The Female Genitalic Region in Basal Earwigs (Insecta: Dermaptera: Pygidicranidae s.l.)

Transcription

1 Entomologische Abhandlungen 61 (2): Museum für Tierkunde Dresden, ISSN , The Female Genitalic Region in Basal Earwigs (Insecta: Dermaptera: Pygidicranidae s.l.) KLAUS-DIETER KLASS Staatliche Naturhistorische Sammlungen Dresden, Museum für Tierkunde, Königsbrücker Landstrasse 159, Dresden, Germany Abstract. The exoskeleton of the female genitalia is described for members of ten genera of basal Dermaptera (paraphyletic Pygidicranidae s.l.), and the musculature for Dacnodes sp. Structural elements are homologized and characters are discussed with an all-insect framework. Pygidicranidae display a surprising structural diversity of the female genitalia. Some have almost complete ovipositors, with well developed gonapophyses and gonoplacs and sclerotizations resembling those in other ovipositor-bearing Pterygota; others show very different combinations of specialization and reduction of the genitalic elements. Most pygidicranids have gonapophyses IX, gonangula, accessory glands, and paired internal tubes arising lateral to the latter, which have all previously not been reported for Dermaptera. The long coxal lobes VIII of some pygidicranids are homonomous with the gonoplacs IX and unique within Pterygota. The structure of the spermatheca shows strong variation, which does not correlate with the number of virgae in the male. The results indicate monophyletic Diplatyinae and possibly Diplatyinae + Karschiellinae. A specific bipartition of coxa IX in Pygidicranidae, Hemiptera, and Hymenoptera could be autapomorphic for a clade Dermaptera + Acercaria + Endopterygota. Some potential genitalic synapomorphies of Acercaria and Endopterygota are additionally proposed. This study also provides the basis for using the genitalia in the taxonomic description of female Pygidicranidae, which have remained difficult to identify. Key words. Accessory glands, Dermaptera, Diplatyidae, evolution, female genitalia, Forficulina, gonangulum, gonapophyses, Karschiellidae, ovipositor, phylogeny, Pygidicranidae, spermatheca Contents 1. Introduction 1.1. The female genitalia in Insecta 1.2. Classification and female genitalia in Dermaptera 2. Material and methods 3. Terminologies, morphological interpretations, and abbreviations 3.1. Elements studied and their interpretation 3.2. Segmental assignment 3.3. Terminologies 3.4. Abbreviations 4. Description of female genitalia and postabdomen of Pygidicranidae 4.1. Echinosoma yorkense and generalities 4.2. Preview of other species 4.3. Dacnodes sp. indet. and Dacnodes caffra 4.4. Tagalina burri 4.5. Anataelia canariensis 4.6. Diplatys macrocephalus 4.7. Haplodiplatys orientalis 4.8. Karschiella buettneri 4.9. Bormansia monardi Esphalmenus basidentatus Pyragra fuscata 5. Discussion of female genitalic region 5.1. The basic design and subsets of female genitalia in Insecta Laterocoxae VIII and IX 5.3. Coxae and gonoplacs (coxal lobes) VIII and IX 5.4. Ventral elements around the segmental border IX/X 5.5. Gonapophyses and gonapophyseal sclerites VIII and IX 5.6. Gonoducts and their openings 5.7. Spermathecae and related sclerites 5.8. Accessory gland and lateral tubes 5.9. Terga VIII X The musculature of the female genitalic region 6. Implications on phylogeny and ovipositor evolution 6.1. Monophyly of Dermaptera 6.2. Relationships between Dermaptera and other Pterygota 6.3. Implications on phylogenetic relationships in Neoptera 6.4. Relationships within Dermaptera 6.5. Evolutionary aspects of the ovipositor in Pygidicranidae 7. Conclusions 8. Acknowledgements 9. References Introduction 1.1. The female genitalia in Insecta The structure of the female genitalic region in insects has always been of great interest to morphologists. The most comprehensive contribution at the all-insect level is that of SNODGRASS (1933). SCUDDER (1957, 1961a, b, 1964) gave some important revisionary notes, while further refinements were contributed by SMITH (1969). Treatments at the ordinal level are numerous, and BITSCH s (1974a) work on Archaeognatha, ROUSSET s (1973) on Zygentoma, MCKITTRICK s (1964) and KLASS (1998) on Dictyoptera, and SMITH s (1970, 1972) on Hymenoptera are among the most prominent. Through the last decades several characters were drawn from female genitalic morphology in support of some of the high-rank lineages within the Hexapoda (e.g., HENNIG 1969, 1981). Most prominent is the presence in the ground plan of Insecta (used here sensu KLASS & KRISTENSEN 2001; Ectognatha sensu HENNIG 1969) of a true ovipositor composed of limb-base elements of abdominal segments VIII and IX. Two pairs of projections are present per segment, the gonapophyses (anterior and median valves, being mo-

2 174 KLASS: Female genitalia in basal Dermaptera dified eversible vesicles) and the (gono)coxal lobes (gonoplacs = lateral valves in segment IX, sclerotized by (gono)coxae = valvifers). The gonoplacs primitively bear styli distally, and the gonapophyses VIII and IX of each side are connected by a tongue-and-groove device for a sliding interlock (olistheter). This entire structure is generally considered one of the most conspicuous (compound) autapomorphies of Insecta, though possible remnants of this pattern have also been reported for Diplura (GRASSI 1888). A variety of additional sclerites can be present in the female genitalia of the different insect subgroups, which have been called the basivalvulae, anterior and posterior intervalvulae, poststernites, spermathecal plates, (eu)sterna, and gonangulum, and which were variously interpreted as limb-base or sternal elements VIII and IX. The gonangulum, which coordinates the movements of the gonapophyses, has become important in phylogeny discussion when it was proposed to be synapomorphic for Zygentoma and Pterygota, which together constitute the Dicondylia (e.g., HENNIG 1969). After BITSCH s (1974a) finding in Archaeognatha of a precursor sclerite to the gonangulum, its larger and multi-articulated condition in Dicondylia appeared to be the autapomorphy. However, also in this modified formulation this autapomorphy has become questionable because all Odonata equipped with a plesiomorphic ovipositor lack a one-piece gonangulum (Epiophlebia superstes (Selys, 1889) is the only known exception). Instead, these Odonata as well as the examined Archaeognatha have two separate sclerites (KLASS in press). Another potential autapomorphy of the Dicondylia is the genital fold formed by the posteriormost part of the abdominal venter VII (see KLASS 1998: 90f, in press), which has been described for Zygentoma ( languette in ROUSSET 1973), Ensifera ( lamina subgenitalis in ANDER 1939), and Dictyoptera ( laterosternal shelf in MCKITTRICK 1964 and KLASS 1998). Furthermore, the modification of the coxal lobes IX (gonoplacs) into a sheath for the gonapophyses VIII and IX (KLASS in press), and the suppression of the corresponding lobes in segment VIII may appear as autapomorphies of the Pterygota. Within the Pterygota the ovipositor is still of wide occurrence, but it has become strongly simplified, reduced, or lost in many subgroups, for instance in the Ephemeroptera (e.g., BIRKET-SMITH 1971), Plecoptera (ZWICK 1973, 1980), and Embioptera (ROSS 2000), and in many anisopteran Odonata (e.g., PFAU 1991) and most Endopterygota (the Hymenoptera being the most prominent exception). While the structure of the most conspicuous components of the female genitalic region (gonapophyses, gonoplacs, valvifers) is well documented throughout the Pterygota, the many accessory components and, in part, the musculature have been strongly neglected. Therefore, except for the gonapophyses and gonoplacs the homologies of the ovipositor elements among order-level taxa are still widely unknown. A hypothesis of the homology of elements between different taxa, however, is needed before any character can be assessed in these taxa (topographic homology hypotheses as defined in KLASS 2001b, preceding primary and secondary homology hypotheses sensu DE PINNA 1991; see also BROWER & SCHAWAROCH 1996). In addition, a convincing morphological interpretation of an element is in many cases helpful in assessing the polarity of related characters. While the interpretation of the three valve pairs, on which there are seemingly endless disputes in the literature, has now probably become settled (BITSCH 1994), many other issues, such as the proper delimitation of the abdominal (eu)sterna and basal podomeres, have remained unclear. Due to these deficiencies in the description and thorough homologization and interpretation of structures, the female genitalic region in insects is still far from being fully available as a source of characters for phylogenetic reconstruction. Furthermore, in some high-rank taxa basal subgroups, which could a priori be supposed to show a comparatively plesiomorphic condition in the female genitalia, have been almost completely ignored in the previous morphological literature. An outstanding example of such insufficient treatment are the Dermaptera, and their most basal subgroup Pygidicranidae (s.l., see below) in particular Classification and female genitalia in Dermaptera The Dermaptera had long been divided into three suborders (e.g., GÜNTHER & HERTER 1974): the Forficulina comprising the many typical earwigs, and the digeneric Hemimerina (Hemimeridae: Hemimerus Walker, 1871; Araeomerus Maa, 1974) and Arixeniina (Arixeniidae: Arixenia Jordan, 1909; Xeniaria Maa, 1974) with few, highly specialized species living in close association with certain rats (Muridae) and bats (Molossidae), respectively (see NAKATA & MAA 1974). However, both Hemimeridae (KLASS 2001a) and Arixeniidae (POPHAM 1985) are more probably deeply subordinate within the Forficulina. The Pygidicranidae, if taken in the wide sense proposed by HINCKS (1955, 1959), POPHAM (1985), and STEINMANN (1986, 1989), comprise the most primitive Forficulina, including also genera such as Karschiella Verhoeff, 1902 and Diplatys Audinet-Serville, 1831, which, however, together with some related genera have both been rightfully assigned family status by some authors (Karschiellidae, Diplatyidae; POPHAM 1965: 38; HAAS 1995). Nevertheless, because the phylogenetic relationships are not yet fully resolved, Pygidicranidae is here used sensu STEINMANN (1986), comprising the Anataelinae, Challinae, Diplatyinae, Esphalmeninae, Blandicinae, Pyragrinae, Karschiellinae, Pygidicraninae, Cylindrogastrinae, Diplatymorphinae, and Echinosomatinae. With this content the Pygidicranidae is surely paraphyletic (see HAAS 1995; HAAS & KUKALOVÁ- PECK 2001). In addition to those mentioned above, the Dermaptera is another taxon to which ovipositor reduction is frequently ascribed in general statements (e.g., KRISTENSEN 1991: 134; WHITING et al. 1997: character 5; WHEELER et al. 2001: character 47). A strong reduction of the ovipositor is clearly true for the derived subgroups of the Dermaptera, including Hemimeridae and Arixeniidae: valves are, at most, represented by very short projections, and the sclerotizations of venters VIII and IX are paired plates that hardly show any further differentiation (KLASS 2001a: figs ; JORDAN 1909a, b). In many of the moderately derived Forficulina, such as the Labiduridae, valves can be distinctly longer, and the basal sclerotizations more strongly differentiated (GILES 1961; BHATNAGAR 1964), but a categorization of the ovipositor as reduced would still apply. KLASS (2001a: 269) indicated that ovipositor reduction in these taxa may be due to a paedomorphic evolution, because the female genitalic area shows attributes otherwise present in the nymphs of Dermaptera and other insects. On the other hand, it has also been known for long that some of the most basal, i.e., pygidicranid earwigs have well-developed ovipositor valves and discrete sclerites around their bases that may be (gono)coxae and gonangula. This is evident from GILES (1963) description of Echinosoma afrum (Palisot de Beauvois, 1805). By applying SNODGRASS (1933) terms, GILES gives topographic homology hypotheses for the basal sclerites in Echinosoma and other Insecta, but these are not very convincing. Unfortunately, the remaining contributions on Pygidicranidae give only crude outlines of the female genitalic region (ZACHER 1911; BURR 1915; HINCKS 1959: figs.

