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Anomalocaris and Other Large Animals in the Lower Cambrian Chengjiang

Fauna of Southwest China

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 GFF volume 117 (1995), pp. 163-183.
Anomalocaris and other large animals in the Lower Cambrian Chengjiang fauna
of southwest China
HOU XIAN-GUANG, JAN BERGSTRÖM, and PER AHLBERG
Hou, X.-g., Bergström, J. & Ahlberg, P., 1995: Anomalocaris and other
large animals in the Lower Cambrian Chengjiang fauna of southwest
China. GFF, Vol. 117 (Pt. 3, September), pp. 163-183. Stockholm. ISSN
1103-5897.
Abstract: Five anomalocaridids are described from the Lower Cam-
brian Chengjiang fauna of China: Anomalocaris saron sp.n., Anoma-
locaris sp., Amplectobelua symbrachiata gen. et sp.n., Cucumericrus
decoratus gen. et sp.n., and Parapeytoia yunnanensis gen. et sp.n. Ven-
tral trunk appendages are reported for the first time. In C. decoratus and
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P. yunnanensis, each appendage has a distal segmented walking leg, and
a wide lateral extension of a long proximal portion forms the structure
previously described as a lateral fin. Furthermore, the structure of the
dorsum is elucidated. The supposed ventrolateral gills are flat scales
covering the back in transverse rows. The mouth was directed back-
wards, as in the Middle Cambrian Opabinia regalis, and not directed
ventrally as in previous reconstructions. The radial arrangement of the
circum-oral sclerites suggests that anomalocaridids were related to
aschelminth worms rather than to arthropods.
Keywords: Anomalocaridids, Aschelminthes, taxonomy, morphology,
functional morphology, Lower Cambrian, Chengjiang fauna, Yunnan,
China.
Hou Xianguang, Nanjing Institute of Geology and Palaeontology, Acade-
mia Sinica, Chi-Ming-Ssu, Nanjing 210008, People's Republic of China,
fax 25/3357026. Jan Bergström, Swedish Museum of Natural History,
Department of Palaeozoology, Box 50007, S-104 05 Stockholm, Swe-
den, fax 08/6664184. Per Ahlberg, Department of Geology, Sölvegatan
13, S-223 62 Lund, Sweden, fax 046/121477. Manuscript received 30
June 1995. Revised manuscript received and accepted 31 July 1995.
Anomalocaridids are a group of fossil animals that includes
Anomalocaris and similar forms from the Burgess shale and other
Cambrian deposits. They were first found as isolated fragments
thought to represent crustaceans, jellyfish, and holothurians. Our
knowledge has been growing successively and the parts have been
brought together into a segmented animal with a pair of large
frontal appendages but with no definitely identified trunk append-
ages (see Whittington & Briggs 1985; Gould 1989; Chen et al.
1994). Specimens from the Burgess shale, described by Whitting-
ton & Briggs (1985), have revealed that the mouth was situated
on the ventral side of the head and surrounded by a circlet of
sclerites. It was previously known only in isolation and interpreted
as the medusoid Peytoia Walcott, 1911. The frontal appendages
are generally preserved in isolation, and they were once believed
to represent the trunk of a crustacean (e.g., Whiteaves 1892; Hen-
riksen 1928; Resser 1929). The segmented body was originally
described as a holothurian echinoderm (Laggania cambria Wal-
cott, 1911). The first nearly complete outlines of anomalocaridids
were only recently reported (Gore 1993; Chen et al. 1994).
For a long time anomalocaridids were known only from Lower
Article
and Middle Cambrian strata in North America (e.g., Briggs &
Mount 1982; Briggs & Robison 1984; Whittington & Briggs
1985). Recent discoveries have shown that they were geographi-
cally widespread, and anomalocaridids have now been described
from the Lower Cambrian of Poland (Dzik & Lendzion 1988),
southern China (Hou & Bergström 1991; Chen et al. 1994), and
South Australia (McHenry & Yates 1993). In addition, an anoma-
locaridid jaw apparatus was recently described from the Upper
Cambrian in the Holy Cross Mountains of Poland (Masiak &
Zylinska 1994).
Anomalocaridids have recently attracted great interest because
of their large size and unusual combination of characters. Al-
though the frontal appendages are similar to those of many ar-
thropods, it is generally agreed that anomalocaridids are unre-
lated to normal arthropods. Chen et al. (1994), however, suggest
that the combination of certain features, such as growth by moult-
ing and true segmentation, indicates affinities to arthropods.
Briggs & Robison (1984) and Whittington & Briggs (1985) fa-
vours the idea that Anomalocaris and related forms represent a
separate phylum, whereas Bergström (1986,1987,1989) and Dzik
& Lendzion (1988) regarded anomalocaridids as representing a
separate trend of arthropodization. In this paper we provide evi-
dence for assignment of the anomalocaridids to the aschelminths
rather than to the arthropods.
The anomalocaridid material described herein was collected
from the famous Chengjiang fauna of southwest China. The fauna
was discovered in 1984 and occurs in the basal and lower parts of
the Eoredlichia Zone of the Lower Cambrian Qiongzhusi (Chiung-
chussu) Formation (e.g. Hou & Sun 1988; Hou & Bergström
1991), a level approximately equivalent to the Atdabanian (Bengt-
son et al. 1990, pp. 16-17). The composition and preservation of
the fauna were reviewed by, e.g., Hou et al. (1991), Chen et al.
(1991), and Chen & Lindström (1991).
When the first soft-bodied fossil (the holotype of Naraoia
longicaudata Zhang & Hou, 1985) was discovered at Maotianshan
in Chengjiang in 1984, new collections aimed for soft-bodied
fossils were made immediately. The fauna is now known from
several locations around Kunming City in Yunnan Province.
The material treated herein was collected from the sections at
Maotianshan (Collection Number M2 and M3), Xiaolantian (Coll.
No. XL1), and Dapotou (Coll. No. DJ1) in Chengjiang (Fig. 1).
Anomalocaridids are rare in the Chengjiang fauna and generally
preserved as isolated fragments. However, collecting over a
number of years has resulted in a substantial number of speci-
mens, and our knowledge on the morphology of anomalocaridids
has been growing in pace with the accumulation of new material.
Five species of anomalocaridids are described from the Cheng-
jiang fauna, three of which seem to represent three new genera,
Amplectobelua, Parapeytoia, and Cucumericrus. The record of
trunk appendages in Parapeytoia yunnanensis gen. et sp.n. and
Cucumericrus decoratus gen. et sp.n. brings perspective to the
 164 Hou et al: Anomalocaris and other large animals in the Chengjiang fauna, China
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claim that anomalocaridids were devoid of such appendages
(Whittington & Briggs 1985; Gore 1993; Briggs 1994; Chen et
al. 1994).
The Chengjiang fauna also includes various spiny structures
(teeth), some of which are preserved in isolation, some preserved
in pairs, and some preserved in groups. Chen et al. (1994) inter-
preted these elongate and generally large sclerites as belonging
to the mouth of anomalocaridids. They were used by Chen et al.
(1994) to estimate a maximum length of about 2 m in anoma-
locaridids.
The material to hand seems to indicate, however, that the mouth
of large anomalocaridids is very similar to the mouth of Peytoia
nathorsti Walcott, 1911, and the spiny sclerites do not seem to
represent anomalocaridid jaw parts. The arrangement of the spiny
sclerites in the Chengjiang fauna is comparable with hooks and
scalids in aschelminths such as priapulids. Thus, the sclerites
appear to have been situated on an introvert and they possibly
represent a priapulid worm (Bergström et al. in prep.).
Terminology
Although the morphology of anomalocaridids includes arthropod-
type features, such as articulated legs, we avoid morphological
terms that have been applied for arthropods. This is because we
regard anomalocaridids as related to aschelminth worms rather
than to arthropods. We also avoid terms which imply functions,
such as fins, gills, and swimmerets, and stick to clearly descriptive
terms without homology implications (Fig. 16C).
Appendage flap. - Wide lateral extension of the propod. It was
previously described as a lateral lobe and interpreted as a lateral
fin along which waves of motion were propagated (Whittington
& Briggs 1985; Briggs 1994).
Endite. - Medial projection of a leg segment.
Oral circlet. - Circle of sclerites surrounding the mouth. In
GFF117(1995)
Fig. 1. Map of small
part of central
Chengjiang County,
Yunnan, southwest
China, showing
localities mentioned
in the text.
x fossil locality
O village
canal
3= roads
anomalocaridids, the structure was originally described as the
'medusa' Peytoia (Walcott 19116).
Propod. - Proximal undivided part of limb that appears to consist
of five fused segments. It extends laterally into an appendage
flap.
Ramipod. - Segmented pediform leg branch attached to the
propod.
Scale. - General term for hardened portion of skin. In Anoma-
locaris, Parapeytoia, Peytoia, Cucumericrus, and Opabinia, long
and narrow lanceolate scales are placed in transverse rows over
the dorsum. These scales were previously recognized as ventro-
laterally situated gills (Whittington 1975; Whittington & Briggs
1985; Chen et al. 1994).
Shaft. - The flexible extension between body and grasping ap-
pendages. This corresponds to the "tubular frontal process" of
Opabinia in Whittington's (1975) terminology.
Sternite. - Sclerotised part of ventral surface of body. In anoma-
locaridids there are segmental median sternites in the space
between the paired limbs.
Systematic palaeontology
Illustrated specimens are deposited in the Nanjing Institute of
Geology and Palaeontology, Academia Sinica (NIGPAS). All
photographs are of dry, uncoated specimens.
Subkingdom Aschelminthes Grobben, 1908
Class Dicephalosomita Sharov, 1966
Remarks. - This class is here interpreted as including extinct
 GFF117(1995) Hou et al: Anomalocaris and other large animals in the Chengjiang fauna, China
marine forms with a radial arrangement of the circumoral sclerites,
a preoral pair of large grasping appendages in cases set on a shared
flexible shaft, successive segmental appendages with a wide lateral
flap, and transverse dorsal sets of flat scales in cases in addition
to segmental tergites. The mouth was situated ventrally on the
head region and appears to have been directed backwards.
Kerygmachela from the Lower Cambrian 'Sirius Passet' fauna
of Greenland has strong similarities with anomalocaridids which
probably indicate close affinities. These similarités include, e.g.,
a pair of large grasping appendages, distinct segmentation with
transverse dorsal sets of elements comparable with the anoma-
locaridid scales, and paired extensions from the tail region (cf.
Chen et al. 1994, p. 1307). The lateral projections from the axial
region of Kerygmachela, interpreted as stout walking append-
ages by Budd (1993), seem to be uniramous and have a lobopodian
appearance.
Groups included. - Opabiniida St0rmer, 1944 with Opabiniidae
Walcott, 1912 and Anomalocarididae Raymond, 1935; probably
also a group containing Kerygmachela Budd, 1993.
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Family Anomalocarididae Raymond, 1935
Remarks. - This family differs from the Opabiniidae Walcott,
1912, in lacking a long shaft of the grasping appendages.
Genera included. - Anomalocaris Whiteaves, 1892; Peytoia
