Fundam. appl. NemaLOI., 1994,17 (4),303-321 Studies on the genus Amphimermis (Nematoda : Mermithidae) : five new species, including four from Orthoptera in southeastern Australia Graeme L. BAKER* and George O. POINAR,jr.** *' Entornology BTanch, Biological and Chemical Research Institute, NSW Agriculture, PM.B. 10, Rydalmere, N.s. W. 2116, Australia and *'*' Division ofEntomology and Parasitology, College of Natural Resources, University of California, Berheley, CA 94720, U.S.A. Accepted for publication 24 September 1993. Summary - Five new species of Amphimerrnis are designated, of which four are described, namely A. buraki n. sp., a parasitoid of the tettigoniid Corwcephalus sp.; A. acridiorum n. sp., A. mirabinda, n. sp. and A. auslraloekgam n. sp., parasitoids of acridids incJuding wingless grasshopper, Phaulacridium villalum (Sj6stedt), eastern plague locust, Oedakus auslra/is (Saussure), and Australian plague locust, ChorLOieeles urminifera (Walker) and, in the case of A. auslraloekgans n. sp., the larval stage of a scarab, Serieeslhis sp. The males of A. buraki n. sp. and A. acridiorum n. sp. resemble A. avolula Rubtsov & Koval but differ in size, configuration of spicule twisting and position of amphids. The male of A. mirabinda n. sp. resembles A. bogongae Welch but is smaller. The male of A. auslraloekgans resembles A. elegam (Hagmeier) and A. LOngaensis Spiridonov, but is larger than both and further differs from A. ekgans in the morphology of the tail and A. LOngaensis in the size of the amphids. Material identified as A. elegam by Artyukhovski and Kharchenko has been designated A. arlyukhovskii n. sp. Keys to males and females are provided in which thirteen previously described species are recognised as valid. Species challenged are A. dolycoris Rubtsov, A. (?) pardonensis Rubtsov and A. elongala Rubtsov which have been declared species inquireruiae. A. ghi/arovi (Polozhentsev & Arryukhovski) has been synonymised with A. elegans (Hagmeier). A. unka, sensu Li has been designated nomen dubium. Résumé - Étude du genre Amphimennis (Nernatoda : Mermithidae) : cinq nouvelles espèces dont quatre provenant d'Orthoptères du sud-est de l'Australie - Cinq nouvelles espèces d'Amphimermis sont désignées dont quatre sont décrites: A. buraki n. sp., parasitoïde de Tettigoniide Conocephalus sp., A. acn'diorum n. sp., A. mirabinda n. sp. et A. auslraloelegans n. sp., parasitoïdes d'Acridides dont Phau/acridium villalum (Sj6stedt), Oedakus auslralis (Saussure) et Chorloiceles lerminifera (Walker), ainsi que, dans le cas d'A. auslraloekgans n. sp., le stade larvaire d'un carabe, Serieeslhis sp. Les mâles de A. buraki n. sp. et de A. acridiorum n. sp. ressemblent à ceux de A. avolzlla Rubsov & Koval mais en diffèrent par la taille, le type de courbure des spicules et la position des amphides. Les mâles de A. mirabinda n. sp. ressemblent à ceux de A. bogongae Welch mais sont plus petits. Les mâles de A. ausLTaloelegans n. sp. ressemblent à ceux de A. ekgans (Hagmeier) et A. LOngaensis (Spiridonov) mais ils sont plus longs dans les deux cas et de plus diffèrent de A. ekgans par la forme de la queue et de A. longaensis par la taille des amphides. Les spécimens identifiés par Arryukhovski et Karchenko comme A. elegans sont considérés comme une nouvelle espèce, A. arlyukhovskii n. sp. Une clé des mâles et une clé des femelles sont proposées dans lesquelles treize espèces déjà décrites sont reconnues valides. Les espèces mises en question sont A. dolycoris Rubtsov, A. ('J pardonensis Rubtsov et A. elongala Rubtsov qui avaient été déclarées species inquirendae. A. ghilorovi (Polozhentrev & Arryukhovski) est synonymisé à A. elegans. A. unka semu Li est considéré comme nomen dubium. Key-words : Mermithidae, Amphimermis, parasitoids, Acrididae, Scarabaeidae. The genus Amphimennis was previously known in Australia only by Amphimennis bogongae Welch, 1963, a parasite of the bogong moth, Agrotis infusa (Boisd.) (Lepidoptera: Noctuidae). The majoriry of species described in this paper are parasitic on orthopterans. Poinar (1975), in a review of nematode-host associations, lists only one species of the genus, Amphimennis elegans (Hagmeier, 1912), parasitising orthopterans, namely Stenobothrus sp. (Acrididae) and Declicus sp. (Tettigoniidae) in Europe. Since the review by Poinar, Amphimennis bonaerensis Miralles & ISSN 1164-5571194/03 S 4.00/ © Gaulhier-Vil/ars - ORS TOM Camino, 1983, has been recorded paraSltlSlng Laplatacris dispar Rehn (Acrididae) in Argentina. Amphimermis spp. appear to be less important as parasitoids of orthopterans as speeies in the genera Mennis, Hexamermis and Agamennis whose role in the control of orthopterans has been reviewed (Webster & Thong, 1984; Street & McGuire, 1990). However, recent stuilles in Australia (Baker, 1986) have indicated that parasitism by Amphimennis spp. is a major influence in the population dynamies of grasshoppers in the moist eastern sub-coastal tablelands. Atthough Mennis and Agamennis are repre303 G. L. Baker & G. O. Poinar Jr. sented in Australia by species which are also parasitoids of orthopterans (Baker, 1986; Baker & Poinar, 1986, 1988), they are generally less abundant or have a restricted distribution rendering them less effective than species of Amphimermis as biological control agents. The genus Amphimermis is here recorded parasitising coleopteran larvae in Australia for the first time. Other mermithid genera, notably Psammomermis, appear far more important in the biological control of coleopterans in this region (A. J. Campbell, pers. comm.). A. elegans is the only species of Amphimermis previously recorded parasitising coleopterans, namely Melolontha melolontha (L.) (Scarabaeidae) (Blunck, 1939) and Lepllnotarsa decemlineata Say (Chrysomelidae) (Kaiser, 1972) in Europe. Materials and methods The specimens used in the descriptions in this paper were collected during a study of the population dynamics of the wingless grasshopper, Phaulacridium viuatum (Sjbstedt). In order to identify the species of nematodes involved, adult specimens suitable for taxonomic study were obtained by rwo methods: i) natural emergence from host and retention in the laboratory in aerated distilled water until after the final moult; il) sampling from soil in pasturelands with a history of parasitism of acridids by mermithid nematodes. Infective juveniles were obtained when possible by hatching eggs laid in the laboratory. Prior ta examination, specimens were genùy heat killed, fixed in 3 per cent formalin and processed to glycerin by slow evaporation of Lee's solution (75 % absolu te ethyl alcohol, 20 % water and 5 % glycerin). During the survey, four species of Amphimermis new to science were recorded and are herein described. The terminology foUows that of Poinar (1983). Marerial has been deposited in the Departmem of Nematology, University of California, Davis, USA (UCD), South Australian Museum, Adelaide, Australia (SAN!.) and Laboratoires de Biologie Parasitaire, Protistologie, Helminthologie, Muséum National d'Histoire Naturelle, Paris, France (MNHN). Amphim.ermis Kaburaki & Imamura, 1932 = Complexomermis Filipjev, 1934 DrAGNOSrS (Kaiser, 1991 emend.) Usually medium to long nematodes, 13-260 mm in length. Mouth opening terminal or with slight ventral shift. Amphids large, usually with porelike aperture. Six head papillae lateral and sub-medial in position. Cuticle thick with clearly visible criss-cross fibres. Longitudinal chords in midbody, six. Pharyngeal tube long, in sorne species reaching 60 % of body length. PharyngeaJ tube does not reach the mouth opening but ends behind it with a ringlike thickening. Tail tip in both sexes blunùy rounded and ventrally curved. Male: with rwo long, rwisted spicules. Genital papillae in three rows, medial 304 row bifurcates around genital opening. Female: vagina elongated, S-shaped, strongly muscularized, often with vulval flap. Preparasites do not amputate tail and have extremely high mobility. Eggs numerous, medium-sized, without byssi. TYPE SPECIES Amphimermis zuimushi Kaburaki & Imamura, 1932. OTHER SPEClES A. acridionlm n. sp. A. artyukhovskii n. sp. =A. elegans aplld Artyukhovski & Karchenko, 1965. A. australoelegans n. sp. A. avolula Rubtsov & Koval, 1975. A. bogongae Welch, 1963. A. bonaerensis Miralles & Camino, 1982. A. bl/raki n. sp. A. elegans (Hagmeier, 1912) WeI ch, 1963 Mermis elegans Hagmeier, 1912. Complexomermis elegans (Hagmeier, 1912) Filipjev, 1934. = Complexomermis ghilamvi Polozhentsev & A.rtyukhovski, 1958. = A. ghilarovi (Polozhentsev & Artyukhovski, 1958) Welch, 1963 n. syn. (male). A. lagidzae Rubstsov, 1975. A. litoralis Artyllkhovski & Karchenko, 1971. A. longiscaplls Rubtsov, 1976. A. maritima Rubstov, 1971. A. rnirabinda n. sp. A. mongolica Rubtsov, 1976. A. tinyi