NEW FOSSIL LACEWINGS AND ANTLIONS … FOSSIL LACEWINGS AND ANTLIONS (INSECTA, NEUROPTERA) FROM THE...

NEW FOSSIL LACEWINGS AND ANTLIONS (INSECTA,

NEUROPTERA) FROM THE LOWER CRETACEOUS

CRATO FORMATION OF BRAZIL

by FEDERICA MENON* and VLADIMIR N. MAKARKIN�*School of Earth, Atmospheric and Environmental Sciences, University of Manchester, Williamson Building, Oxford Road, Manchester M13 9PL, UK;

e-mail: [email protected]

�Institute of Biology and Soil Sciences, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok 690022, Russia; e-mail: [email protected]

Typescript received 8 August 2006; accepted in revised form 22 January 2007

Abstract: Remarkable new fossil taxa of Neuroptera from

the laminated limestone of the Crato Formation, north-east

Brazil, are described: Nuddsia longiantennata gen. et sp. nov.

(Osmylidae, Gumillinae), the first fossil record of this family

from South America, Parapalaeoleon magnus gen. et sp. nov.

(Palaeoleontidae), and Triangulochrysopa formosa sp. nov.

(Mesochrysopidae). A diagnosis of Gumillinae is provided;

Epiosmylidae is considered to be a synonym of this subfam-

ily. A revised diagnosis of Palaeoleontidae is provided, based

on a new interpretation of venational characters evident on

the exceptionally well-preserved specimen of P. magnus; vein

homologies are determined and the fusion of MP and CuA is

discussed. The genus Triangulochrysopa has been known pre-

viously only from the Lower Cretaceous of Las Hoyas, Spain.

Key words: Araripe, Crato Formation, Nova Olinda Mem-

ber, Neuroptera, Osmylidae, Palaeoleontidae, Mesochrysopi-

dae, taxonomy.

The Crato Formation is well known for the diversity and

quality of its fossil insects; three-dimensional features and

colour pattern are often preserved (Martill and Frey 1995;

Heads et al. 2005). The formation ranges across the Ara-

ripe Plateau in the states of Ceara, Pernambuco and Pia-

uı, north-east Brazil. It is composed of a series of finely

laminated limestones that accumulated at the bottom of a

lagoon during the initial phase of the opening of the

Atlantic Ocean (Martill 1993). Its precise age is somewhat

unclear: it is generally considered to be Late Aptian–Early

Albian, 110–120 Ma (Berthou 1994), and has been dated

as Late Aptian on the basis of palynological data (Pons

et al. 1991). The formation is subdivided into three mem-

bers: Nova Olinda, Barbalha and Jamacaru. The basal

Nova Olinda Member is the most fossiliferous: insects are

found together with other arthropods, plants and wood,

fish, pterosaurs and other vertebrates (Maisey 1991).

The order Neuroptera is one of the most significant

and diverse groups of insects found in the Crato Forma-

tion, with 55 species having been described (Martins-Neto

2000, 2003, 2005; Heads et al. 2005; Makarkin and

Menon 2005, 2007; Menon et al. 2005; Nel et al. 2005). A

vast amount of material in a variety of museums remains

unstudied. We describe here two new genera and three

new species belonging to the families Osmylidae, Pala-

eoleontidae and Mesochrysopidae, based on specimens

housed at the Staatlisches Museum fur Naturkunde Stutt-

gart, Germany. No fossil record of Osmylidae was known

hitherto from South America.

MATERIAL AND METHODS

All specimens described here are from the lowest member

of the Crato Formation. They were probably collected by

workers in one of the small mines or stone yards around

Nova Olinda, although the exact locality is unknown.

They are preserved as limonitic replacements after pyrite

(Martill and Frey 1995). The specimens were prepared

on-site by the collectors. An aeroneedle was used, where

necessary, to remove any residual sediment and dust

(Selden 2003).

Drawings were made using a camera lucida attached to

a Wild stereomicroscope; photographs were taken with a

Sony DCS-717 digital camera and a D1X digital camera

attached to a Wild M8 stereo-zoom microscope.

SYSTEMATIC PALAEONTOLOGY

Wing venation terminology follows Comstock (1918), with a few

exceptions in accordance with current usage in neuropterology

(see Archibald and Makarkin 2006; Wedmann and Makarkin

2007). Terminology of wing spaces follows Oswald (1993)

[Palaeontology, Vol. 51, Part 1, 2008, pp. 149–162]

ª The Palaeontological Association doi: 10.1111/j.1475-4983.2007.00740.x 149

Venation abbreviations used in the text and figures are as fol-

lows: 1A–3A, anal veins; bf, anterior Banksian fold; Cu, cubitus;

CuA, anterior cubitus; CuA1, most proximal branch of anterior

cubitus; CuP, posterior cubitus; M, media; MA, anterior branch

of media; mfl, median flexion line; MP, posterior branch of

media; MP1, most proximal branch of MP; R, radius; R1, first

branch of radius; Rs, radial sector; Rs1, most proximal branch of

radial sector; Sc, subcosta.

Institutional abbreviation. SMNS, Staatliches Museum fur

Naturkunde Stuttgart, Germany.

Class INSECTA Linnaeus, 1758

Order NEUROPTERA Linnaeus, 1758

Family OSMYLIDAE Leach, 1815

Subfamily GUMILLINAE Navas, 1912

1912 Gumillini Navas, p. 191 (as a ‘tribe’ of Osmylidae).

1988 Gumillinae Lambkin, p. 455 (as a subfamily of

Osmylidae).

1980 Epiosmylidae Panfilov, in Dolin et al., p. 100, syn.

nov.

Type genus. Gumilla Navas, 1912 (Recent).

Diagnosis. Medium-sized osmylids that may be distin-

guished from other species of the family by the following

combination of character states: (1) antennae exceedingly

elongate, much longer than forewing [much less than

forewing length in other subfamilies]; (2) scape strongly

enlarged [moderately enlarged in other osmylids]; (3)

basal sinuous crossvein r-m in hindwing absent, character

state shared with Stenosmylinae [present in other subfam-

ilies]; (4) outer gradate series of crossveins absent in both

wings, as in Porisminae [present in other subfamilies]; (5)

trichosors limited at most to apical part of wings [nor-

mally present at least in distal half of wings in other os-

mylids].

Included genera. Gumilla (Recent, Brazil; one or two species);

Epiosmylus Panfilov, in Dolin et al. 1980 (Upper Jurassic, Kara-

tau, Kazakhstan, Karabastau Formation; Middle–Upper Jurassic,

Daohugou, Inner Mongolia, China; two species); Nuddsia gen.

nov. (Lower Cretaceous, Upper Aptian, Araripe Basin, Brazil,

Crato Formation; one species).

Remarks. Three taxa with extremely long antennae, at

least 1.5 times longer than the forewing length, are known

within Neuroptera. These include the extant Chrysopidae,

in the majority of Apochrysinae and a few genera of

Chrysopinae, and the Osmylidae, in the subfamily Gumil-

linae and the fossil Epiosmylidae.

The family Epiosmylidae was erected to accommodate

Epiosmylus longicornis Panfilov, 1980 (Upper Jurassic of

Karatau, Kazakhstan). Another species of this genus

(E. panfilovi Ren and Yin, 2002) was added later from the

Middle–Upper Jurassic of China (Ren and Yin 2002).

Since its establishment, Epiosmylidae has been generally

treated as either a synonym of Osmylidae (e.g. Pono-

marenko 1980; Makarkin 1990b; Ren and Yin 2002) or a

taxon whose status is unclear, ‘either an isolated branch

within Osmylidae or else a separate family’ (Makarkin

and Archibald 2003, p. 175). Recently, Ponomarenko

(2003) described another osmylid-like species with extre-

mely long antennae, ‘Nymphites cf. lithographicus’, from

the Upper Jurassic of Solnhofen, Germany, and assigned

it to the Nymphitidae. However, the photograph and

drawings of this specimen provided indicate the possibil-

ity that it belongs to Osmylidae (possibly to Gumillinae

because of its long antennae); its venation is most similar

to that found in this family, and its relation to the true

Nymphites lithographicus Handlirsch, 1906 is unclear, as

its holotype is very poorly preserved.

