A new genus and species of scorpionfly (Mecoptera) from Baltic amber, with an unusually developed postnotal organ

Systematic Entomology (2012), 37, 223–228 DOI: 10.1111/j.1365-3113.2011.00602.x

A new genus and species of scorpionfly (Mecoptera)from Baltic amber, with an unusually developedpostnotal organ

W I E S Ł A W K R Z E M I N S K I1 and A G N I E S Z K A S O S Z Y N S K A - M A J2

1Institute of Systematic and Evolution of Animals, Polish Academy of Sciences, Krakow, Poland and 2Department of InvertebrateZoology and Hydrobiology, University of Łodz, Łodz, Poland

Abstract. A new genus and species of fossil scorpionflies (Mecoptera) Baltipanorpadamzeni gen. et sp.n. is described from two well-preserved male specimens in Balticamber (middle Eocene: Lutetian). The most characteristic feature of the new taxonis an unusually developed postnotal organ on abdominal tergum IV. This is the mostextremely developed example of this organ among Mecoptera and the only observationof notal and postnotal organs among fossil scorpionflies. The following combination ofcharacters are provided to distinguish the new genus from other Panorpidae: Sc, short;R1 and R2 two-branched; A1 joins posterior margin of wing only at same level as forkof vein Rs; unusual shape of abdomen, abdominal segments I–IV strongly reduced,abdominal segment V elongate and widened, segments VII and VIII strongly elongate;notal and postnotal organs present, strongly developed process (postnotal organ) ontergum IV, unknown in all described extant and fossil scorpionflies. Different typesof notal organs of Mecoptera are compared and their function and morphology arediscussed. Morphological analysis of notal and postnotal organs in extant speciespermits us to conclude that B. damzeni sp.n. is characterized by the most developedand complex notal organs in all Mecoptera.

Introduction

Mecoptera, a small, holometabolous order of insects, com-prises about 600 extant species grouped in nine families(Cai et al., 2008; Bicha, 2010), which occupy all regionsof the world except Antarctica. The morphology and scav-enging habits of both adults and larvae are primitive forHolometabola (Grimaldi & Engel, 2005). The largest fam-ily is Panorpidae with approximately 420 species, assignedto four genera: Panorpa Linnaeus; Neopanorpa Weele; Lep-topanorpa MacLachlan; and Sinopanorpa Cai & Hua (Caiet al., 2008).

Mecoptera had a great share in faunas from the LatePermian through the Cretaceous, and constituted a significant

Correspondence: Agnieszka Soszynska-Maj, Department of Inver-tebrate Zoology and Hydrobiology, University of Łodz, ul. Banacha12/16, 90-237 Łodz, Poland. E-mail: [email protected]

element of these ecosystems, being at least three times morewidespread and diverse during the Mesozoic than at present(Krzeminski & Krzeminska, 1996, 2003; Hoell et al., 1998;Novokshonov, 2002). The level of extinction within thisgroup is unusual among holometabolous insects, althoughit is somewhat mirrored among the Neuropterida (Willman,1989; Grimaldi & Engel, 2005). Extant scorpionflies aremostly carnivorous, but examination of fossil species indicatesthat extinct Mecoptera were possibly also important plantpollinators (Ren et al., 2009).

Baltic amber dates from the Lower Eocene–Ypresian(50 Ma) to Lutetian (40 Ma). Until now representatives ofthree families were found in Baltic amber (Weitschat &Wichard, 2002): Bittacidae (five species; Krzeminski, 2007);Panorpidae (two species); and Panorpodidae (two species;Carpenter, 1954, 1955, 1976). The only known species ofPanorpidae in Baltic amber belong to the widely distributedand abundant genus Panorpa: Panorpa obsoleta Carpenter,1954 and Panorpa mortua Carpenter, 1954 (Carpenter, 1954).

