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Praemachiloides iberica n. sp. from Spain (Insecta Microcoryphia)
Pedobiologia 44, 292–299 (2000) © Urban & Fischer Verlag http://www.urbanfischer.de/journals/pedo
PROCEEDINGS OF VTH INTERNATIONAL SEMINAR ON APTERYGOTA, CORDOBA 1998
Praemachiloides iberica n. sp. from Spain (Insecta Microcoryphia) Rosa Ma Mora-Carmona1, Miguel Gaju-Ricart1 and Carmen Bach de Roca2 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 1
Accepted: 30. December 1999
Summary Praemachiloides iberica n. sp. is described and compared with the other four known species of the genus: P. janetschecki, P. insularis, P.tarsispina and P. autumnalis. The main difference of P. iberica n. sp. from the other known congenerics is the presence, in the male, of a field of plaited setae on the ventral surface of the maxillary palp. Key words: Praemachiloides, Spain, new species, Microcoryphia, Apterygota
Introduction In field studies within the “Fauna Iberica” Project, we obtained samples of Praemachiloides sp. that had been studied genetically by Fanciulli et al. (1995), together with other well known species of Praemachiloides. When we studied the morphological characteristics of these Praemachiloides spp. we saw that the specimens of these populations were different, belonging to a new species that we named P. iberica n. sp.
Corresponding author Miguel Gaju-Ricart, e-mail: [email protected]
0031–4056/00/44/03–04–292 $ 12.00/0
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Materials and Methods The specimens were collected with an aspirator, then fixed with 70 % alcohol. In the laboratory, the specimens were dissected and mounted on microscope slides using Tendeiro liquid (see Molero-Baltanás et al. in this volume). The slides were then dried in an oven at 35°C.
Results Praemachiloides iberica n. sp. Studied material Gea de Albarracín (Teruel), 16-04-87, 2 males + 2 females, Ref. M0671; Royuela (Teruel), 16-04-87, 1 male + 1 female, Ref. M0670; Beteta (Cuenca), 16-02-90, 1 male + 1 female, Ref. M0676; Arcos de Jalón (Soria), 29-02-92, 4 females, Ref. M0770; Agost (Alicante), 11-04-92, 1 male, Ref. M0766; Entre Confrides y puerto de Ares (Alicante), 12-04-92, 1 female, Ref. M0772; Alhama de Murcia, Sierra Espuña, 1504,92, 3 males + 2 females, Ref. M0764; Gea de Albarracín (Teruel), 31-03-93, 1 female, Ref. M0531; Beteta (Cuenca), 01-04-93, 1 male holotype + 1 female allotype + 2 females paratypes, Ref. M0533; Arcos de Jalón (Soria), 01-04-93, 1 male, Ref. M0535; Elche de la Sierra (Albacete), 16-01-94, 4 males juv.+ 4 females juv., Ref. M0763; Yecla (Murcia), 29-03-94, 1 female, Ref. M0818; Elche de la Sierra (Albacete), 30-03-94, 4 males + 1 female, Ref. M0844. The holotype male, allotype female and paratypes are held in the “Museo Nacional de Ciencias Naturales” in Madrid; collection number: 8524. Description of the male Body length: 10.5 mm, length of antennae:10 mm, length of paracercus: 10 mm, length of cercus: 4 mm. Head (Figs. 1–2) with pigmented frons, apart from a small central line without pigment between the paired ocelli, extending to the base of the antennae. Clypeus with numerous white setae and little pigment on the lateral side. Pigmented labrum. Compound eyes as long as wide (Fig. 3). Eye ratios: contact line / length (cl/l): 0.36, length / width (l/w): 1.08. The paired ocelli are oval to subtriangular, dark-reddish in colour and located in a submedian position. The antennae are the same length as the body but can be longer. Scapus (Fig. 4) twice as long as wide with diffuse pigment. Flagellus pigmented. Distal chains of the antennae with 17 annuli which are wider than they are long (Figs. 5–6). Although some basiconic sensilla are present (Fig. 6), there is not a regular pattern of sensilla distribution. The maxillary palp (Fig. 7) shows a field of plaited setae (Fig. 9) on the ventral surface of all the segments. Hyaline spines (Fig. 8) appear on the last three segments, their distribution as follows: segment 5: 2 spines; 6: 10-16; 7:11-15. Ratio last / penultimate segments (n/n-1): 0.84. The labial palp is without any special characteristics; the second segment has subparallel sides and the third segment is typically enlarged, with a field of conules present (Figs. 10–11).