Nickle, 1972. A. IOngaensis S piridonov, 1987. A. volubilis Rubtsov & Koval, 1975. SPEClES INQUTRENDAE A. dolycon's Rubtsov, 1974. A. elongala Rubtsov, 1978. A. (?) pardonensis Rubtsov, 1977. REMARKS In reviewing the genus, the male of A. ghilarovi (= Complexornermis ghilarovl) has been designated the lectotype and the species synonymised with A. elegans : the male of A. ghilarovi is similar to A. elegans and the species was differentiated from A. elegans on the basis of differences berween the females, however given the difference in amphid shape berween males and females designated as A. ghilarovi (elegans like in the male and bogongae like in the female) the extreme sexual dimorphism would indicate that males and females have been mistakenly considered conspecific. The females described as A. ghilarovi are not regarded as A. elegans and as a consequence of the proposed synonymy are unassigned. Artyukhoski (1969) transferred female Anzphlnzermis ghilarovi to the genus Amphibiomermis Artyukhoski, 1969 as a n. comb. and indicared the male of Fundam. appl. Nematol. Amphimermis spp. Amphibiomennis ghilarovi was unknown. It is not clear if Artyukhovski (1969) intended male Amphimermis ghilarovi to be unassigned or to remain as a valid species, the females of which were unknown. Species have been declared species inquirendae on the basis that their descriptions were based on post-parasitic juveniles. Amphimennis unka, sensu Li (1981, 1985) is designated nomen dubium being either an erroneous reference to Agamennis unka Kaburaki & Imamura, or an unidentified species of Amphimennis recorded from the same host (Nilaparvala lugens Stal.) by Chen and Yang (1985) and Xia (1989). Agamennis unka has been recorded parasitising N. lugens in China (Wang & Li, 1987) and Korea (Choo el al., 1989). GROUPS Males of the described species of Amphimennis divide readi!y into four groups on the basis of the twisting of the spicule and shape of amphids (Fig. 15). Described species and new species described in this paper have been assigned as foUows : volubilis group: spicule twisted entire length; cup shaped amphids; A. volubilis. avolula group: proximal half of spicule untwisted; cup shaped amphids; A. avohua, A. cun'diorum n. sp. and A. buraki n. sp. bogongae group: spicule twisted loosely distal and proximal half, untwisted in midclle; cup shaped amphids; A. bogongae, A. marùima, A. lùoralis, A. tinyi, A. bonaerensis, A. mirabinda n. sp. elegans group: spicule twisted tightly distal and proximal half, untwisted in midclle; thin-walled, pocket-like amphids; A. elegans, A. zuimushi, A. LOngaensis) A. anyukhovskii n. sp., A. auslraloelegans n. sp. Summaries of the distribution and host range of Amphimennis spp. have been given by Rubstov (1978 a) and Kaiser (1991) and are further elaborated in Table 1. Arnphirnennis acridiorurn n. sp. (Figs 1-5) MEASUREMENTS Hololype (female) : L = 293 mm; mid-body diam. = 340 [Lm; head width (at level of cephalic papillae) = 75 [Lm, (at neck) = 80 [Lm; body diam. at nerve ring = 150 [Lm; cuticle width (at nerve ring) = 20 [Lm, (at midbody) = 50 [Lm; hypodermis width (mid-body) = 15 [Lm; amphid aperture = 5 [Lm; amphid pouch (in dorsal view) = 18 x 12 [Lm; dist. nerve ring/mou th = 310 [Lm; V 50.2; length of vagina (from vulva to junction with uterus in lateral view) = 1.2 mm, (general length in dorsal view) = 670 [Lm; diam. vagina = 160 [Lm; length ofvulval t1ap = 250 [Lm; height ofvulval cone = 70 [Lm; width lateral hypodermal chord = 45 [Lm; ruam. egg in uterus 100 x 105 [Lm. Allolype (male): L = 107 mm; mid-body diam. = 220 [Lm; head diam. (at level of cephalic papillae) = = = Vol. 17, n° 4 - 1994 = 72 J.1.m, (at neck) = 78 [Lm; body diam. at nerve ring 12 J.1.m, (mid120 J.1.m; cuticle width (at nerve ring) body) = 15 J.1.m; hypodermis width (mid-body) = 12 J.1.m; amphid aperture = 6 J.1.m; amphid pouch = 20 x 14 J.1.m (in dorsal view), 20 x 20 J.1.m (in lateral view); dist. nerve ring/mouth = 350 J.1.m; spicule length = 2.7 mm; position of twisting = distal 70 % of length; spicule head width = 25 J.1.m; width mid-shaft = 7 J.1.m; tail length = 350 J.1.m; tail diam. at cloaca = 210 J.1.mj position of proximal genital papilla anterior to cloaca = 835 J.1.m; number of gerutal papillae in medial row (anterior to cloaca) = 35, (posterior to cloaca) = 18. Paralypes (females and males) : see Tables 2 and 3, respectively. Juveniles, SI. 2 (preparasitic; progeny of holotype; n = 10) : L = 1.401 mm (1.27-1.46); mid-body diam. = 14.6 J.1.m (12-16); head width = 9-10 J.1.m; buccal cavity length 38.5 J.1.m (32-44); stylet 24.3 J.1.m (21-26); distance nerve ring mouth = 101 [Lm (92-107); body diam. at nerve ring = 15.3 J.1.m (14-16) j length of stichosorne = from 13.03 % (11.8-14.1) to 61.1 % (59.4-66.6) body length; body width mid-stichosome region 15.4 J.1.m (14-16), mid-trophosome = 10 J.1.m (nil range); taillength = 50-60 J.1.m; tail di am. (50 J.1.m from tip) 4 [Lm (ni! range). = = = = = DESCRlPTION General: Long nematodes (longest species in the genus, cf. A. bogongae), females equal or up to 16 x length of males (males 28-144 mm and females 75-457 mm in length). Cuticle with cross fibres. Head bluntly rounded. Mouth terminal. Four sub-medial cephalic papillae, two lateral cephalic papillae. Operung of lateral cephalic papillae at level of or slightly anterior to level of sub-medial cephalic papillae. Moderate sized amphids, cup shaped, larger in male than female; opening of amphids posterior to opening of lateral cephalic papillae and slightly offset. Six hypodermal chords; lateral hypodermal chord three cells wide. Wall of anterior end of oesophagus thickened to produce a short pharynx. Females: Operung of vulva a transverse slit. Vulval flap present. Vulval cone present; base not muscular. Horse-shoe shaped build up of vulvar concretion often present in mature specimens. Vagina S-shaped, long, muscular; posterior loop equal to or marginally greater than length of anterior loop; bends rounded. Junction of vagina and uterus at level of or slightly posterior to vulva. Tail conical, slightly flattened on ventral surface. Vestigial anus sometimes present. Eggs in single or double row in uterus and two abreast (four in one plane). Eggs unembryonated when laid. Males: Tail curled into ring, conoid, bluntly rounded. Spicule paired; tightly twisted for distal two-thirds of length to within one-tenth oflength from tip; head flared syrnmetrically, bulbous; walls thick, especially in head region; spicule length approximately 12 x body width at 305 G. L. Baker & G. O. Poinar Jr Table 1. Dism'bution and hoSI range of Amphimermis spp. Group/species Distribution volubilis group A. volubilis Ukraine COLEOPTERA : Chrysomelidae Leplinolarsa decemlineata Say Rubtsov & Koval, 1975 avolUla group A. avoluta Ukraine COLEOPTERA : Chrysomelidae Leplinotarsa decemlineata Say ORTHOPTERA: Acrididae Phau/acridium viltalum (Sjostedt) Oedaleus australis (Saussure) ChorlOicetes tenninifera (WaIker) ORTHOPTERA : Tettigonüdae Conocephalus sp. Rubtsov & KovaI, 1975 A. acridiorum n. sp. AustraIia A. buraki n. sp. Australia bogongae group A. bogongae A. marilima A. lilOralis A. linyi Australia Russia (Primorsk Region) Russia (Voronezh Region) USA A. bonaerensis Argentina A. mirabinda n. sp. Australia elegans group A. elegans Germany Austria Kirgizia Russia (Kuibyshev region) Europe A. arlyukhovskii n. sp. A. zuimushi A. tongaensis A. auslraloelegans n. sp. 306 Russia (Voronesh region) Japan Tonga Australia Host Authority Present study Present study LEPlDOPTERA : Noctuidae Agrolis infusa (Boisd.) Unknown Rubstov, 1971 Unknown Artyukhovski & Kharchenko, 1971 ODONATA: Coenagrionidae Ischnura posita (Hagen) Anornalagrion bastatum (Say) ORTHOPTERA: Acrididae Laplatacris dispar Rhen ORTHOPTERA: ACRIDIDAE Phaulacridium vittalum (Sjostedt) WilIis, 1971; NickIe, 1972 ORTHOPTERA: Acrididea Slenobothrus sp. COLEOPTERA : Chrysomelidae Leplirwtarsa decemlineata Say LEPlDOPTERA: Yponomeutidae Yporwmeula maltinella (L.) Yponomeuta padella (L.) COLEOPTERA : EIateridae ? sp. COLEOPTERA : ChrysomeIidae Gaslrophysa sp. Phyllotreta sp. DERMAPTERA : Forficulidae Foificula sp. LEPlDOPTERA : Lymantrüdae Lyman/ria dispar (L.) LEPlDOPTERA : Geometridae Operopht.era brumata (L.) LEPlDOPTERA : Pyralidae Chilo simplex Butler Unknown ORTHOPTERA : Acrididae Phaulacridium vittatum (Sjostedt) Chorloueles tenninifera (WaIker) COLEOPTERA : Scarabaeidae SeTÜeslhis spp. WeJch,1963 Miralles & Camino, 1983 Presesent study Hagmeier, 1912 Kaiser, 1972 Kirjanova el al., 1959 Shimkina, 1978 (as A. ghilarovf) Kaiser, 1991 Kaiser, 1991 Artyukhovski & Kharchenko, 1965 Kaburaki & Imamura, 1932 Spiridonov, 1987 Present study Fundam. appl. Nemalol. Amphimermis spp. Table 2. Morphometrics offemale of new Amphimermis species (dimensions of paratypes; mean and range). Species L (mm) A. acridiornm n. sp. n = 27 A. buraki n. sp. n=5 A. mirabinda n. sp. n=6 A. auslraJoelegans n. sp. n=3 Body width Head width )m~( Amphid lengthl Amphid width l ~ m~ m~ Vagina Nerve ring m~ position (010) dia. lengthl lengthz m~ m~ m~ Vest. anus 3 m~ 241 329 74-457 225-480 68.3 50-85 18.1 13-22 11.7 8-15 317 250-400 50.2 12.8-59.3 126 37-200 697 1142 320-1010 570-1675 30.1 23-B 164 130-187 51 45-54 13.2 10-14 7 249 230-266 56.6 54.6-58 48.7 40-62 210 110-282 324 245-437 161 51-BI 139 91-205 303 270-410 256 220-300 66.8 48-85 66.6 50-85 20.8 15-23 31.6 25-35 14.6 10-\7 15 318 3\2-325 327 272-360 50.1 45.6-53.6 53.4 48-58 110 62-150 73 60-100 628 380-850 780 720-870 1063 770-1500 1350 1200-1500 278 (5) 140-380 197 (2) 190-205 180 (1) lDorsal view. 2Lateral view. 3Disrance of vestigiaJ anus from rail: number in parenthesis. Table 3. Morphometric data fOl" male of new Amphimermis species (dimensions of para types; mean and range). Species L (mm) A. acridiornm n. sp. n = 51 A. !mraki n. sp. n=8 A. mirabinda n. sp. n=8 A. auslraJoelegans n. sp. n = 14 Body width Head width )~( )m~( Amphid lengthl Amphid width 1 )m~( )m~( Nerve ring )m~( L spicule )m~( L tail )m~( 84.3 28-187 188 112-262 64.5 52-87 18.8 12-30 10.8 7-17 2421 302 279 255-342 1775-3100 192-315 22.8 16-31 115 98-140 46.4 39-52 14.6 10-19 6.4 5-7 875 238 211-262 750-975 49.2 25-102 169 145-250 60.7 52-70 20.5 17-25 15.7 12-20 47.4 31-75 163 127-225 64.8 60-70 9 7-10 40.7 32-45 Proximal papilJa 3 Tai! width z )~( )~( Genital PapiUae ant post. N N 696 168 100-225 470-1175 324 100 87-117 250-375 28 20-41 12 7-16 14.5 12-18 7.5 4-9 273 291 1593 250-320 1199-2380 165-310 609 145 100-190 470-915 24 20-40 10 5-15 1675 290 260 255-3\3 1320-2060 205-315 138 753 115-160 575-890 31 24-37 99 9-12 144 130-165 'Dorsal view. 2Tail htd~, measured aI cloaca. 3Disrance of proximal papilla amerior ro cloaca. cloaca, 7 x taillength. Genital papillae arranged in three rows, medial row marginally longer than sub-medial rows; medial row bifurcate immediately anterior and posterior ta cloaca; distance of proximal genital papilla anterior to cloaca = 0.3 x spicule length and 2.4 x tail length. Smaller males (Fig. 4) tend to be aberrant in that the twisted section of the spicule contains a short untwisted portion at about two thirds of the length of the twisted section (Fig. 3 F). Juvenile, st. 2 (preparasitic): Short, slender larvae. Proximal half of body broader than distal half. Cephalic papillae distinct. Stylet barbed on one side; distal end with slight terminal swelling. Oesophagus terminates Vol. 17, n° 4 - 1994 mid-stichosome; sixteen stichocytes present in stichosorne. Trophosome interspaces indistinct and weil spaced (approx. 40-50 !-Lm). Tail attenuated ta a fine point with tip tending to hook shape in Life. A moult occurs within the host during the early stages (510 days) of parasitic development when larvae are 212 mm long (Fig. 5 B, C). A moult during parasitic development has been described for Romanomermis culici-vorax Ross & Smith (Poinar & Otieno 1974; VyasPatel, 1992), and Hominick et al. (1982) suggested the presence of a stylet in sorne parasitic juveniJes of Hexamermis glossinae Poinar, Mondet, Gouteux & Laveissier indicated a moult within the hast. The postparasitic juvenile undergoes a double moult. 307 G. L. Baker & G. O. Poinar Jr. B 50 J1m 100J1m - - CD 100J1m - - E 50J1m - - F AB .C® . \.~ ......... . C '. j .... F Fig. 1. Amphimennis acridiorum n. sp. Female. A: Head, dorsal view; B: Head, laleral view; C: Vagina, ·ventral view; D: Vagina, taleraI view; E: Tm/, laierai view; F: Ulerine egg. c Fig. 2. Amphimermis acridiorum n. sp. Aberrant Jenwles. A : Vagina with cloacal concrelion, ventral view; B . Same, talerai view (Bar: 100 fl.m); C: Tail with veHigial anus (arrow), taleraI view; D : Vesligial anus, laierai view (Bar equivalenr : A, B, C = 100 fl.m; D = 10 fl.m). TYPE HOST AND LOCATION Phaulacridium ·vittalUm (Sjëistedt) (Orthoptera : Acrididae). "Ambleside "[33° 43' S, 149 0 46' El, Oberon, Central Tablelands, New South Wales, Australia. TYPE MATERLAL Holotype (female) and allotype (male) in UCO. Paratypes (one male and one female) deposited in SAM and MNHR. OLAGNOSIS AND RELATIONSHIPS A. acridiorum n. sp. males belong to the avoluta group (Table 1) in having the proximal section of the spicule untwisted. Male A. acn'diorum n. sp. differs from A. avoluta in the position of the amphidial opening in relation to the opening of the lateral cephalic papillae (amphidial opening posterior to lateral cephaLic papillae vs at same level); 308 A. buraki n. sp. in being longer (28-187 vs 16-31 mm), in having a short spicule in relation to body length (2.4 and 87 mm (i.e. 1 : 36) vs 0.8 and 22 mm (i.e. 1 : 27) and in the position of the spicule twisting (distal 62-76 vs distal 51-58 %); relative position of the openings of the lateral and sub-medial head papillae (level vs submedial anterior to lateral) and having proporrionally smailer amphids (30 vs 35 % head width). Female A. acridiomm n. sp. can be distinguished from ail described species except A. bonaerensis and A. bogongae on the basis of the forrn of the vagina; loops anterior and posterior ta vulva of equallength (Fig. 16). A. acridiorum n. sp. differ from A. bonaerensis having a generally shoner vagina in relation ta body (3201010 IJ-m and 75-457 mm vs 400 IJ-m (measured from illustration in Miralles & Camino, 1983) and 100120 mm) and A. bongongae in having a generaily longer Fundam. appl. NernalOl. Amphimermis spp. 200 spicule normal aberrant .. 4 Nor1hern Sdnale~aT * "* HunIer Valley a Central Tablelancts o Soulhern Tablelan<ts &i ~ U ~ <.:1 o....J E E 1 1 l ' 1/ \ 1 1 SOj1m-- H SOj1m----- A 100 j1m - B G • l} l}e \ C D E F ...... 0 . o o· 50 " . .-. .0 i5<.:1 "1 1 1 .. .. ... . 0.-·...& ...&. 100 (fJ 1 . o Z r:.:I ....J o o>< o ~ 10 o 500 1000 1500 2000 2500 3000 3500 SPICULE LENGTH lpm) Fig. 4. Amphimermis acridiorum n. sp. Male. Scaller diagram of spicule length l'egressed against body length. vagina (in one plane) in relation to body length (3201010 fLm and 75-457 mm vs 380 fLm and 160-205 mm for A. bogongae). The uterine egg is of comparable size to A. bogongae but the st. 2 juvenile is only two thirds the length (1.4 mm vs 1. 9 mm). HOST RANGE Oedaleus aUSlrahs (Saussure) (Acrididae) : two males and one female reared From adults, Upper Rouchell, March 25, 1987, K. England. ChorLOiceles lenninifera (Walker) (Acrididae): three males and one female reared From nymphs. Laboratory culture, Rydalmere, NSW, M. Davison; three males reared From adults, " Westbrook ", Singleton, March 16, 1992, R. Pigott; one male reared From adult, " Longarm ", Barraba, February 26, 1992, R. Pigott. DISTRIBUTION Fig. 3. Amphimennis acridiorum n. sp. Male. A .' Head, dorsal view; B.' Head, laleral view; C.' Tml, laleral view; D.' Tail, ventral view; E.' Schema tic representation of genital papillae; F.' Tai!, laleral Vlew (aben'ant fomz); G.' Head, en face view; H.' Mid-body, cross section. Vol. 17, n° 4 - 1994 In addition to the type location, A. acn'diorum n. sp. has been reared From P. VÙlatum collected in the Northern Tablelands (Stonehenge, Hernani, Walcha), CentraI Tablelands (Oberon: " Cormark", " Ambleside ") and Southern Tablelands (Jerangle, Braidwood, Dalgety) and South West Slopes (Tumbarumba). A. acridiorum n. sp. has also been reared From C. lenninifera collected on the North West Slopes and Hunter Valley districts of New South Wales. A. acn'diorum n. sp. appears to be widely distributed throughout the tablelands 309 G L. Baker & G. 0. Po/nar Jr. Amphimermis buraki * n. sp. (Figs 6, 7) MEASUREMENTS Holotype (female) : L = 33 mm; mid-body diam. = 167 fLm; head diam. (at level of cephalic papillae) 53 fLm, (at neck) = 54 fLm; body diam. at nerve ring = 92 fLm; cuticie width (at nerve ring) = 7 fLm, (at midbody) 15 fLm; hypodermis width (mid-body) 6 fLm; amphid aperture = 2 fLm; amphid pouch (in dorsal view) = 13 x 8 fLm; dist. nerve ring/mouth = 230 fLm; V = 56.1; length of vagina (from vulva to junction with uterus) = 265 fLm, (in dorsal view) = 200 fLm; diam. vagina = 62 fLm; length vulval flap = 95 fLm; height of vulval cone 25 fLm; width of lateral hypodennal chord 20 fLm; diam. of uterine egg 75-80 fLm. A l!o type (male): L 16 mm; mid-body v"idth 89 fLm; head width (at level of cephalic papillae) = 39 fLm, (at neck) = 42 fLm; body width at nerve ring = 70 fLm; cuticle width, (at nerve ring) = 13 fLm, (midbody) 15 fLm; hypodermis width (mid-body) = 5 fLm; amphid aperture = 6 fLm; amphid pouch = 15 x 7 fLm (in dorsal view), 17 x 20 fLm (in lateral view); dist. nerve ring/mouth = 245 fLm; spicule length = 810 fLmj position of twisting distal 69 per cent of length; spicule head width = 12 fLm; width mid shaft = 5 fLm; tail length = 142 fLm; tail width at cloaca = 92 fLm; position of proximal genital papilla anterior to cloaca = 307 fLm; number of genital papillae 58. Paratypes (Females and males) : see Tables 2 and 3, respectively. = = = = = = = = = = = DESCRIPTION D Fig. 5. Amphimermis acridiorum n. sp. Parasitic juveniles. A : St. 2 (inJective), whole body; B: St. 2 (parasitic) r.ail with ecdysed GUticle; C: Lace stage St. 2 (parasitic) lail with ecdysed GUticie; D: St. 3 (parasitic). Al! Jrom lWSI Phaulacridium vinatum. (Bar equivalent: A, D: 100 fLm; B, C : 100 fLm). and western slopes of New South Wales in districts receiving both high summer rainfall and districts with marginal summer rainfall and an autumn maximum. 