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Redescription of Catamachilis constricta (Navás, 1905) (Insecta, Apterygota, Microcoryphia)
Pedobiologia 44, 285–291 (2000) © Urban & Fischer Verlag http://www.urbanfischer.de/journals/pedo
PROCEEDINGS OF VTH INTERNATIONAL SEMINAR ON APTERYGOTA, CORDOBA 1998
Redescription of Catamachilis constricta (Navás, 1905) (Insecta, Apterygota, Microcoryphia) Ma José Notario-Muñoz1, Carmen Bach de Roca2, Rafael Molero-Baltanás1 and Miguel Gaju-Ricart1 1 Department of Animal Biology (Zoology), University of Cordoba, C-1 Campus Rabanales, N-IV Km 396A, 14014 Cordoba, Spain 2 Department of Animal Biology, Botany and Ecology, Autonomous University of Barcelona, 08193 Bellaterra, Barcelona, Spain
Accepted: 30. December 1999
Summary The genus Catamachilis was created by Silvestri in 1923 for Machilis constricta Navás, 1905. Because the Navás collection was lost during the Spanish Civil War, material from the type locality was used for redescription of C. constricta, and the establishment of a neotype for the lost holotype of this species. Key words: Catamachilis constricta, neotype, Microcoryphia, Spain
Introduction The genus Catamachilis belongs to Machilidae. It is endemic in the Iberian Peninsula, although once was found in the South of France. The type species of the genus is C. constricta, which was described in 1905 by Navás as Machilis constricta. In 1923, when Silvestri studied the Navás’s material, he examined the holotype of M. constricta and saw that it had characters which differed from those of the genus Machilis. It allowed him to describe a new genus Catamachilis, with C. constricta as the type species. During the Spanish Civil War (1936–1939), the museum where Navás had deposited reference material was destroyed and consequently the Navás collection was lost. Corresponding author Miguel Gaju-Ricart, e-mail: [email protected]
0031–4056/00/44/03–04–285 $ 12.00/0
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Until our work nobody had found any other specimens of C. constricta, so all type material of C. constricta was lost. The “Fauna Iberica Project” allowed us to sample many localities in Spain in order to collect Microcoryphia. One of our sampling sites was in Tudela, the locality where Navás had collected and described C. constricta. We collected one male and one female, whose characteristics agreed with those given by Silvestri in the redescription of C. constricta. In this paper we redescribe this species in detail and establish neotypes. Collection of specimens from a location 160 km from Tudela suggests that the distribution area of this species is larger than had been observed previously.
Materials and Methods The specimens were searched for under stones and crevices in the soil in habitats with sparse vegetation. The insects were collected with an aspirator and kept in 70 % alcohol. In the laboratory the specimens were dissected and mounted in Hoyer’s medium, then dried in an oven at 35°C for four weeks before their examination under an optical microscope (40X to 400X).