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Figs. 1–15. Praemachilodes iberica n. sp., male. 1. Head, frontal view; 2. Head, lateral view; 3. Outline of the eyes and ocelli; 4. Outline of the scapus and pedicellus of the antenna; 5. Shape of one distal antenna chain; 6. Annuli of the antenna chain, showing special sensilla; 7. Outline of the maxillary palp; 8. Last segment of the maxillary palp showing the hyaline spines; 9. Plaited setae on the ventral surface of the maxillary palp; 10. Outline of the labial palp; 11. Sensorial conules of the 3rd segment of the labial palp; 12. Outline of the fore leg; 13.Tarsus of the fore leg showing the spines of the 2nd tarsomere; 14. Outline of the mid leg; 15. Outline of the hind leg. Scale: 0.1 mm
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The three pairs of legs have weak pigmentation. The first pair has the femur enlarged (ratio l/w= 1,88) (Fig. 12). The second tarsomere has short, pigmented, ventral spines (Fig. 13). The second and third pairs of legs (Figs. 14-15) have longer spines than those of the first pair. On the tibia there are some spiniform setae. Length of the tibiae: I: 0.73 mm; II: 0.68 mm; III: 0.91 mm. Urosternites with obtusely-angled sternites; the first to seventh urosternites each posses a pair of eversible vesicles. The coxites have some setae which may vary in number from one coxite to another (Fig. 16). The length ratios between coxite/stylet (without spine)/spine of the stylets are given in Table 1.
Table 1. P. iberica n. sp. Length ratios of coxite, stylet (without spine) and spine for the Vth, VIIIth and IXth urosternites Urosternite V VIII IX
male female male female male female
stylet/coxite 0.58 0.51 0.82 0.84 0.99 0.74
spine/stylet 0.31 0.31 0.26 0.24 0.16 0.16
Figs. 16–20. P. iberica n. sp., male. 16. Vth urosternite; 17. VIIIth urosternite + parameres; 18. Distal part of the VIIIth paramere showing the typical setae; 19. IXth urosternite + parameres and penis; 20. Distal part of the IXth paramere showing the typical setae. Scale: 0.1 mm
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Figs. 21–32. P. iberica n. sp., female. 21. Head, frontal view; 22. Head, lateral view; 23. Outline of the eyes and ocelli; 24. Outline of a distal antenna chain; 25. Outline of the maxillary palp; 26. Ditto, of labial palp; 27. Ditto, fore leg. 28. Outline of the mid leg; 29. Ditto, hind leg; 30. Outline of Vth urosternite; 31. Ditto, VIIth urosternite; 32. Ditto, VIIIth urosternite + gonapophysis. Scale: 0.1 mm
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The VIIIth urosternite has a pair of parameres with 1+6 divisions, with tubular setae only in the apical divisions (Figs. 17–18). The IXth urosternite has the penis and parameres, the latter with 1+6 divisions and tubular setae in all the divisions except the first (Figs. 19–20). The ratio of the length of the basal to terminal parts of the penis is 1.17. Cerci have a forked apex. Description of the female Body length: 11 mm, length of antennae: 9 mm, length of cercus: 4.2 mm, length of paracercus: 9 mm. Head (Figs. 21–23) with pigmented frons apart from an unpigmented transverse band between the paired ocelli and the base of the antennae. Clypeus with little pigment. Compound eyes as long as wide. Eye ratios: cl/l: 0.33, l/w:1.07. The antennae are less robust than those of the male; distal chains with 12 divisions (Fig. 24).
Figs. 33–37. P. iberica n. sp., female. 33. Segments 23 to 27 of the VIIIth gonapophysis; 34. Last segments of the VIIIth gonapophysis; 35. Outline of the IXth urosternite + gonapophysis; 36. Segments 17 to 21 of the IXth gonapophysis; 37. Last segments of the IXth gonapophysis. Scale: 0.1 mm.
