Early Cretaceous Snakefly Larvae in Amber from Lebanon, Myanmar, and France (Raphidioptera)

2007·3.7 MB·English

by  Vincent Perrichot

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3598, 11 pp., 8 figures, 1 table December 12, 2007
Early Cretaceous Snakefly Larvae in Amber from
Lebanon, Myanmar, and France (Raphidioptera)
VINCENT PERRICHOT1 AND MICHAEL S. ENGEL2
ABSTRACT
Snakefly (Raphidioptera) larvae are newly documented from the Early Cretaceous ambers of
Lebanon, Myanmar (Burma), and France. Previously only two Cretaceous larvae had been
documented, one in Late Cretaceous (Turonian) amber from New Jersey and another in Early
Cretaceous (Albian) amber from Myanmar. The specimens discussed herein are likely
representativeoftheextinctfamilyMesoraphidiidae,butdefinitivefamilialassignmentiscurrently
not possible. The new fossil material is described and placed into context with the known larval
morphology of modern and fossil species, as well as with the geological history of the order as
documentedbythe remains of adults.
RE´SUME´
Denouvelleslarvesderaphidiopte`ressontmentionne´esdanslesambresduCre´tace´ infe´rieurdu
Liban, du Myanmar (Birmanie) et de France. Seules deux larves cre´tace´es avaient e´te´ signale´es
jusqu’a` pre´sent, dans l’ambre cre´tace´ supe´rieur (Turonien) du New Jersey et l’ambre cre´tace´
infe´rieur(Albien)duMyanmar.Lesspe´cimensdiscute´siciappartiennentprobablementa` lafamille
fossileMesoraphidiidae,maisuneattributionfamilialecertainedemeureactuellementimpossible.
Lenouveaumate´rielfossileestde´critetplace´enregarddelamorphologiedesespe`cesmoderneset
fossiles, ainsique del’histoire ge´ologique del’ordre telque documente´ parles restesd’adultes.
1Museumfu¨rNaturkundederHumboldt-Universita¨tzuBerlin,Institutfu¨rPala¨ontologie,Invalidenstraße43,D-10115
Berlin, Germany ([email protected]). Present address: Department of Geology, 1475 Jayhawk
Boulevard,LindleyHall,UniversityofKansas,Lawrence,Kansas66045.
2DivisionofInvertebrateZoology,AmericanMuseumofNaturalHistory;DivisionofEntomology(Paleoentomology),
Natural History Museum, and Department of Ecology & Evolutionary Biology, 1501 Crestline Drive-Suite #140,
UniversityofKansas,Lawrence,Kansas66049-2811([email protected]).
CopyrightEAmericanMuseumofNaturalHistory2007 ISSN0003-0082
2 AMERICAN MUSEUMNOVITATES NO. 3598
INTRODUCTION snakefly apomorphies as defined solely on
the basis of more modern species. For now,
Raphidioptera, or snakeflies, are presently the priscaenigmatids remain puzzling and it is
classified into six families, four of which are greatly hoped that more complete material
extinct and known only from the Mesozoic. willeventuallyberecovered.Acatalogueofall
The families Alloraphidiidae Carpenter, described fossil snakeflies was provided by
Baissopteridae Martynova, and Mesoraphi- Engel (2002) to which should be added
diidae Martynov are known exclusively from Sinoinocellia liaoxiensis Wang (1987) and five
the Jurassic and Cretaceous, whereas the fossils described subsequently from the
Inocelliidae and Raphidiidae comprise all Eocene-Oligocene of Florissant (Engel,
extant and some Tertiary species (Engel, 2003), in middle Eocene Baltic amber
2002). Three other fossil families from the (Aspo¨ck and Aspo¨ck, 2004), and the Early
Cretaceous of China—Huaxiaraphidiidae Cretaceous of southern Korea (Engel et al.,
Hong, Jilinoraphidiidae Hong and Chang, 2006). Table 1 summarizes the current classi-
and Sinoraphidiidae Hong—are now consid- ficationoftheorderaswellasthedocumented
ered synonyms of Mesoraphidiidae (Engel, geological ranges of the included lineages.
