F eat 80 0 6 s ne t . . . . . TE I S FO R AD I ES . I llu s t r a t e d .

MA RI E A NTOI NETTE . I llus t ra t e d .

S econ d E d i t i on Cr 8 00 5s . . . . ne t .

F th E d i t i
ou r D my 8 on .
l e 0 0. l 8s n e t . .

Ch e s t e r t on ( G CH A ES DI C E S RL K N
T H E PYR E N E E S P P
. .
’

I ll . us tr a t e d . S econd W i t h t wo or t ra i t s i n h ot ogr a v ur e E i gh th .

E d i t i on . D e my 800 . ms 6d . . ne t . E d i t i on Cr 800 7 s 6 d n et . . . . . .
4 M ET H U EN A ND C O M PA N Y L I M I TED
T H E B A LLA D O F T H E WH I TE H O R S E
F if t h E d i t i 6 t on . s . ne .
. TH E
Cr 80 0
OO D -
as ne t
.
OD F
. .
PR
.
U CI NG GARD E N .

A LL T H I N G S CO N S I D E R E D T th
H I LD L
en
B ea r me r i n IT
.

ale S E OF
’
A C

T R E M E N D OUS T R I F LE S
E d i ti F t 8 6 t
on . ea 00. s ne
R
.

CH I S
. . .

I llu s t r a t ed F ou r th Ed z t i on
F if t h E d i
. .

L a me Cr
‘
.
. 800 és ne t .

A L A R M S A N D D I S CU R S I O N S
. .

KV
ti F t 8
on . 6 t ea . 00. 3 . ne .
‘

S d econ Di ck i n s on ( S e THE G EE I EW R
L
. .

OF I FE
Elev ent h E d i t i on Cr 8.00 . . . .

A M I SCE LL A N Y O F M E N S d . econ

WI N E WA T E R A N D S O N G N i th
E d iti F t 8 6 5 . ne t .
on . ea . 00.
Dltoh fle ld ( P t TH E VI ‘
LL
AGE
CH CH UR
.

n
, ,
.
S econd E d i t ion . . I llu st ra t ed .

E d i t i on . F eat . 8 00 . I s. 6d . ne t .
Cr 800 6s ne t
. . . .

Ge or e) O A A CA D E R Y L MY T H E E G A D O F S H A ES N L N K P E AR E
Cla u sen
L gN P
.

N N
.

E CT U ES A I TI G I llus tra t e d . . I llus t ra t ed Cr 800 6s net . . . . .

Cr 80 0 7 s 6d ne t
F U RT H ER S T U D IES I N
. . .

B
. .

PR Y B OO K C 8 6 m
Don /d e n ( L ) .
Glop na n ( Coltma n ) T H E TO A U RN TH E A ER s
P
r 0 0.
M N
LE
.

Wi t h
. . . . .

E T : I ts e ri od s a nd Ph a s es
P FF
.

OU K S m ( Th E l f) T H E R E P O R T
N
r e face b y CH A s J I llu s tra t ed .
Du r h a e ar o
£
. . .
.

R oy a l 4 t0 2 as net . . .
O N CA A DA Wi th an I ntrod u

mu t ton B
.

De m 8 ; 7c 00
T H O G H TS U
.

- r ock (L ) . ON
TH E WA R . N i n th E di ti on .

V
1 s 6d
. . net ~ .

WH A T I S T H E K I N GD O M O F H E A E N?

Conr a d ( J o sep h ) . TH E I O OF M RR R ‘
Eti enne .
’
A A A N V L LI E U T E N A N T ,

TH E S EA : e mori es a n d
,
I mp r e s si on s M . 1
ne t
91 4
-1
91 8 . I llus tra t e d .
“

6d .

F ea t . 800. 5s . net .
.

Cou lt on (G CHA U CE R AND H I S Fa i r b r ot h er ( W TH E

L N P
.

W
.

N
E G A D I llust r a t e d . . S econ d E d i ti on .
S O H Y OF T H GR E E N . . H
I
LS
D ent}! 800 1 2 s 6d ne t . . . .
E d i ti on Cr Boo 5s ne t . . . . .

P wit h R M UR R
Cowp e r ( Wi ll i a m)
B IL
fi oulk es ( Ch a rle s ) THE A O E
A EY
O E M S Ed i ted ,
R F
.
. .

N AN D H I S C A T I llus tr a t ed R oy a e
'

a n I n tr od u ct i on a nd ot es , b y J C
£
. .
. . .

D e my 80 0 1 2 s 6d 4t o. t 2 ex. n e
I llus tr a t e d ne t .

D E CO R A TI V E I R O N WO R K F mh
. . . . .

ro t e
R A M BL ES U RR E Y
.

IN
t
Cox ( J S x rt h to t he X V I I I t e n t ur y. I l1us tra ted
£
.
. .

I llus tr a t e d R oy a l et a
.

. . 2 25.

7 s 6d ne t
r C R O M WE LL S A R M Y
. . .

R A M B LE S
’
'
Fi t h ( G H “ .
K E NT
.
IN I llu s t r a t e d cr
A H is t ory 0 t he E nglis h S old i e r d u ring t h e
. . .

800 7 s 6d ne t .
Ci v i l Wars , t h e Commo n wea lth ,
BR
the
. . .

Da l (H h) I WI T H I TI S G S H UN P
r ot e ct ora t e I llu s tra t ed S econd Ed zt eon .

L 7 s 6d
.

Cr Boo n et.
I I TA I llus tra t ed Cr 800 8s 6d
. . . .

~ . . . . .

M t
Fi (H A I )
N N UN N R
5 e

TH E
.

sher
l _

RAD I T I O N
. . n .

Da v i s ( H W E N GLA D DE T I N EUR
RMA NS
. .

TH E O N AN D A GE V I S : 75 . 6d . ne t .

1 066 -1 27 2 . F i tt h E d i t i on D emy 800

RUB Y
. .

i tz Ge r a ld ( E d wa rd THE I AT
'

6d net
1 2s . . .

OF O A KHA M RAM ri n t from P

Da y ( H a r r y A ) , S PA D E CR A F T F t h e F i ft h a n d las t E d i t i on Wi th a Com
me n ta ry b H M AT
‘ .
.

O R , H ow T o B E i G A RD E N E R S econd E di B S ON , 1 n d a Bn 1 3 p h
R
. °
.

t i on Cr 80 0 as net i ca l I u tr u ct i on D oss Cr 800

V T A LEU L AULARD
. . . . .
. . . .

6d n e t

IN
.

R OW E E
.

E GEC T E : H ow TO G V C
AN D E N R I ES HI M
’

EyI ema n ( Rose )

AN D C U N T R Y
S, S S, H RB S TO W F AI A ND C
Y
B .

O . S econd E d i t i on . Cr 800 . . N
E S F eaf . .
6 M ET H U E N A ND C O M PA NY L M I I T ED
Hu t ton ( E d wa rd ) TH E CI T I E S F O Kea tn oh n) .
E d i t ed , wit h ntro P OE M S I
UM B R IA I ll
.

d uct i on a nd
.

. us t r a te d F if t h E di t i on o t e s , b y E d e S é LI N CO U R I N
W
. . ’ ‘

F r on t is p ie ce i n Ph otogra v ur e
.

C r 8 .
7 6d 00. t 3. ne it h a
T H E CI T I E S O F L O M B A R D Y
. .

I ll . us Th i r d E d i t i on D e my 80 0 r 6d
r oa net . . .

bWli
. .

d C B 6d
T H E C I T I ES O F R O MA G NA A N D T H E
t ra t e
C H R I S T IAN Y E A R
7 t r oo s. ne
( J oh n )
. . . . .

Ke e THE
I W
.

N
.

MA R C H ES I ll d C 8w 7 6d . u s tr a t e . r. . s. .

L O C K I ll
th
.
an n t r od u cti on a n d
us t ra t e d
ot e s
Thi r d E d i t i on
by
F cap . .
.

F L ORE N C E A N D N OR T HER N T U S
800
’

. ne t .

C AN Y W I T H G E N O A I ll d u s tra t e Ke i yn ack ( T IL ), l D TH E
D R I N K P RO BL E M O F T O DA Y I N
, . . . . .
,
Tl i d E d i i
z r C B 7 6d t t on r. oo s. ne -

S I E NA A N D SO U T H ER N T U S C AN Y AS
. . . .

ITS M E D I CO S QC I O L O G I CA L '

I ll
.

d S
u s t ra t e E d iti C B econ on . r oo 5. PE CT S S econd a nd R evi sed E d i ti on
7
. d . .
. .

6d t ne r o s 6d . ne t .

VE NI C E A N D VE N E T IA I ll
. . .

C 8 m 7
d . u s t ra t e .

Ki d d ( B e nj a mi n)
‘
T HE S C I E N C E O F
6d
P W ER
.

7 2 t s. ne

O
E igh th E d i t i on 7 s 6d
NA PL ES A N D SO U T HER N I T A LY
.

Cr 827 0
. . .

. . . . . .

no
I ll
.

d C 8
u s t ra t e 7 6d t r 71 0. s. ne

RO M E I ll B A RR A C K ROO M
. . .

gA Lfl
. d l i d E d iti C
us t ra t e . r on . r . Ki li n Rud ya r d ) .

8 6d DS h Thous a n d
y
t Cr 8zi0
C O UN T R Y W A L K S A B O U T F LORE N C E
00. 3 ne
g
7 . . )

-
. . . . .

B a ch a m, 7s . ne t A ls o F ca fi Boo
. .
‘
.

I ll
.

d S
us t r a t e d E d iti F } 8. econ on . an . 19 0.
6 t
s . ne .

S q ua r e / fen)
T H E C I T I ES O F S P AIN
800 E a ch 33
I ll d . u s tr a t e .

TH E SEV E N SE A S 4 th Th
. .

r
.

o
ne t .

ou s a nd
N

F ift h E d i t i on Cr 6d n et .
. .

73 Cr 89 0 B h m 6d
. .

Al
.

Ib
F ca p
.

t uc r a ne so
Cl t h 6 t : l t h 7 6d
. .

B R AN D A D m i
, . . .

8m
( He n r i k ) o 3 . ne ea er 3. ne t
s en at c
S i Ed iti
.
ra ,

Al
. , . .

l b y W I LL I AM W I LSON
.

Poe m, t r a n s a t e d .
so Tw
a V e r v ce o n. o olmn es .

F ou r t li E d i t ion Cr 800 ne t
Sq / } 8 E
ua re h 3 t ca 00 . ac 5 . ne

TH E F I VE NA T I O N S
53
. .
. . . . .

C H R I S T IAN M Y S T I C I S M
th T h . 1 20 ou s a nd .

In ge ( W C 8 r. B k m 7 6d 00 . t Al uc r a s. ne so F ea t
( Th e mp L Cl th 6 t i t h 7 6d
. . , . . .

Ba t on f F th e ct ure s o 8 00. o 7. n e ea er , s. ne t .
S i Ed i i
ou r ,

Al
.

