ZooKeys | 30: 9 [I 02 (20 | | ) A peer-reviewed open-access journal doi: 10.3897/zookeys. 130.1422 $Z00Ke y S www.zookeys.o rg Launched to accelerate biodiversity research The smallest Neoptera (Baryshnyalidae fam. n.) from Hagen-Vorhalle (early Late Carboniferous: Namurian B; Germany) Jan-Michael Ilger?, Carsten Brauckmann? Clausthal University of Technology, Institute of Geology and Paleontology, Leibnizstrafse 10, 38678 Clausthal- Zellerfeld, Germany Tt urn:lsid:zoobank. org:author:673444 1 F-1298-4D00-A4A8-C0B33D388E54 * urn:lsid:zoobank. org:author:AIB536B4-6DEF-48C4-980A-9EB8A9467F8B Corresponding author: /an-Michael Ilger (ilger@gmx.de) Academic editor: D. Shcherbakov | Received 20 April 2011 | Accepted 2 August 2011 | Published 24 September 2011 urn:lsid:zoobank.org:pub:F3B128 LA-2E50-4F36-ACAD-39917E39D4F9 Citation: Ilger JM, Brauckmann C (2011) The smallest Neoptera (Baryshnyalidae fam. n.) from Hagen-Vorhalle (early Late Carboniferous: Namurian B; Germany). In: Shcherbakov DE, Engel MS, Sharkey MJ (Eds) Advances in the Systematics of Fossil and Modern Insects: Honouring Alexandr Rasnitsyn. ZooKeys 130: 91-102. doi: 10.3897/ zookeys. 130.1422 Abstract With Baryshnyala occulta gen. et sp. n. (Baryshnyalidae fam. n.) we report a new small species from early Late Carboniferous (Pennsylvanian) deposits in Hagen-Vorhalle. It differs in its unique venation pattern and small size from all other Neoptera known from this Lagerstatte and other contemporaneous loca- tions worldwide. With an estimated wing length of <10 mm it is by far the smallest species of Neoptera from Hagen-Vorhalle and is less than half as long as Heterologopsis ruhrensis Brauckmann & Koch, 1982 (-25 mm). The specimen shows some relations to the earliest Holometabola and may date back the first appearance of holometaboly to the Namurian B (early Bashkirian: Marsdenian). The new species increases the paleo-biodiversity and span of inter-specific variability within the early Neoptera. It shows that very small and tiny specimens and species can easily be overlooked. Keywords Hagen-Vorhalle, Neoptera, Baryshnyala occulta gen. et sp. n., Namurian B, holometaboly Copyright Jan-Michael Ilger, Carsten Brauckmann. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 92 Jan-Michael Ilger & Carsten Brauckmann / ZooKeys 130: 91-102 (2011) Introduction Hagen-Vorhalle is one of the most important Konservat-Lagerstatten of the early Late Carboniferous (Early Pennsylvanian) and has provided remarkable data of supra-re- gional importance about the evolution of the most ancient Pterygota. An extraordinary insect fauna was discovered in 1982 and reported in a brief article by Brauckmann and Koch (1982). Over 16,000 samples were recovered in an exca- vation campaign in 1990-1997 and by private collectors. Insects are represented by ~310 individuals (mostly in dorsal view with part and counterpart). Hendricks (2005) published a complete list of determinable taxa whereas Brauckmann et al. (2010) pre- sented an actual outline of the insect fauna and their relation to other locations. Most of the material is very well preserved, including basal Neoptera Martynovy, 1923 with ~210 specimens in 5 species: Kemperala hagenensis Brauckmann, 1984, Holasicia ras- nitsyni Brauckmann, 1984, Kochopteron hoffmannorum Brauckmann, 1984, Heterolo- gopsis ruhrensis Brauckmann & Koch, 1982, and Baryshnyala occulta gen. et sp. n. Kemperala Brauckmann, 1984 and Holasicia Kukalova, 1958 can be assigned to the family Paoliidae Handlirsch 1906. For diagnosis and included genera see Kukalova (1958), Brauckmann (1984), Brauckmann et al. (1985, 2003, 2010), Prokop and Nel (2007), Ilger and Brauckmann (2008), and Prokop et al. (in press). As shown in the historical compilation by Prokop and Nel (2007), the systematic rank of paoliids and related early Pterygota has been the subject of controversy during the last few dec- ades: Carpenter (1992), for example, traditionally considered them