3 Entomologische Abhandlungen 61 (2) ). These are therefore not useful for comparative morphology, but they show that in Pygidicranidae there is a considerable structural diversity in this body area, the female genitalia thus potentially being highly suitable for reconstructing basal dermapteran phylogeny. There is thus a need for descriptive work on the female genitalic region in Pygidicranidae, as well as for comparison within the Dermaptera and between Dermaptera and other Insecta. Furthermore, in view of what is currently known about pygidicranid female genitalia the question arises whether, or to what extent, ovipositor reduction actually is a groundplan condition of Dermaptera. Indeed, some of the basic components of the dicondylian female genitalia have remained unreported for Dermaptera, such as the posterior gonapophyses, gonangula, and accessory glands of segment IX. However, the finding in a recent preliminary study of Karschiella buettneri (Karsch, 1886) (KLASS 2001a: figs. 15, 16) of all these elements (though the gonapophyses IX are quite small) makes the ovipositor in the dermapteran ground plan now appear much more complete than previously thought. The fact that the female genitalic region in Karschiella differs fundamentally from that described for Echinosoma further demonstrates the need for original morphological work in Pygidicranidae. The objective of the present study is to close the major gaps in the knowledge of the female genitalic region in the basal Forficulina. Through the detailed study in ten genera of the exoskeleton of the female genitalia, including the intimabearing gonoducts, spermathecae, and accessory glands, descriptions are provided that should largely cover the structural diversity of this body region in Pygidicranidae. The genitalic musculature, which is entirely unknown in Forficulina, was examined in Dacnodes Burr, 1907 sp. In order to provide a wider morphological framework the entire postabdominal exoskeleton, which is fairly similar in all Dermaptera here studied, is described for Echinosoma yorkense Dohrn, Based on the findings on structure, topographic homologies (sensu KLASS 2001b: 230f) with other Insecta are proposed, groundplan conditions of the female genitalic components in Dermaptera are argued, and it is tested whether the female genitalia in the dermapteran ground plan actually display reductive (or other) apomorphies as compared to the alleged pterygotan ground plan. BITSCH s (1974a) and ROUSSET s (1973) descriptions of female genitalia in Archaeognatha and Zygentoma, and KLASS (1998, 2001a, in press) comparison of female genitalia between these apterygote insects and Dictyoptera, Hemimerus, and Odonata provide most of the framework for the comparative discussions, but descriptions of other ovipositor-bearing pterygotes are evaluated as well. Based on this all-insect framework, it is furthermore attempted to trace apomorphies shared between Dermaptera and other pterygotan taxa, or between any subgroups of Pterygota. This work provides a sound basis for using in forthcoming studies with a larger taxon sample female genitalic characters for reconstructing the phylogeny of the Dermaptera as well as the relationships between Dermaptera and other insects. In addition, this contribution should also strongly improve the applicability of female genitalic structure in the description and identification of pygidicranid species. The two major handbooks for the identification of Dermaptera (HINCKS 1955, 1959; STEINMANN 1986) both have the shortcoming that below the subfamily level in Pygidicranidae and other Dermaptera there is a strong focus on male structures (external genitalia, cerci, etc.), while females remain difficult to identify. A more consistent reference to the female genitalia, based on an improved knowledge of these structures, would thus be highly desirable. Eventually, the enormous variation in the structure of the female genitalia in Pygidicranidae can be supposed to be reflected in a similar diversity in the habits of copulation and egg laying. The morphological findings here presented is hoped to lead to an increased interest in the reproductive biology in the basal Dermaptera, which so far has been a strongly neglected field. 2. Material and methods The Dermaptera species here studied, their systematic placement, the number of female specimens available for this study, the kind of their preservation, and the depository (with restrictions mentioned below) are listed in Tab. 1. All Pygidicranidae had been determined by acknowledged specialists (A. Brindle, H. Steinmann, F. Haas), but the general uncertainty in the identification of females should be noted. In addition, an unidentified species of Tenthredo Linnaeus (Hymenoptera: Tenthredinidae) and Magicicada septendecim (Linnaeus, 1758) (Auchenorrhyncha: Tibicinidae) were studied for comparative purposes. Of the Pygidicranidae stored in ethanol only the two Dacnodes sp. indet. (from Tanzania) were suitable for a study of the muscles. Hereafter the sampled species are designated by the generic name alone; unless otherwise noted, Pyragra and Dacnodes refer to both studied (sub)species. When data from Tab. 1. Species of Dermaptera herein studied. 2nd column: systematic assignment according to STEINMANN (1986). 3rd column: number of specimens investigated. 4th column: kind of preservation of specimens (eth = ethanol). 5th column: depository, ZMUC = Zoological Museum of the University of Copenhagen, MTD = Museum für Tierkunde Dresden, MNHU = Museum für Naturkunde der Humboldt-Universität zu Berlin, HNHM = Hungarian National History Museum Budapest. Echinosoma yorkense Dohrn, 1869 Pygidicranidae: Echinosomatinae 4 eth ZMUC Dacnodes caffra (Dohrn, 1867) Pygidicranidae: Pygidicraninae 2 dry ZMUC Dacnodes sp. indet. Pygidicranidae: Pygidicraninae 2 eth ZMUC Tagalina burri Hincks, 1955 Pygidicranidae: Pygidicraninae 4 eth MTD Anataelia canariensis Bolivar, 1899 Pygidicranidae: Anataelinae 2 dry ZMUC Diplatys macrocephalus (Palisot debeauvois, 1805) Pygidicranidae: Diplatyinae 1 dry ZMUC Haplodiplatys orientalis Steinmann, 1974 Pygidicranidae: Diplatyinae 1 dry HNHM Karschiella buettneri (Karsch, 1886) Pygidicranidae: Karschiellinae 1 dry MNHU Bormansia monardi Menozzi, 1937 Pygidicranidae: Karschiellinae 1 dry HNHM Esphalmenus basidentatus Brindle, 1984 Pygidicranidae: Esphalmeninae 2 dry ZMUC Pyragra fuscata fuscata Serville, 1831 Pygidicranidae: Pyragrinae 1 dry ZMUC Pyragra fuscata brasiliensis (Gray, 1832) Pygidicranidae: Pyragrinae 1 dry ZMUC Forficula auricularia Linnaeus, 1758 Forficulidae: Forficulinae 3 dry

4 176 KLASS: Female genitalia in basal Dermaptera previous studies are referred to, the names of the respective taxa, mostly species, are specified as (in)completely as in the original papers. Data on Hemimerus consistently refer to H. vosseleri Rehn & Rehn, 1935 as treated in KLASS (2001a). For the study of the exoskeleton, the postabdomen (segments VIIff) was macerated in 10 % KOH. Tracheae were cut near the spiracles and removed. The exoskeleton was examined in 70 % ethanol under a stereo microscope, using magnifications 20x 80x. In some specimens the cuticle was mostly cut down to pieces to trace cuticular thickenings, lines of membrane, and articulations; in case of specimens with genitalia used up in this way, the depository given in Tab. 1 relates only to the non-genitalic parts of the specimen. It is here attempted to indicate the length of the spermathecal tubes, but this is difficult to measure because due to their sclerotization the tubes cannot be fully stretched without breaking. In the muscle studies the postabdomen was dissected in 70 % ethanol. Illustration of the female genitalic region (Figs. 1 62) includes for most pygidicranid species here studied a complete ventral (mainly external) view, a complete dorsal (mainly internal) view, and one or more figures showing details. Sclerotized parts of the exoskeleton are shown darkened, membranes are left white. Undulate lines represent cuts through the cuticle. Setae and cuticular sculpture and thickness are usually not considered, but discrete ridges or patches of thickened cuticle that are distinctly delimited from the thinner surrounding cuticle are indicated by hatched stripes or areas. Of the oviducts and accessory glands all intima-bearing parts are included, as resulting from KOH-maceration; when a loss of cuticulized internal parts of these elements is suspected, this is noted in the descriptions. 3. Terminologies, morphological interpretations, and abbreviations 3.1. Elements studied and their interpretation Generalities. The studies include muscles and two categories of exoskeletal elements, namely the sclerites and the formative elements. The latter comprise all in- and evaginations and thickenings of the body wall, such as processes, apodemes, or ridges (for the usage of apodeme, ridge, and tendon see KLASS 2001a: 253). The designation and interpretation of the elements of the terminal abdomen is explained in KLASS (2001a). The terminology for the ventral (s.l.: sternal plus pleural) elements of the female genitalic segments VIII and IX is based on SCUDDER (1961a, b), but some modifications are applied (see Tab. 2) according to more refined contributions on Archaeognatha (BITSCH 1973, 1974a, b), Zygentoma (ROUSSET 1973), Dictyoptera (KLASS 1998), and Odonata (KLASS in press), and in view of new findings in the present study. SMITH s (1969) interpretations of female genitalic sclerites are here only partly accepted because neither sufficient descriptive/pictorial documentation nor discussions are provided. Furthermore, SMITH s data for Archaeognatha and Zygentoma are partly contradictory to the more extensive documentation in BITSCH (1974a) and ROUSSET (1973) (e.g., BITSCH 1974a: 113), and some of SMITH s observations in Odonata (e.g., presence of a notum of gonapophyses IX) were found incorrect in a recent study (KLASS in press). Ventral sclerites. In Archaeognatha an abdominal segment has, apart from the sclerotizations of the gonapophyses and styli, maximally five discrete ventral sclerites, which in the terminology of BITSCH (1973: fig. 2, 1974a: fig. 1) are the unpaired sternite and intersternite, and the paired coxites, laterocoxites, and precoxites (ste, ist, cox, lac, prc in Fig. 64). In SMITH (1969) the laterocoxite is called subcoxa, the coxite is called the gonocoxa, and the sternite, intersternite, and precoxites are apparently comprised as the coxosternum (hence interpreted as including coxal components). On venters VIII and IX of female Dicondylia four sclerotization areas can presently be distinguished, called here: (1) coxa (= gonocoxa in KLASS 1998; corresponding to BITSCH s coxite); (2) post-laterocoxa (= posterior part of laterogonocoxa/gonangulum in KLASS 1998; corresponding to BITSCH s laterocoxite); (3) ante-laterocoxa (= anterior part of laterogonocoxa/gonangulum in KLASS 1998; corresponding to BITSCH s precoxite); (4) sternum (corresponding to BITSCH s sternite and/or intersternite). The terms (1) (4) are here used like in KLASS (2001a, in press) and including the Archaeognatha. The terminological distinction between ante- and post-laterocoxa, which together constitute the laterocoxa (= gonangulum in segment IX), is based on the corresponding separations between ante- and post-laterocoxa in Odonata (segment IX) and some Pygidicranidae (segments IX and possibly VIII), and between precoxite and laterocoxite in Tab. 2. Terminology of ventral elements of abdominal segments VIII and IX (components of female genitalia) as applied in SCUDDER (1961a, b) and this paper. Terms enclosed in parentheses in 2nd column are not used by SCUDDER but supplemented following his terminological scheme. Segment SCUDDER This paper Abbreviation VIII Sternum VIII ST8 First gonocoxa partim Coxa VIII CX8 First gonocoxa partim Laterocoxa VIII (ante-, post-) LC8 (LCa8, LCp8) Gonapophyseal sclerite VIII GP8 (First gonoplac) Coxal lobe VIII = Gonoplac VIII gl8 First gonapophysis Gonapophysis VIII gp8 (First gonostyle) Stylus VIII sl8 IX Sternum IX ST9 Second gonocoxa Coxa IX CX9 Gonangulum Laterocoxa IX (ante-, post-) LC9 (LCa9, LCp9) Gonapophyseal sclerite IX GP9 (Second) gonoplac Coxal lobe IX = Gonoplac IX gl9 Second gonapophysis Gonapophysis IX gp9 (Second) gonostyle Stylus IX sl9