E d i t i on . Cr . 8 so Tw
a V e rv ce t on . o olu mes .

Sq f ) 8 h 3
TH E Y E A RS
E
A H I S T O R Y O F T HE
ua r e t en 11 0. ac 3 ne

B E T WEE N C
. . .

I nn ( A es r 8 00
B R I T I SH I N I NDIA Wi h M p d
. . . .

t a s an B u ck r a m . 7 s 6d ne t A ls o on t hi n p ayer . . ,

Pl
. .

S
a ns . d E d i ti C 8 7 6d econ on . r. vo . s . . F a x) 800 B lue clot h ,
. n et ; L i nc;
.

t
ne la mbs k i n , 7 s 6d net
i
.

Al
. . .

E N GL AND UND ER TH E T UD ORS so a S er vi ce E d i t on Two .

Wi h M p fi ij t/ E d i t i S gu m fi af 80 0
.

D m] B E a ch 35 ne t
D E P A R T M E N T A L DI TT I ES
t
. . . .
a s. i on . e oo.
1 2 s. 6d . n et . 8 4 th Tha n .

I nn ( M y) S C H O O L S
r Cr 800. B uchr a rn , . 7 5 6d n e t
P AIN T C
. . ' . .

es ar OF A ls o F eat 800 lot h , 65 n et ; lea t h er

I N G I ll t d T hi d
. . . .

u s ra te r E d i t ion Cr 7 s 6d net
S
. . . . . . .

80 0 . net . A lso a er vi ce Ed i ti on Two .

S qu a r e f cap 800 E a ch 3s ne t
J e nk s AN O U TL I N E O F E N G H Y MN B E F O RE A CT I O N I ll umi d
. . . .

LISH L G O VER NM E N T Th i d
(E n a te
.
. .

CAL F ) 6d
R R C K E NSOR C RE C E S S I O NA L I ll mi t d
r t
.
oo . rs. . ne .

E d it i on . e vi s ed b y . . . . r. ‘

. u na e .

A SHOR T H I S T OR Y O F E N GL I SH
6d
T WE N TY P OE M S F R O M R U D YA R
I s. t . ne .

LA W F RO M EA R L IES T T I M ES TO
: T HE
K I PL I N G g6 th Th d F m} 8
THE E OF Y E A R 9 D my B ND THE 1 1 1 . e oo.
I s. net .
. o ou s a n . . 00

6d net

b
1 03 . . .

J oh n s ton B R I T I S H CE N L a m ( Ch d T H E CO M
a r le s a n Ma r y)
L E T E WORKS L UCA S
.

( Si r H IL ) .
T R A L A F R I CA I ll F Ed i t ed b y E V
.
. . .

us t r a t ed
. .
A N ew a nd R evis ed E d it i on i n S i x Volu mes

N EW WO R LD
E d iti C 4t on r o. ne t .
Wi t h F r ont zlrp i eces F cap 89 0 E a ch
T H E N E G RO I N
. .
. . .

TH E n et
I ll C w £
.
.

d u s tra t e . ro n 4 to . 1 1 3 . net .
T he lm vo u es ar e

La d y) of Nor wi ch REVE L A M ISC ELL A NEO U S P ROSE I I EL I A A N

T I O N gO F DI V IN E L OVE E d i
J uli a n 1

F O R C H I LDREN
L AS T ES S AY S EL I A m B
. . . .

te d by T HE or oo x
P LA YS A ND P OE M S
. .

G R AC E W
.

S i tl E d i t i A R RA CK x z on. Cr I v.
L TT E S
. .
.

Boo .
sa nd . v . and VI . E R .
 GE N E R A L LIT E R AT UR E
La ne-Poole ( S ta n l e y) . A H I S T OR Y OF A W AND ERER I N H O LL AND I ll us
E GYPT I N MIDD LE A G ES
.

I ll
THE
ii tr a t e d Si xt eent h E d i t i on Cr 8 vo 8 s 6d
. . . . . . .

d S
u s t ra t e . econd E d t on , R evi s ed . Cr . n et .

8 00 .
os . n et .

A W AND ERER I N L O ND O N I ll us

La nk es te r S C I E N C E F RO M tra te d E igh tee n th E d i t i on , R ev i sed Cr .

)
.

( Sir R
.

AN E A S Y C H AI 13 I ll
.
80 a 8s 6d net
d E igh t h
. . . .

u s t ra t e
L O ND O N RE V I S I T E D I ll
.

E d iti C 8 on
7 6d t r . 00. s. . ne . . u s tr a ted . Th i r d
S C I E N C E F RO M AN E AS Y C H AI R
.

E d i t i on . Cr . 80 0 8s 6d . . . ne t .

I ll A W AND ERER I N P A R I S I ll
.

S d S i
econ d F i t E d i ti
er es . us tra te . rs on . . u s t ra t e d .

C 8 6d t
DI VERS I O N S O F A NA T U R A L I S T
r .
7 00. s. . ne .
T h i r t ee n t h E d i ti on . Cr . Boo . 8s 6d . . n et .

A ls o F cap Boo 6s ne t
I ll
. . . . .

d S
us tra t e
6d
d E d i ti
net
C . 8 econ on . r. 00.
A WAND ERER I N F L ORE N C E . I ll us
7s . . .

t ra t e d . S i x th E d i t i on . Cr . 8o o . 8s . 6d .

Le w i s ( E d wa r d ) E D WA R D C A R P E N ne t .

A WAND ERER I N VE NI C E I ll
A N E X P OSI T ION AN D
.

TE R A AN PPR ECI A
d . us tr a te .

6s S d E diti
econ C 8 8 6d t on r. 00 s. ne
Cr 8290 net
. . . .

TH E O P E N RO AD A L I TT LE B OOK F O R
. . . .

:
L Walter ) P AU L WAY F A RERS Tw ty
ock THE
i
ST th Ed i i en -
s ev en t
M A S R B UI LD ER
. .
, . on .

. Th i r d E d i t ion . F ea t 80 0 os 6d
. . . . ne t . I nd a P ap er , 7 s 6d . .

C r 5 t s . ne . me t .

T H E B I BL E A N D C HR I S T IAN L I F E
.
'

.
A lso l llu s t r a ted . Cr .
4 to . 1 5s . ne t .

Cr 800 6s ne t
TH E F R I E ND LY T OW N A L I TTL E B OOK
F O R T H E U R B ANE
. . . .

N i n t h E d i t i on F ca p
M AN A N D T H E
( S i r Oli ver ) 8m
. . .

U I VE R S E A STU DY O F H I NF LU EN C E és ne t
.

ADVA N C E I N S ENT I F C K NOW S UN S H IN E

. . .

FI RES ID E
T E
OR T HE CI I AND . N i n th
L EDG E UPON U NDERS TA D I N G O UR N or
C HRIS T I A NI T Y N i th E d it i C w 8 . n on. ro n oo. C H A R A CT ER A N D C O M E D Y E igh th .

A S TU D Y I N
E dit i F p 8
on . 6 t ca 0 0. s . ne

T H E S U R V I VA L O F M AN T H E G E N TL E S T A R T
. .

U NREC OG NISED HUMA N F ACULT Y S t h A C HOI C E O F :

E d i ti C 8 6d t
. even L ETT ERS B Y E N T ER TA ININ G H A NDS .

M O D ER N P RO BL E M S C B
on .
7 r 00. s. ne
. . .
T th E d i t i
en F ) 8 6 t on . ed . 00. s . ne .

T H E SE C O ND P OS T
r. o o.
F if th E d i t i
.
. on .
t
ne
F fi 8 6 t
.

R A Y M O ND ;O R L IFE AN D D EAT H I ll ca 90 s ne
. . . .

H E R IN F INI T E V A R I E TY A F E M I INE
, . us
t ra t ed E leven th E d i t i on D emy Soo : N
. . . 1 5s .

P OR T R A I T G ALL ER Y E ighth E d i t i on .

A F T ER S HOR T C HAP
.

F p B
ca 6 t oo s ne
WAR
T ERS ON S U BJECT S S ERIO U S P RACT I CAL
. . . .

TH E AND
or
:
G OO D C O M P AN Y A RA L LY O F M EN :
I MPOR T A ERAG E C I T I EN I N
.

F OR T H E V Z Ar D F ou r t h E d it i on . F eat . 8v o . és . n et .

9 5 O N W A RD
. .

1 1 E ight h E d i ti c p S.
“
on . a .

ON E D A Y A N D AN O T HER S h . even t
Ed i ti on F ea t 811 0 é t s . ne

S i th E d i t i
. . . .

Lore b u r n ( E a r l) . C A PT U RE A T SE A .
O L D L AM P S F OR N EW . x on .

S econ d Ed i i C 8 6d t
t on . r. 720. as . . ne F p 8
ca . 6 t 0a . s . ne .

L O I T ERER S H A R V ES T Th i d E d i t i
.

H OW T H E W A R C AM E Wi h M p ’
. t a a . . r on .

Cr 800 .
7 s 6d . . ne t F p 8
ca . 6 t 00. s . ne .

CL O UD A N D S I L VER Thi d E d i ti
. .

L
r on .
L E TT ERS
.

or i mer ( Ge e e H or a ce F p 8 m 6 t
F RO M A Ss L F MAD M ER C H AN T
ca . . s . ne .

T O H I S S ON I ll d T w ty
-
L I S T E N E R S LU RE A N O B L I QU E N A RR A
’

. us t ra t e . en
T ION Tw lf t h E d i t i F p 8 6
. e on . cez . 9 0. s . n et .

O L D G OR G O N G R A H AM I ll t t d OVER B E M E R T ON S AN EAS Y G OIN G ’

: -

S econd E d i t i on . C r . Boo . 6s
.

. ne t
us ra e
.
.

C HRONI CL E S i t th E d i t i .F t x een on . ea .

B oo. é t s . ne

E d iti
.

Lor i mer ( Norma ) TH E BY W A T ERS M R IN GL ES ID E T t zu on .

E GYPT I ll
. . .

OF . us tra te d Th i r d E d i t i on
. . F } 8ax . 6 t 00. s . ne .

Cr 8120 . .
7s . 6d . net .
L O ND O N L AVE ND ER Tw lfth E d i i . e t on .

L F OF LI E C H A R L ES F p B 6 t oo. s . ne

L ANDMA RK S F if th E d i ti F t 8
ca
u ca s E TH E
. .

LA M I ll
.

us tr a t e d S i x t h E d i t i on . . D enty . on . en . vo.
800 . ms . 6d net . .
8 M ET H U E N A ND CO M A N Y LI MIT E D
P

T HE B R I T I SH S CH OO L A A NEC DO TAL z N T H E B U R G O MA S T ER O F S T I L E
G U IDE T O T HE B RI T ISH P A ND A I NT E R s M O ND E A P LAY IN T HREE A CT S :
‘

PA IN T IN G S IN T HE N AT I ON A L G AL L ERY l d by A L EX A NDER T EIXEIR A DE

.

