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Speciation and Adaptive Radiation in the Fig Wasp Nematode, Parasitodiplagaster (Diplogasteridae : Rhabditida) in Panama George O

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Speciation and Adaptive Radiation in the Fig Wasp Nematode, Parasitodiplagaster (Diplogasteridae : Rhabditida) in Panama George O Speciation and adaptive radiation in the fig wasp nematode, Parasitodiplagaster (Diplogasteridae : Rhabditida) in Panama George O. POINAR,Jr. and Edward A. HERRE Department of Entomological Sciences, University of California, Berkeley, CA 94720, USA and Snzithsonian Tropical Research Institute,Apartado 2072, Balboa, Republic of Panama, SUMMARY The following species of the hereby redefined genus Parasitodiplogaster are described from fruits of Ficus spp. in Panama : P. citrinema n. sp. from F. citrifolia P. Miller, P. duganema n. sp. from F. dugandii Standl., P. mminema n. sp. from F. maxima P. Miller, P. nymphanema n. sp. frorn E nymphaeifolia L., P. obtusinema n. sp. frorn E obtusifolia HBK, P. paranema n. sp. from F. paraensis (Miq.) Miq., P. pertanema n. sp. from F. pertusa L. f., P. popenema n. sp. from E popenoei Standl., P. trigonema n. sp. from E trigonata L. and P. yoponema n. sp. from E yoponensis Desv. Al1 of the above species, which are presumed to have evolved from a free-living diplogasterid stem species, represent alloparric species which are carriedby specific figwasp species to specific fig fruitspecies. The diverse morphological characters (especially those ofthe alimentary tract) exhibited among these ten new species of fig wasp nematodes represent examplesof selective adaptation to physical and nutritional conditions foundin the fruits (sycones) of the different fig species. &SUME Spéciation et divergence adaptative chez le nématode de la guêpe du figuierParasitodiplogaster (Rhabditida :Diplogasteridae) au Panama Le genre Parasitodiplogaster est redéfini et les espèces nouvelles suivantes, provenant de fruits de Ficus spp. au Panama, sont décrites :P. citrinema n. sp. surE citrifolia P. Miller, P. duganema n. sp. surE dugandii Standi., P. maxinema n. sp.sur F. maxima, P. Miller, P. nymphanenza n. sp. sur E nymphaeifolia L., P. obtusinema n. sp. sur F. obtusifolia HBK, P. paranema n. sp. sur F. paraensis (Miq.) Miq., P. pertanema n. sp. sur E pertusa L. f., P. popenenza n. sp. sur F. popenoei Standl., P. trigonema n. sp. sur F. trigonata L. et P. yoponema n. sp. surF. yoponensis Desv. Présumées avoir évoluéa partir d'unrameau de Diplogastérides libres, elles forment un groupe d'espèces alloparriques, transportées par des guêpes spécifiquessur des fruits de figuiers spécifiques. Les variations des caractères morphologiques présentes chez ces dix espèces (en particulier ceux liés au tractus digestif) représentent un exemple d'adaptation sélective aux conditions physiques et alimentaires des fruits (des sycones) des différentes espèces de figuier. The genus Parasitodiplogaster Poinar, 1979, was paraensis (Miq.)Miq., E pertusa L. f., E popenoei described as a parasite of the fig wasp Elisabethiella Standl., E trigonata L. and E yoponensis Desv. by EAH stuckenbergi Grandi (Agaonidae : Hymenoptera) from in the vicinity of the Panama Canal, Republic of Pa- the fruits of Ficus burkei (Miq.) Miq. in Rhodesia. nama. The figs were placed in 70 % alcohol and sent to Several years ago, one of us (EAH) discovered ne- GOP, Jr. for determination. The preserved fruits were matodes in thefruits of different Ficus species in cut open and adult nematodes were removed from the Panama. These nematodes al1 belong to the genus Pa- cavity and processed to glycerin for taxonomic studies. rasitodiplogaster, which is the first record of this group In some cases, juvenile nematodes were also removed in the New World. The present study redefines the from adult fig wasps found inside the fruits. genus Parasitodiplogaster and describes ten new species The anterior pharynx was measured from the tip of from Panamanian fig fruits. the head to theline separatingthe anterior and posterior pharynx; the posterior pharynx was measured from the line separating the anterior and posterior pharynxto the base of the posterior pharynx. Specific names of the Materials and methods nematodes are based on thespecific names of the Ficus spp. in which the nematodes occurred. In illustrations Figfruits were collected from wild trees of Ficus of the pharyngeal regionof the variousParasitodiplogas- citrifolia P. Miller, E dugandii Standl., E maxima P. ter species, the pattern of vertical lines represents strong Miller, E nymnphaez$olia L., E obtusifolia HBK, E musculature,broken vertical linesrepresents weak Revue Nématol. 