5 Entomologische Abhandlungen 61 (2) 177 Archaeognatha (1). Laterocoxa and coxa are designated collectively as the coxopodium (pleural), most of which (the ante-laterocoxa may have to be excluded) is probably formed by limb base sclerotizations. The paired coxopodia and the unpaired sternum together constitute the coxosternum. The pregenital abdominal sterna of Dicondylia are also regarded as such coxosterna (see KLASS 2001a: 272 for details and background). It should be emphasized that the use of these terms is not meant to indicate strict homonomy with parts of the thoracic limbs, and no reference is intended to theories of limb base composition in a large-scale arthropod view; the terminology as used herein indicates only hypotheses of inner-insect abdominal homology and homonomy (= serial homology, homodynamy). Ventral formative elements. Major projections in the female genitalic region of Archaeognatha and Zygentoma are the coxal lobes, the styli located upon them, and the gonapophyses, all present on both segments VIII and IX. A coxal lobe is presumably the projecting body of a true appendage, sclerotized by the coxa, and corresponding to the coxal and possibly more distal parts of the thoracic leg; further distal podomeres are probably represented by the stylus (KLASS & KRISTENSEN 2001: 274). The gonapophyses are likely homonomous with the eversible vesicles (BITSCH 1994), which in Archaeognatha and Zygentoma occur in varied configurations on the abdominal coxal lobes II VII but are (with few possible exceptions: reversals?) absent in Pterygota. Many Pterygota have coxal lobes IX (= gonoplacs of SCUDDER, lateral valves); coxal lobes VIII (not considered in SCUDDER s terminology) are usually suppressed, but strongly developed in some Pygidicranidae. The coxal lobes VIII/IX, here also called gonoplacs VIII/IX, are probably homonomous with the ventral intersegmental folds formed by the posterior part of each of the preceding abdominal segments. Discrete styli in adult female Pterygota occur on segment IX of Odonata (see KLASS in press) and few other insects (e.g., ANDER 1939: fig. 127; MICKOLEIT 1973a; DEITZ et al. 2003: 81), and on segment VIII in the blattarian Cryptocercus (KLASS 1998: fig. 15). They cannot be identified in the Pygidicranidae, where they are either absent or have been incorporated into the distal parts of the coxal lobes. Gonoducts. According to the basic scheme of SNODGRASS (1933: 16ff), in the ontogenetic development of insects each of the segments VII, VIII, and IX forms a median invagination at its ventral hind margin; these are the rudiments of the common oviduct, spermatheca, and accessory gland. The common oviduct later acquires contact with the mesodermal lateral oviducts and their ampullae. Depending on details of this procedure the inner rim of the cuticle (likely the border between ectoderm and mesoderm) lies either within the common oviduct or within the lateral oviducts (review of examples in KLASS in press). In cases where the common oviduct opening develops directly into the definitive genital opening, the intima-bearing median gonoduct in the adult is exclusively constituted by the common oviduct, whose opening is here called a primary gonopore. In cases where a groove forms behind the common oviduct and becomes closed ventrally, the extension of the duct is here called the extended oviduct, the intima-bearing median gonoduct in the adult is constituted by the common plus the extended oviducts, and the opening on segment VIII is the secondary gonopore. In cases where additionally the VIIIthsegmental area bearing the secondary gonopore and the spermathecal opening becomes invaginated anteriorly, the terminal part of the intima-bearing median gonoduct is a vagina (or a genital chamber if it is not a discrete tube), whose opening is the vulva (see SNODGRASS 1933: fig. 4A D; Odc = common oviduct; Odc = common + extended oviduct = median oviduct). The application of this scheme is tentative in cases where either the entire gonoducts develop from an invagination of segment VIII or the development of the ducts is unknown. The term spermatheca is here used in a morphological sense to comprise all surmised derivatives of the spermathecal invagination. It frequently consists of functionally and histologically differentiated partitions (e.g., copulatory pouch and seminal vesicle in Zygentoma, ROUSSET 1973: 60) Segmental assignment For sclerites the segmental assignment is given with respect to secondary segmentation because the latter is established by the sclerites. For formative elements and muscles assignment is given, as far as possible, with respect to primary segmentation. Components assumed to lie between two primary segments or to be of bisegmental origin (e.g., antecostae, anterior tergo-coxosternal muscles) are formally assigned to the respective posterior segment (see KLASS 1999: 7). In segments VII IX the belonging of most components to a certain primary segment is obvious from their positions. For the assignment of elements behind segment IX see discussions in KLASS (2001a) Terminologies Sclerites are designated by terms composed of two upper case letters; their subdivisions are specified by a lower case letter in the third position. Formative elements including the components of the gonoducts are given terms composed of two lower case letters. A number in the last position gives, if needed, the assignment of a sclerite or formative element to a segment. Single lower case letters are used to mark some particular locations in the exoskeleton that are referred to in the descriptions (a d: levels of longitudinal section through the postabdominal terga; e q: specific areas or fusions of sclerites; for meaning of x, y see in 4.1). Muscles are designated by Arabic numerals, and partly by descriptive names in addition. The terminological principles followed, their inherent problems, and the terms used for characterizing the shape of a muscle are explained in KLASS (1999: 5f). Unless otherwise noted, muscles are present as a pair. Generally in this paper, the equal designation of elements expresses the assumption of homology or homonomy, with the restrictions given in the discussions in terms of probability. All terms are given in bold print. Only some terms that BITSCH (1974a) and ROUSSET (1973) used for sclerites of Archaeognatha and Zygentoma and that are also used herein for clear reference are given underlined and in Italics. Terms from other contributions are marked with an asterisk Abbreviations muscles ac (+ number) antecosta (number = segment following) ag accessory gland: ag(p) paired anterior glands, ag(u) unpaired posterior gland an anus AP anal plate (mesal part of paraproct) AS paired sclerites anterior to spermathecal opening at (+ number) tendon or apodeme anterolaterally on coxosternum = sternal apophysis (number = segment following) (1) A terminological subdivision of laterocoxa is preferred to using BITSCH s laterocoxite and precoxite in order to avoid precoxa with its burden of arthropod limb base theories.

6 178 KLASS: Female genitalia in basal Dermaptera bs bulge bearing spermathecal opening CE sclerotization of cercus ce cercus cox coxite (term of BITSCH 1974a and ROUSSET 1973) cp coxal pouch IX CS (+ number) coxosternum (number = segment) ct tendons on cercal base CX (+ l-c-letter coxa (lower case letter = specification, + number) number = segment); ± valvifer on segments VIII and IX df (+ number) dorsal fold of segment overlapping succeeding segment (number = segment) dg dorsal groove of extended oviduct DT dorsal sclerite of telson ec line along which cuticle ends (mesoderm adjoining) fa apodeme anterolaterally on coxa IX fs field of setae anterior to ridge pr9 gb apodeme on dorsal gonoplac base and/or its lateral tendons gf genital fold (of venter VII) gl (+ number) gonoplac = coxal lobe = projecting body of abdominal limb (number = segment) gm mould harboring both gonapophyses IX go anterior border of definitive genital opening gon gonangulum (term of ROUSSET 1973) gp (+ number) gonapophysis (number = segment) GP (+ l-c-letter gonapophyseal sclerite (lower case letter + number) = specification, number = segment) ist intersternite (term of BITSCH 1974a) la apodeme posterolaterally on coxa IX lac laterocoxite (term of BITSCH 1974a) lsc languette sclerite (term of ROUSSET 1973) LC (+ l-c-letter laterocoxa (lower case letter = specification, + number) number = segment); gonangulum on segment IX lg lateral spermathecal grooves LP lateral plate (lateral part of paraproct) lt (+ number) lateral tendon beside coxosternum (number = segment) ma mesal apodeme on cercal base mg median spermathecal groove (posterior to spermathecal opening) ms (+ number) manubrium of spiracle (number = segment) MS mt (+ number) oc oe ol ola pg median sclerite of segment VIII mesal tendon or ridge of coxosternum (part of antecosta; number = segment following) common oviduct (intima-bearing) extended oviduct (intima-bearing) lateral oviduct, intima-bearing part lateral oviduct, mesodermal part pouch above genital fold, possibly a remnant of primary gonopore on hind margin of venter VII pr (+ number) posterior ridge on tergum (number = segment) prc precoxite (term of BITSCH 1974a) ps paired pouches into which several spermathecal tubes open PS poststernite pt (+ number) posterior tendon or apodeme on laterocoxa (number = segment) re rectum sb subanal lobe scs coxal scs-sclerite (term of BITSCH 1974a) sdovi sternite VIII (term of ROUSSET 1973) si (+ number) spiracle (number = segment) sl (+ number) stylus (number = segment) so spermathecal opening SO sclerotization of/around spermathecal opening sp spermatheca st spermathecal tendon ste sternite (term of BITSCH 1974a) tf projection homologous with terminal = caudal filament on tergum XI TG (+ number) tergum (number = segment) tl tp tr va vc vf (+ number) vr (+ number) lateral tubes (opening near accessory gland into tube pouch) tube pouch (posterior to accessory gland opening) transverse internal ridge on telson sclerite DT vagina ventral condyle supporting cercal base (on sclerite LP) ventral fold of segment overlapping succeeding segment (number = segment); subgenital lobe on venter VII longitudinal ventromedian cuticular ridge on coxosternum (number = segment) 4. Description of female genitalia and postabdomen of Pygidicranidae 4.1. Echinosoma yorkense and generalities General condition of cuticle. In the Dermaptera here studied, like in Hemimerus (see KLASS 2001a: 255), cuticle thickness varies strongly along the body surface, and the cuticle consists of two layers that in many body-wall areas are easily peeled apart. The external layer shows the pattern of sclerotization (stiff, brown) and membrane (flexible, transparent); the internal layer, though brownish in some areas, is milky and has a rubber-like texture and flexibility. It is unclear in which way this mechanical subdivision meets some biochemically or ultrastructurally defined layering. A thickening of the cuticle can be due to a thickening of either of these two layers or of both. Survey of postabdomen. Segment VII resembles the midabdominal segments, but its coxosternum CS7 (Fig. 5) is slightly longer. Terga TG8 and TG9 (Figs. 1, 2) are very short and both completely concealed beneath tergum TG7. Venters VIII and IX form the components of the external genitalia, with the gonapophyses gp and gonoplacs gl, some sclerites around their bases, the vagina, and the openings of the oviduct, spermatheca, and accessory gland. Most of these elements are covered ventrally by coxosternum CS7. Tergum TG10 is very long (Fig. 1). Its lateral parts bend far to the ventral side, where sclerites LP and AP follow (Fig. 2). The anus an lies between the sclerites AP. The cerci ce have their bases beneath the lateral hind margin of tergum TG10 (Fig. 1). Tergum TG11 and the dorsal telson sclerite DT are located in between the bases of the cerci. TG11 forms a projection tf, the apparent posterior tip of the body. Coxosternum VII, subgenital lobe, and genital fold. Most of venter VII (Fig. 5) is sclerotized by coxosternum CS7 (subgenital plate). The posterior part of venter VII forms a wide posteriad-directed lobe (subgenital lobe vf7), which also dorsally is weakly sclerotized by a part of CS7. CS7 has a strong antecosta ac7 along its entire anterior margin, which in the midline gives rise to a short longitudinal ridge vr7. Laterally the antecostal thickening occupies stout apodemes at7 with a sclerotized dorsal and a largely membranous ventral wall. The pleural membrane along the lateral margin of CS7 bears a lateral tendon lt7 and a spiracle si7. The latter has a distinct apodeme (manubrium ms; no further details of the spiracles were studied). Venter VII differs from the mid-abdominal venters only in that the fold vf is slightly longer and its hind edge is more convex. This difference in proportion is in all other species here studied much stronger than in Echinosoma. The mid-abdominal folds vf are also weakly sclerotized dorsally. The enclosed space above fold vf7 is here called the vestibulum (like the corresponding space in Dictyoptera: MCKITTRICK 1964; KLASS 1998). The membrane bending from the anterodorsal base of the subgenital lobe (edge y in Figs. 2, 5) posterodorsad towards the female genitalia bears a wide but

7 Entomologische Abhandlungen 61 (2) 179 Figs Echinosoma yorkense. Orientation: anterior. Scale in mm. 1: Dorsal view of postabdomen, segments VIIIff. Dorsal wall removed on left side. Cerci cut on both sides. Terga VIII and IX simplified, with only external layer of cuticle considered (see Figs. 6 9). Spermatheca, accessory gland, and rectum cut near their openings. 2: Ventral view of postabdomen, segments VIIIff. On left side of figure gonapophysis VIII and gonoplac IX cut basally, and window cut into sclerite LP. Cerci cut on both sides. Terga VIII and IX simplified. 3: Dorsal view of base of left cercus. 4: Ventral view of base of right cercus. 5: Coxosternum VII (subgenital plate). Dorsal, predominantly internal view; lateral parts forced downwards. Dorsal wall of subgenital lobe (= ventral fold VII) removed on left side. Genital fold and anteroventral parts of segment VIII included on right side. very short membranous fold (genital fold gf; in Fig. 10 and corresponding illustrations for the other species the membrane forming gf is flapped to the anterior along the line marked x x). Fold gf is a far posterior element of venter VII and peculiar to this segment. Terga VIII, IX, and X. It is frequently claimed that in Dermaptera terga VIII X are fused (e.g., GILES 1963: 134). However, conditions are more complicated. In all species here studied terga VIII and IX overlap their succeeding counterpart through dorsal folds (df8, df9 in Figs. 8, 9). Nevertheless, in some species including Echinosoma all terga VIII X are immovable relative to each other. The kind of their interconnection is understood when the external and internal layers of the cuticle, which are easily peeled apart, are considered separately (Figs. 6 9). Regarding only the external layer, the three terga are all free from each other: There are weak tergal sclerites from whose posterior margins membrane bends anteriad towards the anterior margin of the following tergum (Figs. 8, 9; the difference between sclerotized and membranous is not very conspicuous). The appearance of the terga in an internal view (Fig. 7) is, considering only this layer, just as it is expected from typical