T ra n s a t e

M ATT OS F j 8 5 t
.

F ea t Bo o és n et ea ) 0 0. 3 ne

T HE B E T RO T H A L ; OR T HE B LU E
. . . . . . . .

A B OSW E LL O F B A G H DAD AND

O T H ER ES S AY S Th i d E d it i F p B I R C H O SES T l d b A L X
‘

,
,
D O ra ns a t e E
on
A NDER T E I XEIR A DE M
.
. r . ea .
‘

p B A T T os . ca . oo.

T W I ( T E A GL E AND D O V E Th i d
6 3 t ne

M O UN T AIN P A T HS T l d by A LEX
. .

’ ‘ r
ra n s a t e
.

A NDER T EIXEIR A DE M
.

F p 8 A T T O s. ca . 00.

Lyd e k k cr T HE OX AND I T S M h afl u
a A H I S TOR Y O F E GYP T
'

K IND RE D I ll UND fiR T H E PT O L E MAI C D Y NA S TY

.

d E d iti
u st r a t e d Cr 800 7 s 6d
I ll
. . . . . . .

d S
us t ra t e C 8 . econ on . r . 00 .

ne t .

C R I T I CA L RO MAN C AN O N L AW
H I S T OR I C A L E SS A Y S E d i d b y F
a oa u la y ( Lor d ) Ma lt la nd
C H U R C H O F E N GLAND
.

C M ON TAGU E Th V l m C B
. te .
I N TH E .

. . r ee o u es . r . oo .
R oy a l Svo . I os . 6d . n et.
1 8s . n et . p

l
Ma cd on a d ( J R FL ) A H I S T OR Y O F
F R AN C E
. . .

Th r ee Volu mes Cr Swo

E N GL AN D S IN CE
. . . .

E a ch 6d n et Ma r r i ot t ( J A
W AT ER L OO Wi h M p
1 0s . . . . .

. t S a s . e cond
E d i t i on , R eveked D e m 800 ms 6d
’

McDou ll lllla m) AN I N T R O D U C . . . . ne

TIO T SO C IA L P S YC H O L OGY Ma se fi eld

J
A S AI L OR S G A
.

( J oh n)
’

L AND Sl
.

Tw elft h E d it ion . Cr . 800 . 7s . 6d . ne t . . e e ct ed a nd E d i ted . .

E d i t i on Cr 8m
B O D Y AND MIND A H T ORY A N D A : IS
. . . 65 . n et .

D EFEN C E A NI M IS M F fl E d i t i or ou r i on . Ma s t er ma n (0 F G T E NN YS O
A S A RE L I G I O U S
. . . . .

D e my 80 0 . 1 2s . 6d . ne t . EA CH E R S econd .

E d i t i on . Cr . 8290 .
75 . 6d n et . .

a e t e r lln ck T HE BL U E Ma ur i ce) l OR I G INA L I LL U S T R A

‘

E N GL I S H CO N S TI T U
Med ( D .I
B I R D A A IR Y P LAY IN S A CT S e
.
. .

: Ix
TI O s O
l d b y A L EX A NDER T EIXEIRA DE
‘
.

T ra n s a t e
T I O N A L H I S T OR Y C 8 8 6d r. 00. s.

b k F y
. .

M F p B
A T T os . 6 t Al F t ca . oo. s ne . so ea .

b
.

B oo t Of h 2: ne t e a o ve oo or t
L I F E A F T ER L I F E
. .

E d i t i on s i n b M i le s ( E u s t ace)
MA R Y MA G DA L E N E A P LA Y IN T H REE
on e a ll h a ve ee n i ss ued . .

:
OR T T ,
HE RE I NCA RN AT I ON H EO R Y QO F .

ACT S T l d by A EX A NDER T
. ra n s a t e L E Ix E I RA
C 8 r 3 . 6d
T HE PO WE R O F C O
t
00. s. ne . .

DE M Th i d E d i t i F f
A T T os . 8 r on . ed . 11 0 .

H w o ACQU IRE I T To
,

5s . ne t . A ls o F eap . Boo . as . ne t .
C 8 6 t
D E A T H Tra ns l a t ed b y A LEX A NDE R
r vo. s . ne
T EIX P RE V E N T I O N AND CU RE S
. .

E I RA DE M A T T os F ou th Ed it ion
.
. ecan o
. r . F cai o .

80 0 3 s 6d ne t .

O U R E T ER NI TY T l t d b y A LEX HE A LT H W I T H
. . .

Mi es ( Mu E us tace)
UT M EAT
.
ra ns a e
A NDER TEIXEIR A DE M ATT O S S d S ix t h F cap 8 m
.

i
. .

econ
6d
.

13 net .
E d t i on F 800 6s net
.

p
.
.

e a

T HE L I F E AND L E T
.
:
. .

T H E UN K N O W N G U ES T T l r a ns a t e d
T ERS O F
H i lla ll ( J . (L) .

S I R JOH N E VE R E T I
A LEX A NDER T EIXEIR A DE M
.

MI LL AI S
'

A T T os
h i r d E d i t i on . Cr Boo . . 6s . ne t .
.

. I ll d Th i d E d i t i
us t r a t e . r on .
D emy 80 0 1 2s . 6d n et
P OE M S D Verse b y l
. . .

l e ( J G A H I S T OR Y O F E G Y PT
. one i n to E ng is h
B ERN A RD M Mi
UND ER 13O MAN R U L E I ll
I A LL . S econd E d i ti on Cr . .
n . .

80 0 .
5s . n et . d . us t r a t e .

T HE WR A C K O F T HE S T O R M . Th i r d
R I C HES AND
I
E d i t i on Cr Boo 6s net H on e y ( Si r Le o Ch i oz z a )
P OVER TY
.
. . . . .

E leven th E d i t i on
T H E M I R A C L E O F S T AN T H O N Y A : D e my 800
, gr o . .

P LAY I N O ACT T l d b y A L EX 5s ne t
.
. . .

A NDER T EIXEIR A DE M ATT OS F p B

NE ra n s a t e

D R AMAT I C V A LU ES
.

Mon ta gu
ca o o. e (c
i
. . . .

S econd E d t i on . F eat . 87m .

5s net . .
 Io M ET H U E N A ND CO M A N Y LIMIT E D
P

RE G INA L D IN R U SS IA F ea t 871 0 ASK MAM MA ; T H E R I CH E S I °

C O MM O N ER I N R ENG L AN D I ll
. . .
’

3s 6d . . n et . . us

S sh id r c si tz r
( Ph i i p ) l . R U BB ER . I ll us
t ra t e d
ne t
. S econ d E d i t i on . F ea t . 8 v0 .
7 s 6d .

t ra t e d S econd E d z t i an D ewy 80 0
J AUN T S
.

. . . 1 5s .

J O R R O CK S S AND
'

j OLL I
n et .

T I ES I ll d u s tra te S i x t h E d i t z on F ea t
l T O MM Y S M I I H S
. . .

( Ed m nd )
'
Se
' ‘

ou s u 8 6 t
ANI MA L S I ll
. vo s ne
Si t M R F A C E Y R O M FO R D S HO UND S
. . .

d th E d i . us t ra t e . x ee n ’

I ll
.

T O MM Y S MI T H S O T H ER ANI MA L S
ti F ) 8
on . 3 6d t en . 71 0. s. . ne .
d Th i d E d i ti
us t r a t e F t 8 . r on . ea . 71 0
’

7 6d
s. t ne
I ll H A W B U C K G R AN G E ;O R T H E S P OR T
. . .

d S t h E d i ti
u st r a te F p 8 ev e n on . ea 00.

I N G AD VE N T U RES O F T H O M A S
. .
,

3 6d st ne

T O M M Y S MI T H AT T H E ZOO I ll
. . .

. us S C O TT E Q I ll d F ) 8, S . us t ra t e . ea . v0 .

t ra t e d S econd E d i t i on F ea t 8710 .

PL AI N O R R I N GLE T S I ll
. .

T O MM Y S MI T H A G AIN AT T H E
25 .
9d .
d ? u s t ra t e
2 00 F p 8 ca 7 6d t v0. s ne
I ll H I LL IN G D O N H A LL W h C l d
. . . . .

d F p 8 9d
J A C K S IN S E CT S I ll t d C 8
u s t ra t e . ca . 210 . as . .
It 12 o ou r e
Pl H E AT H d J E LL I
.

6s by W
’
. u s tr a e . r. 00 . . a t es I LD R A K E , , an
ne t . COE . F ea t . 800 .
7s . 6d . n et .

S h a k es p ea re
O U R F O L I OS
(W i lli a m) .
S u s o ( H e n r y) L I FE OF THE TH E
TH E F BL ESSE D H E N R Y S U SO By H I M SE L F
.

1 62 3 ; 1 6 3 2 ;1 66 4 ;
1 6 85 E a ch £4 4 s net , or a
,

co m pl e te se t, T l d b y T F K NOX Wi h I
ra n s a t e
.

t an nt ro
.

£ by D E A N I
. .
. . .

S H AKE
r 2 1 2 s . ne t d i u ct on
P OE M S O F W I LL IA M
.

TH E C 8 6 t 00.
S P E A RE W i h I d i
r s . ne
N
, . .

t an n tr o u ct o n a n d ot e s
t n (E W F UN G I A N D H O W
TO K N O W T H M
Sw
b y G EOR G E WY NDH AM
.

an o
D my 8
I ll
. .
e 00. B u ch
d C 8
.

us t ra t e r v 0.
m, 6d
. .

n et .
.

ra 1 2s.
6d
BRITISH
t
.

r os . ne
P OE M S PL AN T G A LL S C 8
. .

Sh e ll y e ( Pe r cy B ys sh e) Wi t h r.
I 11 0.
. .

A
.

n t r od uc t i on by C L U T I O N B R OC K a nd ’

6d
‘

an ne t
'
.
1 0s . . .

no t es
D e my
by C .

561
D L o co c x . T wo Volu mes
r s . n et .
. .

Ta bA R T ( MW g
or TH E
ar IN
a r et
A b
S AIN T S
b l
i t h t h e ir t t r i u t e s a nd S y m o s
S l a d en gl ) S I C I LY : T N W ( Dou as HE E
A h a b e t ica lly
.

rr a nge d u st ra t ed A I ll
W I T ER R ESO T A E y l md i f
.

g
. .

R n nc c o a o
N
/ r d E d z t zon
S i ily W i h 3 4 I ll t i F eet} 800 5s ne t
.
.

M
m f I ily
. . .

E L E M E N TS O F M E TA
d t 2 u s t ra on s , at a an

f h R ilw y Sy
c
,
T bl
.

P H YS
a a e o t e a a s te o c .
Ta ylor A .

R evi s ed . Cr . 800 .
7s . 6d .
CS . F ou r t h E d i t i on . D ewy 8 00 .

1 zs . 6d . net .