14 (3) :361-374 (1991) 361 G. O. Poinar, Jr. & E. A. Herre musculature and fine wavy lines represents glandular Parasitodiplogaster citrineman. sp. tissue. (Figs lB, 3C) MEASUREMENTS Parasitodiplogaster Poinar, 1979 Adttlts :See Tables 1 and 2. DIAGNOSIS(emended) Diplogasteridae(Micoktzky, 1922) Steiner, 1929. DESCRIPTION Adults variable in size, ranging from 0.5 to 4.0 mm in Females :Medium to large nematodes with a well- length; cuticle smooth or with faint annulations, often developed stoma and very muscular anterior pharynx swollen aroundvulva; six lipspartly or completely wi& inconspicuous valve plates at base; base of anterior fused; amphids pore-like, on lateral lips; stomavariable, , pharynxdistinctly wider than proceedingposterior from reduced -to distinct;glottoid apparatus incon- pharynx; posterior pharynx short,also muscular; baseof spicuous or lacking; pharynx composed of an anterior posterior pharynx withvalve area; pharyngeal-intestinal portion containing faint to distinct crescentvalve plates valve well-developed; excretory pore opposite posterior near the base and a basal portion sometimes terminating pharynx;intestinal lumen distinct, wide, filled with in faintcrescent valve plates; ovaries paired, tips usually viscous material and containing both bacterial rodsand reflexed; four vaginal glands (variable in size) surround- cocci; vaginal glands large, cuticle expanded in region ing vagina; testis single, tip outstretched or reflexed; of vulva, especially on ventral side; tail bearing a fine spicules paired, separate, gubernaculum present, bursa mucron at tip. absent;pre-and post-cloaca1 genitalpapillae present; found in fig fruits (Ficus spp.) and fig wasps (Agaoni- Males : Generalfeatures same as describedunder dae : Hymenoptera). females:, testis outstretched; spicules short and wide; Table 1 Morphometric data for femalesof Parasitodiplogaster species. Species L Body Stoma Stama L. ant.pharynx Past. L. post. Ant. end Ant. end V L Width L Width width width L pharynxpharynx ht, pharynx to E. P. to N. R. tail tail egg egg (Pm)(Pm) (Pm) (Pm) Cm) (Pd (Pm) (Pm) (Pm) (Pm) (Pd (Pm) citrinelna n. sp. 1950 98 5 797 141 3'6 216 177 5032 9160 30 (N = 3) 1 800-2 040 45-120 5-8 75-81138-144 .52-.59 174-180 45-53 88-93 57-6330-36 dugonefnan. sp, 2 080 96 2 5 101 87 36 173 141 53 39 9758 36 (N = 4) 1 728-2 208 79-108 4-6 79-9298-105 31-.90 165-184 130-152 49-54 89-108 35-4157-5931-41 nwineam n. sp. 1 7892.5 47 695 159 .43 175 168 48 32 12654 28 (N = 3) 1 500-2 280 2-3 45-51 4-6 60-75150-168 .40-.47 165-180 159-180 46-51 102-150 24-3048-6024-39 rrvlllphanelm n. sp. 2 136 96 3 724 116 .61 197 137 50 132 35 (N = 10) 1 560-2 880 75-129 2-4 3-5 78-135 60-84 36-.69 135-273 96-159 44-63 75-183 24-48 obtirsinetm n. sp. 2 745 3.5 133 4.5 102 79 .78 256 150 51 81 168 124 48 (N = 2) 2 550-2 940 105-1623-4 4-5 99-105 78-81 ,7432 237-276 141-159 50-52 45-51120-129 72-90 paranerm? n. sp. 920 42 2 723 108 .66 75 129 43 69 20 (N= 1) pertaneau n. sp. 1747 109 , 6 668 127 .52 216 157 5436 8374 35 (N = 10) 1440-2 016 3-988-133 5-11 54-79120-136 .44-.63 159-279 133-174 38-73 63-98 32-3863-8625-48 popenenla n. sp. 2 001 87 4 825 164 .50 190 198 47 132 3237 66 (N = 7) 1740-2 190 3-5 75-93 4-7 144-183 69-102 .39-.60 150-240 159-240 37-52 105-162 33-4260-9024-36 tigonelna n. sp. 2 556 129 2 2 99 14 .73 210 128 5142 16274 52 (N = 10) 2 070-3 330 1-399-156 1-3 66-120 60-90 ,6588 171-279 90-159 48-55 129-180 42-6360-9039-45 yoponema n. sp. 1331 51 4 577 142 .39 122 117 5034 8764 24 (N = 5) 1088-1 600 4-5 47-54 6-8 41-73120-174 .30-.49 108-143 95-143 48-55 76-101 19-2954-7332-38 3 62 Revue Nimatal. 14 (3) :361-374 (1991) Parasitodiplogaster spp. in fig wasps , Table 2 Morphometric data for males of Parasitodiplogaster species. citrinenla n. sp. 1835 56 6 9 134 69 .52 23 1 176 92 46 29 8 18 4 (N = 5) 1 384-2240 45-76 4-8 6-13 126-143 60-82 .48-.57 171-260 153-193 77-10241-48 26-32 7-9 16-19 3-5 dugatmna n. sp. 1888 98 2 4 92 80 .87 187 129 83 42 45 9 20 4 (N = 3) 1760-1 98488-105 3-5 82-98 73-86 .86-.90 127-225 117-146 77-8840-43 39-48 8-10 17-21 3-5 tminema n. sp. 1098 30 3 5 144 65 .46 162 16025 50 36 6 22 4 (N = 5) 750-1350 21-42 2-3 4-6 117-156 57-75 .38-.59 144-198 138-180 19-2939-54 26-41 4-8 15-26 3-5 nylnphanenla n. sp. 761 150 3 4 120 68 36 158 13756 84 78 14 28 4 (N = 10) 960-1520 60-92 2-4 3-5 102-141 42-90 .41-.70 78-210 117-156 71-10551-64 66-84 12-15 22-33 3-6 obusinelna n.
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