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The maxillary palp has weak pigment and lacks the field of plaited setae which is present in males on the ventral surface on the segments (Fig. 25). The labial palp is without special characteristics; a field of conules is present on the last segment, but this is smaller than in the male (Fig. 26). The three pairs of legs are robust (Figs. 27–29), but the female tarsi lack the typical spines of the male; females only have spiniform tarsal setae. Length of the tibiae: I: 0.71mm; II: 0.58mm; III: 0.85mm. Urosternites are without special features (Fig. 30). The medial part of the VIIth urosternite has a process whose length exceeds the posterior main border by a distance longer than the width of the process (ratio l/w: 1.21) (Fig. 31). The length ratios of coxite/stylet (without spine)/spine of stylets are given in Table 1. The ovipositor is of the tertiary type (Sturm & Bach 1993) and exceeds the IXth stylets by 3/5 of its length. Gonapophysis VIII has 1+49 divisions, all of which except the three basal have some setae (Figs. 32–34). Special rounded sensilla can be observed in some of the divisions. The main chaetotaxy is shown in Figs. 33 and 34. Gonapophysis IX has 1+42 divisions (Figs. 35–37), all except the basal ones, with some setae. Special rounded sensilla can be observed on some divisions. The main chaetotaxy is shown in Figs. 36 and 37.
Discussion We have compared the new species with the other four known congenerics: P. tarsispina Janetscheck, 1954, P. janetschecki Bach, 1979, P. insularis Gaju et al., 1995 and P. autumnalis Gaju et al., 1995. The new species can be easily differentiated from P.autumnalis by the special field of short and thick setae on the ventral side of the maxillary palp in the male; these are not arranged in a plaited pattern (Gaju et al. 1995a). With respect to the female, the main difference is the presence of an ovipositor of secondary type in P. autumnalis and of tertiary type in P. iberica. P. janetschecki and P. insularis are closely related and differ from the new species by their pigmentation, which is more diffuse than in P. iberica (Bach 1979; Gaju et al. 1995b). Also the fore femur of these species is, in both sexes, longer and narrower than in P. iberica. The male of P. insularis has hyaline spines on the ventral surface of the femur and tarsi of the fore leg, while P. iberica only has short and pigmented spines on the tarsi. The new species is more related to P. tarsispina, but can be differentiated by the maxillary palp which is less robust in P. tarsispina and lacks the plaited setae field (Janetschek 1954). Moreover the head pigmentation of P. tarsispina is, in both sexes, stronger with a pigmented clypeus. Derivation of the name The new species has been named P. iberica because the samples have been found in the Iberian mountains between the Meseta and the Ebro Bassin. Species habitat The specimens were collected under stones in calcareous places.
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Acknowledgements This work was supported by the Project “Fauna Ibérica III” SEUI-DGICYT PB92-0121. We would like to thank Dr. Geoff Frampton for improving English in the manuscript.
References Bach de Roca, C. (1979) Contribución al conocimiento de los Machilida de España: Descripción de una nueva especie (Insecta Thysanura). Boletín de la Real Sociedad Española de Historia Natural (Biología) 76, 171–176. Fanciulli, P.P., Bach de Roca, C., Gaju, M., Frati, F. (1995) Genetic differentiation within two genera of Microcoryphia (Insecta, Apterygota): Promesomachilis and Praemachiloides. Bulletin Entomologique de Pologne 64, 33–40. Gaju-Ricart, M., Bach de Roca, C., Mora-Carmona, R., Molero-Baltanás, R.(1995a) The genus Praemachiloides in the Iberian Peninsula: description of P. autumnalis sp. n. (Microcoryphia: Machilidae). Bulletin Entomologique de Pologne 64, 5–14. Gaju-Ricart, M., Bach de Roca, C., Molero-Baltanás, R., Mora-Carmona, R. (1995b) Descripción de Praemachiloides insularis sp. n. de las islas Baleares (Microcoryphia: Machilidae). Boletín de la Asociación española de Entomología 19 (3–4), 163–172. Janetschek, H. (1954) Ueber Felsenspringer der Mittelmeerländer (Thysanura, Machilidae). Eos 30 (3–4), 163–314. 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
Praemachiloides iberica n. sp. from Spain (Insecta Microcoryphia) Rosa Ma Mora-Carmona1, Miguel Gaju-Ricart1 and Carmen Bach de Roca2 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 1
Accepted: 30. December 1999
Summary Praemachiloides iberica n. sp. is described and compared with the other four known species of the genus: P. janetschecki, P. insularis, P.tarsispina and P. autumnalis. The main difference of P. iberica n. sp. from the other known congenerics is the presence, in the male, of a field of plaited setae on the ventral surface of the maxillary palp. Key words: Praemachiloides, Spain, new species, Microcoryphia, Apterygota
Introduction In field studies within the “Fauna Iberica” Project, we obtained samples of Praemachiloides sp. that had been studied genetically by Fanciulli et al. (1995), together with other well known species of Praemachiloides. When we studied the morphological characteristics of these Praemachiloides spp. we saw that the specimens of these populations were different, belonging to a new species that we named P. iberica n. sp.
