The Neuropterid Fauna of Dominican and Mexican Amber

60 Pages·2007·15.92 MB·English

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3587, 58 pp., 51 figures, 9 tables September 6, 2007
The Neuropterid Fauna of Dominican and Mexican
Amber (Neuropterida: Megaloptera, Neuroptera)
MICHAEL S. ENGEL1 AND DAVID A. GRIMALDI2
CONTENTS
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Resumen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Materials and Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Systematic Paleontology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Order Megaloptera Latreille. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Family Sialidae Leach. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Order Neuroptera Linnaeus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Family Mantispidae Leach. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Family Coniopterygidae Burmeister . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Family Hemerobiidae Latreille. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Family Chrysopidae Schneider. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Family Myrmeleontidae Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Family Ascalaphidae Rambur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Discussion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Paleoecology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Age of Fauna and Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
1DivisionofInvertebrateZoology,AmericanMuseumofNaturalHistory;DivisionofEntomology(Paleoentomology),
Natural History Museum, and Department of Ecology and Evolutionary Biology, 1501 Crestline Drive–Suite 140,
UniversityofKansas,Lawrence,KS66049-2811([email protected]).
2DivisionofInvertebrateZoology,AmericanMuseumofNaturalHistory([email protected]).
CopyrightEAmericanMuseumofNaturalHistory2007 ISSN0003-0082
2 AMERICAN MUSEUMNOVITATES NO. 3587
ABSTRACT
TheneuropteridfaunaofearlyMioceneDominicanandOligocene-MioceneMexicanamberis
treated. The fauna consists entirely of Megaloptera and Neuroptera while the snakeflies
(Raphidioptera)arenotpresentlyknowninTertiaryambersfromtheNewWorld.Fifteenspecies
are recognized,all inliving generaandsubgenera:Sialidae, Sialis (Protosialis)casca,new species
(Dominican);Ascalaphidae,Ululodespaleonesia,newspecies(Dominican),Amoeaelectrodominica,
new species (Dominican); Chrysopidae, Chrysopa glaesaria, new species (Dominican), C. vetula,
newspecies(Dominican),Leucochrysa(Nodita)prisca,newspecies(Dominican);Coniopterygidae,
Coniopteryx antiquua, new species (Dominican), Spiloconis glaesaria Meinander (Dominican), S.
oediloma, new species (Dominican); Hemerobiidae, Notiobiella thaumasta Oswald (Dominican),
Sympherobius sp. (Dominican); Mantispidae, Dicromantispa electromexicana, new species
(Mexican), D. moronei, new species (Dominican); Feroseta prisca Poinar, nomen emendatum
(Dominican); Myrmeleontidae, Porrerus dominicanus Poinar and Stange (Dominican). Immature
stages of Ascalaphidae (two species of Ululodes; Dominican), Chrysopidae (genus indet.;
Dominican), and Myrmeleontidae (P. dominicanus?; Dominican) are reported. The neuropterid
fossilrecordissummarizedandtheDominicanandMexicanamberfaunasarecomparedtoother
neuropterid amber faunas. The biogeographic and paleoecological implications of the Miocene
fossilsarediscussed.AbundanceanddiversityofNeuropterainambersappeartoberelatedtothe
abundance ofSternorrhyncha, onwhich manyneuropterans feed.