2002). The last family recognized, Pri- Although adult snakeflies are uncommon,
scaenigmatidaeEngel,issegregatedintoasep- theirimmature stagesareexceptionallyrare in
arate, basal suborder in some classifications the fossil record, with specimens previously
and its attribution to Raphidioptera remains known only as inclusions in amber from the
controversial (e.g., Willmann, 1994; Pono- middle Eocene of the Baltic region (Hagen in
marenko, 2002). This family contains only Pictet-Baraban and Hagen, 1856; Weidner,
the enigmatic genera Priscaenigma Whalley 1958; Weitschat and Wichard, 1998) and two
and Hondelagia Bode from the Early Jurassic inamberfromtheCretaceous(Grimaldi,2000;
ofEurope.Mostrecently,Aspo¨ckandAspo¨ck Engel,2002).Hereinweprovidedescriptionsof
(2004) dismissed the priscaenigmatids from the remains of immature snakeflies preserved
the order since these fossils do not exhibit the in Early Cretaceous amber from Lebanon
entire suite of apomorphies in forewing (Neocomian, perhaps Barremian), northern
venation that are used to define higher Myanmar (Late Albian), and southwestern
snakeflies. However, as noted by Engel France(LateAlbian).Severalofthespecimens
(2002) and Grimaldi and Engel (2005), pris- arepreservedonlybytheirheadcapsules,while
caenigmatids are likely stem-group Raphi- intwocasesa singleindividual,inFrenchand
dioptera as evidenced by the hind-wing Burmese amber respectively, also preserves
venation and would, therefore, not be ex- portions of the prothorax. Despite the frag-
pected to possess the complete suite of mentary nature of some of the material, the
TABLE1
Classification andGeological Distributionof Raphidioptera(updated fromEngel, 2002)
OrderRAPHIDIOPTERANava´s
Suborder{PRISCAENIGMATOMORPHAEngel
Family{PriscaenigmatidaeEngel Liassic
SuborderRAPHIDIOMORPHAEngel
Family{MesoraphidiidaeMartynov Liassic-Turonian
Family{BaissopteridaeMartynova Oxfordian-Cenomanian
Family{AlloraphidiidaeCarpenter Hauterivian-Cenomanian
InfraorderNeoraphidiopteraEngel
FamilyRaphidiidaeLatreille
Subfamily{SuccinoraphidiinaeAspo¨ck&Aspo¨ck Eocene
SubfamilyRaphidiinaeLatreille Eocene-Holocene
FamilyInocelliidaeNava´s
Subfamily{ElectrinocelliinaeEngel Eocene
SubfamilyInocelliinaeNava´s Eocene-Holocene
2007 PERRICHOT ANDENGEL: CRETACEOUS SNAKEFLY LARVAE 3
high fidelity of preservation in amber permits fact that in the Cretaceous and Tertiary
adetaileddescriptionofthecephalicstructures. periods, Raphidioptera were not as climati-
Furthermore, characters of the head capsule cally restricted as they are today. Although
alone permit positive identification as the order is today restricted to temperate to
Raphidioptera (see Discussion, below). While cold-temperate habitats or to high elevations
previously the specimen reported by Engel where milder climates occur in otherwise
(2002) in Burmese amber was the oldest warmer regions (e.g., some northern moun-
immature snakefly in amber (other immatures tains of Central America), in the past snake-
arenewlydescribedfromthesedepositsherein, flies were found in a diversity of habitats and
seebelow),thespecimenfromLebanonisnow biogeographic regions (e.g., Engel, 1995,
theoldestknownsnakeflylarva. 2002).