310 General: Short nematades, female 1.3 x length of male. Cuticle with cross fibres. Head bulbous. Cuticle anterior ta head protoplasm extremely broad. Mouth terminal. Four submedial cephalic papillae; t'wo lateral cephalic papillae. Opening of sub-medial cephalic papillae anterior to opening of lateral cephalic papilla, more pronounced in male than female. Large cup shaped amphids, larger in male than female; opening of amphids posterior ta opening of lateral cephalic papillae and slightly offset. Six hypodermal chords. Wall of anterior end of oesophagus thickened to produce a short pharynx. Females : Opening of vulva a transverse slit. Vulval flap present, rim thickened. Vulval cone present; base muscular and fused with terminal segment of vagina. Vagina S shaped, short, muscular; posterior loop x 1.5 length of anterior loop; bends extremely sharp. Junction of vagina and uterus weil posrerior to vulva. Tail conical, slightly curved ventrally. Tail bluntly rounded. No vestigial anus. Eggs in double row in uterus. Eggs unembryonated when laid. burakl~ aboriginal (Yarrowitch dialect) = hidden valley, alluding ta limited distribution. * Fundam. appl. NemalOl. Amphimermis spp. A 50 pm - - - - - - A B 50 pm 100pm-- C 0 E AB 50pm - - E F 100pm - - C 0 A B o • • 0 • 0 0 • 0 o 0 o • ; c • • 0 0 0 • 0 • o 0 00 0 (>00 0 go& ~-o 0 :3% : .. 000 o8g g :sg" o F c Fig. 6. Amphimermis buraki n. sp. Female. A : Head, dorsal view; B: Head, lateral view; C: Vagina, ventral view; D: Vagina, laIerai view; E: Tail, laIerai view; F: Uterine egg. Males: Tail tightly curled into ring, conoid, bluntly round. SpicLÙes paired; loosely twisted for distal two thirds of length (70 %); fused at tip, anenuated to fine point; head gently f1ared with pincer like terminus; wall thick proximal third, thin distal two thirds (twisted section); spicLÙe length x 9 body width at cloaca, x 6 tail length. Genital papillae arranged in three rows, medial row marginally longer than submedial rows; distance of proximal genital papillae from cloaca x OAlength of spicLÙe and x 2.2 tail length. Head more bLÙbous than female. Wall of am phid thin relative to that in female. Ducts of submedial cephalic papillae at acute angle to long axis ofbody, ducts oflateral cephalic papillae transverse. TYPE HOST AND LOCATION Conocephalus sp. (Orthoptera: Tenigoniidae). Kangaroo Flat, [31 0 Il' S, 152 0 07' E], Yarrowitch, Northern Tablelands, New South Wales, Australia. Vol. 17,n o 4-1994 .00 E Fig. 7. Amphimermis buraki n. sp. Male. A: Head, dorsal view; B: Head, lateral view; C: Tail, laIerai view; D: Tail, ventral view; E: Schema tic representalion of genital papillae. TYPE MATERIAL Holotype (female), allolype (male) in UCD. Paralypes (one female and one male) deposited in SAM and MNHN. DIAGNOSIS AND RELATIONSHIPS Male A. buraki n. sp. is distinguished from ail previously described species of Amphimemlis except A. avoluta and A. acridiorum n. sp. on the basis of the twisting configuration of the paired spicules, being twisted for the distal two thirds only, the proximal third being straight. A. buraki n. sp. differs from A. avoluta in that the body length is shorter (16-31 mm vs 57 mm); the spicule length is shorter (750-975 /-Lm vs 2000 /-Lm); the distal twisted portion of the spicules is a greater proportion of the total spicule length (70 vs 45 % ); the spicule head is pincer-shaped rather than f1ared; the head is bulbous rather than bluntly rounded; the opening of the amphids is a substantial distance posterior to the lateral cephalic papillae rather than adjacent and A. acridiorum n. sp. in being shorter (16-31 mm vs 28-187 mm), the spicule length is shorter (750-975 /-Lm vs 17753100 /-Lm); the head is bulbous rather than bluntly rounded. 311 G. L. BakeT & G O. PoinaT JI' The female of A. buraki n. sp. is distinguished from ail previously described species except A. elongaw) A. maritima and A. mongolica on the lack of vestigial anus (excluding intersexuals) and form of the vagina : loop of vagina posterior to \rulva greater length than anterior loop and ju nction of vagina and uterus posterior ta vulva greater length than anterior loop and junction of vagina and uterus posterior to level of vulva. A. buraki n. sp. differs from A. marilima and A. elongala in having a relatively long vagina (greater than body width) and A. mongolica in having thick-walled amphids and the opening of the lateral cephalic papillae in close proximity to the opening of the submedial cephalic papillae. FemaJeA. buraki n. sp. differ from other species in the avolula group in the following respects: A. avolUla in that body length is shorter (31-36 mm vs 125 mm); the head is bulbous rather than blundy rounded; the opening of the amphids is well posterior to the lateral cephalic papillae rather than adjacent; the vagina is shorter (265350 fLm vs 950 fLm); the junction of the vagina with the uterus is posterior ta the vulva rather than anterior; uterine eggs generally smaller (75 x 80 fLm vs 65 x 100 fLm): A. acndiontm n. sp. in being shorter (33 mm vs 241 mm); the form of the head (bulbous vs blundy rounded); the length of the vagina (0.26 mm vs 1.2 mm); and differing in the relative length of the anterior and posterior loops (approximately equaJ vs anterior loop 1.3 x posterior loop). The openings of the submedial and lateral cephalic papillae are on the same level in the females of both A. buraki and A. avolula yet in the male of both species the opening of the lateral cephalic papillae is weil posterior to the submedial cephalic papillae. This sexual dimorphi sm in regard to arrangement of cephalic papillae is not shared by other species of Amphimermis. Arnphirnermis rnirabinda '" n. sp. (Figs 8, 9) MEASUREMENTS Hololype (female) : L = 113 mm; mid-body diam. = 270 fLm; head diam. (at level of cephalic papillae) = 65 fLm, (at neck) = 78 fLm; body diam. at nerve ring = 120 fLm; cuticJe width (at nerve ring) 10 fLm, (at rnidbody) = 27 fLm; hypodermis width (mid-body) = 12 fLm; amphid aperture = 7 fLm; amphid pouch 23 x 14 (in dorsal view); dist. nerve ring/mouth = 312 fLm; V = 49.5; length vagina (from \rulva ta junction with uterus) = 975 fLm, (in dorsal view) = 540 fLm; length vulval flap = 75 fLm; height vulval cone = 45 fLm; width lateral hypodermal chord = 25 fLm; diam. egg in uterus = 77-82 x 100-102 fLm. Allolype (male): L = 36 mm; mid-body diam. = 117 fLm; head diam. (at level of cephalic papillae) = = * mimbinda, aboriginal (Nembo dialect) = place with running warer, alluding ra habitat. 312 A 50J,Jm 100J,Jm 100J,Jm SOJ,Jm A 8 - CD E -- F c F Fig. 8. Arnphirnermis rnirabinda n. sp. Female. A : Head, dOTsaI view; B: Head, lateml view; C: Vagina, ventral view; D: Vagina, laleml view; E: Tail, laieraI view; F: Ulerine egg. 52 fLm, (at neck) = 55 fLm; body diam. at nerve ring = 87 fLm; cuticle width (at nerve ring = 15 fLm), (midbody) = 25 fLm; hypodermis width (mid-body) = 14 fLm; amphid aperture 8 fLm; amphid pouch = 20 x 15 fLm (in dorsal view), 21 x 21 fLm (io lateral view); dist. nerve ring/mouth = 255fLm; spicule length = 1199 fLm; position of twisting from tip = 10.4 to 54.2 and 80.2 ta 97.1 % spicule Iength; spicule head width = 14 fLm; spicule width mid-shaft = 5fLm; tail Iength = 165 fLm; tail diam. at cloaca = 100 fLm; position of proximal genital papilla anterior ta cloaca = 587 fLm; number of genital papillae = 62. Paralypes (Females and males) : see Table 2 and 3, respectively. Juvenile) Si. 2 (pre-pamsùic) (n = 5) : L = 1.44 mm (1.40-1.53); mid body diam. 15 (nil range); head diam. : 12 fLm (nil range); buccal cavity length = 20 fLm (nil range); stylet = 26 fLm (25-28); dist. nerve ring/ mou th = 108 fLm (100-115); body diam. at nerve ring = = = Fundam. appl. NemalOl. Arnphimermis spp. B ::~ .:. :: : ·:.11 " r. ~ .. 50pm 50pm 100 pm DE AB C o o o o 00 o 0 o 1'-+--' D E Females.- Opening of vulva a transverse slit. VuIval flap large, rim thickened to appear bulbous in lateral view. Vagina S-shaped, anterior and posterior loops of equal prominence. Junction of vagina and uterus posterior to vulva. Distal uterus with transversely looped segment immediately posterior ta vagina. Tai! conoid, slightly curved ventrally. Uterine eggs in double row. Eggs unembryonated when laid. Males.