Science in China Series D: Earth Sciences
© 2007
Science in China Press
Springer-Verlag
New fossil eccoptarthrids (Coleoptera: Curculionoidea)
from the Yixian Formation of western Liaoning, China
LIU Ming & REN Dong
College of Life Science, Capital Normal University, Beijing 100037, China
One new genus and three new species of the weevil family Eccoptarthridae (Curculionoidea), Leptocar
polychaetus gen. et sp. nov., Abrocar macilentus sp. nov., Cretonanophyes punctatus sp. nov., are
described and illustrated. They pertain to the Upper Jurassic or Lower Cretaceous Yixian Formation of
western Liaoning Province, China. The host plants to the living eccoptarthrids and the floras in Yixian
Formation indicate that those archaic eccoptarthrids possibly lived on conifers with a phylogenetic
closeness to Cupressaceae. Besides, the early diversification of eccoptarthrids is discussed: originated in Central or East Asia in the Late Jurassic, spread into Western Europe and South America
during the Early Cretaceous, underwent a dramatic decline after the Early Cretaceous which likely was
caused by competitive pressure and the displacement of their host plants.
fossil, weevil, Eccoptarthridae, new taxa, Late Jurassic, Early Cretaceous, Yixian Formation, Liaoning
Eccoptarthridae is a small weevil family with four extant
genera restricted entirely to Australia and South America[1,2]. It is one of the archaic weevils and can date back
to the Late Jurassic. Till now, thirteen species within ten
fossil genera assigned to the family Eccoptarthridae
have been reported from the worldwide sites, of which
two species are known from the Late Jurassic of Kazakhstan and Mongolia[3,4], two from the Late Jurassic or
Early Cretaceous of China[5], seven from the Early Cre―
taceous of Russian, Spain and Brazil[6 10], one from the
Late Cretaceous New Jersey amber [11], and one from the
Oligocene Baltic amber[1].
The Yixian Formation is of disputed Late Jurassic or
Early Cretaceous age. Its insect fauna is diverse and
abundant, but the fossil weevils are relatively rare. In
total, four fossil weevils [5,12,13] have been reported from
this formation so far. Here we describe one new genus
and three new species obtained from the Yixian Formation of western Liaoning Province, China, which adds
considerably to our knowledge of weevil group in this
formation.
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1 Materials and methods
The specimens were examined under a Leica MZ12.5
dissecting microscope and illustrated with the aid of a
camera lucida attached to the microscope. All the type
specimens studied in this paper are housed in the Key
Lab of Insect Evolution & Environmental Changes,
College of Life Science, Capital Normal University,
Beijing, China (CNU; Ren Dong, Curator).
The following standards were used for characters:
body length, measured from the base of rostrum to the
apex of elytra; body width, measured at the base of
elytra; elytra length, measured from the base of elytra to
the apex; elytra width, measured at the widest of a single
elytron.
Received September 1, 2005; accepted November 3, 2006
doi: 10.1007/s11430-007-0030-z
†
Corresponding author (email: [email protected])
Supported by grants from the National Nature Science Foundation of China (Grant
Nos. 30025006, 30370184 and 30430100), the National Nature Science Foundation
of Beijing (Grant No.5032003), the Scientific Research Key Program (Grant No.
KZ200410028013) and the PHR Project of Beijing Municipal Commission of Education
Sci China Ser D-Earth Sci | May 2007 | vol. 50 | no. 5 | 641-648
2 Systematic paleontology
Order Coleoptera Linnaeus, 1758
Superfamily Curculionoidea Latreille, 1802
Family Eccoptarthridae Arnoldi, 1977
Eccoptarthrini Arnoldi, 1977[3]; Caridae Thompson, 1992[14];
Baissorhynchini Zherikhin, 1993[7]; Caridae Zimmerman, 1994[2];
Eccoptarthridae Zherikhin and Gratshev, 1995[15].
Genus Leptocar gen. nov.
Etymology: The name is derived from the Greek prefix lept (meaning “thin”) and the genus Car (the type
genus of this family), gender masculine.
Type species: Leptocar polychaetus gen. et sp. nov.,
by present designation.
Species included: Only the type species: Leptocar
polychaetus gen.et sp.nov.