For the descriptions we follow the format
and terminology of Aspo¨ck et al. (1991) and
Engel (2002). SYSTEMATIC PALEONTOLOGY
Order RAPHIDIOPTERA Nava´s
GEOGRAPHICAL AND
GEOLOGICAL SETTING
FRENCH SPECIMENS
Two specimens(see figs. 1–2) were excavat-
Mesoraphidiidae? indet.
ed from the deposit of Archingeay/Les-
NouillersinCharente-Maritime(southwestern Figures 1–4
France), which already provided numerous
fossils, principally insects (Perrichot, 2004, MATERIALEXAMINED: Larvalsnakeflyhead
2005; Perrichot et al., 2007). The amber is capsule; MNHN-ARC-265.6 (figs. 1, 3); de-
derivedfromalternatinglayersofestuarineor posited in the Laboratoire de Pale´ontologie,
deltaic fine sand and clay containing mixed Muse´um national d’histoire naturelle, Paris,
fragments of fossil plants (cuticles and lignitic France.
wood).Theseamber-bearingstrataweredated Larval snakefly head capsule with protho-
aslatestAlbian(ca.100 Ma)bypalynological racic fragments; MNHN-ARC-328.4 (figs. 2,
studies (Ne´raudeau et al., 2002; Dejax and 4); deposited in the Laboratoire de Pale´on-
Masure,2005).Thecorrespondingpaleoenvir- tologie, Muse´um national d’histoire naturelle,
onmentisestimatedtohavebeenasubtropical Paris, France.
forest in estuarine or mangrove-like context, OCCURRENCE: Archingeay/Les-Nouillers,
underaglobalwarm-and-wetclimatebutwith Charente-Maritime, southwestern France;
possible dry periods (Perrichot, 2004, 2005). Early Cretaceous, Uppermost Albian (Ne´rau-
The densely fossil-packed Burmese amber deauetal.,2002).
has long been considered of Tertiary origin DESCRIPTION: ARC-265.6. Head slightly
but was recently confirmed to be of middle elongate, 0.4 mm in length (excluding mand-
Cretaceous age (reviewed in Zherikhin and ibles),0.3 mminwidth;sixstemmatadisposed
Ross,2000;Grimaldietal.,2002;Cruickshank in circle at base of antenna (fig. 3); ocelli
and Ko, 2003). The deposit is of Late Albian absent;sevenfine,rathershortsetaeonlateral
age(ca.100 Ma)andwasproducedinawarm, marginsofhead;coronalecdysialcleavageline
moist tropical paleoclimate and with the most complete, forking at about head midlength
southerly paleolatitude of the three deposits (fig. 3)(itsfracturingandtheseparationofthe
discussed herein (Grimaldi et al., 2002). head from other body sclerites suggest that
The Lebanese specimen is fossilized in the this was a cast exuvium). Antennae short,
Neocomian amber of Jezzine, which was, about one-half length of head (ratio ‘‘antenna
similar to the French amber, deposited in length/head length’’ 5 0.625), with three
a deltaic plain under a tropical to subtropical cylindrical articles of 0.09/0.10/0.06 mm in
climate (Azar, 2000). length, respectively; only two fine setae at
The occurrence of these snakeflies in trop- apex of second and third articles of right
icalorsubtropicalforestsfurtheratteststothe antenna (fig. 3). Mandibles with three distinct
4 AMERICAN MUSEUMNOVITATES NO. 3598
Figs.1–2. PhotomicrographsoflarvalsnakefliesinFrenchamber.1.HeadcapsuleofspecimenMNHN-
ARC-265.6 in dorsal view. 2. Head capsule and prothorax of specimen MNHN-ARC-328.4. Scale bars 5
0.25 mm.
teeth, third tooth biseriate, incisors asymmet- palpomeres, palpomeres slightly tapering pro-
rical; left apical tooth broken; clypeus and gressivelyinwidth;onlyrightmaxillarypalpus
labrum trapezoidal or nearly subquadrate; preserved, with four short palpomeres, P
4
aseriesofsixfinesetaeonlabrum,twolateral tapering to acute apex. Head with distinct
setae being twice length of other setae; two narrowing and with minute lip posterior to
strong and short setae on each lateral margin this constriction representing raphidiopteran
of labium; labial palpus with three short collar (figs. 1, 3).
