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Ellis G. MacLeod.
The Neuroptera of the Baltic Amber. I. Ascalaphidae, Nymphidae, and Psychopsidae.
Psyche 77:147-180, 1970.

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Pig. 1. Neadtlphw frotae n. sp. Dorsal aspect of holotype.



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PSYCHE
Val. 77 June, 1970
No. 2
THE NEUROPTERA OF THE BALTIC AMBER.
I. ASCALAPHIDAE, NYMPHIDAE, AND
PSYCHOPSIDAE1
INTRODUCTION
Along with the related Megaloptera and Raphidiodea, fossils representing the Neuroptera (Planipennia) are known from the Permian of Russia (Martynova, 1962)) Australia (Riek, 1953), and Kan~as.~
Several of these early fossils, such as those of the Palaemerobiidae and Pern~ithonidae, have a decidedly modern aspect and by the mid-Mesozoic the living families Chrysopidae (Adams, I 967)) Nymphidae (Adams, I 958), and Psychopsidae were already in existence. Other than the Baltic amber, Tertiary deposits have yielded a disappointingly small number of Neuroptera. Among these the Chrysopidae are relatively the most numerous, although in fact they are actually represented by only a small number of fossils from the Florissant shales of Colorado and a few additional specimens from Europe.
Both the Megaloptera and Raphidiodea are known from the Baltic amber (Hagen, 1856; Carpenter, 1956) from a very limited number of specimens, whereas the Neuroptera are much more com- 'Published with the aid of a grant from the Museum of Comparative Zoology at Harvard College and funds from NSF Grant GB-19922 (R. C. Rollins, Principal Investigator, Harvard University). Manuscript received by the editor, July 15, 1970. "epartment of Entomology, University of Illinois, Urbana, Illinois. 'The two insects described as planipennian Neuroptera from the Lower Permian of Kansas by Tillyard (1932, 1937) are now considered as having quite different affinities. One of them, Permobiella perspicua Tillyard, is now recognized as belonging to the Caloneurodea (Martynov, 1938a, 1938b; Carpenter, 1943a) ; and the other, Permoberotha villosa Tillyard has been assigned to the Glosselytrodea (Martynova, 1962), which is regarded by Carpenter (1964) and Sharov (1966) as closely related to the Neuroptera. The collections of the Museum of Comparative Zoology do, however, con- tain a specimen from this deposit which I regard as a true iieuropteran.



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THE NEUROPTEROUS LARVAE OF THE BALTIC AMBER Without formally naming them, Hagen ( 1856) provided detailed descriptions of two larval specimens which he felt belonged to the Neuroptera. The first of these, designated as "Larva Hemerobii", was described as possessing long, scimitar-shaped jaws, a broad head, and round, strongly constricted setigerous tubercles on the thorax. The remains of a trash packet, consisting of stellate plant hairs, was noted to be present in the vicinity of the larva. It has proved impossible to learn of the present whereabouts of this larva and this is unfortunate since it is quite likely that this specimen is a chrysopid, a family, as noted above, otherwise curiously unrepresented in the Baltic amber. Hagen suggested that this "Larva Hemerobii" might possibly belong to one or another of his several species of amber Hemesobiidae, but our knowledge of present-day hemerobiids rules this out as their larvae are now known to be non-trash carrying forms lacking tubercles and with unspecialized setae. The idea that hemerobiid larvae were trash carriers was wide-spread in the last century and for its time Hagen's suggestion, though incorrect, was reasonable. I have seen several specimens of trash-carrying cole- opterous larvae from several amber collections, and it might seem possible that Hagen misidentified one of these as a neuropteran. The mouthparts of these beetle larvae, however, are small and typically coleopterous, whereas Hagen describes the jaws of his larva as ". . . etwa noch einmal so long als der' Kopf, sabelformig mit scharf er Spitze, glatt und zahnlos, und zangenformig gestallt wie bei Hemerobius und Chrysopa." Various species of Ascalaphidae, Myrmeleontidae, and Nymphidae are also known to construct dorsal trash packets, but the larvae of none of these families really have globular, strongly constructed tubercles and, of course, all have toothed mandibles.
Hagen's second larval specimen, designated by him simply as "Larva", has been located in the portion of the Berendt collection now residing in the Museum of Humboldt University, Berlin. It is a member of the Psychopsidae, a family which at the time was unknown to Hagen in the larval stage. This specimen is redescribed below.
In addition to these specimens, larvae, apparently of a myrme- leoritoid facies, were mentioned on a number of occasions in the early literature dealing with the Baltic amber (Berendt, 1830, 1845; 'Burmeister, 1832; Hope, 1834). Upon study, it emerges that only Berendt claimed to have actually seen such a specimen,