as a family within the “Protorthoptera,” a clearly polyphyletic “order”. Kukalova-Peck and Brauckmann (1992) assigned the “paoliid line” together with other early Neoptera to the “hemipter- oid stem assemblage”. Grimaldi and Engel (2005) placed them as Paoliidae directly in Neoptera without indication of any order. On the other hand, Rohdendorf and Ras- nitsyn (1980) and subsequently Rasnitsyn and Quicke (2002) ranked them as Paoliida on ordinal level. Prokop and Nel (2007) did the same, and additionally introduced their Katerinkidae, a closely related new family of the same order. A more general critical discussion of different views of hexapod phylogeny including the Palaeoptera/ early Neoptera problem has been provided by Klass (2007, 2009). In the present con- tribution we avoid a precise higher systematic assignment, because we do not want to anticipate the final results of the current phylogenetic analyses. The systematic position of the Kochopteron Brauckmann, 1984 and Heterologopsis Brauckmann & Koch, 1982—which are morphologically similar to the Paoliidae—is unclear and still under controversial discussion. Brauckmann et al. (2003) assigned them to the family ?Cacurgidae Handlirsch, 1911. In contrast, Béthoux and Nel (2002) placed Heterologopsis in the Archaeorthoptera. Baryshnyala occulta gen. et sp. n. cannot be allocated to any of these groups and is a representative of a new family within the early Neoptera (Baryshnyalidae fam. n.) The smallest Neoptera (Baryshnyalidae fam. n.) from Hagen-Vorhalle (early Late Carboniferous... 93 Methods The specimen WME.N P.21299 is stored in the Hagen-Vorhalle collection of the LWL- Museum fiir Naturkunde, Westfalisches Landesmuseum mit Planetarium in Minster. It was recovered in an excavation campaign in 1990-1991. Investigations were done using a Euromex® ZT-45 zoom trinocular with an at- tached camera tube. Objects were illuminated with a Euromex® EK-1 cold light lamp with polarizing filters to minimize reflections on clay minerals and mica surfaces. Im- aging conditions for photo documentation were optimized by varying illumination levels and angles with polarized and non-polarized light. Photographs were taken by using a Canon PowerShot A470 digital point-and-shoot camera with Super Macro function. Digital images were edited using the computer programs IMAGEJ, COREL PHOTO-PAINT and GIMP. The nomenclature of wing venation follows Kukalova-Peck and Willmann (1990) as well as Kukalova-Peck and Brauckmann (1992). Main vein terminology corresponds to this scheme: ScP— = Subcosta posterior, RA+ = Radius anterior, RP— = Radius poste- rior, Ma+ = Media anterior, MP— = Media posterior, CuA+ = Cubitus anterior, CuP— = Cubitus posterior, AA+ = Analis anterior, arc = arculus (strong cross-vein between MP— and CuA+), cmf = cubito-median fold (nomenclature follows Nel et al. (2007): convex fold between MP— and CuA+). Attached plus and minus indicate the corrugation of the wing; i.e.: + = above plane of projection, — = below plane of projection. Systematics Infraclass Neoptera Martynov, 1923 Family Baryshnyalidae fam. n. urn:lsid:zoobank.org:act: CBBFCO06D-A9 15-4F58-8A9B-EXED57BA9E7 1 http://species-id.net/wiki/Baryshnyalidae Type (and only known) genus. Barys/nyala gen. n., original designation. Diagnosis. Wing small and well rounded, with the following venation pattern: (i) strong cross-vein between MP-— and CuA+ in basal part of wing (arculus), (ii) pronounced convex fold between MP— and CuA+ (cubito-median fold), (iii) CuP-— strongly convex, with 3 terminal branches, (iv) a number of straight cross-veins be- tween main veins. 