Corresponding author Miguel Gaju-Ricart, e-mail: [email protected]
0031–4056/00/44/03–04–292 $ 12.00/0
Praemachiloides iberica
293
Materials and Methods The specimens were collected with an aspirator, then fixed with 70 % alcohol. In the laboratory, the specimens were dissected and mounted on microscope slides using Tendeiro liquid (see Molero-Baltanás et al. in this volume). The slides were then dried in an oven at 35°C.
Results Praemachiloides iberica n. sp. Studied material Gea de Albarracín (Teruel), 16-04-87, 2 males + 2 females, Ref. M0671; Royuela (Teruel), 16-04-87, 1 male + 1 female, Ref. M0670; Beteta (Cuenca), 16-02-90, 1 male + 1 female, Ref. M0676; Arcos de Jalón (Soria), 29-02-92, 4 females, Ref. M0770; Agost (Alicante), 11-04-92, 1 male, Ref. M0766; Entre Confrides y puerto de Ares (Alicante), 12-04-92, 1 female, Ref. M0772; Alhama de Murcia, Sierra Espuña, 1504,92, 3 males + 2 females, Ref. M0764; Gea de Albarracín (Teruel), 31-03-93, 1 female, Ref. M0531; Beteta (Cuenca), 01-04-93, 1 male holotype + 1 female allotype + 2 females paratypes, Ref. M0533; Arcos de Jalón (Soria), 01-04-93, 1 male, Ref. M0535; Elche de la Sierra (Albacete), 16-01-94, 4 males juv.+ 4 females juv., Ref. M0763; Yecla (Murcia), 29-03-94, 1 female, Ref. M0818; Elche de la Sierra (Albacete), 30-03-94, 4 males + 1 female, Ref. M0844. The holotype male, allotype female and paratypes are held in the “Museo Nacional de Ciencias Naturales” in Madrid; collection number: 8524. Description of the male Body length: 10.5 mm, length of antennae:10 mm, length of paracercus: 10 mm, length of cercus: 4 mm. Head (Figs. 1–2) with pigmented frons, apart from a small central line without pigment between the paired ocelli, extending to the base of the antennae. Clypeus with numerous white setae and little pigment on the lateral side. Pigmented labrum. Compound eyes as long as wide (Fig. 3). Eye ratios: contact line / length (cl/l): 0.36, length / width (l/w): 1.08. The paired ocelli are oval to subtriangular, dark-reddish in colour and located in a submedian position. The antennae are the same length as the body but can be longer. Scapus (Fig. 4) twice as long as wide with diffuse pigment. Flagellus pigmented. Distal chains of the antennae with 17 annuli which are wider than they are long (Figs. 5–6). Although some basiconic sensilla are present (Fig. 6), there is not a regular pattern of sensilla distribution. The maxillary palp (Fig. 7) shows a field of plaited setae (Fig. 9) on the ventral surface of all the segments. Hyaline spines (Fig. 8) appear on the last three segments, their distribution as follows: segment 5: 2 spines; 6: 10-16; 7:11-15. Ratio last / penultimate segments (n/n-1): 0.84. The labial palp is without any special characteristics; the second segment has subparallel sides and the third segment is typically enlarged, with a field of conules present (Figs. 10–11).
