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Mendeschilis a new genus of Microcoryphia (Insecta, Apterygota, Machilidae) from Spain
Pedobiologia 44, 257–267 (2000) © Urban & Fischer Verlag http://www.urbanfischer.de/journals/pedo
PROCEEDINGS OF VTH INTERNATIONAL SEMINAR ON APTERYGOTA, CORDOBA 1998
Mendeschilis a new genus of Microcoryphia (Insecta, Apterygota, Machilidae) from Spain Miguel Gaju-Ricart1, Rosa Ma Mora-Carmona1, Rafael Molero-Baltanás1 and Carmen Bach de Roca2 1 Department of Animal Biology (Zoology), University of Córdoba, C-1 Campus Rabanales, N-IV Km 396A, 14014 Córdoba, Spain 2 Department of Animal Biology, Botany and Ecology, Autonomous University of Barcelona, 08193 Bellaterra, Barcelona, Spain
Accepted: 30. December 1999
Summary A new species and genus of Microcoryphia is described as a result of the detailed comparison of specimens from Italy and Spain which had been initially identified as Praemachilis excelsior. The new genus, named Mendeschilis has as its most important distinctive taxonomic characteristic the presence, in the male, of parameres in the VIIIth and IXth urosternites, while Praemachilis (from Italy) has only one pair of parameres in the IXth urosternite. Other anatomical characteristics which allow us to differentiate the new genus and the new species are described. Key words: Mendeschilis escorcai, new genus, new species, Praemachilis excelsior, Majorca, Spain
Introduction In 1904, Silvestri described the genus Praemachilis defined by, among other characteristics, the presence of parameres in the VIIIth and IXth urosternites. As a type species of the genus he described P. excelsior, but only the female. Later, Janetschek (1954) studied the cotypes of Silvestri (only females). In the genus Praemachilis, SilCorresponding author Miguel Gaju-Ricart, e-mail: [email protected]
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vestri included other species, which have since been placed in the genera, Praetrigoniophthalmus and Dilta; these possess parameres only in the IXth or in the VIIIth and IXth urosternites respectively. We cannot confirm that Silvestri really observed parameres in urosternite VIIIth in Praemachilis, because we have not found any male specimens in his material. We studied the cotype material of Silvestri (only females) from the Natural History Museum of Portici (Italy), female specimens collected by us in Pontignano (Siena, Italy), and specimens obtained from to the Museum of Natural History of Verona; the latter collection included a juvenile male of P. excelsior with parameres only in the IXth urosternite. Paclt (1969) reported the presence of P. excelsior from Majorca (Spain) finding two males. Later, Mendes (1981) studying a collection from Majorca, found some specimens which agree with the description given by Silvestri for P. excelsior, and described a male which has parameres in the VIIIth and IXth urosternites. We have studied further specimens from Majorca which has allowed us to compare Italian material (comprising only one male but several females) with Spanish (Majorcan) specimens. We observed morphological differences which prompted us to retain the specimens from Italy as P. excelsior but to create a new genus and a new species for the specimens from Majorca. These we have named Mendeschilis escorcai n.g., n. sp. Mendeschilis n. g. Description: Adult body length: 11-13 mm. Antennae longer than the body, distal chains (Figs. 4 and 20) with many annuli (15 to 33). Flagellum of antennae with scales. Compound eyes slightly wider than long (Fig. 3), paired ocelli large, rounded, submedian in position (Figs. 1–2). Frons swollen between the paired ocelli. Mandibles with four apical teeth. Male maxillary palp modified in shape and chaetotaxy (Fig. 6). Thorax normal, the nota with usual development. Coxal stylets present on legs II and III. Fore femur of male not swollen. Legs without spiniform setae (Fig. 14). Tarsal scopulae absent. Sternites acutely angled, setae present on the coxites (Figs. 15 and 27). 1+1 eversible vesicles present in urosternites I–VII. Parameres on the VIIIth and IXth urosternites, clearly pseudosegmented (Figs. 16 and 18). Male genitalia covered by the IXth coxites (Fig. 18). Penis opening small, apical. Ovipositor (Figs. 29–34) of the tertiary type (Sturm & Bach 1993). Terminal filaments with piliform scales. Type species: Mendeschilis escorcai n. sp. Mendeschilis escorcai n. sp. Studied material Balearic Islands: Pollença, Cabo Formentor (Majorca), 30-05-90, 1 male + 2 females + 3 juveniles, Ref. M0352; Pollença, Puerto de Pollença (Majorca), 30-05-90, 1 female, Ref. M0356; Ferreries, Cala Galdana (Menorca), 05-06-90, 4 females, Ref. M0387; Escorca, Torrent de Pareis, Sa Calobra (Majorca), 27-09-91, 1 male, Ref. M0422; Escorca, Sa Calobra (Majorca), 27-09-91, 1 male holotype + 1 female allotype + 1 male + 5 females paratypes, Ref. M0427; Escorca, Sa Calobra (Majorca), 15-05-92, 1 male, Ref. M0451; Escorca, Sa Calobra (Majorca), 15-09-95, 1 male, Ref. M1522.