Results Catamachilis constricta (Navás, 1905) Studied material: Tudela (Navarra), 22-07-87, 1 male neotype + 1 female, Ref. M0682; Alcorisa (Teruel), 30-07-87, 2 males + 2 females + 1 juv. Ref. M0617. The neotype is held in the Museo Nacional de Ciencias Naturales in Madrid. Collection number: 8527. Redescription of the male Body length: 10.5 mm; length of conserved antennae: 9 mm; cerci and paracercus broken (conserved part: 4mm). Head without pigment, the shape is as in figures 1–3. Eye ratios: contact line / length (cl/l): 0.39; length / width (l/w): 0.87. Antennae with distal chains formed by 16 annuli. The maxillary palp chaetotaxy is dense (Fig. 4); on the ventral side from the second to the seventh segments there is a field of spirally-arranged short setae which are denser in the distal segments (Figs. 4–6); in addition, there are from the second to the sixth segments, some setae which are longer than a third of the width of each segment. Characteristic hyaline spines appear on the last three segments (Fig. 6), their distribution as follows: 5 segment: 1 spine; 6: 11; 7: 11. The length ratio seventh / sixth segments (n/n-1): 0.68. Labial palp modified in shape; the third segment is strongly widened with a field of sensorial conuli on the distal part; among the conuli there are thin and short setae (Fig. 7). The three pairs of legs are morphologically typical of the genus, only the third pair has a coxal stylet (Figs. 8–10). The fore femur long; ratio length / width (l/w): 2.33. Length of tibiae: I: 0.9 mm; II: 0.8 mm; III: 1.02 mm. All of them have ventral spines whose distribution is shown in Table 1. Urosternites (Fig. 11) are typical of the genus, with one pair of coxal vesicles pre-
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Figs. 1–10. Catamachilis constricta (Navás, 1905), male. 1. Head, frontal view; 2. Head, lateral view; 3. Outline of the eyes and ocelli; 4. Outline and chaetotaxy of the maxillary palp; 5. Second segment of the maxillary palp showing the short setae on the ventral side; 6. Shape and setae of the seventh segment of the maxillary palp; 7. Outline and setae of the labial palp. 8. Outline and chaetotaxy of the first leg; 9. Ditto, second leg; 10. Ditto, third leg. Scale: 0.1mm
sent on the first to the seventh. Sternites right-angled. Ratios of the length of coxite, stylet (without spine) and spine of the stylet are given in Table 2. The eighth and ninth urosternites are the genital ones; as is usual in the genus Catamachilis the eighth lacks parameres (Fig. 12). On the ninth there is a penis and a pair of parameres (Fig. 13). The penis opening is oval in shape, a little longer than it is wide and, inside, there is an elongated papillus (Fig. 14). The length ratio of basal and terminal parts of the penis is 0.68. The parameres have 8 divisions each.
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Table 1. Catamachilis constricta (Navás, 1905). Spines on the legs. Brackets, data for C. clipeata from Bach de Roca 1980. Leg I II III
male female male female male female
Femur 0 (–) 2 (1–2) 7 (0) 8 (0–1) 3 (3–4) 3 (4)
Tibia 7 (8) 6 (6) 18 (3) 10 (3–4) 21 (8) 14 (7)
tarsus I 4 (3) 3 (3–4) 6 (3) 7 (3–4) 9 (4) 6 (5)
tarsus II 11 (6) 8 (5) 15 (6) 6 (5) 13 (4) 9 (8)
tarsus III 6 (4) 7 (5–6) 7 (4) 6 (5–6) 7 (3) 8 (7)
Figs. 11–14. C. constricta (Navás, 1905), male. 11. Outline of the Vth urosternite; 12. Ditto, VIIIth urosternite; 13. Ditto, IXth urosternite with parameres and penis; 14. Urosternite IX: parameres and penis. Scale: 0.1 mm
Female redescription Body length: 9 mm; antennae length: 7 mm; cerci and paracercus broken. The head has the same shape as that of the male. Ratio cl/l: 0.28; l/w:0.91. The distal antennal chains have 14 annuli. The maxillary palp lacks secondary sexual characteristics, without speciallized setae (Fig. 15). The distribution of hyaline spines in the last three segments is: 5 segment: 3 spines; 6: 15; 7: 16. Ratio n/n-1: 0.78. The third segment of the labial palp is not modified in shape (Fig. 16). The shape of the legs is similar to that in the male. Length of tibiae I: 0.72 mm; II: 0.60 mm; III: 0.80 mm. The number of ventral spines differs among legs and between sexes (Table 1). Urosternites with obtusely-angled sternites (Fig. 17); the seventh is swollen on its inner part (Fig. 18). The ratios between length of coxite, stylet (without spine) and spine of the stylet are shown in Table 2.