8 180 KLASS: Female genitalia in basal Dermaptera Figs Echinosoma yorkense. No scale. 6, 7: Scheme of terga VIII X, left halves, internal view. Orientation: dorsomesal, ventromesal, anterior. Only external layer of cuticle considered in Fig. 7, entire cuticle considered in Fig. 6. Arrows on top indicate levels of sections shown in Figs. 8, 9. 8, 9: Longitudinal sections through terga VIII X, at the levels indicated in Fig. 6. External (ext) and internal (int) layers of cuticle considered. Orientation: external, internal, anterior. successive terga. The internal layer follows, in the lateral parts of the terga (at a in Fig. 6), all foldings of the external layer (Fig. 9); here the three terga are entirely free and movable. Farther dorsomesally (from b in Fig. 6 onwards) the internal layer fills the space between the dorsal and ventral walls of the folds df and thus renders the inner surface of the cuticle an almost even plane all along terga VIII X (Fig. 8). Still farther mesally (at c in Fig. 6) tergum VIII separates from tergum IX. Close to the midline (at d in Fig. 6) tergum IX separates from tergum X, and a section through this area complies with Fig. 9. Hence, interconnection between terga VIII X is not due to a sclerite fusion (= loss of membrane between sclerites) but to the peculiar condition of the thick internal cuticular layer, which holds all sclerites and membranes in a firm position. KLASS (2001a) called this kind of connection tergal immobilization. In Dermaptera in general this is likely restricted to adult females, and it is in Echinosoma less complete than in, e.g., Hemimerus and Forficula (see KLASS 2001a: 257, 268). There is still more complexity in the details of terga VIII X, which differ quite strongly among the species here studied and might provide useful characters; however, a comparative analysis of this issue is outside the scope of this study. The lateralmost parts of terga IX and X are in all species here studied actually fused, without membrane or overlapping between them (e in Figs. 1, 2, 6); the fusion area in Echinosoma bears some feeble folds (Figs. 10, 11). Each tergum VIII X has ventromesally a mesad-directed extension (f, g, h in Figs. 6, 10, 11). That of TG8 approaches a small sclerite, laterocoxa LC8. That of TG9 articulates with laterocoxa LC9 (gonangulum). That of TG10 has no contacts. The pleural membrane at the ventromesal margin of TG8 forms a small mesad-directed lobe bearing the orifice of spiracle si8 (Fig. 10). Terga TG9 and TG10 bear some internal ridges of thickened cuticle (Figs. 10, 11; no corresponding external grooves visible). Ridges ac9 and ac10 along the anterior tergal margins are likely antecostae. Ventromesally they target the tergal extensions g and h, thus strengthening them; dorsolaterally they obliterate, i.e., continue into a wide and indistinctly bordered thickening of the anterior part of the tergum. Tergum TG8 also has a discrete antecosta ac8 along its anterior margin, but this element is not considered in the present study. Ridges pr9 and pr10 extend posterolaterad from the tergal extensions g and h. pr9 dives beneath the fold between the terga TG9 and TG10 to join ac10. pr10 continues along the margin of TG10 and then crosses to sclerite LP, where it soon obliterates. The homonomy of pr9 and pr10 is doubtful. Elements of segments X and XI and telson. The heavy sclerite LP is completely separated from TG10 by a narrow membrane (Figs. 1, 2); also the ridge pr10 is membranous where it crosses from TG10 to LP (Fig. 10). Posterolaterally LP forms a blunt process vc, which serves as a condyle for the cercus (Fig. 2). Mesal to LP arises the posteromesad-directed subanal lobe sb, which bears a weak ventral sclerite AP (Fig. 2). The anus an lies between the two subanal lobes. The terminal portion of the rectum re is narrow and orientated vertically (Fig. 1). Tergum TG11 joins the median hind margin of TG10 via a narrow membranous hinge-line (Fig. 1). It is shaped like a pyramid, whose base lies vertically between the cercal bases, and whose lateral planes bulge inwards. The pyramid body, called here the projection tf (terminal filament ), forms the apparent posterior tip of the animal. The dorsal telson sclerite DT joins via another narrow membranous hinge-line (Fig. 2) the ventral (actually posterior) margin of TG11 and lies between the ventral parts of the cercal bases and the anus. An internal transverse ridge tr borders a stronger abanal part of DT from a weaker adanal part. In a strict morphological sense, the anus an is the posteriormost area of the body, sclerites TG11 and DT are both dorsal (though DT is entirely located on the ventral side of the body), TG11 is located anterior to DT, and the pyramid tf is a dorsal projection. The cerci ce in Echinosoma are one-segmented claspers as in all adult Forficulina (HINCKS 1959: fig. 124; KLASS 2001a). The bases of the cerci are tightly embraced by scle-

9 181 Entomologische Abhandlungen 61 (2) Figs Echinosoma yorkense. Orientation: anterior (except Figs ). Scale in mm. 10: Ventral view of female genitalic region. Right side of figure with lateral (i.e. ventrally located) parts of terga VIII X and sclerites LP and AP. On left side gonapophysis VIII and gonoplac VIII cut basally. 11: Dorsal view of female genitalic region. Spiracle VIII cut near atrium and simplified. Longer part of spermatheca removed (cut by bars). 12, 13, 14: Transverse sections through vagina, at levels indicated in Fig. 11. Orientation: dorsal, lateral. Thin lines = membrane, thick lines = sclerite, dashes = setose areas. 15, 16: Dorsal view of anteromedian part of female genitalic region. Dorsal parts of vagina successively removed, especially on left sides. Gonapophyses VIII cut. 17: Dorsal view of posteromedian part of female genitalic region. Left gonoplac IX entirely removed. Accessory gland cut longitudinally near midline, lateral tube cut.

10 182 KLASS: Female genitalia in basal Dermaptera rites TG10, LP, TG11, and DT (Figs. 1, 2), part of whose margins fold outward to form projections fitting with sculptural elements of the cercal base. For instance, condyle vc fits into a cercal base ginglymus (for details of the complex cercal articulations in Forficulina see STRENGER 1950). The cercal walls protrude basally into the body cavity to form an inward-directed ring-wall (Figs. 1, 3, 4), which bears four delicate cuticular tendons ct, and whose deeply projecting mesal portion forms a stout cercal apodeme ma. Along the inner base of the ring-wall runs a circular rupture line, along which the cercus easily breaks during dissection leaving behind a narrow sclerite ring (with apodeme ma) surrounding a hole. The hole, however, is almost closed by a wide, delicate cuticular flap, which projects internally from the retained part of the cercal base. This rupture line was only found in Echinosoma. Its presence appears contradictory to the function of the cerci as strong claspers. On the other hand, the cuticular flap seems to be suited for preventing an extensive loss of body fluids in advance to healing and indicates that an autotomy of the cerci could in Echinosoma be a defensive strategy. Ventral elements of segment VIII. The small laterocoxa LC8, located mesal to spiracle si8 (Figs. 10, 11), hinges posteriorly with the front margin of tergum TG9. A membranous tendon at8 originates immediately anterior to LC8, and a membranous bulge gl8 is situated farther mesally (area of obliterated coxa CX8). The gonapophyses gp8, having their bases far mesal to this bulge, bear two sclerites: The vaguely S-shaped GPa8 (Fig. 10) occupies the basal ventral wall of gp8. GPb8 occupies the distal ventral wall of gp8 and bends, where it meets GPa8, sharply into the dorsal wall of gp8. Here it continues (Fig. 15) far anteriad into a deep pouch (vagina va), whose lateral walls are folded longitudinally. Where it enters the vagina, sclerite GPb8 bends dorsad around one of these folds, and it bends further dorsad shortly before it terminates. The vagina va is by the longitudinal folds divided into three chambers one above the other (Figs ); some parts of the chamber walls are densely clothed with fine bristles. The narrow, delicate lateral oviducts ol open into a longitudinal groove oc+oe in the ventral wall of the vagina (as in the following species, only the intima-bearing terminal parts of the lateral oviducts are here considered). The groove oc+oe is here regarded as a very short common oviduct that may additionally include an extended oviduct oe; its lining in the ventral wall of the vagina is thus either a primary or a secondary gonopore. The spermatheca sp opens into the anterior dorsal wall of the vagina. It is a very long (more than 10 mm), unbranched tube with a wider, membranous external portion, a narrower, sclerotized internal portion, and a simple inner end (Fig. 11). The tube forms a dense coil lying on the roof of the vagina. The transverse fold go between the two gonapophyses gp8 is the anterior border of the vaginal opening (= vulva; Figs. 10, 15, 16). Accordingly, the definitive genital opening is located between and behind the bases of the gonapophyses gp8, and hence on segment VIII. Ventral elements of segment IX. The large laterocoxa LC9 (gonangulum) extends from the tergal extension IX g to the ventral base of gonapophysis gp8, where it articulates with sclerite GPa8 (Figs. 10, 11). About midway its hind margin articulates with sclerite CXf9. Thickened cuticle forms an L-shaped ridge on the lateralmost part of LC9 and a short ridge at the articulation with CXf9 (Fig. 11). The L- shaped ridge also supports an apodeme pt9 on the posterolateral corner of LC9. The gonoplacs gl9 are large and spatulate, and entirely sclerotized by CXh9. A transverse fold connects the two gl9 at their bases, and at the dorsal base each gl9 bears a small tendon gb. Sclerite CXh9 has three basal arms (i, j, k in Figs. 10, 11). One (i) extends mesad upon the transverse fold to approach (but not meet) its counterpart of the other side. The second arm (j) extends anteriad to embrace gonapophysis gp9 laterally. The third, lateral arm (k) continues into sclerite CXf9, which approaches LC9 and projects beneath it to form a small apodeme (Fig. 11). A cuticular ridge runs all along CXf9, the sclerite thus being rod-like. At the posterior end of the ridge, CXf9 is flexible upon CXh9. The gonapophyses gp9 are small membranous lobes. Immediately behind their bases open, via two juxtapositional transverse slits, the accessory gland ag and a structure here called the tube pouch tp. The accessory gland is sac-like and internally divided into delicate, finger-like branches. The tube pouch is the very short common outlet duct of a pair of lateral tubes tl with club-shaped internal ends. Differences to GILES (1963). GILES description of Echinosoma afrum differs strongly from my findings in Echinosoma yorkense. The gonapophyses gp9, accessory gland ag, tubes tl, plates AP, and subanal lobes sb are neither illustrated nor mentioned. The part of sclerite GPb8 that lies within the vagina is shown to articulate with the anterior tip of GPa8 and to be disconnected with the distal part of GPb8. The gonoplac sclerites CXh9 lack the arms i and j. The oviduct oc is shown as a tube that straightly continues the vagina to the anterior. The laterocoxae LC8 are interpreted as parts of sternum VIII, the anterior vaginal parts of sclerite GPb8 as valvifers VIII (coxae VIII), and the laterocoxae LC9 (gonangulum) as valvifers IX. These interpretations will be revised in chapter Preview of other species The remaining pygidicranids here studied differ all strongly from Echinosoma in the structure of the female genitalic region. The Pygidicraninae (Dacnodes, Tagalina) and Anataelinae (Anataelia) are fairly similar. They show a very rich differentiation of the female genitalic components and appear as the Dermaptera with the most plesiomorphic ovipositors. Conspicuous differences to Echinosoma lie in the much stronger developement of the coxal lobes VIII and gonapophyses IX, in the smaller size of the coxal lobes IX (gonoplacs IX) and the vagina, and in the lack of the extension of sclerite GPb8 into the vagina. The Diplatyinae (Diplatys, Haplodiplatys), Karschiellinae (Karschiella, Bormansia), Esphalmeninae (Esphalmenus), and Pyragrinae (Pyragra) differ strongly from the foregoing taxa as well as from each other. They show, as compared to, e.g., Dacnodes, very different reductions and modifications. Haplodiplatys and Diplatys are strikingly similar in some respects, but surprisingly different in others. Karschiella and Bormansia are altogether quite similar. Esphalmenus differs strongly from Echinosoma through its extensive development of the coxopodia VIII (LC8 plus CX8), but conspicuously resembles it in the shape and enormous size of the gonoplacs IX. Though most elements of the female genitalia show much variation among the species described in the following, the strong differences in the spermathecae, gonoducts, and laterocoxae VIII are most noteworthy Dacnodes sp. indet. and Dacnodes caffra Elements surrounding the genitalic area. Spiracle si8 opens, as in Echinosoma, on a small lobe (Figs. 18, 23). Terga VIII X have small mesad-directed extensions f, g, h; those of TG8 and TG9 articulate with the laterocoxae of their segments, LC8 and LC9 (Figs. 18, 19, 23, 24); that of TG10 (absent in D. sp. indet.) is, in contrast to Echinosoma, small and weak, not strengthened by cuticular thickening. The tergal ridges resemble those in Echinosoma, but ac10 is excessively developed as a high, massive fold (Figs. 18, 19, 23, 24, left sides), which anterolaterally continues into the infolded anterior edge of tergum TG10, and which posteriorly continues straightly into pr10 (according to condi-

