Aeshna brasiliensis sp. nov. (Odonata: Aeshnidae) from South and Southeastern Brazil, with a Redescription of its Larva

Aeshna brasiliensis sp. nov. (Odonata: Aeshnidae) do Sul e Sudeste do Brasil, com Redescrição de sua Larva

Abstracts

Aeshna brasiliensis (holotype <IMG SRC="http:/img/fbpe/ne/v31n3/a05img01.gif"> Rio Grande do Sul: Rio Tainha, 29º 26'60'' S 50º 34'60'' W, 900 m, MNRJ; allotype <IMG SRC="http:/img/fbpe/ne/v31n3/a05img02.gif"> Rio de Janeiro: Itatiaia, Represa do Planalto de Itatiaia, 22º 30'00"S 44º 40'00"W, 2200 m, MNRJ; nine males and 16 female paratypes) is described and illustrated from 10 males and 17 females from southeastern Brazil. The last larval instar is redescribed and diagnostic characters are illustrated. A. brasiliensis resembles A. variegata and A. peralta in the color pattern of head and abdomen, but differs from them in the thoracic color pattern and shape of cerci, from A. variegata in the shape of anterior lamina spines, and from A. peralta by its larger size and shape of male cerci. The larvae uniquely differ from all the other known Brazilian congeners by lacking lateral spines on abdominal segment VI.

Insecta; Aeshna peralta; A. variegata

SYSTEMATICS, MORPHOLOGY AND PHYSIOLOGY

Aeshna brasiliensis sp. nov. (Odonata: Aeshnidae) from South and Southeastern Brazil, with a Redescription of its Larva

NATALIA VON ELLENRIEDER¹ AND JANIRA M. COSTA²

¹Research Associate, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, CA 90007, USA

e-mail: [e-mail]

2Museu Nacional, Universidade Federal do Rio de Janeiro, Brasil, e-mail: [e-mail]

Aeshna brasiliensis sp. nov. (Odonata: Aeshnidae) do Sul e Sudeste do Brasil, com Redescrição de sua Larva

RESUMO - Aeshna brasiliensis é descrita e ilustrada de 10 machos e 17 fêmeas do Sul e Sudeste do Brasil (holótipo-Rio Grande do Sul: Rio Tainha, 29º26'60" S 50º34'60" W, 900 m, MNRJ; alótipo- Rio de Janeiro: Itatiaia, Represa do Planalto de Itatiaia, 22º30'00" S 44º40'00" W, 2200 m, MNRJ; nove machos e 16 fêmeas parátipos). O último estádio larval é redescrito e caracteres diagnósticos são ilustrados. A. brasiliensis assemelha-se a A. variegata e A. peralta no padrão de cor da cabeça e abdome, porém difere delas no padrão de cor do tórax e forma dos cercos, e de A. variegata pela forma dos espinhos da lâmina anterior do macho, e de A. peralta por ser maior e pela forma dos cercos do macho. A larva se destaca por diferenciar-se de todas as outras congêneres conhecidas do Brasil pela ausência do espinho lateral no VI segmento do abdome.

PALAVRAS-CHAVE: Insecta, Aeshna peralta, A. variegata.

ABSTRACT -Aeshna brasiliensis (holotype Rio Grande do Sul: Rio Tainha, 29º 26'60'' S 50º 34'60'' W, 900 m, MNRJ; allotype Rio de Janeiro: Itatiaia, Represa do Planalto de Itatiaia, 22º 30'00"S 44º 40'00"W, 2200 m, MNRJ; nine males and 16 female paratypes) is described and illustrated from 10 males and 17 females from southeastern Brazil. The last larval instar is redescribed and diagnostic characters are illustrated. A. brasiliensis resembles A. variegata and A. peralta in the color pattern of head and abdomen, but differs from them in the thoracic color pattern and shape of cerci, from A. variegata in the shape of anterior lamina spines, and from A. peralta by its larger size and shape of male cerci. The larvae uniquely differ from all the other known Brazilian congeners by lacking lateral spines on abdominal segment VI.

KEY WORDS: Insecta, Aeshna peralta, A. variegata.