Walcott, 1911; Cassubia Lendzion, 1977; Parapeytoia gen.n.;
Amplectobelua gen.n.; Cucumericrus gen.n.
Genus Anomalocaris Whiteaves, 1892
Type species. -Anomalocaris canadensis Whiteaves, 1892.
Emended diagnosis. -Anomalocaridids with long, distally tapered
grasping appendages consisting of 15-16 segments, junction
between the two proximal segments nearly obscured; each seg-
ment, apart from the most distal and proximal ones, bears a pair
of endites armed with lateral and terminal spines; upper ('anterior')
part of trunk appendage flaps distinctly striated with evenly spaced
lines.
Other species. - A. pennsylvanica Resser, 1929 (p. 12, pi. 5, fig.
5); A.I cf. pennsylvanica of Briggs (1979, pp. 659-660, pi. 81,
figs. 9-11, text-figs. 33-34); A. sp. of McHenry & Yates (1993,
p. 81, figs. 2-7 and 8-9?); A. sawn sp.n. (= A. n.sp. of Chen et al.
1994); A. sp. in this paper.
Anomalocaris saron sp.n.
Figs. 2-6
Synonymy. -»1991 Anomalocaris canadensis - Hou & Bergström,
p. 185 (partim), PI. 2, fig. 2. »1994 Anomalocaris n. sp. - Chen et
al.,p. 1305,figs. l-2.»1994AnomaZocaris-Briggs,upperfigure
on p. 1283.
Etymology. - Greek saron, broom, referring to the numerous
spines on the endites.
Holotype. - An incomplete but well-preserved grasping appendage
165
with 10 preserved segments, NIGPAS Cat. No. 115337a and b
(part and counterpart; Figs. 2C, 2F, and 3B). The specimen shows
the presence of long endites, each of them having two pairs of
lateral spines.
Other material. - NIGPAS Cat. No. 110827 (Figs. 2B and 3A)
from coll. No. DJ1, illustrated by Hou & Bergström (1991, PI. 2,
fig. 2); NIGPAS 115338 (Fig. 2D) from coll. No. XL1, a large
grasping appendage (incomplete); NIGPAS 115336 (Fig. 2A)
from coll. No. Cf5, a nearly complete grasping appendage;
NIGPAS 115339 (Fig. 2E), an incomplete grasping appendage;
NIGPAS 115340 (Figs. 4-5) from coll. No. M2, a specimen with
seven appendage flaps (part and counterpart); NIGPAS 115341
(Fig. 6) from coll. No. M3, a specimen preserved in disorder (part
and counterpart); numerous isolated and incomplete grasping
appendages (unnumbered).
Type locality and type horizon. — Jianbaobaoshan near Dapotou
Village, Chengjiang; levelDJl, lowerpart of the Eoredlichia Zone
of the Lower Cambrian Qiongzhusi Formation.
Diagnosis. - A species of Anomalocaris with grasping appendages
provided with long, slender endites; each endite on grasping
appendage bears two pairs of lateral spines.
Description. - The grasping appendages are long, slender, and
taper distally. As preserved, all specimens are curved at varying
degrees. The appendages are generally incompletely preserved,
but rare specimens indicate that they consist of 16 segments. Each
segment, apart from the most proximal one and the most distal
one, has a pair of long medial endites. In the more proximal seg-
ments of the holotype, the endites are directed more or less normal
to the medial (inner) margin of the appendage, whereas they are
distally directed and inclined to the medial margin in the distal
podomeres. Two pairs of long spines extend from each endite at
an angle of 15-30°. There is apparently some variation in the
position of these 'auxiliary' spines, but in the distal segments
they tend to project from near the base of the endite, whereas
they project from about the mid-length of each endite in the more
proximal segments. The second segment bears a pair of short
endites devoid of spines. The junction between the two proximal
segments is nearly obscured.
The lateral (outer) margin of the more distal segments is pro-
duced distally into a pair of elongate spines (one of the pair is at
an angle to the bedding plain and is usually not seen) extending
parallel to the appendage and overlapping the segment bounda-
ries. In general, there appears to be four or five pairs of such
spines, and they become progressively sturdier and longer to-
wards the distal end of the appendage. The terminal segment car-
ries two pairs of medially curved spines or claws. The outer pah-
is elongate and similar to the lateral spines of the adjacent seg-
ments. The inner pair is short and subtriangular in outline.
Most of the grasping appendages have an estimated total length
ranging from less than 2 to 8 cm. One specimen shows, however,
that they could be considerably larger. In this specimen four seg-
ments are preserved only, each being 1.5-1.7 cm long (Fig. 2D).
Thus, the complete appendage must have been at least 20 cm
long.
One specimen displays seven appendage flaps along one side,
and transverse sets with lanceolate scales (Figs. 4—5). The over-
lap between successive flaps is extensive, each flap overlapping
the one in front. The flaps are flat and widest (exsag.) proximally,
 166 Hou et al: Anomalocaris and other large animals in the Chengjiang fauna, China GFF117(1995)
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wmÊOiî
Fig. 2. Anomalocaris saron sp.n. A. Nearly complete grasping appendage. Coll. No. Cf5 (northwest slope of Maotianshan). NIGPAS 115336. x5.
B. Nearly complete grasping appendage. Coll. No. DJl. Original of Hou & Bergström (1991, pi. 2, fig. 2). NIGPAS 110827. x7. C. Holotype, distal
 GFF117(1995) Hou et al: Anomalocaris and other large animals in the Chengjiang fauna, China
0.5 cm
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becoming gradually narrower towards the distal edge, which is
rounded. Evenly spaced lines, trending transversely, are present
on the anterior and antero-lateral portion of the appendage flaps.
The lines are gently curved and inclined to the antero-lateral
margin of the flap. There are 18-20 lines in the anterior flaps.
The number of lines decreases backwards and there are only 9
lines on the 7th flap.
Another specimen exhibits three striated appendage flaps. They
are incomplete, preserved in disorder, and associated with rows
of scales, sternites, a poorly preserved oral circlet, and an iso-
lated grasping appendage (Fig. 6). The scales are arranged in dis-
tinct bands with an undulating course. In one of the flaps, a pos-
terior line extends obliquely to intersect three of the evenly-spaced
lines.
Remarks. - I n the description above, the directions of the grasping
appendages are given as lateral, medial, proximal, and distal,
assuming that they were attached to the head as indicated on a
specimen figured by Chen et al. (1994, figs. 1-2).
This species was briefly described by Chen et al. (1994) as
Anomalocaris n. sp. In their paper they illustrated a specimen
with a nearly complete outline. No comparisons between this new
species and A. canadensis were made. The strong inclination of
the "fins" of this specimen (Chen et al. 1994, figs. 1-2) makes a
broad, horizontal attachment on the side of the body impossible.
It fits, however, with the present interpretation of the 'fin' as a
part of a ventral appendage.
Anomalocaris saron most closely resembles A. canadensis from
the Middle Cambrian of British Columbia. One major difference
is that the grasping appendages of A. saron have longer medial
endites. Furthermore, the endites of A. saron are provided with
part of grasping appendage showing endites with spines. Counterpart to Fig. 2F. Coll. No. DJ1. NIGPAS 115337b. x3. D. Large grasping append-
age, incomplete. Coll. No. XL1. NIGPAS 115338. xl.5. E. Distal part of grasping appendage showing spines on the lateral (outer) margin. NIGPAS
115339. x4. F. Holotype, a well-preserved grasping appendage with 10 preserved segments. Counterpart to Fig. 2C. Coll. No. DJ1. NIGPAS
115337a. x2.25.
167
Fig. 3. Anomalocaris
saron sp.n. A.
Drawing of speci-
men shown in Fig.
2B. B. Drawing of
holotype (cf. Fig.
2F). C. Reconstruc-
tion of grasping
appendages.
two pairs of lateral spines, whereas the endites of A. canadensis
have only one pair of lateral spines. The appendage flaps of A.
canadensis do not show any lines, but this may be due to the
preservation. Anomalocaris sp. of McHenry & Yates (1993) dif-
fers widely from A. saron in having many small spines on each
endite.
Anomalocaris sp.
Figs. 7-8
Synonymy. -1991 Anomalocaris canadensis - Hou & Bergström,
p. 185 (partim), PL 2, fig. 3.
Material. - NIGPAS 110828 (Figs. 7A and 8A) from coll. M2,
an isolated grasping appendage illustrated by Hou & Bergström
(1991, PL 2, fig. 3); NIGPAS 115342 (Figs. 7B and 8B) from
coll. M3, an isolated grasping appendage (part and counterpart);
NIGPAS 115343 (Fig. 7C-D) from coll. M3, an isolated grasping
appendage (part and counterpart); several incomplete grasping
appendages (unnumbered).
Discussion. - The appendages of Anomalocaris sp. are closely
comparable to those of A. saron sp.n. in having long endites with
two pairs of lateral spines. They differ from those of the latter in
being much smaller (maximum length less than 2.5 cm), more
strongly curved, and less tapered distally. Furthermore, the seg-
ments of A. sp. are shorter than in A. saron. In A. sp. the second
segment bears a long endite with 'auxiliary' spines, whereas the
second segment of A. saron bears a simple, short spine devoid of
lateral spines.
 168 Hou et al: Anomalocaris and other large animals in the Chengjiang fauna, China
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F/g. 4. Anomalocaris saron sp.n. A specimen with seven appendage flaps (part and counterpart). Coll. No. M2. NIGPAS 115340. xl.8.
The appendages are here interpreted as grasping appendages,
and A. sp. and A. saron are regarded as two separate taxa. There
is the possibility, however, that they are conspecific and that the
appendages under discussion represent ventral, pediform legs of
A. saron.
Genus Parapeytoia gen.n.
Etymology. - Greek para, near, and Peytoia, name of a previously
described anomalocaridid.
Type and only known species. - Parapeytoia yunnanensis sp.n.
Diagnosis. - Anomalocaridids with disconnected grasping ap-
pendages provided with a stout and curved proximal segment, a
long terminal finger, and three long and flat serrated endites;
propod with strong medial spines and extended laterally into large
flap; ramipod a distinctly segmented, pediform branch devoid of
distal spines; two pairs of small postoral appendages with seg-
GFF117 (1995)
ments prolonged into endites; medial sternites well developed,
the three anterior ones about half as long as those further back.
Parapeytoia yunnanensis sp.n.
Figs. 9-13,18A, 19
Etymology. - After Yunnan, a province of southern China.
Holotype. - NIGPAS Cat. No. 115334, a specimen with grasping
appendages, a circular mouth, eight sternites, seven pairs of legs,
and lanceolate scales (part and counterpart; Figs. 9-10,18A).
Other material. -NIGPAS Cat. No. 115345 (Figs. 1 ID and 12D)
from coll No. DJ1, an isolated appendage, probably from the
posterior part of the animal (part and counterpart); NIGPAS Cat.
No. 115344 (Figs. 11B-C and 12A-C) from coll. No. M3, an
isolated grasping appendage lacking the proximal segment (part
and counterpart); NIGPAS Cat. No. 115335 (Figs. 11A and 13),
proximal-medial portion of appendage with gnathobases; two
 GFF117 (1995) Hou et al: Anomalocaris and other large animals in the Chengjiang fauna, China 169
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Fig. 5. Anomalocaris sawn sp.n. Drawing of specimen shown in Fig. Fig. 6. Anomalocaris saron sp.n. Drawing of specimen preserved in dis-
4A. order; m, mouth (oral circlet); ga, grasping appendage; af, appendage
flap; sc, sets of lanceolate dorsal scales; s, medial sternite? Coll. No.
M3.NIGPAS 115341.