2007 PERRICHOT ANDENGEL: CRETACEOUS SNAKEFLY LARVAE 5
Fig. 4. Line illustration of specimen MNHN-
ARC-328.4inFrenchamber.Scalebar50.25 mm.
Early Cretaceous, Upper Albian (Grimaldi et
al.,2002;CruickshankandKo,2003).
DESCRIPTION: Bu-507. Head slightly elon-
gate, 1.6 mm in length (excluding mandibles),
1.3 mm in width; seven stemmata disposed in
rough oval at base of antenna (fig. 5); ocelli
absent;scattered,finesetaeonlateralmargins
of head; coronal ecdysial cleavage line com-
plete, forking just apical head midlength
(fig. 5) (its fracturing and the separation of
the head from other body sclerites, like ARC-
265.6, suggest that this was a cast exuvium).
Fig. 3. Dorsal (top) and ventral (bottom) line Antennaeaboutone-halflengthofhead(ratio
illustrations of specimen MNHN-ARC-265.6 in
‘‘antenna length/head length’’ 5 0.456), with
French amber. Scalebar50.25 mm.
three cylindrical articles of 0.26/0.26/0.21 mm
inlength,respectively;afew,finesetaeatapex
ARC-328.4. Nearly twice the size of ARC-
of each antennal article. Mandibles with three
265.6butotherwiseidenticalinfeaturesexcept
distinct teeth, third tooth biseriate, incisors
that the antennae are slightly longer (whether
slightly asymmetrical, space between first and
this is enough to indicate that they were from
second tooth greater on left mandible, apical
different species is uncertain as the length is
toothlargestinbothmandibles;clypeusshort,
within the realm of natural size variation).
transverse;labrumshort,transverse,withafew
Unlike ARC-265.6, ventral part of prothorax
minute setae; labial and maxillary palpi not
preserved (figs. 2, 4), nearly equal in length
visible (ventral aspect of head obscured).
and width to that of head capsule.
Lateral margins of head slightly and gently
convex; head with distinct posterior constric-
tion forming posterior collar (fig. 5).
BURMESE SPECIMENS
Mesoraphidiidae? indet. 1 Mesoraphidiidae? indet. 2
Figure 5 Figure 6
MATERIALEXAMINED: Snakefly larval head MATERIALEXAMINED: Snakefly larval head
capsule, AMNH Bu-507, deposited in the capsule and prothorax, AMNH Bu-644, de-
Amber Fossil Collection, Division of posited in the Amber Fossil Collection,
Invertebrate Zoology, American Museum of Division of Invertebrate Zoology, American
Natural History, New York. Museum of Natural History, New York.
OCCURRENCE: NorthernMyanmar,Kachin, OCCURRENCE: NorthernMyanmar,Kachin,
TanaiVillage,105 kmNorthwestofMyitkyna; TanaiVillage,105 kmNorthwestofMyitkyna;
6 AMERICAN MUSEUMNOVITATES NO. 3598
Figs.5–6. PhotomicrographsoflarvalsnakefliesinBurmeseamber.5.HeadcapsuleofAMNHBu-507
(dorsal view). 6.Head capsuleand prothoraxof AMNHBu-644 (ventralview). Scalebars5 0.5mm.
Early Cretaceous, Upper Albian (Grimaldi et length, about three-quarters length of head
al.,2002;CruickshankandKo,2003). (ratio ‘‘antenna length/head length’’ 5 0.734),
DESCRIPTION: Bu-644. Head slightly elon- with three cylindrical articles of 0.18/0.17/
gate, 0.64 mm in length (excluding mand- 0.12 mm in length, respectively; sparse, fine
ibles), 0.53 mm in width; five stemmata setae at apex of articles (not discernible for
disposed in semicircle at base of antenna; third article). Mandibles strongly angulate in
lateral margins of head with sparse, fine setae apical third, with three distinct teeth, third
on lateral margins. Antennae of moderate tooth biseriate, incisors asymmetrical, apical-
2007 PERRICHOT ANDENGEL: CRETACEOUS SNAKEFLY LARVAE 7
Fig. 7. Photomicrograph of head capsule (dorsal view left, ventral view right) of specimen JG-13 in
Lebaneseamber. Scalebars5 0.25mm.