The new species herein described was previously recorded from Brazil as Aeshna peralta Ris by Calvert (1956) and Santos (1966b). The latter provided a description of the last larval instar and measurements of the adults, based on reared specimens from Brejo da Lapa (Itatiaia). Jurzitza (1990) synonymyzed A. peralta with A. variegata Fabricius, without examining these specimens from Brazil. Muzón & Ellenrieder (1997) described the larva of A. variegata, and noted some differences with the larva described by Santos (1966b) as A. peralta. Examination of the types of A. peralta revealed it to be a good species (von Ellenrieder, unpublished), related to A. variegata and A. brasiliensis.

Material and Methods

Wing terminology follows Riek & Kukalová-Peck (1984), and larval mandibular formula follows Watson (1956). Length of pterostigma was measured from anterior costal end to posterior anal end. All dimensions are in millimeters. Type material of A. brasiliensis is deposited in the following collections: MNRJ: Museu Nacional de Rio de Janeiro, BRAZIL; MLP: Museo de La Plata, Departamento de Entomología, ARGENTINA; RWG: Dr. R.W. Garrison private collection, Azusa, USA.

Aeshna brasiliensis sp. nov.

Aeshna peralta Ris: Calvert (1956) (in part; from Itatiaia); Santos (1966a) (description of larval habitat); Santos (1966b) Figs. 1-3 (description of larva, comments and measurements of HW, abdomen and cerci length of adults); Santos (1988) (mention of larva); Muzón & Ellenrieder (1997) (comparison with larva of variegata); Carvalho & Nessimian (1998) (mention from Rio de Janeiro State); Costa et al. (2000) (mention from São Paulo State).

Aeshna sp. Ellenrieder (2001a, b)

Material examined - Holotype:

Brazil: Rio Grande do Sul: Rio Tainha-900 m, N.D. Santos leg., 20-I-1958 (MNRJ); Allotype:

Brazil: Rio de Janeiro: Itatiaia, Represa do Planalto de Itatiaia, N.D. Santos leg., 13-I-1953 (MNRJ # 21.958); Paratypes: Brazil: Rio de Janeiro: 1

8

same data as allotype (MNRJ # 21.928-31, 21.951, 52, 54, 56) - 1

(MLP # 21.955), 2

1

Itatiaia, Brejo da Lapa, 2200 m, N.D. Santos & J.P. Machado leg., 12-X-1967 (MNRJ) - 2

(MLP, RWG); 1

1

same data except N.D.Santos & Duílio leg., 2-III-1976 (MNRJ); São Paulo: 1

Ypiranga, J.G. Fonseca leg., III-1927 (MNRJ # 21976); 1

Campos do Jordão, Umuarama, 1700 m, Ebert leg., I-1966 (MNRJ); Paraná: 1

Curitiba, XI-1941 (MNRJ # 22011); 1

União da Vitória, V. Staviarski leg., I-1952 (MNRJ # 22046), 1 Bituruna, V. Staviarski leg., III-1966 (MNRJ); Rio Grande do Sul: 1

Rio Grande, Rio Lajeado Grande, 30-I-1982, N.D. Santos, J.M. Costa, L.F. Netto & H. Mesquita leg., 30-I-1982 (MNRJ), 1

Riacho do Arroio, km 113 entre S. Francisco de Paula e Rio Tainha, N.D. Santos leg., 9-XI-1967 (MNRJ), 1

Rio Grande do Sul, S. Francisco de Paula, Rio Tainha, J.A. Petersen leg., 5-X-1957 (RWG). Larvae: Brazil: Rio de Janeiro: Itatiaia, Brejo da Lapa - 2.200 m, 5

1

(larvae), N.D. Santos, J.M. Costa & J.P.Machado leg., 4-VII-1966 (MNRJ); same data except 2

1

(exuviae), 23.VII.1966 (MNRJ)- 2

(larvae), N.D.Santos & J.P.Machado leg. , 12-X-1967 (MNRJ)- 1

2

(exuviae), N.D. Santos leg., 29-VII-1968 (MNRJ)- 1

2

(exuviae), N.D. Santos leg., 10.XI.1968 (MNRJ)- 7

4

(larvae), N.D. Santos leg., 20-IV-1976 (MNRJ)- 1

2

(larvae), N.D. Santos leg., 21-IV-1976 (MNRJ).