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Figs. 15–22. C. constricta (Navás, 1905), female. 15. Outline of the maxillary palp; 16. Ditto, labial palp; 17. Ditto, Vth urosternite; 18. Ditto, VIIth urosternite; 19. Outline of the IXth urosternite with gonapophysis; 20. Ditto, VIIIth urosternite with gonapophysis; 21. Last divisions of the VIIIth gonapophysis; 22. Ditto, IXth gonapophysis. Scale: 0.1 mm
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Table 2. Catamachilis constricta (Navás, 1905). Ratios of the length of coxite, stylet (without spine) and spine for each of three urosternites Urosternite V VIII IX
male female male female male female
stylet/coxite 0.58 0.52 0.86 0.78 1.05 0.60
spine/stylet 0.23 0.38 0.23 0.26 0.14 0.19
The ovipositor is of the primary type (Sturm & Bach 1993), not surpassing the apex of the ninth stylet (Fig. 19). The eighth gonapophysis has 26 divisions (Fig. 20) and the ninth 27 (Fig. 19), their chaetotaxy is shown in Figs. 21–22.
Discussion The neotype of C. constricta (Navás, 1905) is redescribed, allowing us to update the earlier diagnosis and to compare and distinguish it from four other known species of the genus: C. amara Janetschek, 1954; C. clipeata Stach, 1930; C. franzi Janetschek, 1954 and C. ancorata Stach, 1930. The most important character for distinguishing the male of C. constricta from C. amara and C. clipeata is the penis, which has in C. amara and C. clipeata a subparallel shape with a rounded opening (Janetschek 1954 p. 182, Fig. 33) but it is large with an oval opening in C. constricta (Figs. 13, 14). Moreover, on the seventh segment of the maxillary palp of C. clipeata and C. amara there is a field of sensorial setae (see Janetschek 1954 p. 182, Fig. 26 and Bach de Roca 1980, p. 37, Figs. 22, 23) which is absent in C. constricta (Fig. 6). Between C. franzi and C. constricta males, the most important difference is found on the maxillary palp, where the seventh segment is clearly swollen (Janetschek 1954), in addition, the former has a field of sensorial setae (similar to that one of C. amara and C. clipeata). The male of C. ancorata is distinguished from C. constricta by the shape of the maxillary palp, the last segment of which is more elongate in the latter species; There are also differences in chaetotaxy, with an abundant field of long setae from the second to the sixth maxillary palp segments of C. ancorata whereas C. constricta has some long setae and a dense field of short and spirally-arranged setae (compare Fig. 5 with Bach de Roca 1980 p. 34, Fig. 2). Among females differences between species are less apparent than among males, but separation is possible using body pigmentation, the shape of the ovipositors and the spines on the legs. C. ancorata has quite a long ovipositor (Stach 1930 and Bach de Roca 1980) surpassing the IXth stylets, and so completely different to all other species. C. amara and C. franzi have strong pigmentation of head (Janetschek, 1954), so they are different from C. constricta. Finally C. clipeata can be distinguished by the spines on the legs , which are more numerous in C. constricta (Table 1).
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Acknowledgements This work was supported by the Project “Fauna Ibérica III” SEUI-DGICYT PB92-0121. We wish to thank Dr. Geoff Frampton for improving English in the manuscript.
