Devonian Trilobites From the Shogr m Formation Kur Gh Chitral NW Pakistan

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Alcheringa: An Australasian Journal of Palaeontology

ISSN: 0311-5518 (Print) 1752-0754 (Online) Journal homepage: http://www.tandfonline.com/loi/talc20

Devonian trilobites from the Shogrām Formation,Kurāgh, Chitral (NW Pakistan)

Raimund Feist , John A. Talent & Ruth Mawson

To cite this article: Raimund Feist , John A. Talent & Ruth Mawson (2001) Devonian trilobites

from the Shogrām Formation, Kurāgh, Chitral (NW Pakistan), Alcheringa: An Australasian Journal of Palaeontology, 25:1, 107-115, DOI: 10.1080/03115510108619216

To link to this article: http://dx.doi.org/10.1080/03115510108619216

Published online: 27 Nov 2008.

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D evonian trilobites from the Shogr m Form ation,

Kur gh, C hitral N W Pak istan)

R A I M U N D F E IS T , J O H N A . T A L E N T A N D R U T H M A W S O N

FEIST R . TALENT .A., MA wsos, R., 2001:09:15. De vonian trilobites from the Shog r]m

Formation, KurAgh, Chitral (NW Pakistan). Alcheringa 25, 107-115 . ISSN 0311-5518.

Late D evonian trilobites from horizons close to the Frasnian-Famennian boundary in

the S hogram Formation at K ur~gh, Ch itral (NW Pakistan) are described. A new species of

Asteropyginae,Neocalmo nia chitralensis sp. nov ., and a new subpecies,Neocalm onia batillifera

orientalis subsp, nov., are described; hese exten d he range of Asteropyginae eastwards from

Iran and southern Afghanistan. The Upp er Kellwasser Event is located within KUR 19 o f

Talent et aL (1999).

R. Feist*, lnstitut des Sciences de l Evolution, U MR 5554 du C NRS , U niversitd de Mo ntpellier

I I, 3 4 0 9 5 M o n tp e l l i e r C ed ex 0 5, F ra n ce ; J .A . Ta len t a n d R . Ma w so n , C en t re f o r

Ecostratigraphy and Palaeobiology (MUCEP), Department of Earth and Planetary Sciences,

Macquarie University 2109, Australia; rece ived 1 June, 2001; revis ed 10 July, 2001;

*corresponding author; [e-mail: [email protected]].

Keywords: T rilobita, Asteropyginae, La te Devonian, Frasnian, C hitrai, Pakistan.

T H E S H O G R A M F O R M A T I O N ( D e v o n i a n ,

G i v e t i a n - F a m e n n i a n ) a n d a s s o c i a t e d o l d e r

M iddle Pa laeoz oic uni ts----Chanm Qua rtzi te , Lu n

S h a le , C h i lm a r a b a d F o r m a t i o n - - e x t e n d a c ro s s

Chi t ra l and ad jacent nor thwe s te rn Gi lg i t f rom the

u p p e r K a r a m b a r v a l l e y a n d Y a r k h u n v a l le y s i n

t h e n o r t h e a s t t o t h e B f i n i - K u r a g h - S h o g r ~ m -

Banam a r ea and the Lu tk o va l l ey some 200 km to

the sou thw es t ( s ee Ta l en t e t a l . 1999 , f o r su rvey

o f e a r l i e r l i te r a tu r e ), a n d c o n c e i v a b l y b e y o n d

(Bu chro i thner 1998).

Th e t r i lob i te mater ia l descr ib ed in th is repor t

c o m e s f r o m e l e g a n t e x p o s u r e s o f t h e S h o g r ~ m

Form a t ion - - an app rox ima te ly 230 -m- th i ck sha l e -

l imes ton e -a r en i t e succes sion -- - -on t he sou thwes t

f l ank o f t he Ku r~gh Spur , Ch i t r a l (F ig s 1 , 2 ;

Ha yde n 1915 ; Ta l en t M aws on 1979 ; Ta l en t e t

a l . 1982; 1999) . Macrofo ss i l s der iv ed f rom these

exposu re s , bu t co l l e c t ed f rom sc r ee be low the i n

s i t u sequenc e (e.g ., Des io 1966) , fo rm ed the bas i s

o f s eve ra l repo r t s on mac ro fos s i l s ( summ ary i n

Ta l en t

et a l .

