Post on 26-Jan-2023
COPTOCLAVID BEETLES (COLEOPTERA: ADEPHAGA)
FROM THE LOWER CRETACEOUS OF SPAIN: A NEW
FEEDING STRATEGY IN BEETLES
by CARMEN SORIANO* , ALEXANDR G. PONOMARENKO� and
XAVIER DELCLOS**Departament d’Estratigrafia, Palaeontologia i Geociencies Marines, Universitat de Barcelona, Spain; e-mail: carmen.soriano@gmail.com; xdelclos@ub.edu
�Paleontological Institute, Russian Academy of Sciences, Moscow, Russia; e-mail: aponom@palaeo.ru
Typescript received 15 July 2005; accepted in revised form 23 May 2006
Abstract: Currently the beetle family Coptoclavidae consists
of four subfamilies known from the Upper Triassic–Lower
Cretaceous (Aptian). We describe two new subfamilies, three
new genera and five new species from the Las Hoyas (Cuenca
Province) and El Montsec (Lleida Province) localities of Bar-
remian (Early Cretaceous) age from Spain: the first new sub-
family, Hispanoclavinae, is from Las Hoyas and comprises
one new genus, Hispanoclavina, and two new species,
H. diazromerali and H. gratshevi; the second, Coptoclavisci-
nae, is represented at El Montsec by one new species of Cop-
toclavella (C. inexpecta). In addition, at Las Hoyas the
subfamilies Necronectinae and Coptoclavinae are each repre-
sented by one new genus and species: Ovonectes pilosum and
Hoyaclava buscalionae, respectively. All beetles previously
assigned to the family Coptoclavidae have been interpreted
as active hunters. However, we consider Hispanoclavina and
Hoyaclava to be filter-feeding, with forelegs adapted for fil-
tering plankton (probably zooplankton because other mem-
bers of the family are carnivorous) on and beneath the water
surface. This represents a new feeding strategy in beetles
(extant and extinct). The five new species extend the known
geographical distribution of the Coptoclavidae into the west-
ern part of European Barremian deposits. The Las Hoyas
locality now has the highest known diversity of coptoclavid
species.
Key words: Coleoptera, Coptoclavidae, Lower Cretaceous,
planktonic filter-feeding, Spain.
During the last 30 years the Lower Cretaceous (Barre-
mian) localities of Las Hoyas and La Cabrua, commonly
referred to as El Montsec, have yielded thousands of fossil
insect specimens. Most of the insect groups in these col-
lections have been thoroughly studied. At both localities
beetles exhibit the highest diversity, with more than 70
different morphotypes having been identified to date
(Soriano and Delclos 2005), and 32 species from nine
families described (Gomez-Pallerola 1979; Whalley and
Jarzembowski 1985; Alexeev 1993; Zherikhin and Grat-
shev 1997; Gratshev and Zherikhin 2000; Ponomarenko
and Martınez-Delclos 2000; Zherikhin 20031 ; Soriano and
Delclos 2006; Soriano et al. 2006, in press).
Early Cretaceous beetle faunas were composed of a
mixture of relict (Jurassic) groups and modern types
(2 Ponomarenko 1995). This is clear from the composition
of the aquatic beetle faunas, which are typically made up
of coptoclavids (usually the predominant group), dytisc-
ids, gyrinids and hydrophilids. In both Spanish localities
these Jurassic and Early Cretaceous groups are usually
found together in the same outcrops (Soriano and Delclos
2005). This is probably a result of the isolation of the
Iberian Plate during most of the Mesozoic Era.
To date, only two species of coptoclavid have been des-
cribed from these localities (Ponomarenko and Martınez-
Delclos 2000): Megacoptoclava longiurogomphia from Las
Hoyas and Bolbonectus lithographicus from El Montsec,
both in the subfamily Coptoclavinae. This paper describes
new specimens of coptoclavids from both localities.
Coptoclavids have been recorded from Rhaetian–Sin-
emurian deposits in England, the Toarcian of Germany,
the Barremian of Spain and the Aptian of Mongolia
(Ponomarenko 1961, 1987; Ponomarenko and Martınez-
Delclos 2000). Surprisingly, there are no cited records of
this family from the Americas. The family occurs in the
fossil record as larval stages and adult forms. Adults are
usually large predatory beetles (3 Ponomarenko 1995).
They exhibit a general morphological resemblance to
P A L A 6 4 2 B Dispatch: 8.1.07 Journal: PALA CE: Blackwell
Journal Name Manuscript No. Author Received: No. of pages: 12 PE: Raymond
[Palaeontology, Vol. 50, Part 2, 2007, pp. 1–12]
The Palaeontological Association 1
dytiscids (large bodies with raptorial first legs, the sec-
ond and third pairs used for swimming, in some cases
with long natatory hairs), but also show some similarit-
ies to whirligig beetles in terms of both morphology and
ecology (two pairs of eyes adapted for aerial and aquatic
vision, in some cases with oar-shaped second and final
pairs of legs adapted for swimming), although coptoclav-
ids are interpreted as less specialized swimmers than gyr-
inids (Ponomarenko 1961). The larvae are typical of
dytiscoid forms: aquatic predators with well-developed
meso- and metathoracic legs for swimming, and raptor-
like forelegs. Members of this family are interpreted as
hunters of other insects, or even small fishes (Pon-
omarenko 1961).