(H T R AD E UNI O NI S M J C O M IN G
S AIN T S
Ble s s e t . .
Ta ylor ( . TH E OF TH E
. S econd E d i t ion . Cr . Svo . 6s .

S mi t h ( Ad a rn WE A LT H O F net

NA T I O N S é
TH E
.

d it e d b y E D w C A NN A N Th omas ( E d wa rd ) . MAU R I C E MA E
£
. .

Tw V l m o to u es . D eniy 871 0 1 5s . . ne .
TE R L I N C K I llus tr a t e d
S d E d ti . . eeon z on .

S mi t h ( G F H b t) G E M S TO N ES er er - Cr 8v0 6s n e t
A L I T ER A R Y P I LG R I M I N E N GLAND
. . . .

A N D T HE I R DI S T IN CT I VE C H A R A C
. . .

T ERS I ll I ll
.

d S d E d it i
. C us t r a t e . e con on . r . d D my 8
u s t ra t e 6d . e 00. r es . . n et .

6d
( M y W ) DAI LY S T R E N G T H
8 7 00. t s ne

GO LF D O S AND D O N T S
. . .

Ti l t es on ar
F O R D A I LY N EE D S
.

li fl
' ’ ’

St a nc e. .
1 m ty fi f t h . ea -

S i t h E d i ti
x F p 8 t on . ea . v0 . as . ne .
E d it i on M ed i u m 1 6 11 1 0 3s 6d net

b L FE N D W ORKS
. . . . .

T H E L E TT ERS O F
St ev e n s on
RO B ER T O UI S S T EVE N SO N E d i d Toyn ee ( Puge t ) DAN T E A L I G H I ER I
W I ll
.

I A
.

te
H IS ith 1 6
S IDNE Y C O LVIN A N w R us tra
. .
.

b y Si r . e e
t i on s F ou r t h a nd E n la rged E d i ti on Cr

i E d i t i on i n f ou r volu me s
. . .

a r r a nged F ou r t h
8v0 6s ne t .
.
. .

E d i t on fi ca fi 80 0 E a ch 6s net L ea t he r ,
E N GL AND UND E R
. . . . .

e a ch 7 s 6d ne t r ev elya n (G
H AND L E Y C ROSS S T UA R T S Wi h M p d Pl
. . . .

THE t a s an a n s.
(R
.

Eur t ees
D emy 821 0
I ll
.

S even th E d i t i on 6d
.

1 2s . n et
us t r a t e d E igh t h E d i ti on F eat 811 0
. . .

I ni g ) T O W N PL ANNIN G
.

. . . .

6d n et
7s Tr i ggl (H o
P AST P RES EN T AN D P OSSI B L E I ll
. . .

S P O N G E S S P OR T IN G T O U R
. .

MR u s t ra
’

I ll
.
, ,

Wi d e
. .

u s tr a t e d .
A l i on r t h E d z t i 0n - . F ea t . 8 21 0 . t ed . S econd E d i t z on . R oy a l SW .

6d n et . 1 65 ne t
75
. .
. .
 G E N E R AL LIT E R A TU R E I I

Un d er h i ll ( E ve yn ) l M YSTIC I S M A A L I TTL E B OOK O F L I F E A N D

l pm D E AT H N i t h l d i i
. .

S t ud y i n t h e a t u re a n d D e ve N o ent of S m ll n e een t : t on
l C C l t h 6d t
a
M p
. .

S ir i t u a onsc i o u s ne s s S event /z P tt 8
’
an s . o 11 0 . o , 2 s. . no .

D en zy Se a . 1 55 . n et .

W t a ers(W I T A L IAN S C U LPT ORS

HOW TO PL AY G O L F I ll
.

i
Y ar d on ( Ha rr y) d C 8
u s t ra t e 7 6d t r 110. s. ne
I ll
. . . . . .

u s tr a t e d . E levent h E d t i o n . Cr . 811 0 .

W tt ( F n i )
a C AN T ER B U R Y P I L
ra c s
ne t
G R IM S A N D T H E I R W A Y S W i h
.
.

t
F i pi i C l
a

RE ADIN G S I ll
.

V er n on ( Hon W Wa rre n ) ro n t s d h e ce n o ou r a n 1 2 ot er u s t ra
ON IN ER N O O DAN T E With
. . .

THE F F t i ons D e my 8210 1 os . 6d ne t

I M OORE
. . . .
.

an n t r od u ct i on b y t h e R e v D . r. .
We i ga ll ( Ar t h u r G UID E T O E A
T w o Volu mes S econd E d z t i on , R ewr zt ten
AN T I QUI T I ES O F U PP ER
.

TH E
. .

Cr Boo . r ss

RE ADIN G S O N T H E P U RG AT OR I O
. . ne t .

E GYPT F RO M ABY DO S SU D A N : To THE

F RON T IER I ll d S d E d i ti
O F DAN T E W i h I d i by t an nt r o u c t on Cr
'
800 ms
.

6d
u s t ra t e . econ on .

h l D E A N C H U R C H Tw V l m ne t.
.
. . . .

A SHOR T H I S T OR Y
t e ate o o u es .
Wells
.

TI i d E d i ti R i d C t OF
RO M E Wi h 3 M p
z r 5 on , ev s e r 1 5 ne

RE ADIN G S O N T H E P A R ADI SO O F
. . . .

. S i x t een t h E d i t i on . t a s.

DAN T E W i h I d i b y h t an n tro uct on t e

Cr . 6s .

B ISHO P R I PON Tw V l m S d
.

or . o o u es . econ Wi l d e ( Os ca r ) T H E WORKS O F OS C A R
W I L D E Th i t V l m E t 8
.

Ed it ion , R evi s ed . Cr . Boo . 1 53 . net .

r een o u es . co
. . 71 0.

c k e rs E N GL AND I N
( Ken n eth IL )
E h 6 6d
ac t
LORD A RT H U R S
s .

C RI M E AN D
. ne .

L A T ER M IDD L E A G ES Wi h
.
’

TH E t Ae E s
t
POR T RA I T O F M W H I I T
.

Mp
.

THE
S econd E d i t i on , R evi s ed D ewey R.
D UC HESS O F P A D UA I II P OE M S
a s. .
. . . HE
80 0 ms 6d ne t Iv
L AD Y W FAN A W O MA N
. . . .
. . . .
'
I N D E R M E RE S v

L H AS A O F N O I MP R TA N CE V I A N I DE AL H
. .

Wa d d e ll L. A N D I TS O
B A ND VII T I MPOR TA N C E O F B EIN G
us
I ES Wi h R
. .

M Y S TE t e c or d o f t h e E x HE
a
EA RNES T VIII A H O U SE O F P O M E
. .

I ll
.

p e d i t i on o f 9 3 9 u s t ra t ed T/zi r d
1 0 -1 04.
I N T EN T IONS
.

G R A N AT ES
.
.

M ed i u m 89 0 1 2 s 6d net D P Ix x I: RO
F U NDIS AN D P RISON L E TT ERS E SS AYS
. . . .
. . . .

xI
J R AM BL ES I N SA LO M E A F L OREN T INE T R AG ED Y
. .

Wa d e ( G W
.

SO M ER S E T I ll
a nd x11
SA IN T E CO U R T IS A NE
. . . .
, ,

u s tr a t e d . Cr 8 71 0 7 s 6d d
an L A x rv.
S E L ECT ED P RO S E O F O S CA R W I L DE
. . . . . .

ne t

A H O U SE O F P O M E G R ANA T ES I ll
. .

Wa g ner ( Ri ch ar d ) R I C H A RD W A G t ra ted Cr 4 t0
. us

N ER S MU S I C D R AMA S I p ne t
.
. . . 21s . .
’
nt er re t a
mb d yi g W g Wi ld i ng ( An th on y F) O N T H E COURT
B y A L I C E L EI G H T ON C
pl
.

ti w
’

O FF Wi h 58 I ll
o ns , e o n a ne r s o n ex a na .

ti ons LE A I H E R
' ‘
AND t us t r a t ions S e ve n t h
d B A SI L C R UMP i
. . .

an F t 8 E / 4 . ca . 00. d e 1 5. E d t i on . Cr . 80 0 . 6s . n et .

t
ne
R IN G O F
.

N I BE UN G Wi ls on ( Er n e s t IL ) A N A TU RA L I S I I N ’ ‘

WES T ER N C H INA I ll
.

T HE T HE L
d d
.
S . us tr a t e econ
S i tl E d it i V l £
.

x z on .
Ed i t i D my 8 on
LOHEN G RIN P A ND A R S I I’A L.
t . 2 o s. e 00 . 1 r os . ne .

W d ( S i Ev lyn ) F RO M MID S H I P
oo r e
M AN T O F I E LD MA RS H A L I ll
.

T RIS TA N AN D I SO L DE .

t ra t e d F if th E d it i on
-

D emy 871 0
. us
6d
M A S T ERSIN G ERS
. . . 1 2s . .

T A NNH AU SER AND THE

ne t .

REVO LT I N H INDU S TAN (

N U RE M B U R G
or .
THE
I ll
1 8 57

( E l i b th )
d S d E d i ti
u s t ra t e C econ on . r 800
WITH T H E
. . .

W t h
a er ou se za e 7s 6d ne t

S IM PLE H E A R T E D L i l H mili
. . .
.

i Wood ( L i eu t w
- tt e o
E d mon d s ( Col
e s.
B ) a nd
.

1 ni r d E d t i on S ma ll P ott 8 21 0
A H I S T OR Y CIVIL
‘ . . .

J
.

IL ) . OF T H E
. .

WA R I N T H E UNI T E D S T A T ES
.

TH E H O U SEC H ERR Y T REE

B Y TH E Wi h I SPENSER
L l
S ec on d S e r ie s of i t t e H omi i es
l ( 86 t an nt r od uct i o n by
.

A
1

W I LKINSON W i h
1
S ma ll .
. t 24 M p a s an d Pl a ns .
P ott 8 21 0 3 : 6d n et
C O M P ANI O N S O WAY
. . .

F TH E B e ing
l M R Wor d s worth P OE M S Wi h
.

S e ec t i on s for orn i ng a n d E v e n i ng ead t

N OW E LL
an
I d i N
.

in g Cr 871 0 n t ro uct on a nd ot e s by C
7 s 6d ne t
T HO UG H T S O F A T ER T IA R Y S d SM I T H
. . . . . . .

. Th r ee V olu mes . De m
y 8 21 0 . 1 85
econ
.
.

E d iti S m ll P t t 8
on . 6d t a o 71 0. 1 s. ne

I R I SH
. .

VERSES S d E d t i E l g d F t . econ z on , n af e . ca . Yea t s ( W . A B OOK OF

VERSE . Thi r d E d i t i on . Cr . Boo . 6 5 net . .
 [2 M ETH U E N A ND C OM P A N Y LI M IT E D

PA R T I I . A S E L E CT I O N
-
OF S E RI E S

An cie n t Ci t ie s

Ge n e ra l E d i tor, S I R B . C . A . WI N D L E
Cr . 8 71 0 . Or . n et ea ch volu me

With I llus tra tions by E H . . N E W, a n d oth e r A r tis ts

B RIS T O L A lf d H y . re a rv e . E DIN B U R G H M G W i ll i m . . . a son.