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Figs. 1–14: Mendeschilis escorcai n.g., n.sp., male. 1. Head, frontal view; 2. Head, lateral view; 3. Outline of the compound eyes and ocelli; 4. Outline of a distal antenna chain; 5. Three apical annuli of the antenna chain; 6. Outline of the maxillary palp; 7. Ciliary setae of the 4th segment of the maxillary palp; 8. Apical part of the 7th segment of the maxillary palp; 9. Outline of the labial palp; 10. Sensory connules of the 3rd segment of the labial palp; 11. Outline of the fore leg; 12. Ditto, mid leg; 13. Ditto, hind leg; 14. Chaetotaxy of the tarsi of the fore leg. Scale: 0.1 mm
The holotype male, allotype female and paratypes were deposited in the Museo Nacional de Ciencias Naturales in Madrid; collection number: 8526. Description of the male Body length: 11 mm, length of antennae (broken): 13 mm, length of paracercus: 13 mm, length of cercus: 5 mm. Head with scarce pigment, as shown in Figs. 1 and 2. The paired ocelli are rounded and located in a submedian position. The eyes are slightly larger than long (Fig. 3), ratio eyes contact line / eyes length (cl/l): 0.35; ratio eyes length / width (l/w): 0.86.
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The antennae are longer than the body, in well preserved specimens the most distal chains have 29 or more annuli, which are twice as wide as long (Figs. 4–5). The maxillary palp (Fig. 6) has weak and diffuse pigment. The external apophysis (processus triangularis) of the first segment is conical in shape and slightly back curved, the internal one is smaller and also conical (Figs. 6 and 38). The fourth segment is short, approximately half the length of the fifth. The seventh segment is conical in shape. Length ratio of last to penultimate segments (n/n-1): 0.62. Ciliary setae present on the ventral side on the second to fifth segments (Figs. 6, 7). Hyaline spines (Fig. 8) on the dorsal side of the last three segments of the maxillary palp, with the following distribution: segment 5: 2 spines, 6: 14 and 7: 13. The labial palp is without pigment, the second segment with subparallel sides, 3rd segment enlarged, semicircular in shape; with numerous sensory connules occupying the apex of this segment (Figs. 9–10). The three pairs of legs with weak and diffuse pigment. Femur of the fore leg not swollen (Fig. 11), ratio of length to width of the femur: 2.1. Mid leg (Fig. 12) and hind leg (Fig. 13) with coxal stylets. Ventral side of the legs with numerous strong setae (Fig. 14). Length of tibiae I: 0.91 mm; II: 0.80 mm; III: 1.15 mm. Urosternites II–VII with acutely angled sternite; the coxites I–VII with only a pair of eversible vesicles and sometimes also with strong setae (Fig. 15). Table 1 shows the length ratios between stylet (without apical spine) to coxite and stylet apical spine to stylet. The VIIIth urosternite (Fig. 16) has parameres with 1+5 divisions, which are weakly marked, the distal divisions with tubular setae (Fig. 17). IXth urosternite (Fig. 18) with parameres and penis, the former with 1+5 divisions, all except the first of which have tubular setae (Fig. 19); the penis is between the parameres and exceeds them in length. The penis basal (bp) and distal parts (dp) are almost similar in length; ratio bp/dp: 0.89. Cerci with two apical spines. Description of the female Body length: 13 mm, length of antennae: 15 mm, length of paracercus: 9 mm (broken), length of cercus: 5.1 mm. Head as in male. Ratios cl/l: 0.46; l/w: 0.87.
Table 1. Mendeschilis escorcai n.g., n. sp. Ratios between coxite, stylet (without spine) and spine of the stylet in the urosternites V, VIII and IX
Urosternite V
male female Urosternite VIII male female Urosternite IX male female
Stylet (without spine)/Coxite 0.56 0.51 0.79 0.91 1.19 0.89
Spine/Stylet (without spine) 0.30 0.32 0.25 0.16 0.15 0.11
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Figs. 15–23. Mendeschilis escorcai n.g., n. sp., male. 15. Outline of the V urosternite; 16. Urosternite VIII and parameres; 17. Chaetotaxy of the VIIIth parameres; 18. IXth urosternite with parameres and penis; 19. Chaetotaxy of the IXth parameres. M. escorcai n.g., n. sp., female. 20. Outline of a distal antenna chain; 21. Three apical annuli of the antenna chain; 22. Outline of the maxillary palp; 23. Ditto, of the labial palp. Scale: 0.1 mm
The antennae are less robust than in the male, but also longer than the body. In well preserved antennae the distal chains have 23 annuli; each approximately equal in length and width (Figs. 20–21). The maxillary palp (Fig. 22), has weak and diffuse pigment, and lacks the ciliary setae which are present in the male palps. Ratio n/n-1: 0.7. The labial palp is without pigment (Fig. 23). The third segment is smaller than in the male. Fewer sensory connules are present than in the male. The three pairs of legs are robust with weak and diffuse pigment (Figs 24–26). The ventral side of the legs with strong setae as in the male. Length of the tibiae: I: 0.86 mm, II: 0.74 mm, III: 1.01 mm.
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Figs. 24–29. Mendeschilis scorcai n.g., n.sp., female. 24. Outline of fore leg; 25. Ditto, mid leg; 26. Ditto, hind leg; 27. Outline of the Vth urosternite; 28. Ditto, of the VIIth; 29. Ditto, of the VIIIth with gonapophysis. Scale: 0.1 mm
Urosternites II–VII with acutely angle sternites; coxites, sometimes with strong setae (Figs. 27–30). Table 1 shows the length ratios between coxite/stylet (without spine)/spine of the stylet. The female has a tertiary type ovipositor that exceeds the IXth stylet in shorter length than the length of the stylet. Gonapophysis VIII with 1+61 divisions, all except the more basal divisions with setae. The chaetotaxy is shown in figures 31–32. Gonapophysis IX with 1+61 divisions; its chaetotaxy is shown in figures 33–34. Apex of the cerci as in the male.