1999), pr incipal ly rugose and tabulate

c o r a l s a n d b r a c h i o p o d s , b u t in c l u d i n g a n

as t e ropyg ine t ri l ob it e (Reed 1922). T he p r e sence

0311/5518/2001/010107-09

3 . 0 0 ©

AAP

o f f ive long la te ra l margina l sp ines on a pygid iu m

desc r ibed by Reed a s a new spec i e s,

D a l m a n i t e s

( A s t e r o p y g e ) k o r a g h e n s i s , sugges t s i t cou ld be

a s s igned t o H e l i o p y g e Haas M ens ink , 1969.

C o l l e ct in g o f m a c r o f a u n a a n d s a m p l in g f o r

m i c r o f a u n a f r o m t h e i n s i t u s e q u e n c e b y T a l e n t

and col leagu es in 1973 an d 1975, and for t r i lobi tes

by Fe i s t in 1999 , ha s cons ide r ab ly en l a rged t he

macrofoss i l da tabase , inc luding t r i lob i tes ( th i s

r e p o r t ) , a n d h a s p r o v i d e d a b i o c h r o n o l o g i c

f r a m e w o r k , b a s e d o n c o n o d o n t d a t a , a g a i n s t

wh ich t he r anges o f the v a r i ous mac ro fos s i l taxa ,

i nc lud ing t he t r il ob i t e s de sc r i bed he r e in , c an be

a s s e s s e d .

ges

C o n o d o n t d a t a f r o m U n i t K U R 9 , a 1 5 m s h al e

in te rva l (c f . F ig . 2 ) , a re sparse (Ta len t e t a l . 1999;

Tab l e 1 ) bu t i t appea r s t he un i t r ep r e sen t s a

s u b s t a n ti a l p a r t o f F r a s n i a n t i m e . T h e n e w

subspec i e s

N e o c a l m o n i a b a t i l l if e r a o r ie n t a li s

is f r om K U R 9 B . T h e u n d e r ly i n g K U R 9 A h a s

b e e n s u g g e s t e d t o b e c l o s e t o t h e G i v e t i a n -

F ra sn i an bounda ry and KU R 9E , some 8 m h ighe r ,

D o w

n l o a d e d b y [ U n i v e r s i t y o f t h e

P u n j a b ] a t 2 1 : 1 2 2 2 F e b r u a r y

2 0 1 6


3/10

10 8 R . F E I S T , J. A . T A L E N T A N D R .. M A W S O N AL C H ERI N G A

. . . , - ~ R i v e r s

V e h i c l e t r a c k

• . - . . .. . F o o t t r a c k

~ Sa m p le d e c ti on

- R S a m p l e s i t e

/

~ , 4 -

\ O

° ° . -

j c . ~ G o s h

. . . . . . . . ~

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.

.. ... .. : S h o g r & ~ . } ~

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. . ' : . . S h o g r ~ . m ? ~ J . .~ : '~ / L L

. . . .. . . . .. . S ~ ..::':~Za

° . .

i : ' Y ~ ,~

. . . . . . . . . . . . . . . -

~ - ~ i i r i m L a s h t

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

. . . . . .~ R e d ~ . ~ ,: :. . ' a r p i s h

i~ Nol

( ~ S h i t i a r l

°°° .° , '*

• : i ~

: : K u r h g h

l ~ / W & r i n j u n

y ~ ..

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• " . . . . . .

; h a r u n

I I i

s p _ _ u 1

o 2 " e , - o

f g o /

S h ~ d a k

~..o . ~2, o

~ni

3 6 o 1 0 ' I

N

5 9 5 6 m

5 k i n

F i g . 1 .