Diagnosis: Small, shorter than 5 mm, weakly arched
in lateral view. Rostrum longer than head and pronotum
together, relatively straight, nearly parallel-sided; mandibles narrow, shorter than the apical width of rostrum,
without tooth. Antennal socket at the basal half of rostrum laterally. Head coniform, lacking obvious postocular constriction; frons broad; eyes oval, dorsolaterally
positioned; gena long. Pronotum weakly convex, the
base wider than its apex. Fore coxae large and prominent,
originating in the antemedial region of prothorax. Middle coxae apart from the base of pronotum. Legs long;
femora intermediate in size; the base of fore tibiae
slightly curved; middle and hind tibiae with long setae
on lateral side and two small apical spurs; first tarsal
segment long, without obvious inflation; claws paired
and accrete, without tooth. Elytra not shorter than abdomen, subparallel, broadly rounded at apex.
Comparison: The following features of this weevil
place it in the family Eccoptarthridae: body small, rostrum longer than head and pronotum together, antennal
insertion at the basal half of rostrum, legs long, first tarsal segment long, apex of elytra broadly rounded. Till
now, there have been ten fossil genera within Eccoptarthridae. The new genus is distinct for its less convex
body and relatively straight rostrum. Moreover it can be
distinguished from other known fossil genera as following: from the genera Baissorhynchus Zherikhin, 1977[6],
Gobicar Gratshev and Zherikhin, 1999[4], and Abrocar
Liu and Ren, 2006[5] in the rostrum longer than head and
pronotum combined; from the Eccoptarthrus Arnoldi,
1977[3], Cretonanophyes Zherikhin, 1977[6], Emanrhyn642
chus Zherikhin, 1993[7], Cretocar Gratshev and
Zherikhin, 2000[11], and Martinsnetoa Zherikhin and
Gratshev, 2004[10] in the not inflated first tarsal segment;
from the Baltocar Voss, 1953[1] in lacking postocular
constriction and the medium sized eyes; from the genus
Jarzembowskia Zherikhin and Gratshev, 1997[8] in the
long gena and the base of tibiae without emargination.
So, the comparison with the known fossil genera within
this family indicates that our fossil represents a new genus.
Leptocar polychaetus sp. nov. (Figures 1 and 2)
Figure 1 Leptocar polychaetus gen. et sp. nov., line drawing of holotype,
No. CNU-C-LB2006101.
Material: Holotype CNU-C-LB2006101, an impression of nearly complete weevil in ventrolateral position.
Locality and horizon: Yixian Formation, Huangbanjigou, Chaomidian Village, Beipiao City, Liaoning
Province, China.
Etymology: The name is derived from the Greek
polychaetus (meaning “setose”).
Description: Body dark, weakly arched in lateral
view. Rostrum straight, slightly arched on apical 1/4,
nearly parallel-sided, originating from the mid-height of
head, 8 times longer than its apical width, 1.2 times
longer than head and pronotum together; mandibles
narrow, without tooth, half length of the apical width of
rostrum; labrum visible, probably rectangular. Antennal
socket oval, inserted laterally at 1/4 base of rostrum.
Head coniform, lacking obvious postocular constriction,
its base 3 times longer than the apex and 2 times longer
than the length of head; frons broad; eyes oval, the
length similar to basal width of rostrum, dorsolaterally
positioned, close to the base of rostrum; gena long,
slightly shorter than eye. Pronotum weakly convex, 2.5
times longer than head, the apex slightly wider than the
LIU Ming et al. Sci China Ser D-Earth Sci | May 2007 | vol. 50 | no. 5 | 641-648
Figure 2
Leptocar polychaetus gen. et sp. nov., photograph of holotype, No. CNU-C-LB2006101. (a) Body; (b) head; (c) middle tibiae.
base of head, the length same to its basal width and 1.5
times longer than the apical width, notosternal suture
invisible. Fore coxae coniform, large and prominent,
close to the apical prothorax. Fore femora intermediate
in width; fore tibiae 1.2 times longer than femora, its
base slightly curved, gradually widening towards apex;
tarsi as long as its femora, tarsal ratio 4:2:2:5, first
segment trapeziform, 2 times longer than wide, without
obvious inflation, second segment trapeziform, third
segment bilobate, fourth one narrow and long; claws
long, paired and accrete, without tooth. Middle coxae
oval, slightly longer and thicker than fore ones, apart
from the base of prothorax. Middle femora not clear,
probably in same size with fore ones; its tibiae shorter
than fore tibiae, gradually widening towards apex, with
long setae on lateral side and two small apical spurs.
Hind femora not clear, probably longer and wider than
fore or middle femora; its tibiae similar to middle tibiae.