Discussion Before the analysis of the new genus, some information is provided about Praemachilis excelsior Silvestri, 1904, a rarely found species. P. excelsior was first described from Bevagna and Portici (Italy) by Silvestri (1904), and later cited in Monaco and Menton in Southeastern France (Silvestri 1907).
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Figs. 30–34. Mendeschilis escorcai n.g., n.sp., female. 30. IXth urosternite with gonapophysis; 31. Divisions 25 to 30 of the VIIIth gonapophysis; 32. Apical divisions of the VIIIth gonapophysis; 33. Divisions 17 to 21 of the IXth gonapophysis; 34. Apical divisions of the IXth gonapophysis. Scale: 0.1 mm
Originally only the female was described, but the male characteristics (exclusively genitalia) were included in the generic definition and the identification keys only. Janetschek (1954) redescribed the genus and the species using the “cotypes” from Silvestri (only females). Paclt (1969) observed P. excelsior males in Majorca and Mendes (1981) made the formal description of male characteristics. Now we consider that these Balearic specimens do not belong to P. excelsior but to a new species and to a new genus Mendeschilis escorcai n. g., n. sp. To clarify the correct taxonomic positioning of each taxon, some available samples which had been identified as P. excelsior were obtained from Italy, using material held in Natural History Museum of Portici (NHMP), the Museum of Natural History of Verona (MNHV) and the University of Córdoba (UCO): Bevagna, 1 female (cotypus labelled), no date, Silvestri det., (Microscopic slide), NHMP. Portici, 1 female (cotypus labelled), no date, Silvestri det., (Microscopic slide), NHMP.
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Fig. 35–45. 35. Praemachilis excelsior Silvestri, 1904:1st segment of the maxillary palp showing the apophyses, female specimen (cotypus) from Portici (Italy); 36. Ditto, female specimen (cotypus) from Bevagna (Italy); 37. Ditto, male specimen from Santa Eufemia (Aspromonte, Italy); 38. Mesdeschilis escorcai n.g. n. sp. from Majorca (Spain); 39. Mendeschilis sp. from Marettimo (Egadi Islands, Italy); 40. Praemachilis excelsior Silvestri, 1904, from Pontignano (Siena, Italy) Head: frontal view; 41. Head, lateral view; 42. Mendeschilis sp. from Marettimo (Egadi Is., Italy) Head: frontal view; 43. Head, lateral view; 44. Praemachilis excelsior Silvestri, 1904, Urosternite VIII of the male; 45. Ditto, Urosternite IX with the penis and parameres. Scale: 0.1 mm
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Portici, 6 females (labelled as cotypi!!) Giugno 1907, Silvestri det., (Alcohol), NHMP. (This specimens are erroneously labelled as “cotipes”, because the species was described in 1904). Santa Eufemia (Aspromonte), 1 male, 22-7-1957, Bach det. 1983, MNHV. Marettimo (Egadi Is.), 1 female, 23-10-1967, Bach det. 1983, MNHV. Pontignano (Siena), 5 females, 19-10-1995, Gaju det. UCO. (new material) The revision of these samples allows to state the similarities among the Italian specimens, all of which except the female from Marettimo belong to P. excelsior. It was very important to collect and study live specimens from Pontignano, because their facies was very characteristic, and our revision of the Silvestri’s original description suggests they belong to P. excelsior. The specimens show a slightly humped mesonotum, which fits Silvestri’s description “Gibba thoracica magna …”, the pattern of scales in the dorsal part of the body was also consistent with Silvestri’s description with a transverse black band in the middle flanked by white bands (“... tergito tertio nigro-velutino, albo marginato, tergitis 4° et 5° albis …”). Also, the antenna show a characteristic white band in their middle part, recorded by Silvestri as (“… partem medianam versus albo”). Studies on the Silvestri’s specimens and the other material from Italy show that the main morphological characteristics were consistent but some slight differences were observed which could reflect the diverse range of mounting media used (Canada Balsam, Hoyer or Tendeiro liquid). The apophysis of the first maxillary palp segment may be used to taxonomically distinguish the two genera (Figs. 35–37); in all cases in Praemachilis the external apophysis is large, conical, slightly curved back and rounded at the apex, the internal apophysis is short, conical and rounded at the apex. Comparing the same segment of the new genus, the outer apophysis, although similar, is slightly thinner with a pointed ending, and the inner one is clearly conical, also with a pointed ending (Fig. 38). Another distinguishing feature of the two genera is the shape of paired ocelli; they are oval in P. excelsior (Figs. 40–41) (see also drawings of Janetschek 1954, p. 210), but rounded in M. escorcai (Figs. 1–3). Curiously the female from Marettimo (Egadi Islands) studied by Bach de Roca (1983) must belong to Mendeschilis genus because of the last two characteristics (Figs. 39 and 42–43). In relation to the male characteristics, as the specimens referred to by Silvestri have not been found, we suppose they could belong to another species and genus, because the original Praemachilis genus include, in addition to P. excelsior, the following species: P. meticulosa, (now = Praetrigoniophthalmus meticulosa (Silvestri 1904)) whose male has parameres in the IXth urosternite only; P. italica (now = Dilta italica (Grassi, 1887)), whose male has parameres in the VIIIth and IXth urosternites and P. aurea (Giardina 1900) a species insufficiently described and considered by Janetschek (1954) as not valid until new material provides a new diagnosis. So, in the same genus Silvestri included species with different numbers of parameres and which today are recognised as belonging to three different genera. Study of the specimen from Aspromonte (Santa Eufemia) suggests that it belongs to the genus Praemachilis according to the characteristics of its maxillary palps (Fig. 37). Unfortunately the specimen is not completely adult and the part preserved in alcohol was dried and its head characteristics are not clearly visible. The VIIIth and