fragmentary grasping appendages (unnumbered); five fragmentary
appendage flaps with gnathobases (unnumbered) are tentatively
assigned to the species.
Type locality and type horizon. - Maotianshan (Maotian Hill) in
Chengjiang County; level M2, lower part of the Eoredlichia Zone
of the Lower Cambrian Qiongzhusi Formation.
Diagnosis. — As for genus.
Description. - The grasping appendages consist of five segments.
The proximal segment is stout and curved. The three successive
segments are extended into long blade-shaped endites. The most
proximal endite is serrated on one edge, whereas the two others
are serrated on both edges. The terminal segment is extended into
a long, slightly curved finger, which is serrated on the inner edge.
The trunk appendages consist of a stout proximal part (propod)
and a distal segmented branch (ramipod). The medial edge of the
former carries strong spines arranged on five weakly defined
gnathobases, indicating that the propod probably consists of five
segments. These supposedly fused segments extend laterally Into
a wide lateral flap, which is separated into anterior and posterior
fields by a line. The proximal field is striated in the area adjoin-
ing the appendage-body attachment (Figs. 11A and 13). The
ramipod is pediform and consists of eight cylindrical segments
plus a conical terminal element. Each podomere is almost sur-
rounded by a zone of spines.
The holotype displays a compressed mouth with radially ar-
ranged sclerites. This oral circlet is probably preserved in its origi-
nal position, and it is directed backwards. This is at high angle
with the bedding and is not likely to have been so if the mouth
had originally been facing ventrally. Two pairs of small postoral
cephalic appendages with segments prolonged into endites are
present in the holotype. The medial sternites are well developed.
The three anterior ones are short (sag.) and subquadratic in out-
line. The successive sternites are considerably longer and sub-
rectangular In outline. In the holotype there are also patches of
dorsal lanceolate scales characteristic of anomalocaridids.
Remarks. - The holotype shows much of the underside of this
 170 Hou et al.: Anomalocaris and other large animals in the Chengjiang fauna, China GFF117(1995)
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Fig. 7. Anomalocaris sp. Grasping appendages. A. NIGPAS 110828. Coll. No. M2. Original of Hou & Bergström (1991, pi. 2, fig. 3). x6. B. NIG-
PAS 115342. CoU. No. M3. x8. C-D. NIGPAS 115343 (part and counterpart). Coll. No. M3. x4.

