Taeniogonalos woodorum Smith, sp. n.

Smith, David R., Janzen, Daniel H., Hallwachs, Winnie & Smith, M. Alexander, 2012, Hyperparasitoid wasps (Hymenoptera, Trigonalidae) reared from dry forest and rain forest caterpillars of Area de Conservacion Guanacaste, Costa Rica, Journal of Hymenoptera Research 29, pp. 119-144 : 125-129

Taeniogonalos woodorum Smith, sp. n. Figures 4-7

Diagnosis.

Posterior postocellar area with two yellow oblique stripes; mandible mostly yellow, teeth reddish brown; gena mostly yellow, yellow extending dorsally and onto occiput; mesoscutellum with anterior third yellow; second metasomal tergum with narrow yellow band continuous along lateral and posterior margins. Female armature absent. Male with medial flattened area on sternum 2; genitalia with parameres about half as long as gonostipes.

Female. Length, 5.0-12.0 mm (holotype 8.0 mm). Antenna black. Head black, with following yellow ( Figs 4, 5 ): Interantennal area, supraclypeal area, clypeus, labrum, mandible except reddish-brown apex, mouthparts, two oblique stripes at posterior or postocellar area, broad stripe on gena behind eyes extending dorsally onto occiput. Mesosoma black with following yellow markings ( Figs 4, 5 ): anterolateral spot on mesoprescutum, axillae, anterior third of mesoscutellum, metascutellum, stripe on upper pronotum and lower pronotum, broad oblique stripe on mesopleuron, broad oblique stripe on metapleuron, large lateral spots on propodeum. Legs black with inner surfaces of coxae, trochanters, extreme bases of femora, and outer surfaces of tibiae and tarsi yellow. Metasoma black with following yellow markings ( Figs 4-6 ): broad stripe on posterior margin of tergum 1, narrow continuous stripe on posterior and lateral margins of tergum 2; narrow stripe on posterolateral margins of terga 3 and 4, small spot on posterior lateral margin of sternum 2. Wings hyaline, black anterioapically, mostly in radial cell; veins and stigma black. Head and body covered with fine, short, white hairs. Head: Covered with short, white hairs. Shiny, evenly punctate, punctures mostly separated by rounded ridges, interspaces less than puncture diameters; punctures on gena less dense, farther apart than those on vertex and frons, and with flat shiny interspaces. Antenna with 24 flagellomeres, length about twice head width. Clypeus with median length about .3 × width. Inner margins of eyes subparallel, lower interocular distance 0.7 × eye length; malar space about 0.15 × eye length. Distance between eye and margin of lateral ocellus about 3.0 × distance between inner margins of hind ocelli. Area between torruli slightly concave ( Fig. 5 ). Antennal carinae low, sharp. Occipital carina complete. Mesosoma: Covered with short, white hairs. Shiny, evenly punctate with punctures similar to those on vertex and frons, most separated by rounded ridges, with interspaces less than puncture diameters; punctures on propleuron farther apart than those on mesonotum and separated by broader, flat interspaces; dorsoposterior section of mesepisternum, posterior downturned margin of mesoscutellum, and metapleuron, except lower margin, almost impunctate; punctures on propodeum small, denser than those on rest of mesosoma. Prescutum elevated above lateral lobes. Notaulus deep, with large punctures posteriorly; transverse row of large punctures anterior to mesoscutellum. Propodeal foramen shallowly concave at center. Hind coxa about as long as wide, with longitudinal carina on outer surface; hind basitarsomere subequal to length of remaining tarsomeres combined. Metasoma: Covered with fine, white hairs. Shiny, rather evenly punctate, punctures separated by rounded ridges mostly less than puncture diameters. Tergum 1 depressed at center. Armature absent from sternum 2 ( Figs 4, 6 ). Sheath directed downward, rounded at apex in lateral view.

Male. Length, 4.0-7.5 mm. Color and structure similar to female. Tyloids present, long and narrow, middle tyloids longer than half length of flagellomeres. Male genitalia with parameres about half-length of gonostipes ( Fig. 7 ); sternum 2 with medial flattened area on apical half.

Type material.