Middle and hind tarsi and claws similar to fore leg. Abdomen not longer than elytra. Elytra 2.5 times longer
than pronotum, subparallel, broadly rounded at apex.
Measurements (in mm): Body length: 3.3. Rostrum
length: 1.5, width: 0.2; mandible length: 0.08. Head
length: 0.35, eye length: 0.2. Pronotum length: 0.9;
width: base 0.9, apex 0.6. Fore leg length: coxae 0.4;
femora 0.8; tibiae 0.95; tarsi 1―4: 0.2, 0.1, 0.1, 0.25;
claw 0.15. Middle leg length: coxae 0.45, femora 0.8?;
tibiae 0.85?, spur 0.04; tarsi 1―4: 0.2, 0.1, 0.1, 0.25;
claw 0.15. Hind leg length: femora 1?; tibiae 0.9, spur
0.04; tarsi 1―4: 0.2, 0.1, 0.1, 0.25; claw 0.12. Elytra
length: 2.3, width: 1.1.
Genus Abrocar Liu and Ren, 2006
Type species: Abrocar brachyorhinos Liu and Ren,
LIU Ming et al. Sci China Ser D-Earth Sci | May 2007 | vol. 50 | no. 5 | 641-648
643
2006, Upper Jurassic or Lower Cretaceous, Yixian Formation of China.
Species included: Two species: A. brachyorhinos
(type species, Upper Jurassic or Lower Cretaceous,
Yixian Formation of China); the second is the new one
described below.
Diagnosis: Small, shorter than 5 mm. Rostrum parallel-sided, nearly in same length with the head and
pronotum together. Antennae inserted laterally at the
basal half of rostrum, not exceeding hind margin of head,
club distinct. Eyes oval, dorsolaterally positioned. Fore
coxae large and prominent, originating in the antemedial
region of prothorax. Legs long, narrow; femora intermediate in size; tibiae slightly curved at the base; tarsi long,
first tarsal segment without inflation. Elytra not shorter
than abdomen, broadly rounded at apex, with rows of
distinct rounded punctation.
Abrocar macilentus sp. nov. (Figures 3 and 4)
Figure 3 Abrocar macilentus sp. nov., line drawing of holotype, No.
CNU-C-LB2006102.
Material: Holotype CNU-C-LB2006102, an impression of nearly complete weevil in ventrolateral position.
Locality and horizon: Yixian Formation, Huangbanjigou, Chaomidian Village, Beipiao City, Liaoning
Province, China.
Etymology: The name is derived from the Greek
macilentus (meaning “slim”).
Diagnosis: Rostrum strongly curved, originating from
the lower height of head; antennal insertion slightly before the middle rostrum; eyes close to the underside of
head; pronotum short, transverse, the base arcuate; tibiae
with one straight longitudinal carina from the base to the
apex; elytra narrow, punctation medium sized.
Description: Body dark arched in lateral view.
644
Figure 4 Abrocar macilentus sp. nov., photograph of holotype, No.
CNU-C-LB2006102.
Rostrum strongly arched, parallel-sided, originating
from the lower height of head, as long as head and
pronotum together, 6 times longer than its apical width.
Antennal insertion laterally and slightly before the middle rostrum; antenna probably exceeding hind margin of
head; scape longer than other visible segments respectively except terminal one; the visible funicular segments narrow, each segment longer than wide; club distinct, each segment oval, terminal one acuminate at its
apex. Head coniform, lacking postocular constriction;
frons broad, convex; eyes oval, large, longer than the
basal width of rostrum, dorsolaterally positioned, close
to the underside of head; gena much shorter than eyes.
Pronotum gently convex, 1.6 times longer than head; its
base arcuate, 1.3 times longer than the apex, 1.4 times
longer than the length of pronotum; notosternal suture
invisible. Fore femora intermediate in width; fore tibiae
curved at the base, gradually widening towards apex, 1.2
times longer than femora, with one straight longitudinal
carina from the base to the apex; first tarsal segment
long, rectangular, without inflation, second one rectangular. Hind femora broader and longer than fore ones;
tibiae curved, 0.9 times as long as the femora, with one
straight longitudinal carina from the base to nearly apex;
tarsi long, tarsal ration 3:2:2:4, first segment trapeziform, second segment rectangular, third segment bilobate, fourth one narrow and long; claw long, without
tooth. Elytra not shorter than abdomen, 3.6 times longer
than pronotum, narrow, subparallel on basal 3/4, slightly
tapering caudad on apical 1/4, broadly rounded at apex;
punctation rounded, distance between punctations in
rows 4 times longer than their diameter, interstrial
LIU Ming et al. Sci China Ser D-Earth Sci | May 2007 | vol. 50 | no. 5 | 641-648
spaces 6 times longer than the diameter of punctation.