most incisor greatly elongate and sharply no visible stemmata or ocelli; a series of fine
acute at apex, nearly twice as long as second setae on lateral margins of head, two longer
incisor; base of mandibles unusually broad, setae at base of antenna. Three antennal
giving mandibles a stout appearance. articles, articles progressively increasing in
Mouthparts poorly preserved such that max- length from base to apex (lengths 0.07/0.08/
illary and labial palpi are indiscernible. 0.1 mm, respectively) and together slightly
Lateral margins of head relatively parallel, shorter than head capsule (ratio ‘‘antenna
giving head quadrate appearance; head with length/head length’’ 5 0.625); a minute but
strong posterior constriction and distinct lip distinct spine on outer apex of second
representing raphidiopteran collar (fig. 6). antennal article (visible on right antenna);
Prothoraxwithproportionslikethoseofhead aseriesoffinesetaeoneacharticle.Mandibles
capsule (fig. 6), with sparse setae (unlike the tridentate, teeth distinct, third tooth biseriate,
specimens previously discussed, Bu-644 ap-
pearstobeadeceasedlarvaratherthanacast
exuvium: fig. 6).
LEBANESE SPECIMEN
Raphidioptera: Raphidiomorpha indet.
Figures 7–8
MATERIALEXAMINED: Larvalsnakeflyhead
capsule; JG-13; provisionally deposited in the
Laboratoire de Pale´ontologie, Muse´um
National d’histoire Naturelle, Paris, France.
OCCURRENCE: Jezzine, southern Lebanon;
Fig. 8. Dorsal (left) and ventral (right) re-
Early Cretaceous, Upper Neocomian (Barre-
constructionofspecimenJG-13inLebaneseamber.
mian?).
Ventral surface partly covered by dust and micro-
DESCRIPTION: JG-13. Head elongate and scopic bubbles (indicated by stippled areas). Scale
narrow, 0.4 mm in length, 0.28 mm in width; bar5 0.25 mm.
8 AMERICAN MUSEUMNOVITATES NO. 3598
incisors slightly asymmetrical; labrum rectan- more subquadrate labra of other species).
gular and longer than wide; ventral surface Naturally, there are just as many differences
partly covered by dust and microscopic suchasthevestigialcollar,thegreatlyelongate
bubbles so that base of labial and maxillary antennal articles (lacking a minute outer,
palpi hidden, with three and four palpomeres apical spine on the second article), and the
visible, respectively. Distinct collar present; elongate posterior setae in the Burmese
prothorax preserved by only a minute frag- species.TheBurmesespecimenwasconsidered
ment of its anterior margin. a putative mesoraphidiid and additional wing
fragmentssubsequentlydiscoveredinBurmese
amber are all of mesoraphidiids although of
DISCUSSION
species larger and differing in venation from
The head capsules are undoubtedly of the adult described by Engel (2002) (Engel,
raphidiopteran larvae as evidenced by the unpubl. data). These observations of only
following characters: shape elongate and mesoraphidiid fragments in Burmese amber
relatively narrow (more extreme in the further strengthen the possibility that the
Lebanese specimen and in the species de- larvae are of the same family. By extrapola-
scribed by Engel, 2002); antenna with three tion, if the shared features between Engel’s
articles; head with fine, long setae near the (2002)BurmeseamberlarvaandtheLebanese
lateral margins; distinct collar present. In specimen described herein are indicative of
addition, the specimens share the asymmetric somedegreeofrelationship,thenitispossible
development of the mandibles and relatively that the latter is also representative of
similar antenna/head ratios (i.e., antenna Mesoraphidiidae.Asimilarheadshapeisalso
length/head length). seen in the New Jersey amber (Turonian)
The Lebanese individual is not without its larval snakefly described by Grimaldi (2000)
own peculiarities, differing from all other and which was assuredly a mesoraphidiid.