Holotype female, "Voucher: D. H. Janzen & W. Hallwachs, DB: http://Janzen.sas.upenn.edu, Area de Conservation Guanacaste, Costa Rica," “10-SRNP-22162,” “DHJPAR0041177.” (USNM). Paratypes: Same data as for holotype except caterpillar rearing code (yy-SRNP-xxxx) and parasitoid individual code (DHJPARxxxxxxx); one specimen for each caterpillar collection and parasitoid rearing code. 98-SRNP-6785, DHJPAR0010613 (♂); 98-SRNP-7262, DHJPAR0016904 (♂); 98-SRNP-7361, DHJPAR0016911 (♀); 98-SRNP-15545, DHJPAR0016916 (♀); 98-SRNP-15545, DHJPAR0016899 (♀); 98-SRNP-15545, DHJPAR0016888 (♂); 99-SRNP-5503, DHJPAR0016895 (♀); 99-SRNP-5508, DHJPAR0016897 (♂); 99-SRNP-12098, DHJPAR0016891 (♀); 99-SRNP-12098, DHJPAR0016892 (♂); 99-SRNP-12098, DHJPAR0016896 (♀); 99-SRNP-12761, DHJPAR0010612 (♀); 99-SRNP-12852, DHJPAR0010604 (♂); 99-SRNP-13819, DHJPAR0010611 (♂); 99-SRNP-13823, DHJPAR0016909 (♀); 01-SRNP-3507, DHJPAR0010598 (♀); 01-SRNP-3507, DHJPAR0010599 (♀); 01-SRNP-5325, DHJPAR0010597 (♂); 01-SRNP-5932, DHJPAR0010605 (♀); 01-SRNP-9359, DHJPAR0010607 (♀); 01-SRNP-25186, DHJPAR0010600 (♂); 02-SRNP-7679, DHJPAR0010596 (♀); 02-SRNP-7978, DHJPAR0010595 (♀); 03-SRNP-6738, DHJPAR0010588 (♂); 03-SRNP-10070, DHJPAR0010585 (♀); 03-SRNP-11855, DHJPAR0010591 (♂); 03-SRNP-11855, DHJPAR0010592 (♂); 03-SRNP-11855, DHJPAR0010593 (♂); 03-SRNP-11855, DHJPAR0010594 (♂); 03-SRNP-20157, DHJPAR0010590 (♀); 03-SRNP-20236, DHJPAR0028047 (♀); 03-SRNP-21817, DHJPAR0010586 (♀); 04-SRNP-30072 [no barcode] (♀); 04-SRNP-41595, DHJPAR0010574 (♂); 04-SRNP-55214, DHJPAR0010571 (♀); 04-SRNP-55214.1, DJHPAR0010572 (♀); 04-SRNP-55215, DHJPAR0010573 (♀); 04-SRNP-56432, DJHPAR0010581 (♀); 04-SRNP-56458, DHJPAR0010582 (♀); 05-SRNP-4939, DHJPAR0010559 (♂); 05-SRNP-7881, DHJPAR0010551 (♂); 05-SRNP-33818, DHJPAR0010569 (♂); 05-SRNP-34358, DHJPAR0010562 (♂); 05-SRNP-42584, DHJPAR0010563 (♂); 05-SRNP-42827, DHJPAR0010570 (♂); 05-SRNP-70122, DHJPAR0010560 (♂); 05-SRNP-70325, DHJPAR0010561 (♀); 06-SRNP-6781, DHJPAR0016876 (♀); 06-SRNP-6781, DHJPAR0016877 (♀); 06-SRNP-6781, DHJPAR0016878 (♂); 06-SRNP-6781, DHJPAR0016884 (♀); 06-SRNP-30294, DHJPPAR0010443 (♂); 06-SRNP-30295, DHJPAR0010554 (♂); 06-SRNP-33412, DHJPAR0016873 (♀); 06-SRNP-34200, DHJPAR0016875 (♀); 06-SRNP-34577, DHJPAR0016886 (♀); 06-SRNP-34579, DHJPAR0016887 (♂); 06-SRNP-42284, DHJPAR0010555 (♀); 06-SRNP-42284, DHJPAR0010556 (♂); 06-SRNP-42284, DHJPAR0010557 (♂); 06-SRNP-42814, DHJPAR0016882 (♀); 06-SRNP-42819, DHJPAR0016883 (♀); 06-SRNP-43560, DHJPAR0016874 (♀); 06-SRNP-65304, DHJPAR0016885 (♂); 08-SRNP-2414, DHJPAR0027762 (♂); 08-SRNP-2414, DHJPAR0027763 (♂); 08-SRNP-6017, DHJPAR0030373 (♂); 08-SRNP-32269, DHJPAR0030377 (♀); 08-SRNP-41835 [no barcode] (♀); 08-SRNP-42172, DHJPAR0030374 (♀); 08-SRNP-42172, DHJPAR0030375 (♀); 08-SRNP-42172, DHJPAR0030376 (♀); 09-SRNP-2888, DHJPAR0036406 (♀); 09-SRNP-5008, DHJPAR0036682 (♀); 09-SRNP-32681, DHJPAR0038544 (♀); 09-SRNP-32752, DHJPAR0038545 (♀); 09-SRNP-32752, DHJPAR0038546 (♀); 09-SRNP-33385, DHJPAR0038543 (♂); 09-SRNP-69541, DHJPAR0036404 (♂); 09-SRNP-70610, DHJPAR0036405 (♀); 09-SRNP-73449, DHJPAR0037846 (♂); 09-SRNP-80526, DHJPAR0037847 (♂); 10-SRNP-1030, DHJPAR0039355 (♀); 10-SRNP-4609, DHJPAR0041181 (♀); 10-SRNP-22641, DHJPAR0041178 (♂); 10-SRNP-32041, DHJPAR0041179 (♂); 10-SRNP-42215, DHJPAR0041180 (♀); 10-SRNP-73124, DHJPAR0041176 (♀); 10-SRNP-73289, DHJPAR0041174 (♀); 10-SRNP-80666, DHJPAR0041175 (♂); 11-SRNP-2784, DHJPAR0045823 (♀); 11-SRNP-2784, DHJPAR0045824 (♂); 11-SRNP-2859, DHJPAR0044983 (♀); 11-SRNP-2911, DHJPAR0045822 (♂); 11-SRNP-71666, DHJPAR0045825 (♂); 11-SRNP-80954, DHJPAR0044984 (♀). Deposited in INBio, USNM, CNC, BMNH.