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19701 MacLeod - Baltic Amber Neuroptera 15 1 the notations of Burmeister and Hope being obviously secondary sources. Such a larva was, curiously, not mentioned by Hagen and, aside from the later bibliograhic compilations of Scudder ( I 886,
1891 ) , which do not note that the earlier references pertained to a larva, this larval type was not mentioned again until 1910. In that year Klebs, again without formal description, mentioned the exist- ence of this type of larva in his extensive personal collection and, without reaching any final conclusions, discussed the opinions of several persons whom he had contacted as to whether it was an ascalaphid or a myrmeleontid. Klebs' collection was subsequently acquired by Albertus University, Konigsberg (Andrie, 1937), while, as noted above, portions of Berendt's collection are now located in Berlin; however, a recent search of the rema-ins of these collections has failed to produce any specimens on which these records might have been based. Both Handlirsch ( 1906, 1925) and Bachofen-Echt ( 1949)) again apparently misled by the earlier references of the nineteenth century, have also dutifully recorded the existence of a myrmeleontid from the amber.5
Weidner (1958) has finally given a formal description of a myr~neleontoid larva and his specimen has been available to me for restudy. In addition, a magnificent specimen of an ascalaphid larva from the Haren Collection of the Museum of Comparative Zoology will also be described. As noted below, it is possible that this latter specimen is the one which was once owned by Klebs. FAMILY ASCALAPHIDAE SCHNEIDER, I 845
The present distribution of this small family includes all major zoogeographic regions, although there is a pronounced concentration of the major taxa in the tropical and subtropical regions of the New and, particularly, the Old World. Two fossil ascalaphids, both adults, have been described from Tertiary deposits of Europe: AscaIaphus proavus Hagen (1858), from the brown coal near Linz, West Germany, and A. edwardsi Oustalet (1870) from Saint- - .--
^Like Scudder, the listings of both Handlirsch and Bachofen-Echt suggest that they are based on adult fossils. Handlirsch, however, is clearly citing the old reports of Berendt (1830, 1845) and Burmeister (1832) which deal with a larval specimen. Handlirsch and Bachofen-Echt have introduced additional confusion by claiming not one myrmeleontid species from the amber, but three. The three "species" of Handlirsch's account trace back to the three early papers just noted, all of which deal with a single (larval) specimen. Bachofen-Echt has simply repeated Handlirsch's error.



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Psyche
Fig. 2.
Neadelphus protae n. sp. Dorsal aspect of holotype. The body, which is slightly curved in the specimen, is here depicted as straightened out.




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19701 MacLeod - Baltic Amber Neuroptera 153 Gerand le Puy, France. Both of these deposits are referred to Upper Oligocene horizons. Navis ( 1913) has erected a new genus for each of the fossil species, Borgia and Ricartus respectively, but this treatment, as well as their original assignment to Ascalaphus, has no present value as the fossils have never been restudied in the light of the modern classification of the family. Weidner ( 1958) described a larva from the Baltic amber which he felt was an ascalaphid. My study of this specimen had indicated that it is actually a nymphid and it is dealt with below under that family. The collection of the Museum of Comparative Zoology contains a beautifully preserved small larva which is without question an ascalaphid. Because this specimen provides the earliest geological record for a member of this family, a formal description of the larva is presented here.
Neadelphus new genus
(figs. I-4)6
Description. Head capsule: quadrate, parallel-sided, with cor- date postero-lateral margins; surface generally smooth, raised bases of setae imparting only a slightly rugose texture to surface. Ocular tubercles large, prominent, approximately parallel-sided, each with the usual seven stemmata, six visible in dorsal view, the seventh located ventrally. Antenna1 tubercle very small. Jaws very long and slender, nearly straight for most of their length, with the three true teeth of the medial mandibular surface beyond the mid-point of the mandible.
Labial palpi short, slender.
Body: prothorax approximately elliptical, with a bilateral pair of small, globular setigerous tubercles antero-laterally; protboracic spi~ades only slightly produced as a low cone, elliptical in outline. Meso- and rnetathorax broader and shorter, each with two elongate setigerous scoli on each lateral margin, the posterior member of each pair distinctly smaller than the anterior one. Legs as in fig. 4, the tarsi quite distinct from the tibiae on the pro- and mesothoracic segments, the metathoracic tarsi fused to their tibiae and showing no indication of a line of fusion.
Abdominal segments I-VIII with a bilateral pair of elongate, unflattened, setigerous scoli, the pairs located on segments I-VII 'The
specimen upon which these descriptions are based has previously been illustrated, as a color photograph, on p. 21, of the volume of the Life Nature Library entitled The Insects (1962. New York: Time Inc.).



