Taeniogonalos fasciatipennis (Cameron)

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 : 129-132

Taeniogonalos fasciatipennis (Cameron) Figures 8 14

Trigonalys fasciatipennis Cameron 1897: 271.

Taeniogonalos gundlachii (in part): Carmean and Kimsey 1998: 67.

Discussion.

This species is distinguished as follows: vertex black; mandible outer surface black, upper surface yellow, teeth reddish brown; clypeus with large yellow spot on each side; narrow yellow line on gena on hind orbit ( Figs 8 , 10 ); mesoscutellum entirely yellow or with thin medial black stripe; second metasomal tergum with broad posterior transverse band, not extended laterally ( Figs 8, 10 ). Female armature present, in ventral view with lobes short and with shallow central depression ( Fig. 12 ), in lateral view, posterior and ventral sides perpendicular, not directed downward at apex ( Figs 8 , 11 ); male genitalia with paramere long, about three-quarters length of gonostipes ( Fig. 14 ); paramere in lateral view with dorsal margin straight ( Fig. 13 ), without slight depression.

Taeniogonalos fasciatipennis was described by Cameron (1897) from two Mexican specimens, a female from "Atoyac in Vera Cruz" and a male from "Venta de Zopilote in Guerrero." The male from the State of Guerrero was chosen as the lectotype by Carmean and Kimsey (1998). This lectotype (BMNH, examined) is morphologically identical to males reared as this species from ACG. It is noteworthy that the region around the type locality in Mexico is ecologically the same kind of tropical dry for est ecosystem as is the dry forest of ACG and shares a very large number of species of plants and insects with it.

The female syntype (not examined) from Veracruz was treated by Carmean and Kimsey (1998) as the “same” as Taeniogonalos gundlachii . It may not be the same species as the lectotype, and some of the specimens treated as Taeniogonalos fasciatipennis by Carmean and Kimsey (1998) are probably the species we here describe as Taeniogonalos woodorum .

This and the species from North America and Cuba have long been regarded as the " gundlachii " group, the members of which are distinguished from others by having similar color and the presence of distinctive female armature. However, Taeniogonalos fasciatipennis can be distinguished from North American Taeniogonalos gundlachii by the female armature and male parameres as described above. Taeniogonalos fasciatipennis is morphologically separable from Taeniogonalos woodorum by the former having the female armature present and the male paramere short in relation to the gonostipes.

The DNA barcodes of Taeniogonalos fasciatipenni sDHJ01 and Taeniogonalos fasciatipennis DHJ02 are 5.66% divergent from each other in the COI barcode region and both are 9% divergent from Taeniogonalos woodorum .

Distribution.

Mexico and Costa Rica.

Specimens examined.

Taeniogonalos fasciatipennisDHJ01 53, 42 barcoded; Taeniogonalos fasciatipennisDHJ02 15, 13 barcoded. Deposited in USNM, INBio, CNC, BMNH.

Hosts and biology.

In the absence of genetic information, Taeniogonalos fasciatipennis appears to be one morphologically distinctive species that occurs throughout ACG dry forest (85 m to about 1300 m elevation) and does not occur in adjacent ACG rain forest or cloud forest. It is a hyperparasitoid of a wide variety of caterpillar and primary host species (see below). However, DNA barcoding has revealed that this morphologically-distinct species is divided into two distinct mitochondrial types in the ACG dry forest. One, baptized here as Taeniogonalos fasciatipennisDHJ01 ( Figs 8, 9 , 11-14 ) is an interim name in the inventory website database (http://janzen.bio.upenn.edu/caterpillars/database.lasso) and occurs throughout the ACG dry forest (85 to 700 m elevation). The other, interim name Taeniogonalos fasciatipennisDHJ02 ( Fig. 10 ), has a very distinctive microgeographic distribution. Twelve of the 13 records are from the intermediate elevation southwest facing slope of the volcanic massif of Rincon de la Vieja (325-1276 m elevation in Sector Mundo Nuevo of ACG). The single other rearing record (two individuals from tachinid fly puparia from the same caterpillar) is from a site that is an ecogeographic analogue in biota, elevation, and climate to the Sector Mundo Nuevo site (600 m on the southwest facing slope of Volcan Cacao in Sector Cacao of ACG). Taeniogonalos fasciatipennisDHJ01 is probably the same as the lectotype from western (dry forest) Mexico, but since there is no present way to know for certain, both Taeniogonalos fasciatipennisDHJ01 and Taeniogonalos fasciatipennisDHJ02 have to be treated as interim names, and are therefore not italicized (and see below).