CAN I '

J C C
E R B U RY . . . ox . L IN C O L N E M ] S ymp . . a nse so n.

C HES T ER S i B C A W i d l . r . . . n e. S HREWS B U RY T A d . . u e n.

D U B L IN 8 A 0 F i p i k
. . . . tz a tr c . W E LL d G LA S T ON B U RY T S
S an . . . H ol mes .

Th e Ant i quary B oo ks
’
s
Gene ra l Ed i tor , J . C HA R L E S CO X

D emy 8 00 . 1 0s . 6d . n et each volu me

Wi th N u merous I llustra ti ons

A N C IEN T P A I N T ED G L ASS IN E N GLA ND E N GL IS H C OS TUM E F m P ro re h is t or c i Ti m

Ph l i p N l
. .

e s on . t ho E d f h
t E igh e n o t e t e e nt h Ce n t ur
G g Cli h
I
i
e or e nc
A R C H EO LOG Y A ND F ALSE A N T IQU I T IES
.

R M.
1
u nr o .
.

E N GL ISH M L IFE C ON A S T I C . ar d i na lG a sq u c
fi ou r tk E d i ti on
B E LL S OF E N GLA ND T HE C
.

J J a non
E N GL ISH S E AL S J H y B l m
R
. . .
,

a ve n S d E d i ti
. econ on .
. . a r ve oo .

B R A SSES OF E N GLA ND T HE H b t W er er
F O L K L ORI;
S G L G mm
- A N H IS T ORI CAL S As CI E NC

M k l i Th i d E d i ti
. .
,
ir . . o e.
ac n r on .

C A S TL ES A ND WALLED T O W NS OF E N GLA ND
T HE A H y
,
G I L DS A ND C O MPA NIES OF LONDON T H
G e or ge U n w n. i
,

H ER M I T S A ND A N C HORI TES OF E
. . a r ve .

N G LA NI
C E LT I C A RT IN P AGA N C HRIS T I A N AND T HE R h M y C l y ot a ar a

T I M ES J R mi lly A ll S d E d i ti
. .

o en. econ on .
M A NOR A ND M A NORI AL R
C H U R C H W A RDENS A CC O U N T S J C C
. .

'
. . . ox .
N h i l at an e

D O M ESD AY I N QU ES T T HE A d l ph B ll d
M EDI V AL H OS P I TALS
E E NGLA ND T H or
, . o us a ar .
R h M y Cl y
ot a ar a .
,

E N GL ISH C H U RC H F U RNI TU RE J C C ox O E N GL ISH I N S T R UM EN TS OF M

LD us I t
F W G lpi
. . .

. S d E d it i
. a n. econ on.
I4 M ET H U E N A ND CO M A N Y LIMIT E D
P

Cla s si cs of Art — con ti nued

G HIR LA ND A IO G l d S D i S d . era . a v es . econ R B ENS Ed w d D ill

U . ar on . 3os . n e t.
E d i ti t
ons .

L AWREN C E S i W l A m g 5 t
r s. n e

r
.

a te r r s t r on s . ne
T IN T ORE TT O E l y M . ve n a r ch Ph i i ll pp s. 16
. . 2 .
net .

M I C HE L A N G ELO G ld S D i s . er a . a v e s. 1 s.
T I T I A N Ch l Ri k t 6 t ar es c et s. 1 s . ne

RAP H A E L A P O p pe 5
. .

T U RNER S S KE TC HES AN D D R AW IN G S A
i
'

. t . . . 1 s . ne .
. .

Fi nb e rg S econd E d t i on ne t .
R E M B R A ND T S ETC HIN G S A M H i d ’
. n .
. . 1 ss .

Tw V l m
. .

o s o u t es . a s . ne . VE LA ZQUE z . A . d e B e r u e te . x5s . ne t.

Th e Co mp l e t e ’
S erie s
F u lly I llu s tr a t ed . D emy 8vo

C O MP LE T E A MAT E U R B OXER T HE J G C O MPL E T E L AWN T ENNIS P LAY ER T H

Ly h A W lli M y
. . .
,

C O MPLE T E AS SO I AT ION F OO T B AL LER T H E

,
B ho un 6d t nc . 1 0s . . ne . . a F th E d i t i
s 6 e rs . ou r on . 1 2s .

CO MPL E T E M O T ORI ST T H E F i l
C
B S E
.

d C E H ghve r s D i an . . u es - a v es .
Y s on ou
W G A
. .

, .

r os . 6d t . ne d
an R i d E d iti s t on. ev s e o
C O MPL ET E A T H L E T I C T R A I NER T HE S A
.
. .

xos 6d t ne
M bi i
. . . . . .
,

us sa 6d n t r os . ne .
C O MPL E T E M O U N TA INEER T HE C
C O MPLET E B I LL I A RD P LAY ER T HE Ch l
. .

, . .

Ab h m S
Rb , . ar e s ra d E d i ti
a .6 t econ on . 1 s . ne .

o e r ts 6d
C O MPL E T E COOK T HE L i l i
. t 1 2s

Wh i l i g
. . ne .

C O MPL ET E OA RS MA N T HE R C L hm , . . . e an

C O MP LE T E P HO T OG R AP HER T H E R Chi
an t n
6d t
1 2s. ne
. .
, . .

C O MP LE T E C RI C KE T ER T HE Al b E
1 0s . 6d t . ne .

KNI G H T S d E d i ti
,

B yl y
. .

e rt
F ij i / E d i ti R i d
. .
,
a e I on , ev s e 1 2 s.
6d t
. .

econ on . 1 0s . . ne .

t
C O MPLE T E F OXH U N T ER T HE C h l R i h
.

ne
C O MPL E T E RUG Y F OO T B A LLER ON
.

,
. a r es c
N B THE
ZE ALA ND SY S T E M T HE D G ll h
,
ar d S
s on. d E d i ti é t
econ on . r s . ne .
a a er an
C O MPLE T E GO L FER T HE H y V d W J S d S
.
, .

, . arr ar on . . . d E d i ti
t ea m 6d . econ on . s. .

F ift th E d i t i
een R i d m 6d t on , ev s e . s. . ne .
CO MPL E T E S HO T T HE G T T d eas a
C O MPLE T E H OC KE Y P LAY ER T HE E , . . .

- u s t a ce B k ll T / i d E d i ti 6 t
I
, . uc e z r on. 1 s ne

F S h
. . .

C O MPLE T E H ORSE MA N T H E W S h
E Wh i
. S d E d i ti
te . 6d t econ on . 1 0s . . ne .
C O MPLE T E S WI MM ER T , H E. . ac s .

, . . car t 6d t . ne .

Di S
x on . d E d i ti 6decon on . 1 2s . . ne t .
C O MP LE T E Y T S MA N T HE B H k A CH ec s ta
T HE W H G d
.
, .

C O MPL E T E J J U I T S UA N , . . . a rr u . S mi h d l d t B l y S d E d iti
an u ou a . econ o
R evis ed . ne t .

Th e Co nn o i ss e ur
’
s Libra ry

Wi t lz nu mer ou s I llu str a t i on s . Wi d e R oy a l 8 va

'

. a ss . net ed e k volu me
E N GL ISH C O LO U RED B OOKS M i H . art n a r d ie . M A N U SC RI PT S J A
I L L UI I I N A T E D . . . H er b
e

E N GLIS F U RNI TU RE F S R b i
H . . . o nso n .

I ORIES Alf d M k ll
V . re as e .

ETC HI G S S i F W d
N . r . e rn o re . S econd E d i t i on .
J EW E LL ERY H C li ff d S mi h . . or t .

Ed i i t on .
EU RO E A N E N AM E LS
P . H e nr y H . Cu nyn g
M Cy i l D p
E ZZO T I N I S.
‘ ‘
a ve n or t .
b me
r

F INE
a .

B OOKS A W P ll d o ar
M INI ATU RES D d l y H h . u e ea t .

P OR C E LA IN E d w d D i ll
. . . .

G LA SS E d w d D ill ar o n.
. ar on.

S E AL S \V l d G y B i h
.

GO L DS M I T HS S LV S M I T HS W ORK ’ ' . a t er e ra rc .

AND I ER
W OOD SCULPTU RE A l f d M k ll
' .

N l D w e so n S d Ed i i a s on . econ t on . . re as e .
 G E N E R A L LIT E R ATU R E
Hand b oo ks of En gi sh Ch urch Hi st o ry l
E d i ted b y J . H . B U RN . Cr own 8va .
5 s . net eacfi v olu me

F O U ND AT IONS O F T H E E N GL ISH C H U RC H THE R EFOR MAT I ON P ERIOD T HE H y G e nr e e.

J H M a ud e S T R UGGLE WI T H P U RI TA NIS M T HE B
. .
, ,

. . .

SA XON C H U RC H AN D N OR MA N C ON QU EST THE

Bl l d
, . ru ce

T HE C T C w ll ,
ax a n

P APACY TH E
.

. . . ru t t e .

M EDI V AL C H U RC H AN D
E THE C H U R C H O F E N GLA ND I N E T HE I G H T E E N T II
A C J C EN TU RY T HE A l f d Pl mm
.
,
.
g . e nn i n s. , . re u e r.

Han d boo ks of T h eology

D emy 8va

D O CT RINE O F I NCA RN AT ON T HE R L THE I

’

I N T ROD U I ON CT H I S T OR Y O F To T HE TH E
O l y F if h E d i ti C REED S A N A E B
. . .
,
tt e .
5 t t on . 1 5 . ne .
, 6d . . . u rn . 1 2s . .

P HI LOSO P HY O F R E L I G ION I N E N GLA ND

net .

H IS T OR Y O F EA R LY C HRIS T I A N D OCT I NE A
I5 t
R A ND

J F B h Bk A M ERI CA T HE Alf d C l d
.
,

. . et u ne a er . s . ne .
, 6d . re a e cot t . 1 2s . .

net .

I N T ROD UCT ION T O H I ST OR Y O F R E L G ION T HE I X X X I X A R T I CL ES O F C H U RC H OF E THE NG

AN F B J T HE Ed i d b y E C S G ib
.

th
I
. S
. . e vons . even . te . . . so n .
ne t . E d i t i on .
5s . n et .

He al th Se rie s
2 s. 6d . n et

B A B Y T HE A h S d rt ur aun e r s. H OW T O L IVE LON G J W l C a te r a rr .

B ODY T HE F C
. . .
,

C A RE OF T HEgh a va n a P REVEN ION O F T C O MM ON C O L D T HE THE

0 K W i ll i m
. .
, .

C A RE OF T EE T H T HE A T Pi
, .