Description

Holotype Male. Head. Labium pale brown; labrum yellow, with basal and distal margins dark brown; clypeus and frons yellow, top of frons with T-spot complete (Fig. 1A), with stem slightly widened basally; a narrower area of yellow followed by a dark-gray rounded spot on each side of the T-spot; wide black stripe along frontoclypeal groove (Fig. 1B); frontal carina (anterior margin of T-spot) angulate in frontal view (Fig. 1B). Blackstripe along ocular groove, abruptly widening towards frontoclypeal groove (Fig. 1C); vertex yellow with black margins; occipital triangle yellow, rear of head black.

Thorax.Pterothorax reddish brown with pale stripes and spots (Fig. 1D): Mesepisternum with incomplete, yellow and narrow antehumeral stripe at basal 0.5, mesepimeron with a stripe at basal 0.75, yellow, sinuous, and narrowed at its medial length, and with a round small yellow spot at its upper anterior edge; metepimeron with a stripe divided into yellow basal round spot and distal T-shaped white spot. Venter reddish brown with a medial round yellow spot. Legs: black, basal 0.60 of femora red brown. Wings (Fig.1H): Hyaline, venation black except for yellow costa. Pterostigma dark brown dorsally and yellow ventrally, covering three cells. Membranula basal half white, distal half black.

Venation. (range; holotype values in parenthesis). Antenodal crossveins FW: 12-17 (15); antenodal crossveins HW: 8-11 (11); cells triangle FW: 4-5 (5); cells triangle HW: 4-5 (4); crossveins supratriangle FW: 1-3 (2); crossveins supratriangle HW: 0-3 (2); cells subtriangle FW: 1-2 (2); cells subtriangle HW: 2; crossveins of Cu-A space FW: 3-4 (4); bridge crossveins FW: 2-4 (4 right, 3 left); bridge crossveins HW: 2-4 (3); row of cells enclosed in IRP2 fork at pterostigma level FW: 2-4; row of cells enclosed in IRP2 fork at pterostigma level HW: 2-4 (3-4); cells anal loop: 7-10 (9 right, 10 left); row of cells in anal loop: 3, 2 on costal side; cells anal triangle: 3.

Abdomen. Reddish brown with black areas and pale maculation (color not recognizable due to bad preservation), as shown in Fig. 2 A-B. Ventral terga as shown in Fig. 2E. Medial tubercle of sternum I well developed, bearing small denticles; auricles of segment II each with 2 teeth; genital lobes triangularly ridged, with 15-20 small denticles on posterior end (Fig. 3A); anterior process of anterior hamulus very short, forming a small mesally pointed tip (Fig. 3B paratype); spine of anterior lamina laterally compressed surpassing caudal end of hamular fold, with pointed tip and ventral margin approximately linear (Fig. 3C paratype); distal segment of vesica spermalis with well developed latero-posterior and dorsal folds (Fig. 3EF). Cerci gradually widening at 0.50 of length; maximum width at distal 0.30; tips mucronate (Fig. 4A); subbasal tooth small; dorsal crest well developed at distal 0.2 (Fig. 4B).

Variation (Paratypes). Similar to holotype, pale stripes of pterothorax variable in extension (Fig. 1D, F, G), mesepisternal stripe extending from basal 0.25 to 0.50; mesepimeral from basal 0.50 to 0.75, and metepimeral continuous or divided into two spots. In some specimens the color of pale abdominal spots is recognizable: Dorsal spots light blue except those from abdominal segment X, which are yellow.

Measurements (mm) (Mean and standard deviation; number of specimens; range in square brackets; holotype values in parenthesis): Head maximum width: 8.45 ± 0.12, N=7 [8.3- 8.6] (8.3); frons max. width: 4.23 ± 0.12, N= 8 [4.1-4.5] (4.2); forewing length: 39 ± 0.97, N= 9 [38-40] (40); hindwing length: 37.73 ± 1.03, N= 9 [35.7-39.2] (38.5); forewing width: 9.65 ± 0.33, N= 10 [9.3-10.2] (9.6); hindwing width: 12.04 ± 0.43, N= 10 [11.5-12.7] (11.7); forewing pterostigma length: 3.16± 0.21, N= 10 [2.8-3.4] (3.33); hindwing pterostigma length: 3.04 ± 0.18, N= 10 [2.8-3.3] (3.1); cerci length: 4.77 ± 0.16, N= 9 [4.58-5.1] (4.75); epiproct length: 2.37 ± 0.16, N= 10 [2.16-2.66] (2.16); abdominal segment III length: 7.02 ± 0.36, N= 9 [6.5-7.66] (7.66); total length: 61.13 ± 0.97, N= 8 [59.9-63] (63).