The subfamily Gumillinae has not been formally

defined. The ‘tribe’ Gumillini was created by Navas

(1912) within Osmylidae for the genus Gumilla Navas,

including the two extant Brazilian species, G. adspersa

Navas, 1912 and G. longicornis (Walker, 1853), which

should perhaps be considered synonyms (Adams 1977).

Navas (1912) grouped all other osmylids in the ‘tribe’

Osmylini; thus the actual rank of these ‘tribes’, according

to Navas (1912), could be considered at the subfamily

level. Gumilla was subsequently excluded from the family

by Kruger (1913, 1915), then included again by Adams

(1969, 1977). Adams (1969, p. 2) mentioned that this

genus is highly aberrant and ‘is too poorly known for

meaningful discussion’. He correctly concluded that Nav-

as’ (1912, fig. 24) drawing of the forewing is reasonably

accurate, and provided some additional diagnostic fea-

tures of the genus (Adams 1977). Unfortunately, the char-

acters of the male terminalia are largely unknown, beyond

the short description of Navas (1912, p. 57) (‘cercis cylin-

dricis, pilosis, brevibus, externe convexis’), and they have

never been illustrated.

It is reasonable to assume that Epiosmylidae and

Gumillinae might be the same, as they share some impor-

tant character states (see diagnosis, above), even though

Epiosmylus and Gumilla, their type genera, differ quite

considerably by the configuration of MA, MP, CuA and

CuP in the forewing, which are straighter with branching

that is fairly regular in the former and strongly irregular

in the latter (see also note of Lambkin 1988, p. 455).

Examination of the photograph of the holotype of G. ad-

spersa reveals that its forewing venation is somewhat

anomalous (the venation of right and left wings is

strongly asymmetrical). The forewing venation of Nuddsia

(e.g. configuration of MP, CuA, CuP) is also irregular to

150 P A L A E O N T O L O G Y , V O L U M E 5 1

some extent (Text-fig. 2), providing additional support

for placing it in Gumillinae. The hindwing venation of all

species (including the holotype of G. adspersa and the

new species) does not differ in its main character states.

Ocelli are absent in the extant Gumilla (present in all

other osmylids) but are not detectable in fossil taxa

because of poor preservation or as the head is missing.

Gumillinae is considered here to be a subfamily of

Osmylidae because its venation generally agrees well with

that of this family, and to be the sister group of all other

osmylids because of the clear differences between them

(see diagnosis above). The loss of the basal sinuous cross-

vein r-m in the hindwing and the absence of the outer

gradate series of crossveins in both wings, shared by

Gumillinae with Stenosmylinae (the former) and Porismi-

nae (the latter) might be convergent.

Genus NUDDSIA gen. nov.

Type and only species. Nuddsia longiantennata sp. nov.

Derivation of name. After Dr John Nudds, Head of the Palaeon-

tology Research Group at the University of Manchester. Gender

feminine.

Diagnosis. May be distinguished from other genera of the

subfamily by the following combination of character

states: (1) antennae nearly 2.5 times as long as forewing

[not more than 1.5 times in Epiosmylus, Gumilla]; (2) M

in forewing forked highly proximal to origin of Rs1

[slightly distal in Gumilla]; (3) distal half of CuA in fore-

wing arched [parallel to hind margin in Epiosmylus].

Remarks. Nuddsia gen. nov. represents the first fossil

record of this family from South America. Interestingly,

the single living genus of this subfamily (Gumilla) also

occurs in Brazil, and the wing venation of both genera is

quite similar.

Nuddsia longiantennata sp. nov

Text-figures 1–2

Derivation of name. Latin, longus, long, and Medieval Latin

antenna, sail yard (translation of Greek keraia, insect feeler,

yard-arm), in reference to the extremely long antennae of this

species.

Holotype. SMNS 66000 ⁄ 263. An almost complete specimen

exposed in ventral aspect.

Type locality and horizon. Brazil, Ceara, Araripe Basin (Chapada

do Araripe), vicinity of Nova Olinda; Crato Formation (Nova

Olinda Member), Lower Cretaceous (Upper Aptian).

Diagnosis. As for the genus.

Description. Head twice as wide as long (1 mm long, 2 mm

wide); large compound eyes. Antennae extremely long (60 mm),

filiform; scape very large, bulky; pedicel elongate, rounded, larger

than flagellar segments. Prothorax nearly quadrate, c. 1 mm

long. Mesothorax not clearly visible, apparently wide. Legs not

preserved. Abdomen incomplete, 4 mm long as preserved. Head,

thorax and abdomen strongly compressed. Exoskeleton of most

portions of head, thorax and abdomen not preserved.

Forewing elongate, 25.5 mm long, c. 6.5 mm wide; apex sub-

acute. Trichosors prominent, restricted to apical portion of wing.

Costal space 0.6 mm maximum width, narrow basally, slightly

dilated near proximal third, then narrowed towards apex. Sub-

costal veinlets simple, straight, nearly perpendicular to Sc in

basal two-thirds of costal space, increasingly more oblique dis-

tad; veinlets of Sc+R1 closely spaced, strongly oblique. Subcostal

space narrow, dilated near fusion of Sc, R1. Sc, R1 fused 4 mm

from wing apex. Sc+R1 entering margin before wing apex. Rs

originating near wing base. Stem of Rs zigzagged, forked api-

cally, entering margin slightly before wing apex. R1 space wider

than costal space, 0.8 mm, strongly narrowed apically, with 23

more or less regularly spaced crossveins proximad fusion of Sc,

R1, one distad (left wing). Rs with six branches, distal branch

deeply forked. Origin of Rs1 at approximately one-third wing

length, c. 8 mm from base (left wing); origins of Rs1–Rs3 widely

spaced, other three branches restricted to apical portion of wing.

Radial crossveins numerous, regularly spaced (except for apical

portion of space, where crossveins rare), not forming gradate

series. Fork of M considerably distal to origin of Rs but much

proximal to origin of Rs1. MA straight, with four pectinate, obli-

que branches distally. MP basally straight, parallel to MA, con-

figuration of distal portion somewhat irregular with 2–3

branches, most proximal of which long, dichotomously

branched. Fork of Cu not preserved, apparently near wing base.

Configuration of distal portion of CuA rather irregular, dichoto-

mously branched. CuP poorly preserved; distal half clearly

arched; short, simple distalmost branches preserved. Anal veins

not preserved. Crossveins rather regularly spaced, except for

most distal portion of radial space, where crossveins are scarce.

Gradate series of crossveins not detected.

Hindwing elongate, 24.5 mm long, 5.5 mm wide, with sub-

acute apex. Trichosors restricted to apical portion. Nygmata

not detected. Costal space very narrow, 0.5 mm wide basally

(left wing), slightly dilated in pterostigmal region. Subcostal

veinlets simple, perpendicular to Sc in basal half of costal

space, increasingly more oblique distad; veinlets of Sc+R1 clo-

sely spaced, strongly oblique. Subcostal space very narrow for

most of its length, slightly dilated near fusion of Sc, R1. Sc, R1

fused at 3.5 mm from apex. Sc+R1 entering margin slightly

before apex. Rs originating near wing base. R1 space 0.8 mm

wide medially, strongly narrowed apically; with 21 more or less

regularly spaced crossveins before fusion of Sc, R1, long hypo-

stigmal cell after. Rs strongly zigzagged, forked apically, enter-

ing margin at apex, with 6–7 branches, mostly zigzagged, with

deep marginal forks. Origin of Rs1 at approximately one-third

wing length. Radial, medio-radial crossveins numerous, regu-

larly spaced, not forming gradate series. Basal crossveins m-r

M E N O N A N D M A K A R K I N : C R E T A C E O U S F O S S I L L A C E W I N G S A N D A N T L I O N S F R O M B R A Z I L 151

not detected, probably absent. Origin of M, its division into

MA and MP not preserved; MA straight for entire preserved

length, pectinately branched distally, with five branches, most

proximal of which forked twice. MP zigzagged, with three dis-

tal branches, most proximal of which long, pectinately

branched (two branches). CuA long, zigzagged, parallel to hind

margin, pectinately branched, with at least 12 rather short,

simple branches. Crossveins in medial, medio-cubital spaces

numerous, more or less regularly spaced. CuP, anal veins not

preserved.