The weak sclerotization of the ventral part of the body
in coptoclavids, as found in modern gyrinids or dytiscids,
makes it possible to analyse some internal structures of
these fossil beetles, including dorsal structures such as the
mesonotum and metanotum, and the insertion of hind
wings and genitalia (Ponomarenko 1961). This is partic-
ularly clear in Hoyaclava buscalionae gen. et sp. nov. It is
advantageous because it allows us to study structures not
commonly preserved in fossils, but is disadvantageous as
it may also mask the true ventral structures.
The phylogenetic position of this extinct family with
respect to other adephagan families remains unclear, par-
ticularly because of the difficulties in determining the true
synapomorphies of the group and the polarization of
some characters. Nevertheless, some analyses based on
adult coptoclavids and extant beetles suggest that they
were related to gyrinids (Balke et al. 2003). Although
most of the characters used in the study of extant forms
are not preserved in fossils, it is necessary to carry out a
detailed phylogenetic analysis using both the larval and
the adult stages of fossil and recent forms in order to
clarify this issue.
GEOLOGICAL SETTING
The Las Hoyas site is located in the Serranıa de Cuenca
(Cuenca Province) and the La Cabrua site in the El
Montsec Mountains (Lleida Province). Both consist of
laminated mudstones formed by deposition of carbona-
ceous muds in lakes of varying depth (El Montsec) or in
a wetland ecosystem (Las Hoyas) (Fregenal-Martınez and
Melendez 1995; Buscalioni and Fregenal 2003). These
inundated areas developed in Early Cretaceous grabens
that formed during an important stage of rifting that
affected the Iberian Peninsula. Particularly during the
Barremian, sedimentation in these depressed areas was
dominated by the accumulation of lacustrine ⁄palustrine
carbonates and marls, associated with palaeosols (Frege-
nal-Martınez 2000)4 in a subtropical, semi-arid climate
with alternating wet and dry seasons (Fregenal-Martınez
and Melendez 1995).
Aquatic insect association. A comparison of the sites at
Las Hoyas and La Cabrua reveals differences between
their aquatic palaeoentomological content. At Las Hoyas
aquatic insects are predominantly belostomid heteropter-
ans (particularly the single species Iberonepa romerali, rep-
resented by four juveniles and one adult) with a
nektobenthonic habit (Martınez-Delclos et al. 1995). At
La Cabrua the most common aquatic insect is Mesopalin-
gea leridae, an ephemeropteran larva possibly with a
benthonic burrowing habit (Whalley and Jarzembowski
1985; Martınez-Delclos 1991a).
In the nektobenthonic zone of Las Hoyas coptoclavids
shared the ecological space with belostomids, with legs
having different morphofunctional structures compared
with Iberonepa romerali (Martınez-Delclos 1991a; Martı-
nez-Delclos et al. 1995). The benthos was dominated by
the crayfish Austropotamobius, but with a few representa-
tives of unclassified odonatan and trichopteran larvae,
small gastropods and bivalves, ostracods, peracarids
(Delclos et al. 2004)5 , and coptoclavid (Megacoptoclava
longiurogomphia, a medium-sized larva morphologically
convergent with plecopteran larvae, with raptor-like fore-
legs and cursorial meso- and metathoracic legs), dytiscid
and gyrinid coleopterans (Soriano and Delclos 2003,
2004a). Three detritivorous mayfly families (Potaman-
thidae, Euthyplociidae and Leptophlebiidae) are also
represented (Martınez-Delclos 1991a). Species of Euthyp-
lociidae and Potamanthidae were burrowers, both with
forelegs that are morphologically similar to those of the
members of Hispanoclavinae subfam. nov. described
herein, whereas representatives of the Leptophlebiidae were
crawlers on the bottom and climbers amid vegetation.
The pleuston was dominated in the water by the cop-
toclavid genera Hispanoclavina gen. nov., Hoyaclava gen.
nov., Ovonectes gen. nov. and Coptoclavella Ponomarenko,
1980 (Soriano and Delclos 2003), and by chresmodids on
the water-surface (Martınez-Delclos 1989; Nel et al.