Female (Allotype). Head and thorax as in holotype except mesepisternal stripe at basal 0.30, and metepimeral stripe narrow at mid-length but not divided into two spots, distal half light blue (Fig. 1E).

Abdomen. As in holotype (Fig. 2C-D); ventral terga as shown in Fig. 2F. Cerci lanceolate; maximum width at medial 0.30; tips mucronate (Fig. 4E).

Variation (Paratypes). Similar to allotype (Fig. 1E), pale stripes of pterothorax variable in extension, mesepisternal stripe extending from basal 0.25 to 0.50; mesepimeral from basal 0.50 to 0.75, and metepimeral continuous or divided into two spots. In some specimens the color of pale abdominal spots is recognizable: Dorsal spots are light blue and lateral spots yellow.

Measurements (mm) (Mean and standard deviation; number of specimens; range in square brackets; allotype values in parenthesis): Head maximum width: 8.65 ± 0.21, N= 16 [8.3-9] (8.6); frons max. width: 4.37 ± 0.13, N= 17 [4.1-4.5] (4.5); forewing length: 41.02 ± 0.9, N= 17 [39.5-43] (40); hindwing length: 39.37 ± 0.84, N= 17 [38-41] (39.5); forewing width: 9.98 ± 0.3, N= 17 [9.24-10.7] (9.7); hind wingwidth: 13.09 ± 0.41, N= 17 [12-14] (12.7); forewing pterostigma length: 3.69 ± 0.23, N= 17 [3.2-4] (3.6); hindwing pterostigma length: 3.5 ± 0.25, N=[3.1-4]; cerci length: 5.15 ± 0.3, N= 14 [4.6-5.6] (5.2); cerci max. width: 1.21 ± 0.04, N= 14 [1.15-1.3] (1.2); abdominal segment III length: 6.86 ± 0.25, N= 17 [6.5-7.4] (6.7); total length: 63.03 ± 1.61, N= 13 [60.2-65.5] (62.3).

Venation (range; allotype values in parenthesis): Antenodal crossveins FW: : 13-17 (15 right, 14 left); antenodal crossveins HW: 8-12 (10); cells triangle FW: 3-5 (4); cells triangle HW: : 4-6 (4); crossveins supratriangle FW: 0-3 (2 right, 3 left); crossveins supratriangle HW: 0-2 (1 right, 2 left); cells subtriangle FW: 1-2 (2); cells subtriangle HW: 1-2 (2); crossveins of Cu-A space FW: : 3-4 (4); : 3-5 (4 right, 3 left); crossveins of Cu-A space HW: : 2-4 (4 right, 3 left); bridge crossveins FW: 2-4 (3); bridge crossveins HW: 2-4 (3 right, 2 left); row of cells enclosed in IRP2 fork at pterostigma level FW: 2-4; row of cells enclosed in IRP2 fork at pterostigma level HW: 2-4 (3-4); cells anal loop: 7-13 (9 right, 10 left); row of cells in anal loop: 3, 2 on costal side.

Redescription of the Last Larval Instar

Head (Fig. 5A): Approximately one and a half to two times wider than long. Occipital margin concave, with five pilose patches on each side. Antennae (Fig. 6E) 7-segmented, the third antennomere the longest. Labium reaching caudad posterior margin of second coxae; prementum as wide as 0.9 its length, border of medial lobe with fringe of setae and with two small tubercles, one on each side of median cleft, both shorter than setae. Labial palp (Fig. 6G) with a small infraapical tooth, inner margin serrated, with 16 denticles; movable hook 1.5 times as long as the inner margin. Mandibular formula (Fig. 6A-D): L 1234 0 a b k/ R 1234 y a a' b k

Thorax: Prothoracic supracoxal processes with pointed apex, base of the anterior one as broad as posterior one (Fig. 5BC); wing pads nearly parallel, the external reaching caudad basal third of abdominal segment V; femora and tibiae without dark annular bands (Fig. 5A).