94 Jan-Michael Ilger & Carsten Brauckmann / ZooKeys 130: 91-102 (2011) Baryshnyala gen. n. urn:lsid:zoobank.org:act:B3D7979B-92A 1-4ED8-B55C-A4858E47861B http://species-id.net/wiki/Baryshnyala Type (and only known) species. Baryshnyala occulta gen. et sp. n., original designation. Diagnosis. Very small (length <10 mm) and compact wing with well rounded apex. The venation pattern shows a combination of the following unique characters: (i) RP— branches far before reaching the mid-wing, (ii) well pronounced cubito-me- dian fold, (iii) CuP — very strongly convex, (iv) posterior branches of MP— and CuA+ curved backwards (strongly convex), (v) no archedictyon but a number of cross-veins mainly in distal half of the wing. Remarks. Despite the small size of the wing the regular shape precludes that it is a nymphal wing as it shows no kind of a typical strong backward flexion. Etymology. The genus name is a combination of the Russian “baryshnya“ for young, unmarried woman and Latin “ala” for wing; gender feminine. Baryshnyala occulta gen. et sp. n. urn:lsid:zoobank.org:act:A096877A-8 14A-4E30-B782-5E7 LACFOFE5C http://species-id.net/wiki/Baryshnyala_occulta Fig 1A-B, 2A—F Holotype (and only known specimen). Specimen no. WMf.N P.212999, left meta- thoracic wing, with slightly damaged apex and lacking wing base. Stored in the col- lection of the LWL-Museum ftir Naturkunde, Westfalisches Landesmuseum mit Plan- etarium in Minster (Germany). Type locality. Former brickyard quarry near Hagen-Vorhalle, North Rhine- Westphalia, Germany (topographic map 1 : 25,000 sheet no. 4610 Hagen/Westfalen; 51°22.88'N; 007°26.77'E, -115 ma.s.L). Type stratum. Early Late Carboniferous (Early Pennsylvanian): early Bashkirian, late Namurian B, late Marsdenian, ammonoid zone R2c, Ziegelschiefer Formation. Diagnosis. As for the genus (due to the temporarily monospecific status). Length >9.6 mm. Preservation. ‘The isolated wing is quite well preserved, though it lacks the proxi- mal tenth with articulation and most of the anal area. Apical wing margin is also slight- ly damaged. ‘The corrugation is easily discernible but flattened by diagenetic compres- sion. The anterior mid-wing is slightly damaged by preparation marks which obscure the exact reconstruction of terminal ScP— and RA+ branches—especially whether they do or do not fuse. Also visible in the apical area (Fig. 1), tiny prodissoconchs of young bivalves are attached to the dorsal wing membrane. These embryonic shells are common in basal Neoptera from Hagen-Vorhalle but are absent in all other Pterygota from the same locality. Most recently this has led to new interpretation of the taphonomy of the The smallest Neoptera (Baryshnyalidae fam. n.) from Hagen-Vorhalle (early Late Carboniferous... 95 Lear Figure |. Baryshnyala occulta gen. et sp. n.; holotype, isolated left metathoracic wing (WME£.N P.21299); early Late Carboniferous (Early Pennsylvanian: Namurian B, Marsdenian): A superimposed drawing with interpretation of the wing venation pattern; black arrow indicates arculus (arc) B photograph under polar- ized light; white arrows indicate attached bivalve prodissoconchs. Scale bars: 1 mm. Lagerstatte——A manuscript by Ilger is currently in review, Ilger and Brauckmann (2009) gave a short overview.— The model implies a stage of drifting along a pycno- cline in a well stratified water body. During this period the corpses were affected by free-swimming bivalve larvae. Measurements. Preserved length: 8.7 mm; estimated length: 29.6 mm; maxi- mum width (at most posterior branch of MP—): 4.2 mm. Approximated ratio length/ width: ~2.3. 96 Jan-Michael Ilger & Carsten Brauckmann / ZooKeys 130: 91-102 (2011) Figure 2. Baryshnyala occulta gen. et sp. n.; holotype, isolated left metathoracic wing (WME£.N P.21299); early Late Carboniferous (Early Pennsylvanian: Namurian B, Marsdenian): A-F The same wing in differ- ent illumination settings. Scale bar: 1 mm. Description. There is only one isolated wing known. Due to its shape and the existence of a cubito-median fold we suppose it is a metathoracic wing (Figs 1-2). Costal margin in mid-wing straight, apex well rounded. Costal area strongly pectinate with wide-standing cross-veins. ScP— simple and most likely reaching costal margin. R stem proximally thick and strong, branching in proximal