COPTOCLAVID BEETLES (COLEOPTERA: ADEPHAGA) FROM THE LOWER CRETACEOUS OF SPAIN: A NEW FEEDING...

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: [email protected]; [email protected]

�Paleontological Institute, Russian Academy of Sciences, Moscow, Russia; e-mail: [email protected]

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.


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.



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.


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.


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.


PLATE 1

SORIANO et al., Hispanoclavina, Hoyaclava

1

3 4

2

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.


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.


PLATE 2

SORIANO et al., coptoclavid beetles

1

2

5

3

4

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.


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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COPTOCLAVID BEETLES (COLEOPTERA: ADEPHAGA) FROM THE LOWER CRETACEOUS OF SPAIN: A NEW FEEDING...

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