A

B

TEXT -F IG . 1 . Nuddsia longiantennata gen. et sp. nov. A, photograph, and B, line drawing of the holotype, SMNS 66000 ⁄ 263. Scale

bar represents 10 mm.


Family PALAEOLEONTIDAE Martins-Neto, 1992

1992 Palaeoleontinae Martins-Neto, p. 810 (as a subfamily

of Myrmeleontidae).

1997 Palaeoleontidae; Dobruskina et al., p. 93.

Type genus. Palaeoleon Rice, 1969, from the Albian–Cenomanian

of Labrador, Canada.

Revised diagnosis. Large myrmeleontoids (forewing 36–

70 mm long) with relatively long antennae, unusually

dense crossvenation in the majority of genera, easily dis-

tinguished from other myrmeleontoid families by the fol-

lowing combination of character states: (1) antennae

long, filiform, neither dilated nor clavate distally [dilated

distally in Myrmeleontidae, Araripeneuridae; clavate in

Ascalaphidae]; (2) origin of Rs situated near wing base in

both wings [removed from base in Myrmeleontidae,

Araripeneuridae, Ascalaphidae, Nemopteridae, Babinskaii-

dae]; (3) presectorial crossveins absent in both wings

[present in Myrmeleontidae, Ascalaphidae, Nemopteridae,

Babinskaiidae]; (4) ‘oblique vein’ present in forewing

[absent in Nymphidae].

Revised description of venation. In forewing, origin of Rs close to

wing base; without presectorial crossveins; Rs1 convex, profusely

branched distally; single well-developed median longitudinal fold

present from wing base to apex; fork of M far removed from wing

base (Palaeoleon, Baisopardus); MA strongly concave, simple;

basal part of MP represented by ‘oblique vein’, distal MP fused

with CuA; Cu divided into CuA, CuP close to wing base; CuA,

MP+CuA convex, both parallel to MA; MP+CuA, MP+CuA1 and

TEXT -F IG . 2 . Nuddsia longiantennata

gen. et sp. nov., holotype, SMNS

66000 ⁄ 263. A, left forewing. B, right

forewing. C, left hindwing. D, right

hindwing. Scale bar represents 5 mm.


its basalmost branch form large triangular area characteristic of

higher myrmeleontoids; CuP relatively short, pectinately branch-

ing, not fused with 1A; 1A from short, nearly simple to long,

pectinately branched. In hindwing, origin of Rs close to wing base,

without presectorial crossveins; Rs1 profusely branched distally;

M forked very close to wing base; MA simple; MP with long

pectinate branches; Cu forked close to wing base; CuA parallel to

hind margin basally, arched, pectinately branched distally; CuP

short, close, parallel to hind margin, pectinately branched; anal

area very restricted by short simple veins.

Included genera. Baisopardus Ponomarenko, 1992a (Lower Cre-

taceous, Lower Valanginian, Baissa, Transbaikalia, Russia; Crato

Formation; 3–4 species); Parapalaeoleon gen. nov. (Crato Forma-

tion; monotypic); Neurastenyx Martins-Neto and Vulcano, 1997

(Crato Formation; monotypic); Paraneurastenyx Martin-Neto,

1998 (Crato Formation; monotypic); Palaeoleon (Lower–Upper

Cretaceous, Albian–Cenomanian, Labrador, Canada; monotypic);

Samsonileon Ponomarenko, in Dobruskina et al. 1997 (Upper

Cretaceous, Lower Turonian, Israel; monotypic); Metahemerobius

Makarkin, 1990a (Upper Cretaceous–Palaeogene, Maastrichtian–

Danian, Antibes, Siberia, Russia; monotypic).

Remarks. The diagnosis of this family provided by

Dobruskina et al. (1997) and Heads et al. (2005) is par-

tially erroneous, being based on misinterpretation of the

venation in more poorly preserved specimens, particularly

concerning M and CuA. The new material provides the

opportunity to clarify this. According to these authors,

‘MP’ in the forewing is forked into ‘MP1’ and ‘MP2’ in

the proximal portion of the wing, with ‘MP1’ profusely

branched distally, and ‘MP2’ simple. However, this fork is

not detected confidently in any species. The well-pre-

served venation of the forewing of Parapalaeoleon magnus

(Text-fig. 3C) shows that this ‘MP1’ is actually Rs1. The

homology of Rs1 in all species is unequivocal: this vein is

always convex, and is followed posteriorly by a strongly

concave simple vein (we interpret this as MA). Compari-

son of Palaeoleon ferrogeneticus Rice, 1969 and Baisopar-

dus cryptohymen Heads et al., 2005 shows that the

structure of Rs2 is also similar in these species: convex

basally and concave distally.

Examination of photographs of these specimens reveals

that what the authors interpreted as the proximal part of

‘MP1’ (Heads et al. 2005, fig. 3A) and ‘M1’ (Rice 1969, fig.

2) is actually the proximal part of a longitudinal vein-like

fold running between Rs and M. Its course is completely

coincident with that of the median flexion line in some

insects with anteroposteriorly symmetric wings, such as Si-

alis Latreille, 1802 (Wootton 2002, fig. 3A), running longi-

tudinally from the wing base to the wing apex between the

radial and medial vein systems and distally crossing the

branches of Rs (Wootton 2003, fig. 5). The wings of many

extant Myrmeleontidae that have anteroposteriorly asym-

metric wings appear to possess the distal part of this line

(appearing as a fold), which is often accompanied by a

false longitudinal vein formed by the bending of branches

of Rs and the crossveins connecting them, called the ‘ante-

rior Banksian line’ (Tillyard 1916; Krivokhatsky 1998).

This fold will be called herein ‘the anterior Banksian fold’

as its homology with the distal part of the median flexion

line of species with the anteroposteriorly asymmetric wings

is unclear. In Parapalaeoleon magnus, the median flexion

line appears as a true flexion line running (as in other

Neuroptera with anteroposteriorly asymmetric wings:

Wootton 2002, fig. 4B) between MA and Rs, then Rs1, ter-

minating far distal to the origin of the anterior Banksian

fold, which begins slightly distal to Rs1 (i.e. distal part of

the median flexion line and proximal part of the anterior

Banksian fold are divergent). In other species (e.g. Palaeo-

leon ferrogeneticus, Baisopardus cryptohymen) in which the

origin of Rs1 is situated very near the origin of Rs, the

median flexion line crosses Rs1 near its base and continues

further as a concave fold towards the wing apex. In these

species the median flexion line proximally and the anterior

Banksian fold distally form a single straight line, called

here the median longitudinal fold. Basally, this line

appears to be vein-like in compression fossils, as in P. fer-

rogeneticus. Such secondary vein elements are frequently

reformed from membrane (Kukalova-Peck 1991).