RESUMEN
Sedescribelafaunadeneuropte´ridosencontradaena´mbardominicanodelMiocenobasalyen
a´mbarmexicanodelOligoceno-Mioceno.Lafaunaesta´ compuestaenteramenteporMegaloptera
yNeuroptera,mientrasquehastalafechanoseconocenregistrosdeRaphidiopteraenyacimientos
de a´mbar del Terciario en el Nuevo Mundo. Se reconocen 15 especies, todas pertenecientes
a ge´neros y subge´neros vivos en la actualidad: Sialidae, Sialis (Protosialis) casca, especie nueva
(Dominicana); Ascalaphidae, Ululodes paleonesia, especie nueva (Dominicana), Amoea electro-
dominica, especie nueva (Dominicana); Chrysopidae, Chrysopa glaesaria, especie nueva
(Dominicana),C.vetula,especienueva(Dominicana),Leucochrysa(Nodita)prisca,especienueva
(Dominicana); Coniopterygidae, Coniopteryx antiquua, especie nueva (Dominicana), Spiloconis
glaesaria Meinander (Dominicana), S. oediloma, especie nueva (Dominicana); Hemerobiidae,
Notiobiella thaumasta Oswald (Dominicana), Sympherobius sp. (Dominicana); Mantispidae,
Dicromantispaelectromexicana,especienueva(Me´xico),D.moronei,especienueva(Dominicana);
FerosetapriscaPoinar,nom.emend.(Dominicana);Myrmeleontidae,PorrerusdominicanusPoinar
yStange(Dominicana).Secitanestad´ıosinmadurosdeAscalaphidae(Ululodesspp.;Dominicana),
Chrysopidae (ge´nero no determinado; Dominicana) y Myrmeleontidae (P. dominicanus?;
Dominicana). Se comparan estas faunas extintas con otras de neuropte´ridos en a´mbar y se
discutenlasimplicacionesbiogeogra´ficasypaleontolo´gicasdelostaxonestratados.Laabundancia
ydiversidaddeNeuropteraenyacimientosdea´mbarpareceestarrelacionadaconlaabundancia
deStenorrhyncha, delos cualesse alimentan muchos neuro´pteros.
INTRODUCTION paired diverticulum of the proventriculus,
and the caudally bifid mediolongitudinal
The superorder Neuropterida comprises suture on the first abdominal tergum
three distinctive orders of holometabolous (Mickoleit, 1973; Achtelig, 1975; Kristensen,
insects: the Megaloptera, the Neuroptera, 1991). Although such characters are obscure,
and the Raphidioptera. These three orders eachofthemajormonophyleticgroupswithin
form a monophylum defined by the fusion of the Neuropterida has a distinctive wing
the third valvulae in the ovipositor, the venation. Venation has allowed interpretation
medially divided metapostnotum, the un- of fossils, and in fact the abundant record
2007 ENGEL AND GRIMALDI: TERTIARY AMBER NEUROPTERIDA 3
of wing impressions in rocks gives the and is the most intensively studied of amber
Neuropterida among the most extensive fossil Lagersta¨tte;however,veryfewmodernstudies
recordofallHolometabola.Togetherwiththe have investigated inclusions of Neuropterida,
Coleoptera, their closest living relatives and comprehensive revisionary work is need-
among the Holometabola (Kristensen, 1991, ed.Evenbasicdescriptiveworkofnewspecies
1995; Ho¨rnschemeyer, 1998; Whiting et al., in Baltic amber is ongoing (e.g., Wichard,
1997; Carpenter and Wheeler, 1999; Wheeler 1997; Engel, 1999a; Dobosz and Krsemin´ski,
et al., 2001; Grimaldi and Engel, 2005), the 2000; Wichard and Engel, 2006). Among
neuropterids are an ancient offshoot of Cretaceous ambers, the fauna of the
endopterygoteinsectswithgeologicalhistories Turonian New Jersey amber is only slightly
extending back to the Early Permian lessdiversethanthatofBalticamber(table 2),
(Kukalova´-Peck, 1991; Carpenter, 1992; particularly when accounting for the much
GrimaldiandEngel,2005).Unlikethebeetles, smaller size of the deposits of the former. The
however, the neuropterid orders are not very onlyotherCretaceousamberwithasignificant
diverse and some, like the families of neuropterid fauna is the Late Albian amber
Raphidioptera,areperhapsindeclineglobally from Myanmar (Grimaldi et al., 2002; Engel,
(Aspo¨ck, 1998; Engel, 2002a). In fact, a re- 2002a, 2004a, 2004b, in prep.). Other amber
markable diversity of extinct neuropterid deposits have revealed a few other
lineages is known from the Mesozoic and, to Neuropterida (table 3), principally of families
a lesser degree, earlier deposits (e.g., Schlu¨ter, welldocumentedinthemajordeposits.Lastly,
1986; Carpenter, 1992; Grimaldi and Engel, the Cenozoic amber deposits of North
2005). Owing to the combination of their America have similarly revealed only sparse