© 2011 The AuthorsSystematic Entomology © 2011 The Royal Entomological Society 223

224 W. Krzeminski and A. Soszynska-Maj

Most Mecoptera (especially Panorpidae) are characterizedby interesting sexual and mating behaviour, including nuptialfeeding, pheromone emission or forced copulation (Thornhill,1973, 1980, 1981, 1990; Thornhill & Sauer, 1991; Kocket al., 2009). The structures important for the sexual behaviourof some panorpids are processes occurring in the posteriorarea of the third and fourth abdominal tergites of the males.These used to be called, respectively, notal (= notorganus,notorgan) and postnotal organs (Crampton, 1931; Mickoleit,1971), ‘upper/lower part of the notal organ’ (Chu & Byers,1978), wing-clasping organ (Penny, 1975) and clamp-likestructure (Byers & Thornhill, 1983), or just notal organ(Thornhill, 1991). The basic role of the organ is to hold thewings of the female, thereby prolonging or forcing copulation(Byers & Thornhill, 1983; Thornhill, 1990; Thornhill & Sauer,1991; Kock et al., 2009). However, Felt (1896) proposed thatthe notal organ was a source of male pheromone licked upby females (after Crampton, 1931). This was later rejectedby Mickoleit (1971) and Thornhill (1973), who found thatthe pheromone gland is positioned in the genital bulb. Theprocess of the third tergum is sclerotized, projects backwardsand its length differs in individual species from half ofone tergite to even several tergites of the abdomen. Thesecond process located on the fourth tergum is always small,more sclerotized than the former process, conical or hookshaped (Byers & Thornhill, 1983; Thornhill, 1990). Thisorgan was found in four families of Mecoptera: Eomeropidae(= Nothiothaumidae), Panorpidae, to a limited extent in thePanorpodidae (strongly reduced) and in a few European speciesof Boreidae (a rudimentary process, only on third tergite)(Mickoleit, 1971; Penny, 1975). The greatest development ofthese structures is observed in the ‘living fossil’ scorpionflyNotiothauma reedi MacLachlan (Eomeropidae) and in thefamily Panorpidae, with the exception of Sinopanorpa, wherethis organ is not developed (Mickoleit, 1971; Cai et al.,2008).

The first discovery of exceptionally well-developed notaland postnotal structures in fossil Mecoptera is presented here,along with the description of a new monotypic genus andspecies of Panorpidae from Baltic amber.

Material and methods

The study was based on two inclusions in Baltic amber (datedto the middle Eocene, ∼45 Ma, Lutetian) deposited in themuseum collection of the Institute of Systematic and Evolutionof Animals, Krakow, Poland (MP ISEA). The specimens werestudied with the use of a Leica MZFLII stereomicroscope,under reflected and transmitted light. Photographs were madewith a Leica DFC295 camera attached to the microscope.Drawings were made from the photographs. The terminologyof wing venation, the notal organ and genitalia follows thatof Chau & Byers (1978), Crampton (1931) and Webb et al.,(1975), respectively.

Systematic paleontology

Order Mecoptera Hyatt & Arms

Family Panorpidae Stephens

Genus Baltipanorpa gen.n.

Type species. Baltipanorpa damzeni gen. et sp.n.

Diagnosis. Both wings about as long as abdomen. ForewingSc short, not extending beyond one-half wing length; R1

forked before pterostigmal band to R1a and R1b; R2 two-branched; A1 joins posterior margin of wing only at samelevel as fork of vein Rs; three crossveins between A1

and A2 present. Abdominal segments I–IV strongly reduced;terga III and IV with long, narrow processes, extending totwo-thirds the length of segment VII, both set together andforming a functional entity; process on tergum IV curvedupwards at apex; abdominal segment V elongate and widened;segments VII and VIII strongly elongate, very thin at base,in second half strongly widening and sinusoidally curved;tergum VII in dorsal part produced in wide lobe; segment VIIIwidened to a goblet shape, without additional lobe.

Etymology. Combination of Balti- for the occurrence of thenew genus in Baltic amber and Panorpa, type genus of thefamily Panorpidae. The name is feminine.

Baltipanorpa damzeni gen. et sp.n.(Figs 1, 2)

Material examined. Holotype: male, no. MP/3076; wellpreserved and complete, except for the wings, which arecrumpled and the venation is partially poorly visible. Paratype:male, no. MP/1/4/552/04; specimen without head, remainderwell preserved, wings complete with clearly visible venation,except for the distal part. Both specimens in Baltic amber arehoused in the ISEA, Krakow, Poland.

Description. (Figs 1, 2). Male. Body length ∼16 mm.Head: eyes round, separated by width of rostrum, bare,

facets of equal size, three ocelli present; antennae filiformcomposed of 40 segments equal in length, covered by short,thick setae; first flagellomere as long as next four combined,last flagellomere rounded; scape wide and short, pedicel thickerand rounded; rostrum long, narrow, typical for Panorpidae;palpi not visible.