Fusion of MP and CuA in the forewing. This fusion has

apparently occurred independently many times in the

evolution of the Neuroptera. It is found in a few genera

of the extant Hemerobiidae (Nusalala Navas, 1913, Micro-

mus Rambur, 1842), Dilaridae (Nallachius Navas, 1909),

and in a Lower Cretaceous psychopsid-like genus from

the Baissa locality (Makarkin, unpublished). In two speci-

mens of the extant Gerstaeckerella chilensis (Hagen, 1859)

(Mantispidae) examined in this study, MP and CuA are

fused abnormally for short distance at the crossvein 2m-

cu (which is consequently lost) in the manner that occurs

in Parapalaeoleon magnus. Interestingly, CuA in Gers-

taeckerella Enderlein, 1910 (as in other mantispids) is

short. In Mesochrysopidae, all intermediate stages are

present, from MP and CuA widely spaced to touching

(Makarkin and Menon 2005; Nel et al. 2005). The fullest

fusion of these veins, however, occurs in the forewing of

the myrmeleontoid lineage (Palaeoleontidae, Araripeneu-

ridae, Myrmeleontidae, Ascalaphidae, Nemopteridae, and

probably Babinskaiidae), in which MP resembles a cross-

vein (‘oblique vein’) often not distinguished from true

crossveins (see discussions in Tillyard 1916, 1918; Com-

stock 1918; Adams 1996). The venation of P. magnus

(Text-figs 3B, 4B) rather clearly demonstrates that this

full fusion might have resulted from touching or short

fusion of a long pectinate MP and short CuA, almost

exactly in the way predicted by Tillyard (1916, fig. 8b; his


‘Cu2’ is, however, actually CuA1). Moreover, MP and

CuA in the forewing of Paraneurastenyx ascalaphix Mar-

tins-Neto, 1998 are almost touching but not fused (Mar-

tins-Neto 1998, fig. 1A), as in some Mesochrysopidae

(Makarkin and Menon 2005, fig. 3B). Based on this

hypothesis, it is reasonable to assume that the most prox-

imal pectinate branch of MP+CuA (i.e. MP+CuA1) repre-

sents the distal portion of CuA (Text-fig. 3B); the next

branch, MP+CuA2 (fused with MP+CuA1 distally in this

specimen), might have originated from the most proximal

branch of MP (MP1), and other branches of MP+CuA all

originate from MP. Some species of the extant Myr-

meleontidae show a remarkable reversal of the conditions

of these veins, indirectly supporting this hypothesis. In

the Australian Mjoberbia fulviguttata Esben-Petersen, 1918

and Protoplectron pallidum Banks, 1910, MP and CuA are

not fused, and CuA has few branches (New 1985, figs

263, 290, but see intraspecific variation, e.g. fig. 265).

Notes on generic and specific composition of Palaeoleonti-

dae. Palaeoleon was erected for P. ferrogeneticus, repre-

sented by a single, well-preserved, apical two-thirds of a

wing. We examined photographs of the holotype and

found that the original description and figures of Rice

(1969, p. 3, figs 1, 3; pl. 1) appear to be fairly adequate,

although the venation could possibly be interpreted dif-

ferently: Rice’s M1 is the median flexion line (basally)

and Rs2 (distally), M2 is Rs1, Cu1 is MA, Cu2 is MP (see

reasoning above).

According to Martins-Neto (1997) the genus Neuraste-

nyx includes N. gigas Martins-Neto and Vulcano, 1997

(type species), N. polyhymnia Martins-Neto, 1997 and N.

araripensis (Martins-Neto 1992). All of these species were

placed in Baisopardus (see below), along with Baisopardus

gigas, by Heads et al. (2005) who consequently synonym-

yzed these two genera. However, in our opinion, Neurast-

enyx is valid and consists of N. gigas only. The holotype

of the type species is poorly preserved (Martins-Neto and

Vulcano 1997, fig. 7), but it possesses a combination of

the character states that distinguish the species from other

palaeoleontids: large size (forewing length about 70 mm)

and very widely spaced crossveins.

Baisopardus is considered to include four species

(Heads et al. 2005). The type species, B. banksianus Pon-

omarenko, 1992a, is known from two isolated hindwings,

the holotype and paratype, from the Lower Cretaceous of

Baissa, Transbaikalian Russia (Ponomarenko 1992a). The

genus was included by Dobruskina et al. (1997) in this

family with some doubt, whereas Heads et al. (2005)

listed it among the genera of Palaeoleontidae with cer-

tainty. In the original description, however, the venation

was misinterpreted by Ponomarenko (1992a, p. 49, fig.

5A): his ‘MA’ is our Rs1, ‘MP’ is MA, ‘CuA’ is MP, ‘CuP’

is CuA, ‘1A’ is CuP.

Palaeoleon araripensis was the first species of this family

to be described from the Crato Formation (Martins-Neto

1992). Later, it was transferred to Neurastenyx by Martins-

Neto (1997), and to Baisopardus by Dobruskina et al.

(1997). The holotype is an almost complete specimen, but

lacks the abdomen, and has incomplete hind- and fore-

wings, which are overlapped pairwise. Ponomarenko (in

Dobruskina et al. 1997) noted that the hindwing venation

of the type specimen was partially incorrectly drawn by

Martins-Neto (1992, p. 811, fig. 5a): the proximal branches

of his CuA [actually MP, see above] belongs to CuP [actu-

ally CuA]; therefore, the configuration of MP and CuA are

similar to those of the type species (see also below).

Baisopardus cryptohymen is represented by a beautifully

preserved, complete, single specimen. The species has a

hindwing shape and venation similar to those of the type

species, B. banksianus, so we find it quite reasonable that it

is assigned to Baisopardus. The single major difference

between these species is that the former possesses closely

spaced subcostal crossveins, which the latter lacks. The

interpretation of the venation in the original description

and figures of Heads et al. (2005, fig. 3) is partially incor-

rect (see reasoning above) and should be emended. In the

forewing, their MA is Rs1 (basally) and Rs2 (distally),

MP1 is the median flexion line (basally) and Rs1 (distally),

MP2 is MA, CuA is MP+CuA, CuA1 is MP+CuA, CuA2 is

MP+CuA1; in the hindwing, their MA is Rs1 (basally) and

Rs2 (distally), MP1 is the median flexion line (basally) and

Rs1 (distally), MP2 is MA, CuA1 is MP, CuA2 plus CuP is

CuA, 1A is CuP (see above for reasoning).

All wings of ‘Neurastenyx’ polyhymnia are so strongly

overlapped that few details of the venation may be clearly

identified (Martins-Neto, 1997, fig. 7). Its systematic posi-

tion, therefore, may possibly be determined confidently

only after close re-examination of the type. The assign-

ment of this species to Baisopardus (as well as to Neurast-

enyx) should be considered tentative, based on the

general impression of its venation rather than on analysis

of particular character states.

Samsonileon and Metahemerobius are apparently closely

related genera; this is indicated by their similar forewing

venation, in particular, MP+CuA1 and CuP run parallel

and very close for a considerable distance, and 1A is

long and pectinately branched. Unfortunately, both gen-

era are represented by incomplete wings, lacking basal

portions in known specimens. Makarkin’s (1990a, fig. 5)

designations of the veins of Metahemerobius kalligram-

mus Makarkin, 1990a are partially incorrect: his MA is

Rs2, anterior branch of MP is Rs1, posterior branch of

MP is MA, CuA is MP+CuA, CuP is MP+CuA1, A1 is

CuP. The venation of Samsonileon fragmentatus Pon-

omarenko, in Dobruskina et al. 1997 is interpreted simi-

larly here: Ponomarenko’s MA is Rs1 (basally) and the

anterior Banksian fold or Rs2 (distally), MP1 is probably


the median flexion line (basally) and Rs1 (distally), CuA

is MP+CuA, CuP and A1 are CuP and 1A respectively.

Cretoleon Ponomarenko, 1992b is a monotypic genus

represented by a forewing of C. acanthoclysoides Pono-

marenko, 1992b from the Lower Cretaceous (Aptian)

deposits of Bon-Tsagan, Mongolia (Ponomarenko 1992b).

It was considered possibly to belong to Palaeoleontidae by

Ponomarenko (in Dobruskina et al. 1997), and listed

among the genera of this family by Heads et al. (2005). It is

tentatively excluded from the family here because it differs

in the following features of the forewing: the costal space is

broad; Rs originates relatively distant from the wing base;

the anterior Banksian fold is absent; the subcostal veinlets

in the pterostigmal region are very closely spaced and sim-

ple; crossveins are rare in the distal portion of the wing.

Genus PARAPALAEOLEON gen. nov.

Type and only species. Parapalaeoleon magnus sp. nov.

Derivation of name. Greek, para, near, and Palaeoleon, a generic

name. Gender masculine.

Diagnosis. May be distinguished from other genera of the

family by the combination of the following forewing char-

acter states: (1) origin of Rs1 far removed from origin of

Rs [near origin of Rs in Baisopardus, Palaeoleon]; (2) Rs1

shallowly branched, highly distal to the Banksian fold

[deeply branched, at most slightly distal to the Banksian

fold in Baisopardus, Palaeoleon]; (3) maximum width of

wing at mid-point [in distal portion in Baisopardus and

in proximal portion in Palaeoleon].

