Journal of Hymenoptera Research 35: 17–31, doi: 10.3897/JHR.35.5496
Food plants and life histories of sawflies of the families Argidae and Tenthredinidae (Hymenoptera) in Costa Rica, a supplement
David R. Smith 1,†, Daniel H. Janzen 2,‡, Winnie Hallwachs 2,§
1 Systematic Entomology Laboratory, Agricultural Research Service, U. S. Department of Agriculture, c/o National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, MRC 168, Washington, DC 20013-7012
2 Department of Biology, University of Pennsylvania, Philadelphia, PA 19104

Corresponding author: David R. Smith (

Academic editor: Stefan Schmidt

received 9 May 2013 | accepted 3 July 2013 | Published 25 October 2013

(C) 2013 David R. Smith. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

For reference, use of the paginated PDF or printed version of this article is recommended.

Citation: Smith DR, Janzen DH, Hallwachs W (2013) Food plants and life histories of sawflies of the families Argidae and Tenthredinidae (Hymenoptera) in Costa Rica, a supplement. Journal of Hymenoptera Research 35: 17–31. doi: 10.3897/JHR.35.5496


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.


Symphyta, Area de Conservación Guanacaste, ACG, fern, tropical dry forest, tropical rain forest


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 and in Janzen (2000, 2001), Burns and Janzen (2001), and Janzen and Hallwachs (2011). Most adult specimens have been DNA barcoded (see acknowledgments).

The format follows Smith and Janzen (2003a, 2003b).


Female (Figs 1–3). Length, 4.0 mm. Head black, labrum and mandible whitish. Antenna black with first and second antennomeres dark orange. Thorax black with pronotum, tegula, mesoprescutum, and mesonotal lateral lobes red; mesosternum dark orange. Legs mostly white; tarsi black, tibiae with apex black and spot of black near base, black more distinct on hind tibia. Abdomen black. Wings darkly infuscated, slightly lighter toward apex; veins and stigma black.

Figures 1–3.

Atomacera josefernandezi. 1 Lateral 2 Dorsum of head and thorax 3 Face, front.

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.

Male. Unknown.

Type material.

Holotype female, labeled “Voucher: D. H. Janzen & W. Hallwachs, DB:, Area de Conservación Guanacaste, Costa Rica, 11-SRNP-20098, ” “legs away for DNA” (USNM). Paratypes: Same data except for voucher numbers, 10-SRNP-22258 (♀), 10-SRNP-22259 (1 ♀), 10- SRNP-22260 (1 ♀), 10- SRNP-22263 (1 ♀), 10- SRNP-22264 (1 ♀), 10- SRNP-22265 (1 ♀), 10- SRNP-22266 (1 ♀), 10- SRNP-22269 (1 ♀), 10- SRNP-22272 (1 ♀), 10- SRNP-22273 (1 ♀), 10- SRNP-22274 (1 ♀), 11- SRNP-20096 (1 ♀), 11- SRNP-20099 (1 ♀), 11- SRNP-20104 (1 ♀), 11- SRNP-20105 (1 ♀), 11- SRNP-20109 (1 ♀), 11- SRNP-20110 (1 ♀) (USNM, INBio).


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.

Food plant and biology.

All specimens of Atomacera 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 Atomacera 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 Hampea 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.


The combination of the following characters will distinguish Atomacera 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 Atomacera josefernandezi by the presence of a short interantennal 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 Atomacera nama Smith, but Atomacera 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), Atomacera 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 Atomacera debilis Say feeds on Desmodium sp. (Fabaceae), and Atomacera decepta Rohwer feeds on Hibiscus sp. (Malvaceae) (Smith 1969).

Eriglenum tristum Smith

Figs 16, 18
Eriglenum tristum Smith, 1992: 53.

This species was described from a single female from Cartago, Turrialba, Costa Rica. Another species, Eriglenum 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.


Known only from Costa Rica.

Food plant and biology.

Nine specimens from two broods were reared, with voucher codes 03-SRNP-11949, 03-SRNP-11951, 03-SRNP-11952, 03-SRNP-11953, 08-SRNP-70547, 08-SRNP-70548, 08-SRNP-70549 (Fig. 16), 08-SRNP-70550, and 08-SRNP-70551. They were reared from larvae feeding on leaves of Machaerium seemannii Benth. Ex Seem. (Fabaceae). The tough, oval cocoon was formed on a leaf (Fig. 18).

Ptenos leucopodus (Cameron)

Sericocera leucopoda Cameron, 1883: 48, pl. 3, fig. 3.
Hemidianeura leucopoda: Smith 1992: 120.
Ptenos leucopodus: Taeger et al. 2010: 169.