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IXth urosternites with the genitalia are shown in figures 44–45. The degree of development of the penis and IXth parameres suggests that the VIIIth parameres would be perfectly visible if they existed, in spite of being a subadult; so we believe this specimen did not possess any VIIIth paramera. We therefore suggest that members of the genus Praemachilis only posses parameres in the IXth urosternites. In Machilis Latreille, 1832, there are many species with parameres in the VIIIth and IXth urosternites, but only two species have parameres only in the IXth urosternite; for these two species the subgenus Pseudomachilis was created (Janetscheck 1955), based only upon the number of parameres. In our case, although the number of parameres is one difference between Praemachilis and Mendeschilis, other morphological differences exist which permit both genera to be distinguished (Table 2). Finally, we have two additional remarks concerning P. excelsior. The first is to suggest that probably the species can reproduce partenogenetically in some places, because of the low number of males found; in fact, in Pontignano we have found more than 15 specimens and all were females. The second point is to suggest the possible synonimy between the genera Praemachilis and Wygodzinskylis Janetschek, 1954, because comparison of the studied specimens of P. excelsior with the description of Wygodzinskilis klinocellata Janetschek (1954) a further redescription (Janetschek 1959), and two specimens from L.F. Mendes’ Collection, yielded no obvious differences between these genera (as discussed above, we consider that both genera have parameres only in the IXth urosternite). Other potential differences between the genera (such as the shape of the apophysis of the first maxillary palp segment and the characteristics of the antennae) were not evident. However, until new and abundant material can be studied, we prefer to maintain the status of both genera.
Table 2. Differences between Mendeschilis n. g. and Praemachilis Silvestri, 1904 Characteristics
Mendeschilis n.g.
Body shape Ocelli
Mesonotum rounded Rounded
Antennae Distal antennae chains First segment of the maxillary palp: External apophysis Ditto, internal apophysis Legs (ventral side of tibia and tarsi) Coxites of urosternites Parameters Ratio bp/dp of penis
Longer than the body Until 30 divisions Conical and apically pointed (Figs. 6 and 38)
Praemachilis Silvestri,1904 Slightly humped Oval (twice as long as wide) Shorter Usually 9 divisions Conical to Subcilindrical and apically rounded
Pointed (Figs. 6 and 38) With strong setae
Rounded (Figs. 35–37) With spiniform satae
With some setae VIII and IX 0.89
Without setae Only IX (Fig. 44–45) 1.5
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Derivation of the name The new genus is named Mendeschilis and is dedicated to our colleague and friend Luis F. Mendes from Lisbon (Portugal). The species name is taken from the site of sampling in Escorca (Majorca, Spain).
Acknowledgements This work had been supported by the Project “Fauna Ibérica III” SEUI-DGICYT PB92-0121. The authors are deeply grateful to the Museum of Natural History of Verona and the Natural History Museum of Portici (Italy) for sending us the specimens for this study, as well as to Dr. Geoff Frampton for improving English in the manuscript.
References Bach de Roca, C. (1983) Contribución al conocimiento de los Microcoryphia de Italia (Insecta, Apterygota). Bollettino del Museo Civico di Storia Naturale di Verona, [1982], 523–629. Janetschek, H. (1954) Ueber Felsenspringer der Mittelmeerländer (Thysanura, Machilidae). Eos 30, 163–314. Janetschek, H. (1955) Felsenspringer aus Sardinien, Korsika und den Ostpyrenäen (Ins., Thysanura). Österreichische Zoologische Zeitschrift 5, 407–441. Janetschek, H. (1959) Weitere Machilida aus dem Balkan. Acta Musei Macedonici Scientiarum Naturalium 6 (58), 120–141. Mendes, L.F. (1981) Contribution à la connaissance des Machilides (Microcoryphia: Apterygota) de l’Ile de Majorque, avec description de deux nouvelles espèces. Revue Suisse de Zoologie 88(2), 413–432. Paclt, J. (1969) Neue Beiträge zur Kenntnis der Apterygoten-Sammlung des Zoologischen Staatsinstituts und Zoologischen Museums Hamburg. III. Meinertellidae und Machilidae (Thysanura). Entomologische Mitteilungen aus dem Zoologischen Staatsinstitut und Zoologischen Museum Hamburg 3(63), 269–272. Silvestri, F. (1904) Nuovi generi e specie di Machilidae. Redia 2, 4–9. Silvestri, F. (1907) Catalogue des Machilidae de la collection du Muséum. Note sui Machilidae. Redia 3, 32–34. Sturm, H., Bach de Roca, C. (1993) On the Systematics of the Archaeognatha (Insecta). Entomologia Generalis 18(1/2), 55–90.