References Bach de Roca, C. (1980) Contribución al conocimiento de los Microcoryphia (Insecta, Apterygota) de España: Género Catamachilis Silvestri, 1923. Miscellania Zoologica 6, 33–39. Janetschek, H. (1954) Ueber Felsenspringer der Mittelmeerländer (Thysanura, Machilidae). Eos 30, 163–314. Navás, L. (1905) Mis excursiones durante el verano de 1904. Boletín de la Sociedad aragonesa de Ciencias Naturales 4 (4–5), 1–35. Silvestri, F. (1923) Due nuovi generi e una nuova specie di Machilidae della Spagna. Revista de la Academia de Ciencias de Zaragoza 6, 123–131. Stach, J. (1930) Apterygoten aus dem nördlichen und östlichen Spanien, gesammelt von D. F. Haas in den Jahren 1914–1919. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 42 (1), 1–83. 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
Redescription of Catamachilis constricta (Navás, 1905) (Insecta, Apterygota, Microcoryphia) Ma José Notario-Muñoz1, Carmen Bach de Roca2, Rafael Molero-Baltanás1 and Miguel Gaju-Ricart1 1 Department of Animal Biology (Zoology), University of Cordoba, C-1 Campus Rabanales, N-IV Km 396A, 14014 Cordoba, Spain 2 Department of Animal Biology, Botany and Ecology, Autonomous University of Barcelona, 08193 Bellaterra, Barcelona, Spain
Accepted: 30. December 1999
Summary The genus Catamachilis was created by Silvestri in 1923 for Machilis constricta Navás, 1905. Because the Navás collection was lost during the Spanish Civil War, material from the type locality was used for redescription of C. constricta, and the establishment of a neotype for the lost holotype of this species. Key words: Catamachilis constricta, neotype, Microcoryphia, Spain
Introduction The genus Catamachilis belongs to Machilidae. It is endemic in the Iberian Peninsula, although once was found in the South of France. The type species of the genus is C. constricta, which was described in 1905 by Navás as Machilis constricta. In 1923, when Silvestri studied the Navás’s material, he examined the holotype of M. constricta and saw that it had characters which differed from those of the genus Machilis. It allowed him to describe a new genus Catamachilis, with C. constricta as the type species. During the Spanish Civil War (1936–1939), the museum where Navás had deposited reference material was destroyed and consequently the Navás collection was lost. Corresponding author Miguel Gaju-Ricart, e-mail: [email protected]
0031–4056/00/44/03–04–285 $ 12.00/0
286
Ma José Notario-Mun˜oz et al.
Until our work nobody had found any other specimens of C. constricta, so all type material of C. constricta was lost. The “Fauna Iberica Project” allowed us to sample many localities in Spain in order to collect Microcoryphia. One of our sampling sites was in Tudela, the locality where Navás had collected and described C. constricta. We collected one male and one female, whose characteristics agreed with those given by Silvestri in the redescription of C. constricta. In this paper we redescribe this species in detail and establish neotypes. Collection of specimens from a location 160 km from Tudela suggests that the distribution area of this species is larger than had been observed previously.
Materials and Methods The specimens were searched for under stones and crevices in the soil in habitats with sparse vegetation. The insects were collected with an aspirator and kept in 70 % alcohol. In the laboratory the specimens were dissected and mounted in Hoyer’s medium, then dried in an oven at 35°C for four weeks before their examination under an optical microscope (40X to 400X).