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154 Psyche [June
subequal in length, the pair on segment VIII much shorter; with no trace of a ventral series of scoli. Only abdominal spiracles 1-11 visible in fossil, these located ventrally, directly beneath base of corresponding scolus (fig. 4, sp I, sp 2). Posterior margin of ninth sternum lacking short, stout, "digging" setae. Head and body covered with short black setae which are longer along the lateral margins of the head capsule, down the midline of the thorax and abdomen and, particularly, on the scoli; longer setae narrowly lanceolate in shape.
Type species, the following -
Neadelphus protae new species
Description. Setae along lateral margins of head only moderately elongated ; central mandibular tooth the la,rgest of the three, distinctly nearer the anterior tooth than the posterior tooth, the posterior tooth the smallest; right antenna with 14 sub-segments distal to the scape, the left antenna with 12.
Maximum width of head capsule immedia,tely behind ocular areas - 1.40 mm. Body length from anterior -most clypeal margin to tip of IX abdominal segment - 3.6 mm.
Holotype: No. 5848, in the Haren Collection of Baltic amber of the Museum of Comparative Zoology. The name for this insect is derived from the Greek Phaethon myth and translates literally as "Prota's new brother".
The specimen, undoubtedly a first-instar larva, is contained in a nearly square block of pale-yellow amber with all important taxo- nomic details easily visible.
The block has been mounted on an
oblong piece of glass by some previous owner. This glass bears a label with the printed notation "Coll. Dr. Klebs", to which some- one has added in India ink "N27-". To this I have added the MCZ type number in blue ink and a new label with the designations "Neadelphus protae MacLeod" and "Holotype". It seems certain that this was the larva mentioned by Klebs in 1910, and it is possible that the references to a larva from the early nineteenth century also pertain to this specimen. The route by which it finally reached the Haren Collection is not known.
Generic diagnosis and discussion. The cordate posterior ma,rgins of the head capsule and large ocular tubercles with seven stemmata combined with the distinct fusion of the tibia and tarsus of the metathoracic leg show, without question, that this larva belongs to the Ascalaphidae.




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19701
MacLeod - Baltic Amber Newoptera
Fig. 3. Neadelphus protae n. sp. Detail of ocular tubercle (OT), an- tennal tubercle (AT) and antenna, jaw base, and chaetotaxy of holotype, dorsal view.
From the time of Hagen's pioneering synthesis of 1873, a number of different types of larval ascalaphids have been described (sum- marized in MacLeod, 1964 and MS in reparation). A careful reading of this literature, however, reveals few cases in which larvae have been associated with their adults either through rearing or by hatching from eggs laid by a captive female. Most often these associations have been made by a loose, deductive process of elimina- tion from a list of the species which are known as adults from the general area from which the larva was obtained. In view of the continuing poor state of our knowledge concerning the distribution of the species of the Ascalaphidae, this procedure is of no real value. Larval-adult associations which I believe to be reliable have only been achieved for the genera Ascalaphus, ~Heli~omit~us, Pseudoptynx, Suhpalacsa, and UZuZodes, all of which belong to the subfamily Ascalaphinae. One might add to these the description by Froggatt (1902) of the rearing of a species now placed in the genus Acmo- notus, but his unillustrated account is too general to be of any present use. In addition to the genera just noted, I can add to the list of associated forms the larvae of the genus Ascalofitynx, repre- senting the subfamily Ascaloptynginae, which I have reared. For strictly nomenclatorial reasons, the larvae of the genera Neulatus



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Psyche [June
Figs. 4, 5.
Larvae of Nymphidae and Ascalaphidae.
Fig. 4. Neadelphus protae n. sp. Ventral view of holotype showing position of first two abdominal spiracles (spl, sp2) and leg segmentation, setae omitted.
Fig. 5.
Larvae of ?Pronymfhes mengeanus (Hagen), dorsal view, show- ing details of lateral scoli; setae of dorsal head capsule and of body proper omitted.