species, demonstrating the type of structures that belong together.
The somewhat distorted arrangement in the specimen is interpreted
as indicating that it is an ecdysial moult. Only remains of a few
sets of dorsal scales are seen. The posterior part is missing, but
an isolated appendage (Figs. 11D and 12D) may represent the
enlarged appendages of the tail region.
Parapeytoia yunnanensis is most similar to Peytoia nathorsti
Walcott, 1911, and Cassubia infercambriensis (Lendzion, 1975)
in having long endite blades extending from the segments of the
grasping appendages. It differs, however, in having a long termi-
nal finger, whereas the terminal segment is very short in the other
species, and is provided with strong claws, at least in P. nathorsti.
Furthermore, P. yunnanensis has only five segments and three
blade-shaped endites in the grasping appendage, whereas the cor-
responding numbers are 11 and 9 in P. nathorsti and apparently
still higher in C. infercambriensis, at least 19 and 12 according to
the reconstruction (Dzik & Lendzion 1988, fig. 3A). Except for
the most proximal endite and the terminal finger, the endite blades
are serrated on both edges in P. yunnanensis, whereas C. infer-
cambriensis has one smooth and one serrated edge, and P. nat-
horsti one smooth and one strongly spinous edge. The unique
morphology of the grasping appendage of P. yunnanensis justi-
fies recognition as a separate genus.
Genus Amplectobelua gen.n.
Etymology. - Latin amplecto, embrace, and belua, monster.
Type and only known species. -Amplectobelua symbrachiata sp.n.
Diagnosis. - Anomalocaridids with grasping appendages provided
 GFF117 (1995) Hou et al: Anomalocaris and other large animals in the Chengjiang fauna, China 171

Diagnosis. - As for genus.

Description. - Two pairs of grasping appendages are well-pres-

erved with the medial margins facing each other (Figs. 14A-B
and 15A-B). The appendages are parallel along the two proximal
segments, curved outward from the 3rd and 4th segment, and
then curved inward distally.
The grasping appendages consist of 15 segments (numbered
herein 1-15 from the proximal end). Segments 1-2, and 4 in par-
ticular, are comparatively long, whereas segments 3 and 5-15
are short; segments 5—15 becoming gradually shorter distally.
Each segment except for the most proximal one bears a pair of
long, pointed endites. They are frequently broken, but a rounded
structure ventro-medially or dorso-medially on the segments in-
dicates the site of attachment. The endites alternate in length on
successive segments, but they generally become smaller towards
the distal end of the appendage. The endite is exceptionally long
(one-third to nearly half as long as the length of the appendage)
and stout on segment 4, which also carries a pair of short spines
projecting from near the base of the main endite. The endite on
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segment 2 is distally directed and inclined to the medial margin