Specimens in USNM from Venezuela bear a label “caterpillar on Inya sp.” (Smith 1992). This is probably a misspelling for Inga. Ptenos occurs from southwestern United States and Dominica to Argentina; about 31 species are known, which were keyed by Smith (1992) under the name Hemidianeura. Host plants are not known for the other species.


Costa Rica, Ecuador, Guyana, Honduras, Mexico (Chiapas, San Luis Potosi, Tabasco, Veracruz), Panama, Surinam, Venezuela (Smith 1992).

Food plant and biology.

Nine ACG specimens were reared, with voucher codes 99-SRNP-17561, 99-SRNP-17562, 01-SRNP-22641, 06-SRNP-31799, 06-SRNP-31799.1, 06-SRNP-31799.2, 06-SRNP-31799.3, 06-SRNP-317799.4, and 06-SRNP-31799.5. Larvae were feeding on foliage of Inga vera Willd. (ACG dry forest) and Inga oerstediana Benth (Fabaceae) (ACG rain forest).

Ptilia peleterii (Gray)

Schizocerus Peleterii Gray, 1831 in Gray 1831–1832: 403, pl. 66, fig. 1.
Ptilia peletieri: Smith 1992: 137.

This species was reared from Cnestidium rufescens Planch. (Connaraceae) in Panama, and the larva was described and biological notes given by Kimsey and Smith (1985). Ptilia occurs from Mexico to Brazil; seven species are known (Smith 1992).


Brazil (Amazonas, Mato Grosso); Colombia; Costa Rica; Guyana; Panama; Surinam; Trinidad and Tobago (both islands); Venezuela (Smith 1992).

Food plant and biology.

One ACG specimen was reared (03-SRNP-20546), feeding on very young foliage of the rain forest woody vine Cnestidium rufescens, confirming the report by Kimsey and Smith (1985). A related species, Ptilia versicolor (Klug), feeds on Rourea glabra Kunth (Connaraceae) in ACG (Smith and Janzen 2003a).

Scobina lepida (Klug)

Fig. 19
Hylotoma lepida Klug, 1834: 239.
Scobina lepida: Smith 1992: 26.

This is one of the more common species of Scobina in Central America, but the host plant was not known. Scobina includes about 50 species from Mexico to Argentina, and they were keyed by Smith (1992). Host information was known for only three species, Scobina guatemalensis (Dalla Torre), Scobina consobrina (Norton), and Scobina notaticollis (Konow), all of which fed on the foliage of Sida (Malvaceae).


Costa Rica, El Salvador, Guatemala, Honduras, Mexico (Chiapas, Oaxaca, Veracruz), Nicaragua, Panama (Smith 1992).

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).

Scobina notaticollis (Konow)

Fig. 20
Labidarge notaticollis Konow, 1899: 310.
Scobina notaticollis: Smith 1992: 29.

Costa Rica is the northernmost record for this species.


Bolivia; Colombia, Costa Rica, Ecuador, Panama, Peru, Venezuela (Smith 1992).

Food plant and biology.

One larva (Fig. 20) was found feeding on foliage of ACG rain forest Sida rhombifolia (06-SRNP-44107).

Dochmioglene crassa (Cameron), comb. n.

Figs 4–6
Selandria crassa Cameron, 1883: 19, pl. 1, fig. 5.

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 Dochmioglene crassa except for the lack of a black mark on the mesoprescutum. This mark, however, is faint to absent in some other specimens examined.

Figures 4–6.

Dochmioglene crassa. 4 Lateral 5 Dorsum of head and thorax 6 Face, front.

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.


Costa Rica, Guatemala, Panama, Surinam (Smith, unpublished).

Food plant and biology.

One specimen, 09-SRNP-32388, was found feeding on the foliage of an ACG rain forest fern, Lomariopsis vestita E. Fourn. (Lomariopsidaceae).

Dochmioglene Smith03

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 Dochmioglene crassa or another species. Since the host plant is different than that of Dochmioglene 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.


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)

Monophadnus laetus Cameron, 1883: 22.

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: Waldheimia laeta and Waldheimia lucianocapellii are treated here. Waldheimia suturalis (Cameron) feeds on very young leaves of Cissus rhombifolia Vahl, Waldheimia fascipennis (Norton) feeds on very young leaves of Cissus pseudosicyoides Croat (Vitaceae), and Waldheimia interstitialis (Cameron) feeds on leaves of Hamelia patens Jacq. (Rubiaceae) (Smith and Janzen 2003b).