PROCEEDINGS OF VTH INTERNATIONAL SEMINAR ON APTERYGOTA, CORDOBA 1998
Mendeschilis a new genus of Microcoryphia (Insecta, Apterygota, Machilidae) from Spain Miguel Gaju-Ricart1, Rosa Ma Mora-Carmona1, Rafael Molero-Baltanás1 and Carmen Bach de Roca2 1 Department of Animal Biology (Zoology), University of Córdoba, C-1 Campus Rabanales, N-IV Km 396A, 14014 Córdoba, Spain 2 Department of Animal Biology, Botany and Ecology, Autonomous University of Barcelona, 08193 Bellaterra, Barcelona, Spain
Accepted: 30. December 1999
Summary A new species and genus of Microcoryphia is described as a result of the detailed comparison of specimens from Italy and Spain which had been initially identified as Praemachilis excelsior. The new genus, named Mendeschilis has as its most important distinctive taxonomic characteristic the presence, in the male, of parameres in the VIIIth and IXth urosternites, while Praemachilis (from Italy) has only one pair of parameres in the IXth urosternite. Other anatomical characteristics which allow us to differentiate the new genus and the new species are described. Key words: Mendeschilis escorcai, new genus, new species, Praemachilis excelsior, Majorca, Spain
Introduction In 1904, Silvestri described the genus Praemachilis defined by, among other characteristics, the presence of parameres in the VIIIth and IXth urosternites. As a type species of the genus he described P. excelsior, but only the female. Later, Janetschek (1954) studied the cotypes of Silvestri (only females). In the genus Praemachilis, SilCorresponding author Miguel Gaju-Ricart, e-mail: [email protected]
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vestri included other species, which have since been placed in the genera, Praetrigoniophthalmus and Dilta; these possess parameres only in the IXth or in the VIIIth and IXth urosternites respectively. We cannot confirm that Silvestri really observed parameres in urosternite VIIIth in Praemachilis, because we have not found any male specimens in his material. We studied the cotype material of Silvestri (only females) from the Natural History Museum of Portici (Italy), female specimens collected by us in Pontignano (Siena, Italy), and specimens obtained from to the Museum of Natural History of Verona; the latter collection included a juvenile male of P. excelsior with parameres only in the IXth urosternite. Paclt (1969) reported the presence of P. excelsior from Majorca (Spain) finding two males. Later, Mendes (1981) studying a collection from Majorca, found some specimens which agree with the description given by Silvestri for P. excelsior, and described a male which has parameres in the VIIIth and IXth urosternites. We have studied further specimens from Majorca which has allowed us to compare Italian material (comprising only one male but several females) with Spanish (Majorcan) specimens. We observed morphological differences which prompted us to retain the specimens from Italy as P. excelsior but to create a new genus and a new species for the specimens from Majorca. These we have named Mendeschilis escorcai n.g., n. sp. Mendeschilis n. g. Description: Adult body length: 11-13 mm. Antennae longer than the body, distal chains (Figs. 4 and 20) with many annuli (15 to 33). Flagellum of antennae with scales. Compound eyes slightly wider than long (Fig. 3), paired ocelli large, rounded, submedian in position (Figs. 1–2). Frons swollen between the paired ocelli. Mandibles with four apical teeth. Male maxillary palp modified in shape and chaetotaxy (Fig. 6). Thorax normal, the nota with usual development. Coxal stylets present on legs II and III. Fore femur of male not swollen. Legs without spiniform setae (Fig. 14). Tarsal scopulae absent. Sternites acutely angled, setae present on the coxites (Figs. 15 and 27). 1+1 eversible vesicles present in urosternites I–VII. Parameres on the VIIIth and IXth urosternites, clearly pseudosegmented (Figs. 16 and 18). Male genitalia covered by the IXth coxites (Fig. 18). Penis opening small, apical. Ovipositor (Figs. 29–34) of the tertiary type (Sturm & Bach 1993). Terminal filaments with piliform scales. Type species: Mendeschilis escorcai n. sp. Mendeschilis escorcai n. sp. Studied material Balearic Islands: Pollença, Cabo Formentor (Majorca), 30-05-90, 1 male + 2 females + 3 juveniles, Ref. M0352; Pollença, Puerto de Pollença (Majorca), 30-05-90, 1 female, Ref. M0356; Ferreries, Cala Galdana (Menorca), 05-06-90, 4 females, Ref. M0387; Escorca, Torrent de Pareis, Sa Calobra (Majorca), 27-09-91, 1 male, Ref. M0422; Escorca, Sa Calobra (Majorca), 27-09-91, 1 male holotype + 1 female allotype + 1 male + 5 females paratypes, Ref. M0427; Escorca, Sa Calobra (Majorca), 15-05-92, 1 male, Ref. M0451; Escorca, Sa Calobra (Majorca), 15-09-95, 1 male, Ref. M1522.
