Food plants and life histories of sawflies of the families Argidae and Tenthredinidae ( Hymenoptera ) in Costa Rica , a supplement

Food plants and information on life history are presented for six species of Argidae and four species of Tenthredinidae in Costa Rica. The Argidae include cocoons of Atomacera josefernandezi Smith, sp. n., found on Hampea appendiculata (Donn. Sm.) Standl. (Malvaceae) and likely feeding on its leaves before pupation, and larvae of Eriglenum tristum Smith feeding on Machaerium seemanii Benth. Ex Seem. (Fabaceae), Ptenos leucopodus (Cameron) feeding on Inga oerstediana Benth. and I. vera Willd. (Fabaceae), Ptilia peleterii (Gray) feeding on Cnestidium rufescens (Connaraceae), and Scobina lepida (Klug) and S. notaticollis (Konow) feeding on Sida rhombifolia L. (Malvaceae). The Tenthredinidae include larvae of Dochmioglene crassa (Cameron) feeding on the fern Lomariopsis vestita E. Fourn. (Lomariopsidaceeae), Dochmiogleme Smith03 feeding on Blechnum occidentale L. (Blechnaceae), Waldheimia laeta (Cameron) feeding on Cissus alata Jacq. (Vitaceae), and Waldheimia lucianocapellii Smith, sp. n., feeding on Davilla nitida (Vahl) Kubitzki (Dilleniaceae). Waldheimia lucianocapellii is described from specimens from both Panama and Costa Rica. Selandria crassa Cameron, 1883 is a comb. n. in Dochmioglene.


Introduction
Two previous papers presented food plants and life history notes on twelve species of Argidae (Smith and Janzen 2003a) and five species of Tenthredinidae and six species of Pergidae (Smith and Janzen 2003b).Since, several other species have been reared during the ongoing caterpillar inventory of Area de Conservación Guanacaste (ACG) in Guanacaste Province in northwestern Costa Rica.Here, we report on six additional species of Argidae and four additional species of Tenthredinidae.

Materials and methods
Acronyms used are: USNM, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA; INBio, Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica; BMNH, The Natural History Museum, London, U.K.
Images 1-15 were obtained using an EntoVision Imaging Suite that included a firewire JVC KY-75 3CCD digital camera mounted to a Leica M16 zoom lens via a Leica z-step microscope stand.Multiple focal planes were merged using Cartograph 5.6.0 (Microvision Instruments, France) software.Images 16-22 are from the ACG website, cited below.
Voucher codes associated with each reared adult are expressed as, for example, "99-SRNP-4547" (year-Santa Rosa National Park-unique rearing number for that year).This code is unique for the event of finding and rearing the caterpillar, irrespective of what it produces.Full details of the voucher records and associated images may be obtained at http://janzen.sas.upenn.eduand in Janzen (2000Janzen ( , 2001)), Burns and Janzen (2001), and Janzen and Hallwachs (2011).Most adult specimens have been DNA barcoded (see acknowledgments).
The format follows Smith andJanzen (2003a, 2003b).Head smooth and shining, without punctures or other sculpture.Antennal length 1.3× head width.Lower interocular distance about 1.2× eye height.Distances between eye and hind ocellus, between hind ocelli, and between hind ocellus and posterior margin of head as 1.0:1.2:0.5.Clypeus with shallow central emargination.Interantennal area rounded, without carina.Malar space about 1.3× diameter of front ocellus.Postocellar area very short, almost non-existent, sloping downward just behind lateral ocelli; without lateral postocellar grooves.Forewing with 4 cubital cells, first cubital crossvein may be weak.Hind basitarsomere 0.9× length of remaining tarsomeres combined.Sheath uniformly slender in dorsal view, straight above and rounded below in lateral view.Lancet (Fig. 12) with serrulae moderately deep, with 1 or 2 anterior and 4-5 posterior subbasal teeth; annuli slightly curved in basal half, straighter in apical half; short hairs on annuli.Etymology.This species is named in honor of Jose Fernandez-Triana of Ottawa, Canada, in recognition of his outstanding efforts to describe and otherwise clarify the taxonomy of the many hundreds of species of microgastrine braconid wasps being reared by the same inventory that discovered this new sawfly.