ancient age, highly apomorphic relict taxa, material of a few neuropterids. Previously
and numerous extinct lineages, the only three species in the families Myrmeleon-
Neuropterida is one of those groups whereby tidae, Coniopterygidae, and Hemerobiidae
inclusion of fossil taxa into cladistic studies is have been described from Dominican amber
critical (Gauthier et al., 1989). It will be while none has been described from Mexican
interesting to explore the effects of such fossil amber. Herein we report on new neuropterids
taxa on our understanding of neuropterid discoveredinDominicanandMexicanamber;
phylogeny at all hierarchical levels. Most these include the first New World fossil of an
neuropterid fossils, however, are compression alderfly(Sialidae)andthefirstgreenlacewings
fossilsthatpreserveonlywingsorverylimited, (Chrysopidae) and the first adult owlfly
structural details. Preservation in amber, (Ascalaphidae) in amber. In total the fauna
however, is unsurpassed, preserving not only has now increased to 13 species distributed
the external morphology in essentially lifelike among seven families (table 4). The age of
detail but often internal tissues (Grimaldi et Dominican amber has been unfortunately
al., 1994; Grimaldi and Engel, 2005). Amber confused, but it is certainly no older than
inclusionsofferagreateropportunitytomake EarlyMiocene inorigin(Iturralde-Vinentand
meaningful comparisons between Recent and MacPhee, 1996, 1999). Mexican amber is of
fossil taxa as well as simultaneous cladistic a similar age, originating from the Late
analyses of paleontological and neontological Oligocene–EarlyMiocene(Langenheim,1966).
data (e.g., Grimaldi and Cumming, 1999; With such a long geological duration of
Engel, 2001). neuropteridan history (Permian to present),
The most diverse neuropterid fauna in Tertiary fossils are of little phylogenetic
amber is from the Baltic region, which has consequence regarding higher taxa. The sig-
classically receivedthegreatest attention from nificance of Dominican and Mexican amber
systematists. There are presently 33 described fossils lies in (1) the preservation of amber,
speciesinBalticamber(table 1).Thisamberis allowing close species comparisons; (2) in-
middle Eocene in age (dating reviewed in formation for West Indian biogeography,
Engel, 2001; Weitschat and Wichard, 2002). particularlyassessinganygeographicalextinc-
Balticamberisthelargestdepositofamberin tions; and (3) paleoecology, or taphonomic
the world. It has been studied for centuries inferences provided by the taxa that are
4 AMERICAN MUSEUMNOVITATES NO. 3587
TABLE1
Named Neuropterida inBaltic Ambera
Taxa Reference
ORDERNEUROPTERALINNAEUS
(5Planipennia)
—FamilyASCALAPHIDAERambur
NeadelphusprotaeMacLeod MacLeod,1970
—FamilyBEROTHIDAEHandlirsch
ProberothapriscaKru¨ger Kru¨ger,1923
—FamilyCONIOPTERYGIDAEBurmeister
ArchiconiocompsapriscaEnderlein Enderlein,1910
ArchiconiselectricaEnderlein Enderlein,1930
Coniopteryxtimidus(Hagen) Pictet-Baraban&Hagen,1856;Meinander,1972,1975
HeminiphetiafritschiEnderlein Enderlein,1930
HemisemidaliskulickaeDobosz&Krzemin´ski Dobosz&Krzemin´ski,2000
HemisemidalissharoviMeinander Meinander,1975
—FamilyDILARIDAENewman
CascadilareocenicusEngel Engel,1999a
—FamilyHEMEROBIIDAELatreille
HemerobitesantiquusGermar Germar,1813
Prochlaniusresinatus(Hagen) Pictet-Baraban&Hagen,1856;Kru¨ger,1923
ProphlebonemaresinataKru¨ger Kru¨ger,1923
Prospadobiusmoestus(Hagen) Pictet-Baraban&Hagen,1856;Kru¨ger,1923
—FamilyNEVRORTHIDAENakahara
Rophalisrelicta(Hagen) Pictet-Baraban&Hagen,1856;Nel&Jarzembowski,1997
—FamilyNYMPHIDAERambur
Pronymphesmengeanus(Hagen) Pictet-Baraban&Hagen,1856;Kru¨ger,1923;MacLeod,1970[larva]
—FamilyOSMYLIDAELeach
Protosmyluspictus(Hagen) Pictet-Baraban&Hagen,1856;Kru¨ger,1923
—FamilyPSYCHOPSIDAEHandlirsch
PropsychopsishageniMacLeod MacLeod,1970
PropsychopsishelmiKru¨ger Kru¨ger,1923
PropsychopsislapicidaMacLeod MacLeod,1970
—FamilyRHACHIBEROTHIDAETjeder
Whalferavenatrix(Whalley)b Whalley,1983;Engel,2004b
—FamilySISYRIDAEHandlirsch
Sisyra(?)amissaHagen Pictet-Baraban&Hagen,1856
ORDERMEGALOPTERALATREILLE
—FamilyCORYDALIDAELeach
ChauliodespriscaPictet Pictet,1854
ChauliodescarsteniWichard Wichard,2003
—FamilyCORYDASIALIDAEWichardetal.
