Family Affinity of the Genus Palaeopsychops Andersen With

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SYSTEMATICS Family Afﬁnity of the Genus Palaeopsychops Andersen with Description of a New Species from the Early Eocene of British Columbia, Canada (Neuroptera: Polystoechotidae)

1 2 VLADIMIR N. MAKARKIN AND S. BRUCE ARCHIBALD

Ann. Entomol. Soc. Am. 96(3): 171Ð180 (2003) ABSTRACT Palaeopsychops dodgeorum sp. n. from the Early Eocene Okanagan Highlands of Quilchena, British Columbia, Canada is described. The systematic position of the genus Palaeopsychops Andersen, 2001 is discussed, interpreting this as most closely associated with Polystoechotidae. Osmylites protogaea (Hagen 1862) is considered as nomen nudum and an objective synonym of Osmylites excelsa (Oppenheim, 1888), syn. n.

KEY WORDS Eocene, Polystoechotidae, Neuroptera, Okanagan Highlands, fossil insects

THE FAMILY POLYSTOECHOTIDAE is now a relict group very likely belong to this family (Makarkin, personal represented by only four species belonging to three observation). genera: Polystoechotes Burmeister, 1839, Platysto- Late Cretaceous and Tertiary Polystoechotidae echotes Carpenter, 1940, and Fontecilla Nava´s, 1932, have hitherto been unknown, because the single de- which are restricted to the New World (Oswald 1998). scribed species (Polystoechotes piperatus Cockerell, Although fossils of this family are few and controver- 1908 from Late Eocene of Florissant) was considered sial, occurrences in the Mesozoic of insects referable to be a psychopsid (Carpenter 1943, MacLeod 1970). to Polystoechotidae indicate that this family may have However, examination of the holotype shows that a been more diverse and widespread. The Early Jurassic polystoechotid afÞnity is most probable, although this species Mesopolystoechus apicalis Martynov, 1937, de- species certainly doesnÕt belong to the genus Polysto- scribed from Tajikistan, Central Asia, has been con- echotes. Unfortunately, the sole specimen of this spe- sidered the single member of the family Mesopolys- cies is an incomplete and poorly preserved forewing, toechotidae Martynova, 1949. Lambkin (1988) and its systematic position is not clear. Another Ter- implied his opinion of possible synonymy of Me- tiary genus, Palaeopsychops Andersen, 2001, was re- sopolystoechotidae and Polystoechotidae when he recently described from the Late Paleocene/Early Eo- ferred to M. apicalis as a polystoechotid (p. 455) and cene of Denmark in the family Psychopsidae, “probable Polystoechotidae” (p. 457). He also re- however, we consider this genus as most closely as- viewed the other four fossil species from the Late sociated withPolystoechotidae(see “Determination Triassic to Late Jurassic probably belonging to Polys- of Family AfÞnity”). toechotidae: Lithosmylidia lineata Riek, 1955 (Late A large lacewing was recently Þgured along witha Triassic of Australia); Kasachstania fasciata PanÞlov, short description by Archibald and Mathewes (2000, 1980, Osmyliodea distincta PanÞlov, 1980, and Ptero- Figs. 6A, 20). The authors brießy discussed its possible calla superba PanÞlov, 1980 (all from the Late Jurassic family afÞnity, comparing it withOsmylidae and of Kazakhstan). Another two species were assigned to Polystoechotidae. In this paper we provide a detailed Mesopolystoechotidae by Whalley (1988) description of this specimen and discussion of its sys- [Megapolystoechus magniﬁcus Tillyard, 1933 (Late tematic position. We interpret this as a new species of Triassic of England)] and Hong (1983) [Mesopolys- the genus Palaeopsychops. toechus wangyingziensis Hong, 1983 (Middle Jurassic of China)]. Apart from these occurrences, there are Materials and Methods undescribed species from the Late Jurassic of Kaza- khstan and Early Cretaceous of southern Siberia that The specimen is from the Early Eocene lacustrine shales of the “Coldwater beds” exposed along the banks of Quilchena Creek at Quilchena in south-cen- 1 Institute of Biology and Soil Sciences, Far Eastern Branchof the tral British Columbia, Canada. The Quilchena locality Russian Academy of Sciences, Vladivostok, 690022, Russia (e-mail: is the oldest of the “Okanagan Highlands” series of [email protected]). 2 Department of Organismic and Evolutionary Biology, Harvard fossiliferous Eocene shale and amber deposits in Brit- University, Museum of Comparative Zoology, 26 Oxford Street, Cam- ish Columbia and Washington State. This locality bridge, MA 02138 (e-mail: [email protected]). bears numerous fossil insect, Þsh, plant, and bird

0013-8746/03/0171Ð0180$04.00/0 ᭧ 2003 Entomological Society of America 172 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 3