11 Entomologische Abhandlungen 61 (2) 183 Figs Dacnodes sp. indet. Orientation: anterior. Scale in mm. 18: Ventral view of female genitalic region. Right side of figure with lateral parts of terga VIII X and sclerites LP and AP; gonapophysis VIII cut near midlength. On left side gonapophysis VIII and gonoplac VIII cut basally, and most of the excessively developed ridge ac10 retained. 19: Dorsal view of female genitalic region. Spiracle VIII cut near atrium and simplified. Longer part of spermatheca removed (cut by bars). Internal branchings of accessory gland largely retained. Left lateral tube of accessory gland cut near its opening. 20: Dorsal view of anteromedian part of female genitalic region. Tips of gonapophyses VIII cut. Left side with further dorsal parts of vagina removed. 21: Dorsal view of posteromedian part of female genitalic region. Dorsomedian wall of accessory gland removed, lateral tubes cut. tions in Echinosoma, extension h in D. caffra is considered as marking the border between ac10 and pr10). In front of pr9 lies a conspicuous field fs of short setae. Sclerites LP and TG10 are entirely separated by membrane. The membranous genital fold gf on the hind margin of venter VII is narrow but relatively long, hence quite conspicuous (Figs. 18, 23). Ventral elements of segment VIII. The small laterocoxa LC8, located mesal to spiracle si8 (Figs. 18, 19, 23, 24), articulates anteriorly with tergum TG8 and posteriorly with TG9. An internal ridge runs along LC8 and joins, traversing the articulation, ridge ac9. The membranous tendon at8, originating shortly mesal to LC8, is much larger in D. sp. indet. than in D. caffra. Another tendon mt8, which is lacking in Echinosoma, is located mesal to at8; only in D. sp. indet. it is sclerotized by another individualized part of laterocoxa LC8. Coxa CX8 is represented by the weak sclerotization of the large coxal lobe gl8. The gonapophyses gp8 bear two sclerites: GPa8 (Figs. 18, 23), a narrow, curved ribbon, lies in the basal ventral wall. GPb8 occupies the distal ventral wall; in contrast to Echinosoma, it does not extend further basad from its articulation with GPa8 (Fig. 25). In the midline the two gonapophyses gp8 are transversely connected by a short fold go. The go-area forms either a pair of small sclerites (D. sp. indet.) or a single median sclerite (D. caffra) MS. Above the fold go and the bases of the gonapophyses gp8 lies the entrance into the vagina va (Figs. 19, 20, 24, 25). Through longitudinal folds in its lateral wall (Figs. 19, 20, 24, 25) the vagina is, like in Echinosoma, divided into three chambers one above the other (more distinct in D. caffra). However, the dorsal chamber, which receives the spermatheca, is represented

12 184 KLASS: Female genitalia in basal Dermaptera Fig. 22. Dacnodes sp. indet. Dorsal view of female genitalic region with muscles. Muscles striped according to course of their fibers. Insertion areas indicated by dashed lines (if muscle shown) or by white areas (if muscle not shown). Some muscles cut (by undulate lines), and opposite insertion areas not shown (see descriptions). Mesodermal parts of left lateral oviduct included. Scale in mm. only in the posterior half of the vagina. The delicate lateral oviducts ol unite to form a tube oc+oe, which is likely composed of a common and an extended oviduct. This tube opens into the anteroventral wall of the vagina. The spermatheca sp opens through the mid-dorsal vaginal wall into the anterior part of the dorsal chamber. It is a very long (not measured), unbranched tube with a wider, membranous external portion (strongly curved in D. caffra, Fig. 24), a narrower, sclerotized internal portion, and a simple inner end. The spermatheca forms one dense coil above the vagina and (as observed in D. sp. indet.) beneath the terminal abdominal ganglion, which probably is composed of neuromeres VIIff. Ventral elements of segment IX. Laterocoxa LC9 extends from tergal extension IX g to the ventrolateral base of gonapophysis VIII gp8, where it articulates with sclerite GPa8 (D. caffra, Fig. 23) or is synsclerotic with it (D. sp. indet., Fig. 18). The lateral part of LC9 is strengthened by some ridges, and it bulges inward to form a heavily sclerotized anteriad-directed fold mt9 (Figs. 19, 24). About midway LC9 articulates posteriorly with sclerite CXf9. The posterolateral corner of LC9 forms a small apodeme pt9. The moderately sized gonoplacs gl9 are entirely sclerotized by CXh9. A transverse fold connects the two gl9 basally, and from the dorsal base of the fold a transverse infolding gb projects anteriorly. The mesal arms i of CXh9, upon the fold, are narrowly separated dorsally (Figs. 19, 24) and ventrally (Figs. 18, 23). The lateral anterior margin of CXh9 hinges with sclerite CXf9. The posteromesal corner of CXf9 articulates, in turn, with a small sclerite GP9 in the basal lateral wall of gonapophysis gp9. The anterior part of CXf9, strengthened by a short (D. caffra) or a long (D. sp. indet.) internal ridge, approaches sclerite LC9 for articulation. The gonapophyses gp9 (Figs. 18, 21, 23, 26) are much larger than in Echinosoma. Between the bases of the two gp9 open the accessory gland ag and the lateral tubes tl (Figs. 19, 21, 24, 26). The accessory gland resembles that in Echinosoma, but the tubes tl are much longer than in Echinosoma and have simple inner ends; only in D. caffra the tl have a short common outlet pouch tp. Musculature. The lateral and ventral muscles of the female genitalic area as well as some muscles of segment VII (ventral muscles) and of the postgenital abdomen were studied in Dacnodes sp. indet. (Fig. 22). In the following list Figs Dacnodes caffra. Orientation: anterior. Scale in mm. 23: Ventral view of female genitalic region. Right side of figure with lateral parts of terga VIII X and sclerites LP and AP. On left side gonapophysis VIII and gonoplac VIII cut basally, and most of the excessively developed ridge ac10 retained. 24: Dorsal view of female genitalic region. Spiracle VIII cut near atrium and simplified. Longer part of spermatheca removed (cut by bars). Internal branchings of accessory gland mostly not retained. Left lateral tube of accessory gland cut near its opening. 25: Dorsal view of anteromedian part of female genitalic region. Left side with further dorsal parts of vagina removed. Tips of gonapophyses VIII cut. 26: Dorsal view of posteromedian part of female genitalic region. Dorsomedian wall of accessory gland and tube pouch removed, lateral tubes cut.

13 Entomologische Abhandlungen 61 (2) 185

14 186 KLASS: Female genitalia in basal Dermaptera means from... to..., and the numbers of the presumably homologous muscles of Hemimerus (KLASS 2001a: 260ff) are given in brackets, (--) meaning the absence of a homologue in Hemimerus. Muscles of segment VII. 1 (9): Coxosternum CS7, anterior margin Laterocoxa LC8, on tendon mt8. Internal ventral muscle. One moderately strong, compact sheet. 2 (10): Coxosternum CS7, anterior part Laterocoxal area VIII, on and mesal to tendon at8. Part of external ventral muscle. One strong, compact sheet. 3 (20+21): Coxosternum CS7, anterior margin Vagina, dorsal wall. Part of external ventral muscle. One moderately strong, compact sheet. 4 (--): Coxosternum CS7, posterolateral part Membrane of far anterior dorsal wall of subgenital lobe vf7. Part of external ventral muscle. One moderately strong, compact sheet. Muscles of segment VIII. 5 (part of 23): Vagina, dorsolateral wall Laterocoxa LC8 on tendon mt8. One delicate, compact sheet. 6 (part of 23): Vagina, lateral wall Membrane shortly lateral to vagina. Diffuse groups of few fibers. 7 (--): Longitudinal and transverse fibers in dorsal wall of vagina. Delicate and diffuse. 8 (26): Laterocoxa LC8 on tendon mt8 Laterocoxa LC9 on apodeme mt9. Internal ventral muscle. One compact sheet. 9 (27): Manubrium ms of spiracle si8 Posteromesal wall of atrium of spiracle si8. Likely a spiracle occlusor. One small, compact bundle. 10 (--): Manubrium ms of spiracle si8 Ventromesal part of tergum TG8, anterior to spiracle. Likely a spiracle dilator. One small, compact bundle. Muscles of segment IX. 11 (32): Laterocoxa LC9 on apodeme mt9 Membrane anterior to sclerite AP, on a small tendon. Internal ventral muscle. One narrow, compact sheet. 12 (33): Laterocoxa LC9 on apodeme mt9 Membrane anteromesal to sclerite AP. Internal ventral muscle. One narrow, compact sheet. 13 (25?): Tergum TG9, lateral part Laterocoxa LC9 on apodeme pt9. Intrasegmental tergo-laterocoxal muscle. One strong, compact bundle. 14 (--): Tergum TG9, lateralmost part Coxa CXf9, lateral margin. Intrasegmental tergo-coxal muscle. One strong, compact bundle. 15 (part of 34): Transversely on venter IX, connecting apodemes mt9 of laterocoxae LC9. Unpaired. One slender, compact bundle. 16 (part of 34): Transversely on membranous venter IX, connecting points shortly anteromesal to coxae CXf9. Unpaired. Slightly diffuse. 17 (--): Coxa CXf9, anterior part Membrane posteromesal to base of gonapophysis gp9. Coxo-gonapophyseal muscle. One moderately strong, compact bundle. 18 (--): Coxa CXf9, posterior part Coxa CXh9, anterior part. A delicate sheet of fibers in which no striation was found by light microscopy; possibly a degenerated muscle. Muscles of postgenital abdomen and rectal muscles. 19 (--): Tergum TG10, lateralmost part, along mesal flank of ridge ac10 Apodeme gb, dorsal face. One strong, compact bundle. 20 (36?): Sclerite LP, anterior part Dorsal telson sclerite DT. One strong, compact bundle. 21 (41): Tergum TG10, lateral part, along posterolateral flank of ridges ac10 and pr10 (the part of the insertion extending farther up on tergum TG10 was not traced) Cercus base, lateral part. 22 (46): Laterocoxa LC9 on apodeme mt9 A longitudinal line in ventrolateral wall of rectum. Ventral extrinsic rectal muscle. Compact near laterocoxal insertion but spreading fanwise and becoming diffuse towards rectal insertion. 23 (49?): Sclerite LP Anterior lateral wall of rectum. One slightly diffuse bundle. 24 (48?): Tergum TG10, lateral margin Posteriormost lateral wall of rectum. One narrow, compact bundle Tagalina burri Elements surrounding the genitalic area. Spiracle si8 opens on a small lobe (Fig. 27). Terga VIII and IX (not X) have small mesad-directed extensions f, g (Figs. 27, 28); that of TG9 articulates with the laterocoxal sclerite LCp9. The tergal ridges resemble those in Echinosoma, but the ventromesal ends of ac9 and pr9 are separated, and ac10 continues straightly into pr10 like in Dacnodes. In front of pr9 lies a field fs of short setae. Where pr10 crosses to sclerite LP, LP is synsclerotic with TG10. The genital fold gf of venter VII is wide and short like in Echinosoma (Fig. 27). Ventral elements of segment VIII. Laterocoxal sclerites LC8 are absent (Figs. 27, 28; cf. Figs. 18, 19). Tendon at8 originates shortly mesal to spiracle si8, and a short tendon mt8 arises farther mesally. Coxa CX8 is represented by the weak sclerotization of the long coxal lobe gl8. The gonapophyses gp8, which originate far mesal to the coxal lobes and have a stout base and a slender distal part, bear two sclerites (Fig. 27): The weakly S-shaped GPa8 lies in the basal ventral wall of gp8. GPb8 occupies the distal and mesobasal ventral wall of gp8; the part that extends parallel to GPa8 to the base of the gonapophysis takes the same position as MS in Dacnodes and is here labeled MS. Between the anterior tips of the MS the two gonapophyses are transversely connected by a short fold go. Above this fold and above the bases of the gonapophyses gp8 lies the entrance into the oval vagina va (Figs. 28, 29). Through longitudinal folds in its lateral wall (Figs. 28, 29) the vagina is divided into three chambers one above the other. The delicate lateral oviducts ol unite to form a tube oc+oe, which is likely composed of a common and an extended oviduct. This tube opens into the ventral wall of the vagina. The spermatheca sp opens into the anterior dorsal wall of the vagina. It has a wide, membranous, and curved external portion, which soon narrows and becomes weakly sclerotized. Farther internally the spermatheca divides by two dichotomies into four equal, long branches with simple inner ends (Fig. 28; a large portion of one branch was measured 14 mm). All parts of the spermatheca form together one dense coil. Ventral elements of segment IX. Laterocoxa LC9, which altogether extends from tergal extension IX g to the base of gonapophysis gp8, is divided into an anteromesal sclerite LCa9 and a posterolateral sclerite LCp9 (Figs. 27, 28). LCa9 articulates with sclerite GPa8, LCp9 with tergum TG9, and both sclerites articulate posteriorly with sclerite CXf9. Small membranous tendons at9 and pt9 (Fig. 28) arise from the articulation area between TG9 and LCp9 and from the posterolateral corner of LCp9. A ridge strengthens the lateral margin of LCp9. The narrow, moderately sized gonoplacs gl9 are entirely sclerotized by CXh9 (Figs. 27, 28). A transverse fold connects the two gl9 basally, and from the dorsal base of the fold a wide but short transverse infolding gb projects anteriorly (Fig. 28). The mesal arms i of CXh9, on the transverse fold, are narrowly separated dorsally (Fig. 28) and ventrally (Fig. 27). The anterior margins of CXh9 are narrowly connected with sclerite CXf9. CXf9 has a posteromesal extension that is continuous with sclerite GP9 in the lateral wall of gonapophysis gp9 (Fig. 27). The anterior part of CXf9 approaches the sclerites LC9 for articulation. An internal ridge runs all along CXf9 and continues posteriorly into a distinct dark line crossing to sclerite CXh9. The moderately sized gonapophyses gp9 resemble the gp8 in shape. Between the bases of the two gp9 open the accessory gland ag and the tube pouch tp (Figs. 28, 30, 31). The latter bears moderately long tubes tl with simple inner ends.