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Figs. 1–14: Mendeschilis escorcai n.g., n.sp., male. 1. Head, frontal view; 2. Head, lateral view; 3. Outline of the compound eyes and ocelli; 4. Outline of a distal antenna chain; 5. Three apical annuli of the antenna chain; 6. Outline of the maxillary palp; 7. Ciliary setae of the 4th segment of the maxillary palp; 8. Apical part of the 7th segment of the maxillary palp; 9. Outline of the labial palp; 10. Sensory connules of the 3rd segment of the labial palp; 11. Outline of the fore leg; 12. Ditto, mid leg; 13. Ditto, hind leg; 14. Chaetotaxy of the tarsi of the fore leg. Scale: 0.1 mm
The holotype male, allotype female and paratypes were deposited in the Museo Nacional de Ciencias Naturales in Madrid; collection number: 8526. Description of the male Body length: 11 mm, length of antennae (broken): 13 mm, length of paracercus: 13 mm, length of cercus: 5 mm. Head with scarce pigment, as shown in Figs. 1 and 2. The paired ocelli are rounded and located in a submedian position. The eyes are slightly larger than long (Fig. 3), ratio eyes contact line / eyes length (cl/l): 0.35; ratio eyes length / width (l/w): 0.86.
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The antennae are longer than the body, in well preserved specimens the most distal chains have 29 or more annuli, which are twice as wide as long (Figs. 4–5). The maxillary palp (Fig. 6) has weak and diffuse pigment. The external apophysis (processus triangularis) of the first segment is conical in shape and slightly back curved, the internal one is smaller and also conical (Figs. 6 and 38). The fourth segment is short, approximately half the length of the fifth. The seventh segment is conical in shape. Length ratio of last to penultimate segments (n/n-1): 0.62. Ciliary setae present on the ventral side on the second to fifth segments (Figs. 6, 7). Hyaline spines (Fig. 8) on the dorsal side of the last three segments of the maxillary palp, with the following distribution: segment 5: 2 spines, 6: 14 and 7: 13. The labial palp is without pigment, the second segment with subparallel sides, 3rd segment enlarged, semicircular in shape; with numerous sensory connules occupying the apex of this segment (Figs. 9–10). The three pairs of legs with weak and diffuse pigment. Femur of the fore leg not swollen (Fig. 11), ratio of length to width of the femur: 2.1. Mid leg (Fig. 12) and hind leg (Fig. 13) with coxal stylets. Ventral side of the legs with numerous strong setae (Fig. 14). Length of tibiae I: 0.91 mm; II: 0.80 mm; III: 1.15 mm. Urosternites II–VII with acutely angled sternite; the coxites I–VII with only a pair of eversible vesicles and sometimes also with strong setae (Fig. 15). Table 1 shows the length ratios between stylet (without apical spine) to coxite and stylet apical spine to stylet. The VIIIth urosternite (Fig. 16) has parameres with 1+5 divisions, which are weakly marked, the distal divisions with tubular setae (Fig. 17). IXth urosternite (Fig. 18) with parameres and penis, the former with 1+5 divisions, all except the first of which have tubular setae (Fig. 19); the penis is between the parameres and exceeds them in length. The penis basal (bp) and distal parts (dp) are almost similar in length; ratio bp/dp: 0.89. Cerci with two apical spines. Description of the female Body length: 13 mm, length of antennae: 15 mm, length of paracercus: 9 mm (broken), length of cercus: 5.1 mm. Head as in male. Ratios cl/l: 0.46; l/w: 0.87.
Table 1. Mendeschilis escorcai n.g., n. sp. Ratios between coxite, stylet (without spine) and spine of the stylet in the urosternites V, VIII and IX
Urosternite V
male female Urosternite VIII male female Urosternite IX male female
Stylet (without spine)/Coxite 0.56 0.51 0.79 0.91 1.19 0.89
Spine/Stylet (without spine) 0.30 0.32 0.25 0.16 0.15 0.11
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Figs. 15–23. Mendeschilis escorcai n.g., n. sp., male. 15. Outline of the V urosternite; 16. Urosternite VIII and parameres; 17. Chaetotaxy of the VIIIth parameres; 18. IXth urosternite with parameres and penis; 19. Chaetotaxy of the IXth parameres. M. escorcai n.g., n. sp., female. 20. Outline of a distal antenna chain; 21. Three apical annuli of the antenna chain; 22. Outline of the maxillary palp; 23. Ditto, of the labial palp. Scale: 0.1 mm
The antennae are less robust than in the male, but also longer than the body. In well preserved antennae the distal chains have 23 annuli; each approximately equal in length and width (Figs. 20–21). The maxillary palp (Fig. 22), has weak and diffuse pigment, and lacks the ciliary setae which are present in the male palps. Ratio n/n-1: 0.7. The labial palp is without pigment (Fig. 23). The third segment is smaller than in the male. Fewer sensory connules are present than in the male. The three pairs of legs are robust with weak and diffuse pigment (Figs 24–26). The ventral side of the legs with strong setae as in the male. Length of the tibiae: I: 0.86 mm, II: 0.74 mm, III: 1.01 mm.
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Figs. 24–29. Mendeschilis scorcai n.g., n.sp., female. 24. Outline of fore leg; 25. Ditto, mid leg; 26. Ditto, hind leg; 27. Outline of the Vth urosternite; 28. Ditto, of the VIIth; 29. Ditto, of the VIIIth with gonapophysis. Scale: 0.1 mm
Urosternites II–VII with acutely angle sternites; coxites, sometimes with strong setae (Figs. 27–30). Table 1 shows the length ratios between coxite/stylet (without spine)/spine of the stylet. The female has a tertiary type ovipositor that exceeds the IXth stylet in shorter length than the length of the stylet. Gonapophysis VIII with 1+61 divisions, all except the more basal divisions with setae. The chaetotaxy is shown in figures 31–32. Gonapophysis IX with 1+61 divisions; its chaetotaxy is shown in figures 33–34. Apex of the cerci as in the male.