quarter of wing length. RA+ simple and generally straight, terminal development not well preserved but probably reaching costal margin in distal fifth. RP— well and strongly developed, with 9 termi- nal branches. MP— strong, with 5 terminal branches, first branching clearly before mid-wing and fanning at basal third of wing length, posterior branches with strongly curved backwards at posterior wing margin. Strong arculus (= cross-vein between MP- and CuA+) present in proximal eighth of the wing. CuA+ slightly reduced, slightly concave in the middle part before branching, with 3 terminal branches, all very strongly curved backwards. Well developed and probably sclerotizied cubito-me- dian fold (between MP- and CuA+) running close to CuA+, especially in the proximal part. CuP— with 3 terminal branches, CuP,— very strongly convex. First AA+ straight. The smallest Neoptera (Baryshnyalidae fam. n.) from Hagen-Vorhalle (early Late Carboniferous... 97 A number of thin and straight cross-veins mainly in the sectors of RP— and MP- and distal of mid-wing. Etymology. Latin occultus, -a, -um (adjective) meaning arcane. The holotype specimen was stored for twenty years in the collection without being identified as a new species. Discussion Small basal Neoptera of the Late Carboniferous Baryshnyala occulta gen. et sp. n. can be easily separated from all other Neoptera from Hagen-Vorhalle by its very small size (length: >9.6 mm, width: 4.2 mm). In com- parison to other wings within this group the specimen is about half as long as Het- erologopsis ruhrensis (length: -25 mm, width: ~8 mm). Holasicia rasnitsyni (length: 36 mm, width: 13 mm) and Kochopteron hoffmannorum (length: 41 mm, width: 13 mm) are distinctly larger, and Kemperala hagenensis (length: 61 mm, width: 22 mm) is by far the largest species of basal Neoptera from Hagen-Vorhalle (Fig. 3). Baryshnyala occulta fits very well in this set of proportions: all species show a ten- dency towards a length/width ration about 3 (l/w = 2.8 in B. occulta). K. hagenensis and K. hoffmannorum seem to vary mostly in the wing length whereas the width is more consistent. In contrast, the wings of H. rasnitsyni vary more in their width and less in length. The variation in H. rubrensis and B. occulta is unknown because they are monotypic. Further small Late Carboniferous (Pennsylvanian) basal Neoptera from localities in northwestern Germany and thus close to Hagen-Vorhalle are for example (arranged by their stratigraphical age): Metropatorites kassenbergensis Keller, 1934 (ord. et fam. inc. sed.), Namurian C (Bashkirian: Yeadonian): preserved length: 9 mm, estimated total length: ~22 mm, preserved width: 5 mm, estimated maximal width: -9 mm. Micropalentomum minusculum Schmidt, 1962 (Micropalentomidae), late West- phalian A (Bashkirian: Langsettian): length: ~3,5 mm [sic!], width: ~1 mm. Eodelopterum priscum Schmidt, 1962 (Miomoptera: Archaemiopteridae), early Westphalian B (late Bashkirian: Duckmantian): length: <5 mm, width: -2.3 mm. Heterologellus teichmuellerae Schmidt, 1962 (Protophasmatidae), early Westphalian C (Moskovian: Bolsovian): length: -18 mm, width: ~7 mm. Controversala miomopteroides Brauckmann & Herd, 2005 (ord. et fam. inc. sed.), Westphalian D (late Moskovian): preserved length: 19 mm, estimated total length: ~20 mm, width: 8 mm. Worldwide there are known many other small Neoptera from the Late Carboniferous and Permian. Especially Early Permian Protomeropidae Tillyard, 1926 are very small. For example Westphalomerope maryvonneae Nel et al., 2007 from the early Langsettian (Late Carboniferous: Bashkirian, Westphalian A) of France is twice smaller than B. occulta. 98 Jan-Michael Ilger & Carsten Brauckmann / ZooKeys 130: 91-102 (2011) 25 5 Kemperala hagenensis fe ———— (n = 16) ; 20 - Holasicia eee (n =7) Pe = 15:5 as SN = AEE? = Cr Kochopteron hoffmannorum s Breast n= 13) o x = "10 = a _@ Heterologopsis ruhrensis pee (n =1) 5 “| @ 4 . _” Baryshnyala occulta or ihig=