CorydasialisinexpectatusWichardetal. Wichardetal.,2005
—FamilySIALIDAELeach
Sialis(Protosialis)balticaWichard Wichard,1997
Sialis(Protosialis)herrlingiWichard Wichard,2002
Sialis(Protosialis)voigtiWichard&Engel Wichard&Engel,2006
SialisgroehniWichard Wichard,1997
ORDERRAPHIDIOPTERANAVA´Sc
—FamilyINOCELLIIDAENava´s
Electrinocelliapeculiaris(Carpenter) Carpenter,1956;Engel,1995
FiblacarpenteriEngel Engel,1995
Fiblaerigena(Hagen) Pictet-Baraban&Hagen,1856;Carpenter,1956
SuccinofiblaapertaAspo¨ck&Aspo¨ck Aspo¨ck&Aspo¨ck,2004
2007 ENGEL AND GRIMALDI: TERTIARY AMBER NEUROPTERIDA 5
TABLE1
(Continued)
Taxa Reference
—FamilyRAPHIDIIDAELatreille
RaphidiabalticaCarpenter Carpenter,1956
SuccinoraphidiaexhibiensAspo¨ck&Aspo¨ck Aspo¨ck&Aspo¨ck,2004
aUndescribedimmaturesarerecordedforSialidae(Weidner,1958),Raphidiidae(Berendt,1856),Inocelliidae(Weidner,
1958),Hemerobiidae(Pictet-BarabanandHagen,1856),andNevrorthidae(WeitschatandWichard,1998,2002).
bThisspecimenwasdescribedfrom‘‘British’’amber(andisthereforealsolistedintable3),butthisisthesameasBaltic
amberandismerelymaterialfromtheBalticthathaswashedupontheBritishIsles.
cBecauseNava´s(1916)wasthefirsttousetheordinalnameinthisform,wehaveattributedauthorshiptohimeven
though several earlier authors had also considered the snakeflies as a distinct order (e.g., Burmeister, 1839: as
Raphidiodea).Nava´s(1916)appearstohavefollowedShipley’s(1904)recommendationforacommon-pteraendingfor
insect ordinal names by emending the name to Raphidioptera. Although the name Raphidioptera is somewhat
meaninglessinitsoriginalGreek(raphidos,meaning‘‘needle’’,areferencetotheelongateovipositor;andpteron,meaning
‘‘wing’’),itisinalmostuniversalusageforthisgroup,andthereseemstobelittlereasontochangeitbacktoRaphidiodea,
Raphidiida,ortosomethingmoredescriptiveinGreeksuchasOphidiodera(ophidion,or‘‘snake’’;dere,or‘‘neck’’).
preserved. The Dominican and Mexican am- al., 1991; Nel, 1993; Engel, 2002a, 2003) and
ber faunas are relatively similar to tropical snakeflies are present in mid-Eocene ambers
neuropteridfaunasintheseregionstoday.The of Europe (Carpenter, 1956; Engel, 1995,
fauna consists entirely of Megaloptera and 2002a; Weitschat and Wichard, 1998, 2002;
Neuroptera while the snakeflies (Raphi- Aspo¨ck and Aspo¨ck, 2004). However, on the
dioptera) are presently unknown in Tertiary whole the Cenozoic record of this order is
ambers of the New World. Tertiary compres- insignificant and it is in the Mesozoic that
sion fossils of Raphidioptera are known from a remarkable diversity of Raphidioptera from
North America and Europe (e.g., Aspo¨ck et throughout the world is known (e.g., Aspo¨ck,
1998; Grimaldi, 2000; Engel, 2002a, unpubl.
data; Engel et al., 2006; Perrichot and Engel,
TABLE2 in press).