Atomacera josefernandezi
Food plant and biology.All specimens of A. josefernandezi were found as tiny spun cocoons on the leaves of Hampea appendiculata (Donn.Sm.) Standl.(Malvaceae).While the cocoons of all other species of sawflies reared by the ACG inventory have been ovoid in shape, the cocoons of A. josefernandezi are conspicuously squared off at both ends, making them look like small bricks (Fig. 16).The larvae were not seen, and this may not be the food plant.However, we suspect that it is the food plant because the many cocoons were all on one individual of H. appendiculata and not scattered over other individual plants, and were accompanied by large areas of strongly skeletonized leaves.There were two sets, apparently broods, of wild-caught cocoons (2010 and 2011) in the same area on the same species of plant, again suggesting that this really is the food plant.
Remarks.The combination of the following characters will distinguish A. josefernandezi from other species of Atomacera: tegula, pronotum, and mesonotum (except mesoscutellum) red; mesosternum dark orange; tibiae mostly white with some black at base and apex; interantennal area rounded, without a carina; clypeus, supraclypeal area, and areas surrounding antennae smooth, shiny, without sculpture; very short postocellar area, lacking lateral furrows; and lancet (Fig. 12) with distinct serrulae and short hairs on the annuli.With the red pronotum, tegula, and most of the mesonotum and mostly white tibiae, this species will run to couplet 10, Atomacera ebena Smith, in the key to Atomacera (Smith 1992).Atomacera ebena is separated from A. josefernandezi by the presence of a short in-terantennal carina, the clypeus, supraclypeal area, and interantennal area punctate, the postocellar area defined by lateral furrows, a red mesoscutellum, most of the hind tarsi white, and the lancet with very low serrulae and lacking annular hairs (Fig. 13).This species is also similar to A. nama Smith, but A. nama has the legs black, metascutellum and metascutellum orange, and flatter serrulae on the lancet (Smith 1992: fig. 113).
Atomacera josefernandezi DNA barcodes (Janzen and Hallwachs 2011) are very distinctively different from all other species of Argidae and Tenthredinidae reared to date in ACG.However, it is noteworthy that the 2010 rearing consistently differed by what appears to be 1 basepair from the 2011 rearing, a very shallow split that needs to be analyzed with a larger sample size.All the specimens found within a year, to date, are likely to be sibs, and therefore in one sense we have DNA barcoded only two specimens (= two broods).
The New World genus Atomacera includes about 32 species, 30 of which are Neotropical and were keyed by Smith (1992).It occurs from Canada to Argentina.Only five species apart from the one recorded here have been associated with food plants.Atomacera pubicornis (Fabricius) from northern South America feeds on Ipomoea sp.(Convolvulaceae) (Smith 1992), A. petroa Smith has been reared from Miconia calvescens DC (Melastomataceae) in Costa Rica (Badenes-Perez and Johnson 2007), and Atomacera raza (99-SRNP-4547), also reared by the ACG inventory (Smith and Janzen 2003a), was feeding on leaves of Malvaviscus palmanus (Malvaceae).The Nearctic species A. debilis Say feeds on Desmodium sp.(Fabaceae), and A. decepta Rohwer feeds on Hibiscus sp.(Malvaceae) (Smith 1969).Smith, 1992: 53.Remarks.This species was described from a single female from Cartago, Turrialba, Costa Rica.Another species, E. crudum Konow, was reared from Maechaerium acuminatum Kunth (Fabaceae) in ACG (Smith and Janzen 2003a).Eriglenum occurs from Mexico to northern Argentina; see Smith (1992) for separation of the four species.
Food plant and biology.One larva (07-SRNP-40019) was found eating mature leaves of rain forest Sida rhombifolia L. (Malvaceae), a common pasture and roadside woody herb.The fibrous, oval cocoon was on a leaf (Fig. 19).
Food plant and biology.One larva (Fig. 20) was found feeding on foliage of ACG rain forest Sida rhombifolia (06-SRNP-44107) .Remarks.Dochmioglene includes about 11 species distributed from Central America to Argentina (Smith, unpublished).The genus has not been revised.The reared specimen (Figs 4 -6) agrees with D. crassa except for the lack of a black mark on the mesoprescutum.This mark, however, is faint to absent in some other specimens examined.