2005), whereas the plankton seems to have been domin-
ated by dipteran chaoborid-like and chironomid-like
pupae (Delclos et al. 2005), although they are not well
enough preserved to allow detailed study. In the nektob-
enthonic zone, coptoclavids shared the ecological space
with other very small aquatic heteropterans (at La Cab-
rua) and with caridean decapods such as Delclosia. As at
Las Hoyas, the benthos was dominated by the crayfish
Austropotamobius, Mesopalingea and two mayfly families
(Euthyplociidae and Leptophlebiidae), the most com-
monly found aquatic insects, with several ontogenetic sta-
ges represented. The benthos also included representatives
of anisopteran-like odonatans (Palaeaeschna vidali), small
gastropods and bivalves, ostracods, and some unidentified
2 PALAEONTOLOGY, VOLUME 50
polyphagan coleopterans (Martınez-Delclos 1991b; Sori-
ano and Delclos 2004b).
None of this group of aquatic coleopterans is found in
the nearby Spanish Lower Cretaceous amber localities
(Alonso et al. 2000; Perrichot 2004).
MATERIAL AND METHODS
Material from the Las Hoyas outcrop is housed in the
MCCM (Museo de las Ciencias de Castilla La Mancha),
Cuenca, Spain (LH, Las Hoyas collection). Material from
the La Cabrua outcrop is housed in the IEI (Institut d’Es-
tudis Ilerdencs), Lleida, Spain (LC, La Cabrua collection).
All measurements are in mm. Drawings were made under
incident light with a camera lucida attached to a Leica
MS5 stereomicroscope. Photographs were taken with a
Nikon Coolpix 4500 digital camera attached to a Leica
MS5 stereomicroscope.
SYSTEMATIC PALAEONTOLOGY
Order COLEOPTERA Linnaeus, 1758
Suborder ADEPHAGA Emery, 1886
Family COPTOCLAVIDAE Ponomarenko, 1961
Type species. Coptoclava longipoda Ping, 1926, Lower Cretaceous,
China, Siberia and Mongolia.
Diagnosis (sensu Ponomarenko 1961). Large to medium-
sized aquatic beetles. Head with divided eyes, with dorsal
and ventral part. Metepisternum not reaching mesocoxae.
Metacoxae not broadened anteriorly, transverse metaster-
nal suture absent. Meso- and metathoracic legs modified
for swimming, tibiae broadened.
Composition. Five subfamilies: Charanoscaphinae, Upper Juras-
sic, Kazakhstan; Hispanoclavinae subfam. nov., Lower Creta-
ceous, Spain; Coptoclavinae, Upper Jurassic–Lower Cretaceous,
China, Russia, Spain, Angola; Necronectinae, Jurassic, Mongolia,
Russia, Kazakhstan, Kyrgyzstan, Germany, Algeria (Lower Creta-
ceous, Spain); Coptoclaviscinae subfam. nov., Lower Cretaceous,
Russia, Mongolia, England, Spain.
Subfamily CHARANOSCAPHINAE Ponomarenko, 1977
Type species. Charanoscapha Ponomarenko, 1977, Upper Juras-
sic, South Kazakhstan.
Diagnosis. Large to medium-sized beetles. Metacoxae
without femoral plates. Metatibiae broadened and flat-
tened, tarsi flattened but not broadened, narrower than
tibiae. Metatibiae and tarsi with long natatory hairs.
Composition. Charanoscapha grossa Ponomarenko, 1977;
C. ovata Ponomarenko, 1977; Charanoscaphidia elongata Pono-
marenko, 1977; Upper Jurassic, South Kazakhstan.
Subfamily HISPANOCLAVINAE subfam. nov.
Type genus. Hispanoclavina gen. nov., Barremian, Las Hoyas,
Spain.
Diagnosis. Body large. Dorsal eyes anterior to ventral eyes.
Pronotum comparatively narrow and long. Metacoxae
very large, about 3–4 times as long as wide, with long
femoral plates. Elytra densely striated, with 12–18 striae
on each. Meso- and metathoracic legs with hairs on tibiae
and tarsomeres. Profemora and protibiae with very robust,
large, dense bristles on exterior margin. Meso- and meta-
thoracic legs with natatory hairs on femur, tibia and tarsi;
tibiae not wider than femora; tarsi narrower than tibiae.
Composition. Hispanoclavina diazromerali gen. et sp. nov. and
H. gratshevi gen. et sp. nov.
Genus HISPANOCLAVINA gen. nov.
Derivation of name. Hispania, the Roman name for Spain and
family name.
Type species. Hispanoclavina diazromerali sp. nov., Barremian,
Las Hoyas, Spain.
Diagnosis. As for subfamily.
Description. Head as long as wide, dorsal eyes larger than vent-
ral. Procoxae quite short, prosternal process c. 1Æ5 times as long
as procoxae. Mesosternum shorter than metasternum, with deep
medial fossa to receive prosternal process. Mesocoxae suboval.
Metacoxae very large, c. 3–4 times as long as wide; bilobed at
base; femoral plates large. Elytra widest in first third, then
roundly narrowing to anterior; propleura quite narrow. Elytra
with 12–18 dense striae, not joined at the apex. Profemora and
protibiae elongated, with a row of very robust, large, dense bris-
tles on exterior margin, denser and longer on tibiae. Meso- and
metathoracic legs with natatory hairs on femur, tibiae and tarsi;
tibiae not broader than femora; tarsi narrower than tibiae.