Other specimens.

03-SRNP-38118, DHJPAR0010587 (metasoma missing); 06-SRNP-6781, DHJPAR0016872 (metasoma missing); 09-SRNP-72860, DHJPAR0040090 (broken).

Specimens examined.

100; 98 submitted for DNA barcoding, 89 of which yielded complete DNA barcodes publically available from BOLD.

Etymology.

Taeniogonalos woodorum is named in honor of Monty and Grace Wood of Ottawa, Canada in recognition of their three decades of intense and enthusiastic taxonomic and spiritual participation in the tachinid fly inventory of Area de Conservación Guanacaste and in INBio’s inventory of the flies of Costa Rica.

Barcode.

The DNA barcodes of the 89 Taeniogonalos woodorum specimens longer than 400 bp have less than 1% intraspecific divergence (0.702% avg, max 2.523%, min, 0%). They are 9% divergent from the DNA barcodes of Taeniogonalos fasciatipennis DHJ01 and Taeniogonalos fasciatipennis DHJ02.

Discussion.

The mostly black color with yellow maculation, as described and illustrated, and lack of female armature on metasomal sternum 2 help distinguish this species from most other New World species of Taeniogonalos . The females of Taeniogonalos fasciatipennis , Taeniogonalos gundlachii , Taeniogonalos lugubris (Westwood), and Taeniogonalos ornata (Smith) have distinct armature on sternum 2, and the latter three are mostly yellow or reddish-brown with black maculation. The females of Taeniogonalos enderleini (De Santis) and Taeniogonalos jucunda (Westwood) from South America lack female armature. Taeniogonalos enderleini occurs in southeastern Brazil, is mostly black with some yellow maculation, but the posterior lower part of the mesopleuron and the metapleuron are reddish brown and the postocellar area lacks yellow marks. Taeniogonalos jucunda (Westwood) was described from “Amaz.”, and the color was described as mostly reddish brown, head yellow, and the scutellum black, all of which differ from the color of Taeniogonalos woodorum .

Hosts and biology.