Abdomen: Widest on segments V-VIII . Dorsal color pattern as in Fig. 5A. Lateral spines present on segments VII-IX,the VIII the longest. Female gonapophyses (Fig. 5E) not reaching the posterior margin of segment IX. Cerci (Fig. 5D) slightly shorter than epiproct, epiproct with middorsal ridge and two apical short spines; male basal lamina with blunt tip, as long as 0.45 of epiproct.

Habitat: The larvae were collected at small lotic pools formed by cascading waters characteristic of the highlands within the region of Itatiaia. These pools contained abundant aquatic vegetation, with abundant Amphipoda (Hyallela sp.) and larval Trichoptera, Ephemeroptera, Aeshna punctata Martin (Aeshnidae) and Oxyagrion simile Costa (Coenagrionidae).

Measurements (mm) (Mean and range in square brackets, N= 17

14

; unless stated, measurements are for both

and

): Total length (without caudal appendages):

27.5 [27-28];

: 28.5 [27-30]; head maximum width: 7.5 [7-8], head maximum length:

4.5 [4-5],

4.25 [4-4.5]; prementum maximum length: 5.5 [5-6]; prementum maximum width: 4.75 [4.5-5]; palp movable hook: 1.8 [1.6-2]; antennomere I: 0.5; II: 0.7; III: 1.7; IV: 0.6; V: 0.8; VI: 0.6; VII: 1.0; femur I length: 3.52 [3.3-3.75], femur II: 4.75 [4.5-5], femur III: 6.1 [5.6-6.6]; tibia I length: 4.2 [4-4.4], tibia II: 4.65 [4.3-5], tibia III: 5.1 [5- 5.2]; internal wing pads length: 7.9 [7.5-8.3], external: 7.7 [7.2-8.2]; maximum length of abdominal segment V: 1.8, VI: 1.7, VII: 1.7 , VIII: 1.7, IX: 1.3, X: 1.2; lateral spines (inner margin) on segment VII: 0.5 , VIII: 0.9, IX: 0.7; epiproct length: 3.1; cerci (external margin): 2.8; paraprocts: 4; inner gonapophyses (inner margin): 2.

Distribution (Fig. 7): Brasil: Rio de Janeiro: Itatiaia, Represa do Planalto de Itatiaia; Itatiaia, Brejo da Lapa, 22º30'00"S 44º40'00"W, 2200 m. São Paulo: Ypiranga, 23º32'52"S 46º38'07"W, 760 m; Campos do Jordão, Umuarama, 22º 44'45"S 45º 34'47"W, 1700 m. Paraná: Curitiba, 25º 25'00"S 49º 15'00"W, 913 m; União da Vitoria, 26º 13'00"S 51º 04'60"W, 752 m; Bituruna, 26º 10'00"S 51º 34'00"W, 914 m.Rio Grande do Sul: Rio Tainha-900 m; San Francisco de Paula, Rio Tainha; Riacho do Arroio, km 113 entre S. Francisco de Paula e Rio Tainha; Rio Lajeado Grande, 29º 26'60"S 50º 34'60"W, 900 - 922 m.

Discussion

Aeshna brasiliensis was previously recorded as Aeshna peralta (Calvert 1956, Santos 1966b) probably because of their overall similarity. It is easily distinguished from A. peralta and A. variegata by the shape of male cerci (gradually widened in brasiliensis with maximum width at distal 0.30 - >abruptly so in peralta and variegata with maximum width at basal 0.30; Fig. 4A, 4C), female cerci (tips mucronate in brasiliensis – rounded in A. peralta and A. variegata), and by the thoracic color pattern (mesepimeral and metepimeral stripes narrowed or interrupted at mid-length and pale spot at upper anterior edge of mesepimeron in brasiliensis - stripes not narrowed at midlength in peralta and variegata and no pale mesepimeral spot). It differs also from A. peralta by its larger size (Ellenrieder, unpublished): Total length