raphidiopteran larvae by the absence of Whileplacementwithin Baissopteridae,which
stemmata,butalsointerestingforitsrelatively would certainly seem possible from a biogeo-
shortantennaeandthemoredevelopedcollar. graphicperspective(e.g.,Aspo¨ck,1998;Engel,
The absence of stemmata assuredly excludes 2002), cannot be excluded, it currently lacks
placement in the Neoraphidioptera as all any evidence. Naturally, an assignment to
known larvae of Raphidiidae and Inocel- Mesoraphidiidae for the Lebanese specimen
liidae have stemmata (and furthermore these remains largely speculative but is not without
familiesarenotknownpriortotheTertiaryas some ancillary evidence, as we have noted
evidenced by relatively decent record of adult (unlike the baissopterid possibility). Allora-
Raphidioptera: table 1). The Lebanese speci- phidiidae likely renders Mesoraphidiidae
men differs most notably from the species in paraphyletic and is, therefore, not a viable
French amber by the absence of stemmata, ‘‘alternative’’ familial assignment as this
aminutespineattheouterapexofthesecond wouldlikelymerelymakethelarvaaparticular
antennal article (observable on the right form of mesoraphidiid.
antenna),longermaxillarypalpi,andthemore The new larval material from Burmese
elongate head. In some of these attributes the amber differs considerably from the peculiar
Lebanese specimen is similar to the species species described by Engel (2002) and the
described by Engel (2002) from Burmese remains are more typical of the species in
amber. Both species have more strongly Frenchamber.Allcanbereadilydistinguished
elongate heads, albeit even more strongly from that species by the greatly elongate head
developed in the Burmese specimen. In addi- andantennaeintheformerspecies(seeEngel,
tion, the coronal ecdysial cleavage lines are 2002). Bu-507 is, of course, easily differentiat-
weak (nearly imperceptible or absent in the ed from Bu-644 as well as the French species
Burmese species: see Engel, 2002) or incom- by the significantly larger size, being nearly
plete (in the Lebanese specimen) and the three times the length of the others. Aside
labrum is more rectangular with its longest from size, Bu-507 is generally quite similar to
axis oriented longitudinally (rather than the the French material differing most notably in
2007 PERRICHOT ANDENGEL: CRETACEOUS SNAKEFLY LARVAE 9
the slightly more elongate antennal articles, and intersegmental membranes as suggested
the short and transverse clypeus and labrum by Grimaldi (2000: 263). Unfortunately,
(these are more subquadrate in the French mouthparts and all but one stemma also are
species), and the presence of an additional hidden in this fossil by the froth of micro-
stemma in the larval eye (seven in Bu-507, six scopic bubbles, which prohibit a more precise
in ARC-265.6). The second Burmese species comparison with our specimen. The French
(Bu-644), while of more diminutive propor- species is also quite similar to the snakefly
tions, has some additional differences from head capsule newly recorded herein from
Bu-507 indicating that it was perhaps a sepa- Burmese amber (fig. 5) as detailed in the
rate species. Only five stemmata can be preceding paragraph.