Hispanoclavina diazromerali sp. nov.
Plate 1, figure 1; Text-figure 1
Derivation of name. After Mr Armando Dıaz-Romeral, Cuenca,
Spain.
Type specimens. LH16264, part and counterpart of a beetle lack-
ing antennae and protarsi (Text-fig. 1A), and paratype,
SORIANO ET AL . : EARLY CRETACEOUS COPTOCLAVID BEETLES FROM SPAIN 3
LH23856, only part of an impression of a beetle lacking anten-
nae and distal part of legs (Text-fig. 1B).
Type locality and horizon. Las Hoyas fossil site, close to the
village of La Cierva, Serranıa de Cuenca, Cuenca Province;
second lithosome of finely laminated limestones of the La
Huerguina Formation, Barremian (Fregenal-Martınez and
Melendez 2000).
Other material. Same locality: LH15070, 16315, 22299, 23859.
Diagnosis. Body elongated. Profemora wider than meso-
and metafemora. Metacoxae longer than femoral plates.
Elytra with 15–18 striae.
Description. Total length 31Æ0–31Æ8 mm; maximum width 12Æ2–
12Æ9 mm; elytra length 22Æ4 mm. Body comparatively large and
elongated. Head transverse, about 1Æ4 times as wide as long.
Dorsal eyes large, somewhat larger than ventral eyes (Text-
fig. 1), inserted laterally and anterior to ventral eyes. Pronotum
quite long, c. 1Æ3 times as long as wide, widest at middle and
roundly narrowed to anterior and posterior. Prosternum about
twice as long as wide, length before procoxae much greater than
length of procoxae. Procoxae comparatively large and suboval,
about twice as long as wide; prosternal process narrow, about
half as wide as procoxae, and extended behind them. Mesoster-
num short, c. 0Æ75 times as long as prosternum, and with deep
medial fossa in last third. Mesocoxae large and rounded, with
quite large trocanter. Metasternum strongly transverse, about
four times as wide as long. Metacoxae large, about four times as
long as wide, with two strongly marked basal lobes, trochantin
fairly large. Elytra densely striated, with 15–18 striae along its
length; exterior margins smooth. Legs moderately developed.
Profemora exterior margin widened in last third, and as long as
mesofemora, bearing a row of dense, robust hairs. Protibiae nar-
row, about five times narrower than profemora at its widest
part, bearing a row of very dense, thick bristles (Text-fig. 1).
Mesofemora and mesotibiae bear natatory hairs on interior mar-
gin. Mesotibiae slightly narrower than, and as long as, mesofe-
mora, widest on posterior edge. Mesotarsi narrower than
mesotibiae, first tarsomere 1Æ5 times longer than second. Metafe-
mora slightly longer than mesofemora, roundly narrowed to
anterior and posterior. Metatibiae as long and wide as metafe-
mora, widest at base and bearing two short spurs. Metatarsi nar-
rower than metatibiae; first tarsomere about twice as long as rest
of tarsomeres.
Hispanoclavina gratshevi sp. nov.
Plate 1, figure 2; Text-figure 2
Derivation of name. After the Russian palaeoentomologist Dr
Vadim G. Gratshev.
Type specimen. LH15961, part and counterpart, impression of
beetle lacking antennae and distal part of legs.
Type locality and horizon. As for H. diazromerali.
Other material. Same locality: LH15747, 23000.
Diagnosis. Body suboval, metacoxae shorter than femoral
plates. Elytra with no more than 12 striae.
TEXT -F IG . 1 . Hispanoclavina diazromerali gen. et sp. nov., Las Hoyas fossil site, La Huerguina Formation, camera lucida drawings.
A, holotype, LH16264, part, dorsal and ventral views. B, paratype LH23856, part, dorsal and ventral views. Br, bristles; De, dorsal eyes;
Fp, femoral plates; Nt, natatory hairs; Ve, ventral eyes. Scale bars represent 10 mm.
4 PALAEONTOLOGY, VOLUME 50
Description. Total length 14Æ2–15Æ0 mm; maximum width 7Æ2–
7Æ5 mm; elytra length 11Æ0–11Æ5 mm. Body medium-sized and
suboval. Head transverse, about twice as wide as long. Dorsal eyes
quite large, inserted in exterior margins of head. Pronotum com-
paratively short, about twice as long as wide, widest at middle and
roundly narrowed to anterior and posterior. Procoxae short and
rounded; prosternal process about as wide as procoxae and exten-
ded behind them. Mesosternum quite long, about 1Æ5 times as
long as metasternum, and with deep medial fossa in last third.
Mesocoxae narrow and elongated, about twice as long as wide.
Metasternum transverse, about three times as wide as long. Metac-
oxae long, about four times as long as wide, with two basal lobes,
trochantin fairly large. Elytra striated, with about 12 striae along
its length; exterior margins smooth. Legs moderately developed.