Measurements (in mm): Body length: 2.5. Rostrum
length: 0.8, width: 0.15. Antennal segment 1―11 length:
0.10, ?, ?, ?, 0.07, 0.04, 0.07, 0.05, 0.09, 0.09, 0.14.
Head length: 0.3, eye length: 0.2. Pronotum length: 0.5;
width: base 0.8, apex 0.6. Fore leg length: femora 0.6;
tibiae 0.7; tarsi 1―2: 0.15, 0.1. Hind leg length: femora
0.7; tibiae 0.6; tarsi 1―4: 0.15, 0.1, 0.1, 0.2; claw 0.1.
Elytra length: 1.8, width: 0.7; punctation diameter: 0.01.
Comparison: The following features of this fossil
weevil place it in the family Eccoptarthridae: body small
and convex, rostrum not shorter than head and pronotum
together, antennal insertion near the middle rostrum,
antennal club distinct, legs long, apex of elytra broadly
rounded. We assign this new species to the genus Abrocar by the agreement with the diagnosis of this genus.
The new one can be distinguished from A.
brachyorhinos by the following characters: rostrum
strongly curved, originating from the lower height of
head; antennal insertion slightly before the middle rostrum; eyes oval, large, close to the underside of head;
pronotum short, transverse; tibiae with one straight longitudinal carina from the base to apex; elytra narrow.
Genus Cretonanophyes Zherikhin, 1977
Type species: Cretonanophyes longirostris Zherikhin,
1977, Lower Cretaceous, Transbaikalia of Russia.
Species included: Four species: C. longirostris (type
species, Lower Cretaceous, Transbaikalia of Russia); C.
rugosithorax Gratshev and Zherikhin, 2000 (Lower
Cretaceous, Sierra del Montsec of Spain); C. zherikhini
Liu and Ren, 2006 (Upper Jurassic or Lower Cretaceous,
Yixian Formation of China); the fourth is the new one
described below.
Diagnosis: Body arched in lateral view. Rostrum
nearly parallel-sided, longer than head and pronotum
together. Antennal insertion at the basal half of rostrum,
antenna exceeding hind margin of head; scape long, not
reaching base of rostrum; club distinct. Fore coxae
reaching fore margin of prothorax. Legs long, narrow;
bases of tibiae curved distinctly; first tarsal segment inflated, nearly triangular; claws free, paired, without
tooth.
Cretonanophyes punctatus sp. nov. (Figures 5 and 6)
Material: Holotype CNU-C-LB2006103, an impression of nearly complete weevil in dorsoventral position.
Locality and horizon: Yixian Formation, Huangban-
Figure 5 Cretonanophyes punctatus sp. nov., line drawing of holotype,
No. CNU-C-LB2006103. (a) Dorsal view; (b) ventral view.
jigou, Chaomidian Village, Beipiao City, Liaoning
Province, China.
Etymology: The name is derived from the Greek
punctatus (meaning “punctate”).
Diagnosis: Antennal insertion at 1/4 base of rostrum;
eyes oval, large, close to base of rostrum; fore tibiae
with two small apical spurs; elytra with narrow epipleuron, punctation large.
Description: Rostrum long, 1.1 times longer than
head and pronotum together, nearly parallel-sided, with
weakly widened base and apex, 8 times longer than its
apical width; mandibles narrow, without tooth, half
length of the apical width of rostrum. Antennal socket
oval, inserted laterally at 1/4 base of rostrum, reaching
middle prothorax; terminal funicular segment narrow,
longer than wide; club distinct, loose, first two segments
rounded, terminal one longest, oval, acuminate at its
LIU Ming et al. Sci China Ser D-Earth Sci | May 2007 | vol. 50 | no. 5 | 641-648
645
Figure 6
Cretonanophyes punctatus sp. nov., photograph of holotype, No. CNU-C-LB2006103. (a) Body; (b) head; (c) fore tarsi.