discerned in Bu-644 (a lateral view is not According to Aspo¨ck et al. (1991: 74), the
possible as the specimen is situated in a flow French specimen would fall into the modern
plane),thelateralmarginsoftheheadcapsule familyRaphidiidaeratherthantheInocellidae
are relatively straight and parallel, and the owing to the absence of a short sclerite
mandibles aremore strongly angulateapically between the basal two articles of its antenna
(this latter feature also serves to distinguish and the presence of six stemmata gathered at
the species from all other Cretaceous snakefly the base of its antenna (in contrast with the
larvae). four present in inocelliids). However, each of
It is interesting to note that the Burmese these traits is considered plesiomorphic and
amber fauna consists of three distinct larval likely represents the primitive condition for
types (two described herein and a third by the clade Raphidiidae + Inocelliidae (i.e., the
Engel,2002)inadditiontoatleasttwospecies Neoraphidioptera: table 1), if not for raphi-
documented from adults (Nanoraphidia elec- diomorphan snakeflies as a whole (interest-
troburmica Engel and another known only ingly, the family Raphidiidae itself is largely
from wing fragments: Engel, unpubl. data). It defined by plesiomorphies relative to
would appear that the Burmese amber fauna Inocelliidae and the former may indeed
mayeventuallyrivalinnumberofspeciesthat eventually prove to be paraphyletic). Though
of the more well-studied Baltic amber fauna the French specimen has strong similarities
(Carpenter, 1956; Engel, 2002; Aspo¨ck and with some raphidiid larvae (cf. figs. 53–55 in
Aspo¨ck, 2004). Aspo¨ck et al., 1991), the preservation of the
The French amber species is quite distinc- head alone does not allow definitive assign-
tive from the putative mesoraphidiid larva ment to this family. Moreover, larvae are
describedpreviouslyfromBurmeseamber(see unknown for the extinct families Allora-
Engel,2002:fig.4)byitssmallersize,asmaller phidiidae and Baissopteridae, which are also
ratio ‘‘antenna length/head length’’ (1.18 vs. known to have occurred during the Creta-
0.625 in Burmese and French specimens ceous. We therefore cannot make a definitive
respectively), the presence of six stemmata, assignment as to family for our fossil.
the more typical head shape that is slightly However, based on the biogeographic distri-
longer than wide (greatly elongate in the bution of alloraphidiids, it is unlikely that the
Burmese species), the more subquadrate la- species belongs to this family (Carpenter,
brum(elongateintheBurmesespecimen),and 1968; Engel, 2002) (plus the aforementioned
the shorter antennae that just surpass the possibility that alloraphidiids are derived
apices of the mandibles (the antennae are mesoraphidiids partly vitiates this concern).
exceedingly elongate in the Burmese species, Similarly,Baissopteridaeareknownonlyfrom
with the first article alone surpassing the the Late Jurassic of eastern Asia (Martynova,
mandibular apices: see Engel, 2002). The 1961) and the Early Cretaceous of Brazil
French amber material is more similar to the (Martins-Neto and Vulcano, 1989; Oswald,
putative mesoraphidiid larva described from 1990;Neletal.,1990).Thus,itisperhapsmost
New Jersey amber (see Grimaldi, 2000: fig. 2) likely that the French specimen is representa-
by its similar head and antennal sizes. A tiveoftheMesoraphidiidae,particularlygiven
distinct collar is not visible in the New Jersey that raphidiids are entirely confined to the
specimen, although it islikely hidden by froth Cenozoic, the oldest being those preserved in
10 AMERICAN MUSEUMNOVITATES NO. 3598
middleEoceneBalticamber(Carpenter,1956; Raphidiopteren (Insecta: Neuropteroidea).
Aspo¨ck and Aspo¨ck, 2004). Mesoraphidiids, Krefeld: Goecke andEvers, 730pp.[vol. 1].
by contrast, are known from the Jurassic and Aspo¨ck,U.,andH.Aspo¨ck.2004.Twosignificant
CretaceousofEurasia(e.g.,Martynov,1925a, new snakeflies from Baltic amber, with discus-
sion on autapomorphies of the order and its
1925b; Martynova, 1947; Hong, 1982, 1992;
included taxa (Raphidioptera). Systematic
Hong and Chang, 1989; Ren, 1997; Engel,
Entomology 29(1):11–19.
2002; Engel et al., 2006) and the Late
Azar, D. 2000. Les ambres me´sozo¨ıques du Liban.
Cretaceous of North America (Grimaldi,
Ph.D.dissertation,Universite´ParisXI,164pp.
2000).Takentogether,itseemsmostplausible Carpenter, F.M.1956. The Balticamber snakeflies
that our specimen belongs to the Mesora- (Neuroptera). Psyche 63:77–81.
phidiidae. Carpenter, F.M. 1968. Cretaceous insects from
Labrador. 2. A new family of snake-flies
ACKNOWLEDGMENTS (Neuroptera: Alloraphidiidae). Psyche 74:
270–275.