15 Entomologische Abhandlungen 61 (2) 187 Figs Tagalina burri. Orientation: anterior. Scale in mm. 27: Ventral view of female genitalic region. Right side of figure with lateral parts of terga VIII X and sclerites LP and AP. On left side gonapophysis VIII and gonoplac VIII cut basally. 28: Dorsal view of female genitalic region. Spiracle VIII cut near atrium and simplified. Of spermatheca, longer part of one branch and most of three branches removed (cut by bars). Left lateral tube of accessory gland cut near its opening. 29: Dorsal view of anteromedian part of female genitalic region. Most of vagina removed on left side. Tips of gonapophyses VIII cut. 30, 31: Dorsal view of posteromedian part of female genitalic region. In Fig. 30 dorsal wall of accessory gland and of tube pouch removed. In Fig. 31 accessory gland and tube pouch cut longitudinally near midline.

16 188 KLASS: Female genitalia in basal Dermaptera 4.5. Anataelia canariensis Elements surrounding the genitalic area. Spiracle si8 opens on a small lobe (Fig. 32). Terga VIII and IX (not X) have distinct mesad-directed extensions f, g (Figs. 32, 33); that of TG9 articulates with laterocoxa LC9. The tergal ridges and posterior elements are like in Tagalina, with the mesal ends of ac9 and pr9 separated, with ac10 continuing straightly into pr10, with a field of setae fs in front of pr9, and with TG10 and LP synsclerotic where ridge pr10 crosses to LP. The genital fold gf of venter VII is virtually absent (Fig. 32, cf. Fig. 23). Ventral elements of segment VIII. Laterocoxal sclerites LC8 are absent (Figs. 32, 33). Tendon at8 originates shortly mesal to spiracle si8, and a tendon mt8 arises farther mesally. Coxa CX8 is represented by the weak sclerotization of the bulge-shaped coxal lobe gl8 (Fig. 32). The gonapophyses gp8 bear two sclerites: GPa8 (Fig. 32), weakly S-shaped (but in the opposite way as compared to Echinosoma and Tagalina), lies in the basal ventral wall. GPb8 occupies the distal ventral wall and extends neither to the ventral gonapophysis base (compare MS-portion in Tagalina and MS in Dacnodes) nor into the dorsal base of the gonapophysis (as in Echinosoma). Medially the two gonapophyses gp8 are transversely connected by a short fold go. Above this fold and above the bases of the gp8 lies the entrance into the vagina va (Figs. 33, 34). Through longitudinal folds in its lateral wall (Figs. 33, 34) the vagina is divided into three chambers one above the other. The delicate lateral oviducts ol unite to form a tube oc+oe, which is likely composed of a common and an extended oviduct. This tube opens into the anteroventral wall of the vagina. The spermatheca sp (Fig. 33) opens into the anterior dorsal wall of the vagina, which is thickened and weakly melanized (sclerite SO). It is a very long (at least 22 mm), unbranched tube with a slightly wider, weakly sclerotized external portion, a narrower, more strongly sclerotized internal portion, and a simple inner end. The spermatheca forms one dense coil. Ventral elements of segment IX. Laterocoxa LC9 extends from tergal extension IX g to the ventral base of gonapophysis gp8, where it articulates with sclerite GPa8 (Figs. 32, 33); the opposing tips of LC9 and GPa8 are thickened. Far laterally LC9 is deeply notched; the LC9 portions on the two sides of the notch are, like the fully separated sclerites in Tagalina (Fig. 27), called LCa9 and LCp9. In the notch area the posterior margin of LC9 articulates with sclerite CXf9. A long ridge extends between the LC9-articulations with TG9 and CXf9, and a short ridge extends anteriad from the latter articulation. A thin membranous tendon pt9 arises from the posterolateral corner of LC9 (Fig. 33). The long, narrow gonoplacs gl9 are only ventrally sclerotized by CXh9 (Figs. 32, 33). A transverse fold connects the two gl9 basally, and from the dorsal base of the fold a transverse infolding gb projects anteriorly, which has discrete lateral tips and a median keel. CXh9 lacks a mesal arm i, the left and right CXh9 thus being widely separated (Fig. 32). Anteriorly CXh9 undergoes a constriction with a weak point upon a small bulge, beyond which sclerite CXf9 follows. The posteromesal corner of CXf9, within the constriction, articulates with sclerite GP9 in the lateral wall of gonapophysis gp9. The anterior part of CXf9, strengthened by a long ridge, approaches sclerite LC9 for articulation. The gonapophyses gp9 are quite large, with a stout base and a thread-like distal part. Between the bases of the two gp9 open the accessory gland ag and the wide tube pouch tp, the latter receiving a pair of tubes tl (Figs. 33, 35; the internal parts of both ag and tl were not found) Diplatys macrocephalus Elements surrounding the genitalic area. Spiracle si8 opens on a small lobe (Fig. 36). Only tergum IX has a small mesad-directed extension g, which articulates with laterocoxa LC9. The tergal ridges (Figs. 36, 38) are as in Tagalina and Anataelia, with the mesal ends of ac9 and pr9 separated, with ac10 continuing straightly into pr10, and with a field fs of setae in front of pr9. TG10 and LP are, as in Echinosoma and Dacnodes, separated where pr10 crosses to LP. The genital fold gf of venter VII is virtually absent (Fig. 36). Ventral elements of segment VIII. Laterocoxal sclerites LC8 are absent (Figs. 36, 38). The membranous tendon originating mesal to spiracle si8 is rather at8 than mt8 (cf. Fig. 32). A coxal sclerite CX8 is also lacking, but the short lobe found in the corresponding area represents a small coxal lobe gl8. The short and stout gonapophyses gp8, shaped like a bird s head, bear only one sclerite GP8 (Figs. 36, 39), which occupies the lateral parts. Anteromedially the two gonapophyses gp8 are transversely connected by a fold go. Above this fold and above and between the bases of the gp8 lies the entrance into a wide tube oc+oe, which likely includes a common and an extended oviduct (Figs ). It bears a pair of laterally directed ventral folds but otherwise lacks the complicated longitudinal folding present in the vagina of the foregoing species. Line ec in the illustrations shows the inner end of the cuticle as observed after maceration; since the line was symmetrical in the specimen, there were probably no parts lost but the cuticle actually ends here. Then, intima-bearing lateral oviducts ol are absent. The paired spermathecae sp open through a pair of longitudinal grooves so on a large unpaired sclerite SO (Figs. 37, 38). The spermathecal openings are located distinctly farther posteriorly than in the foregoing species and thus visible in a ventral view (Fig. 36; this is the reason to regard tube oc+oe as a median oviduct rather than a vagina). The heaviness of sclerite SO decreases from posterior to anterior. The posterior part of SO is overfolded by the membrane behind it (Fig. 36), and the anterior part lies on a low bulge with a median anteriorly produced tip. Immediately in front of SO lies a pair of weak sclerite ribbons AS (Figs. 37, 38). From each spermathecal opening so arise three spermathecal tubes, the anterior of which further divides into three branches (Fig. 38). Each of the resulting five tubes per side is whitish and flexible, hence probably membranous, for most of its length; only the innermost parts, which gradually widen and then abruptly form a globular, easily detaching bulb, are heavily sclerotized. The spermathecal tubes form together a single dense coil. Ventral elements of segment IX. Laterocoxa LC9 extends from tergal extension IX g to the base of gonapophysis gp8, where it articulates with sclerite GP8 (Figs. 36, 38). About midway LC9 articulates posteriorly with sclerite CX9. At this articulation LC9 is notched from anterolaterally and posteromesally. Like the fully separated sclerites in Tagalina, the two portions of LC9 are here called LCa9 and LCp9. Long ridges extend between the LCp9-articulations with TG9 and CX9 and along the mesal part of LCa9. The short, narrow gonoplacs gl9 are entirely but very weakly sclerotized by CX9. Sclerite CX9 is in no way subdivided (compare CXf9 and CXh9 in the foregoing species). A transverse infolding immediately behind the two gonoplacs bears a pair of small tendons gb? (Fig. 38). CX9 lacks a mesal arm i, the left and right CX9 thus being widely separated (Figs. 36, 38). The anterior tip of CX9, strengthened by a short ridge, approaches sclerite LC9 for articulation. The conspicuously heavier anteromesal part of CX9 forms an arm p towards sclerite GPb9 of gonapophysis gp9. The gonapophyses gp9 are relatively large, rounded flaps (Fig. 36). Both together fit tightly into a mould gm formed by the membrane behind their bases. Besides sclerite GPb9 in its basal ventral wall, gonapophysis gp9 has a rod-like sclerite GPa9 in its lateral wall, which corresponds with the GP9 in the foregoing species. A transverse fold ag (Figs. 38, 40) invaginates immediately behind the gp9 bases; it could be a vestige either of the accessory gland ag or of the tube pouch tp, which are otherwise both absent.

17 Entomologische Abhandlungen 61 (2) 189 Figs Anataelia canariensis. Orientation: anterior. Scale in mm. 32: Ventral view of female genitalic region. Right side of figure with lateral parts of terga VIII X and sclerites LP and AP. On left side gonapophysis VIII and gonoplac VIII cut basally. 33: Dorsal view of female genitalic region. Spiracle VIII cut near atrium and simplified. Longer part of spermatheca removed (cut by bars). Internal branchings of accessory gland not retained. Left lateral tube of accessory gland cut near its opening, right tube shown to the extent it could be observed. 34: Dorsal view of anteromedian part of female genitalic region. Left side with further dorsal parts of vagina removed. Tips of gonapophyses VIII cut. 35: Dorsal view of posteromedian part of female genitalic region. Dorsomedian parts of accessory gland and tube pouch removed.