(feather, coprolite) remains. Although most shale sopolystoechus by Hong (1983), that is represented from Quilchena is easily worked, unfortunately the only by an incomplete forewing. particular nature of the matrix of this fossil (relatively In the drawing of the holotype of Palaeopsychops large-grained, consolidated to the fossil, and not ßak- latifasciatus (Fig. 10: Andersen 2001), the subcosta, ing away withneedles) makes preparation unusually anterior radial trace (“R1”), and anterior sectoral trace problematic. (“Rs”) all terminate together, however, the photo- Terminology of forewing venation and wing spaces graph (his Fig. 3) shows that whereas Sc and R1 do employed here mainly follows Oswald (1993a, 1998), probably fuse, Rs runs separately to the wing margin, in particular withregards to themedian vein: “MA” of as in our specimen. We Þnd it most probable that in authors (e.g., Carpenter 1940; MacLeod 1970) is the other species of Palaeopsychops the conÞguration of most proximal branchof Rs, and “MP”Õ is simply M. these veins is the same, and therefore, we assume that there is no “vena triplica ” in this genus (see Oswald 1993b: 41 for discussion of this character). Because the Taxonomy distal nygma (not mentioned in the text of Andersen (2001), but clearly seen in the photograph, Fig. 3) is situated in the neuropteran forewing between the Palaeopsychops Andersen 2001 most proximal two branches of Rs, it follows then that

Palaeopsychops Andersen 2001: 422. AndersenÕs M1 is actually the most proximal branch of Diagnosis. Distinguished from other polystoechtid Rs, his M2 is M1, his Cu1 is M2 (Andersen 2001: Fig. 10), genera by the following combination of forewing his Cu2 is CuA, and his A1 is CuP (Andersen 2001: Fig. characters: (1) broad-subtriangular in shape [more or 9). less elongate in all other genera, except for Fontecilla]; (2) hind margin of wing not falcate [falcate in Fon- Palaeopsychops dodgeorum New Species tecilla]; (3) M2 strongly pectinately branched (shared by fossil genus Pterocalla) [few pectinate branches in Neuroptera incertae sedis: Archibald and Mathewes all other genera]; (4) outer gradate series of crossveins 2000: 1445, 1448, Figs. 6A, 20. in radial space well developed (shared by all extant Diagnosis. Distinguished from other species of ge- genera) [absent in fossil genera, except hindwings of nus by numerous, irregularly arranged crossveins in Mesopolystoechus apicalis and Osmyliodea distincta radial space (in all other species of Palaeopsychops (but see discussion)]; (5) transverse bands (fasciae) these crossveins form inner gradate series), somewhat distinct [absent in all extant genera and never de- more elongate shape of forewing. tected in any fossil genus]. Description. Forewing broad-subtriangular in Discussion. The genus Palaeopsychops was de- shape, length 45 mm. Trichosors indistinct, visible only scribed from the Late Paleocene/Early Eocene of at hind wing margin where margin well-preserved Denmark (“Mo-clay”), withfour species: P. latifascia- (see Fig. 5, not shown in Fig. 1). Distal nygma distinct, tus Andersen (type species), P. abruptus Andersen, P. appearing as small bulge in dilation between most angustifasciatus Andersen, and P. maculatus Andersen. proximal two branches of Rs (see Fig. 4). Costal space The latter species is possibly a synonym of one of the basally expanded, narrowed toward apex, withcom- other three, as the holotype is a hindwing, not forew- plete gradate series of crossveins parallel to anterior ing as in Andersen (2001) (the costal space not ex- wing margin. Subcostal veinlets always forked, some- panded basally as in forewings). Withthepresent times several times. Subcostal space rather wide along description of Palaeopsychops dodgeorum, the genus entire length, crossveins not preserved. Apical portion expands its distribution to NorthAmerica. of Sc not preserved (possibly obscured by matrix). R1 ϩ Palaeopsychops represents the single described Ter- (or Sc R1) entering margin apparently at wing apex, tiary genus in the family. A number of genera were withbrancheslong and forked. R 1 space somewhat erected from the Late Jurassic to Late Triassic (see narrower than subcostal space, with Ϸ10 crossveins. introduction) and none from the Cretaceous. Palaeo- Origin of Rs not preserved (possibly obscured by psychops is easily distinguished from all extant genera matrix), but clearly close to base of wing. Rs with (see “Diagnosis”). Of the Jurassic-Triassic genera only numerous (33Ð34 in number) pectinate branches, the genus Mesopolystoechus Martynov, 1937 is similar sometimes fusing withone another;many irregularly to Palaeopsychops, as seen from the venation of a spaced crossveins between them in central part of hindwing of Palaeopsychops maculatus (Andersen wing. Outer gradate series in radial space distinct, 2001: Fig. 12). The type species (M. apicalis) is rep- nearly parallel to posterior margin, continuing radio- resented by a well preserved, but incomplete hind- medially to anal space. Fork of M not detected (ob- wing (Martynova 1962: Fig. 860). The single known scured by matrix); anterior branchof M (M 1) con- specimen of Osmyliodea distincta is also a hindwing. cave, unbranched before outer gradate series;