Bfini-Kur~gh-Shogr~tm-Banamarea of Chitral showing location of the Kuragh section.

as no o lde r t han La t e hassi Zone ( Ta l en t et al.

1999; Fig . 7) . The bes t approxima t ion for the ag e

o f N . batillifera orientalis appears therefore to

be ear ly (poss ib ly very ea r ly) Frasnian .

N. chitralensis i s f rom a recess ive in te rva l o f

da r k sha l e s w i t h i r r egu l a r , t r i l ob i t e - bea r i ng

l im es t one nodu l e s i den t if i ed by one o f u s ( JA T)

t o b e K U R 1 9 o f T a l e n t

et al.

(1999) , but the

subs tantial cono don t da ta f rom th i s hor izon f rom

sampling in 1975 (Talent etal. 1999, Ta ble 1, Fig.

7) inc lude Palm atolepis tenu ipunctata and Pal.

minuta minuta sugges t ing th i s in te rva l i s par t ly

o r e n t i re l y o f e a r l i e st F a m e n n i a n a g e . T h e

conod ont faun a f rom KU R 18, inc identa lly , does

n o t p r o v i d e c o m p e l l i n g d a t a r e l a t i v e t o t h e

F r a s n i a n - F a m e n n i a n b o u n d a r y . A c c o r d i n g t o

da t a f r om e l s ew he r e , the A s t e r opyg i nae d i ed ou t

w i t h t he U ppe r K e l l w asse r Even t a t t he end o f

t he F r a sn i an ( end o f t he linguiformis Z o n e ) . W e

suspec t therefore tha t the co nodo nt sample t aken

in 1975 was f rom abo ve the in te rva l f rom w hich

t he spec im ens o f N . chitralensis were col lec ted ,

and t ha t the U ppe r K e l l w asse r Even t i s l oca t ed

w i t h i n U n i t K U R 1 9 , b u t w i t h o u t a s a l i e n t

s e d i m e n t a r y s ig n a t u r e . T h o u g h a s t r i k in g

sed i m en t a r y s i gna t u r e i s a s soc i a t ed w i t h t h i s

even t in s equences be t w een M or occo and Sou t h

China , there a re im por tant sec tions , such as those

of the Cann ing Bas in o f Western Aus t ra li a , where

there i s no sed imentary s igna ture assoc ia ted wi th

t h is even t a t t he F r a sn i an - F am e nn i an bounda r y ,

t h o u g h i t c a n b e d e t e c t e d n e v e r t h e l e s s o n

isotopic data (Joachims ki & B ugg isch 1996). Th e

bes t app r ox im a t i on f o r t he age o f N . chitralensis

a p p e a r s t h e r e f o r e t o b e l ingui formis Z o n e ,

conce ivably l a te in tha t zone .

alaeobiogeogra ph ic im plications

The d i s cove r y o f F r a sn i an r ep r e sen t a t i ve s o f

Neoca lmonia i n N or t h Pak i s t an en l a r ges t he

d i s t r i bu t i on o f a g r oup p r ev i ous l y cons i de r ed

endem ic to Iran a nd A fghanistan (Pil let 1969; Haa s

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LCHERING

D E V O N I A N T R I L O B I T E S F R O M N W P A K I S T A N 1 9

te becoming f iner bedded and w i th som e si ltstones upwards

Mth occasional strophomenidines

tben -l ike recessive interval of shales and minor si l tstones a nd arenites

:y arenite

• pping interval of f laggy quartzarenites w ith some ripple marks

silty quartzarenites with rubbly bedding

dip no an ular calcareous si l tstone and impure l imestone with cyrtospiri ferids

Nith solitary rugosa ns in upper part

ded l imestone

nterval of nodular\shaley l imestone

50 m ropping impure l imestone

interval of nodular~shaley limestone -

Upper Kellwasser Event

[ l iferous corall ine l imestone w ith comp ound rugosans

often impure and rubbly bedded with cyrtospiri ferids

40 ss-bedd ed quartzarenite with abunda nt cyrtospiri ferids and yphoterorhynchus

and calcareous arenaceous si l tstones, often rubbly bedded, with

:yrtospiri ferids

=ite

h subordinate l imestone

shale-dom inated interval uppermost interval of unit b of Hayde n (1915)