- Tail-shaped (a feature shared with A. bogongae), conoid, terminus bulbous. Spicules long, fine, paired; loosely twisted for proximal fifth of length and distal half of length; tips fused, anenuated to a fine point; head gently flared with pincer like terminus; spicule length 12 x body width at cloaca, 7-9 x tail length. Genital papillae arranged in three rows, medial row marginalJy larger than submedial rows; distance of proximal genital papillae from cloaca 0.3-0.48 x length of spicule and 2.6-3.5 x taillength. Juvenile, sl. 2 (preparasiLic) .- Short, siender. Proximal half of body broader than distal half. Cephalic papillae distinct. Stylet barbed on one side. Trophosome interspaces indistinct. Tail anenuated to a fine point. Similar to A. acridiorum n. sp. preparasitic juvenile in many respects differing in the slightly greater length of the stylet, a more pronounced thickening of the tip of the stylet and elongated stichocytes. TYPE HOST AND LOCATION c Phaulacn'dium viualum (Sj6stedt) (Orthoptera : Acrididae). Jingera [35 0 45' S, 1490 26' El, Southern Tablelands, New South Wales, Australia. TYPE MATERIAL Fig. 9. Arnphimermis mirabinda n. sp. Male. A : Head, dorsal view; B.' Head, La le rai view; C: Tail, laIerai view; D.' Tail, ventral view; E: Schematic represenration of genital papillae. Hololype (female) and alfolype (male) in UCD. Paralypes (one female and one male) deposited in SAM and MNHN. DIAGNOSIS AND RELATIONSHIPS 15 (ni! range); position of stichosome : proximal end = 13.7 % (13.3-14.8), distal end = 62 % (60-63) body length; body diam. mid-stichosome = 17 fLm (nil range); mid-trophosome = 15 fLm (nïl range); length of tail = 56.6 fLm (44-70); tail diam. (50 fLm from tip) = 6 fLm (ni! range). DESCRlPTION General.- Medium length nematades, females 1.23.1 x length of male. Cuticle with cross fibres. Mouth terminal. Four submedial cephalic papillae; two lateral cephalic papillae. Opening of submedial cephalic papillae posterior ta opening of lateral cephalic papillae in both males and females. Enormous amphids, larger in male than female; opening of amphids posterior to opening of lateral cephalic papillae. Six hypodermal chords. Wall of anterior end of oesophagus thickened to produce a short pharynx. Vol. 17, n° 4 - 1994 Male A. mirabinda n. sp. differs from ail species in the bogongae group (A. bogongae, A. Linyi, A. mariLima, A. liLoralis, A. bonaerensis) by having enormous amphids (width approximately half head diameter). The body length of A. mirabinda is comparable ta that of A. mariLima and A. liLoralis. However, the spicule is considerably shorter (1199-2380 vs 2200 and 3400 fLm, respectively) and position of proximal papilla as a proportion of spicule length (30-48 vs 19 %). A. mirabinda n. sp. also differs from A. bogongae in having larger amphids and a more bulbous tail. Female A. mirabinda n. sp. can be distinguished from ail other species except A. bonaerensis and A. bogongae by the form of the vagina (anterior and posterior loops of equal prominence and junction of vagina and uterus posterior ta level of vulva). A. mirabinda n. sp. differs from A. bonaerensis in lacking a vulval /lange, though this may not be a good diagnostic character (see discussion), and from A. bogongae in length (51-113 vs 160313 G. L. Baker & G. 0. Poinar JI'. 205 mm), egg diameter (102 vs 140 f..lm) and width of lateral hypodermai chord (25 vs 100 f..lm). D1STRlBlJTION Apart from the type location, A. mirabinda n. sp. has been reared from P. villalum collected in the Central Tablelands (Oberon) and has been collected ex soil in the Hunter Valley (Gundy, April 1989, R. Pigott). ~ Arnphinwrrnis australoelegans n. sp. (Figs 10-11) l MEASUREMENTS Hololype (female) : L = 123 mm; mid-body diam. = 250 f..lmi head diam. (at level of cephalic papillae) = 85 f..lm, (at neck) = 83 f..lm; body diam. at nerve ring = 117 f..lm; cuticle width, (at nerve ring) = 10 f..lm, (at mid-body) = 12 f..lm; hypodermis width, mid-body = 25 f..lm; amphid aperture = 8 f..lm; amphid pouch: (in lateral view) = 35 x 5 f..lm, (in dorsal view) = 35 x 25 f..lm; dist. nerve ring/mouth = 350 f..lm; V = 48.8 length vagina (from vulva to junction with uterus) = 1500 J.Lm, (in dorsal view) = 870 J.Lmi length vulval t1ap = 105 J.Lm; height vulval cone = 50 f..lm; width lateral hypodermal chord = 37 \Lm; diam. uterine egg = 65-70 f..lm. Allolype (male): L = 62 mm; mid-body diam. = 165 J.Lm; head diam. (level of cephalic papillae) = 63 J.Lm, (at neck) = 65 f..lm; body diam. at nerve ring = 100 J.Lm; cuticle width (at nerve ring) = 18 f..lm, (midbody) = 23 f..lm; hypodermis width (mid-body) = 8 J.Lm; amphid aperture = ill defined; amphid pouch = 43 x 12 J.Lm (in dorsal view), 43 x 32 J.Lm (in lateral view) i dist. nerve ring/mouth = 295 J.Lm; spicule length= 1711 J.LmiPositionoftwisting= 15.3-59.1 and 69.4-94.9 % of length; spicule head width = 23 J.Lm; width mid-shaft = 9 f..lm; tail length = 260 J.Lm; tail at cloaca = 155 J.Lm; position of proximal genital papilla anterior to cloaca = 887 f..lm; number of genital papillae in medial row (anterior to cloaca) = 37, (posterior to cloaca) = 10. Paralypes (Females and males) : see Tables 2 and 3, respectively. DESCRIPTION General: Medium sized nematodes, female 2 x Iength of males. Cuticle with cross fibres. Head blunùy rounded. Mouth terminal. Head with four submedial cephalic papillae and two lateral cephalic papillae. Opening of submedial cephalic papillae posterior ta opening of lateral cephalic papillae. Amphids large, thin walled, irregular shaped cuticular incursion into head protoplasm. Opening of amphids indistinct. Six hypodermal chords. Fernales: Opening of vulva a transverse slit. Vulval t1ap thickened ta form a semicircular rim anterior to vulva and extending posteriorly around edge of vulva. Post laying, cytoplasmic extrusions may modify the 314 1 50 pm 50pm 100pm - j,l' A 8 F C D D Fig. 10. Amphimermis ausrraloelegans n. sp. Female. A: Head, dorsal view; B: Head, lateral view; C: Vagina, ventral view; D: Vagirza, laLeral view; E: Tail, laleral view; F: Utenne egg. form of the ring to produce a t1ange. Vagina S-shaped and relatively long in relation to body width; posterior loop 4 x length of anterior loop; walls uniformly thin. Junction of vagina and uterus slightly posterior to the position of the \ruJva. Tail tapered to a point. No vestigial anus. Eggs in up to four rows in uterus. Eggs relatively small, unembryonated when laid. Males: Tail curled into a ring, t1at ventrally, pointed. Spicules paired, tighùy twisted except for one sixth of length from tip and a small section at one third of length from proxima! end; head t1ared, ventral t1are shorter than dorsal t1are; walls thick uniformly along length; spicule length approximately Il x body width at cloaca, 6.5 x taillength, 27 % body length. Genital papillae arranged in three rows median row marginally longer than sub-medial rows; medial row bifurcate anterior and Fundam. appl. NemalOl. Amphimermis spp. SOJ.im and A. artyukhovski n. sp. A. auslraloelegans n. sp. differs from A. elegans in regard the form of the tail (conoid with pointed terminus in A. auslraloelegans vs bluntly rounded in A. elegans), having greater number of anal papillae anterior to cloaca (24-37 vs 13), in form of spicule twisting (proportionally short straight section mid-spicule vs long straight section mid-spicule) and spicule tip (attenuated to a fme point vs relatively blunt); A. zuimushi in having a relatively short body length in relation to spicule length (x 23-30 vs 40-60) and with regard to the position of the proximal anal papilla (approximately 30 % of spicule length in A. auslraloelegans n. sp. vs approximately 60 % in A. zuimushl); A. tongaensù in greater body length (31-64 mm vs 20-32 mm) and form of the amphids (sensilla core centrally placed vs on outer rim) and size of the amphids in lateral view (one third diameter of head vs two thirds diameter of head) and A. artyukhovskii n. sp. in the shape of the tail (pointed vs bluntly rounded). The female of A. australoelegans n. sp., like the male, has characteristic amphids which separate it from al! species except A. elegans, A. zuimushi and A. artyukhovskii n. sp. The female of A. tongaensis is unknown. A. australoelegans n. sp. differs from A. elegans with regard to range of body length (91-205 mm in A. australoelegans n. sp. vs 145-260 mm in A. elegans (Hagmeier, 1912; Kirjanova et al. 1959); A. zuimushi with regard to vagina length in relation to body width (780 and 256 f.Lm respectively in A. australoelegans n. sp. vs 800 and 350 f.Lm respectively