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 Waldheimia suturalis and Waldheimia fascipennis (Smith and Janzen 2003b). This sawfly did not spin a cocoon, but rather pupated naked in the leaf litter.


Female (Figs 7–9). Length, 7.0 mm. Antenna black with antennomeres 1 and 2 and basal 2/3 of 3rd yellow. Head yellow with area just in front of anterior ocellus to posterior margin of head and hind orbits above eyes black; apex of mandible reddish. Thorax yellow with triangular black spot on mesoprescutum. Abdomen yellow with apical segments and sheath black. Legs yellow, apex of foretarsus and apex of midtarsus infuscated; apical 2/3 of hind tibia, apical 2/3 of hind basitarsomere, and apical 4 hind tarsomeres black. Wings fasciated, area apical to apex of stigma and area basal to intercostal veins darkly infuscated, center yellow; extreme base of costa yellow; stigma yellow; veins in yellow area, yellow; veins in infuscated area, black.

Figures 7–11.

Waldheimia lucianocapellii. 7 Lateral 8 Dorsal 9 Face, front 10 Male genital capsule, ventral 11 Male penis valve, lateral.

Antenna 2.0× head width, apical 4 segments longer than broad and shorter than segments 4 and 5 combined. Lower interocular distance 0.8× eye length; distances between eye and hind ocellus, between hind ocelli, and between hind ocellus and posterior margin of head as 1.0:1.0:1.0. Malar space absent. Hind basitarsomere longer than following tarsomeres combined. Sheath rounded at apex in lateral view. Lancet (Fig. 14) with serrulae flat, each with 7-9 fine subbasal teeth; annular hairs dense, short.

Male. 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.

Type material

. Holotype female labeled, “Barro Colorado, CZ, Pan., R.C. Shannon, VII-19-23” (USNM). Paratypes: Panama, Canal Zone, Barro Colorado Is., 9°9'N, 79°51'W, 8–15 Feb. 1995, J. Pickering (1 ♀); same data except 3 May, 1995 (1 ♀), 19–26 Jan. 1994 (1 ♀); Panama, Canal Zone, Barrow Colorado Is., 9 May 1978, in Malaise trap, 20 m up in Tachigaua, Rainer Krell (1 ♂) (USNM).

Other specimen.

Costa Rica, Area de Conservación Guanacaste, lat11.01926 long -85.40997, 14 Oct 2005, 05-SRNP-33935 (1 ♀, USNM) (


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.

Food plant and biology.

One specimen, 05-SRNP-33935), was reared from Davilla nitida (Vahl) Kubitzki (Dilleniaceae). The larva (Figs 21, 22) is very similar to that of Waldheimia interstitialis, illustrated by Smith and Janzen (2003b: fig.3).


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 Waldheimia 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 3rd, 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 Waldheimia fascipennis, which also has the wings broadly black at the base and apex and yellow at the center; however, Waldheimia fascipennis has an entirely orange head, entirely white hind basitarsomere, and deeper serrulae on the lancet (Fig. 15).

Figures 12–15.

Lancets. 12 Atomacera josefernandezi 13 Atomacera ebena 14 Waldheimia lucianocapellii 15 Waldheimia fascipennis.

Figures 16–22.

16 Adult of Eriglenum tristum 17 Cocoon of Atomacera josefernandezi on leaf 18 Cocoon of Eriglenum tristum on leaf 19 cocoon of Scobina lepida 20 Larva of Scobina notaticollis 21, 22 Larvae of Waldheimia lucianocapellii.


Michele Touchet, Systematic Entomology Laboratory, USDA, Washington, DC, helped with Figures 115. We gratefully acknowledge the team of ACG parataxonomists (Janzen et al. 2009) who found and reared the specimens used in this study, and the team of biodiversity managers who keep alive the ACG forests that host these sawflies. The study has been supported by U.S. National Science Foundation grants BSR 9024770 and DEB 9306296, 9400829, 9705072, 0072730, 0515699, and grants from the Wege Foundation, International Conservation Fund of Canada, Jessie B. Cox Charitable Trust, Blue Moon Fund, Guanacaste Dry Forest Conservation Fund, Area de Conservación Guanacaste, and University of Pennsylvania (DHJ). Laboratory analyses of the DNA barcode sequences were funded by the Government of Canada through Genome Canada and the Ontario Genomics Institute (2008-OGI-ICI-03), thanks to P. D.N. Hebert, M. Hajobabae, and S. Naik. USDA is an equal opportunity provider and employer.

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