Results Catamachilis constricta (Navás, 1905) Studied material: Tudela (Navarra), 22-07-87, 1 male neotype + 1 female, Ref. M0682; Alcorisa (Teruel), 30-07-87, 2 males + 2 females + 1 juv. Ref. M0617. The neotype is held in the Museo Nacional de Ciencias Naturales in Madrid. Collection number: 8527. Redescription of the male Body length: 10.5 mm; length of conserved antennae: 9 mm; cerci and paracercus broken (conserved part: 4mm). Head without pigment, the shape is as in figures 1–3. Eye ratios: contact line / length (cl/l): 0.39; length / width (l/w): 0.87. Antennae with distal chains formed by 16 annuli. The maxillary palp chaetotaxy is dense (Fig. 4); on the ventral side from the second to the seventh segments there is a field of spirally-arranged short setae which are denser in the distal segments (Figs. 4–6); in addition, there are from the second to the sixth segments, some setae which are longer than a third of the width of each segment. Characteristic hyaline spines appear on the last three segments (Fig. 6), their distribution as follows: 5 segment: 1 spine; 6: 11; 7: 11. The length ratio seventh / sixth segments (n/n-1): 0.68. Labial palp modified in shape; the third segment is strongly widened with a field of sensorial conuli on the distal part; among the conuli there are thin and short setae (Fig. 7). The three pairs of legs are morphologically typical of the genus, only the third pair has a coxal stylet (Figs. 8–10). The fore femur long; ratio length / width (l/w): 2.33. Length of tibiae: I: 0.9 mm; II: 0.8 mm; III: 1.02 mm. All of them have ventral spines whose distribution is shown in Table 1. Urosternites (Fig. 11) are typical of the genus, with one pair of coxal vesicles pre-
Redescription of Catamachilis constricta
287
Figs. 1–10. Catamachilis constricta (Navás, 1905), male. 1. Head, frontal view; 2. Head, lateral view; 3. Outline of the eyes and ocelli; 4. Outline and chaetotaxy of the maxillary palp; 5. Second segment of the maxillary palp showing the short setae on the ventral side; 6. Shape and setae of the seventh segment of the maxillary palp; 7. Outline and setae of the labial palp. 8. Outline and chaetotaxy of the first leg; 9. Ditto, second leg; 10. Ditto, third leg. Scale: 0.1mm
sent on the first to the seventh. Sternites right-angled. Ratios of the length of coxite, stylet (without spine) and spine of the stylet are given in Table 2. The eighth and ninth urosternites are the genital ones; as is usual in the genus Catamachilis the eighth lacks parameres (Fig. 12). On the ninth there is a penis and a pair of parameres (Fig. 13). The penis opening is oval in shape, a little longer than it is wide and, inside, there is an elongated papillus (Fig. 14). The length ratio of basal and terminal parts of the penis is 0.68. The parameres have 8 divisions each.
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Table 1. Catamachilis constricta (Navás, 1905). Spines on the legs. Brackets, data for C. clipeata from Bach de Roca 1980. Leg I II III
male female male female male female
Femur 0 (–) 2 (1–2) 7 (0) 8 (0–1) 3 (3–4) 3 (4)
Tibia 7 (8) 6 (6) 18 (3) 10 (3–4) 21 (8) 14 (7)
tarsus I 4 (3) 3 (3–4) 6 (3) 7 (3–4) 9 (4) 6 (5)
tarsus II 11 (6) 8 (5) 15 (6) 6 (5) 13 (4) 9 (8)
tarsus III 6 (4) 7 (5–6) 7 (4) 6 (5–6) 7 (3) 8 (7)
Figs. 11–14. C. constricta (Navás, 1905), male. 11. Outline of the Vth urosternite; 12. Ditto, VIIIth urosternite; 13. Ditto, IXth urosternite with parameres and penis; 14. Urosternite IX: parameres and penis. Scale: 0.1 mm
Female redescription Body length: 9 mm; antennae length: 7 mm; cerci and paracercus broken. The head has the same shape as that of the male. Ratio cl/l: 0.28; l/w:0.91. The distal antennal chains have 14 annuli. The maxillary palp lacks secondary sexual characteristics, without speciallized setae (Fig. 15). The distribution of hyaline spines in the last three segments is: 5 segment: 3 spines; 6: 15; 7: 16. Ratio n/n-1: 0.78. The third segment of the labial palp is not modified in shape (Fig. 16). The shape of the legs is similar to that in the male. Length of tibiae I: 0.72 mm; II: 0.60 mm; III: 0.80 mm. The number of ventral spines differs among legs and between sexes (Table 1). Urosternites with obtusely-angled sternites (Fig. 17); the seventh is swollen on its inner part (Fig. 18). The ratios between length of coxite, stylet (without spine) and spine of the stylet are shown in Table 2.