However, the venation of hindwings in many Neu- posterior branchof M (M 2) convex, pectinately roptera greatly differs from that of forewing, and so branched in distal portion (with Ϸ8Ð10 long, parallel does not allow comparison between genera described and rather oblique branches). Cu divided into CuA from only forewings and those described solely from and CuP close to base of wing. CuA strongly convex hindwings. Therefore, there is insufÞcient reason to basally, distally slightly concave and pectinately consider congeneric another species assigned to Me- branched (with 11Ð12 long and rather oblique May 2003 MAKARKIN AND ARCHIBALD:POLYSTOECHOTIDAE FROM THE EARLY EOCENE 173

Fig. 1. Palaeopsychops dodgeorum, forewing venation of the holotype Q-0422a, b (composite drawing from part and counterpart). 1AÐ3A, anal veins; CuA, anterior cubitus; CuP, posterior cubitus; M1,M2, anterior and posterior branches of media; n, distal nygma; R1, Þrst branch of radius; Rs, radial sector; Sc, subcosta. Demarcation lines show a question mark where precise limits are not certain.

branches parallel to that of M2). CuP pectinately oblique lighting, clearly shows pigmentation pattern branched, branches (four in number) strongly (see Fig. 3). Furthermore, absence of coloration in a oblique; 1A comparatively short, dichotomously fossil wing does not necessarily indicate absence in branched; 2A pectinately branched, with branches life; the type specimen of P. maculatus, a hindwing rather long; 3A poorly preserved, few-branched. Jugal (above), has clearly preserved color patterns; and the lobe, if present, not preserved (possibly obscured by basally broadened costal region in our specimen make matrix). Wing pattering represented by maculations this readily identiÞable as a forewing. throughout, evident on the part (Q-0042a) (see Fig. The distal portion of M2 on our specimen may ap- 3). Near costal margin fasciae more conspicuous, pear to be a continuation of CuA obscured by matrix oblique and wide. Outer gradate series clearly shaded. in middle of M2-CuA. Concavity and convexity rela- Type Material. HOLOTYPE: a rather well pre- tionships of veins indicate, however, that M2 is entirely served, almost complete left forewing, Quilchena, (except most basally) strongly convex, and the distal BritishColumbia, Canada, Early Eocene. Depository: part of CuA is slightly concave (Fig. 2), which allows fossil collection of Simon Fraser University (Burnaby, this to be considered as the distal portion of M2, not BritishColumbia, Canada) (Q-0422 a, part; Q-0422 b, CuA. Because of incomplete preservation of venation counterpart). in this portion of the wing, this interpretation is ten- Etymology. The speciÞc epithet is in honor of Ken- tative. neth Dodge and his father Ken Dodge, who collected and generously donated this specimen to the Simon Determination of Family Afﬁnity Fraser University collection; the Latin sufÞx -orum shows genitive case of plural nouns. The genus Palaeopsychops is of considerable interest Discussion. We assign this new species to the genus because of its large size and dense venation, particu- Palaeopsychops by agreement withall principal char- larly P. dodgeorum. At Þrst glance, this and another acters of the diagnosis: forewing venation, similar size, similar specimen from Quilchena (Archibald and and color pattering. We treat the numerous crossveins Mathewes, 2000, Figs. 6B and 21), as well as possibly scattered in the radial space of our specimen as a others in the Okanagan Highlands (unpublished), speciÞc character, and the fusing of some of the should be placed with psychopsid-like neuropterans. branches of Rs with one another as possibly an indi- Nevertheless, we consider this genus as most closely vidual abnormality of this specimen. Such anomalous associated withPolystoechotidae.Suchlarge neurop- specimens occur at times in extant species of various terans as Palaeopsychops dodgeorum often have dense families (Makarkin, personal observation). P. dodgeo- venation and numerous crossveins presumably result- rum is known only from the holotype forewing. ing from the structural demands of large wing size, but Andersen stated that this particular specimen is “pre- not always, as in Mesopsychopsis hospes (Germar sumably a hindwing 4.5 cm long, as it is without pig- 1839), and may appear then superÞcially similar to mentation pattern” (Andersen 2001: 433). No colora- psychopsid-like neuropterans. The systematics of fos- tion of this specimen was reported in Archibald and sil neuropterans is in need of revision, including phy- Mathewes (2000), nor evident in the accompanying logenetic analysis along withextant Neuroptera. De- photograph of the counterpart (Q-0042b), however, scriptions of taxa that are often obsolete or incomplete subsequent examination of the part, particularly with and in need of redescription are scattered throughout 174 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 3