Remarks. The new genus is most similar to Baisopardus

and Palaeoleon; other genera are very distant from it.

Parapalaeoleon and Baisopardus both occur in the Crato

Formation and are clearly distinguished from each other

(see diagnosis above). In general, all three genera differ in

the shape of the wing: wide basally with a somewhat elon-

gated apex in Palaeoleon, narrower basally and widest at

mid-length in Parapaleoleon; maximum width after the

mid-wing in Baisopardus. The wing shape of Baisopardus

araripensis, however, is more similar to that of Parapalaeo-

leon magnus, than to other species of Baisopardus, but it

possesses other character states of Baisopardus, e.g. Rs1

originates near the base of Rs and is deeply branched.

Parapalaeoleon magnus sp. nov.

Text-figures 3–4

Derivation of name. Latin adjective, magnus, large, in reference

to the large size of the species.

Holotype. SMNS 66000 ⁄ 268. An incomplete specimen exposed

in lateral aspect, including head, thorax with appendages, and

right forewing.

Type locality and horizon. As for Nuddsia longiantennata above.

Diagnosis. As for the genus.

Description. Head 4 mm long. Compound eyes 2.5 mm in diam-

eter. Antennae 25 mm long, composed of 71 preserved segments:

scape not preserved; first visible segment wider than other flagel-

lar segments, which are square basally, transverse distad. Thorax

not complete (9 mm long as preserved), compressed, exoskeleton

mostly not preserved; prothorax apparently comparatively short.

Foreleg: coxa 2 mm long; femur rather stout, 6 mm long; tibia

stout, 5 mm long, armed with apical spine; tarsus 4 mm long,

five-segmented, distalmost segment approximately as long as four

basal segments together; claws very long, 2 mm. Midleg: coxa

2 mm long; femur 6 mm long; tibia 4.5 mm long, apical spines

not detected; tarsus as in forelegs. Hind legs not preserved.

Forewing widest at mid length, 67–68 mm long (estimated), c.

20 mm wide. Costal space narrow, 1.5 mm wide in basal part,

slightly narrowed toward fusion of Sc, R1, strongly expanded

beyond this. Subcostal veinlets simple, perpendicular to Sc in

basal half of costal space, oblique, forked distally. Veinlets of

Sc+R1 long, very oblique, usually dichotomously forked, con-

nected by occasional crossveins not forming a regular gradate

series. Sc and R1 fused at 12 mm from wing apex, entering wing

margin apparently well after apex (apical portion of wing not

preserved). No crossveins in subcostal space. Pterostigma absent

or not discernible. Stem of Rs straight, not zigzagged for entire

length; its origin situated near wing base, inclined at a very acute

angle to R1. No presectorial crossveins detected. R1 space most

dilated at middle portion, strongly narrowed basad and distad;

85 very closely spaced crossveins proximal to fusion of Sc, R1.

Long hypostigmal cell absent. Rs with 13 branches, somewhat

sigmoid in configuration, branched dichotomously distally. Ori-

gin of Rs1 located 21 mm from origin of Rs; branches of Rs

dichotomously branched distally; Rs1 branched much distal to

Banksian fold. Rs2, Rs3 appear fused. Median flexion line dis-

tinct, running between MA, Rs, then Rs1 from wing base to Rs3.

Anterior Banksian fold distinct, beginning slightly distal to Rs1,

directed to wing apex (somewhat posterior to it), unaccompa-

nied by false longitudinal vein formed by bending of branches of

Rs and crossveins connecting them. Radial crossveins very

numerous, not forming gradate series. Origin of M not pre-

served, probably not fused with R at wing base. Fork of M

rather distinct, with origin of MA forming shallow angle with

stem of M, and MP (‘oblique vein’) stouter than crossveins in

median space. MA simple, with only shallow marginal fork,

straight, curved posteriorly distally. ‘Oblique vein’ well devel-

oped, located slightly proximal to most proximal branch of

MP+CuA. CuA, MP+CuA strongly convex, both parallel to MA,

dense crossveins for entire length between these and MA.

MP+CuA with 12 branches, of which most proximal branch

(MP+CuA1) pectinately branched; next branch (MP+CuA2)

fused with it; MP+CuA3 with deep fork; other branches shal-

lowly forked. All branches of MP+CuA connected by numerous


crossveins; near posterior margin crossveins connected by addi-

tional crossveins forming short irregular pseudo-longitudinal

veins. Crossveins between CuA (also MP+CuA1), CuP numer-

ous, dense. CuP concave basally, rather short, with shallow mar-

ginal fork, seven simple branches, crossveins between them rare.

1A not fused basally with CuP. Two short unbranched anal veins

preserved (1A, 2A).

Family MESOCHRYSOPIDAE Handlirsch, 1906

Genus TRIANGULOCHRYSOPA Nel et al., 2005

Type species. Triangulochrysopa sanzi Nel et al., 2005, by original

designation.

A

C

B

TEXT -F IG . 3 . Parapalaeoleon magnus


66000 ⁄ 268. A, photograph of the

specimen. B, close-up of forewing

showing fusion of MP and CuA (the

names of the veins are partly

hypothetical; see text for detail). C,

portion of forewing showing median

flexion line (mfl) and anterior Banksian

fold (bf). Scale bars represent 10 mm.

TEXT -F IG . 4 . Parapalaeoleon magnus


66000 ⁄ 268. A, anterior part of body. B,

forewing. Scale bars represent 5 mm.


Diagnosis. Prothorax somewhat elongated. Forelegs

rather short, probably raptorial. Forewing: MP touching

(or almost touching) CuA; venation in radial to cubital

spaces highly reticulated. Hindwing triangular in shape

with distinct tornus, rather long (0.67 times forewing

length); apex of wing acute; MP appears to be continua-

tion of M; MA arising from M at right angle or nearly

so.

Included species. Triangulochrysopa sanzi (Lower Cretaceous, Bar-

remian, Las Hoyas, Spain, La Huerguina Formation), T. formosa

sp. nov. (Lower Cretaceous, Upper Aptian, Araripe Basin, Brazil,

Crato Formation).

Remarks. Triangulochrysopa is most similar to the Lower

Cretaceous genera Allopterus Zhang, 1991 and Karenina

Martins-Neto, 1997 in the structure of the pronotum,

and its foreleg and forewing venation. It may be easily

distinguished from these by hindwing size, shape and

venation: e.g. the hindwing is short and broad, c. 0.4 of

forewing length, with rounded apex and reduced venation

in Allopterus, and relatively narrow, without tornus, 0.6–

0.8 of forewing length, with different venation (in partic-

ular, Rs inclined at a more acute angle to R1, with Rs1

originating more distally) in Karenina (Makarkin and

Menon 2005; Nel et al. 2005).

According to Nel et al. (2005), Triangulochrysopa (with

Allopterus, Karenina and Armandochrysopa Nel et al.,

2005) belongs to the family Allopteridae in the superfam-

ily Chrysopoidea. They proposed the following phyloge-

netic relationship: (Cratochrysa Martins-Neto, 1994 +

(Liassochrysidae + (Paralembochrysa Nel et al., 2005 +

((Allopteridae + Mesochrysopidae + Tachinymphidae +

Mesotermes Haase, 1890) + (Limaiidae + Chrysopi-

dae))))). We have previously regarded Mesotermes and

the genera of Allopteridae, Mesochrysopidae and Tachi-

nymphidae as forming the single, monophyletic family

Mesochrysopidae (Makarkin and Menon 2005). We are

still of the opinion that the elevation of these groups

(and Limaiidae) to family rank is unjustified.

Triangulochrysopa formosa sp. nov.

Text-figures 5–6

2001 Unnamed neuropteran; Bechly et al. p. 55, fig. 45.

Derivation of name. Latin adjective, formosus, graceful, beautiful,

with reference to the appearance of the holotype.