Wing venation: general model of wing venation similar tovenation of Panorpa; forewings (Fig. 3) ∼15 mm long, as longas abdomen, membrane slightly infuscate, with clearly visibledarker apical and pterostigmal bands; Sc short, not extendingbeyond one-half wing length; crossvein sc-r at four times itslength before tip of Sc; R1 forked before pterostigmal band intoR1a and R1b; one crossvein between R1 and R2 and secondcrossvein between R1 and R2–3; R2 two-branched; A1 joins

© 2011 The AuthorsSystematic Entomology © 2011 The Royal Entomological Society, Systematic Entomology, 37, 223–228

Baltipanorpa damzeni gen. et sp.n. 225

Fig. 1. Baltipanorpa damzeni gen. et sp.n. – holotype.

Fig. 2. Baltipanorpa damzeni gen. et sp.n. – abdomen of paratype.

posterior margin of wing only at same level as fork of veinRs; three crossveins between A1 and A2 (Fig. 4A, the completereconstruction of wing venation is impossible because only thebasal part of the forewings and cubital and anal parts of thehindwings are well preserved; Fig. 4B).

Legs: long, tibia with two spurs, basitarsus a little shorterthan remaining tarsomeres, second tarsomere almost 1.5× as

Fig. 3. Distal part of forewing of paratype of Baltipanorpadamzeni gen. et sp.n.

Fig. 4. Anal parts of wings of holotype of Baltipanorpadamzeni gen. et sp.n.: A, forewing; B, hindwing.

long as succeeding tarsomeres combined, a row of spines atdistal part of all tarsomeres; two terminal, pretarsal clawsvisible.

Abdomen (Fig. 5): abdominal segments I–IV narrow; pro-cess on segment III (notal organ) strongly widened at base,projecting backwards, distal half with numerous short spineson inner surface and four pairs of thin, long setae; firstseta longest, following pairs gradually shorter, last pair onlyinsignificantly longer than width of tergite III process; seg-ment IV with long, thin and bare process (postnotal organ);both processes almost equal in length, tergum IV process a lit-tle longer and curved upwards at apex, abdominal segment Velongate and widened, corresponding sternite almost one-thirdshorter than tergite V; sternites and tergites of remaining seg-ments fused; segment VI narrow in basal part, much wideneddistally, dorsal part elongated in wide lobe; part of segment VIinside segment V in holotype, clearly visible in paratype; seg-ments VII and VIII strongly elongate, considerably thinner atbase, in distal half strongly widened and sinusoidally curved;segment VII in dorsal part elongate in wide lobe, almost asin previous segment; segment VIII widened in goblet shape,without additional lobe; genital bulb elongate, very narrow atbase, which looks like pedicel; gonostylus in male genitaliashort and wide; inner structures of male genitalia not visible.

Diagnosis. The same as for the genus.



Fig. 5. Abdomen, lateral view, of Baltipanorpa damzeni gen. et sp.n.

Etymology. The specific name is derived from JonasDamzen, who found both inclusions and recognized theirimportance.

Discussion

Baltipanorpa gen.n. differs significantly from all known extantand fossil Mecoptera. The following combination of charac-ters enable us to distinguish the genus from all other describedgenera of Panorpidae: differences in wing venation in anal andradial part (Figs 3, 4); distinctive shape of abdomen, abdominalsegments I–IV strongly reduced, abdominal segment V elon-gate and widened, segments VII and VIII strongly elongate(Fig. 5); strong development of postnotal organ on tergum IV(which is even longer than notal organ on tergum III), unlikeanything observed in any previously described scorpionflies.

Characteristic differences in wing venation are present inanal and distal parts. Short Sc in forewings, not extendingbeyond one-half wing length, is similar to Panorpa alpina(Rambur), and is in opposition to other Panorpa species andother panorpid genera. Baltipanorpa gen.n. has two-branchedR1, whereas in several previously described genera R1 is notforked. R2 is two-branched, as in Panorpa, rather than three-branched, as in Sinopanorpa (Cai et al., 2008; Tillier, 2008).A1 joins posterior margin of wing only at same level as fork ofvein Rs, as in Neopanorpa, in a different way to Sinopanorpaand Panorpa. There are three crossveins between A1 and A2

in Baltipanorpa gen.n., whereas in existing genera only one(Neopanorpa) or two crossviens (Sinopanorpa and Panorpa)are described (Cai et al., 2008).