18 190 KLASS: Female genitalia in basal Dermaptera Figs Diplatys macrocephalus. Orientation: anterior. Scale in mm. 36: Ventral view of female genitalic region. Right side of figure with lateral parts of terga VIII X and sclerites LP and AP. On left side ventral wall of gonapophysis VIII and entire gonoplac VIII removed. 37: Ventral view of anteromedian part of female genitalic region, with focus on sclerite of spermathecal opening; gonapophyses VIII almost entirely removed. 38: Dorsal view of female genitalic region. Spiracle VIII cut near atrium and simplified. Left branches of spermatheca removed (cut by bars). 39: Dorsal view of anteromedian part of female genitalic region. On left side further dorsal parts of gonopore area removed. 40: Dorsal view of posteromedian part of female genitalic region.

19 Entomologische Abhandlungen 61 (2) 191 Figs Haplodiplatys orientalis. Orientation: anterior. Scale in mm. 41: Ventral view of female genitalic region. Of terga only lateral tip of tergum IX included. On left side of figure gonapophysis VIII and gonoplac VIII cut basally. 42: Ventral view of anteromedian part of female genitalic region, with focus on spermathecal openings; gonapophyses VIII entirely removed. 43: Dorsal view of female genitalic region. All left branches and some of the right branches of spermatheca removed (cut by bars). Internal branchings of accessory gland mostly not retained. Left lateral tube of accessory gland cut near its opening, right tube shown to the extent it could be observed. 44: Dorsal view of anteromedian part of female genitalic region. Many dorsal parts removed. 45: Dorsal view of posteromedian part of female genitalic region. Dorsal wall of accessory gland removed, lateral tubes cut.

20 192 KLASS: Female genitalia in basal Dermaptera 4.7. Haplodiplatys orientalis Ventral elements of segment VIII. Tendons mt8 and at8 were not found (Fig. 43). Coxa CX8 and the coxal lobe gl8 are, in contrast to Diplatys, strongly developed (Fig. 41). Laterocoxa LC8 is either absent, or possibly fused with sclerite CX8 to form its anterior part. The very short gonapophyses gp8 bear a single ventral sclerotization GP8, which is fused with laterocoxa LC9; a short ridge possibly marks the border. Anteromedially the two gonapophyses gp8 are transversely connected by a wide fold go. Above this fold and above and between the bases of the gp8 lies the entrance into a wide tube oc+oe, which for the same reason as in Diplatys is here considered the common plus the extended oviduct (= median oviduct) rather than a vagina (Figs. 41, 44). The structure of oc+oe could not be clearly observed, but probably, like in Diplatys, it lacks any complicated longitudinal folding and terminates anteriorly with a single wide opening (line ec in Figs. 41, 43, 44: inner end of cuticle), intima-bearing lateral oviducts being absent. The spermathecal openings so are paired like in Diplatys, but are located somewhat farther anteriorly, though still visible ventrally (Fig. 41). The area bearing them is overfolded by the membrane behind it (Figs ). Each opening so leads into a large membranous pouch ps, which bears a small, heavy sclerite SO in its distal ventrolateral wall (Figs. 42, 43). The sclerites of the spermathecal opening area are thus, in contrast to Diplatys, present as a pair. A thin spermathecal tube sp arises from the bottom of each pouch ps and soon divides successively into several moderately long branches; these are brownish and fairly stiff (therefore categorized as sclerotized). Five tubes were counted on the left side, seven on the right side (only three shown in Fig. 43). The internal end of each tube widens and then forms a pear-shaped, sclerotized terminal bulb; the bulb is less sharply than in Diplatys demarcated from its tubes (though the degree varies), and it is not so easily detached. The spermathecal tubes form together a single dense coil, but the left and right portions are easily separated. Ventral elements of segment IX. Laterocoxa LC9 extends from the ventromesal tip of TG9, which lacks a discrete extension g, to the base of gonapophysis gp8, where it is fused to sclerite GP8 (Fig. 41). The mesal part of LC9, which is located on a posteriad-directed fold, is much weaker than the lateral part. About midway LC9 articulates posteriorly with sclerite CX9. A heavy ridge extends between the LC9-articulations with TG9 and CX9. The long, narrow gonoplacs gl9 (Figs. 41, 43, 45) are entirely sclerotized by CX9. Like in Diplatys, CX9 is in no way subdivided, but the sclerite is altogether fairly weak. A low, posteriorly arched transverse fold connects the bases of the two gl9; tendons or an infolding gb (cf. Figs. 11, 28, 33) were not observed. The mesal arm i of CX9 is very short, the left and right CX9 thus being widely separated (Figs. 41, 43). The anterior tip of CX9, strengthened by a very short ridge, approaches sclerite LC9 for articulation. Anteromesal arms of sclerite CX9 (which in Diplatys contact sclerites GPb9) as well as sclerites GPb9 of the gonapophyses gp9 are apparently absent, though it cannot be excluded that both are present in a very weak condition. The gonapophyses gp9 (Figs. 41, 45) are larger and more elongate and pointed than in Diplatys, but like in the latter they are lobe-like and both together fit into a mould gm formed by the membrane behind their bases. Gonapophysis gp9 is partly sclerotized by GPa9, which occupies its lateral wall and much of its dorsal wall (Fig. 45). Like in Diplatys, the mesal edge of the left gp9 is in a ventral view covered by the mesal edge of the right gp9 (Figs. 36, 41). In addition, however, the mesal edge of the left gp9 is sclerotized and held within a groove along the mesal edge of the right gp9. The two gp9 are thus loosely interlocked. They may work as a guide for the secretions of the accessory gland ag and the lateral tubes tl, which, in contrast to Diplatys, are fully developed (Fig. 43). Of the accessory gland some internal branches were observed. The tubes tl originate lateral to the accessory gland, immediately behind the bases of the gp9, without a common tube pouch tp; they are very long but could not be traced entirely (Fig. 43 shows the minimum length) Karschiella buettneri Elements surrounding the genitalic area. Spiracle si8 opens upon even membrane (Fig. 46). The hind margin of TG8 and the front margin of TG9 are connected by a ribbon-like melanization m (Figs. 46, 47), which is lacking in all foregoing taxa. Only TG8 has a distinct mesad-directed extension f, which approaches laterocoxa LC8. TG9 articulates bluntly with laterocoxa LC9. In the tergal ridges, the mesal ends of ac9 and pr9 are connected, and ac10 continues straightly into pr10; pr9 is close to ac10 and lacks a field fs of setae in front of it. TG10 and LP are separate where pr10 crosses to LP. The area of the genital fold gf of venter VII was not preserved in the specimen. Ventral elements of segment VIII. Laterocoxa LC8 is composed of three arms (Figs. 46, 47); two of them extend onto tendons at8, which is located anteromesal to spiracle si8, and mt8, which is located farther mesally; the third closely approaches, like LC8 in Echinosoma and Dacnodes, the anterior margin of tergum TG9. Coxa CX8 is represented by the weak sclerotization of a coxal lobe gl8 (Fig. 46). The gonapophyses gp8 have two sclerites in the ventral wall: the large GP8, which extends all along the gonapophysis, and the small MS at the mesal gonapophyseal base. Ventromedially the two gonapophyses gp8 are transversely connected by a fold go. Dorsal to go the mesal walls of the gonapophyses form asymmetrical longitudinal folds (details in Figs. 46, 48, 49). Above fold go, in between the longitudinal folds, and above the bases of the gp8 lies the entrance into the vagina va (Figs ). The dorsal part of the vagina va is a simple wide sac; the ventral part is divided into several (asymmetrical and communicating) stories by the longitudinal folds. The delicate lateral oviducts ol unite to form a short common plus extended oviduct oc+oe (Figs. 46, 49), which then opens into the ventral part of the vagina. The spermathecae sp, present as one pair, open into the anterior wall of the vagina. Each opening is surrounded by a ringwall of thick, sclerotized cuticle (with sclerites SO; Figs. 48, 50; thickening not considered). The unbranched spermathecal tube (right tube in Fig. 47 complete) is membranous near its opening, strongly sclerotized and stiff in the longer internal part, and widest at midlength, and it has a simple inner end. Each tube forms a coil like that shown in Fig. 47. Ventral elements of segment IX. The strongly angled laterocoxa LC9 extends from TG9 to the base of gonapophysis gp8, where it articulates with sclerite GP8 (Figs. 46, 47). The opposing tips of LC9 and GP8 are thickened. At its posteriormost point LC9 articulates with sclerite CXf9. A short ridge on LC9 extends anteriad from this articulation. The anterolateral corner of LC9 forms a small apodeme at9 (Fig. 47). The wide, short gonoplacs gl9 are entirely but weakly sclerotized by CXh9 (Figs. 46, 47). A transverse fold connects the two gl9 basally. The membrane shortly behind the dorsal base of the gonoplacs bears short tendons gb?. The CXh9 have short mesal arms i upon the transverse fold, which remain widely separated medially (Fig. 46). Anteriorly CXh9 is delimited by a distinct weak line from sclerite CXf9 (Figs. 46, 47). CXf9 closely contacts sclerite GP9 of gonapophysis gp9 with its posteromesal corner, while its anterolateral part, which is strengthened by a thick ridge, articulates with sclerite LC9. The stout, moderately sized gonapophyses gp9 bear sclerite GP9 in their basal lateral wall. Between the bases of the two gp9 opens the acces-

21 Entomologische Abhandlungen 61 (2) 193 Figs Karschiella buettneri. Orientation: anterior. Scale in mm. 46: Ventral view of female genitalic region. Right side of figure with lateral parts of terga VIII X and sclerites LP and AP. On left side gonapophysis VIII and gonoplac VIII cut basally. 47: Dorsal view of female genitalic region. Spiracle VIII cut near atrium and simplified. Of spermatheca right branch complete, left branch largely removed (cut by bar). Internal parts of accessory gland not retained. Left lateral tube of accessory gland cut near its opening, right lateral tube shown to the extent it could be observed. 48, 49: Dorsal view of anteromedian part of female genitalic region. Entire vagina (Fig. 49) or its left half (Fig. 48) removed. 50: Spermathecal opening on wall of vagina. Orientation as in Fig. 48. Dorsal wall of spermathecal bulge partly removed. 51: Dorsal view of posteromedian part of female genitalic region. Dorsal wall of accessory gland removed. Lateral tubes of accessory gland cut. sory gland ag (Figs. 47, 51). Probably two tubes tl (only the right one was found) enter the externalmost dorsal wall of the gland duct, without forming a common pouch tp (the internal parts of ag and tl were not found in the specimen studied) Bormansia monardi The female genitalia in Bormansia closely resemble those in Karschiella. In the single available specimen the sclerotizations were generally not very marked, and a complete dissection was not possible. The differences to Karschiella, as observable under these restrictions, are here described (cuticular ridges and tergal connection VIII-IX m not examined). Laterocoxa LC8, shaped like in Karschiella, is weakened, and thus indistinctly subdivided, in the area marked with an arrowhead in Fig. 47. The coxal lobes gl8 are slightly wider than in Karschiella. The gonapophyses gp8 are basally more slender (not bulging laterally) and have the tips widened, altogether being shaped like a laterad-directed ax. Sclerites MS were not observed. The vagina is shorter and not bilobed anteriorly (the oviducts could not be reliably identified). The spermathecal openings so, paired as in Karschiella, are close together near the midline and located in the anteriormost dorsal wall of the vagina, hence approaching conditions in Haplodiplatys (Fig. 43); they lack sclerotized ringwalls. The dorsal wall of the vagina that receives the spermathecae is bulged dorsally and demarcated