in A. zuimushl) and thickness of wall of anterior loop of vagina (thin vs thick) and A. artyukhovskii n. sp. in relative thickness of vagina wall (thin vs thick). A BFG l00J.im- C DE F Fig. 11. Amphimermis australoelegans n. sp. Male. A " Head, dorsal view; B.' Head, laierai view; C.' Tail, laierai view; D,' Tail, ventral view; E. Schematic presentation of genùal papillae; F,' Head, en face; G.' Mid-body, cross section. HOST RANGE posterior to cloaca; distance of proximal genital papiUa anterior to cloaca = 0.5 x spicule length and 3.4 x tail length. In addition to the type host, A. auslraloelegans n. sp. has been recorded from the fol!owing hosts : ChorlOiceles tenninifera (Walker) (Orthoptera : Acrididae) : one male and one female reared from adult host, Weldon Lane, Moree, May 10, 1988, R. Pigon, and Sericeslhù sp. (Coleoptera: Scarabaeidae) : Melolonthinae) : one female reared from late instar host larva, Hernani, March 14, 1985, A. J. Campbell. TYPE HOST AND LOCATION DISTRlBUTION Phaulacridium 'villalUm (Sjôsredt) (Grthoptera : Acrididae). Kangaroo Flat [0 Il S, 007' El, Yarrowitch, Northern Tablelands, New South Wales, Australia. f TYPE MATERlAL Holotype (female) and alfotype (male) and 3 paratype males in UCD. Paratypes (one female and two males) deposited in SAM and MNHN. DIAGNOSIS AND RELATIONSHIPS Male A. australoelegans n. sp. has characteristic amphids and twisting of the spicule which separates it from ail species except A. elegans, A. zuimushi, A. tongaensis Vol. 17, n° 4 - 1994 Apart from the type locality A. australoelegans n. sp. has been recorded from Hernani (ex Sericesthis sp.) and Moree (ex C. tenninifera). The species would appear ta be restricted to relatively warm districts with high summer rainfall. A»lphitnerrnis artyukhovskii n. sp. =A. elegans apud Artyukhovski & Kharchenko, 1965. A. elegans, sensu Artyukhovski & Kharchenko (1965), is considered a distinct species on the basis of the high number of genital papillae anterior to cloaca, and the more rounded (bulbous) tail. The hosts also differ sig315 G. L. Baker & G. O. Poinar Jr. nificantly (Lepidoptera vs Orthoptera). The material is assigned the new name arlyukhovskii n. sp. The male of A. auslraloelegans n. sp. differs from A. arlyukhovskii n. sp. in having fewer genital papillae anterior ta cloaca, and a tail with a more pointed terminus. The female of A. auslraloelegans n. sp. differs from A. arlyukhovskii n. sp. in the form of the vagina (long and thin walled 'vs relatively short and thick walled). The male of A. arlyukhovskii n. sp. most closely resembles A. zuùnushi differing from this species only in the relative length of the spicule in relation ta body length. The male A. elegans illustrated in Kiryanova el al. (1959) has a rounded tail similar ta that illustrated for A. elegans in Artyukhovski and Kharchenko (1965), which is much more rounded than that illustrated by Hagmeier (1912) for A. elegans. However, it is similar ta A. elegans in respect ta number of genital papillae and position of proximal genital papillae, differing from the material of Artyukhovski and Kharchenko (1965) and A. auslraloelegans n. sp. in bath these respects. The differences are not considered sufficient ta erect a new species as has been done for the material of Artyukhovski and Kharchenko (1965). o Key to males of the genus Arnphirnerrnis (Figs 12-15) 1. - Spicule twisted entire length (= volubilis group) ......................................................................... volubilis 2 - SpicLÙe twisted for only part of length ... 2. - Proximal half of spicLÙe unrwisted (= auolma group) 3 - Spicule twisted distal and proximal ends with srraight section in middle 5 3. - Body short « 30 mm), spicule length 750-975 f1.m, terresrrial . tlUraki n. sp. 4. - Amphid opening at level of lateral cephalic pa pilla avolula - Amphid opening posterior to lateral cephalic pa pilla acridiorum n. sp. 5. - Amphid indistinct, amphidial pore (opening) small: a thin walled pocket in cuticle (= elegans group) ......... 6 - Amphid distinct, amphidia] pore medium-large: a thick waIJed, cup-shaped incursion into head cytoplasm (= bogongaegroup) 10 6. - Amphid diameter greater than two thirds head width LOngaensis - Amphid diameter Jess than half head width 7 7. - Distance proximal anal papi!lae to cloaca> 60 % spicLÙe length.......... . 8 - Distance proximal anal papi!lae ta cloaca < 60 % spicLÙe length 9 8. - Spicule 1020-] 450 f1.m. Body 42-88 mm ZUlmushi -Spicule 1600-1800 f1.m. Body 26-53 mm. Tail conoid, rounded artyukhovskii n. sp. 9. - Tai! bluntly rounded - Tai! conoid, poinred terminus.... elegans australoelegans n. sp. 10. - Body short (11-17 mm), spicule 700-860 f1.m, aquatic .. 316 tinyi Fig. 12. Amphimerrnis spp. Males; Head, dorsal Vlew showing form of amphid and position of amphidial pore (arrow). A : A. acridiorum n. sp.; B: A. buraki n. sp.; c: A. mirabinda n. sp.; D : A. ausrraloelegans n. sp. (Bar = 10 f1.m). 11. 12. 13. - 14. - Body medium-long (30-123 mm), terresrrial .... . Il Body short (30-70 mm) ." .... 12 13 Body long (70-123 mm) Spicule long (2900-3600 f1.m) in relation to body length lita ralis (45-70 mm) Spicule medium (1200-2200 f1.m) 14 Tail bluntly rounded, distance of proximal papillae from cloaca = 20 % of spicLÙe length bogongae Tai! pointed, distance proximal papillae to cloaca"" spicule length bonaerensis SpicLÙe 1200-1500 f1.m. Body 35-42 mm mirabinda sp. n. - Spicule 2,200 f1.m. Body 53 mm ..... maritima Key to females of the genus Arnphirnerrnis (Figs 1 B; 10 B, C, D; 16) 1. - Amphids indistinct, thin-walJed pear shaped pocket in cuticle 2 Fundam. appl. Nernatol. Arnphimermis spp. A:avoluta group 200 A.lwo/u/a - a- 100 "'- buraki n.a:p. A. aCrJrJlOtVffl n.sp. 50 ~ H 2J w. Ô :3 E ~ -A :r: E-< / 10 500 0 1000 1500 2000 3000 3500 4000 2500 3000 3500 4000 2500 3000 3500 4000 2500 B: bogongae group 200 A. 100 -loi. ::::- / ~ ~ A manI/ma -e- 50 "ny' A bogongao :Ill A mlmb,nrja n sp 0 Ô Z W H :>< Cl 0 ~ I~ /~ 10 0 1000 500 1500 2000 C : elegans group 200 Ou"'''''.'o.ons n.sp. _ A. 4- A zu;mush, J", 1 • 100 • 50 == : ::::::./, n.sp. 1" /i/ D Fig. 13. Amphimermis spp. Males; Proximal lips of spicules (large arrow) in relalion 10 position of proximal genital papillae (small arrow). A : A. acridiorum n. sp.; B: A. buraki n. sp.; C: A. mirabinda n. sp.; D: A. ausrraloelegans n. sp. (Bar = 100 Il.)' - Amphids distinct, thick-waUed cup shaped incursion inta head protaplasm 4 2. - Vagina long (1.5 mm), thin waUed ................................................ auslraloeœgans n. sp. - Vagina shon-medium length (0.5-0.9 mm), thick waUed 3 3. - Vagina medium length (0.9 mm) zuimushi - Vagina shon (0.5 mm) 15 4. - Anterior loop of vagina of equal or greater length than posterior loop . 5 - Anterior loop of vagina shorter than posterior loop 8 5. - Anterior loop greater length than posterior loop 6 Vol. 17, n° 4 - 1994 10 500 1000 1500 2000 SPICULE LENGTH (pm) Fig. 14. The relationship belween body lenglh and spicule lenglh for male Arnphimermis sp-p. A : avoluta group; B: bogongae group; C: elegans group. - Anterior loop equal to posterior loop 7 6. - Vagina medium (0.5 mm) avoluta - Vagina long (0.7 mm) volubilis 16 7. - Vagina shon and broad (in lateral view) - Vagina long and narrow (in lateral view) acridioruTll n. sp. 8. - Posterior loop of vagin a less than or equal to 3 fold length of anterior loop 9 - Posterior loop of vagina greater than 3 fold length of anterior loop 14 9. - ]unction of vagina and uterus at level of vulva 10 - ]unction of vagina and uterus posterior ta level of vulva .. 11 317 G. L. Baker & G. O. Poinar Jr. 10. - Body length 30 mm, aquatic - Body length 110 mm, terresrrial .. II. - Vagina length equal to mid-body width . - Vagina length greater than mid-body width .... tinyi li/oralis 12 13 12. - Junction of vagina and uterus at an acute antero-venrral angle ...... elongata - Junction of vagina and uterus at an obruse antero-venrral angle maritima 13. - Amphids thin-walled, siruated in neck region with opening weil posrerior tO lateral head papillae . mongolica - Amphids thick-walied, siruated anterior to neck region with opening immediately posterior to lateral head papiJlae buraki n. sp. 14. - Amphids siruated anterior to neck region with opening immediately posterior to lateral head papillae. Body 43 mm longiscapus - Amphids siruated in neck region with opening well posterior to lateral head papillae. Body 74 mm... lagidzae 15. - Body length 36-190 mm - Body length 195-260 mm 16. - Vulva with flanges - Vulva without flanges bonaerensis 17 17. - Medium sized amphids in relation to head diameter, thick-walied bogongae - Large amphids in relation to head diameter, thin-walled .. miratrinda n. sp. B Fig. 15. Variations in the fonn of the spicule in Amphimermis males. A: A. volubilis (after Rubtsov and Koval, 1975); B: A. avoluta (after Rubtsov and Koval, 1975); C: A. elegans (after Hagmeier, 1912) (Bars = 100 f.Lm). A B c D Fig. 16. Variations in the fonn of the vagina in Amphimermis females. A: A. avoluta (after Rubtsov and Koval, 1975); B: A. bogongae (after Welch, 1963); C: A. mongolica (after Rubtsov, 1976 b); D: A. longiscapus after Rubtsov, 1976 a) (Bars = 100 f.Lm). 