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Figs. 15–22. C. constricta (Navás, 1905), female. 15. Outline of the maxillary palp; 16. Ditto, labial palp; 17. Ditto, Vth urosternite; 18. Ditto, VIIth urosternite; 19. Outline of the IXth urosternite with gonapophysis; 20. Ditto, VIIIth urosternite with gonapophysis; 21. Last divisions of the VIIIth gonapophysis; 22. Ditto, IXth gonapophysis. Scale: 0.1 mm
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Table 2. Catamachilis constricta (Navás, 1905). Ratios of the length of coxite, stylet (without spine) and spine for each of three urosternites Urosternite V VIII IX
male female male female male female
stylet/coxite 0.58 0.52 0.86 0.78 1.05 0.60
spine/stylet 0.23 0.38 0.23 0.26 0.14 0.19
The ovipositor is of the primary type (Sturm & Bach 1993), not surpassing the apex of the ninth stylet (Fig. 19). The eighth gonapophysis has 26 divisions (Fig. 20) and the ninth 27 (Fig. 19), their chaetotaxy is shown in Figs. 21–22.
Discussion The neotype of C. constricta (Navás, 1905) is redescribed, allowing us to update the earlier diagnosis and to compare and distinguish it from four other known species of the genus: C. amara Janetschek, 1954; C. clipeata Stach, 1930; C. franzi Janetschek, 1954 and C. ancorata Stach, 1930. The most important character for distinguishing the male of C. constricta from C. amara and C. clipeata is the penis, which has in C. amara and C. clipeata a subparallel shape with a rounded opening (Janetschek 1954 p. 182, Fig. 33) but it is large with an oval opening in C. constricta (Figs. 13, 14). Moreover, on the seventh segment of the maxillary palp of C. clipeata and C. amara there is a field of sensorial setae (see Janetschek 1954 p. 182, Fig. 26 and Bach de Roca 1980, p. 37, Figs. 22, 23) which is absent in C. constricta (Fig. 6). Between C. franzi and C. constricta males, the most important difference is found on the maxillary palp, where the seventh segment is clearly swollen (Janetschek 1954), in addition, the former has a field of sensorial setae (similar to that one of C. amara and C. clipeata). The male of C. ancorata is distinguished from C. constricta by the shape of the maxillary palp, the last segment of which is more elongate in the latter species; There are also differences in chaetotaxy, with an abundant field of long setae from the second to the sixth maxillary palp segments of C. ancorata whereas C. constricta has some long setae and a dense field of short and spirally-arranged setae (compare Fig. 5 with Bach de Roca 1980 p. 34, Fig. 2). Among females differences between species are less apparent than among males, but separation is possible using body pigmentation, the shape of the ovipositors and the spines on the legs. C. ancorata has quite a long ovipositor (Stach 1930 and Bach de Roca 1980) surpassing the IXth stylets, and so completely different to all other species. C. amara and C. franzi have strong pigmentation of head (Janetschek, 1954), so they are different from C. constricta. Finally C. clipeata can be distinguished by the spines on the legs , which are more numerous in C. constricta (Table 1).
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Acknowledgements This work was supported by the Project “Fauna Ibérica III” SEUI-DGICYT PB92-0121. We wish to thank Dr. Geoff Frampton for improving English in the manuscript.
References Bach de Roca, C. (1980) Contribución al conocimiento de los Microcoryphia (Insecta, Apterygota) de España: Género Catamachilis Silvestri, 1923. Miscellania Zoologica 6, 33–39. Janetschek, H. (1954) Ueber Felsenspringer der Mittelmeerländer (Thysanura, Machilidae). Eos 30, 163–314. Navás, L. (1905) Mis excursiones durante el verano de 1904. Boletín de la Sociedad aragonesa de Ciencias Naturales 4 (4–5), 1–35. Silvestri, F. (1923) Due nuovi generi e una nuova specie di Machilidae della Spagna. Revista de la Academia de Ciencias de Zaragoza 6, 123–131. Stach, J. (1930) Apterygoten aus dem nördlichen und östlichen Spanien, gesammelt von D. F. Haas in den Jahren 1914–1919. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 42 (1), 1–83. Sturm, H., Bach de Roca, C. (1993) On the Systematics of the Archaeognatha (Insecta). Entomologia Generalis 18 (1/2), 55–90.