In extant Mecoptera the posterior median process of the thirdtergum is sclerotized, broad and flat in Panorpa, or elongateand rod-shaped in Neopanorpa, and projects backwards (Issiki,1933). Crampton (1931) proposed that this structure elongatedduring the evolution of the Mecoptera. He suggested thatthis is an orthogenetic tendency that probably started fromthe most primitive and archaic scorpionfly, Notiothauma reedi

MacLachlan, where the notal organ is a pincer-like structure. Itwould reach its culmination in Panorpa takenouchii Miyake,which has the longest notal organ among extant species.Panorpid genera, as well as species, differ in the length ofthe notal organ (Fig. S1). In European species of Panorpa thenotal organs are slightly produced and barely visible or evencompletely reduced in Panorpa cognata Rambur (Engqvist &Sauer, 2002). However, a few Panorpa species from easternAsia have very long processes, reaching almost the endof abdomen Panorpa leucoptera (Uhler) and P. takenouchii(Issiki). Leptopanorpa is characterized by a well-developedbut small, clamp-like structure (Lieftinck, 1936), and theorgan is not developed in Sinopanorpa (Cai et al., 2008). Ingeneral, the longest notal organs in Mecoptera are found inNeopanorpa spp., which was an important reason to separatethem from Panorpa (Weele, 1909), although the monophylyof these groups relative to each other requires rigorous testing.Neopanorpa species from the Oriental region are characterizedby short notal organs, reaching to only one half of the fourthabdominal segment, e.g. in Neopanorpa byersi Webb & Penny(Webb & Penny, 1979), whereas the longest reaching almostto half of the seventh segment are observed in Neopanorpasubreticulata Miyamoto & Makihara (Miyamoto & Makihara,1979; Fig. S1). In the latter it is a bit shorter than inB. damzeni sp.n.

In extant scorpionflies the second process (postnotal organ)located on the fourth tergum is always very small, moresclerotized than the notal organ, conical or hook shaped andprojects upwards. In species with very elongate processes ofthe third tergum, the postnotal process is usually flat, barelyvisible and reduced or only present as a small elevation witha dense set of setae, as in P. leucoptera and P. takenouchii(Issiki, 1933). No examples of any elongation of the processon the fourth tergite appear in extant Mecoptera. The elongateprocess on abdominal tergite IV, reaching two-thirds oftergum VII, is only known from B. damzeni sp.n., and is anunusually developed postnotal organ among Mecoptera.

As mentioned earlier, the function of notal and postnotalorgans is generally related to copulation and mating behaviour.This was described in detail for Panorpa spp. (Mickoleit,1971; Thornhill, 1973, 1980, 1981, 1990; Thornhill & Sauer,1991). During mating in the European species Panorpacommunis Linnaeus, the male and female forms a V-shapedposition. The male first grasps the female’s leg or wing withhis genital claspers, and then repositions her to catch thecosta of the hindwing with his notal organ (Mickoleit, 1971;Byers & Thornhill, 1983). It was proved that the function ofnotal and postnotal organs is to enforce (without nuptial giftor salivary mass) and prolong copulation (with nuptial gift orsalivary mass) to transfer more sperm, minimize the costs ofsalivary mass production and avoid interruption of copulationby an intruder. Between the third and fourth segments thereare muscles bringing the two processes together (Thornhill &Sauer, 1991). Kock et al. (2009) proposed that the functionof the notal organ is to initiate and stabilize females duringmating in the V-shaped position. When the clamp-like structureis reduced (as in P. cognata) the mating strategy is different.


Baltipanorpa damzeni gen. et sp.n. 227

The pre-mating period is prolonged to ensure that the femaleis motivated to mate, the male looks for a sheltered locationand only one large salivary mass is produced to occupy thefemale and prolong copulation (Engqvist & Sauer, 2002).