Fig. 2. Palaeopsychops dodgeorum, Q-0422b. the literature. Examples include classiÞcations of the Nevrorthidae, Kalligrammatidae, Mesoberothidae, order and diagnoses of its families done by Martynova Mesithonidae. Of these, the following are too small in (1962) and Carpenter (1992). Therefore, in our anal- size to include these large insects, and are strongly ysis of family afÞnity we will consider all neuropteran different in the conÞguration of their principal veins: families. Conioptegyridae, Berothidae, Rhachiberothidae, Di- There are 35Ð47 families currently recognized in laridae, Sisyridae, Mesoberothidae, Mesithonidae, the order Neuroptera [ ϭ Planipennia] depending on Nevrorthidae. The others have characters that auto- synonymy. We have here divided them into three matically exclude them from consideration: Hemero- groups withregard to thegenus Palaeopsychops: biidae have at least two “radial sectors” (ϭoblique (1) Those families that we exclude because of fun- radial branches of Oswald 1993a), and are too small damental venational difference. The following fami- (here and below, only forewing venation is consid- lies have no species reasonably resembling Palaeopsy- ered, if otherwise not indicated); in Kalligrammatidae, chops: Chrysopidae, Conioptegyridae, Hemerobiidae the entire wing is Þlled with closely spaced crossveins (Promegalomidae), Mantispidae (Promantispidae), (including the subcostal space); the venation of Chry-

Berothidae, Rhachiberothidae, Dilaridae, Sisyridae, sopidae is strongly distinct (M1,M2, CuA, and CuP

Fig. 3. Palaeopsychops dodgeorum, Q-0422a. May 2003 MAKARKIN AND ARCHIBALD:POLYSTOECHOTIDAE FROM THE EARLY EOCENE 175