EY ES O F OU R C H I L DREN T HE N B i h p P LAGU E N B i h p H m
THE . . . t ts .
. . a s on .
,

s o STAY IN G THE s o ar an .

T HRO AT AN D E T RO U B L ES M l d
. .
.

H m
.
,

ar an AR
M IDD LE-AG ED S ym Y l y l i d Edi i
ac eo
H E A L T H FOR
. .

THE e o ur ea r s e r t on
l Thi d E d i ti
. . .

T a y or . r on.
T U B ER CUL OSIS Cli R i i
A W O MA N T HE R M y
H E AL T H O F
ve v e re
H EALT H
. .

or urr a
L li C HI LD T 0 H il
. .
,

es e. THE HE ton
S KIN T HE G g F t
, .

H E ALT H OF
. .

T HE ,
. e or e e r ne . S econd E d i t i on . as . ne t .

The Ho me Li fe S eri e s

I llu str a ted .

H O M E L IFE I N A M ERI CA K th . a e r i ne G . H O M E L IFE I N I TALY Li . na Dufi ’

G ord o n .

B b yus Se ed E d i ti . 6d con on . r es . . n e t. l i r d E d it i on . 1 z s. 6d . net .

H O M E L IFE I N C HIN A 1 T yl o H . . a r ea d a n d . l H O M E L IFE I N N OR W AY H K Da ni e l s

6d t
. . . .
r es. n e .

S e cond E d i t i on 6d
.

H O M E L IFE I N F R A N C E M i B h m . ss et a
. 1 2s . . n et .

Ed w d S i tl E d ti
ar 7 6d
s t
H O M E L IFE I N G ER MA N Y M A S id gwi k
. x z z on . s. . ne .
H O M E L IFE I N SPA IN . S L . . B e ns usa n .

. rs . . c .
S econ d E d i t i on . 1 25 . 6d . ne t .
Tb i d d ti
r 6d t on . 1 2. ne

B AL K A N H O M E L IFE L
. .

H O M E L IFE I N H O LL ND D S M l d m A . . . e ru . . u cy M J G . . a r ne t t .

S econd E d i t i on . 1 2s . 6d . net . r e s. 6d . ne t .
 M ET H U E N A ND C M O PA N Y LIMIT E D

Le ad e r s of l
Re i gi o n
E d i te d b y H . C . B EE CH I N G . Wi th Por tr a i ts

Cr own 8 vo .
3s . net each volu me
A UGU S T I NE OF C A N T ER B U RY E L C . . . u t ts . J OHN KE L E W lt L k S th E d i ti
E . a er oc . even o

B ISHO P B UTL ER W A S p . . . oo n e r . J OHN K NOX F M C S .d E d i ti . ac u nn . econ o

B ISHO P W I L B ERFOR CE G W D i ll . . . an e .
J OHN W ES LE Y J H O t . . . ve r on.

C A RDIN AL M A NNIN G A W H S . . . u t t on . econd L A N C E LO T A NDREWES R L O t l y S

E d i t i on
. . . t e . eco
.

C RDIN AL N E W MA N R H i H
A . . u tt on . L AT I M ER R M d A J C l yl
. . . an . . ar e.

C H A R LES S I M EON H C G M l . . . . ou e .
T HO MA S C H ALM ER S M O l i ph . r s. a n t. S eco
E d zt i on

T H O MA S C RA N M ER A J M
.

G EORG E F T HE QUA KER T H

ox , . . od gk i n .

. - . . as on.

J OHN D ONNE A g t J u us u s e s so p T HO MA S K EN F A Cl k . . . ar e.

W I LL I AM L AU D W H H
. .

u t t on. F ou r
J OHN H OW E R F H t
. . .

. . . or on.

Th e Libra ry of De v o t i on
With I n tr od uctions a nd ( wh ere necessa ry ) N ot es

S ma ll Pot t 8 vo, cloth , 3s . n et ;a lso me volu mes i n lea t her ,

so

3s 6 d . . net each volu me

B ISHO P W I L SON S SAC R A P R I V ATA

’
L I G H T L I FE a nd LOVE A S l e e c t io n fr c
My i
.
.
, ,
th e G ma n
B OOK OF D EVO T IONS A S d E d i ti econ on .
er s t cs .

L I TT LE B OOK OF H E AVEN LY W ISDO M

.
,
'

C HRIS T I A N Y E A T F if th E d i t i
R, H E. on. A S l i f m th E gli h M y i
e e ct on ro e n s s t cs.
,

C ONFESSIONS OF S T A UGU S T INE T HE .

, . L Y R A A POS T O L I CA .

N i n t h E d i t i on 3s 6d ne t
LY R A I
. . . .

Thi d E d i t i
N N O CE N T I UM . r on .
DA Y B OOK FRO M T HE SA IN T S A ND F AT HERS
A .
,
LY R A SAC R A A B k f S d V . oo o a cr e e ra

D E AT H A ND I MM OR TAL I TY
M A N UAL OF C ONSO LAT ION FRO M T HE S A I N
.

D EVO T IONS FRO M T HE A P OC RYP H A .

A ND F AT HERS A , .

D EVO T IONS OF ST A NSE LM T HE .

, .
O T HE LOVE OF G OD
N .

D EVO T IONS FOR E VER Y D IN T HE WEEK AY

P RE C ES P
A ND T HE G R AT F ES T IV ALS
E
R I VA T A E .

G R AC E AB O U NDIN G T HE C HIEF OF S IN P S ALM S OF D AVID T HE

.

.
,
To
NERS S ERIO U S C AL L A D EVO UT A ND H O To
L IFE A F ift h E d i t i
.

G U IDE T O ET ERNI TY A , . on .

S ON G OF S ON G S T HE
.
,

H OR A E M ADyB k f m h
S P IRI TUA L C O M B AT T HE
.
,
Y S T I CA E a oo ro t e
W i i g f My i f M y N i
.

r t n s o s t cs o an a t ons .
, .

I M I TAT ION OF C HRIS T T HE E igh th d i t i , . E on . SP I RI TUAL G U IDE T HE Th i d E d it i , . r on .

I NNER WAY T HE Thi d E d i ti

, . r on . T E MPLE T HE S d E d i t i
, . e con on .

i T HE D EVO UT L
N T R O D UCT I O N T o AN I FE , . T HO UG H T S OF P A S CAL T HE S d E d i ti , . econ o
 M ET H U E N A ND CO M P A N Y LIMIT E D
Th e Li t tle Gui d es— con ti n ued

B E D RO RD S H I RE AN D HU N T IN G DONSHIRE H N OR T H AMPT ONSHIRE W D y S

W r eco
. . . .

M a ck lm
.

. .

B E R K S H I RF . F G . . B ra b a nt . N OR T H UM B ER LA ND J E M i . . . orr s.

S R
N O TT IN G H AM SHIRE L G il f d
ne t
B UC K I N G I I A M S H I R E . E . . oscoe . S econd
.

E d i t i on , R evi s ed .
. . u or .

C AM B RIDG ESHIRE J C Cox O XFORDSHIRE F G B b S d E d i t i ant

S HRO P SHIRE J E A d S d E i i
. . . . . . . ra . econ e

C HESHIRE W M Ga lli ch a n t

S O M ERSE T G W d J H W d F
. . . . . . . u e n. econ d e

CORN WAL L A L . . . S lm
a on . S econd E d i t i on . . . . an . . a e. ou ;

D R BY SHIRE J C C S d E d i t i
S TA FFORDSHIRE C M fi ld S d E
E . . . ox . econ on .

D EVON S B i g G l d F t h E d i ti
. . ar n - ou . ou r on .
. . ase e . econ 4

D ORSE T F R H h F t h E d i ti S U FFO L K W A D S d E d i ti
D U R H AM J E H d gki
. . . ea t . ou r on .

. . . utt . e con on .
. . . o n.
SU RREY J C C Th i d E d i t i ox . r on ,
E SSEX J C C S d E d i ti
. . .

ox econ on .

SU SSEX F G B b
. . . .

G LO UC ES T ERSHIRE J C C S d . . . ox . econ . F if t/ E d i t i . . ra a nt . f
z on.

WA R W I C KSHIRE J C C ox
HAMPSHIRE J C C Thi d E d i t i
. . .

.
~
. . ox . r on .

W I LT SH I RE F R H h Th i d E d i t i ea t r c
H EREFORDSHIRE G W d J H W d Y OR SHIRE T HE EA S T R IDIN G J
. . . .

. . . an . . a e.
K
H ER T FORDSHIRE H W T mpk i M
, . .

Y ORKSHIRE T HE N OR T H R IDIN G J
. . . o ns. orri s .

K EN T J C C S d E d it i R
. . . ox . econ on , e , . .

w i tt
r en .
M i orr s .

K ERR Y C P ( b. S d E d it i
. . a ne . econ on . Y ORKSHIRE T HE W ES T R I DIN G J
M i 5 t , . .

L EI C ES T ERSHIRE AN D RUTLA ND A H y a r ve or r s . 9 . ne

d V B C w h
.
. .

an . B y . ro t er e n on .

L IN C O L NSHIRE J C C ox.
B RI TTA N Y S B i g G l d
. . .

M J B Fi h ar n - ou S econd E d z t i c

M ON M O UT H SHIRE G W d J H W d
. . .
I D D LE S E x . rt

N OR MA ND Y C S d m
. . .

. . cu a or e. S econd E d i t i
. . . an . . a e.

N ORFO L K W A D F h E d i ti ut t . ou r t on ,
R O M E C G E ll b y
. . . a .

SI IL F H J k
. . .

R evi s ed . C
'

Y. . . ac s o n.

The Li tt e Lib r ar y l
Wi th I ntrod uct on i N , otes , a n d Photo g ra vu re F ronti spi e ces

Sn
z a ll F all 871 0 . E a ch Volu me, clot h , 25 6d net ;also . . so me volu mes
i n lea t her a t 35 6d n et . .

A n on L I TTL E B O OK O F E N GL I SH
“

A l SE L E CT I O N S FR O
B la k e ( Wi li a m) .
LY R I C S S d E d i ti 3 6d t WORKS O F W I LL IAM BL A K
.

. econ on . 5 . . ne . TH E
A t n (J n )
us e P R ID E A N D PR EJ U
a e
B wn i n gmb t ) SE L E C T I O N S F R O
DI C E Tw V l m
.
ro o er
T H E E A R LY P OE M S O F RO B E
.
o o u es .

N O R T H A N G E R A BB E Y
.

B RO W NIN G
ESS A Y S O F
.
.

B n (F n i ) TH E ra
SE L E C T I O N S F R O
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MM AN U E L UR
S d E d i ti C . econ on . I

B ei loc ( IL) E B DE , N 80 0 75 . ne t .

M E RC H AN I ll
.

d T . u s tra t e . S econd E d i
t i on Cr ne t .
W I I LL M A I N TA I N
B owen ( Ma rj or i e)
i
. .
75 .
.

N i nt h Ed t i on Cr 8m . . .
7 5 ne t. .

B enn et t ( Ar n o d ) l . C L A Y H A N GE R .
D EF E N D ER OF TH E F AITH . S even t
T w elf t h E d i t i on . Cr . 800 . 85 . ne t.

H I LD A LE S S WA YS . E ig hth E d i t i on . W I LL I A M BY T H E , G RAC E OF GO D
S econ d E d i t ion Cr 800 . . .
75 . ne t .
26 M E T H UE N A ND CO M PA N Y L IM ITED

F E LI X
E d i ti
HR Y
C 8 7 t A LIF
: T EE
‘
E A RS I N E. S ev en t h J aco s ( b .
‘

Th i r ty -t hi r d E d i t i on
w M AN Y CA R GO E
on . r . vo. s . ne .
. Cr .

T H E WO M AN W I T H TH E F AN
C
A lso Cr

U RC H I N S
80a 6d
C
23 , ne t.
E igh t h
. . .

E d i t i on . r . 8 vo .
~
7s . n et . SEA . N i n eteent h E d i t ion .

BY E WA Y S Cr 7 3 . n e t.
N

NI llO
. .

TH E GA R D E
' '

i t/ E d i t i
s x i
M A S
on .
TE O F CR A F T I ll u t
A LL A H
F
us tr a t e d
Tw en ty
Cr 80 0 85 6d
.
A . s ra t e
El t h E i ti C 8 5 t
. . . . .
even an . r. 00 . 5 . ne .

L I G H T F R E I G H T S I ll d F ift
T H E C A LL O F T H E B LOO D
t . us tr a te . ee n
N i th E d i ti C B 5 t . n on . r. oo . 5 . ne .

T H E S K I PPE R S WOO I N G
E d i ti C 8 8 6d
on . t r. vo. 3 . . ne .
'

B A R B ARY S H E E P S o n
d Ed i ti
c t/I
t ‘
. u n

A T S U N WI C H POR T I ll
C E d i ti C 8 t . ec on. r. on . r . vo. ne .

8 vo. 6 t 5 . ne .

t d El t . u s t ra e . even

T H E D WE LL E R ON TH ET H R E S H OL D
D I A LSTON E LA N E
.

I ll t d E igh t
'

C 8 r .
7 t 00. 3. ne .
. u s tr a e .

T H E WAY O F A M B I T
i
t on. C 8 t r. OD D CR A FT
00 I ll t d F if th E d i t i ne . u s ra t e .
'

I N T H E W I L D E R N ESS
. .

C 8 t r. 00 . ne . o h ! I

T H E L A D Y O F TH E B A R GE
Thi d E d i ti
I ll
'
r on . ‘

t t us ra e
C 8 t
.
r. 7 vo 5 . ne .

Tent h E d i t i on
.

. Cr 800 . . net .

C H AN GE AI R
l
H op e A S A LT H A V E N F ou rt h E d i t i w
( An th ofly) OF .
. .

S i x th E d i t i on Cr Boo ne t
Cr . 8 00 . ne t .

S A I LORS K N OTS
. .

I ll
. .
_ 1

O F M AR K
’
u s t ra ted . S ix t
A M AN S event h E d i t i on Cr
.

S H ORT CR U I S E S
. . .

C
C H RO N I CL E S O F CO U NT A N
. Th i r d E d i t ion .
‘

TH E Boo net .
O N I O S i th Ed iti
.

T C 8
.
7 x on . r. 00. 3.
“
L IF TED VE I L
[

t
ne .
Ki n! (B u ll) TH E C
PH ROSO I ll t d N i t h E d i ti n C
.
.

. us tr a e . n o . r. 8000 75 . 1

S I M O N D A L E I ll t d mm 5 d . u s t ra e .
° L b
e th
H RO
dg
E
ri e
Cr
( S y il
Boo
b ne t .
ONE WO M AN ’

C 8 t
. . .
r. 7 00. 5 . ne .

TH E KI NG S MI RROR
C Lon d on ( J a ck ) . WH I TE F AN G
’
.
. N i nt
r.

LO D G E
I A LOGU E S
net .
Lown d es ( Mu . Belloc) TH E .
”
T H E D O L LY D Cr 800 Th ir d E d i t ion Cr 300 .
73 .
. .
. . .

L I S TE N E R S L U R E A
I I ' "

T WO PE O PL E
' I
4 l

TA L ES
C O B L I QU N A RR A T IO N Tw lfth E di ti w
Luca s ( E.
'

OF . Thi r d E di :
E

S E RV AN T O F T H E PU B L I C I ll
t i on 80 0 e
7 3 . net .
.
. r . .

F p 8 ca t 00. ne
A
. .

OV E R B E ME R TON S A E AS COI N
CH RO N I C L S i t th E d i t i
. us
th E d i t i
’

d F C : N Y-
tr a t e . t ou r on . r. ne .

F
M RS M AX O N PRO TE S T S Thi d E d i
. . r 8 00. 6 t 5 . ne
E .

.
x een on. ca

A YO U N G MAN S Y E A R S ’
M R I NG L ES I D E Thi t th Ed t i
. . r een i o
d E d iti . econ on . F t B ea t. oo. ne .

Cr 800. .
7 3 ne t . .

LO N D O N L A VE N D E R Tw l fi h E d i t i . e o
I p F eat 800 ne t .
M R H ORROC K S
U RS
H yne ( C J
. .

R
Gu tcli ffe )
i L AN M AR K S
. . . .
,
P E F ift h E d t i on Cr SW D C 8 . F if th E d i ti on '
r 9
73 .
. .
. . .

t ne

C T H E V E R MI L I O N B OX
ne t .
.

FI R E M E N F th H OT ou r E d i t i on . r.
C SW r. t
F if t h E d i t i
ne
. o

8 t
. .
71 0. s ne

C A PT A I N K ETT L E O N
7

Ly ll ( E d ) D E RR I C K V AU G H A N
. .

WA R
PA T H Th i d E d i t i C . r on . r
TH E
8v o 7 s
. . . ne t.
a
N O V E L I S T 44 t h Th d C
na
8m
.

ousa n r.
R E D H E RR I N GS
. .

(3 8 . 7 1 00. 6: . net . 53 . ne t .
 F I CT I O N 27

McKe nn a ( Bte p l e n) SO N I A B E WE E T N TH E V A N I S H E D M E SS E N GE RJ l S eamid

W O R L D S gi t
:
C
.

5"T WO . x eent h E d i tion . r. 89 0 . E d i t i on . Cr . 8w . not .

TH E H I LL M AN . Cr . 8vo .
7t . h ot . .

Ox enh a m ( J oh n) A W E AV E R OF
W S I ll
.

EB d F ift h E d i t i
. C u st ra te . on . r .

AN D L O S S S i th E d i t i
8 0 0. t ne
PR O FI T
.

- r x on

R AN D J AN E
.
’

Macna
H E SO NG O F H YA C LN T H AN D OTH R
ta n PE T E C B
r. t oo. ne
C
.

E d i ti on . r. 80 0 . ne t .

E
S T O RI S
,

S d E d it i E C 8 on
L ORY S I R 7 econ r 00. 5.
HI
. . .

Ha let ( 1 HE
'

u ca s ).
ST OF ne t
R C HAR A R O M AN C
.

i
D CA L M A D Y
- I

S event h E d i t on Cr 800 no t
E .

L A U R I S TO N S .
‘
F ou r t h E i t i on d . b y. ao o .

WAG S
. . . .

n et .
TH E E OF S I N S i x tee nt h E d i t ion
CO I L O F C A R N E S i th E d i t i n
. .

TH E . x o .

TH E C A R I SS IM A F ift h Ed i t i
T H E QU E S T O F T H E G OL D E N RO S E
Cr 800
'

ne t .
. on . co .
. .

T H E GA TE L E S S B A RR I E R
.

'
F if t h Ed i C 8 m
M A RY A LL A LO N E Thi d E d ti C
. F t h E d i ti
ou r t on . r . 7 . ne .

ti on Cr 800 ne t .
1

. .
-
.

. r z on. 7 .

8 t
CLEM E N T I NA ne
00.

B RO K E N S HA C K L E S
.

Ma s on
'

(A E
F t h E d i ti
I ll
. . . '

ou r on
us tra t e d . N i nt h E d i ti on . Cr . Boo .
. .

I l
1914 . T hi r d E d i ti on . Cr . goo . ne t .

P l I R
TH E
ti a n
U
G A RD E D F LA M E
Cr 8 00 net .
ne t .

. S even t h Edi
a r k er
PE
t
ne
O PLE
.
( Gi b ert )
.
P E RE .

S event h E d i ti on .
A ND H IS

L N S S d E d i ti C
. .

M R S F A LC HIO N
.

ODD E GT H . econ on . r. 8m . . . F iflh E d i t i on . Cr .

t ne Boo net
H I LL R I S E F th E di t i
. . .

C 8 ou r on .
R AN SL AT I O N SA V A OE
'

r. 00.
TH E
.

OF
’
T A

TH E R E S T CU RE
.

F th E d i t i ou r on. Cr
R AI L H WOR I ll
. .

TH E T OF T E S D . us
tr a t ed Tent h E d i ti on Cr Spo ne t .

AY L AY
. . . .

Mi lne ( l L ). S P S ix t h
WH E N VA L M O ND C A M E T O PO N T I A C
’
. TH E D .

E d i t i on Cr Boo net
ST O RY O F A LOS T N A PO L ON S th
O N CE A iTHE
. . . .

WE E ct soo K ne t
E . even

AN AD VE N T U R E R O F T H E N OR T H
. . . .
C 8
E d t i on t . r. 00. ne .

AL S M E AN L A S T A D V N T U R S O F PR TT
( Ar th ur ). T
R
E OF
i
‘
T
ST R S EET S event h E d i t i on
. .
PE
HE
I R E.
’
M h Ed ti on
E
.
E
Cr 800 . .
E
ne t
Y
.

ne t
A C H I LD O F T H E J AG O S i th Ed i ti A S OF MI G H T Y I ll
.

. x on. TH E S E T TH E . us
tr a t ed Twent i e th E d i t i on
. . Cr . 89 0 .

T H E H OL E I N T H E W A LL
ne t
th
.

F ou r
ATT L E
O F T H E S T RO NG A
.

TH E B :
D I V E RS V AN I T I ES C B t . r. oo. no .
R O M AN C O F Tw K I N G D O M S I ll d E o . u s t ra t e .

S event h E d i t i on . Cr . 8 00 . ne t .

Op p h i m ( E Ph i ll i p )
en e M AS TE R O F . s . .
THE P P OF T H E
'
OM L A V I L E TT E S .

M EN F if t h E d i t i C 8 7 t on. r. 00. 3 . ne T h i r d E d i t i on Cr 871 0 ne t

T H E MI S S I NG D E LOR A
. .

I ll
. . . .

d
N OR T H E R N
us tr a t e
. .

I GH TS
[ H E D O U B L E L I FE O F M R A L F R E D
L . F ou r t h E d i t i on .