Holotype. SMNS 66000 ⁄ 271. A complete well-preserved speci-

men exposed in ventral aspect, with all wings wide open; disar-

ticulated fragments of legs are visible along the abdomen in the

matrix.

Type locality and horizon. As for Nuddsia longiantennata above.

Diagnosis. Triangulochrysopa formosa may be distin-

guished from T. sanzi by the following forewing charac-

ters: forewing more elongate, length ⁄ width ratio 4.0 [3.3

in the holotype of T. sanzi], pterostigma distinct [absent

(or not detectable) in T. sanzi], MP touching CuA [con-

nected by short crossvein in T. sanzi], 1A with two

branches [one in T. sanzi].

Description. Head oval, wider than long, 4.5 mm long, 2.6 mm

wide with large compound eyes, each 1 mm in diameter. Anten-

nae incomplete, 13 mm long (estimated), filiform; scape not pre-

served, flagellum composed of at least 40 equal segments.

Prothorax elongate, 3.5 mm long, 2.3 mm wide; mesothorax and

metathorax oval, robust, 8 mm long, 4 mm wide. Legs only

partly preserved: fragmentary coxae on all three pairs of legs;

right foreleg: femur rather stout, 3.5 mm long, tibiae 3.7 mm

long; disarticulated tarsus (belonging probably to left midleg)

with five cup-shaped segments (total length 3 mm), bearing two

distinct claws. Abdomen composed of nine segments, 22 mm

long.

Forewing: elongate with sub-acute apex, 44 mm long, 11 mm

wide (length ⁄ width ratio 4.0). Costal space narrow, slightly

dilated towards pterostigma. Subcostal veinlets proximal to

pterostigma simple (one forked, obviously an anomaly), 36 in

number, perpendicular to Sc or inclined to apex, becoming

increasingly inclined, closely spaced to pterostigmal region; ante-

rior tip of most basal subcostal (humeral) veinlet inclined

towards base. Veinlets of Sc, Sc+R1 distal to pterostigma mostly

forked, connected by 1–2 crossveins, forming 2–3 rows of cells

between C, Sc+R1. Pterostigma dark coloured, 3.5 mm long,

covering seven cells. No crossveins in subcostal space detected.

Sc, R1 fused distal to pterostigma. Sc+R1 entering wing margin

at apex. Stem of Rs straight, zigzagged distally; its origin 4.5 mm

from wing base. R1 space dilated at origin of Rs1, 1.3 mm

(wider than width of costal space), narrowed towards apex, with

28 crossveins; Rs with 20 zigzagged branches not forked before

marginal forks, each with 1–2 crossveins connected to branches

of fork. Venation in radial space highly reticulated, not forming

distinct gradate series. In radial space two longitudinal conver-

gent folds present. No crossvein between stem of Rs, M. Origin

of M at 3 mm from wing base. M moderately convex, divided

into MA, MP at 8 mm from wing base and at relatively acute

angle (c. 45 degrees). MA smooth, slightly convex, entering mar-

gin well before wing mid-point, with marginal fork. MP zig-

zagged, without marginal fork, with two long zigzagged

branches. Crossveins in medial space not arranged in regular ser-

ies. Basal crossvein m-cu located slightly distal to origin of M,

oblique, long. Cu not fused with R basally, divided into CuA,

CuP distal to crossvein m-cu. CuA strongly convex, straight,

only slightly bent at MP touching, ending in very shallow mar-

ginal fork and with single short branch (it is difficult to identify

these with certainty as the venation in medial and cubical spaces

is highly reticulated; CuA could be also strongly zigzagged in

distal half, with two branches having each wide marginal fork).

CuA touching MP. CuP short, with two short branches. Basal


crossvein cua-cup oblique, long, situated slightly proximal to

meeting point of MP, CuA; distal crossvein cua-cup oblique,

rather short, situated far distal to meeting point of MP and

CuA. At least one distal crossvein between CuP, 1A. 1A, 2A

nearly parallel to hind margin; 1A with one branch; 2A not

branched. 3A not detected.

Hindwing: subtriangular in shape with somewhat pointed

apex, 28 mm long, 8.5 mm maximum width. Costal space

equally narrow throughout length (0.5 mm). Subcostal veinlets

simple, forming 31 cells proximal to pterostigma. Pterostigma

short (1 mm long), covering three cells, dark coloured. Sc and

R1 fused distal to pterostigma. No crossveins in subcostal space

detected. Origin of Rs at 5.7 mm from wing base. R1 space

1 mm at its widest point, with 17 crossveins; hypostigmal cells

short. Stem of Rs zigzagged, with 12 zigzagged branches, simple

(distal branches) or with shallow, wide marginal fork (proximal

branches). Crossveins in radial space numerous, not forming

distinct gradate series. M dividing into MA, MP at 4 mm from

wing base; MP appears to be a continuation of M, MA arising

from M nearly at right angle. MA smooth, only slightly arched,

with marginal fork entering wing margin at mid-point (15 mm

from wing base). MP zigzagged, divided into two long branches.

CuA strongly convex basally, touching MP at nearly half of its

length, strongly zigzagged distally, with seven short branches.

Venation between MA, CuA strongly reticulated (especially dis-

tally, making it impossible to discriminate with certainty longi-

tudinal veins from crossveins). CuP, anal veins not detected.

Remarks. Both species of the genus, T. sanzi and T. for-

mosa, are very similar to each other. T. sanzi was

described from five specimens from the Barremian of Las

Hoyas, Spain, which differ somewhat in size (forewing

length ranges from 27.5 to 39.2 mm) and venation. Nel

A

B

TEXT -F IG . 5 . Triangulochrysopa formosa sp. nov. A, photograph, and B line drawing of the holotype, SMNS 66000 ⁄ 271. Scale bar

represents 10 mm.


et al. (2005) considered these differences insufficient to

separate them; however, some may be significant at a spe-

cific level. For example, the hindwing venation of the

specimens 92 ⁄ 2 ⁄ 3 (Nel et al. 2005, fig. 7.5) and LH-18572

(Nel et al. 2005, fig. 7.4) of T. sanzi is very different; they

are 19.2 mm and 21.6 mm long respectively, whereas the

hindwing of the holotype is 31.6 mm long (unfortunately,

this wing was not figured). The hindwing ⁄ forewing length

ratio ranges from 0.55 to 0.66 in these paratypes, and is

0.95 in the holotype. The photograph of the holotype

does not reveal such long hindwings; the measurement

should probably be 21.6, not 31.6 mm. In any case, after

this discovery of another very similar species from a

South American locality distant from southern Europe, all

material previously identified as T. sanzi should be

revised.

Acknowledgements. We thank Dr Gunter Bechly and Dr Gerd

Dietl (SMNS) for allowing us to study the material we have

described; Dr Andre Nel (Museum national d’Histoire naturelle,

Paris) and Prof. Dr Rafael Martins-Neto (Universidade Federal

de Juiz de Fora, Brazil) for providing us with advance copies of

their papers in press; Prof. Dr Ulrike Aspock and Dr Harald

Schillhammer (Naturhistorisches Museum Wien) for providing

the photograph of the holotype of Gumilla adspersa; Jean

Dougherty (Geological Survey of Canada, Ottawa, Canada), and

Bruce Archibald (Museum of Comparative Zoology, Harvard

University) for providing the photograph of the holotype of Pal-

aeoleon ferrogeneticus; Mr Robert Loveridge (University of Ports-

mouth) for providing the photograph of the holotype of

Baisopardus chryptohymen; Bruce Archibald for helpful com-

ments and for suggestions on the English; James Jepson (Univer-

sity of Manchester) for correcting our English; Dr Gunter Bechly

and Dr Rafael Giois Martins-Neto for reviewing the manuscript;

and Prof. David Batten (University of Manchester) for editorial

work. The visit to Stuttgart by FM was funded by the Deutscher

Akademischer Austausch Dienst (DAAD).

REFERENCES

A D A M S , P. A. 1969. A new genus and species of Osmylidae

(Neuroptera) from Chile and Argentina, with a discussion of

planipennian genitalic homologies. Postilla, 141, 1–11.