It is easy to imagine how males of species with smallnotal organs, like P. communis or N. byersi catch the female’sforewing costa using the notal organ as a pincer. It is moredifficult to understand how such elongated notal organs inMecoptera (as in Neopanorpa species or in P. takenouchii )operate if the process on the fourth tergite is usually barelydeveloped. Issiki (1933) stated that the male can holdthe wings of females between the elongate notal processand fourth tergite. The only description of other speciesusing the elongated organ comes from Thornhill (personalcommunication), who observed flying males of species withvery elongated notal organs grabbing stationary females beforemating. In B. damzeni sp.n. the situation is more complicatedbecause not only is the process of the third tergite long,but also the process on the fourth abdominal segment isstrongly elongated (even a bit longer than the process ontergite III), and is extended to about two-thirds of abdominaltergum VII. Males of B. damzeni sp.n. could hold the wholesurface of the female’s wing with four pairs of long seta onthe notal process, thereby immobilizing the female’s wing.Taking into consideration Thornhill’s observations (personalcommunication), we proposed another hypothesis. Males mightuse the strongly elongated notal and postnotal organ to interrupta female mating with another male, or even to abduct her.

Some functional similarity of a plier-like organ ofB. damzeni sp.n. may be observed with structures on the tho-rax in Boreidae. Their metamorphosed wings form four narrow,hooked straps. Males of Boreus spp. hold the wingless femalelegs on their backs during copulation (Byers & Thornhill,1983). Even though these structures have evolved on differ-ent body parts, their morphological similarity is considerable.However, the absence of such developed notal structures inany living scorpionfly makes any interpretation of their usevery speculative.

The shape of the abdomen of B. damzeni sp.n. differsfrom other described Panorpidae genera. The abdominalsegment V of Baltipanorpa gen.n. is the longest amongPanorpidae. The abdominal segments VI–VIII are elongated,similar to Panorpa species with a long notal organ, andNeopanorpa species, however not so extremely long as inLeptopanorpa spp. (Lieftinck, 1936). Additionally, VI–VIIIsegments are markedly thinner at the base what is commonto species of Sinopanorpa spp. (Cai et al. 2008). Accordingto the unusual development of notal and postnotal organs,a change in general plan of the abdomen was necessary.The most elongated notal organs are positioned on a smallpostnotal process, and lie on the abdomen in extant Mecoptera.In the case of an elongation of the postnotal organ inB. damzeni sp.n., the processes do not lie on the abdomenbut protrude from the body, forming an angle of about 45◦.The terminal segments of the abdomen are thinner at thebase and elongated, as in Sinopanorpa (Cai et al., 2008), butare recurved upwards, as in all Panorpidae. However, the

long and massive segment V is unique among Panorpidae.It is interesting if the elongated postnotal organ induced thepeculiar shape of the abdomen. One hypothesis may be thatmales could use the massive segment V to support the femaleduring copulation. On the other hand, in the extant Panorpidaeanother side-by-side V-shaped mating position was observed(Mickoleit, 1971; Byers & Thornhill, 1983). Nevertheless, suchsupport would be necessary if the females of Baltipanorpawere wingless or if their wings were reduced.

Penny (1975) hypothesized that the notal organ was probablyacquired by a Mesozoic mecopteran ancestor some 200 millionyears ago. Willmann (1987, 1989) suggested an earlieracquirement of this structure and monophyly of Mecoptera.However, Whiting et al. (1997) and Whiting (2002) proposedthe paraphyly of this order based on molecular studies. Mostrecently, Grimaldi & Engel (2005) confirmed Mecoptera tobe paraphyletic. From the topology of the phylogenetic treethey proposed and taking into consideration the presence ofa rudimentary notal organ in Boreidae, we can assume thatthe notal organ was acquired by a mecopteroid ancestor in theEarly Triassic, more than 240 million years ago.

Supporting Information

Additional Supporting Information may be found in the onlineversion of this article under the DOI reference:10.1111/j.1365-3113.2011.00602.x

Figure S1. The examples of notal and postnotal organs inextant Mecoptera.

Please note: Neither the Editors nor Wiley-Blackwellare responsible for the content or functionality of anysupporting materials supplied by the authors. Any queries(other than missing material) should be directed to thecorresponding author for the article.

Acknowledgements

We would like to thank to Michał Grabowski (University ofŁodz) and Ewa Krzeminska (MP ISEA, Krakow) for theirsuggestions and help in preparing the earlier versions of thearticle. Special acknowledgments are due to Lars Vilhelmsenfor his very useful comments on the submitted article, and tothe anonymous reviewer for an extensive and valuable review.

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Accepted 1 August 2011First published online 28 September 2011


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A new genus and species of scorpionfly (Mecoptera) from Baltic amber, with an unusually developed postnotal organ