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Figs. 1–15. Praemachilodes iberica n. sp., male. 1. Head, frontal view; 2. Head, lateral view; 3. Outline of the eyes and ocelli; 4. Outline of the scapus and pedicellus of the antenna; 5. Shape of one distal antenna chain; 6. Annuli of the antenna chain, showing special sensilla; 7. Outline of the maxillary palp; 8. Last segment of the maxillary palp showing the hyaline spines; 9. Plaited setae on the ventral surface of the maxillary palp; 10. Outline of the labial palp; 11. Sensorial conules of the 3rd segment of the labial palp; 12. Outline of the fore leg; 13.Tarsus of the fore leg showing the spines of the 2nd tarsomere; 14. Outline of the mid leg; 15. Outline of the hind leg. Scale: 0.1 mm
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The three pairs of legs have weak pigmentation. The first pair has the femur enlarged (ratio l/w= 1,88) (Fig. 12). The second tarsomere has short, pigmented, ventral spines (Fig. 13). The second and third pairs of legs (Figs. 14-15) have longer spines than those of the first pair. On the tibia there are some spiniform setae. Length of the tibiae: I: 0.73 mm; II: 0.68 mm; III: 0.91 mm. Urosternites with obtusely-angled sternites; the first to seventh urosternites each posses a pair of eversible vesicles. The coxites have some setae which may vary in number from one coxite to another (Fig. 16). The length ratios between coxite/stylet (without spine)/spine of the stylets are given in Table 1.
Table 1. P. iberica n. sp. Length ratios of coxite, stylet (without spine) and spine for the Vth, VIIIth and IXth urosternites Urosternite V VIII IX
male female male female male female
stylet/coxite 0.58 0.51 0.82 0.84 0.99 0.74
spine/stylet 0.31 0.31 0.26 0.24 0.16 0.16
Figs. 16–20. P. iberica n. sp., male. 16. Vth urosternite; 17. VIIIth urosternite + parameres; 18. Distal part of the VIIIth paramere showing the typical setae; 19. IXth urosternite + parameres and penis; 20. Distal part of the IXth paramere showing the typical setae. Scale: 0.1 mm
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Figs. 21–32. P. iberica n. sp., female. 21. Head, frontal view; 22. Head, lateral view; 23. Outline of the eyes and ocelli; 24. Outline of a distal antenna chain; 25. Outline of the maxillary palp; 26. Ditto, of labial palp; 27. Ditto, fore leg. 28. Outline of the mid leg; 29. Ditto, hind leg; 30. Outline of Vth urosternite; 31. Ditto, VIIth urosternite; 32. Ditto, VIIIth urosternite + gonapophysis. Scale: 0.1 mm
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The VIIIth urosternite has a pair of parameres with 1+6 divisions, with tubular setae only in the apical divisions (Figs. 17–18). The IXth urosternite has the penis and parameres, the latter with 1+6 divisions and tubular setae in all the divisions except the first (Figs. 19–20). The ratio of the length of the basal to terminal parts of the penis is 1.17. Cerci have a forked apex. Description of the female Body length: 11 mm, length of antennae: 9 mm, length of cercus: 4.2 mm, length of paracercus: 9 mm. Head (Figs. 21–23) with pigmented frons apart from an unpigmented transverse band between the paired ocelli and the base of the antennae. Clypeus with little pigment. Compound eyes as long as wide. Eye ratios: cl/l: 0.33, l/w:1.07. The antennae are less robust than those of the male; distal chains with 12 divisions (Fig. 24).
Figs. 33–37. P. iberica n. sp., female. 33. Segments 23 to 27 of the VIIIth gonapophysis; 34. Last segments of the VIIIth gonapophysis; 35. Outline of the IXth urosternite + gonapophysis; 36. Segments 17 to 21 of the IXth gonapophysis; 37. Last segments of the IXth gonapophysis. Scale: 0.1 mm.