Nemopteridae), withsizes comparable to our specimen (sometimes almost identical in shape and size, e.g., in myrmeleontid Palparinae), but upon close examination, their venation has basic features very different from that of Palaeopsychops. In all of these families (except Nymphidae) trichosors are absent, subcostal veinlets (at least in basal half) are entirely simple, the costal space is narrow and not expanded basally, and most have branches of Rs, M and CuA more or less zig-zagged because of numerous crossveins (except Crocidae, Mesochrysopidae, Allopteri- dae, and Babinskaiidae for the latter character), and many other features characteristic of each family show Fig. 4. Palaeopsychops dodgeorum, Q-0422b. The distal conclusively that Palaeopsychops does not belong to nygma, between the most proximal two branches of Rs (ar- this family group. Only Nymphidae bears some exter- row). nal similarity with Palaeopsychops, in that there is dilation of the costal space (e.g., in Osmylops Banks, 1913 and Myiodactylus Brauer, 1866) but this extends never have strong pectinate branching); and Man- along the entire length, except near the wing base tispidae have an easily visible pterostigma, if not, the where this is narrowed, and the subcostal veinlets are subcostal space is distally broad. sometimes forked. However, in all species of this fam- All of the Permian families (Permithonidae, Palae- ily, humeral vein is not branched, and the branches of merobiidae, Parasisyridae, Permegalomidae, Permo- Rs are somewhat zig-zagged because of the presence psychopsidae, Permosisyridae, Sialidopsidae, and Ar- of numerous crossveins (including in R space) as in cheosmylidae) are now considered to be synonyms of 1 other members of Myrmeleontoidea. the Permithonidae (Whalley 1988, Novokshonov The family Osmylidae comprises many genera both 1996). Within Permithonidae, Permorapisma Tillyard, in the recent and fossil fauna, with occurrences re- 1926 (Late Permian of Australia) superÞcially most corded from the Early Jurassic to the Miocene. Os- resembles Palaeopsychops in having comparatively mylids have rather conservative venation, which in- large size (Ϸ20 mm), dense venation, and numerous cludes suchfeatures as thecostal space tapering crossveins (see Carpenter 1992: Fig. 192.2), however, toward the wing base; the basal subcostal veinlets are that genus as well as all other genera of Permithonidae always simple; Rs and its branches somewhat zig- may be easily distinguished from Palaeopsychops by zagged (in most Kempyninae Rs is straight, only its the following characters: costal space tapering toward branches are slightly zig-zagged); crossveins occur wing base, origin of Rs shifted distally, M dichoto- throughout almost the entire body of the wing; CuA mously branched and occupying a comparative large and CuP are almost always straight (except for Poris- area, CuA withfew distal pectinate branches,CuP mus McLachlan, 1868 and Gumilla Nava´s, 1912), par- simple or rarely withonly marginal end-twigging. “ ” allel to hind margin of the wing, and pectinately These traits exclude this family from consideration. branched. For these reasons, Osmylidae may be The families Myrmeleontidae, Palaeoleontidae, As- clearly ruled out. calaphidae, Nymphidae, Nemopteridae, Crocidae, (2) Those whose statuses are doubtful or unclear. Babinskaiidae (comprising the superfamily Myrme- The family Nymphitidae was established by Handlir- leontoidea), Mesochrysopidae, and Allopteridae all sch(1906Ð1908) for threeLate Jurassic genera: Nym- have wings narrow to elongate-oval (except for phites Haase, 1890, Gigantotermes Haase, 1890, and SialiumWestwood, 1854. Martynova (1949) added three other Mesozoic genera (Epigambria Handlirsch, 1939, Chrysoleonites Martynov, 1925, and Me- sonymphes Carpenter, 1929) and excluded Gigantot- ermes from the family. Later she added the Early Jurassic Sogjuta Martynova, 1958 and synonymized this family with the extant Nymphidae (Martynova 1962). Subsequently, two other genera were referred to Nymphitidae: Minonymphites Hong, 1980 (Middle Triassic of China) and Baissoleon Makarkin, 1990a (Early Cretaceous of Siberia). At present, the composition and status of this family remains one of most unresolved within Neuroptera. Of the recent authors who have considered it as a separate family (Whalley 1988, Makarkin 1990a, Carpenter 1992), only the latter gives a diagnosis, although this is Fig. 5. Palaeopsychops dodgeorum, Q-0422b. Detail of the very short. Carpenter considered this family as con- wing hind margin showing trichosors (arrows). taining four genera: Nymphites, Sialium, Chrysoleo- 176 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 3 nites, and Sogjuta. Of these, Sogjuta appears to be an Kalligrammatidae, e.g., very closely spaced crossveins osmylid (Lambkin 1988), and Chrysoleonites is be- throughout the wing including subcostal space, mod- lieved to be a member of Mesochrysopidae (PanÞlov erately expanded costal space for entire lengthwith 1980). The type genus includes three species: Nym- oblique subcostal veinlets connected by numerous phites priscus(Weyenbergh, 1869) (type species), N. crossveins, Sc and R1 merged very close to the wing braueri Haase, 1890, and N. lithographicus Handlirsch, apex; it is possible that it is a kalligrammatid. Makar- 1906. Unfortunately, the type species is represented by kinia, with M. adamsi (Martins-Neto, 1992), described an incomplete and rather poorly preserved specimen, from the Early Cretaceous of Brazil, also shows strong which is in need of redescription, and