318 artyukhovskii n. sp. elegans Discussion Stage and sex of the host may have a substantial impact on subsequent adult length (Herron & Baker, 1991) even within a single host species. Similarly, the physical environment may have an indirect impact on development (Baker & Holmes, 1986). Such plasticity impacts on many aspects of morphology and reduces the availability of stable diagnostic characters. Given the intraspecific variation which may occur, the species described in this paper are further discussed. The body and spicule length of A. buraki n. sp. is at the low extremity of a continuum which includes all species in the avoluta group (Fig. 14). As such this species could be regarded sim ply as small specimens of A. acridiontm n. sp., the small size induced by development in a small alternative host. A. buraki n. sp. was only recorded in the 1984-85 season which received above average rainfall resulting in rank pastures which favoured grass-seed feeding tettigoniids such as Conocephalus sp. but was coincident with very low densities of acridids following the collapse of the 1979-1982 outbreak of P. vittaLUm (Baker, 1992). The coincidence of a temporal change in abundance with an inversion in the relative abundance of two potential alternative hosts implies a host induced change in size of a single species. However, A. hl/raki n. sp. has only been recorded from the Kangaroo Flat area of the Northern Tablelands. If sim ply a host induced aberrant sized A. acridiorwn n. sp. then both forms would be expected to have a similar Fundam. appl. NemalOl. Amphimermis spp. distribution. However, des pite equally exhaustive sampling at other sites within the distribution of A. aen'diorum n. sp. the smaller A. buraki n. sp. has not been recorded. Differences possessed by A. buraki n. sp. which are not readily attributable to host size are the much steeper slope of the relationship between spicule length and body length and morphological characters such as head form and relative position of head papillae openings. A. acridiorum n. sp. is closely related to A. buraki n. sp. and represents the upper timit of a continuum in regard many morphological features. However, the configuration of the spicule in the aberrant form of A. aendiorum n. sp. tends towards that of A. bogongae and the aberrant form of A. acridiontm n. sp. may represent a sibling species which is the phylogenetic precursor of both A. bogongae and A. bonaerenis as weil as being the phylogenetic link between the avoluta and bogongae groups of Amphimermis. Three specimens designated as A. acridiorum n. sp. in Fig. 4, wruch were collected from soil in the Hunter Valley, have not been included in the range of dimensions given in Table 3. Their extremely short spicules sets them apart from other specimens of A. acridiorum n. sp. and they may represent a new species. In many respects, the difference between A. mirabinda n. sp. and A. bogongae) both males and females, is one of size only, A. bogongae being the larger species. However, A. mirabinda n. sp. is considered as a distinct species as the width of the lateral hypodermal chord is disproportionately broad in A. bogongae. Also, the head protoplasm in A. bogongae is more extensive and the head papillae much less pronounced. The size difference between A. mirabinda n. sp. and A. bogongae could be attributed to development of A. bogongae in a smaller host. However, mitigating against their being conspecific in the fact that differences in host size, although affecting female length is rarely a limiting factor in determining the length of males. A. mirabinda n. sp. is restricted to the Southern Tablelands and Hunter Valley, both districts which have relatively low summer rainfall. In A. mirabinda n. sp. the relationship between the body length and spicule length is at the lower end of the range for A. bogongae and overlaps to sorne extent. However, the species are separated by habitat, A. bogongae being found in hibernating moths occuring only in rock crevices and caves at high altitudes. In May 1990 the acridid Kosiuscola cognatus Rehn was collected at the type locality of A. bogongae (Mt Gingara) ACT) Australia) and found ta be parasitised by a species of Amphimermis which unfortunate]y was not reared to the adult stage. Comparison of the DNA of the parasitic juveniles with that of A. bogongae and A. mirabinda n. sp. adults should confirm if A. bogongae has acridids as alternative hosts of if A. mirabinda n. sp. has a wide geographic range or a further unVol. 17, n° 4 - 1994 described species of Amphimermis occurs at high altitude and has an acridid host G. Curran pers. comm.). The genus Amphimermis) as with the majority of mermithid genera, is ubiquitaus being represented on ail continents except Africa and Antarctica. The apparent discontinuity in the distribution of the closely related A. elegans and A. australoelegans n. sp. with dissimilar species in the intervening geographic regions (A. artyukhovskii n. sp. in Central Asia, A. zuimushi in South-East Asia and A. LOngaensis in Tonga) may indicate that A. auslraloelegans n. sp. represents convergent host adaptation by a formerly widespread elegans group progenitor in the Asian region. No species belonging to the elegans group have been recorded from North and South America or Africa and it is likely the group originated in Central Asia and spread to Australia after the fusion of the Australian and Indonesian tectonic plates in the mid Miocene period. As A. auslraloelegans n. sp. is distributed in temperature regions with a summer rainfall maxima, colonisation of Australia by the species progenitor was most likely from the north and may have coincided with invasion of large tropical Acridinae (i.e. Locusta) Gastrimargus) and Crytacanthacridinae (i.e. Nomadacris) wruch taok place in recent (Pleistocene) times (Key, 1959). Of the species groups erected in this paper) endemicity is greatest in the primitive avoluta group. The avoluta group is considered the most primitive group because of the relatively simple arrangement of the spicule twisting. The coincidence of both primitiveness and a high level of speciation in this group in Australia could indicate an Australian origin. The bogongae group is onJy slighùy more evolved than the avoluta group but again shows a high level of speciation and specialisation of host and habitat in Australia. Two species in this group) A. bonaerensis and A. mirafyinda n. sp. are parasitoids of Orthopterans in South America and Australia respectively. This could again indicate origin in Australia with a spread from Australia during the Upper Cretaceous period as far as Eurasia with adaptation to Orthoptera taking place prior ta this spread with more recent adaptation ta alternative hosts in Eurasia. The speciation of both the avoluta and bogongae groups in Australia may have been in response to conditions favouring both host and nematade during the Tertiary period. There was a proliferation of grasslands during the Eocene and Oligocene epochs of the Tertiary period and it was during this period that the acridid fauna proliferated, especially the subfamily Catantopinae (Key, 1959). The Catantopinae today are represented by species restricted to moist, alpine habitats (Key, 1986) coincident with the distribution of Amphlmermis spp. The increasing aridity during the Pliocene, Pleistocene and Holocene epochs would be expected to have restricted the distribution of both the acridid and mermithid fauna to refugia. The rugh level of speciation in 319 G. L. Baker & G. 0. Poinar Jr. the avoluta group in Australia with orthopterans as host was most probably the result of disparate evolution in the few isolated grasslands present in the mois t, upland areas of south eastern Australia. The currenùy evident sympatry between endemic Amphimermis spp. may have been achieved post-European settlement, following the creation of contiguous grasslands through the extensive clearing of forest for grazing. Under this scenario, competition between formerly allopatric species may ultimately reduce species richness. The implicit associated loss of biological diversity could reduce the capability of residual Amphimermis spp. to adapt to changing climatic conditions and consequenùy impair their ability to suppress acridid host populations. Acknowledgrnents The mermithid nematodes on which the descriptions are based were collected during a study of the population dynamics of the acridid P. villalUm supponed by an Ausrralian Meat Research Commirtee grant (DAN 19 S). The authors are grateful to Messrs. R Pigort and A.]