Taeniogonalos woodorum is the most frequently reared of the ACG Trigonalidae , known only from ACG rain forest, and superficially resembles Taeniogonalos fasciatipennis and Taeniogonalos gundlachii (see below). It is the only species of trigonalid that has been reared from the sample of more than 220,000 wild-caught ACG rain forest caterpillars. This microgeographic and ecosystem separation from the parapatric and adjacent ACG dry forest of Taeniogonalos fasciatipennis (see below) allows first-pass species-level identification of Taeniogonalos woodorum even if key morphological traits are invisible and DNA barcodes have not been obtained from the specimen, such as when the reared wasp escapes or the abdomen is broken off and lost or consumed in analysis. This method of ecology-based identification cannot, however, be used for specimens from the narrow ecotone between ACG dry forest and rain forest, where both species of Taeniogonalos have been reared from caterpillars found in the same hectare. The presence of Taeniogonalos woodorum was first noticed in 2006 by its strikingly different (15%) DNA barcode from that of Taeniogonalos fasciatipennis (also called Taeniogonalos gundlachii at that time). Adult Taeniogonalos woodorum , as is the case with the other ACG Taeniogonalos , is a Batesian and Mullerian mimic of Polybia wasps ( Vespidae ; abundant in ACG) in body size, color pattern, and flight/walking behavior.

Taeniogonalos woodorum has been reared 97 times from 14 caterpillar families ( Arctiidae , Crambidae , Elachistidae , Geometridae , Hesperiidae , Lasiocampidae , Noctuidae , Notodontidae , Nymphalidae , Pyralidae , Saturniidae , Sphingidae , Thyrididae , Uraniidae ), parasitized by Braconidae ( Bassus , Dolichogenidea , Glyptapanteles , Meteorus , Stantonia , Zelomorpha ), Ichneumonidae ( Agrypon , Charops , Dusona , Eiphosoma , Hyposoter , Leurus , Microcharops , Xiphosomella , Zaglyptomorpha ) and Tachinidae (at least Anoxynops , Agryrochaetona , Argyrophylax , Belvosia , Calolydella , Campylochaeta , Chrysotachina , Drino , Eucelatoria , Eujuriniodes , Eumea , Genea , Houghia , Hyphatrophaga , Lespesia , Patelloa , Phytomyptera , Winthemia ). The host caterpillars of these primary parasitoids may all be characterized as exposed leaf feeders (even these pa rticular species of leaf rollers and tiers, Crambidae , Elachistidae , Pyralidae , Thyrididae , venture out of their leaf-silk nests to feed on fully exposed leaf blades), and no one species dominates this diverse list. While it is evident that Taeniogonalos woodorum can develop in a very wide variety of host caterpillars and primary parasitoids, experience with other apparent “generalists” in the ACG inventory warns that when much larger sample sizes have accumulated, it may become evident that certain taxa and ecologies are either being avoided by ovipositing wasps or the eggs/larvae do not survive their tour in the host or primary parasitoid.

It remains a mystery as to why this hyperparasitoid remains microgeographically restricted to ACG rain forest and does not venture into the extensive adjacent dry forest with its many thousands of species of potential caterpillar and primary parasitoid hosts only a few hundred meters away. Indeed, there is a single record of Taeniogonalos woodorum (DHJPAR0016846) well into the microgeographic distribution of Taeniogonalos fasciatipennisDHJ02 (see below), emphasizing the parapatric nature of the distribution of these two species of Taeniogonalos . However, in remaining restricted to rain forest, it is representative of the thousands of other species of ACG Lepidoptera , Hymenoptera , and Diptera which are faithful to their respective ecosystems, even at the time when the intense six month rainy season turns the adjacent dry forest in a very wet ecosystem.

Figures

Figures 4 - 7. Taeniogonalos woodorum. 4 Lateral view, female 5 Dorsal view, female 6 Metasoma, lateral view, female 7 Male genitalia.

Figures 8 - 10. Taeniogonalos fasciatipennis, female. 8 Taeniogonalos fasciatipennisDHJ 01, lateral view 9 Taeniogonalos fasciatipennisDHH 01, dorsal view 10 Taeniogonalos fasciatipennisDHJ 02, lateral view.

Figures 15 - 20. Taeniogonalos gundlachii. 15 Lateral view, female 16 Dorsal view, female 17 Metasoma, lateral view, female 18 Female armature, sternum 2, ventral view 19 Male paramere, lateral view 20 Male genitalia.