appears to be in afÞnity withKalligrammatidae. Osmylogramma, with too poor condition to allow determination of family O. martinsoni Ponomarenko, 1992 from Early Creta- afÞnity withcertainty. Thisfamily is in strong need of ceous of Mongolia, resembles Palaeopsychops at Þrst revision, and its true status will become clear only after glance, but when examined in detail, these genera are its type species is redescribed. different bothin venation (in Osmylogramma, imme- ϩ Palaeopsychops, however, is very far from the type diately after Sc R1 merge they are rather sharply genus of Nymphitidae, which possesses the features curved anteriad, the cubital space occupies a broader characteristic of the families of Myrmeleontoidea and area, closely spaced crossveins Þll the entire body of Osmylidae, i.e., the groups that we have ruled out the wing), and in coloration (in Osmylogramma the (forewings elongate, the costal space narrow, the sub- membrane is entirely very dark pigmented). costal veinlets in basal half of wing simple, CuA is The family Glottididae was established for the single strongly parallel to the hind margin). species Glottidia multivenosa Bode, described from The Epiosmylidae PanÞlov, 1980 contain only one the Early Jurassic of Germany (Bode 1953) and re- species, Epiosmylus longicornis PanÞlov, 1980 (Late cently redescribed by Ponomarenko (1995), who re- Jurassic of Karatau, Kazakhstan). This family has hith- ferred it to the Osmylopsychopidae. Although the erto been considered a synonym of Osmylidae (Lam- apical two-thirds of the wing is well preserved, it is bkin 1988, Makarkin 1990b). Examination of the ho- hard to assign this species to a family with certainty, lotype, however, shows that this synonymy is possibly as its venation resembles that of Osmylopsychopidae, unwarranted, as it lacks two characters found within Brongniartiellidae, and Prohemerobiidae. At any rate, almost all Osmylidae: trichosors and a sigmoid vein Palaeopsychops is unlike Glottidia in that Glottidia connecting R and M basally in the hind wing. It may lacks crossveins, M2 is not pectinately branched, and be interpreted, however, that this species represents CuP is rather strongly pectinately branched. either an isolated branch within Osmylidae or else a Seven genera from the Late Triassic to Early Cre- separate family; it remains difÞcult at present to de- taceous have been referred to the family Osmylitidae termine between these possibilities. In any case, (Osmylites Haase, 1890, Kirgisellodes Martynov, 1937, Palaeopsychops is very far from Epiosmylus (e.g., in the Petrushevskia Martynova, 1958, Tetanoptilon Bode, latter, the forewings are elongate, the costal space 1953, Mesosmylina Bode, 1953, Sinosmylites Hong, 1983 narrow, the subcostal veinlets in basal half of the wing and Sinosmylites Hong, 1996 [non Hong, 1983]) (Mar- are simple and widely spaced, and CuA and CuP are tynova 1962; Hong 1983, 1996), but all of these vary too strongly parallel to the hind margin). greatly from each other in their venation to comprise Handlirsch(1939) erected Epigambriidae for the same family. The only known species of the type Epigambria longipennis (a small species, 8 mm long, genus is Osmylites excelsa (Oppenheim, 1888) [ ϭ from the Early Jurassic of Germany), and was of the Osmylites protogaea (Hagen, 1862), syn. n.] from Late opinion that another two species from Late Jurassic of Jurassic of Germany. It is represented by a single Germany and England possibly belong to this family: specimen named (but neither described nor illus- Osmylites protogaeus (Hagen, 1862)(see below under trated) by Hagen (1862) as Chrysopa protogaea. Later Osmylitidae) and Osmylopsis buplicata(Giebel, 1856) this specimen was reexamined by Oppenheim (1888), (now familia incertae sedis: Carpenter 1992). Subse- who illustrated and brießy described it as Chrysopa quent authors (Martynova 1949, 1962; Ross and Jar- excelsa. When he established the genus Osmylites, zembowski 1993) placed the Epigambria within Nym- Haase (1890) included this species as “Osm[ylites] phitidae, synonymizing Nymphitidae and (Chrysopa) protogaea Hag. (ϭexcelsa Opp. nec Hag.”). Epigambriidae, however, as the only known specimen Thus the gender of this genus is feminine (not mas- is an incomplete hindwing with characters undeÞned culine) (Article 30.1.4.4 of the International Code of and insufÞcient to determine even placement in any Zoological Nomenclature, 4thed.), and Chrysopa pro- superfamily, synonymy is not justiÞed. It would be togaea Hagen, 1862 must be considered as nomen nu- better to consider Epigambria “Neuroptera incertae dum, and an objective synonym of Chrysopa excelsa sedis” as treated by Carpenter (1992). Oppenheim, 1888, although the name O. protogaea is Three genera have been assigned to PanÞloviidae: incorrectly used in the literature to the present (e.g., Panﬁlovia Makarkin, 1990c, Makarkinia Martins-Neto, Lambkin 1988, Ross and Jarzembowski 1993), and the 1997, and Osmylogramma Ponomarenko, 1992. How- name O. excelsa was hitherto considered a junior syn- ever, the status of this family is uncertain. Wings of the onym of O. protogaea. The holotype of Osmylites ex- type species of Panﬁlovia [P. acuminata (PanÞlov celsa, an overlapping fore and hindwing, has not yet 1980), from the Late Jurassic of Karatau, Kazakhstan] been redescribed. The Þgures in Oppenheim (1888: Pl. are elongate-oval withvenation resembling thatof 30, Fig. 2), Haase (1890: Fig. 10) and Handlirsch May 2003 MAKARKIN AND ARCHIBALD:POLYSTOECHOTIDAE FROM THE EARLY EOCENE 177