at an angle of 15-20°. The endites on segments 3 to 12 become
progressively more inclined, and on segments 11-12 they extend
B from the medial margin at an angle of 70-80°. The endites on
segments 13 and 14 are directed slightly backwards or normal to
the medial (inner) margin. There is no evidence for articulation
of the endites.
Fig. 8. Anomalocaris sp. A. Drawing of specimen shown in Fig. 7A. B. The lateral (outer) margin of segments 10-14 is produced into
Drawing of specimen shown in Fig. 7B.
an elongate, distally directed spine that extends more or less par-
allel to the appendage. One specimen (Figs. 14C and 15C) shows
that these spines occur in pairs. The terminal segment carries two
medially curved spines or claws, which are subequal in size and
with long, spine-like endites generally devoid of 'auxiliary' spines; similar to the outer (lateral) spine of the adjacent segment.
endite of 4th segment stout and exceptionally long, one-third to In the grasping appendages preserved in pairs there are plate-
nearly half as long as the length of the appendage. shaped structures between the two pairs of proximal segments.
The plates are elongate and at the posterior end they were possi-
bly attached to the head. The holotype (Figs. 14A and 15A) is
Amplectobelua symbrachiata sp.n. three-dimensionally preserved and exhibits a large plate under-
Figs. 14-15 neath the elongate plates. Posterior to these plates there is an oval,
convex structure and a large, transverse plate. Further back, there
Synonymy. -1994 New anomalocaridid animal 2 from Chengjiang is a pair of large, oval plates which seem to belong to the animal.
- Chen et al., p. 1305, figs. 3-4. A convex, striated structure, consisting of a series of narrow
sclerites and a wide sclerite, is present posterior to one of the
Etymology. — Greek syn, together, and brachion, arm, referring appendages in the holotype. It is similar to the mouth oí Anoma-
to the preservation of the grasping appendages in pairs. locaris, Peytoia, and Parapeytoia, and it probably represents an
oral circlet
Holotype. - A pair of nearly complete grasping appendages,
NIGPAS Cat. No. 115346 (part only; Figs. 14A and 15A). Remarks. - The reconstruction of the grasping appendages (Fig.
15E) is based on two well-preserved pairs. It appears likely that
Other material. - NIGPAS 115347 (Figs. 14B and 15B) from the endites were directed medially and faced each other. As noted
coll. XL1, a pair of nearly complete grasping appendages (part by Chen et al. (1994, p. 1306), the long and stout endite on segment
and counterpart); NIGPAS 115348 (Figs. 14C and 15C) from 4 could interact with the endites on the distal segments. There is
coll. DJ1, an isolated grasping appendage (part and counterpart); no evidence that the endites were directed downwards as in the
NIGPAS 115349 (Fig. 14D) from coll. M3, an incomplete reconstructions of Anomalocaris (e.g. Gore 1993, pp. 122-123;
grasping appendage (part only); NIGPAS 115350 (Figs. 14E and Briggs 1994, lower figure on p. 1283).
15D) from coll. M3, a complete grasping appendage (part and The striation on the appendage flaps of Amplectobelua sym-
counterpart); numerous incomplete grasping appendages (un- brachiata is similar to that of Anomalocaris saron, but there ap-
numbered) from coll. M2 and M3. pear to be fewer Unes in the former species, at least on the ante-
rior flaps. Thus, there are 18-20 parallel lines on the anterior
Type locality and type horizon. - Maotianshan (Maotian Hill) in flaps in A. saron (Fig. 5), but less than 15 subparallel lines on the
Chengjiang County; level M2, lower part of the Eoredlichia Zone anterior flaps in À symbrachiata (see Chen et al. 1994, fig. 3B).
of the Lower Cambrian Qiongzhusi Formation. In A. saron, three of the parallel Unes are cut by the oblique line
 172 Hou et al.: Anomalocaris and other large animals in the Chengjiang fauna, China GFF 117 (1995)
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 GFF117(1995) Hou et al: Anomalocaris and other large animals in the Chengjiang fauna, China
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marking the boundary between the striated and smooth fields at
an angle of about 20°. In A. symbrachiata at least seven of the
subparallel lines are cut by an oblique line at an angle of about
50°.
Both Anomalocaris canadensis and Peytoia nathorsti carried a
Peytoia-Uke mouth (Whittington & Briggs 1985). A closely com-
parable mouth was also present in Anomalocaris saron sp.n. and
Parapeytoia yunnanensis gen. et sp.n., and probably also in
Amplectobelua symbrachiata. A specimen of A. symbrachiata,
173
Fig. 10. Parapeytoia yunnanensis gen. et sp.n.
Drawing of holotype (cf. Fig. 9). Composite
view of part and counterpart. Numbers 1-8, ap-
pendages, starting with the grasping append-
ages; m, mouth of peytoiid type; s, medial
sternite; sc, lanceolate scales characteristical-
ly placed in rows over the back in anoma-
locaridids. The compressed mouth is probably
preserved in its original position, directed back-
wards as in Opabinia (see Whittington 1975).
illustrated by Chen et al. (1994, fig. 3B) as "new anomalocaridid
animal 2", is associated with sets of spines or teeth. Each set con-
sists of at least six spines, and they were interpreted as belonging
to the jaw apparatus of the species under discussion (Chen et al.
1994). The material described herein suggests, however, that the
mouth of the Chengjiang anomalocaridids is similar to that of
Peytoia nathorsti, and, as noted in the introductory chaper, the
spiny sclerites do not seem to belong to an anomalocaridid.
 174 Hou et al: Anomalocaris and other large animals in the Chengjiang fauna, China GFF117 (1995)

^ ^ * nX* ~fy 11. * S ^ ^ " ^ ï&^ îïB^kiiJlCjLÎM'*** ^ * . "^f • „ ^fr —S .. ^ * .L «*••*( . 1.Í ^
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t» _•** : - . i " < V •** *v 5 * 'J

D
F/^. 7i. Parapeytoia yunnanensis gen. et sp.n. A. Proximal-medial portion of appendage showing five gnathobases and striation close to the
attachment margin. NIGPAS 115335. x2.5. B-C. Isolated grasping appendage (part and counterpart). Coll. No. M3. NIGPAS 115344. x2.5. D.
Isolated appendage, probably from the posterior part of the animal. Coll. No. DJl. NIGPAS 115345. x2.

Genus Cucumericrus gen.n. Diagnosis. - Anomalocaridids with at least five rounded gnatho-
c.. , T .. . , . , . . bases on propod; proximal part of ramipod with poorly defined
S r h f ' " H CMC"7Wf " U T ' C
™ ' S> ng
segmentation, distal part distinctly segmented; up£er ('anterior')
to the shape and ornament of the legs. partof appendage flap distinctly süiaíd with fairry widely spaced
Type and only known species. — Cucumericrus decoratus sp.n.
 GFF117(1995) Hou et al: Anomalocaris and other large animals in the Chengjiang fauna, China 175
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Fig. 12. Parapeytoia yunnanensis gen. et sp.n. A-B. Drawings of part and counterpart of grasp- Fig. 13. Parapeytoia yunnanensis gen et sp.n.
ing appendage shown in Fig. 11B-C. C. Composite drawing of grasping appendage shown in Fig. Drawing of specimen shown in Fig. 11 A; proxi-
11B-C. D. Drawing of trunk appendage shown in Fig. 1 ID; rp, ramipod; pp, propod; af, append- mal-medial portion of appendage with typical
age flap. striation close to the attachment margin (at) of
the propod (pp), five gnathobases (gnl-gn5),
and the basal part of the ramipod (rp). Most of
the appendage flap is missing.

The propod has at least 5 rounded gnathobases without spines.

Cucumericrus decoratus sp.n.
Fig. 16
Etymology. —Latin decorus, beautiful, referring to the beautifully
wrinkled surface of the proximal part of the ramipod and to the
wrinkled skin of the body.
Holotype. - NIGPAS Cat. No. 115352 from coll. No. M3, a
specimen consisting of a series of trunk appendages from the right
side of the body, and patches of wrinkled skin and of dorsal scales
(part and counterpart; Fig. 16B, D, E).
Other material. - NIGPAS Cat. No. 115351 from coll. No. M2,
an isolated ramipod (part and counterpart; Fig. 16A).
Type locality and type horizon. - Maotianshan (Maotian Hill) in
Chengjiang County; lower part of the Eoredlichia Zone of the
Lower Cambrian Qiorigzhusi Formation.
Diagnosis. - As for genus.
Description. - The appendages consist of a fused multisegmented
propod extending into a wide lateral flap and a leglike ramipod.
The upper ('anterior') part of the appendage flap is distinctly
striated. There are about 10 lines on the anterior flap. These are
fairly widely spaced and trend roughly transversely. Posteriorly,
four lines are cut by an oblique line. The ramipod tapers regularly
distally. It is distinctly segmented only distally, whereas the
surface is irregularly wrinkled in the proximal part.
The specimen shown in Fig. 16B has low relief and a colour
pattern, which makes it very difficult to illustrate photographi-
cally. However, it is most important in demonstrating the pres-
ence of soft skin over the back, under the dorsal scales. Some
skin fragments may also stem from the ventral side of the animal.
The skin is ornamented (Fig. 16D-E). Parts are tightly striated
with about ten narrow furrows in one millimetre. In other parts
the furrows are more widely spaced, leaving about 0.5 mm wide,
elongate areas between the furrows. Such elongate areas may be
flat, but occasionally have central thickenings. The animal thus
is fundamentally soft-bodied, devoid of sclerite plates, but pro-
vided with tranverse sets of dorsal scales and with stiff ventral
appendages.
Remarks. - Virtually only the appendages are known of Cucum-
ericrus decoratus, which makes it difficult to compare it with
other species. The appendage flap is broadly similar to that of
 176 Hou et al: Anomalocaris and other large animals in the Chengjiang fauna, China GFF117(1995)
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J
^Wfc«n*«mrií »^ — ... J * ^ . — »*y irfV"- ^"TiaftM>/'1J**"fii •***-*^-&& ^*Z£:..&*,'£M1¡
 GFF 117(1995) Hou et al: Anomalocaris and other large animals in the Chengjiang fauna, China \11