59.9-63 versus 51.75-56.5,

60.2-65.5 versus 51.5-57; cerci length

4.58-5.1 versus 3.75-4.5; cerci length

4.6-5.6 versus 3-4.5, and by the shape of female cerci (inner and outer margins converging gradually in distal half in peralta, approx. parallel in brasiliensis). The shape of the spines of the male anterior lamina separate it also from A. variegate (ventral margin straight in brasiliensis, convex in variegata; Fig. 3CE), as well as the degree of development of the subbasal tooth of the male cerci (small in brasiliensis - prominent in variegata; Figs 4B, 4D). The last larval instar of Aeshna brasiliensis differs from that of all the other known Aeshna larvae from Brazil (A. bonariensis, A. confusa, A. cornigera and A. punctata) and from A. variegata by the absence of lateral spines on abdominal segment VI; it differs from A. variegata also by the absence of dark annular bands in the legs (present in variegata); its smaller size (body length 32.8/37 mm in variegata – 27/ 30 mm in brasiliensis), shorter cerci (2-2.55 mm in variegata – 2.8 mm in brasiliensis) and length of anterior/ posterior ocular margin (0.66-0.8 in variegata – 0.8/0.82 in brasiliensis). The larva of A. peralta is still unknown.

Aeshna brasiliensis is thus far know from southeasternt and south Brazilian localities (Fig. 7), all within the Paranaense biogeographic province (Cabrera & Willink 1980). Records comprise a south-eastern strip from 22º30' to 29º 26' S and 44º40' to 51º34'W, between 750 and 2200 m above sea level.

Acknowledgments

We thank Dr. R.W. Garrisson for critically reading the manuscript and the Fundación Antorchas (Argentina), the CNPq and FAPERJ (Brazil) for their support.

Literature Cited

Recebido 20/08/01. Accepted 15/07/02.

• Cabrera, A.L. & A. Willink. 1980 Biogeografía de América Latina. Secretaría General de la O.E.A. (Biol.), Washington D.C.,13: vi + 122p.
• Calvert, P.P. 1956 The neotropical species of the "subgenus Aeshna" sensu Selysii 1883 (Odonata). Mem. Am.Entomol. Soc. 15: 1-251.
• Carvalho, A.L & J.L. Nessimian. 1998 Odonata do Estado do Rio de Janeiro, Brasil: hábitats e hábitos das larvas. Ecol. Insetos Aquáticos 5: 3-28
• Costa, J.M., A.B.M. Machado, F.A.A.Lencioni & T.C. Santos. 2000 Diversidade e distribuiçăo dos Odonata (Insecta) no estado de Săo Paulo, Brasil: Parte I- Lista das espécies e registros bibliográficos. Publ. Avul. Mus.Nac. Rio de Janeiro 80: 1-27.
• Ellenrieder, N. von. 2001a A synopsis of the Patagonian species of the genus Aeshna Fabricius (Anisoptera: Aeshnidae). Odonatologica 30: 299-235.
• Ellenrieder, N. von. 2001b The Larvae of Patagonian Aeshna Fabricius species (Anisoptera: Aeshnidae). Odonatologica 30: 423-434.
• Jurzitza, G. 1990 Aeshna peralta Ris, 1918, ein Synonym von A. variegata Fabricius, 1775 (Anisoptera: Aeshnidae). Odonatologica 19: 385-393.
• Muzón, J. & N. von Ellenrieder. 1997 Estadios larvales de Odonata de la Patagonia. I. Descripción de Aeshna variegata Fabricius (Odonata: Aeshnidae). Rev. Soc. Entomol. Argent. 56: 143-146.
• Riek, E.F. & J. Kukalová-Peck. 1984 A new interpretation of dragonfly wing venation based upon early Carboniferous fossils from Argentina (Insecta: Odonatoidea) and basic characters states in pterygote wings. Can. J. Zool. 62: 1150-1166.
• Santos, N.D. 1966a Notas sobre Aeshna (Hesperaeschna) punctata Martin, 1908 e sua ninfa (Odonata, Aeshnidae).Atas Soc. Biol. Río de Janeiro 10: 97-100.
• Santos, N.D. 1966b Notas sobre Aeshna (Hesperaeschna) peralta Ris, 1918 e sua ninfa (Odonata, Aeshnidae). Atas Soc. Biol. Río de Janeiro 10: 123-124.
• Santos, N.D. 1988 Catálogo bibliográfico de ninfas de odonatos neotropicais. (Acompanhado de relaçăo alfabética de autores e seus trabalhos). Acta Amazonica 18: 265-350.
• Watson, M.C. 1956 The utilization of mandibular armature in taxonomic studies of anisopterous nymphs. Trans. Amer. Entomol. Soc. 81: 155-205.

Publication Dates

• Publication in this collection
  27 Nov 2002
• Date of issue
  July 2002

History

• Accepted
  15 July 2002
• Received
  20 Aug 2001