(1906Ð1908: Pl. 48, Fig. 4) show the venation to be referred to the genus Prohemerobius, but the majority poorly preserved. The characters visible in the forew- of them were synomynized by Ponomarenko (1995). ing are: all basal subcostal veinlets simple, widely We have included Prohemerobiidae in this group only spaced and nearly at a right angle to Sc; the costal because of basal dilation of costal space, forking of space is rather narrow with a somewhat expanded subcostal veinlets, and similar shape of forewing in basal region; CuA has few pectinate branches; and the Prohemerobius. However, all genera of this family anal veins are short. No other characters are certain. (even considered in the broadest sense) are unlike This species may belong to a number of families: Nym- Palaeopsychops in other respects. phidae, Mesochrysopidae, even Chrysopidae or Polys- Extant species of Psychopsidae, a small family with toechotidae. Osmylitidae is therefore a grab bag taxon, an Afro-Oriento-Australian distribution, have rather whose members should be reevaluated, and likely as- conservative forewing shape (always very broad) and signed to other families. Placing Palaeopsychops in this venation, with the costal space wide along the entire group therefore would only compound a problem length, the humeral vein branched and proximal hu- awaiting resolution. Furthermore, of the genera as- meral trace recurrent, crossveins in the radial space signed to the Osmylitidae, only Kirgisellodes [with K. are arranged mostly in 2Ð3 gradate series, branches of ornatus(Martynov 1925), the single species in the ge- Rs are nearly parallel to the hind margin of the wing, nus described from the Late Jurassic Karatau and and prominent trichosors are present. All of these possibly belonging to Polystoechotidae], resembles characters occur in some fossil genera, e.g., Propsy- Palaeopsychops in having a somewhat similar vena- chopsis Kru¨ ger, 1923 (Eocene Baltic amber), Baiso- tional pattern (see Carpenter 1992: Fig. 192.3). Palaeo- psychops Makarkin, 1997 (Early Cretaceous of Sibe- psychops may however be easily distinguished from K. ria). However, most other genera from the Late ornatus, in which the subcostal veinlets are usually Triassic to Late Oligocene (see partial list in Oswald simple, there is an absence of gradate series, and the 1993b) referred to this family do not possess most of branches of M2 and CuA are muchshorter. these traits, in particular they have the costal space The family Solenoptilidae, erected for Solenoptilon expanded basally, narrowing toward the wing apex, kochi (Geinitz, 1887), is represented only by the apical the body of the wing is Þlled with numerous cross- portion of a wing from the Early Jurassic of Germany veins, and the branches of the Rs are dichotomously (Handlirsch 1906Ð1908). Subsequently, three other branched (e.g., Embaneura G. Zalessky, 1953, and species were included in the family (Martynova 1949, Grammapsychops Martynova, 1954: Carpenter 1992: Bode 1953), although Makarkin (1998) objected to Figs. 195.1, 195.2). Recently Andersen (2001) treated this placement. This latter author described the mo- Osmylopsychopidae, Brongniartiellidae, and even notypic genus Oligogetes Makarkin from the Late Eo- Kalligrammatidae only as groups within “the Psychop- cene/Early Oligocene of the Russian Far East, based sidae lineage.” If the family is broadened to such an again on apical portions of wings and placed it tenta- undeÞned status, then this allows assignment to Psy- tively in this family. Data on the Solenoptilidae is very chopsidae of genera very unlike modern psychopsids. fragmentary. The only precise knowledge on this This family is strongly in need of reevaluation and group is the structure of the apical portion of wing revision of bothextinct and extant taxa. Palaeopsy- (i.e., the costal space strongly narrow [narrowest in chops differs from these insects even when this family the order among middle and large-sized neuropter- is considered in the broadest sense (sensu lato)inthat ans], Sc and R1 apically not fused, and the crossveins M2 is not pectinately branched and 1A is multi- of radial space widely spaced). All these characters are branched and occupies more area, and so it cannot obviously do not present in Palaeopsychops. reasonably be considered a psychopsid. (3) Those families that are possibilities, at least The family Brongniartiellidae is comprised of sev- theoretically, for Palaeopsychops. There are two su- eral Mesozoic genera (Martynova 1949, 1962; Whalley perfamilies to which Palaeopsychops may be possibly 1988) and is only a theoretical possibility for Palaeo- assigned: Psychopsidoidea or psychopsid-like neurop- psychops. The genus Brongniartiella Meunier, 1897 is terans (Prohemerobiidae, Brongniartiellidae, Psy- poorly known because its type species (Ricania gigas chopsidae, Osmylopsychopidae, and Kalligrammati- Weyenbergh, 1869 [ ϭ Brongniartiella problematica dae: Martynova, 1949) and Ithonoidea Meunier, 1897] from the Late Jurassic of Germany) (Polystoechotidae, Ithonidae, and Rapismatidae: needs to be redescribed. The works in which it was Tauber & Adams, 1990). The family Kalligrammatidae illustrated were published Ͼ100 yr ago (Weyenbergh is excluded, as it has markedly different venation. 1869, Meunier 1897). The photograph in Meunier Prohemerobiidae is the furthest theoretical possi- (1897) shows that the specimen is apparently a pair of bility for Palaeopsychops in this group. The Prohe- large, broadly subtriangular overlapping wings, with merobiidae are poorly deÞned. A variety of genera venation similar to that of such genera as Mesopsy- from the Late Triassic to Late Cretaceous have been chopsis Handlirsch, 1906, Embaneura, Grammapsy- referred to this family, making it to a considerable chops, Kagapsychops Fujiyama, 1978, i.e., the genera extent a grab bag taxon. Until a comprehensive revi- very distant venationally from Palaeopsychops (see sion is done (which is strongly necessary), we propose Carpenter 1992: Figs. 195.1, 195.2, 196.5). The genera restricting this family to only the type genus, as in Epactinophlebia Martynov, 1927 and Actinophlebia Carpenter (1992). Approximately 30 species from Handlirsch, 1906 have been referred to Brongniartiel- Early Jurassic of Germany and England have been lidae by some authors (e.g., Carpenter 1992). They 178 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 3 resemble Palaeopsychops in having a rather similarly of Mesopolystoechotidae and Polystoechotidae, and shaped forewing, a basal dilation of the costal space, consider them together here. The venation of extant forking of the subcostal veinlets, M is forked into M1 species is described by Carpenter (1940), Lambkin and M2 parallel to each other, and CuA has long (1988), and Oswald (1998). This completely conforms pectinate branches; they are otherwise unlike Palaeo- withthevenation of Palaeopsychops, as is evident in psychops, e.g., M2 is not pectinately branched, CuP is these fossils. Also, when the forewing venation of parallel to the hind margin of the wing and pectinately Polystoechotes punctatus Fabricius is examined with branched, and crossveins are almost entirely absent. special reference to concavity and convexity relation-