—— 1977. Taxonomy of United States Leucochrysa (Neuroptera:

Chrysopidae). Psyche, 84, 92–102.

—— 1996. Venational homologies and nomenclature in Chrysopi-

dae, with comments on the Myrmeleontoidea (Insecta: Neuro-

ptera). 19–30. In C A N A R D , M., A S P OC K, H. and

M A N S E L L , M. W. (eds). Pure and applied research in neur-

opterology. Proceedings of the Fifth International Symposium on

Neuropterology. Privately printed, Toulouse (France), 414 pp.

A R C HI B A L D, S. B. and M A K A R K I N , V. N. 2006. Tertiary

giant lacewings (Neuroptera: Polystoechotidae): revision and

description of new taxa from western North America and

Denmark. Journal of Systematic Palaeontology, 4, 119–155.

B A N K S , N. 1910. Myrmeleonidae from Australia. Annals of the

Entomological Society of America, 3, 40–44.

B E CH L Y , G., H A A S , F., S CH A W A L L E R , W., S C H M A L -

F US S , H. and S C H M I D, U. 2001. Ur-Geziefer – Die faszini-

erende Evolution der Insekten. Stuttgarter Beitrage zur

Nuturkunde, Serie C, 49, 1–96.

B E R T HO U , P. Y. 1994. Relationship between the ostracod

fauna and the oxic or anoxic character of the Aptian–Albian

strata of the Araripe basin (NE Brazil). Proceedings of the 14th

Sedimentological Congress, 14, G11.

C OM S T OC K , J. H. 1918. The wings of insects. Comstock

Publication Company, Ithaca, NY, 430 pp.

D OB R US KI N A , I. A., P ON O M A R E N KO , A. G. and

R A S N I T S Y N , A. P. 1997. Fossil insects found in Israel.

Paleontologicheskii Zhurnal, 1997 (5), 91–95. [In Russian].

D OL I N , V. G., PA N FI L OV , D. V., P ON OM A R E N KO ,

A. G. and PR I T Y KI N A , L. N. 1980. Fossil insects of the

Mesozoic. Naukova Dumka, Kiev, 135 pp. [In Russian].

E N D E R L E I N , G. 1910. Klassifikation der Mantispiden nach

dem Material des Stettiner Zoologischen Museums. Stettiner

Entomologische Zeitung, 71, 341–379.

TEXT-F IG . 6 . Triangulochrysopa formosa

sp. nov., holotype, SMNS 66000 ⁄ 271. A,

left forewing. B, left hindwing. Scale bar

represents 5 mm.


E S B E N - P E T E R S E N , P. 1918. Results of Dr. E. Mjoberg’s

Swedish scientific expeditions to Australia 1910–1913. 18. Neu-

roptera and Mecoptera. Arkiv for Zoologi, 11(26), 1–37.

H A A S E , E. 1890. Bemerkungen zur Palaeontologie der Insec-

ten. Neues Jahrbuch fur Mineralogie, Geologie und Palaontolo-

gie, 2 (1), 1–33.

H A N D L I R S C H, A. 1906–08. Die fossilen Insekten und die Phy-

logenie der rezenten Formen. Ein Handbuch fur Palaontologen

und Zoologen. Engelmann, Leipzig, ix + 1430 pp., 51 pls.

H A G E N , H. A. 1859. Beitrag zur Kenntniss der Neuropteren.

Stettiner Entomologische Zeitung, 20, 405–412.

H E A D S , S. W., M A R TI L L , D. M. and L O V E R I D GE , R. F.

2005. An exceptionally preserved antlion (Insecta, Neuroptera)

with colour pattern preservation from the Cretaceous of Bra-

zil. Palaeontology, 48, 1409–1417.

K R I V O K HA T S K Y , V. A. 1998. Antlions (Neuroptera, Myr-

meleontidae) of the Palaearctic Region (morphology, classifi-

cation, zoogeography). Unpublished PhD thesis, Zoological

Institute, Russian Academy of Sciences, St. Petersburg, 364 pp.

[In Russian].

K R UG E R , L. 1913. Osmylidae. Beitrage zu einer Monographie

der Neuropteren-Familie der Osmyliden. III. Literatur und

Katalog. Stettiner Entomologische Zeitung, 74, 193–224.

—— 1915. Osmylidae. Beitrage zu einer Monographie der Neur-

opteren-Familie der Osmyliden. VIII. Anhang II. Stettiner En-

tomologische Zeitung, 76, 60–87.

K UK A L O V A - PE CK , J. 1991. Fossil history and the evolution

of hexapod structure. 144–179. In N A UM A N N , D. (chief

ed.). The insects of Australia, Volume 1. Second edition. Mel-

bourne University Press, Carlton, 752 pp.

L A M BK I N , K. J. 1988. A re-examination of Lithosmylidia Riek

from the Triassic of Queensland with notes on Mesozoic

‘osmylid-like’ fossil Neuroptera (Insecta: Neuroptera). Mem-

oirs of the Queensland Museum, 25, 445–458.

L A T R E I L L E , P. A. 1802. Histoire naturelle, generale et particu-

liere de Crustaces et des Insectes. Volume 3. Dufart, Paris, 467 pp.

L E A C H , W. E. 1815. Entomology. 57–172. In B R E W S T E R ,

D. (ed.). Edinburgh Encylopaedia 9 (1). Edinburgh.

L I N N A E US , C. 1758. Systema natura per regna tria naturae

secundum classes, ordines, genera, species, cum characteribus,

differentiis, synonymis, locis. Tenth edition. Laurentius Salvius,

Holmiae, 824 pp.

M A I S E Y , J. C. (ed.) 1991. Santana fossils: an illustrated atlas.

Tropical Fish Hobbist Publications, Neptune City, NJ, 459 pp.

M A KA R K I N , V. N. 1990a. New lacewings (Neuroptera) from

the Upper Cretaceous of Asia. 63–68. In A K I M O V , I. A.

(ed.). News of faunistics and systematics. Naukova Dumka,

Kiev, 184 pp. [In Russian].

—— 1990b. A new fossil genus and species of Osmylidae from

the Lower Cretaceous of East Siberia (Neuroptera). Deutsche

Entomologische Zeitschrift, 37, 101–103.

—— and A R CH I B A L D , S. B. 2003. Family affinity of the

genus Palaeopsychops Andersen with description of a new spe-

cies from Early Eocene of British Columbia (Neuroptera:

Polystoechotidae). Annals of the Entomological Society of Amer-

ica, 96, 171–180.

—— and M E N ON , F. 2005. New species of the Mesochrysopi-

dae (Insecta, Neuroptera) from the Crato Formation of Brazil

(Lower Cretaceous), with taxonomic treatment of the family.

Cretaceous Research, 26, 801–812.

—— —— 2007. First record of fossil rapismatid-like Ithonidae

(Insecta, Neuroptera) from the Lower Cretaceous Crato

Formation of Brazil. Cretaceous Research, 28, 743–753.

M A R TI L L , D. M. 1993. Fossils of the Santana and Crato For-

mations, Brazil. Field Guide to Fossils No. 5. The Palaeonto-

logical Association, London, 159 pp.

—— and F R E Y , E. 1995. Colour patterning preserved in Lower

Cretaceous birds and insects: the Crato Formation of N. E.

Brazil. Neues Jarhbuch fur Geologie und Palaontologie, Monats-

hefte, 1995, 118–128.

M A R TI N S - N E T O , R. G. 1992. Neuropteros (Insecta, Plani-

pennia) da Formacao Santana (Cretaceo Inferior), Bacia do

Araripe, nordeste do Brasil. VII. Palaeoleontinae, nova sub-

famılia de Myrmeleontidae e descricao de novos taxons. Revi-

sta Brasileira de Entomologia, 36, 803–815.

—— 1994. Neuropteros (Insecta, Planipennia) da Formacao

Santana (Cretaceo Inferior), Bacia do Araripe, nordeste do

Brasil – IX – Primeiros resultados da composicao da fauna e

descricao de novos taxons. Acta Geologica Leopoldensia, 17

(39 ⁄ 1), 269–288.