Fig. 14. Amplectobelua symbrachiata gen. et sp.n. A. Holotype, a pair of nearly complete grasping appendages. Coll. No. M2. NIGPAS 115346.
x2.5. B. A pair of nearly complete grasping appendages. Coll. No. XL1. NIGPAS 115347. x2.5. C. Nearly complete grasping appendage with
endites in pairs. Coll. No. DJl. NIGPAS 115348. x3. D. Incomplete grasping appendage showing the long and stout endite on fourth segment. Coll.
No. M3. NIGPAS 115349. x2.5. E. Nearly complete grasping appendage. Coll. No. M3. NIGPAS 115350. x6.
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Fig. 75. Amplectobelua symbrachiata gen. et sp.n. A. Drawing of holotype (cf. Fig. 14A). B. Drawing of grasping appendages shown in Fig. 14B.
C. Drawing of grasping appendage shown in Fig. 14C. D. Drawing of nearly complete grasping appendage shown in Fig. 14E. E. Reconstruction of
grasping appendages.
 178 Hou et al.: Anomalocaris and other large animals in the Chengjiang fauna, China
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1cm
rp
c & Briggs 1985).
Fig. 16. Cucumericrus decoratus gen. et sp.n. A. Detached ramipod.
Coll. No. M2. NIGPAS 115351. B. Simplified drawing of holotype show-
ing a series of striated appendage flaps (af), patches of dorsal scales (sc),
and an appendage. The latter shows a propod (pp) with four complete
gnathobases (gn), and a ramipod (rp). Only the left part of the specimen
is shown. Coll. No. M3. NIGPAS 115352. C. Reconstruction of append-
age. D-E. Ornamentation of the skin of the body of the holotype. D xl6,
Ex8.
GFF117 (1995)
Anomalocaris saron sp.n. and Amplectobelua symbrachiata gen.
et sp. n., but the propod endites and the ramipod are too different
from their grasping appendages to suggest any identity. In ad-
dition, C. decoratus has a distinctive striation on the appendage
flaps. In the anterior flaps of A. saron there are 18-20 lines, which
are parallel and evenly spaced. Posteriorly, three of these lines
are cut by an oblique line. In A. symbrachiata there are less than
15 subparallel lines on the anterior flaps, and posteriorly at least
seven of these are cut by an oblique Une. In C. decoratus there
are about 10 fairly widely spaced Unes on the anterior flaps. Four
of these are cut by an oblique Une, behind which there is an
additional short Une that is absent in A. saron andA symbrachiata.
Family Opabiniidae Walcott, 1912
Genus Opabinia Walcott, 1912
Opabinia regalis Walcott, 1912
Remarks. - This Burgess shale species is included for comparison.
It has been thoroughly described by Whittington (1975). Some
structures were reinterpreted by Bergström (1986,1987). Opab-
inia differs from anomalocaridids in having a long shaft of the
grasping appendages and in having pleural extensions of tergites
overlapping backwards through much of the body. However, there
are also striking similarities which convincingly show that Opab-
inia and anomalocaridids are closely related (Bergström 1986,
1987; Dzik & Lendzion 1988; Chen et al. 1994). Common
characters include the presence of segmentally arranged transverse
sets of scales over the whole width of the dorsum, a simple
intestinal canal extending from a ventral mouth to a terminal anus,
an interconnected pair of preoral grasping appendages, in Opab-
inia set on a shared flexible shaft, and probably a ring of sclerites
surrounding the mouth which is directed backwards (for Opabinia,
see Whittington 1975, figs. 4, 7, 55, 59). The appendages of the
body are not clearly seen in Opabinia specimens, which is proof
of nothing: we must remember that it has been considered a fact
that Anomalocaris and Peytoia were devoid of legs (Whittington
Interpretation of structures in anomalocaridids
Dorsal side
The presence of segmental tergites is quite clear in Opabinia, but
not in anomalocaridids. The alternation between tergites and sheets
of lanceolate scales is also clear in Opabinia (Bergström 1986).
However, the mode of attachment of the lanceolate scales is not
known, except that they attach along one single transverse line or
axis in each segment. Transverse sets of lanceolate scales are also
present across the dorsal surface in Peytoia nathorsti (see Whit-
tington & Briggs 1985, fig. 34; Bergström 1986; Fig. 17A herein)
and the possibly identical Amiella ornata (see Walcott 191 la, pi.
5, fig. 4; Fig. 17B herein). In the Chengjiang material, lanceolate
scales are seen in Anomalocaris saron, Parapeytoiayunnanensis,
and Cucumericrus decoratus. They are attached at one end only
along transverse Unes in A. saron. In this species several more or
less complete bands of lanceolate sclerites occur together on a
surface without any evidence of associated tergites (Fig. 6). The
undulating course of the bands shows that the axis was not stiff.
 GFF 117(1995) Hou et al.: Anomalocaris and other large animals in the Chengjiang fauna, China
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A ,
Fig. 17. A. Drawing oí Peytoia nathorsti (Walcott, 1911) to show trans-
verse sets of lanceolate scales (sc) on the dorsal surface. Based on
U.S.N.M. 274142, illustrated by Whittington & Briggs (1985, fig. 34).
B. Drawing of holotype of Amiella omata Walcott, 1911, showing two
appendage plates at the top and beneath them five transverse rows of
dorsal scales. A. ornata may be a senior synonym of Peytoia nathorsti,
but the identity is not considered certain. The specimen, USNM 57499,
was illustrated by Walcott (191 \a, pi. 5, fig. 4) and Bruton (1981, figs.
34,38).
This indicates that the axis of attachment was part of a soft body
surface and not of stiff tergites.
Grasping appendages
Grasping appendages are frequently found in associated pairs
despite the low number. This is the case in Amplectobelua
symbrachiata and Parapeytoia yunnanensis. Only in the rare C.
decoratus the grasping appendages are not known. In A. sym-
brachiata and Anomalocaris sawn, the endites of the grasping
appendages were directed medially and faced each other (cf. Chen
etal. 1994, figs. 1B-C.2).
Circumoral ring
This is seen in situ in Parapeytoia yunnanensis (Fig. 18A), and
there is one in a dismembered specimen of Anomalocaris sawn
(Fig. 6). Furthermore, an oral circlet is present posterior to the
grasping appendages in a specimen of Amplectobelua sym-
brachiata. A few isolated specimens have also been found. In
179
Fig. 18. A. Enlargement of area around mouth in holotype of Parapeytoia
yunnanensis gen. et sp.n.; m, oral circlet (peytoiid type mouth); s, me-
dial sternite; 1, portion of serrated endite of grasping appendage; 2-3,
successive limbs. B. Oral circlet of Peytoia nathorsti (Walcott, 1911),
for comparison with the oral circlet of P. yunnanensis. Drawing based
on U.S.N.M. 274155, illustrated by Whittington & Briggs (1985, fig.
73). C. Larva of kinorhynch Haploderas showing distinct segmentation
and peytoiid-hTce sclerites around mouth. Redrawn from Nyholm (1947&).
addition, it should be mentioned that there is indication of the
presence of a circumoral ring in Opabinia regalis (Whittington
1975, figs. 4,7,55,59). The posture of the ring in P. yunnanensis
is as in Opabinia, and is strong indication that the mouth was
directed backwards. Occasional backwards inclination of the
mouth in Peytoia nathorsti indicates a similar arrangement in that
species (see, e.g., Whittington & Briggs 1985, PL 16, fig. 73;
Fig. 18B herein).
Body appendages
These are known from four Chengjiang species. The structure of
the appendages are well-known from Parapeytoia yunnanensis,
where a series of legs are seen more or less in situ. This species
has a high degree of tagmosis. Thus, the legs are completely
different from the grasping appendages in lacking endite blades,
and the two postoral appendages are small and morphologically
unique (Fig. 19). The legs of Cucumericrus decoratus differ from
the others in being irregularly folded and not so clearly segmented
proximally, and no comparison can be made with the grasping
 180 Hou et al.: Anomalocaris and other large animals in the Chengjiang fauna, China
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Fig. 19. Reconstruction oî Parapeytoia yunnanensis gen. et sp. n. in ventral view.
GFF117(1995)
appendages since these are unknown.
The material of P. yunnanensis and C. deco-
ratus convincingly shows that the proximal part
of the trunk appendages extends laterally into
a large flap. This structure was previously de-
scribed as a lateral lobe or "fin" (Whittington
& Briggs 1985). The flap is generally distinctly
striated in the upper ('anterior') part. In P.
yunnanensis the evidence for striation is weak,
but one specimen shows the presence of lines,
at least in the area adjoining the appendage-
body attachment (Figs. 11A and 13).
The anomalocaridid appendage apparently
evolved as a broad flap. A ramipod was ulti-
mately formed on the distal side, and later still
this branch became segmented. This is in sharp
contrast with appendage morphogeny in arthro-
pods. In the most generalized early Cambrian
arthropods, the appendage consists of a single
stout, multisegmented ramus, obviously corre-
sponding to the endopod of derived arthropods.
A thin exopod flap is merely a lateral outgrowth
at the base, technically an exite. Thus, in anoma-
locaridids it is the broad unsegmented flap that
forms the proximal structure, whereas in the
earliest arthropods it is the segmented endopod.
Superficial similarity between the two types
increased with convergent evolution.
Functional morphology
Previous interpretations have been based on the
assumption that anomalocaridids had no se-
rial appendages (Whittington & Briggs 1985;
Briggs 1994). The new finds described herein
prove the ability to form serially repeated
appendages. Despite the find of limbs in anoma-
locaridids, however, the functional morphology
and life habits are still not obvious. We can
judge with some certainty that propulsion was
caused by movements of the limbs. The pres-
ence of a pediform branch (ramipod) suggests
that the animals could walk on the bottom. The
presence of a wide appendage flap makes it hard
to exclude the possibility that they could also
swim.
The mouth appears to have been directed
backwards, as in Opabinia. This means that
anomalocaridids could not attack very large
prey, but only prey that could be brought up
between the legs to the mouth by the grasping
appendages or by the legs. This fits with our
reconstruction of the grasping appendages fac-
ing each other, rather than being curved down-
ward as in previous reconstructions. The new
reconstruction is in accordance with the pos-
ture of the grasping appendages as they are
found in pairs, and it is also a close parallel to
the posture in eurypterids. Presumably the
grasping appendages could be extended for-
wards or backwards through a rocking move-
 GFF117(1995) Hou et al: Anomalocaris and other large animals in the Chengjiang fauna, China 181