. Campbell and Ms. H. M. Holmes, NSW Agriculture, for assistance with field sampling, Mr. A. Westcort, NSW Agriculture, for translation of sorne Russian literarure and Mr. L. Turton, NSW Agriculture, for photographic assistance. A rravel grant to fmalise the taxonomie aspects of the study was provided by the Tablelands Wingless Grasshopper Cornmirtee. References ARTYUKHOVSKl, A. K. (1969). [New genera of the family Mermithidae (Enoplida, Nematoda).] Zool. Zh., 48 : 13091319. ARTYUKHOVSKl, A. K. & KHARCHENKO, N. A. (1965). [On knowledge of mermithids of the Srreletskaya steppe]. Tntdy Tsentral'nogo-Chernozemnogo gos. Zapov., 9 : 159-185. ARTYUKHOVSKl, A. K. & KHARCHENKO, N. A. (1971). [New mermithids (Mermithidae, Nematodea) from the Cenrral Chernozem zone of the USSR] Trudy Tsentral'nogo-Chernozemnogo gos. Zapov., 11 : 109-131. BAKER, G. L. (1986). The ecology of mennithid nematode parasites of grasshoppers and locusts in south-eastern Ausrralia. ln: Samson, R. A., Vlak, J. M. & Peters, D. (Eds). Fundamental and applied aspects of invertebraLe pathology. Wageningen, the Netherlands, Foundation 4th ICTP : 277280 BAKER, G. L. (1992). Current outbreak and pest status of wingless grasshopper, Phaulacn"dium villatum, in southeastem Ausrralia. ln: Delfosse, E. S. (Ed.) Pests of pasture. Melbourne, CSIRO: 121-127. BAKER, G. L. & HOLMES, H.l'vl. (1986). Prorracted parasitic development of mermithid parasites of Orthoptera under drought conditions. Revue Nématol., 9: 287. BAKER, G. L. & POINAR, G. O.,Jr. (1986). Mermis quirindiensis n. sp. (Nematoda: Mermithidae), a parasite of locusts and grasshoppers (Orthoptera : Acrididae) in south-eastern Ausrralia. Revue NémaLOl., 9 : 125-134. BAKER, G. L. & POtNAR, G. O. Jr. (1988). A description of male and redescription of female Mermis alhysanota Steiner, 320 1921 (Nematoda : Mermithidae). Revue NémaLOI., Il : 343350. BLUNCK, H. (1939). Natürliche Feinde und biologische Bekiimpfung der Maikiiferengerlinge. Z. PflSchulZ 49: 338381. CHEN, L. G. & YAt'lG,]. S. (1985). [Efficiency of Amphimermis sp. for conrrolling and forecasting the brown plant hopper, Nilaparvata lugens stal.] Acta PhyLOphyl. Sinica, 12: 151-157. CHOO, H. Y., KAYA, H. K. & K1M, J. B. (1989). Agamermù unka (Mermithidae) parasitism of Nilaparvata lugens in rice fields in Korea. J. Nematol., 21 : 254-259. HAGMELER, A. (1912). Beirriige zur Kennmis der Mermithiden. 1. BioJogische Notizem und systematische Beschreibung einiger alter und neuer Arten. Zool. Jahrb. Syst., 32 : 521-612. HERRON, G. A. & BAKER, G. L. (1991). The effect of host stage and temperature on the development of Hexamermis sp. (Nematoda : Mermithidae) in the Ausrralian plague 10cust ChorLOiceles aref~nimrel (Walker) (Orthoptera: Acrididae). NemaLOlogica, 37: 213-224. HOMfNlCK, W. M., CROfT, S. L. & KUZOE, F. A. S. (1982). Description of parasitic stages of Hexamermis glossinae (Nematoda: Mermithidae) from Glossina in West Africa, with observations on the host parasite association. Canad. J. Zool., 3370-3376. KABURAKl, T. & lMAMuRA, S. (1932). A new mermithidworm parasitic in the rice borer with notes on its life history and habits. Proc. bnp. Acad. Japan, 8 : 109-1 12. KAlSER, H. (1972). Mermithidae (NemaLOda) ais Parasùen des KarlOffelkiifers (Leptinotarsa decemlineata Say.) in de?· SLeiemark. Ph. D. Thesis, Karl Franzens University, 137 p. KAISER, H. (1991). Terresrrial and semiterresrrial mermithidae. ln: Nickie, W. R (Ed.). Manual ofagricultural nemalOlogy. New York, Marcel Dekker : 899-965. KEy, K. H. L. (1959). The ecology and biogeography of Ausrralian grasshoppers and locusts. ln: Keast, A., Crocker, A. L. & Christian, C. S. (Eds). Biogeography and ecology in Auslralia. The Hague, W. Junk: 192-210. KEY, K. H. L. (1986). A provisional synoptic list of the Australian Acridoidea (Orthoptera). Technical Paper, Division of EnlOmology, CSIRO 24, 47 p. KIRJANOVA, E. S., KARAvAEVA, R. P. & ROMANENKO, K. E. (1959). [Mermithids (Mermithidae, Nematodes)- parasites of Hyponomeuta matinella and H. padella in Southem Kirgiz]. Trudy Kirgiz. lesn. opYt. Sta, 2: 195-240. KOVAL, Yu. V. (1977). [parasites of the Colorado beetle]. Vesl. sel'-Khoz Nauk1~ Alma-Ata, 9 : 38-44. LI, F. C. (1981). [A study of a species of Mennithoidea parasitising the planthoppers.] Nal. Enemies lnsects, 3 : 46-50. LI, H. K. (1985). Entomopathogenic microorganisms of ri ce planthoppers and leafhoppers in China. lnt. Rice Res. News/., 10: 13-14. MIRALLES, D. A. B. De & CAMINO, N. B. (1983). Una nueva especie de Merrnithidae: Amphimermis bonaerensis n. sp. (Nematoda : Enoplida). Neolropica, 29 : 153-156. Fundam. appl. NemalOl. Amphimermis spp. NIClQE, W. R. (1972). A contribution ro our knowledge of the Mermithidae (Nemaroda). J. NemaLOI., 4 : 113-146. POINAR, G. O., Jr. (1975). Emomogenous nemaLOdes. Leiden, E. J. Brill, 317 p. POfNAR, G. O., Jr. (1983). The nalUral hisLOry of nemaLOdes. New Jersey, Prentice-Hali, 323 p. POINAR, G. O.,Jr. & OTlENO, W. A. (1974). Evidence of four moults in the mermithidae. NemaLOlogica, 20: 370. POINAR, G.O., Jr. & WELCH, H. E. (1981). Parasites of invertebrates in the terres trial environments. In. Slusarski, ';X,!. (Ed.), Review of advanees in parasùowgy. Warszawa, PWNPolish Scientific Publ. : 947-954. POLOZHENTSEV, P. A. & ARTHYUKHOVSKJ, A. K. (1958). [New mermithid species]. Zool. Zh., 37: 997-1005. RUBTSOV, 1. A. (1971). [New species of mermithids from the southern Primorsk region]. Zool. Zh., 50 : 1143-1 153. RUBTsov,1. A. (1974). [A new merrnithid species from Lentaromidae in Azerbaidzhan]. Dokl. Akad. Nauk azerb. SSR, 30: 63-66. RUBTsov,1. A. (1975). [Merrnithids from the Georgian SSR]. ln: Sublropieheskie kul-tury. Vsesoyuznyi nauehno-issledovar.el'skii Inslùul ehaya i Sublropieheskikh Kuhur. Makharadza, Georgian SSR. Mirusterstvo Sel'skogo Khozyaisrva USSR : 109-112. RUBTsov,1. A. (J 976 a). [Merrnithids from Kirgizia]. ln : EnLOmowgicheskie issledovaniya v Kirgizii. "flim " Frunza, Il : 87-100. RUBTsov,1. A. (1976 b). [Mermithids (Nematoda, Merrnithidae) parasites of insects in Mongolia]. In: Nasekomye Mongolii, Akademiya Nauk SSR, Zoologieheskii Inslùul, No. 4, Lerungrad : Izdatel'srvo " Nauka ", Leningradskoe Otdelenie: 596-614. RUBTSOV, 1. A. (1977). [New species of mermithids from spiders (Pardosa) and earwigs (Forficula)]. In : Taksony fauny Sibiri. Novosibirsk, USSR; " Nauka " Sibirskoe OtdeJenie : 16-22. RUBTSOV, 1. A. (1978 a). [MenniLhidae, elassi{icaLion, imporLance, Ulilisalion]. Leningrad, Nauka, 207 p. RUBTSOV, 1. A. (1978 b). [New species of merrrùthids from BuJgaria]. Khelmimologiya, Sofia, 5 : 97-106. RUBTSOV, 1. A. (1979). [New species of mermithids from arthropods in Azerbaidzhan). fzv. Akad. Nauk azerb. SSR, biol. Nauki. 6 : 89-95 Vol. 17, n° 4 - 1994 RUBTSOV,1. A. & KOVAL, Yu. V. (1975). [Mermithids from the Colorado beetle] Trudy Vses. nauehno-issled. Ins/. Zasfu:.h. Ras/., 44: 126-153. SCHUUIU,,!ANS STEKHOVEN, J. H. (1950). Némarodes des grottes et des eaux souterraines de Roumanie. Bull. Insl. Roy. Sei. na/. Bruxelles, 26, 61 : 2-3. SHIMKJNA, M. A. (1978). [Re1ationships berween metmithids (Nematoda: Merrnithidae) and soil arthropods in the Ravine Forests of Prisamar'ya]. Probl. poehv. Zool., M.insk, USSR: 277-278. SPIRIDONOV, S. E. (1987). [Soil merrnithids of Tonga and Western Samoa]. Trudy Gel'mimh. Labo. (Mor/ol., Taks. Ekol. Gel'mim. Zhivo. i Rasl.), 35 : 135-139. STREET, D. A. & McGUlRE, M. R. (1990). Pathogeruc diseases of grasshoppers. In: Chapman, R. F. & Joern, A. (Eds) : Biowgy ofgrasshoppers. New York, J. Wiley & Sons: 483-516. VYAS-PATEL, N. (1992). The comparative deveJopment of the cuticle of Romanomermis eulieivomx in susceptible and resistant hosts. Fundam. appl. NemaLOl., 15 : 141-147. WANG, J. A. & LI, L. Y. (1987). Entomogenous nematode research in China. Revue NémaLOI., 10 : 483-489. WEBSTER,J. M. & THONG, C. H. S. (1984). Nemarode parasites of orthopterans. In : Nickle, W. R. (Ed.). Plam and inseel nemaLOdes. New York, l\1arcel Decker: 697-726. WELCH, H. (1963). Amphimermis bogongae sp. nov. and Hexamermis eavicola sp. nov. from the Australian bogong moth, AgrOlis infusa (Boisd.), \Vith a review of the genus Amphimermis Kaburaki & Imamura, 1932 (Nematoda: Mermithidae). Parasùology, 53 : 55-62. WlLLlS, O. R. (1971). A mermithid nematode in naiads of damselflies (Odonata : Coenagriorudae). Fior. Enlomologisl, 54: 321-324. XIA, K. X. (1989). [Distinguishing berween t\vo merrnithid nematodes (A mphimermis and Agamermis) in the brown p1anthopper Nilaparvala lugens]. Nal. Enemies Inseels, 3 : 148-149. 321