3 0 - -

bed of calcareni te

lite

l iverse brachiopod fauna with rare tri lobites

20 , richly fossiLiferous interval

cyrtospiri fedds with occasional tri lobites

'opping quartzarenite passing upwards into calcarenite with tabulate

corals and minor brachiopods

minor thin arenite

1 0

arenite

ninent bed of calcareni te o r arenaceous l imestone

:areous shale with tabulate and rugose corals, often crushed

ninent bed of massive l imestone

0 m -- r ly bedded impure l imestone

rly bedded impure l imestone with corals and sparse brachiopods

f rom Charun Quartzi te) Coa rse calcareous areni te wi th rare corals and brachiopods

C HA sequence Uni t d of Hayden (1915)

Q U R T

( ~ = Intervals with tri lobites

F i g 2

L o w e r 8 0 m o f K u g h s e c ti o n fromT a l ¢ ~

e t a l

I 9 9 9 ) f r o m w h i c h t r il o b i te s a n d c o n o d o n t d a ta h a v e b e e n

o b t a i n e d .

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5/10

110 R. FEIST, J .A. TA LEN T AN D R. MAW SON A L C H E R I N G A

& M ensink 1969; Ha as 1970; M orzade c 1992).

Indeed, atter their last occurrence in the early

G i v e t i a n o f M y a n m a r ( R e e d 1 9 08 ), n o

Asteropyginae were known far ther east a long

the Nor th Gondw ana marg in than the Afghan

H i n d u K u s h . A m o n g L a t e D e v o n i a n

A s t e r o p y g i n a e , N e o c a l m o n i a s e e m s t o b e

restr icted to G ond wa nan terrains o f the M iddle

East whereas the western European areas are

charac te r i sed by

B r a d o c r y p h a e u s

and al l ies

(Smeenk 1983; Arbizu 1977, and revis ions in

Lieberman & Kloc 1997) . Adap ted to shal low

wate r habitats, species

o f Neocalmonia are

useful

for discriminating local palaeogeographic terrains

dur ing ear ly La te Devon ian t imes . Thus the

S h o gr ~ m F o r m a t i o n o f t h e B O n i - K u r a g h -

Shogr~n-Banam area is palaeogeographical ly

an d en v i r o n m en ta l l y c lo s e ly r e l a t ed t o t h e

Dasch t -e -Nawar a rea , 500 km to the SW in

Afghanis tan .

System atic pa laeontology R .F.)

T h e f i g u r e d s p e c i m e n s a n d s u p p l e m e n t a r y

trilobite material are housed in the Australian

M u s e u m ( S y d n e y ) ; f i g u r e d s p e c i m e n s a r e

catalogued as AM F 118247-118257; locality data

for each sample can be obtained b y refere nce to

Figs 1 and 2.

Superfamily ACA STOIDEA Delo, 1935

Family ACA STIDAE Delo, 1935

SubfamilyASTEROPYGINAE De lo, 1935

Neoealm onia P illet, 1969

Type species: Neocalmonia quadricosta

Pillet,

1969

eoca lm onia ch i t ra l ens is sp. nov. (Figs 3A -L )

Holotype

Pygidium, specimen A M F 118250, Fig.

3, 4G -I, from Unit KU R 19 of Talente ta l (1999),

SW f lank o f Kur~gh Spur in s i tu ab o v e t h e

Chitral-Ma stuj road , 1.5 km S of Kur~gh village,

Chitral in recess ive interval o f dark shales w ith

irregular, trilobite-bearing limestone nodules.

Mater ia l One com plete cephalon, 6 f ragmentary

cephalic remains, 4 incomplete thoracic segme nts

and 7 pygidia.