Discussion Before the analysis of the new genus, some information is provided about Praemachilis excelsior Silvestri, 1904, a rarely found species. P. excelsior was first described from Bevagna and Portici (Italy) by Silvestri (1904), and later cited in Monaco and Menton in Southeastern France (Silvestri 1907).
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Figs. 30–34. Mendeschilis escorcai n.g., n.sp., female. 30. IXth urosternite with gonapophysis; 31. Divisions 25 to 30 of the VIIIth gonapophysis; 32. Apical divisions of the VIIIth gonapophysis; 33. Divisions 17 to 21 of the IXth gonapophysis; 34. Apical divisions of the IXth gonapophysis. Scale: 0.1 mm
Originally only the female was described, but the male characteristics (exclusively genitalia) were included in the generic definition and the identification keys only. Janetschek (1954) redescribed the genus and the species using the “cotypes” from Silvestri (only females). Paclt (1969) observed P. excelsior males in Majorca and Mendes (1981) made the formal description of male characteristics. Now we consider that these Balearic specimens do not belong to P. excelsior but to a new species and to a new genus Mendeschilis escorcai n. g., n. sp. To clarify the correct taxonomic positioning of each taxon, some available samples which had been identified as P. excelsior were obtained from Italy, using material held in Natural History Museum of Portici (NHMP), the Museum of Natural History of Verona (MNHV) and the University of Córdoba (UCO): Bevagna, 1 female (cotypus labelled), no date, Silvestri det., (Microscopic slide), NHMP. Portici, 1 female (cotypus labelled), no date, Silvestri det., (Microscopic slide), NHMP.
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Fig. 35–45. 35. Praemachilis excelsior Silvestri, 1904:1st segment of the maxillary palp showing the apophyses, female specimen (cotypus) from Portici (Italy); 36. Ditto, female specimen (cotypus) from Bevagna (Italy); 37. Ditto, male specimen from Santa Eufemia (Aspromonte, Italy); 38. Mesdeschilis escorcai n.g. n. sp. from Majorca (Spain); 39. Mendeschilis sp. from Marettimo (Egadi Islands, Italy); 40. Praemachilis excelsior Silvestri, 1904, from Pontignano (Siena, Italy) Head: frontal view; 41. Head, lateral view; 42. Mendeschilis sp. from Marettimo (Egadi Is., Italy) Head: frontal view; 43. Head, lateral view; 44. Praemachilis excelsior Silvestri, 1904, Urosternite VIII of the male; 45. Ditto, Urosternite IX with the penis and parameres. Scale: 0.1 mm
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Portici, 6 females (labelled as cotypi!!) Giugno 1907, Silvestri det., (Alcohol), NHMP. (This specimens are erroneously labelled as “cotipes”, because the species was described in 1904). Santa Eufemia (Aspromonte), 1 male, 22-7-1957, Bach det. 1983, MNHV. Marettimo (Egadi Is.), 1 female, 23-10-1967, Bach det. 1983, MNHV. Pontignano (Siena), 5 females, 19-10-1995, Gaju det. UCO. (new material) The revision of these samples allows to state the similarities among the Italian specimens, all of which except the female from Marettimo belong to P. excelsior. It was very important to collect and study live specimens from Pontignano, because their facies was very characteristic, and our revision of the Silvestri’s original description suggests they belong to P. excelsior. The specimens show a slightly humped mesonotum, which fits Silvestri’s description “Gibba thoracica magna …”, the pattern of scales in the dorsal part of the body was also consistent with Silvestri’s description with a transverse black band in the middle flanked by white bands (“... tergito tertio nigro-velutino, albo marginato, tergitis 4° et 5° albis …”). Also, the antenna show a characteristic white band in their middle part, recorded by Silvestri as (“… partem medianam versus albo”). Studies on the Silvestri’s specimens and the other material from Italy show that the main morphological characteristics were consistent but some slight differences were observed which could reflect the diverse range of mounting media used (Canada Balsam, Hoyer or Tendeiro liquid). The apophysis of the first maxillary palp segment may be used to taxonomically distinguish the two genera (Figs. 35–37); in all cases in Praemachilis the external apophysis is large, conical, slightly curved back and rounded at the apex, the internal apophysis is short, conical and rounded at the apex. Comparing the same segment of the new genus, the outer apophysis, although similar, is slightly thinner with a pointed ending, and the inner one is clearly conical, also with a pointed ending (Fig. 38). Another distinguishing feature of the two genera is the shape of paired ocelli; they are oval in P. excelsior (Figs. 40–41) (see also drawings of Janetschek 1954, p. 210), but rounded in M. escorcai (Figs. 1–3). Curiously the female from Marettimo (Egadi Islands) studied by Bach de Roca (1983) must belong to Mendeschilis genus because of the last two characteristics (Figs. 39 and 42–43). In relation to the male characteristics, as the specimens referred to by Silvestri have not been found, we suppose they could belong to another species and genus, because the original Praemachilis genus include, in addition to P. excelsior, the following species: P. meticulosa, (now = Praetrigoniophthalmus meticulosa (Silvestri 1904)) whose male has parameres in the IXth urosternite only; P. italica (now = Dilta italica (Grassi, 1887)), whose male has parameres in the VIIIth and IXth urosternites and P. aurea (Giardina 1900) a species insufficiently described and considered by Janetschek (1954) as not valid until new material provides a new diagnosis. So, in the same genus Silvestri included species with different numbers of parameres and which today are recognised as belonging to three different genera. Study of the specimen from Aspromonte (Santa Eufemia) suggests that it belongs to the genus Praemachilis according to the characteristics of its maxillary palps (Fig. 37). Unfortunately the specimen is not completely adult and the part preserved in alcohol was dried and its head characteristics are not clearly visible. The VIIIth and