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19701 MacLeod - Baltic Amber Neuroptera I57 and Sodirus, which Navis (1913) erected for two unassociated larval forms, must also be noted here.
Although long experience
with Navis' taxonomic methods teaches one to anticipate novel procedures, I am hard put to understand this particular action since I believe that there is every reason to anticipate the discovery that these larvae produce adults which belong to long-established genera.
By comparison with these larvae and with a number of additional, unassociated forms available to me for study or described by pre- vious workers, Neadelphus appears to differ by the following combination of characters : I ) the quadrate, parallel-sided head capsule; 2) the relatively narrow, produced jaws which are curved only near their tips and which bear the three mandibular teeth distal to the midpoint of the jaw; 3) the elongate, unflattened shape of the twelve pairs of lateral scoli which are unaccompanied by the development of small, additional anterior or ventral scoli or tubercles on the abdominal segments; 4) the ventral position of the first two pairs of abdominal spiracles (and presumably of the remaining six posterior pairs). Various larvae of living species show one or a few of these features, but none show this unique combination. With particular respect to the larval-based genera Neulatm and Sodirus, Neadelphus differs in the form of its prominent, parallel-sided ocular tubercles. These are small and distally narrowed in Matus and rather short and hemispherical in Sodirus. In addition, this latter genus has flattened scoli and a small tubercle on several of the abdominal segments, immediately anterior to the scolus, which are features lacking in Neadelphus.
The ventral location of all eight airs of abdominal spiracles, which Neadelphus shares with several living genera including Ululodes, is presumably a generalized feature derived from the lateral position of these openings in unflattened ancestral nymphids. Such additional features of Neadelphus as the cylindrical shape of the scoli and the non-falcate development of the jaws, though also shared with some living forms, are probably additional examples of generalized character states. In contrast, the presence of only a dorsal series of abdominal scoli must be considered as a specializa- tion from the nymphid double series (see below). Thus, in this respect, the Oligocene Neadelphus appears already specialized by comparison to several living larval forms which retain a trace of the ventral series of scoli either as short projections beneath the main dorsal series on abdominal segments I and I1 (as in an un-



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associated larva which I have seen from Tanzania), or perhaps as the small tubercles anterior to the main scoli on abdominal seg- ments III-VII (as in Ascaloptynx and several unassociated forms from. Central and South America which I have studied). Other than these general observations, I do not believe that the inadequate state of our knowledge of the larvae of this family permits any more definite taxonomic assignment of N. protae at this time. FAMILY NYMPHIDAE RAMBUR, IS@
This phylogenetically important family is presently confined to the Australian Region where a small number of species, classified into six genera, are known. The group is obviously of great antiquity as the very closely related Nymphitidae is known from Triassic strata of Russia (Martynova, 1949) and the species Mesonymphes hageni 'Carpenter from the Bavarian Jurassic is already so similar to the living forms that Adams (1958) concluded that it should be included in the Nymphidae itself.
The only apparent Tertiary record of the family8 has been an adult specimen described from the Baltic amber by Hagen (1856) as Nymphes mengeanus and redescribed as representing a new genus, Pronymphes, by Kruger (1923). This specimen, originally from Menge's collection, retained only the basal portion of the wings and was missing its abdomen. Kruger's reexamination of the speci- men provided a few additional details and corrected several minor errors in Hagen's drawing of the wings. Kruger noted that the specimen was at that time contained in the collections of the Danzig Provincial Museum and, as it has proven impossible to locate, it may not have survived World War 11. My redescription of the genus Pronymphes is, then, derived from the accounts of Hagen and Kruger and may require future corrections. I will also here redescribe the larval specimen which Weidner ( 1958) has treated as a member of the Ascalaphidae. For reasons to be given shortly, I am convinced that the specimen is actually a nymphid, which I will tentatively refer to Pronymphes. I n my treatment of this family I am tentatively following the conclusions of Adams (1958) who presented important reasons against the separation of the Myiodactylidae from their close relatives in the Nymphidae. I have previously suggested (1964 and MS in preparation) that a fuller knowl- edge of the structure and ecology of the larvae of1 this group may force a reevaluation of this idea.
'Bachofen-Echt (1949) notes seven species as having been described from the Baltic amber. I am unable to determine the origin of this error.