apex. Head small, coniform, with slight postocular constriction; eyes large, longer than basal width of rostrum,
dorsolaterally positioned, close to the base of rostrum;
gena much shorter than eye. Pronotum trapeziform, with
slightly rounded lateral sides; 2 times longer than head;
its base 1.8 times longer than its apex, 1.4 times longer
than length of pronotum. Fore femora intermediate in
width; fore tibiae narrow, widening sharply on apical 1/3,
longer than femora, left tibiae with two small apical
spurs, right tibiae with one visible apical spur; fore tarsi
0.8 times longer than the tibiae, tarsal ration 2:1:1:2,
first tarsal segment inflated, trapeziform, 2 times wider
than second one, third segment bilobate, fourth one narrow; claws large, paired and free, without tooth. Middle
coxae oval, apart from the base of prothorax, apart from
646
each other; middle femora in same width with fore ones;
tibiae 0.9 times as long as fore ones; tarsi and claws
similar to fore ones. Hind coxae transverse. Abdomen
with five visible sternites, first sternite longest, last four
ones nearly in same length. Elytra slightly longer than
abdomen, 3 times longer than pronotum, subparallel on
basal 4/5, tapering caudad on apical 1/5, its apex somewhat pointed; epipleuron on right elytron visible, narrow,
extending to 4/5 of elytron; left elytron slightly longer
than right one (might be the result of taphonomic factors); punctation rounded, distance between punctations
in rows slightly longer than their diameter, interstrial
spaces 4 times longer than the diameter of punctation.
Measurements (in mm): Body length: 3.2, width:
1.1. Rostrum length: 1.25, width: 0.15; mandible length:
LIU Ming et al. Sci China Ser D-Earth Sci | May 2007 | vol. 50 | no. 5 | 641-648
0.08. Antennal segment 8―11 lengths: 0.08, 0.10, 0.10,
0.16. Head length: 0.4, eye length: 0.26. Pronotum
length: 0.75; width: base 1.1, apex 0.6. Fore leg length:
femora ?; tibiae 0.9, spur 0.02; tarsi 1―4: 0.2, 0.1, 0.1,
0.2; claw 0.1. Middle leg length: coxae 0.4; femora 0.6;
tibiae 0.8; tarsi 1―4: 0.2, 0.1, 0.1, 0.2; claw 0.1. Hind
coxae length: 0.65, width: 0.3. Abdomen length: 2.0,
width: 1.5. Elytra length: 2.3, width: 0.8; punctation
diameter: 0.02.
Comparison: The following features of this weevil
indicate an affinity with the family Eccoptarthridae:
body small, the rostrum distinctly longer than head and
pronotum combined, antennal insertion at the basal part
of rostrum. We assign this new species to the genus
Cretonanophyes by the agreement with diagnosis of this
genus. Till now there are three species within this genus.
The new species is distinct for its relatively shorter rostrum and two small apical spurs on fore tibiae. Moreover,
the new one can be distinguished from C. longirostris by
the smaller and oval eyes, longer gena; from C.
rugosithorax by its antennal insertion, oval eyes, long
claws; from C. zherikhini by the smaller body, smaller
head and longer pronotum.
nated by conifers and cycadophytes[16], which indicate
that the eccoptarthrids from the Yixian Formation possibly lived on some conifers with a phylogenetic closeness
to Cupressaceae.
Until now the doubtless Late Jurassic eccoptarthrids
were collected from the Karatau Range of South Kazakhstan and Shar-Teg of Mongolia. Two eccoptarthrids
were collected from the Yixian Formation of disputed
Late Jurassic or Early Cretaceous age. These three sites
are relatively close to each other and shared the same
subtropical and seasonal arid or semi-arid climate[16,17].
It suggests that eccoptarthrids may have originated in
Central or East Asia in the Late Jurassic and subsequently spread into Western Europe and South America
during the Early Cretaceous. The Early Cretaceous has
been claimed to be an important time in the evolutionary
history of eccoptarthrids because of their worldwide
distribution and relatively high proportion among the
weevils. After the Early Cretaceous the eccoptarthrids
underwent a dramatic decline from the fossil record.
Competitive pressure from new weevils and the displacement of their host plants during the angiosperm
floristic “radiation” were likely to have contributed to
this.
3 Discussion
The extant eccoptarthrids are associated with Cupressaceae[2], however, the certain members of this plant
family did not appear until the Late Cretaceous. The
elements of the floras in Yixian Formation were domi-
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3
We sincerely thank Professor Zhang Runzhi (Institute of Zoology, Chinese
Academy of Sciences) and Dr. Conrad Labandeira (Smithsonian Institution, National Museum of Natural History, USA) for their endowment of
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