NamedNeuropterida in NewJerseyAmber
(from Grimaldi, 2000; Engel,2002b:all are placed
in extinct genera) MATERIALS AND METHODS
All measurements were made using an
ORDERNEUROPTERALINNAEUS
ocular micrometer and should be considered
–FamilyBEROTHIDAEHandlirsch
approximate since the optimal angle for any
JersiberothaluzziiGrimaldi
JersiberothasimilisGrimaldi given metric was not always achievable owing
NascimberothapictaGrimaldi to the uneven surface of the amber. The
–FamilyCONIOPTERYGIDAEBurmeister acronyms AMNH and MACT are used,
ApoglaesoconisackermaniGrimaldi respectively, for the American Museum of
ApoglaesoconischerylaeEngel Natural History, New York, and the Morone
ApoglaesoconisluzziiGrimaldi Amber Collection, Turin, Italy.
ApoglaesoconisswolenskyiGrimaldi
GlaesoconisnearcticaGrimaldi
–FamilyMANTISPIDAELeach SYSTEMATIC PALEONTOLOGY
MantispidipteraenigmaticaGrimaldi
ORDERMEGALOPTERALATREILLE
MantispidipterahenryiGrimaldi
FAMILYSIALIDAELEACH
–FamilyRHACHIBEROTHIDAETjeder
RhachibermissasplendidaGrimaldi
The Sialidae, or alderflies, are one of two
ORDERRAPHIDIOPTERANAVA´S living families of Megaloptera. Sialid larvae,
–FamilyMESORAPHIDIIDAEMartynova like those of all megalopterans, are aquatic
MesoraphidialuzziiGrimaldi
predators, living in anything from small
aAnundescribedlarvaofthisgroupwasalsorecorded. streams, rivers, ponds, lakes, or even phyto-
6 AMERICAN MUSEUMNOVITATES NO. 3587
TABLE3
Named Neuropterida inLebanese, Burmese, French, Siberian, HatCreek, ‘‘British’’,aandParisian Ambersb
(allare inextinct genera)
Taxa Deposit Reference
ORDERNEUROPTERALINNAEUS
—FamilyBEROTHIDAEHandlirsch
BanoberothaenigmaticaWhalley Lebanese(Neocomian) Whalley,1980
MicroberothamacculloughiArchibaldandMakarkin HatCreek(Eocene) Archibald&Makarkin,2004
Plesiorobius?anadensisKlimaszewskiandKevan Canadian(Campanian) Klimaszewski&Kevan,1986
—FamilyCONIOPTERYGIDAEBurmeister
GlaesoconiscreticaMeinander Siberian(Santonian) Meinander,1975
GlaesoconisbaliopteryxEngel Burmese(Albian) Engel,2004a
Libanoconisfadiacra(Whalley) Lebanese(Neocomian) Whalley,1980;Engel,2002a
PhthanoconisburmiticaEngel Burmese(Albian) Engel,2004a
LibanosemidalishammanaensisAzaretal. Lebanese(Neocomian) Azaretal.,2000
GallosemidaliseocenicaNeletal. Parisian(Eocene) Neletal.,2005a
AlboconiscretacicaNeletal. Lebanese(Neocomian) Neletal.,2005a
—FamilyRHACHIBEROTHIDAETjeder
ParaberothaacraWhalley Lebanese(Neocomian) Whalley,1980
RetinoberothastuermeriSchlu¨ter French(Cenomanian) Schlu¨ter,1978
EorhachiberothaburmiticaEngel Burmese(Albian) Engel,2004b
Whalferavenatrix(Whalley) ‘‘British’’(Eocene) Whalley,1983;Engel,2004b
AlloberothapetruleviciiNeletal. French(Cenomanian) Neletal.,2005b
ChimerhachiberothaacrasariiNeletal. Lebanese(Neocomian) Neletal.,2005b
SpinoberothamickaelacraiNeletal. Lebanese(Neocomian) Neletal.,2005b
Oiseacelinea(Neletal.) Parisian(Eocene) Neletal.,2005b
ORDERMEGALOPTERALATREILLE
—FamilySIALIDAELeach
EosialisdorisiNeletal. Parisian(Eocene) Neletal.,2002
ORDERRAPHIDIOPTERANAVA´S
—FamilyMESORAPHIDIIDAEMartynovc
NanoraphidiaelectroburmicaEngel Burmese(Albian) Engel,2002a
a‘‘British’’amberisthesameasBalticamberandismaterialfromtheBalticthathaswashedupontheBritishIsles.
ThesedepositsaremiddleEoceneinage(seediscussionofdatinginEngel,2001;GrimaldiandEngel,2005).
bAnundescribedspecimenfromCanadianamberhasbeenreportedasapossibleberothid(McAlpineandMartin,1969;
Pike,1995).
cAn undescribed larva of this group was recorded by Engel (2002a), and a further account of various Cretaceous
snakeflyimmaturesissummarizedbyPerrichotandEngel(inpress).
telmata and under stones or where dead plant [1994] and New and Theischinger [1993],
material accumulates. Larvae emerge onto respectively), South Africa (Leptosialis),
land to pupate in vegetation or debris. Australia (Austrosialis and Stenosialis), and
Adultsareshort-lived,occuralongwatersides, Southeast Asia (Indosialis). In addition, 10
andapparentlymayfeedonpollenandnectar, extinct species have been previously proposed
if at all. Generally little is known of adult (table 5), almost all in Tertiary deposits. The
megalopteran biology, particularly so for primitive Dobbertinia reticulata (Handlirsch)
Sialidae. from the Liassic of Germany is the oldest
Today the family is represented by approx- record of the family. This species is clas-
imately66livingspecies(mostinthenominate sified in its own basal subfamily, the
genus Sialis) that are distributed worldwide Dobbertiniinae.
butareprincipallyHolarcticwithafewspecies Herein we describe a new species of Sialis
in the New World tropics (Sialis s.l.: includes (Protosialis) in amber from the Dominican
Protosialis and Nipponosialis after Whiting Republic, a group which is today unknown
2007 ENGEL AND GRIMALDI: TERTIARY AMBER NEUROPTERIDA 7
TABLE4
Neuropterida in DominicanandMexicanaAmber
Taxa Reference
ORDERMEGALOPTERALATREILLE
—FamilySIALIDAELeach
Sialis(Protosialis)cascaEngel&Grimaldin.sp. Presentstudy
ORDERNEUROPTERALINNAEUS
—FamilyASCALAPHIDAERambur
AmoeaelectrodominicanaEngel&Grimaldin.sp. Presentstudy
UlulodespaleonesiaEngel&Grimaldin.sp.b Presentstudy
—FamilyCHRYSOPIDAESchneider
ChrysopaglaesariaEngel&Grimaldin.sp. Presentstudy
ChrysopavetulaEngel&Grimaldin.sp. Presentstudy
Leucochrysa(Nodita)priscaEngel&Grimaldin.sp. Presentstudy
—FamilyCONIOPTERYGIDAEBurmeisterb
ConiopteryxantiquuaEngel&Grimaldin.sp. Presentstudy
SpiloconisglaesariaMeinander Meinander,1998a
SpiloconisoedilomaEngel&Grimaldin.sp. Presentstudy
—FamilyHEMEROBIIDAELatreille
NotiobiellathaumastaOswald Oswald,1999
Sympherobiussp. Presentstudy
—FamilyMANTISPIDAELeach
DicromantispamoroneiEngel&Grimaldin.sp. Presentstudy
DicromantispaelectromexicanaEngel&Grimaldin.sp. Presentstudy
FerosetapriscaPoinar Poinar,2006
—FamilyMYRMELEONTIDAELatreille
PorrerusdominicanusPoinar&Stangeb Poinar&Stange,1996
aDicromantispaelectromexicanaispresentlytheonlydescribedneuropteridfromMexicanamber.
bImmatures(allreporteduponherein)arerecordedforAscalaphidae,Chrysopidae,andMyrmeleontidae.
fromtheWestIndiesexceptforonespecieson ings; antennae black; head with black mark-
the island of Cuba, that is, Sialis (Protosialis) ings along margins of eye and extending
bifasciata Hagen (Penny, 1977; Contreras- posteriorly near to posterior border of head,
Ramos, 1999). black extended medially but not reaching
midline, borders of markings rounded (not
flamelike). Apical segments of maxillary
Sialis (Protosialis) casca, new species
palpi with medioapical tooth. Ocelli absent.
figures 1–3 Pronotum apparently twice as wide as long,
remainder of thorax dark brown. Protibia,
DIAGNOSIS: The orange coloration of the mesofemur, mesotibia, and metatibia black,
head and pronotum and narrowed costal area remaining leg segments whitish; fourth tar-
with a reduced number of costal crossveins somere bilobed. Wing membranes fuscous;
indicate this to be a species of the subgenus veins dark brown to black; forewing with
Protosialis. For Protosialis species the pres- costal area distinctly narrowed just before
ence of only three costal crossveins is unique middle of wing; three costal crossveins,
while the black markings on the head are distal two veins distinctly oblique (i.e., not
similar to that seen in Sialis (Protosialis) perpendicular to C or Sc); distal r-rs cross-
flammata(Penny)butareroundedinthefossil vein meeting R slightly distad to R /
2+3 2+3
(not flamelike as in S. flammata). R fork; posterior branch of R /R sepa-
4+5 4 5
DESCRIPTION: Forewing length (preserved) rating near wing apex (i.e., distad R2/R3
7.7 mm, (estimated: from bent forewing) branching); outer series of gradate cross-
8.4 mm; total body length 7.6 mm (fig. 1). veins confluent (figs. 2, 3). Abdomen
Head and pronotum orange with black mark- slightly distended and bent ventrally near
8 AMERICAN MUSEUMNOVITATES NO. 3587
TABLE5
Named Fossil Sialidae
Taxa Deposit Reference
Dobbertiniareticulata(Handlirsch) Jurassic,Germany Ansorge,2001
EosialisdorisiNeletal. Parisianamber Neletal.,2002
IndosialisbeskonakensisNel Miocene,Turkey Nel,1988a
ProindosialiscantalensisNel Miocene,France Nel,1988a
SialisstrausiIllies Pliocene,Germany Illies,1967
SialisgroehniWichard Balticamber Wichard,1997
SialismuratensisNel Miocene,France Nel,1988a
Sialis(Protosialis)balticaWichard Balticamber Wichard,1997
Sialis(Protosialis)herrlingiWichard Balticamber Wichard,2002
Sialis(Protosialis)cascan.sp. Dominicanamber Presentstudy
Sialis(Protosialis)voigtiWichard&Engel Balticamber Wichard&Engel,2006
midpoint; apparently light brown; apex of ORDERNEUROPTERALINNAEUS
abdomen curled under, with genitalia pointed FAMILYMANTISPIDAELEACH
anteriad.
Themantispidsareperhapsoneofthemost
HOLOTYPE: Male; MACT-2090 (fig. 1), instantly recognizable of neuropteran families
Miocene amber of the Dominican Republic.
and consist of approximately 400 species
ETYMOLOGY: The specific epithet is taken worldwide. Their convergent form with man-
from the Latin word cascus (meaning ‘‘old’’) tises(Mantodea)ofanelongateprothoraxand
and is a reference to the age of the specimen. raptorial forelegs is remarkable. Adults tend
Fig. 1. Photomicrograph ofholotype of Sialis (Protosialis)casca,new species(MACT-2090).
2007 ENGEL AND GRIMALDI: TERTIARY AMBER NEUROPTERIDA 9
Figs. 2, 3. Holotype of Sialis (Protosialis) casca, new species (MACT-2090). 2. Dorsal aspect. 3. Left
lateralaspect.
to be diurnal and are often found on flowers, ators although the larvae of at least one
although a few lineages are nocturnal or studiedspeciesareambulatory.Thesubfamily
crepuscular.Somespeciesarebrightlycolored Drepanicinaeisentirelyunknownbiologically.