Der iva tion o f name In r e fe rence to the type

locality, C hitral, the north ernm ost part o f the

North-West Fro ntier Province, Pakistan.

Diagnosis

Cephalon w ith moderately vaulted low

glabella, protruding triangular praefrontal area,

s t r o n g u p r a i s e d o c c i p i t a l s p i n e a n d

protu bera nce s on postero-lateral borders; edge s

of antero-lateral borders f rame d with prom inent

ridge. Pygidium broad, with central nodes on axial

rings; axis triangular in cross section, slightly

constricted at fifth axial ring; posterior lappet

triangular, pro trudin g behind.

Description Cephalon b road ly pen tagonal in

outline (length:width = 1:1.3). Praefrontal area

triangular with blunt, slightly upturned anterior

tip and straight antero-lateral borders, narrower

than the occip i tal r ing (sag .) , subdivided by

suture into two parts o f equal width (sag.). Glabella

mo derately vaulted (tr.) with uninflated ellipsoidal

frontal lobe (length:width = 1:1.6), separated from

lateral glabellar lobes by de eply incise d straight

and adaxially blindly ending 3 furrows. L3 and

L2 unsw ollen, lowe r than flattened central f ield,

incom pletely separated by 2 not reach ing to the

d o r s a l f u r r o w . S 1 a n d i n t e r m e d i a t e r i n g

inconspicious. Dorsal furrows m ode rately deep,

diverging behind, stron gly ben t arou nd adaxially

p ro t rud ing an te r io r par t o f pa lpebra l lobes .

Occipital furro w shallow , straight, d eep eni ng

behind L1. O ccipital r ing mo dera tely inflated,

m uch high er than glabella, raised posteriorly, and

me dially running into a strong upward ly directed

spine. Palpebral lobe f lat- topped, higher than

glabella, having a continuous groove running

paral le l to i ts edge. Inner par ts of f lx igenae

modera te ly in f la ted , adax ia l ly inc l ined . Eye

separated f rom deep poster ior border fur row by

extrem ely narrow , vertical portion o f f ixigenae.

Genal angle offix igen ae inflated.Posterior borders

semicylindrical, run ning distally into prom inen t,

b lunt protuberances . Eyes rather h igh; v isual

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A L C H E R I N G A D E V O N I A N T R I L O B I T E S F R O M N W P A K I S T A N 111

Fig 3 A - L Neocalmonia chitralensis sp. nov. A -D AM F 118247 cephalon external mould; A dorsal view x3.5;

B lateral view x3.1; C poster ior view x3.5; D frontal view x3.2. E AM F 118248 pygidium ext. mould partly

exfoliated dorsal view x5.3. F AM F 118249 juven ile pygidium ext. mould x7.4. G -I Holotype AM F 118250

pygidium ext. mould; G dorsal view x4.1; H lateral view x4.1; I posterior view x3.9. J-L AM F 118251 pygidium

ext. mould tip o f central marginal spine broken; J dorsal view x7; K posterior view x7.4; L lateral view x7.2.

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1 12 R . F E I ST , J . A . T A L E N T A N D R . M A W S O N A L C H E R I N G A

surface steeply inclined in front and laterally,

vertical behind, ca rrying 2 7 dorsoven tal lens files

with a maximum of 7 lenses. Eye s ocle surrounded

by a thick, sw ollen crest separating i t from the

a d j a c e n t , c o n c a v e , s t e e p l y s l o p i n g o u t e r

l i b r i g e n a e . L a t e r a l m a r g i n s w i t h o u t b o r d e r

furrows, fra me d by a strong, con tinuous ridge.

Outer slope of margins vertical , pro vided w ith a

continuous g roo ve running parallel to the margin.

Genal sp ines broad-based, ha l f as long as the

cephalon (sag.), carrying on their inner third a

prominent c rest s ta r t ing in the gena l angle .

Sculpture: sparse drop-like nodules; marginal

a r e a s a n d o c c i p i t a l l o b e s m o o t h . T h o r a c i c

segments with roun ded pleural ends, carrying a

stout node on the distal posterior margin.