oral sclerites
— radial mouth Gastrotricha
— segmentation Tardigrada
segm. appendages Nematoda
I r. medial tergites
— moulting
2-ra Palaeoscolecida
-Nematomorpha
Loricifera
+ + ± + Tardigrada (6)
+ + + + 1 1 +1
+ + + + 1 11

Priapulida
Priapulida (4) Kinorhyncha
Kinorhyncha (4) Kerygmachela
Anomalocaris Opabinia
Lobopodia (4) Anomalocaris
'Arthropoda' (3) Peytoia
Linguatulida (5) Parapeytoia
A B
Fig. 20. A.Character comparison between Anomalocaris and certain extant groups. Numbers within parantheses indicate numbers of characters
shared between these groups and Anomalocaris. B. Possible relationships and phylogeny of anomalocaridids. 1, Eunemathelminthes, mouth termi-
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nal, pharynx muscular, exocuticle, adhesive glands associated with small size; la, Gastrotricha, aquisition of blood cells, cilia usually confined to
two ventral bands, anterior sensory organs, and midgut; lb, loss of ciliation, introduction of ecdysis; lc, Tardigrada, oligomery, 'enterocoely', and
unsegmented lobopodia; Id, Nematoda, narrow body, circular muscles virtually lost; 2, Cephalorhyncha, true proboscis with scalids, exocuticle
with serially repeated lorical scutes; 2a, Palaeoscolecida, posterior hooks, circular muscles löst; 2b, Nematomorpha, adult parasitic, loss of cuticular
scutes; 3, fat body with circular muscles; 3a, Priapulida, true lorica (at least in larvae), circular muscles expanded to complete layer, acquisition of
blood; 3b, Loricifera, miniaturization, loss of complexity; 3c, bilateral symmetry, development of true segmentation, with segments covered by
scutes (sclerites), formation of 'Peytoia'-like sclerite ring surrounding mouth; 3d, Kinorhyncha, miniaturization; 4, Dicephalosomita, moderate to
large size, transverse dorsal rows of scales, lobopodian ventral appendages, first pair modified as grasping appendages; 4a, Kerygmachela, dorsal
axis with distinct morphology; 4b, basal portions of grasping appendages parallel, mouth secondarily directed backwards, surrounded by stiff ring
of sclerites; 4c, Opabinia, basal portions of grasping appendages fused; 4d, Anomalocarididae, appendages segmented, long propod with five
gnathobases and extending laterally into wide flap; 4e, Anomalocaris, grasping appendages with narrow spiny endites; 4f, peytoiids, grasping
appendages with long flat endites except in distal spiny element; 4g, Peytoia, as 4f; 4h, Parapeytoia, distal element of grasping appendage without
the ordinary spines but with long flat endite.