22 194 KLASS: Female genitalia in basal Dermaptera Figs Esphalmenus basidentatus. Orientation: anterior. Scale in mm. 52: Ventral view of female genitalic region. Right side of figure with lateral parts of terga VIII X and sclerites LP and AP. On left side ventral wall and tip of gonapophysis VIII removed. 53: Dorsal view of female genitalic region. Spiracle VIII cut near atrium and simplified. Of spermatheca one branch complete, two branches cut near their openings (by bars). Left lateral tube of accessory gland cut near its opening. 54: Dorsal view of spermathecal bulge and associated elements. Bulge cut along its base. Branches of spermatheca cut near their openings. 55: Dorsal view of anteromedian part of female genitalic region. On left side dorsal parts of common oviduct removed. 56: Dorsal view of posteromedian part of female genitalic region. Lateral tubes of accessory gland cut. from the vaginal chamber below it by lateral longitudinal folds again a condition reminiscent of Haplodiplatys with its pouches ps. The unbranched spermathecal tubes are coiled in a similar manner as in Karschiella, but they are much thinner and longer and less strongly sclerotized. The laterocoxae LC9 are less strongly angled. The gonoplacs gl9 are more broadly rounded than in Karschiella. The mesal arms i of sclerites CXh9 are longer and almost meet each other in the midline. The weak line separating sclerites CXh9 and CXf9 is indistinct, and the two thus almost form a onepiece sclerite CX9 as in the Diplatyinae. The tubes tl, very long and thin with a slightly widened base, and the accessory gland ag have their openings located as in Karschiella Esphalmenus basidentatus Elements surrounding the genitalic area. Spiracle si8 opens on a small lobe (Fig. 52). Similar to Karschiella, the hind margin of TG8 and the front margin of TG9 are connected by a weakly melanized, thickened ribbon m. Only TG8 has a distinct mesad-directed extension f, which distally has no contact (Figs. 52, 53). TG9 articulates bluntly with laterocoxa LC9. In the tergal ridges, the mesal ends of ac9 and pr9 are separated; pr9 obliterates shortly more laterally, and there is no field fs of setae in front it; ac10 continues straightly into pr10. TG10 and LP are separate where pr10 crosses to LP. A noteworthy feature of Esphalmenus is the weak development of tergal immobilization: TG8 is entirely free from TG9, which is, apart from the far lateral fusion e, only in a narrow dorsolateral area attached to TG10. The genital fold gf of venter VII is inconspicuous, wide and short (Fig. 52). Ventral elements of segment VIII. Laterocoxa LC8 and coxa CX8 constitute together a large, heavy sclerite (Figs. 52, 53). The anterior part (interpreted as LC8) forms two arms extending onto tendons at8, which is located anteromesal to spiracle si8, and mt8, which is located farther mesally. The larger posterior part (interpreted as CX8) forms a plate which bears posteriorly the slender coxal lobe gl8. The short gonapophyses gp8 bear two sclerites (Figs. 52, 55): GPa8 in the basal lateral wall, and GPb8 in the distal dorsal wall. A transverse fold is absent between the bases of the gonapophyses gp8 (compare go in, e.g., Fig. 36), but there is a fold go far in front of them (Fig. 52). Above this fold lies the entrance into a wide pouch oc, which is likely a common oviduct (Figs. 53, 55). The two lateral oviducts ol open into its bottom, but otherwise pouch oc shows no further differentiations. The spermathecal opening so is, like in Diplatys, in a far posterior position and thus visible in a ventral view (Fig. 52), but it is unpaired. Three spermathecal tubes sp open one behind the other into a longitudinal median groove that is situated on a low, membranous spermathecal bulge bs (Fig. 54; one tube completely shown in Fig. 53). A triangular membranous tendon-like structure st is seated posteriorly on the groove. Apart from their externalmost parts the tubes are sclerotized (Figs. 53, 54); they have each a simple inner end, and they form together one dense coil.

23 Entomologische Abhandlungen 61 (2) 195 Ventral elements of segment IX. Laterocoxa LC9 extends as a large, even plate from TG9 to the base of gonapophysis gp8, where its thickened mesal tip articulates with sclerite GPa8 (Figs. 52, 53). The posteromesal margin of LC9 is in close vicinity to sclerite CXf9, but there is no discrete articulation. LC9 has a long internal ridge extending from its articulation with TG9 to its mesal margin; since the ridge is, except for its lateralmost part, membranous, it provides a hinge and almost completely divides LC9; the portions of LC9 anterior and posterior to the hinge are called LCa9 resp. LCp9. A minute sclerite lies between the posterior tip of LC9 and the margin of TG10. The gonoplacs gl9 are large and spatulate as in Echinosoma and entirely sclerotized by CXh9. A very short transverse fold connects the two gl9 basally. Though the two sclerites CXh9 lack mesal arms, they are medially in close contact. The dorsal base of each gonoplac bears laterally a long, thin tendon gb (Fig. 53) and near the midline a small, weak sclerite PS9. Anteriorly CXh9 is delimited from sclerite CXf9 by a slight constriction and a weak line (Figs. 52, 53). CXf9, which is strengthened by an internal ridge, has an orientation very different from that in the foregoing species: it extends anteromesad, paralleling the margin of LC9, and its anterior tip is thus not articulated upon LC9. The gonapophyses gp9 are minute membranous flaps (Figs. 52, 56). Immediately posteromesal to their bases open two very long tubes tl (Figs. 53, 56). The accessory gland ag and the tube pouch tp are both absent Pyragra fuscata Elements surrounding the genitalic area. Spiracle si8 opens upon even membrane (Fig. 57). Like in Karschiella, the hind margin of TG8 and the front margin of TG9 are connected by a melanized ribbon m. Terga VIII and IX (not X) as well as plate LP have distinct mesad-directed extensions f, g, n; that of TG8 approaches laterocoxa LC8; that of TG9 articulates with laterocoxa LC9. In the tergal ridges (Figs. 57, 60) the mesal ends of ac9 and pr9 are connected; pr9 is probably largely fused with ac10 (cf. Karschiella, Fig. 47, with these ridges being closely approximate), and the ridge behind them that joins ac10+pr9 in the area e is peculiar to Pyragra. Similar as in Dacnodes, ac10 (more anteriorly ac10+pr9?) is excessively developed as a high, massive fold (Figs. 57, 60, left sides), which anterolaterally continues into the infolded anterior edge of TG10. In front of ac10+pr9 lies a field fs of short setae. Posteriorly ac10 continues straightly into pr10. TG10 and LP are separate where pr10 crosses to LP. The genital fold gf of venter VII is wide and short (Fig. 57), but the infolding pg above it is conspicuously deep, shaped as a wide, simple pouch in P. f. fuscata, and as a narrow, deep, and internally bifid pouch in P. f. brasiliensis (pouches shown to equal scale in Figs. 58, 59). Ventral elements of segment VIII. Laterocoxa LC8 forms two sclerites (Figs. 57, 60): The triangular lateral one hinges posteriorly upon the anterior margin of tergum TG9 and extends anteriorly onto tendon at8; the disc-shaped mesal one lies behind the base of tendon mt8. A very short lobe with a mesal prominence represents a vestigial coxal lobe gl8, but a coxal sclerotization CX8 is lacking (Fig. 57). The gonapophyses gp8 bear one sclerite GP8, which occupies the entire ventral wall and forms two oval cuticular thickenings anterolaterally. Medially the bases of the two gonapophyses gp8 are transversely connected by a wide fold go. Above this fold and between the bases of the gp8 lies the entrance into what is here considered an extended oviduct oe (Figs. 57, 60, 61). It divides internally into very delicate, paired dorsolateral sacs and an unpaired ventromedian sac. For the former it has remained unclear whether they are anteriorly closed by cuticle or open-ended (i.e., with mesodermal elements adjoining along line ec? in Figs. 60, 61), though a closed condition appears more likely. The ventromedian sac bears dorsally a longitudinal groove dg (seen as a ridge in the internal view in Fig. 61) and receives anteriorly an open-ended (along line ec in Fig. 61) tube. The latter tube is likely the common oviduct oc, cuticulized lateral oviducts ol then being absent or not preserved in the specimens. It is an unlikely alternative that the dorsolateral sacs are the lateral oviducts; this would leave the interpretation of the median tube unresolved. The membrane behind the dorsal wall of the extended oviduct has a complicated architecture related to the spermathecal opening so (Figs. 57, 60, 61): The unpaired spermatheca opens immediately behind the posterior end of groove dg outside the extended oviduct and hence visible in a ventral view (so in Fig. 57). From the orifice so a membranous groove mg winds posteriad and then forks into lateral grooves lg, which laterally curve back anteriad. Some membranous folds run behind and alongside the folds lg (details in Fig. 61), one of them bearing a conspicuous setation (fine dots in Fig. 61). The spermatheca sp is a very long (at least 18 mm), unbranched tube with a slightly widened, membranous external portion, a narrower, sclerotized internal portion, and a simple inner end (Fig. 60). It forms one dense coil. Ventral elements of segment IX. Laterocoxa LC9 extends from tergal projection IX g to the base of gonapophysis gp8, where it articulates with sclerite GP8 (Figs. 57, 60). A long L-shaped ridge extends between the two articulations of LC9. The gonoplacs gl9 are small membranous flaps transversely connected by a wide fold. Coxal sclerites CX9 as well as tendons gb on the dorsal gonoplac base are absent. The two gonapophyses IX are probably represented by the minute, bilobed process gp9 (Fig. 57), which is associated with a very peculiar internal cuticular structure (tp(+ag?) in Fig. 62): A short duct with lateral sclerotizations enters internally an oval bulb and protrudes somewhat into it, the lateral sclerotizations forming prominent bulges. On its posterodorsal face the bulb receives two long, narrow tubes tl (Figs. 60, 62). The bulb and its outlet duct are interpreted here as the internal and external portions of a highly elaborate tube pouch tp, though parts of the accessory gland ag, which otherwise is entirely absent, may have contributed. Unfortunately, the exact position of the orifice on process gp9 could not be observed. 5. Discussion of female genitalic region 5.1. The basic design and subsets of female genitalia in Insecta Generalities. Like in the Pygidicranidae described above, the female genitalic region in many Insecta (= Ectognatha) displays a complex composition of a variety of formative elements, sclerites, and muscles, and of the gonoducts, spermathecae, and accessory glands, which belong to venters VIII and IX and to the posterior part of venter VII. There is a shared basic pattern of the female genitalic region in the Archaeognatha, the Zygentoma, and the various lineages of the Pterygota, which, for instance, includes the lengthening of the eversible vesicles VIII and IX in order to form mutually interlocked gonapophyses and an egg channel in between. Such an ovipositor built of gonapophyses is likely autapomorphic for the Insecta (e.g., KRISTENSEN 1991: 129), though possible vestiges have also been reported for Diplura (GRASSI 1888: figs. 50, 52; MARTEN 1939; GUSTAFSON 1950: 53). However, the pattern shared within the Insecta also includes specifities in the many other, less conspicuous components of the female genitalic region. The high complexity of this body area, the presence of a common pattern in Insecta, and the additional occurrence of much structural modification renders the

24 196 KLASS: Female genitalia in basal Dermaptera Figs Pyragra fuscata (mostly ssp. fuscata). Orientation: Figs. 57, 60, 61 anterior, Figs. 58, 59 posterior, Fig. 62 posteroventral. Scale in mm. 57: Ventral view of female genitalic region. Right side of figure with lateral parts of terga VIII X and sclerites LP and AP. On left side dorsal wall and tip of gonapophysis VIII removed, gonoplac VIII cut, the excessively developed ridge pr9+ac10 retained. 58: Dorsal view of genital pouch of Pyragra fuscata fuscata (compare gf in Fig. 57). 59: Dorsal view of genital pouch of Pyragra fuscata brasiliensis. 60: Dorsal view of female genitalic region. Spiracle VIII cut near atrium and simplified. Longer part of spermatheca removed (cut by bars). Left lateral tube of accessory gland cut near its opening. 61: Dorsal view of anteromedian part of female genitalic region. On left side further dorsal parts of oviduct removed and spermathecal area cut longitudinally. 62: Posterodorsal view of accessory gland area. Window cut into dorsal wall of tube pouch. female genitalia a very rich character system for phylogeny reconstruction. For exploiting the female genitalic region as a source of phylogenetically informative characters, one important step is to analyse the topographic homologies of the various elements among the different insect lineages, i.e., to find corresponding parts that are legitimately assessed within the same character (KLASS 2001b: 230). Only afterwards different conditions of corresponding parts can be defined as different character states and scored throughout the taxa. An analysis of the female genitalic region in the Pterygota and its subgroups, such as the pygidicranid Dermaptera, has to be based on conditions in Archaeognatha and Zygentoma. As a part of this work KLASS (1998, 2001a, in press)