Cruickshank, D., and K. Ko. 2003. Geology of an
We thank Prof. Didier Ne´raudeau
amber locality in the Hukawng Valley, north-
(Universite´ Rennes 1) who coordinated the
ernMyanmar.JournalofAsianEarthSciences
fieldexplorationforFrenchCretaceousamber
21(5):441–455.
anddiscoveredtheamberpiececontainingthe Dejax, J., and E. Masure. 2005. Analyse palynolo-
snakeflydescribedherein.Wearealsograteful gique de l’argile lignitife`re a` ambre de l’Albien
toEricDepre´ fordonatingthepieceofFrench terminal d’Archingeay (Charente-Maritime,
amberwiththesecondspecimen;toDr.Andre´ France). Comptes Rendus Palevol 4(1–2):
Nel(Muse´umnationald’histoirenaturelle)for 53–65.
bringing the Lebanese specimen to our atten- Engel, M.S. 1995. A new fossil snake-fly species
tion; to Dr. Dany Azar (University of Beirut) from Baltic amber (Raphidioptera: Inocel-
liidae). Psyche 102(3–4):187–193.
for its loan and permission to study; and to
Engel, M.S. 2002. The smallest snakefly
Dr. Antonio Arillo and an anonymous re-
(Raphidioptera: Mesoraphidiidae): A new spe-
viewerforcommentsonanearlierdraftofthe
ciesinCretaceousamberfromMyanmar,with
manuscript. This paper is dedicated to Prof.
a catalog of fossil snakeflies. American
Dr.HorstAspo¨ckandDr.UlrikeAspo¨ck,the Museum Novitates 3363:1–22.
world’s authorities on the systematics of Engel, M.S. 2003. A new Eocene-Oligocene snake-
Raphidioptera (indeed, of the Neuropterida fly from Florissant, Colorado (Raphidioptera:
as a whole). Partial support for this work was Raphidiidae). Transactions of the Kansas
provided by the Alexander von Humboldt Academyof Science 106(3–4):124–128.
Foundation (to VP), and by NSF DEB- Engel, M.S., J.-D. Lim, and K.-S. Baek. 2006.
0542909 and a Guggenheim Fellowship from Fossil snakeflies from the Early Cretaceous of
southern Korea (Raphidioptera: Mesoraphi-
the John Simon Guggenheim Memorial
diidae). Neues Jahrbuch fu¨r Geologie und
Foundation (both to MSE). This is contribu-
Pala¨ontologie Monatshefte2006(4): 249–256.
tion No. 3476oftheDivisionofEntomology,
Grimaldi, D. 2000. A diverse fauna of
University of Kansas Natural History
Neuropterodea in amber from the Cretaceous
Museum.
ofNewJersey.In:D.Grimaldi(editor),Studies
onfossilsinamber,withparticularreferenceto
REFERENCES theCretaceousofNewJersey:259–303.Leiden:
Backhuys,viii+498pp.
Aspo¨ck,H.1998.Distributionandbiogeographyof Grimaldi, D., and M.S. Engel. 2005. Evolution of
the order Raphidioptera: updated facts and the insects. Cambridge: Cambridge University
anewhypothesis.ActaZoologicaFennica209: Press, xv+755pp.
33–44. Grimaldi,D.A.,M.S.Engel,andP.C.Nascimbene.
Aspo¨ck,H.,U.Aspo¨ck,andH.Rausch.1991.Die 2002. Fossiliferous Cretaceous amber from
Raphidiopteren der Erde. Eine monogra- Myanmar (Burma): its rediscovery, biotic di-
phische Darstellung der Systematik, Taxon- versity, and paleontological significance.
omie, Biologie, O¨kologie und Chorologie der American Museum Novitates 3361: 1–72.
rezenten Raphidiopteren der Erde, mit einer Hong, Y.-C. 1982. Mesozoic fossil insects of
zusammenfassenden U¨bersicht der fossilen Jiuquan Basin in Gansu Province. Beijing: