A tamerican museum Novitates 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 division of Invertebrate Zoology, American Museum of Natural History; Division of Entomology (Paleoentomology), Natural History Museum, and Department of Ecology and Evolutionary Biology, 1501 Crestline Drive-Suite 140, University of Kansas, Lawrence, KS 66049-2811 ([email protected]). 2Division of Invertebrate Zoology, American Museum of Natural History ([email protected]). Copyright © American Museum of Natural History 2007 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3587 ABSTRACT The neuropterid fauna of early Miocene Dominican and Oligocene-Miocene Mexican amber is treated. The fauna consists entirely of Megaloptera and Neuroptera while the snakeflies (Raphidioptera) are not presently known in Tertiary ambers from the New World. Fifteen species are recognized, all in living genera and subgenera: Sialidae, Sialis (Protosialis) casca, new species (Dominican); Ascalaphidae, Ululodes paleonesia, new species (Dominican), Amoea electrodominica, new species (Dominican); Chrysopidae, Chrysopa glaesaria, new species (Dominican), C. vetula, new species (Dominican), Leucochrysa {Nodita) prisca, new species (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 fossil record is summarized and the Dominican and Mexican amber faunas are compared to other neuropterid amber faunas. The biogeographic and paleoecological implications of the Miocene fossils are discussed. Abundance and diversity of Neuroptera in ambers appear to be related to the abundance of Sternorrhyncha, on which many neuropterans feed. RESUMEN Se describe la fauna de neuropteridos encontrada en ambar dominicano del Mioceno basal y en ambar mexicano del Oligoceno-Mioceno. La fauna esta compuesta enteramente por Megaloptera y Neuroptera, mientras que hasta la fecha no se conocen registros de Raphidioptera en yacimientos de ambar del Terciario en el Nuevo Mundo. Se reconocen 15 especies, todas pertenecientes a generos y subgeneros 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, especie nueva (Dominicana), Leucochrysa (Nodita) prisca, especie nueva (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, Dicromantispa electromexicana, especie nueva (Mexico), D. moronei, especie nueva (Dominicana); Feroseta prisca Poinar, nom. emend. (Dominicana); Myrmeleontidae, Porrerus dominicanus Poinar y Stange (Dominicana). Se citan estadios inmaduros de Ascalaphidae {Ululodes spp.; Dominicana), Chrysopidae (genero no determinado; Dominicana) y Myrmeleontidae (P. dominicanus?; Dominicana). Se comparan estas faunas extintas con otras de neuropteridos en ambar y se discuten las implicaciones biogeograficas y paleontologicas de los taxones tratados. La abundancia y diversidad de Neuroptera en yacimientos de ambar parece estar relacionada con la abundancia de Stenorrhyncha, de los cuales se alimentan muchos neuropteros. 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 each of the major monophyletic groups within 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 Lagerstatte; however, very few modern studies record of all Holometabola. Together with the have investigated inclusions of Neuropterida, Coleoptera, their closest living relatives and comprehensive revisionary work is need¬ among the Holometabola (Kristensen, 1991, ed. Even basic descriptive work of new species 1995; Hornschemeyer, 1998; Whiting et al., in Baltic amber is ongoing (e.g., Wichard, 1997; Carpenter and Wheeler, 1999; Wheeler 1997; Engel, 1999a; Dobosz and Krseminski, 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 endopterygote insects with geological histories Turonian New Jersey amber is only slightly extending back to the Early Permian less diverse than that of Baltic amber (table 2), (Kukalova-Peck, 1991; Carpenter, 1992; particularly when accounting for the much Grimaldi and Engel, 2005). Unlike the beetles, smaller size of the deposits of the former. The however, the neuropterid orders are not very only other Cretaceous amber with a significant diverse and some, like the families of neuropterid fauna is the Late Albian amber Raphidioptera, are perhaps in decline globally from Myanmar (Grimaldi et al., 2002; Engel, (Aspock, 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., Schliiter, well documented in the major deposits. 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 discovered in Dominican and Mexican amber; phylogeny at all hierarchical levels. Most these include the first New World fossil of an neuropterid fossils, however, are compression alderfly (Sialidae) and the first green lacewings fossils that preserve only wings or very limited, (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 inclusions offer a greater opportunity to make Early Miocene in origin (Iturralde-Vinent and 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-Early Miocene (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 received the greatest attention from nificance of Dominican and Mexican amber systematists. There are presently 33 described fossils lies in (1) the preservation of amber, species in Baltic amber (table 1). This amber is allowing close species comparisons; (2) in¬ middle Eocene in age (dating reviewed in formation for West Indian biogeography, Engel, 2001; Weitschat and Wichard, 2002). particularly assessing any geographical extinc¬ Baltic amber is the largest deposit of amber in tions; and (3) paleoecology, or taphonomic the world. It has been studied for centuries inferences provided by the taxa that are 4 AMERICAN MUSEUM NOVITATES NO. 3587 TABLE 1 Named Neuropterida in Baltic Ambera Taxa Reference ORDER NEUROPTERA LINNAEUS (= Planipennia) —Family Ascalaphidae Rambur Neadelphus protae MacLeod MacLeod, 1970 —Family Berothidae Handlirsch Proberotha prisca Kruger Kruger, 1923 —Family Coniopterygidae Burmeister Archiconiocompsa prisca Enderlein Enderlein, 1910 Archiconis electrica Enderlein Enderlein, 1930 Coniopteryx timidus (Hagen) Pictet-Baraban & Hagen, 1856; Meinander, 1972, 1975 Heminiphetia fritschi Enderlein Enderlein, 1930 Hemisemidalis kulickae Dobosz & Krzeminski Dobosz & Krzeminski, 2000 Hemisemidalis sharovi Meinander Meinander, 1975 —Family Dilaridae Newman Cascadilar eocenicus Engel Engel, 1999a —Family Hemerobiidae Latreille Hemerobites antiquus Germar Germar, 1813 Prochlanius resinatus (Hagen) Pictet-Baraban & Hagen, 1856; Kruger, 1923 Prophlebonema resinata Kruger Kruger, 1923 Prospadobius moestus (Hagen) Pictet-Baraban & Hagen, 1856; Kruger, 1923 —Family Nevrorthidae Nakahara Rophalis relicta (Hagen) Pictet-Baraban & Hagen, 1856; Nel & Jarzembowski, 1997 —Family Nymphidae Rambur Pronymphes mengeanus (Hagen) Pictet-Baraban & Hagen, 1856; Kruger, 1923; MacLeod, 1970 [larva] —Family Osmylidae Leach Protosmylus pictus (Hagen) Pictet-Baraban & Hagen, 1856; Kruger, 1923 —Family Psychopsidae Handlirsch Propsychopsis hageni MacLeod MacLeod, 1970 Propsychopsis helmi Kruger Kruger, 1923 Propsychopsis lapicida MacLeod MacLeod, 1970 —Family Rhachiberothidae Tjeder Whalfera venatrix (Whalley)b Whalley, 1983; Engel, 2004b —Family Sisyridae Handlirsch Sisyra (?) amissa Hagen Pictet-Baraban & Hagen, 1856 ORDER MEGALOPTERA LATREILLE —Family Corydalidae Leach Chauliodes prisca Pictet Pictet, 1854 Chauliodes carsteni Wichard Wichard, 2003 —Family Corydasialidae Wichard et al. Corydasialis inexpectatus Wichard et al. Wichard et al., 2005 —Family Sialidae Leach Sialis (Protosialis) baltica Wichard Wichard, 1997 Sialis (Protosialis) herrlingi Wichard Wichard, 2002 Sialis (Protosialis) voigti Wichard & Engel Wichard & Engel, 2006 Sialis groehni Wichard Wichard, 1997 ORDER RAPHIDIOPTERA NAVASC —Family Inocelliidae Navas Electrinocellia peculiaris (Carpenter) Carpenter, 1956; Engel, 1995 Fibla carpenteri Engel Engel, 1995 Fibla erigena (Hagen) Pictet-Baraban & Hagen, 1856; Carpenter, 1956 Succinofibla aperta Aspock & Aspock Aspock & Aspock, 2004 2007 ENGEL AND GRIMALDI: TERTIARY AMBER NEUROPTERIDA 5 TABLE 1 ( Continued) Taxa Reference —Family Raphidiidae Latreille Raphidia baltica Carpenter Carpenter, 1956 Succinoraphidia exhibiens Aspock & Aspock Aspock & Aspock, 2004 aUndescribed immatures are recorded for Sialidae (Weidner, 1958), Raphidiidae (Berendt, 1856), Inocelliidae (Weidner, 1958), Hemerobiidae (Pictet-Baraban and Hagen, 1856), and Nevrorthidae (Weitschat and Wichard, 1998, 2002). bThis specimen was described from “British” amber (and is therefore also listed in table 3), but this is the same as Baltic amber and is merely material from the Baltic that has washed up on the British Isles. cBeeause Navas (1916) was the first to use the ordinal name in this form, we have attributed authorship to him even though several earlier authors had also considered the snakeflies as a distinct order (e.g., Burmeister, 1839: as Raphidiodea). Navas (1916) appears to have followed Shipley’s (1904) recommendation for a common -ptera ending for insect ordinal names by emending the name to Raphidioptera. Although the name Raphidioptera is somewhat meaningless in its original Greek (raphidos, meaning “needle”, a reference to the elongate ovipositor; andpteron, meaning “wing”), it is in almost universal usage for this group, and there seems to be little reason to change it back to Raphidiodea, Raphidiida, or to something more descriptive in Greek such as Ophidiodera (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 neuropterid faunas in these regions today. The of Europe (Carpenter, 1956; Engel, 1995, fauna consists entirely of Megaloptera and 2002a; Weitschat and Wichard, 1998, 2002; Neuroptera while the snakeflies (Raphi¬ Aspock and Aspock, 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., Aspock et throughout the world is known (e.g., Aspock, 1998; Grimaldi, 2000; Engel, 2002a, unpubl. data; Engel et al., 2006; Perrichot and Engel, TABLE 2 in press). Named Neuropterida in New Jersey Amber (from Grimaldi, 2000; Engel, 2002b: all are placed in extinct genera) MATERIALS AND METHODS All measurements were made using an ORDER NEUROPTERA LINNAEUS ocular micrometer and should be considered Family Berothidae Handlirsch approximate since the optimal angle for any Jersiberotha luzzii Grimaldi Jersiberotha similis Grimaldi given metric was not always achievable owing Nascimberotha picta Grimaldi to the uneven surface of the amber. The -Family Coniopterygidae Burmeister acronyms AMNH and MACT are used, Apoglaesoconis ackermani Grimaldi respectively, for the American Museum of Apoglaesoconis cherylae Engel Natural History, New York, and the Morone Apoglaesoconis luzzii Grimaldi Amber Collection, Turin, Italy. Apoglaesoconis swolenskyi Grimaldi Glaesoconis nearctica Grimaldi -Family Mantispidae Leach SYSTEMATIC PALEONTOLOGY Mantispidiptera enigmatica Grimaldi ORDER MEGALOPTERA LATREILLE Mantispidiptera henryi Grimaldi FAMILY SIALIDAE LEACH -Family Rhachiberothidae Tjeder Rhachibermissa splendida Grimaldi The Sialidae, or alderflies, are one of two ORDER RAPHIDIOPTERA NAVAS living families of Megaloptera. Sialid larvae, -Family Mesoraphidiidae Martynov a like those of all megalopterans, are aquatic Mesoraphidia luzzii Grimaldi predators, living in anything from small aAn undescribed larva of this group was also recorded. streams, rivers, ponds, lakes, or even phyto- 6 AMERICAN MUSEUM NOVITATES NO. 3587 TABLE 3 Named Neuropterida in Lebanese, Burmese, French, Siberian, Hat Creek, “British”,3 and Parisian Ambersb (all are in extinct genera) Taxa Deposit Reference ORDER NEUROPTERA LINNAEUS —Family Berothidae Handlirsch Banoberotha enigmatica Whalley Lebanese (Neocomian) Whalley, 1980 Microberotha macculloughi Archibald and Makarkin Hat Creek (Eocene) Archibald & Makarkin, 2004 Plesiorobius ?anadensis Klimaszewski and Kevan Canadian (Campanian) Klimaszewski & Kevan, 1986 —Family Coniopterygidae Burmeister Glaesoconis cretica Meinander Siberian (Santonian) Meinander, 1975 Glaesoconis baliopteryx Engel Burmese (Albian) Engel, 2004a Libanoconis fadiacra (Whalley) Lebanese (Neocomian) Whalley, 1980; Engel, 2002a Phthanoconis burmitica Engel Burmese (Albian) Engel, 2004a Libanosemidalis hammanaensis Azar et al. Lebanese (Neocomian) Azar et al., 2000 Gallosemidalis eocenica Nel et al. Parisian (Eocene) Nel et al., 2005a Alboconis cretacica Nel et al. Lebanese (Neocomian) Nel et al., 2005a —Family Rhachiberothidae Tjeder Paraberotha acra Whalley Lebanese (Neocomian) Whalley, 1980 Retinoberotha stuermeri Schluter French (Cenomanian) Schluter, 1978 Eorhachiberotha burmitica Engel Burmese (Albian) Engel, 2004b Whalfera venatrix (Whalley) “British” (Eocene) Whalley, 1983; Engel, 2004b Alloberotha petrulevicii Nel et al. French (Cenomanian) Nel et al., 2005b Chimerhachiberotha acrasarii Nel et al. Lebanese (Neocomian) Nel et al., 2005b Spinoberotha mickaelacrai Nel et al. Lebanese (Neocomian) Nel et al., 2005b Oise a celinea (Nel et al.) Parisian (Eocene) Nel et al., 2005b ORDER MEGALOPTERA LATREILLE —Family Sialidae Leach Eosialis dorisi Nel et al. Parisian (Eocene) Nel et al., 2002 ORDER RAPHIDIOPTERA NAVAS —Family Mesoraphidiidae Martynovc Nanoraphidia electroburmica Engel Burmese (Albian) Engel, 2002a a“British” amber is the same as Baltic amber and is material from the Baltic that has washed up on the British Isles. These deposits are middle Eocene in age (see discussion of dating in Engel, 2001; Grimaldi and Engel, 2005). bAn undescribed specimen from Canadian amber has been reported as a possible berothid (McAlpine and Martin, 1969; Pike, 1995). cAn undescribed larva of this group was recorded by Engel (2002a), and a further account of various Cretaceous snakefly immatures is summarized by Perrichot and Engel (in press). 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 Adults are short-lived, occur along watersides, Southeast Asia (Indosialis). In addition, 10 and apparently may feed on pollen and nectar, 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¬ imately 66 living species (most in the nominate sified in its own basal subfamily, the genus Sialis) that are distributed worldwide Dobbertiniinae. but are principally Holarctic with a few species 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 TABLE 4 Neuropterida in Dominican and Mexican3 Amber Taxa Reference ORDER MEGALOPTERA LATREILLE —Family Sialidae Leach Sialis (Protosialis) casca Engel & Grimaldi n.sp. Present study ORDER NEUROPTERA LINNAEUS —Family Ascalaphidae Rambur Amoea electrodominicana Engel & Grimaldi n.sp. Present study Ululodes paleonesia Engel & Grimaldi n.sp.b Present study —Family Chrysopidae Schneider Chrysopa glaesaria Engel & Grimaldi n.sp. Present study Chrysopa vetula Engel & Grimaldi n.sp. Present study Leucochrysa (Nodita) prisca Engel & Grimaldi n.sp. Present study —Family Coniopterygidae Burmeisterb Coniopteryx antiquua Engel & Grimaldi n.sp. Present study Spiloconis glaesaria Meinander Meinander, 1998a Spiloconis oediloma Engel & Grimaldi n.sp. Present study —Family Hemerobiidae Latreille Notiobiella thaumasta Oswald Oswald, 1999 Sympherobius sp. Present study —Family Mantispidae Leach Dicromantispa movonei Engel & Grimaldi n.sp. Present study Dicromantispa electromexicana Engel & Grimaldi n.sp. Present study Feroseta prisca Poinar Poinar, 2006 —Family Myrmeleontidae Latreille Porrerus dominicanus Poinar & Stangeb Poinar & Stange, 1996 & Dicromantispa electromexicana is presently the only described neuropterid from Mexican amber. bImmatures (all reported upon herein) are recorded for Ascalaphidae, Chrysopidae, and Myrmeleontidae. from the West Indies except for one species on 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) but are rounded in the fossil vein meeting R slightly distad to R + / 2+3 2 3 (not flamelike as in S. flammata). R4+5 fork; posterior branch of R4/R5 sepa¬ 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 AMERICAN MUSEUM NOVITATES NO. 3587 TABLE 5 Named Fossil Sialidae Taxa Deposit Reference Dobbertinia reticulata (Handlirsch) Jurassic, Germany Ansorge, 2001 Eosialis dorisi Nel et al. Parisian amber Nel et al., 2002 Indosialis beskonakensis Nel Miocene, Turkey Nel, 1988a Proindosialis cantalensis Nel Miocene, France Nel, 1988a Sialis strausi lilies Pliocene, Germany lilies, 1967 Sialis groehni Wichard Baltic amber Wichard, 1997 Sialis muratensis Nel Miocene, France Nel, 1988a Sialis (Protosialis) baltica Wichard Baltic amber Wichard, 1997 Sialis (Protosialis) herrlingi Wichard Baltic amber Wichard, 2002 Sialis (Protosialis) casca n.sp. Dominican amber Present study Sialis (Protosialis) voigti Wichard & Engel Baltic amber Wichard & Engel, 2006 midpoint; apparently light brown; apex of ORDER NEUROPTERA LINNAEUS abdomen curled under, with genitalia pointed FAMILY MANTISPIDAE LEACH anteriad. The mantispids are perhaps one of the most 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) of an elongate prothorax and and is a reference to the age of the specimen. raptorial forelegs is remarkable. Adults tend Fig. 1. Photomicrograph of holotype 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 lateral aspect. to be diurnal and are often found on flowers, ators although the larvae of at least one although a few lineages are nocturnal or studied species are ambulatory. The subfamily crepuscular. Some species are brightly colored Drepanicinae is entirely unknown biologically. 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 sites of spiders, and the first-instar larva either 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 Calomantispinae appear to be generalist pred¬ searches directly for an egg sac. Later larval 10 AMERICAN MUSEUM NOVITATES NO. 3587 TABLE 6 Named Fossil Mantispidae3 Table Deposit Reference Climaciellal henrotayi Nel Oligocene, France Nel, 1988b Dicromantispa electromexicana n.sp. Mexican amber Present study Dicromantispa moronei n.sp. Dominican amber Present study Feroseta prisca Poinar Dominican amber Poinar, 2006 Gerstaeckerella asiatica Makarkin Late Cretaceous, Kazakhstan Makarkin, 1990 Liassochyrsa stigmatica Ansorge Early Jurassic, Germany Ansorge & Schliiter, 1990; & Schulter Wedmann & Makarkin, 2007 Mesomantispa sibirica Makarkin Early Cretaceous, Siberia Makarkin, 1996 Promantispa similis Panfilov Late Jurassic, Karatau Panfilov, 1980 Prosagittalata oligocenica Nel Oligocene, France Nel, 1988b Symphrasites eocenicus Wedmann & Eocene, Germany Wedmann & Makarkin, 2007 Makarkin Vectispa relicta (Cockerell)b Eocene, England Cockerell, 1921 aAlthough Willmann (1994) transferred the Cretaceous amber Retinoberotha and Paraberotha to Mantispidae, these are rhachiberothids. Similarly, Whalfera venatrix (Whalley, 1983) is excluded owing to its likely position in Rhachiberothidae (e.g., Engel, 2004b; Wedmann and Makarkin, 2007). bOriginally proposed in Mantispa and then transferred to Promantispa Jarzembowski (1980: nomen praeoccupatum, nee Panfilov, 1980) and finally to Vectispa Lambkin (1986: nomen novum pro Promantispa Jarzembowski, 1980). instars are relatively immobile. Pupation oc¬ mantispid is Feroseta priscus Poinar in curs within the host egg sac. Adults are actively Miocene Dominican amber (Poinar, 2006). predaceous on a variety of other insect groups. 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 Aspock and Mansell therefore perhaps not surprising that their (1994) and Aspock et al. (2001) by considering parasites would also eventually be discovered. this group as allied to the Berothidae. 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 the Baltic amber deposits of 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, major profemoral spine longer than the genus is very likely a rhachiberothid protarsus, fewer c-sc crossveins, and smaller (Aspock 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