Profemora widest at mid-length of exterior margin, and approxi-
mately as long as mesofemora, bearing a row of very robust hairs
(Text-fig. 2). Protibiae narrow, about six times narrower than
profemora at widest part, bearing a row of very dense, thick, com-
paratively short hairs. Mesofemora c. 1Æ5 times narrower than
profemora at widest part. Mesotibiae quite narrow, about three
times narrower than mesofemora, bearing a row of very dense,
fine natatory hairs. Mesotarsi narrow and long. Metafemora lon-
ger than mesofemora. Metatarsi narrow, last four tarsomeres
approximately equal in length and slightly wider than mesotarsi.
Subfamily COPTOCLAVINAE Ponomarenko, 1961
Type species. Coptoclava longipoda Ping, 1926, Lower Cretaceous,
East Asia.
Diagnosis (sensu Ponomarenko 1961). Large beetles.
Metacoxae without femoral plates. Meso- and metatarsi
flattened and broadened, not narrower than tibiae and
without natatory hairs.
Composition. Bolbonectes intermedius Ponomarenko, 1987, Upper
Jurassic, Russia; B. occidentalis Ponomarenko, 1993, Upper Juras-
sic, Russia; Coptoclava africana Teixeira, 1975, Lower Cretaceous,
Angola; C. longipoda Ping, 1926, Lower Cretaceous, China,
Siberia, Mongolia; Megacoptoclava longiurogomphia Pon-
omarenko and Martınez-Delclos, 2000, and Hoyaclava buscalio-
nae gen. et sp. nov., both Lower Cretaceous, Spain.
Genus HOYACLAVA gen. nov.
Derivation of name. After the fossil site of Las Hoyas and the
family name.
Type species. Hoyaclava buscalionae gen. et sp. nov.
Diagnosis. Beetle with large eyes and prognathous head.
Pronotum subquadrate, about twice as long as head.
Metacoxae short, slightly longer than femoral plates.
Profemora and protibiae bearing a row of thick, long
hairs on upper edge. Meso- and metatibiae wider than
tarsomeres.
Hoyaclava buscalionae sp. nov.
Plate 1, figures 3–4; Plate 2, figures 1–2; Text-figure 3
Derivation of name. After palaeontologist Angela D. Buscalioni,
Madrid.
Type specimens. LH24508, part and counterpart, impression of
partially disarticulated beetle, lacking distal part of legs; Las Ho-
yas (Text-fig. 3A). Paratypes: LH7269 (dorsal view of beetle,
with meso- and metathoracic legs preserved; Text-fig. 3B);
LH23060 (disarticulated beetle with well-preserved fore and
meso legs); LH24517 (disarticulated specimen with well-pre-
served legs; Text-fig. 3C–D).
Type locality and horizon. As for Hispanoclavina diazromerali.
Other material studied. Same locality: LH2229, 7098, 7380, 8066,
8109, 9036, 13247, 13518, 13533, 13581, 13643, 15217, 15299,
15777, 16554, 17028, 17168, 21000, 21174, 21250, 22273, 23056,
23181, 23247, 23429, 23431, 23505, 23618, 23633, 23695, 23719,
23803, 23865, 23867, 23870, 23925, 23973, 24503, 24506, 24507,
24510, 24515, 24520, 24521, 24523, 927063.
Diagnosis. As for genus.
Description. Total length 18Æ5–18Æ7 mm; maximum width 9Æ3–
9Æ6 mm; elytra length 11Æ0–11Æ2 mm. Medium-sized beetle, with
oval body. Head transverse, c. 1Æ5 times as wide as long, deeply
retracted into anterior margin of pronotum. Ventral eyes large
TEXT -F IG . 2 . Hispanoclavina gratshevi gen. et sp. nov., Las
Hoyas fossil site, La Huerguina Formation. Holotype LH15961,
part, camera lucida drawings of dorsal and ventral views. Scale
bar represents 5 mm.
SORIANO ET AL . : EARLY CRETACEOUS COPTOCLAVID BEETLES FROM SPAIN 5
and contiguous, about as long as mid-length of pronotum.
Pronotum subquadrate, 1Æ2 times as wide as long, roundly
tapered to anterior and posterior from its mid-length. Proster-
num anterior to procoxae, noticeably longer than procoxae.
Prosternal process comparatively wide, not narrowed api-
cally. Mesocoxae transverse, comparatively large. Metasternum
strongly transverse, c. 3Æ8 times as wide as long, strongly narrow-
ing anteriorly. Metacoxae short, about twice as wide as long.
Medial raised part of coxae almost 1Æ6 times as long as wide.