Pygidium broadly semi-elliptical (length:width

= 1:1.85). Axis as wid e as one pleural f ie ld (tr .) ,

triangular in cross section, horizontal in side view,

provided wit h 12 + 2 straight axial rings that carry

blunt , an te r ior ly sp iny cent ra l nodes. Dorsa l

furrows convergent, sl ightly constricted at the

level o f he 5th axial ring. Posterior end o f the axis

elevated above narrow post-axial f ie ld, extended

in to a t i ny pos t -ax ia l r idge in t e rrup t ing the

marginal furrow. Pleural f ie lds with 5 ribs, the

anterior three being slightly bent, the posterior

ones straightly divergent from the dorsal furrow.

Pleural bands thick, ofs uprad evon ica-type H a a s

1970), provided with stout spines on the distal

third o f the posterior bands. Bord er furrow deep,

sepa ra t ing s lop ing p leu ra l f i e ld f rom b road ,

horizontal margin. Pleural spines large, sloping

below the m argin, provid ed with crest-l ike ridges

on their external margin. Central spine triangular,

very large at base, shorter than lateral spines

( s a g . ) , s l i g h t l y u p t u r n e d . S c u l p t u r e : t i n y ,

elongated nod ules on the spines o f the axial rings

and p leura l bands as wel l as on the exte rna l

borders o f marg inal spines.

R e m a r k s . The new spec ie s i s c lo se s t t o N .

b a t i l l if e r a a n d N . k a k a r i desc ribed by Haas

Mensink (196 9) from Dascht-e-Nawar in eastern

Afghanistan. In fact, the cep halo n is clos er to

that ofN. bat i l l i f era as bo th share the particular,

adaxially protruding praefrontal area; the latter

remains rath er narro w in N. k a k a r i . H o w e v e r t h e

shape o f this area is m ore triangular with a m ore

pointed tip and straight anterolateral borders in

t h e n e w s p e c i e s . F u r t h e r m o r e , t h e s u t u r e

separates the praefrontal area into two parts of

equal width (sag.) wh ereas the outer part is m uch

larger i n N . ba t i l l i f era . In the new spec ies the

uninflated lateral glabellar lobes remain bel ow the

top o f the central area of the glabella , and the y

are not separated fro m it by an exsagittal furro w

as in the Afghan species. The pygidia contrast

mainly in the shape o f the axis: triangular in cross-

section and slightly constricted laterally in N.

c h i t r a l e n s i s . In addition, the axial rings carry

centra l nodes, a fea ture tha t i s absent in N.

bat i l l i f era . In contrast, these characters oc cu r in

N . k a k a r i , but the pygidium o f the la tter m ay be

discriminated b y its equilateral triangular outline

and the posteriorly lower, dow n-turned axis.

N e o c a l m o n i a b a t i i l i f e r a

Haa s Mensink, 1969

N e o c a l m o n i a b a t i l l i f e r a o r i e n t a l i s subsp, nov.

(F igs 4A-J )

H o l o t y p e . Pygidium, specimen A M F 118256 Fig.

4G, H , from the Shogram Formation, Unit KU R 9B

of Talent e t a l . (1999) , SW f lank of Kur~gh Spur

i n s i t u abov e the Chi t ra l-Mastu j road, 1 .5 km S

o f uraghvil lage, Chitral in dark grey muddy

limestone w ith brachiopods.

M a t e r i a l . 7 cepha la , an incomple te tho rac ic

segment, and 7 pygidia.

F i g 4

A - J .

N e o c a l m o n i a b a t i l l i f e r a o r i e n t a l i s

subsp nov . A-B AM F 118252 cephalon fragmentary la tex cast o f

ext . mou ld; A x4.4; B posterior view x4.3. C AM F 118253 cephalon latex cast of ext . mou ld dorsal view x4.2.