ment at the attachment with the body. The endites and spines of
the legs may have been involved in handling the prey. The con-
struction of the mouth makes it unlikely that it could bite off pieces
of trilobites and other arthropods with a hard exoskeleton, as has
been suggested (e.g., Rudkin 1979; Briggs & Mount 1982;
Babcock & Robison 1989; Babcock 1993). It is more likely that
anomalocaridids fed from worms and other soft-bodied animals.
The endites appear to have been poorly sclerotized and therefore
weak, which indicates poor crushing ability. In detail the arma-
ment of the mouth is different from that of extant aschelminths,
which makes a functional comparison difficult. It may be com-
pared with the echinoid lantern. This structure is used for biting
or nibbling during grazing. The grasping appendages of anoma-
locaridids make it clear that these animals did not feed on sea-
weed but on prey. Still, comparison with the lantern may indicate
biting ability. Given the active swimming ability suggested above,
it is possible that prey was caught during swimming. Alterna-
tively, it could easily be caught on the bottom, which is where
most of the possible prey must have been. Swimming may have
been used for locomotion also in this case. Proceeding beyond
this basic level of interpretation at present would simply be specu-
lation.
Affinities of anomalocaridids
It is difficult to compare anomalocaridids with other animal groups
because of the insufficient knowledge of the former. Only a limited
number of external features can be compared (Fig. 20A). The
mere existence of ventral appendages, but nothing else, may
indicate arthropod affinities (Chen et al. 1993, p. 1308), and from
a functional point of view anomalocaridids were arthropods.
However, the oral circlet with radially arranged sclerites and the
peculiar dorsal rows of scales are alien to arthropods. Even the
trunk appendages with a wide lateral flap may be interpreted as
unique, but that may be a matter of opinion. The development of
a wide, flattened flap probably indicates a modification for
swimming and perhaps for gas exchange.
In the most plesiomorphic Early Cambrian arthropods com-
pound eyes were anteroventral, and they still are so in most crus-
taceans. In anomalocaridids, however, they were dorsal, and ap-
pears to have numbered from two to five.
Key characters of arthropods are segmentation, articulated ap-
pendages, and an exoskeleton. Segmentation of the body has
evolved independently several times in the kingdom Bilateria: in
some Platyhelminthes, in the Tardigrada, Kinorhyncha, Annelida,
Lobopodia, Arthropoda (twice?), the genus Annulonemertes
 182 Hou et al.: Anomalocaris and other large animals in the Chengjiang fauna, China
among the Nemertina, and in the Chordata. The development of
tergites and, in particular, paired appendages may be more or
less connected with the presence of segmentation. Appendages
are widely distributed and have developed in the Tardigrada,
Annelida, Lobopodia, Arthropoda (twice?), and Chordata. An
exoskeleton is typical of the Aschelminthes and Arthropoda. Ani-
mals obviously were evolutionarily very flexible when phyla came
into existence. The presence of segmentation, rudimentary exo-
skeleton and superficially similar limbs therefore cannot be taken
as proof that anomalocaridids belong to the true arthropods. The
strong dissimilarity particularly in the mouth armament, as well
as the cover of soft skin and the lack of both tergites and sternite
plates in anomalocaridids, must be taken seriously as an argu-
ment.
Looking for circular mouth armaments among animals, they
are mostly found among the aschelminth worms, particularly in
the phyla of the Eunemathelrninthes. A special case is the pycno-
gonid proboscis. The two other cases are the echinoderms and
cyclostome fishes. However, while aschelminth and anoma-
locaridid sclerites are clearly ectodermal, the echinoderm lantern
Downloaded by [Yale University Library] at 23:09 18 July 2012
is mesodermal, and the cyclostome dentition is not really compa-
rable. The only real similarity is therefore with aschelminths, in
which the circular arrangement is the rule rather than the excep-
tion.
Accepting this fact by face value, and the fact that aschelminth
phyla have commonly developed exoskeletal sclerites and have
proven their ability to develop segmentation and moulting (Tar-
digrada, Kinorhyncha) and limbs (Tardigrada), the anomalocaridid
combination of characters indicate aschelminth affinities more
than anything else.
This leads to the question of a possible position for anoma-
locaridids within the Aschelminthes. They may or may not be
close to tardigrades or kinorhynchs. Both of these groups have
their mouths armed with radially arranged, circum-oral sclerites.
The circle of placids around the neck in kinorhynchs looks sur-
prisingly similar to the oral circlet of sclerites in anomalocaridids
(Fig. 18C). The placids can close the mouth (Nyholm 1947a, fig.
2, 1947b, figs. 1, 3, 4). Like in Peytoia nathorsti, there may be
broad and narrow sclerites regularly arranged (Nyholm 1947a,
fig. 1). Unlike anomalocaridids but like arthropods, the kinorhynch
cuticle has segmentally arranged sclerites as well as bristles or
spines. The ventral side is divided into a narrow median band
and wide lateral sclerite-covered fields, which is at least a super-
ficial similarity to the division between narrow median sternites
and broad lateral appendage flaps in anomalocaridids. The
kinorhynch body is triangular in cross-section (Nyholm 1947Z»,
pi. 1, fig. 2), as it appears to have been in anomalocaridids. One
difference is the size: kinorhynchs (as well as tardigrades) are
less than 1 mm long. Another is the position of the mouth, which
is terminal in kinorhynchs and other aschelminths. Whatever their
affinities, the posterior orientation of the mouth in anomalocaridids
is an adaptation to the mode of feeding with the aid of append-
ages, as in crustaceans, and is a secondary phenomenon which
could have evolved from the aschelminth condition. The dorsal
scales are alien to arthropods, but find some kind of counterpart
in the lateral spines of kùiorhynchs.
It is therefore tempting to suggest that anomalocaridids are clos-
est to kinorhynchs, although neither group needs to have been
derived from the other (Fig. 20B). Most aschelminths are usually
fairly small and devoid of a vascular system. The large size of
Anomalocaris and related forms indicates that they most prob-
ably must have had a vascular system. This is still compatible
GFF117(1995)
with placing them within the aschelminths, since the largest among
the latter, the priapulids, have also developed a vascular system.
Presumptive anomalocaridids may have involved rocking move-
ment of the paired lateral sternites to improve locomotion on the
bottom. Such sternites may eventually have turned into (the ante-
rior surface of) flat appendages such as those of anomalocaridids.
With the formation of a segmented limb (leg or arm), the food
could be caught and manipulated with the appendages, which
would make it reasonable to have the mouth ventrally instead of
terminally. The introvert function would probably have disap-
peared with the bending of the pharynx. In extant aschelminths
the terminal position of the mouth is also connected with func-
tions. In kinorhynchs, for instance, the introvert is a locomotory
organ.
In summary, anomalocaridids are reminiscent of arthropods in
having ventral appendages. However, they differ from the first
arthropods in the Cambrian (1) in the branching pattern of the
appendages; (2) in having their own evolution of segmentation
of the appendages, starting from a non-segmented level; (3) in
having two to five dorsal compound eyes, while the first arthro-
pods had a pair of ventral eyes; (4) in being soft-skinned despite
the large size; (5) in having transverse rows of long, slender dor-
sal scales covering the back; (6) in having the mouth armed with
a circle of sclerites. They do not fit into the pattern of early ar-
thropod evolution, and we regard it more likely that they repre-
sent an arthropodization event among the aschehninth worms.
Acknowledgements. - We are grateful to Dieter Walossek (Ulm, Germany) for
valuable discussions. Robert R. Hessler (La Jolla, California), Dan-Eric Nilsson
(Lund), and Nils Spjeldnss (Oslo) provided constructive comments on the manu-
script. Christer Erséus (Stockholm) kindly supplied data on some aschelminths.
We also express our gratitude to Uno Samuelson (Stockholm) for producing pho-
tographs and Javier Herbozo (Stockholm) for drawings. Britt Nyberg (Lund) as-
sisted in the drawing of Fig. 1. The Swedish Natural Science Research Council
(NFR) and the Crafoord Fund (Royal Swedish Academy of Sciences) provided
financial support.
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