In general terms, Taeniogonalos fasciatipennisDHJ01 conspicuously ranges from the driest parts of ACG dry forest to its intermediate-elevation intersection with cloud forest and rain forest, and Taeniogonalos fasciatipennisDHJ02 is restricted to the upper, cooler, moister portion of this range. To emphasize the cooler and moister aspect of this very small range, there is even a single specimen (DHJPAR0016846) of the rain forest specialist, Taeniogonalos woodorum , from the very center of the range of Taeniogonalos fasciatipennisDHJ02. While Taeniogonalos fasciatipennisDHJ01 has not been found in the Sector Mundo Nuevo exact site occupied by Taeniogonalos fasciatipennisDHJ02, and thus they are parapatric, Taeniogonalos fasciatipennisDHJ01 is absolutely sympatric with the single record of Taeniogonalos fasciatipennis DHJ02 on the southwestern slopes of Volcan Cacao.

This situation creates two different taxonomic problems. First, since the two apparent ACG species within what appears morphologically to be Taeniogonalos fasciatipennis currently cannot be distinguished by any morphological trait, there is no way to know which of the two matches the lectotype from the State of Guerrero, Mexico. It is also possible that neither does. Second, the presence of two sympatric/parapatric species of " Taeniogonalos fasciatipennis " in the small area of ACG raises the spectre that this species of wasp may be a complex of sibling (cryptic) species. In contrast to parallel cases with extremely host-specific Microgastrinae Braconidae (e.g., Smith et al. 2008) or Tachinidae ( Smith et al. 2007), the unambiguously generalist host-selection and/or larval-survival ability of Taeniogonalos fasciatipennis excludes the use of host records as a way to verify or predict the presence of cryptic species.

If Taeniogonalos fasciatipennisDHJ02 were not distinctive by barcode from Taeniogonalos fasciatipennisDHJ01 ( Fig. 2 ), it would not have been noticed. Taeniogonalos fasciatipennisDHJ01 and Taeniogonalos fasciatipennisDHJ02 have the same extremely varied lists of caterpillar and primary parasitoid hosts. In brief, Taeniogonalos fasciatipennisDHJ01 has been reared and barcoded 48 times from nine Lepidoptera families: Crambidae , Hesperiidae , Megalopygidae , Noctuidae , Nymphalidae , Papilionidae , Saturniidae , Sphingidae , and Uraniidae . While 53 more specimens of Taeniogonalos fasciatipennis have been raised, until (and if) those specimens are successfully barcoded, we cannot categorize them as Taeniogonalos fasciatipennisDHJ01 or Taeniogonalos fasciatipennisDHJ02.

In all cases, the primary parasitoid host was Tachinidae : Acantholespesia , Belvosia , Blepharipa , Calolydella , Carcelia , Chetogena , Drino , Eucelatoria , Hemisturmia , Houghia , Hyphantrophaga , Leschenaultia , Lespesia , Patelloa , Winthemia , and Zizyphomyia . If we add to this the other specimens of " Taeniogonalos fasciatipennis " that were not successfully barcoded but occupy the ACG ecosystem occupied by Taeniogonalos fasciatipennisDHJ01 a few more Lepidoptera families and tachinid genera are added to these lists, as well as three large-bodied genera in the Ichneumonidae ( Enicospilus , Pedinopelte , Trogus ).

Taeniogonalos fasciatipennisDHJ02 has been reared 13 times from parasitoids of Crambidae , Hesperiidae , Noctuidae , Notodontidae , Nymphalidae , Riodinidae , Saturniidae , and Sphingidae . The primary host genera are Tachinidae ( Blepharipa , Drino , Houghia , Lespesia , Patelloa ), Ichneumonidae ( Enicospilus ) and Braconidae ( Macrocentrinae ).

Figures

Figure 2. A neighbor-joining tree (NJ) built using Kimura 2 Parameter distance and including 201 sequenced trigonalid specimens from the ACG and North America that have COI sequence greater than 200 bp. Note the divergence between the ACG Taeniogonalos and the North American specimens - and within the dry forest Taeniogonalos fasciatipennisDHJ 01 and Taeniogonalos fasciatipennisDHJ 02 - all are clearly differentiated by mitochondrial DNA.

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 11 - 14. Taeniogonalos fasciatipennis DHJ 01. 11 Metasoma, lateral view, female 12 Female armature, sternum 2, ventral view 13 Male paramere, lateral view 14 Male genitalia.