Tenthredinidae
Dochmioglene belongs in the subfamily Selandriinae, which includes many species associated with ferns.This rearing, and that of the following species, confirms that larvae of many Neotropical selandriine species may be feeding on fern foliage.Cameron (1883) described the female from "Guatemala, San Isidro 1600 feet" and mentioned that a male from El Tumbador may be the male of the species.Because Cameron did not designate a holotype, apparently included a male, and did not state how many specimens were available, a lectotype is here designated.The female of Selandria crassa in in BMNH, labeled "Type H.T.," "B.M. type :Hym.1.266," "B.C.A. Hymen.I, Selandria crassa Cam.," "San Isidro, 1600 ft., Champion" and "Selandria crassa Cam., type, BCA Hy 1, 10" is designated the lectotype.The combination in Dochmioglene has not been published and is based on study of the type by DRS.

Dochmioglene Smith03
Remarks.Only parts of the reared adult are stored in a gelatin capsule.They are sufficient to place it in Dochmioglene, but there is not enough left to tell if this is the same as D. crassa or another species.Since the host plant is different than that of D. crassa and we cannot be sure it is the same, we here consider it as a second reared species of the genus in ACG, with the interim name of Dochmioglene Smith03.
Distribution.Known only from the single specimen from ACG.Food plant and biology.One specimen, 09-SRNP-21528, was found feeding on the foliage of a rain forest fern, Blechnum occidentale L. (Blechnaceae).

Waldheimia laeta (Cameron)
http://species-id.net/wiki/Waldheimia_laeta Monophadnus laetus Cameron, 1883: 22. Remarks.Waldheimia is a large genus of nearly 100 species and occurs from southwestern United States to Argentina.Waldheimia laeta is separated by its black head, wings yellow with apices black, abdomen orange with apex black, and the very short, triangular lancet with indiscernible serrulae.
The host plants of five species of Waldheimia in Costa Rica are now known: W. laeta and W. lucianocapellii are treated here.Waldheimia suturalis (Cameron) feeds on very young leaves of Cissus rhombifolia Vahl, W. fascipennis (Norton) feeds on very young leaves of Cissus pseudosicyoides Croat (Vitaceae), and W. interstitialis (Cameron) feeds on leaves of Hamelia patens Jacq.(Rubiaceae) (Smith and Janzen 2003b).
Distribution.Bolivia, Colombia, Costa Rica, Panama, Peru, Trinidad, Venezuela (Smith, unpublished) Food plant and biology.One specimen, 09-SRNP-6456, found feeding on very young leaves of ACG dry forest Cissus alata Jacq.(Vitaceae), the same host genus as W. suturalis and W. fascipennis (Smith and Janzen 2003b).This sawfly did not spin a cocoon, but rather pupated naked in the leaf litter.
Male Etymology.This species in named in honor of Luciano Capelli of San Jose, Costa Rica, for his intense and generous efforts to publicize ACG conservation with outstanding video and photographs for the Costa Rican and international public.
Remarks.The specimen reared from Costa Rica is not complete and is not included in the type series.However, enough is present, including the lancet, to show it  is the same as the specimens from Panama.It DNA barcodes as far from the other 4 species of Waldheimia that were DNA barcoded, as they do from each other.
This species is similar to Waldheimia duckei Konow of South America, but W. lucianocapellii is separated by the mostly black mesoprescutum, white base of the hind tibia and base of the hind basitarsomere, yellow antennomeres 1 and 2 and base of 3 rd , head pale from above antennae to clypeus, and low flat serrulae of the lancet (Fig. 14).The only other species with which it might be confused in Costa Rica is W. fascipennis, which also has the wings broadly black at the base and apex and yellow at the center; however, W. fascipennis has an entirely orange head, entirely white hind basitarsomere, and deeper serrulae on the lancet (Fig. 15).
. Length, 7.0 mm.Similar in color to female.Male genitalia in Figs 10, 11; harpe tapering to rounded apex; parapenis rounded on inner margin, almost straight on outer margin; valviceps of penis valve with long, rounded dorsal lobe.