D AM F 118254 cephalon fragmentary ext . m ou ld par t ia l ly exfo l ia ted fronta l v i ew x6 .5 . E-F AM F 11855

cephalon ext . mou ld partial ly exfol iated E lateral view x4.4; F dorsal view x4.1. G-H Holotype AM F 118256

pygidium fragmentary ext . mou ld G dorsal view x3.2; H lateral view x3.0. I-J A M F 118257 pygidium fragmen-

tary ext . mou ld of smal l specimen I dorsal view x7.3; J x6.9.

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LCHERING

D E V O N I A N T R I L O B I T E S F R O M N W P A K I S T A N 3

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114 R . FEIST, J . A . TAL ENT AN D R . M AW SON ALCHERINGA

D e r i v a t i o n o f n a m e . I n r e f e r e n c e t o it s o c c u r r e n c e

e a s t o f t h e t y p e l o c a l i t y o f t h e n o m i n a t e

s u b s p e c i e s .

Diagnosis

P r e g l a b e l l a r a r e a n a r r o w e r t h a n

occ ip i ta l r ing (sag . ) , not e leva ted anter ior ly ;

posterior margin behind cheeks withou t spines.

Pos te r io r pyg id ia l p l eu rae f l exed backwards

adaxially, wit hou t distal protuberances. First tw o

ax ia l r ings o f pyg id ium wi th med ia l nod es ;

pygidial bord er narrow w ith stout anterior lappets

and laterally ex pan ded terminal lappet.

Remarks The

new material is ver y close to the

nominate subspec ies in the degree o f g label lar

v a u l t in g - - - - c o n s id e r a b l y b e l o w t h e e y e s - - a n d i n

t h e o u t l i n e o f t h e s l i g h tl y f u s e d 3 - S 1 f u r r o w s

a n d t h e s w o l l e n p o s t e r i o r g l a b e l l a r l o b e . T h e

c r e s ts o f t h e d i s ta l p o s t e r i o r m a r g i n s b e h i n d t h e

c h e e k s a r e o n l y s l i g h t l y sw o l l e n , b u t d o n o t e x h i b i t

p r o m i n e n t p r o t u b e r a n c e s a s s e e n i n t h e n o m i n a t e

s u b s p e c i e s . T h e a n t e r i o r a r e a i n f r o n t o f th e

g l a b e ll a i s fi a t a n d p r o t r u d e s f o r w a r d m e d i a l l y t o

a l e s s e r d e g r e e . T h e p y g i d i u m i s c h a r a c t e r i s e d

b y s p i n e s o n t h e m e d i a l a n t e r i o r a x ia l r in g s ; t h e s e

a r e a l s o s e e n i n N . c h i t r a l e n s i s b u t , i n c o n t r a s t t o

t h e l a tt er , t h e p r e s e n t f o r m l a c k s p r o t u b e r a n c e s

o n t h e d i s t a l p o s t e r i o r p l e u r a l b a n d s . A

c h a r a c t e r i s t i c f e a t u r e o f th e n e w t a x o n i s t h e

p r o n o u n c e d b a c k w a r d b e n d i n g o f t h e a n t er i o r

p l e u r a e ; t h i s o c c u r s i n b o t h t h e n o m i n a t e

s u b s p e c i e s a n d N . c h i t r a l e n s i s . T h e r a t h e r b r o a d

a n d s h o r t o u t li n e o f th e t r i a n g u la r t e r m i n a l l a p p e t

i s a n o t h e r p a r t i c u l a r t r a i t o f N .

o r i e n t a l i s .

cknowledgments

The work w as supported by a grant to RF by the

French CN RS flSEM , U M R 5554, publ. 2001-000)

and grants from the Australian Research Grants

C o m m i t t e e t o J A T ( 1 9 7 3 - 7 5 ) . T h i s i s a

contribution to IGCP 421 no r th Gondw ana mid-

Palaeozoic b ioevent /b iogeography pa t te rns in

relation to crustal dynamics .

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Devonian Trilobites From the Shogr m Formation Kur Gh Chitral NW Pakistan