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IXth urosternites with the genitalia are shown in figures 44–45. The degree of development of the penis and IXth parameres suggests that the VIIIth parameres would be perfectly visible if they existed, in spite of being a subadult; so we believe this specimen did not possess any VIIIth paramera. We therefore suggest that members of the genus Praemachilis only posses parameres in the IXth urosternites. In Machilis Latreille, 1832, there are many species with parameres in the VIIIth and IXth urosternites, but only two species have parameres only in the IXth urosternite; for these two species the subgenus Pseudomachilis was created (Janetscheck 1955), based only upon the number of parameres. In our case, although the number of parameres is one difference between Praemachilis and Mendeschilis, other morphological differences exist which permit both genera to be distinguished (Table 2). Finally, we have two additional remarks concerning P. excelsior. The first is to suggest that probably the species can reproduce partenogenetically in some places, because of the low number of males found; in fact, in Pontignano we have found more than 15 specimens and all were females. The second point is to suggest the possible synonimy between the genera Praemachilis and Wygodzinskylis Janetschek, 1954, because comparison of the studied specimens of P. excelsior with the description of Wygodzinskilis klinocellata Janetschek (1954) a further redescription (Janetschek 1959), and two specimens from L.F. Mendes’ Collection, yielded no obvious differences between these genera (as discussed above, we consider that both genera have parameres only in the IXth urosternite). Other potential differences between the genera (such as the shape of the apophysis of the first maxillary palp segment and the characteristics of the antennae) were not evident. However, until new and abundant material can be studied, we prefer to maintain the status of both genera.
Table 2. Differences between Mendeschilis n. g. and Praemachilis Silvestri, 1904 Characteristics
Mendeschilis n.g.
Body shape Ocelli
Mesonotum rounded Rounded
Antennae Distal antennae chains First segment of the maxillary palp: External apophysis Ditto, internal apophysis Legs (ventral side of tibia and tarsi) Coxites of urosternites Parameters Ratio bp/dp of penis
Longer than the body Until 30 divisions Conical and apically pointed (Figs. 6 and 38)
Praemachilis Silvestri,1904 Slightly humped Oval (twice as long as wide) Shorter Usually 9 divisions Conical to Subcilindrical and apically rounded
Pointed (Figs. 6 and 38) With strong setae
Rounded (Figs. 35–37) With spiniform satae
With some setae VIII and IX 0.89
Without setae Only IX (Fig. 44–45) 1.5
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Derivation of the name The new genus is named Mendeschilis and is dedicated to our colleague and friend Luis F. Mendes from Lisbon (Portugal). The species name is taken from the site of sampling in Escorca (Majorca, Spain).
Acknowledgements This work had been supported by the Project “Fauna Ibérica III” SEUI-DGICYT PB92-0121. The authors are deeply grateful to the Museum of Natural History of Verona and the Natural History Museum of Portici (Italy) for sending us the specimens for this study, as well as to Dr. Geoff Frampton for improving English in the manuscript.
References Bach de Roca, C. (1983) Contribución al conocimiento de los Microcoryphia de Italia (Insecta, Apterygota). Bollettino del Museo Civico di Storia Naturale di Verona, [1982], 523–629. Janetschek, H. (1954) Ueber Felsenspringer der Mittelmeerländer (Thysanura, Machilidae). Eos 30, 163–314. Janetschek, H. (1955) Felsenspringer aus Sardinien, Korsika und den Ostpyrenäen (Ins., Thysanura). Österreichische Zoologische Zeitschrift 5, 407–441. Janetschek, H. (1959) Weitere Machilida aus dem Balkan. Acta Musei Macedonici Scientiarum Naturalium 6 (58), 120–141. Mendes, L.F. (1981) Contribution à la connaissance des Machilides (Microcoryphia: Apterygota) de l’Ile de Majorque, avec description de deux nouvelles espèces. Revue Suisse de Zoologie 88(2), 413–432. Paclt, J. (1969) Neue Beiträge zur Kenntnis der Apterygoten-Sammlung des Zoologischen Staatsinstituts und Zoologischen Museums Hamburg. III. Meinertellidae und Machilidae (Thysanura). Entomologische Mitteilungen aus dem Zoologischen Staatsinstitut und Zoologischen Museum Hamburg 3(63), 269–272. Silvestri, F. (1904) Nuovi generi e specie di Machilidae. Redia 2, 4–9. Silvestri, F. (1907) Catalogue des Machilidae de la collection du Muséum. Note sui Machilidae. Redia 3, 32–34. Sturm, H., Bach de Roca, C. (1993) On the Systematics of the Archaeognatha (Insecta). Entomologia Generalis 18(1/2), 55–90.