and appear to mimic social, aculeate The largest subfamily, the Mantispinae and
Hymenoptera. The most basal mantispids, the subfamily to which the fossils described
the Symphrasinae, have sedentary larvae that below belong, are specialized, obligate para-
are generalist predators found in the nests of sitesofspiders,andthefirst-instarlarvaeither
social Hymenoptera, particularly vespids, or boards a female spider and enters the egg sac
burrows of scarab beetle larvae. Similarly, the as it is constructed, to feed on the eggs, or it
Calomantispinaeappeartobegeneralistpred- searches directly for an egg sac. Later larval
10 AMERICAN MUSEUMNOVITATES NO. 3587
TABLE6
Named Fossil Mantispidaea
Table Deposit Reference
Climaciella?henrotayiNel Oligocene,France Nel,1988b
Dicromantispaelectromexicanan.sp. Mexicanamber Presentstudy
Dicromantispamoronein.sp. Dominicanamber Presentstudy
FerosetapriscaPoinar Dominicanamber Poinar,2006
GerstaeckerellaasiaticaMakarkin LateCretaceous,Kazakhstan Makarkin,1990
LiassochyrsastigmaticaAnsorge EarlyJurassic,Germany Ansorge&Schlu¨ter,1990;
&Schu¨lter Wedmann&Makarkin,2007
MesomantispasibiricaMakarkin EarlyCretaceous,Siberia Makarkin,1996
PromantispasimilisPanfilov LateJurassic,Karatau Panfilov,1980
ProsagittalataoligocenicaNel Oligocene,France Nel,1988b
SymphrasiteseocenicusWedmann& Eocene,Germany Wedmann&Makarkin,2007
Makarkin
Vectisparelicta(Cockerell)b Eocene,England Cockerell,1921
aAlthoughWillmann(1994)transferredtheCretaceousamberRetinoberothaandParaberothatoMantispidae,theseare
rhachiberothids.Similarly,Whalferavenatrix(Whalley,1983)isexcludedowingtoitslikelypositioninRhachiberothidae
(e.g.,Engel,2004b;WedmannandMakarkin,2007).
bOriginallyproposedinMantispaandthentransferredtoPromantispaJarzembowski(1980:nomenpraeoccupatum,nec
Panfilov,1980)andfinallytoVectispaLambkin(1986:nomennovumproPromantispaJarzembowski,1980).
instars are relatively immobile. Pupation oc- mantispid is Feroseta priscus Poinar in
curswithinthehosteggsac.Adultsareactively Miocene Dominican amber (Poinar, 2006).
predaceousonavarietyofotherinsectgroups. Herein we report two further, definitive
Redborg (1998) has reviewed the available mantispids in Cenozoic amber from the
information on the remarkably specialized Western Hemisphere. Spiders are common in
biology of Mantispinae. Although Willmann Tertiary ambers of North America (e.g.,
(1990) included the Rhachiberothinae in Wunderlich, 1988; Penney, 1999), and it is
Mantispidae, we follow Aspo¨ck and Mansell therefore perhaps not surprising that their
(1994)andAspo¨cketal.(2001)byconsidering parasites would also eventually be discovered.
thisgroupasalliedtotheBerothidae.
The geological history of the Mantispidae Dicromantispa electromexicana, new species
has only recently begun to come together.
figures 4–8
Previously there were nine fossil species de-
scribed from deposits ranging from the Early
Mantispa sp.:Engel, 2004c:184.
Jurassic to the Miocene (table 6: Wedmann
Mantispa sp.: Grimaldi and Engel, 2005: 354,
and Makarkin, 2007), although the exact
fig. 9.35.
systematic position of some of these taxa is DIAGNOSIS: This species is superficially
not entirely certain. Only two mantispid similar to Zeugomantispa minuta (Fabricius)
species had been discovered in amber until but differs by the more elongate pronotum
this time, but one is likely not a member of that apparently lacks markings or bumps and
this family. Walfera venatrix (Whalley) was has exceedingly sparse, minute pubescence,
discovered in amber that had washed up on and by the structure of the ectoprocts. It can
the eastern shore of Britain and presumably be separated from D. moronei in Dominican
was derived from theBalticamberdepositsof amber (see below) by the more elongate and
the blaue Erde. The identity of Whalfera, unexpanded pronotum, absence of pronotal
however, has been questioned by many, and markings,majorprofemoralspinelongerthan
the genus is very likely a rhachiberothid protarsus, fewer c-sc crossveins, and smaller
(Aspo¨ck and Mansell, 1994; Engel, 2004b; body size. The species can be separated from
Grimaldi and Engel, 2005; Wedmann and all modern Dicromantispa by the absence of
Makarkin, 2007). The only other amber the ventromedian lobe of the ectoproct. Such