Abdomen narrowing towards second visible ventrite. Elytra
smooth and oval, epipleura not broadened to anterior, with a
longitudinal stripe on interior margin. Profemora broadened at
mid-length of its anterior edge. Protibiae narrow, c. 3Æ5 times
narrower than profemora at its maximum width, with a keel on
its anterior edge, bearing a row of long, thick hairs. Mesofemora
slightly longer than profemora. Tibiae and tarsomeres of meso-
and metathoracic legs flattened into an oar-shape, widest in last
third, anterior edge rounded; tibial spurs about as long as first
tarsomere, which is considerably narrower than tibiae, without
natatory hairs.
Remarks. Hoyaclava buscalionae is not only the most
common coptoclavid, but one also of the most abundant
insect remains found in Las Hoyas, second only to the
aquatic heteropteran Iberonepa romerali. It is usually
found partially articulated, with at least the two elytra still
joined and part of the prothorax and mesothorax. The
abdomen is not usually preserved in this form, probably
owing to the low degree of sclerotization of the body, a
feature usually observed in other fossil aquatic beetles
(Ponomarenko 1961).
Subfamily NECRONECTINAE Ponomarenko, 1977
Type species. Timarchopsis (¼ Necronectes Ponomarenko, 1977)
czekanowskii Brauer et al. 1889, Lower–Middle Jurassic, Russia.
Diagnosis (sensu Ponomarenko 1977). Large to medium-
sized beetles. Metacoxae with small femoral plates.
Metatibiae and tarsi long and slender, not broadened.
Metatibiae and metatarsi with long, slender natatory hairs.
Composition. Actea sphinx Germar, 1842, Upper Jurassic, Ger-
many; Ditomoptera dubia Germar, 1839, Upper Jurassic, Ger-
many; D. minor Deichmuller, 1886, Upper Jurassic, Germany;
Exedia plana Ponomarenko, 1977, Upper Jurassic, Kazakhstan;
Pseudohydrophilus avitus Heyden, 1847, Upper Jurassic, Ger-
many; Stygeonectes jurassicus Ponomarenko, 1977, Upper Juras-
sic, Mongolia; S. jurassicus Ponomarenko, 1977, Lower Jurassic,
Russia; Timarchopsis czekanowskii Brauer et al., 1889 (¼ aquati-
cus Ponomarenko, 1977), Lower–Middle Jurassic, Russia;
T. cyrenaicus Ponomarenko, 1977, Lower Cretaceous,
Algeria; T. gigas Ponomarenko, 1977, Upper Jurassic, Kazakh-
stan; T. latus Ponomarenko, 1977, Middle Jurassic, Kyrgyzstan;
T. mongolicus Ponomarenko, 1985, Middle Jurassic, Mongolia;
T. gobiensis Ponomarenko, 1987, Upper Jurassic, Mongolia; T.
sainshandensis Ponomarenko, 1987, Middle Jurassic, Mongolia;
Ovonectes pilosum gen. et sp. nov., Lower Cretaceous, Spain.
Genus OVONECTES gen. nov.
Derivation of name. After the oval shape of the body of this bee-
tle and Greek, nectes, oarsman.
Type species. Ovonectes pilosum sp. nov.
Diagnosis. Body small and oval. Pronotum transverse,
with rounded edges. Legs long and slender, with long,
dense natatory hairs on tibiae and tarsi.
Ovonectes pilosum sp. nov.
Plate 2, figures 3–4; Text-figure 4
Derivation of name. After the long natatory hairs of this species.
Types. LH23853, part and counterpart, impression of well-pre-
served beetle, lacking distal part of meta legs (Text-fig. 4A).
Paratype, LH24514, only the part of a disarticulated beetle, with-
out part of legs (Text-fig. 4B).
Type locality and horizon. As for Hispanoclavina diazromerali.
Diagnosis. As for genus.
Description. Total length 9Æ7 mm; maximum width 5Æ4 mm; ely-
tra length 6Æ5 mm. Body convex and oval, not flattened. Head
comparatively large and long, retracted into anterior edge of
pronotum, about twice as wide as long without mandibles. Dor-
sal eyes small, about twice as short as head without mandibles;
separation greater than their diameter. Pronotum transverse,
tapering roundly to anterior, 2Æ2 times as wide as long on its pos-
terior edge, pronotal angles rounded. Propleura wide, narrowing
to posterior. Prosternum longer than procoxae and prosternal
process. Prosternal process as long as procoxae. Mesosternum
comparatively long, 1Æ7 times as wide as long, with deep longi-
EXPLANATION OF PLATE 1
Fig. 1. Hispanoclavina diazromerali gen. et sp. nov., holotype, LH16264, part.
Fig. 2. Hispanoclavina gratshevi gen. et sp. nov., holotype, LH15961, part.
Figs 3–4. Hoyaclava buscalionae gen. et sp. nov. 3, holotype, LH24508, part. 4, paratype LH24517, part.
All specimens from the Las Hoyas locality, La Huerguina Formation. Scale bars represent 5 mm.