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19701 MacLeod - Baltic Amber Veuroptera I59 Pronymphes Icriiger
/ ' ? ~ O ~ I Y I ~ ~ ~ C S Kriigei, 1923, Stett. Ent. Zeit. 84: 75-80. Type species (by original designation) : Nymphes mengeanus Hagen. Bachofen-Echt, 1949, Der Bernstein und seine Einschluse, p. 136.
Di'scrip~ion. Wings narrowly elongate. Fore wing: costal space tiarnw, not abruptly widened at base, costal crossveins unforked; 13
i\s+ A IA originating near base of wing, fused for a rather long distance before separation of MA; MP deeply forked at a level which is distinctly proximal to separation of MA from Rs, the two branches of MP running nearly parallel, the posterior branch sepa- rate from CuA; basal branches of Cu running close together and nearly parallel, CuA unforked in portion preserved, CUP with pectinate branches to hind margin of wing, lacking crossveins between branches basally, perhaps with a series of irregular cross- veins distally.
Hind wing: costal space and crossveins as in fore wing; origin of Rs+ MA much
further distal than in fore wing; MP deeply forked to base of wing, MP2 with wide fork slightly distal to origin of Rs+ MA; Cu very short, with only a few pectinate branches to hind margin of wing, unconnected by crossveins. Discussion. The elongate, narrow form of the wings of Pro- nymphes is quite unlike the short, broad wings with abrupt basal dilations of the costal space to be found in the living genera Mywdactylus and Nymphidrion and I believe that these genera bear no close phylogenetic relationship to the fossil genus. There are such additional differences in venational details as the unforked MP in the fore wing of Myiodactylus to reinforce this conclusion. The wings of the species of Osmylops, though somewhat narrower in proportion to their width, are still distinctly broader than those of Pronymphes and the costal space of Osmylops also has a pro- nounced dilation near the base.
In terms of their overall similarity in wing shape and venation, Pronymphes is obviously close to the living genera Nymphes, Aus- tronymphes, and, particularly, Nesydrion. Unfortunately, attempts to infer the probable details of the interesting phylogenetic inter- relationship of these four genera, from which I believe their phenetic similarities derive, will be somewhat frustrated until specimens show- ing the complete wings of Pronymphes are discovered. It is especially critical to learn the detailed structure of CuA in the fore wing and, particularly, whether this vein was forked or not. Certain similarities



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Psyche
Fig. 6. Larva of Wrofiymfhc~ mrngcanus (Hagen), dorsal view,



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19701 MacLeod - Baltic Amber Neuropfera i 61 and differences between Pronymphes and these living genera are, however, obvious.
Thus, in the fore wing, the rather far distal separation of MA from Rs is a feature common to all of these genera, while Pronymphes shares the deep, basal fork of MP in this wing with both Nymphes and Nesydrion. In both Nymphes and some of the species of Nesydrion this forking is distal to the separation of MA from RS but in other species of Nesydrion the fork of MA is as far proximal as it is in Pronymphes. Austronymphes differs from the other three genera in having MP unforked in the fore wing. The three living genera show various stages in the evolution of the prominent fork of CuA in the fore wing, which is characteristic of all the higher myrmeleontoid families, with this fork nearly marginal in Nesydrion, just proximal to the middle of the wing in Austronymphes and far proximal to mid wing in Nymphes where it is closer to the base than the fork of MP or the separation of MA from Rs. The exact development of this important venational feature cannot yet be determined in Pronymphes, although it is obvious that if a fork of CuA was present it certainly was considerably more distal than in Nymphes and, probably, Austrony~hes. The rather distal point of origin of Rs + MA in the hind wing of Pronymphes is quite like the condition of this vein in Nesydrion and this feature contrasts sharply with the distinctly more basal origin of this vein in all other living genera of the family. Finally, in the weak, submarginal development of the fork of MP2 in the hind wing, Pronymphes is similar to the condition found in Austronymphes and Neyydrion and


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