—— 1997. Neuropteros (Insecta, Planipennia) da Formacao San-

tana (Cretaceo Inferior), Bacia do Araripe, nordeste do Brasil.

X. Descricao de novos taxons (Chrysopidae, Babinskaiidae,

Myrmeleontidae, Ascalaphidae e Psychopsidae). Revista Uni-

versidade de Guarulhos, Serie Ciencias Exactas e Technologicas,

2 (4), 68–83.

—— 1998. Neuropteros (Insecta, Planipennia) da Formacao

Santana (Cretaceo Inferior), Bacia do Araripe, nordeste do

Brasil. XI. Descricao de novos taxons de Myrmeleontidae

(Paleoleontinae e Pseudonymphinae). Revista Universidade de

Guarulhos, Serie Ciencias Biologicas e da Saude, 3 (5), 38–

42.

—— 2000. Remarks on the neuropterofauna (Insecta, Neuro-

ptera) from the Brazilian Cretaceous, with keys for the

identification of the known taxa. Acta Geologica Hispanica, 35,

97–118.

—— 2003. The Santana Formation paleoentomofauna reviewed.

Part I – Neuropteroida (Neuroptera and Raphidioptera): sys-

tematics and phylogeny, with description of new taxa. Acta

Geologica Leopoldensia, 25 (55) (for 2002), 35–66.

—— 2005. New Neuroptera (Nymphidae and Araripeneuridae)

from Santana Formation (Lower Cretaceous, Araripe basin,

northeast Brasil). GAEA, 1, 5–10.

—— and V UL C A N O, M. A. 1997. Neuropteros (Insecta,

Planipennia) da Formacao Santana (Cretaceo Inferior), Bacia

do Araripe, Nordeste do Brasil. VIII. Descricao de novas taxas

de Myrmeleontidae, Ascalaphidae e Nemopteridae. Revista

Universidade de Guarulhos, Serie Ciencias Biologicas e da Saude,

2 (5), 64–81.

M E N O N , F., M A R TI N S - N E T O , R. G. and M A R T I L L ,

D. M. 2005. A new Lower Cretaceous nymphid (Insecta, Neu-

roptera, Nymphidae) from the Crato Formation of Brazil.

GAEA, 1, 11–15.

N A V A S , L. 1909. Monografıa de la familia de los Dilaridos

(Ins. Neur.). Memorias de la Real Academia de Ciencias y Artes

de Barcelona, 3, 7, 619–671, 2 pls.


—— 1912. Insectos neuropteros nuevos o poco conocidos. Mem-

orias de la Real Academia de Ciencias y Artes de Barcelona, 3,

10, 135–202.

—— 1913. Nevropteres. 69–77, 1 pl. In: Mission du Service Geo-

graphique de l’Armee pour la mesure l’un arc de meridien equa-

torial en Amerique du Sud sous le controle scientifique de

l’Academie des Sciences 1899–1906. Tome 10, Fascicle 1. Paris.

N E L , A., D E L CL O S , X. and H U TI N , A. 2005. Mesozoic

chrysopid-like Planipennia: a phylogenetic approach (Insecta:

Neuroptera). Annales de la Societe Entomologique de France,

41, 29–69.

N E W , T. R. 1985. A revision of the Australian Myrmeleontidae

(Insecta: Neuroptera). I. Introduction, Myrmeleontini, Proto-

plectrini. Australian Journal of Zoology, Supplementary Series,

104, 1–90.

O S W A L D , J. D. 1993. Revision and cladistic analysis of the

world genera of the family Hemerobiidae (Insecta: Neuroptera).

Journal of the New York Entomological Society, 101, 143–299.

P A N F I L O V , D. V. 1980. New representatives of lacewings

(Neuroptera) from the Jurassic of Karatau. 63–68, 7 pls. In

DO L I N , V. G., P A N F I L O V , D. V., P ON O M A R E N KO ,

A. G. and P R I T Y K I N A , L. N. Fossil insects of the Mesozoic.

Naukova Dumka, Kiev, 135 pp. [In Russian].

P ON O M A RE N KO , A. G. 1980. Superorder Myrmeleontidea

Latreille, 1802. 84–99, pl. 3, fig. 11. In R O H D E N D O R F ,

B. B. and R A S N I T S Y N , A. P. (eds). The historical develop-

ment of the class Insecta. Trudy Paleontologicheskogo Instituta,

175, 269 pp. [In Russian].

—— 1992a. Neuroptera (Insecta) from the Lower Cretaceous of

Transbaikalia. Paleontologicheskii Zhurnal, 1992 (3), 43–50,

Moscow. [In Russian: English translation, Paleontological

Journal, 1992, 26 (3), 56–66].

—— 1992b. New lacewings (Insecta, Neuroptera) from the

Mesozoic of Mongolia. 101–111, pls 27–29. in G R UN T ,

T. A. (ed.). New taxa of fossil invertebrates of Mongolia. Trans-

actions of the Joint Soviet-Mongolian Paleontological Expedi-

tion, 41. Nauka Press, Moscow, 142 pp. [In Russian].

—— 2003. On some Neuroptera (Insecta) from Upper Jurassic

Solnhofen Limestone. Annals of the Upper Silesian Museum

(Entomology), 12, 87–100.

P ON S , D., BE R T H O U, P. Y. and C A M P OS , D. A. 1991.

Quelques observations sur la palynologie de l’Aptien Superieur

et de l’Albien du Bassin d’Araripe (N. E. du Bresil). 241–252.

In C A M P OS , D. A., V I A N N A , M. S. S., B R I T O , P. M.

and B E UR L E N , G. (eds). Atas do I Simposia sobre a Bacia

do Araripe e Bacias Interiores do Nordeste, Crato, 14–16 Junho

de 1990. Departmento Nacional de Producao Mineral 10�Distritio (DNPM), Universidade Regional do Cariri

(URCA) e Sociedade Brasileira de Paleontologia (SBP), Crato,

405 pp.

R A M BU R , J. P. 1842. Histoire naturelle des insects. Neuropteres.

Fain et Thunot, Paris, xviii + 534 pp., 12 pls.

R E N , D. and Y I N , J. 2002. A new Middle Jurassic species of

Epiosmylus from Inner Mongolia, China (Neuroptera: Osmyli-

dae). Acta Zootaxonomica Sinica, 27, 274–277. [In Chinese,

English summary].

R I C E , H. M. A. 1969. An antlion (Neuroptera) and a stonefly

(Plecoptera) of Cretaceous age from Labrador, Newfoundland.

Geological Survey of Canada, Department of Energy, Mines and

Resources, Paper, 68-65, iv + 11 pp., 2 pls.

S E L D E N , P. A. 2003. A new tool for fossil preparation. The

Geological Curator, 7, 337–339.

T I L L Y A RD , R. J. 1916. Studies in Australian Neuroptera. No.

I. The wing-venation of the Myrmeleontidae. Proceedings of

the Linnean Society of New South Wales, 40 (for 1915), 734–

750.

—— 1918. Studies in Australian Neuroptera. No. 5. The

structure of the cubitus in the wings of the Myrmeleontidae.

Proceedings of the Linnean Society of New South Wales, 43,

116–122.

W A L K E R , F. 1853. List of the specimens of neuropterous insects

in the collection of the British Museum. Part II. British

Museum, London, pp. 193–476.

W E D M A N N , S. and M A K A R KI N , V. N. 2007. A new genus

of Mantispidae (Insecta: Neuroptera) from the Eocene of

Germany, with a review of the fossil record and palaeobio-

geography of the family. Zoological Journal of the Linnean Soci-

ety, 149, 701–716.

W O O TT O N , R. J. 2002. Design, function and evolution in the

wings of holometabolous insects. Zoologica Scripta, 31, 31–40.

—— 2003. Wings. 1186–1192. In RE S H, V. H. and CA R D E ,

R. T. (eds). Encyclopedia of insects. Academic Press, London,

1266 pp.

Z H A N G , J. 1991. A new family of Neuroptera (Insecta) from

the late Mesozoic of Shandong, China. Science in China, Series

B, 34, 1105–1112.


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