6 PALAEONTOLOGY, VOLUME 50
tudinal groove for prosternal process. Mesocoxae oblique, dis-
tance between them equal to mesocoxae width. Metasternum
short, width more than 3Æ2 times distance between meso- and
metacoxae. Metacoxae with femoral plate 1Æ8 times as wide as
long; part of metacoxae projecting over abdomen for almost one-
third length of entire coxae. Femoral plates transverse, roundly
tapered laterally. Ventrites subequal in length, last ventrite
slightly longer than those preceding it. Elytra smooth, without
noticeable striae, epipleura narrow. Profemora three times as long
as wide, not widened medially. Protibiae slightly longer than
profemora, with a medial keel. Tibial spurs as long as first tarso-
mere. Protarsomeres subequal in length, except last, which is
about twice as long. Mesotibiae as wide as protibiae, with medial
keel. Mesotarsomeres with long, dense natatory hairs; last tarso-
mere twice length of others; claw as long as third tarsomere.
Subfamily COPTOCLAVISCINAE subfam. nov.
Derivation of name. After Coptoclavisca Ponomarenko, 1987,
which clearly defines the characters of this new subfamily.
Type species. Coptoclavisca nigricollinus Ponomarenko, 1987,
Lower Cretaceous, Mongolia.
Diagnosis. Small beetles, with femoral plates comparat-
ively widened. All legs without noticeable swimming
hairs.
Composition. Coptoclavisca nigricollinus Ponomarenko, 1987;
Coptoclavella elegans Ponomarenko, 1980; C. minor Pon-
omarenko 1980; C. striata Ponomarenko, 1986; C. vittata Pon-
omarenko, 1986: all Lower Cretaceous, Mongolia; and
C. purbeckensis Ponomarenko et al., 2005, Lower Cretaceous,
England; C. inexpecta gen. et sp. nov., Lower Cretaceous, Spain.
Genus COPTOCLAVELLA Ponomarenko, 1980
Type species. Coptoclavella elegans Ponomarenko, 1980, Lower
Cretaceous, Mongolia.
Coptoclavella inexpecta sp. nov.
Plate 2, figure 5; Text-figure 5
Derivation of name. Latin, inexpecta, unexpected.
Type specimen. LC3955, part and counterpart of an impression
of an articulated beetle, lacking part of the legs (Text-fig. 5).
TEXT -F IG . 3 . Hoyaclava buscalionae gen. et sp. nov., Las
Hoyas fossil site, La Huerguina Formation, camera lucida
drawings. A, holotype, LH24508, part. B, paratype, LH7269,
counterpart. C, paratype, LH24517, part, anterior leg, arrows
indicate the position of the bristles along the exterior margin
of profemur and protibia. D, paratype LH24517, part, meso-
leg. Br, bristles; De-Ve, dorsal and ventral eyes superimposed;
Ft, fused metatarsomeres. Scale bars represent 2 mm.
EXPLANATION OF PLATE 2
Figs 1–2. Hoyaclava buscalionae gen. et sp. nov., Las Hoyas locality, La Huerguina Formation. 1, paratype, LH7269, counterpart. 2,
paratype, LH23060, part. Scale bars represent 5 mm.
Figs 3–4. Ovonectes pilosum gen. et sp. nov., Las Hoyas locality, La Huerguina Formation. 3, holotype, LH23853, part. 4, paratype,
LH24514, part. Scale bars represent 1 mm.
Fig. 5. Coptoclavella inexpecta gen. et sp. nov., El Montsec locality, ‘Calcaires lithographiques a Plantes et Vertebres de la Pedrera de
Rubies’ Formation, holotype, LC3955, counterpart. Scale bars represent 5 mm in 1–2, and 1 mm in 3–5.
8 PALAEONTOLOGY, VOLUME 50
Type locality and horizon. La Cabrua outcrop, near village of
Santa Maria de Meia, Sierra del Montsec, Lleida Province; ‘Calc-
aires lithographiques a Plantes et Vertebres de la Pedrera de
Rubies Formation’, Barremian (Martın-Closas and Lopez-Moron
1995).
Material. Only the holotype.
Diagnosis. Head and pronotum are the shortest in the
genus. Elytra with dark band along the median line.
Description. Total length 6Æ2 mm; maximum width 2Æ7 mm;
elytra length 3Æ8 mm. Body dorsally flattened. Head short and
retracted into anterior edge of pronotum, c. 1Æ3 times as wide as
long. Dorsal eyes small, twice as short as head, separated by
more than 3Æ5 times their width. Pronotum transverse, 3Æ7 times
as wide as long, roundly tapering to anterior and posterior,
pronotal angles rounded. Propleura wide, not narrowed to pos-
terior. Prosternum slightly longer than procoxae and prosternal
process, which is approximately as long as procoxae. Mesoster-
num slightly longer than mesocoxae, with deep longitudinal
groove for prosternal process. Mesocoxae oblique, distance
between them shorter than for procoxae. Metasternum short,
c. 1Æ7 times as wide as long. Metacoxae short, with femoral pla-
te 1Æ3 times as wide as long. Femoral plates transverse, roundly
tapering laterally. Ventrites subequal in length, last ventrite c. 1Æ3
times longer than that preceding it. Elytra with shallow striae
and a white central stripe; epipleura narrow. Mesofemora short,
c. 1Æ2 times as long as wide. Metafemora as long as mesofemora.