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The maxillary palp has weak pigment and lacks the field of plaited setae which is present in males on the ventral surface on the segments (Fig. 25). The labial palp is without special characteristics; a field of conules is present on the last segment, but this is smaller than in the male (Fig. 26). The three pairs of legs are robust (Figs. 27–29), but the female tarsi lack the typical spines of the male; females only have spiniform tarsal setae. Length of the tibiae: I: 0.71mm; II: 0.58mm; III: 0.85mm. Urosternites are without special features (Fig. 30). The medial part of the VIIth urosternite has a process whose length exceeds the posterior main border by a distance longer than the width of the process (ratio l/w: 1.21) (Fig. 31). The length ratios of coxite/stylet (without spine)/spine of stylets are given in Table 1. The ovipositor is of the tertiary type (Sturm & Bach 1993) and exceeds the IXth stylets by 3/5 of its length. Gonapophysis VIII has 1+49 divisions, all of which except the three basal have some setae (Figs. 32–34). Special rounded sensilla can be observed in some of the divisions. The main chaetotaxy is shown in Figs. 33 and 34. Gonapophysis IX has 1+42 divisions (Figs. 35–37), all except the basal ones, with some setae. Special rounded sensilla can be observed on some divisions. The main chaetotaxy is shown in Figs. 36 and 37.
Discussion We have compared the new species with the other four known congenerics: P. tarsispina Janetscheck, 1954, P. janetschecki Bach, 1979, P. insularis Gaju et al., 1995 and P. autumnalis Gaju et al., 1995. The new species can be easily differentiated from P.autumnalis by the special field of short and thick setae on the ventral side of the maxillary palp in the male; these are not arranged in a plaited pattern (Gaju et al. 1995a). With respect to the female, the main difference is the presence of an ovipositor of secondary type in P. autumnalis and of tertiary type in P. iberica. P. janetschecki and P. insularis are closely related and differ from the new species by their pigmentation, which is more diffuse than in P. iberica (Bach 1979; Gaju et al. 1995b). Also the fore femur of these species is, in both sexes, longer and narrower than in P. iberica. The male of P. insularis has hyaline spines on the ventral surface of the femur and tarsi of the fore leg, while P. iberica only has short and pigmented spines on the tarsi. The new species is more related to P. tarsispina, but can be differentiated by the maxillary palp which is less robust in P. tarsispina and lacks the plaited setae field (Janetschek 1954). Moreover the head pigmentation of P. tarsispina is, in both sexes, stronger with a pigmented clypeus. Derivation of the name The new species has been named P. iberica because the samples have been found in the Iberian mountains between the Meseta and the Ebro Bassin. Species habitat The specimens were collected under stones in calcareous places.
Praemachiloides iberica
299
Acknowledgements This work was supported by the Project “Fauna Ibérica III” SEUI-DGICYT PB92-0121. We would like to thank Dr. Geoff Frampton for improving English in the manuscript.
References Bach de Roca, C. (1979) Contribución al conocimiento de los Machilida de España: Descripción de una nueva especie (Insecta Thysanura). Boletín de la Real Sociedad Española de Historia Natural (Biología) 76, 171–176. Fanciulli, P.P., Bach de Roca, C., Gaju, M., Frati, F. (1995) Genetic differentiation within two genera of Microcoryphia (Insecta, Apterygota): Promesomachilis and Praemachiloides. Bulletin Entomologique de Pologne 64, 33–40. Gaju-Ricart, M., Bach de Roca, C., Mora-Carmona, R., Molero-Baltanás, R.(1995a) The genus Praemachiloides in the Iberian Peninsula: description of P. autumnalis sp. n. (Microcoryphia: Machilidae). Bulletin Entomologique de Pologne 64, 5–14. Gaju-Ricart, M., Bach de Roca, C., Molero-Baltanás, R., Mora-Carmona, R. (1995b) Descripción de Praemachiloides insularis sp. n. de las islas Baleares (Microcoryphia: Machilidae). Boletín de la Asociación española de Entomología 19 (3–4), 163–172. Janetschek, H. (1954) Ueber Felsenspringer der Mittelmeerländer (Thysanura, Machilidae). Eos 30 (3–4), 163–314. Sturm, H., Bach de Roca, C. (1993) On the Systematics of the Archaeognatha (Insecta). Entomologia Generalis 18(1/2), 55–90.