B U RT O N
Cr . 800 . ne t .

S d E di t i C 8
C H AR M
. econ on . r. 00.
Perr ln ( Ali ce) TH E
'

net . . F if t h
A PE O PL E S M AN Thi d E d i t i
.

’
C . r on . r. E d i t i on . C7 . 8 00 . ne t .

8 00. t ne

C I LD R E NO F T H E
.

M R G R E X O F M O N T E C A RLO Th i d Ph i ll ott s ( Ed e n) H
g
. . . r .

E d i ti on . Cr 800 . . ne t. M I T S i x th E d i ti on . . Cr 800 . . n et .
28 M ET H U E N A ND CO M PA NY L I M I T ED

TH E H B OY
S even t h E d t i on
U M AN
Wit h a F ron tis ie ce
Cr Boo i
. p . W lle
Cr
s
800
(H . GI) B E AL BY . F m} E d i ti
; on .

ne t . . . . . .
7s . ne t .

SO N S O F TH E M OR N I N G . Second Edi
Wi l li a mson ( 0 N a nd
1

K BL ) . TH E
t i on Cr 82m 7 3 ne t.
L I G H T N I NG C O NDU C T OR
S T RAN A E N T R ES
. . .
. . . .

: TEE
A Mb
t
I llus t r at ed
GE
.
DV
T wen y -second t
U or
Ed i i on tTO
.

A M ER I CA N R I SO N E R ' 800 7 s ne
. . .

TH E P F ou r t h
PR N C S S P A SS ES
M OTO R I ll st rat ed N t
.
'

TH E I d E A R u A Ncn

U
:

DE M ETE R S ’
D A G HTE R Thi r d Ed i
OF A
Cr 8 00
.

ne t
u . i n h E d i t i on .

i on
.

LA D Y B Y A C RO SS H W A T E R
. . .

t Cr Boo n et .

U M AN
. . .
73 .
’

E TT T E
TH E H B O Y A N D TH E WA R
.

C
.

SC A RL ET R U N N E R I ll
Thi r d E d ati on . r 800 7 3 ne t
. . . .

. us t ra t e d .

R d ge ( w. Pott)
i A S ON O F T H E
S A LORD LOVE LA N D D ISC OV ERS
.

T TE Th i r d E d i ti on Cr 800 75
A M ER I C A I l l u s t ra t ed S d E d i ti
. . . . .

econ on

R E MI NG T O N S E N TEN C E
. .

TH E Thi r d Cr 8 220
.
73 ne t

S I LE N CE
. . .
.

Il l u t at d
M AD AM E PR I N C E S W E d i t i on Cr
T H E GO L D E N
E iglz t lz E d i i on t Cr am
.

7:
s r e

R L
. . .

C
. .

G U E S TS
.

OF H E ES I ll us
TOP S PE E D . S econd E a dm . Cf . Boo .
TH E
t at
r ad . F ou r t /z E d i i on t . Cr
U
. 8 00 .
.

7s . ne t

R F OR M A N C ES
'

u rate d
'

I T H A PPE N E D I N E G YPT I ll
'

S PE C I AL PE Cr .
'

8w .
S even h E d i i ontCr 80 0 t .

net .
st

A SOLD IE R
. . .

US L I N G H O U RS
net .

O F T H E L E GI O N S eca m
‘

TH E B T . Cr . Soo .
73 .
.

TH E S H O P GI RL A C 8 7 t
L I G H TNI N G CON D U C T R E SS
. r. vo . s . ne .

TH E
Thi r d E d i t i on Cr 800 73 ne t .
S I TA N M YS T E R I E S
. . . .

TH E S eao nd
t i on . Cr . 8 00 .
7 5 . net
. .

S ECR E T H I S T ORY C 8 . r. 00. 75 . ne t .

O F S E CR E T E GYPT T H E LO VE PI R A TE I ll u s tr a t ed . Th i n
' ‘
I LE S
g t
I . Cr . 800 .
.

s .

T H E ORC H A RD OF T E ARS
CR U C IFI X COR N E R C 8 6 t
. . Cr . 800 .

. r. 00 5 . ne .

Swi Iin or t oE( m OU S M A R T I N SCH U LER

'

S H O PS A N D H Wi l n
‘

SE . so
Cr 800. .
, 7 s ne .

B ook s for Boy s and Gi r ls

TT I N G W LL O F D ORO T
1

G H Y, THE Mr s M AS T R R E V OY AG OC K A F E I I A R S
'
E. W Cl ar

R ll
E
K C l iff d 6 t
E . . . . .

GR AN G T
. or . 3 . ne . u ss e .

M
G RL OF A L M R ED M I E, H E. r s. O C SW OI l h)
'

ON C A PR I NC
P T ea d e.
TH R
T HE E O PI E ,
I
M
HONO U R A B L M I SS h
. . .

EW E AS E E M rs
L M
.

M
. .

T e a d e.
'

E t i E . . . a nn .
,
.30
M ET H U E N A ND CO M PA N Y L I M I TED

Meth uen l Ch ea p Nov els— ’

co nt i nued .

JA N M a i e C rel l i o N E S T HIR T Y W P e tt R i d ge
N E To IX -

OC E A N S L E UT H T HE
E r

J OH A NN A B M C ok er
. . .

M aurice D rak e
.

J OSE P H F k D anb y S I L VER M yr tl e R eed

. . . r . , . .

ra n
O LD R OSE AN D . .

P AT HS OF T HE P R U DEN T T HE J S Fl h
. .

J SH UA D A IDS ON C O MMU NIS T E Ly

O V nn
P AT H W AY OF T HE P IO EER T HE D
. . . . e tc e

L n to n
. .
,

J OSS T HE Ri h d M a sh
i N , . o
Wyll d
.

ar e.
c ar r
PEGG Y OF T HE B AR T ONS B M Crok er
K INS MA N T HE M Alf ed S id gwick
. .
,
. . . .

P EO P LE S M A N A E P h i ll ips Oppen h eim

'

K NI G H T O S PA IN A M a j ori e B owe n
,
. rs . r . ’
.
, .

PE T ER A ND JA NE S M
LAaDnYd AB EMTTYWAillCiROSS T HE W AT ER C N
F r
gh ta n
.
, .
. . a cn a u .

P O MP T HE L OR T HE S ir Gi lb e A V I LE T T ES ,
amso n P ark e
. . .
.

L ALAG E s LO ERS G eor ge A B irm i n gh am G LOR Y T H E M rj ori e B owe n

. . .
r.

QU E T
LAwicN TkE B EA R RS T HE M Alfr d S id g
’ '
V a

QU ES T OF T HE GO LDEN R OSE T HE J l
. . .
S or , . .

RN E , . rs . e , . o

R EG E T T H E A rn ol d B e nn ett
. 0 xen h m a .

John O h m
L AVENDER O L ACE M yr tl e R eed RERMi dINgGTe ON S EN T EN CE T W P
LA U R I S T ON S . xe n a .
N , . .

HE e
A ND , .
LD
.

REST CU RE T HE W B M a well
. .

L I G H T F REIG H T S W W Ja ob s c
.

REBTUu rou
. . . .

x
L ODG ER T M rs B e l l o Low n des
. . . .
,

RN OF T A R A N T HE Ed gar Ri
LON G R O AD T HE Joh O h m
H E. c
Z
. .
,

gh s
.
,
'

LOVE LO U IS A E M aria Alb an i RO U ND T H E R ED L AMP S i A Co an B yl

. n xe n a .
r .
,

LOVE R OYA L GEORG IE S B aring-G ld

AND es r n o

P IR AT E T HE C N an d A M
. .
. . .

S ATD T HE F ISHER MA N M arm d k e P ic

"
. . ou .

W l l iam s on
. . . . .
,

M A R Y A LL A LONE Joh n O h m
i .
. a u
,

t h n

M A S T ER OF T HE V INEY A RD M y tl R ed
- a

S AL L Y D o oth ea C o n yer s
. x en a .
.

M A S T ER S V IO L I T HE M y tl e R e d
r

S AL IN G OF A D ERE L I CT T HE M aur i
. r e e . .
. .

M A ;C
'
V
N, r e
D rak e
.
. . ,

) Er n est B amah
A R R A DO S r
.

S A ND Y M A RRIED D oro th ea Co n ye s
M AY OR OF T RO Y T HE
. .

S E A C APTA IN T HE H C B ai l ey
r
Q
. .
”
‘
. .
,
’

M ESS D E C T W F S ha nn o
. . .
, .

S E A L A DY T H G W ll s
M IG H TY ATO M T H E Mari e Core l li
K, HE . . . n.
HE. e

S E A R C H P A R TY T
. . .
,

g A Bi mi gh
'
G HE
M IR AG E E T emp l e T h sto n S EC RE T AG E T T ) Jos ep h Co n rad :
, . .
, . e or e . r n a

ur N HE

S E C RE T H IS T OR Y C N an d A M Wi ll ia
. . . .
,

M ISSIN G D E L OR A T HE E Ph i ll ips O ppe n

h eim
. . . . .

son
. . .
,

M R G OF M ON T E C A R L O E Ph i ll ips
.

S EC RE T W O MA N T HE Ede n P h i l l potts
.

S ET IN S I L VER C N an d A M W il l iar
'

RE x
.
. . , .

O ppe n h eim
.

M R W A H IN GTO M aijerk;Bowen
. . . . . .

- s on
M f M A ON P
.

S N.
S EV A ST O PO L A ND O T HER S T ORIES L
. .

: A n th o n y H ope
T o l s t oy
, . 4
Rs X R O T ES T s

S EVERINS TH E M A l fr d S id g wick
.

M S P E T ER H wm : M a y E M a nn
.

o n r

W Cl a k
R . . .

rs. e
M Y D A NISH S W EE T H EA R T S HOR T C R U ISES W W Jacob s
.
, .

r
Ru s e ll
. .

MY F RIEND T HE C H AU FFE U R C N S I F M S ERI ES T HE S R h mé

' . . . .

s .

- AN Y T ax o r
d
.
, .

an
A M W i ll i mso n S PA NISH GOLD G eorge A B irmi n gh am
. . .

a . . . .

S P INNER IN T HE S U N A M yrtl e Reed

. .

MY H L eo T ol s t oy
US BA N D A N D . . , . .

M Y L A D Y OF S H A DO WS Jo hn O h m xe n a S T REE T CALL ED S T R A I G H T T HE B , . ac

M YS T ER Y OF D F M A N C H U T HE S King
. .

S UP R M E C RI M E TH E D oro th ea G er ard
’

U-
.

R. , . ax ‘

R oh mcr E

T AL ES OF M E A N S T REE T S A r h M i
.
, . .

MY S T ER Y OF T HE G REEN HEART , T HE . . t ur or r s o
M a x Pe mb er t on
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P RIN T ED BY M O ISON
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 1 4 DAY
RET URN TO D ESK FR O M

EA RT H S C I EN
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