Metatibiae 2Æ5 times narrower than metafemora, without notice-
able natatory hairs.
DISCUSSION
Coptoclavids were the most common aquatic beetles dur-
ing the Jurassic and Early Cretaceous. In the Bon-Tsagan
locality (Lower Cretaceous, Mongolia) adults and larvae
of Coptoclava longipoda are represented in some horizons
by several specimens. At Las Hoyas the most common
coptoclavid, Hoyaclava buscalionae, is also of the sub-
family Coptoclavinae. In contrast, only two specimens of
coptoclavids have been recorded from El Montsec, each
belonging to a different genus and species: Bolbonectes
lithographicus and Coptoclavella inexpecta.
Despite the relative abundance of coptoclavids in Late
Jurassic and Early Cretaceous deposits, their diversity is
limited: 23 species of 13 genera. The most diverse fossil
sites are Las Hoyas with seven species and Karatau (Upper
Jurassic, Kazakhstan) with six. The variety of coptoclavids
found at Las Hoyas, together with the abundance and
diversity of other predaceous aquatic insects, is one of the
main features of the aquatic entomofauna of this outcrop
(Soriano and Delclos 2005); the predaceous component
may explain why the first and only filter-feeding copto-
clavids and beetles, Hispanoclavina and Hoyaclava, are
encountered there. The long bristles in the femur and
tibia-tarsus of the forelegs in these types (Text-figs 1, 3C),
TEXT -F IG . 5 . Coptoclavella inexpecta gen. et sp. nov.,
El Montsec outcrop, ‘Calcaires lithographiques a Plantes et
Vertebres de la Pedrera de Rubies’ Formation, camera lucida
drawings. Holotype, LC3955, counterpart. A, dorsal view. B,
ventral view. Scale bars represent 2 mm.
TEXT -F IG . 4 . Ovonectes pilosum gen. et sp. nov., Las Hoyas
fossil site, La Huerguina Formation, camera lucida drawings. A,
holotype, LH23853, part, dorsal and ventral views. B, paratype,
LH24514, part, dorsal and ventral views. Scale bars represent
2 mm.
10 PALAEONTOLOGY, VOLUME 50
the elongation of the protibiae (consistent with the pres-
ence of euthyplociid mayflies in the deposits), and the
presence of divided eyes typical of the family (four eyes,
two specialized for underwater vision and two open to the
air) suggest that these new species filtered plankton.
Owing to the carnivorous habit of the family, they prob-
ably fed on zooplankton on the water surface (essentially
composed of chaoborid- and chironomid-like larvae and
pupae) rather than the benthos. From the morphology of
their forelegs and the design of their eyes, and given the
feeding strategy among known coptoclavids, we suggest
that their trophic habit was exclusively carnivorous, rather
than suspensivorous or detritivourous as for mayflies.
The leg structure of Hispanoclavina and Hoyaclava, and
the proposed feeding strategy for these genera are unique
among the hyper-diverse group of coleopterans, both
extant and extinct. This new habit for aquatic beetles at
Las Hoyas may be related to the enormous diversity of
predaceous forms in the lake, including other insects
(belostomatids, chresmodids and odonatans) and verte-
brates (fish and amphibians) (Delclos et al. 2004). These
new coptoclavids would have hunted or filtered on and
under the water surface, using their divided eyes with the
dorsal part adapted for aerial vision (which would have
allowed them to monitor the presence of aerial predators,
such as birds, or other pleustonic insects, such as Chres-
moda), and the ventral part with aquatic vision, allowing
them to search for food and monitor aquatic predators,
such as belostomatids, fish or amphibians. In this way,
coptoclavids developed two feeding strategies: hunting, as
for the rest of the group, and zooplankton filtering, in
direct competition with aquatic and non-aquatic forms.
Acknowledgements. We thank all of our colleagues in the Russian
Academy of Sciences, Moscow, for their support, especially Dr
A. P. Rasnitsyn for making possible our study of the collections
housed in this institute and Dr D. E. Shcherbakov for his help
with photography. We also thank Mr Armando Dıaz-Romeral
(Cuenca) for allowing us to study specimens from his personal
collection (now housed in the Museo de las Ciencias de Castilla
La Mancha in Cuenca). Aspects of this research were supported
by grants BOS2001-0173 and CGL2005-00046 ⁄ BOS from the
Ministry of Science and Technology of Spain. We are also grate-
ful for the correction and improvement of the first draft of the
manuscript by two reviewers, and of the revised manuscript by
Profesor D. J. Batten.
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12 PALAEONTOLOGY, VOLUME 50
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