The family Osmylopsychopidae was originally es- ships, it is evident that M2 is entirely (except most tablished for the genus Osmylopsychops Tillyard, 1923. basally) strongly convex and CuA is strongly convex in The type species (O. spillerae Tillyard, 1923 from the the proximal half, i.e., very similar to Palaeopsychops Late Triassic of Australia) was at Þrst known from two dodgeorum. wing fragments. Further, fragmentary occurrences of However, there is difÞculty in the determination of O. spillerae were published by Riek (1955) and Lam- family for the fossil genera assigned to this family, as bkin (1992). Lambkin reexamined them in greater there are no autapomorphic characters of the vena- detail, and published a precise new reconstruction of tion, and so assignment to this family cannot be clearly the forewing venation, signiÞcantly revising that of determined by wings alone without some doubt. The Tillyard (1923). Later, Riek (1956) and Ponomarenko basal dilation of the costal space, the presence of a (1986, 1995) assigned several genera from the Late recurrent proximal humeral trace and a branched hu- Triassic of Australia and the Jurassic of Germany and meral vein, and a strongly pectinately branched CuA England to this family, including some that had been are possibly found throughout Polystoechotidae (dis- previously placed in Brongniartiellidae (e.g., Pterino- tinguishing them from Permian neuropterans), how- blattina Scudder, 1885, Actinophlebia Handlirsch, ever, the family shares these with many other families 1906). Osmylopsychops spillerae superÞcially resem- (e.g., the second character is present also in Hemero- bles Palaeopsychops dodgeorum in that it has similar biidae, Mesithonidae, some Mesozoic Berothidae, wing shape, heavy basal dilation of the costal space, primitive Mantispidae, Psychopsidae, Prohemerobi- CuA and CuP are arranged similarly, but in the former idae, Brongniartiellidae, and Ithonidae), and thus the origin of Sc is shifted distad, M is well-developed these characters are not diagnostic. Most other fea- with several dichotomous branches occupying a larger tures characteristic of the family are plesiomorphic, area, 1A and 2A are well-developed and multi- e.g., the presence of nygmata, the presence of prom- branched, and no gradate series is present. Osmylo- inent trichosors, and Sc and R1 are apically fused (in psychopidae is the most likely family designation for Platystoechotes these are free). Therefore assignment Palaeopsychops within the Psychopsoidea, but the of Palaeopsychops to this family is formally prelimi- characters mentioned above make such possibilities nary, however, taken together as a whole, those fea- signiÞcantly problematic. tures shown to be present suggest Polystoechotidae as Since 1923, when Rapismatidae was named, the most likely family designation. Ithonidae and Rapismatidae have been considered Thus, although there is a theoretical possibility of separate families, but recently Penny (1996) synony- assigning the genus to Psychopsidoidea, the possibility mized them, describing the genus Adamsiana, which is greater for Ithonoidea, and greater still that it be- has a mixture of character states found in both families. longs to Polystoechotidae. In the recent fauna there are Ϸ40 known species, which are distributed in Australia, mountainous areas of the Oriental Region (Rapisma McLachlan 1866), Acknowledgments and in the western hemisphere from Central America We thank Kenneth and Ken Dodge, who have provided to southern North America (Carpenter 1951, Riek such pleasurable companionship in fossil collecting through 1974, Barnard 1981, Penny 1996). Fossil occurrences the years; Rolf Mathewes, who granted access to the Simon are unknown, but Riek (1974) believed that Rapisma Fraser University collection and for his support over many is the only living representative of the family Brong- years; Guy Rose, owner of the site, for continued support of niartiellidae. Ithonidae (Rapismatidae) share all basic scientiÞc researchat Quilchena;Stig Andersen, Geological characters of the forewing venation with Polysto- Museum, Copenhagen, for providing us with an advance echotidae. However, in this family M is not pecti- copy of his paper in press; Bushra Hussaini and David 2 Grimaldi, American Museum of Natural History, for loan of nately branched (except for Oliarces Banks, 1908, in the type of Polystoechotes piperatus; and Stephen Gaimari and which this vein has a few pectinate branches), a gra- an anonymous reviewer for helpful comments that improved date series of crossveins is never present, and R1 and this paper. Funding (S.B.A.) was provided in part by a Nat- Rs are usually widely spaced apically. Thus, although ural Science and Engineering Research Council scholarship there is a theoretical possibility that Palaeopsychops and a Putnam Expeditionary Grant. belongs to Ithonidae, this seems less likely than that it belongs to Polystoechotidae. Polystoechotidae (Mesopolystoechotidae). See In- References Cited troduction for the family composition and its geo- Andersen, S. 2001. 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