Sawflies (Hymenoptera: Argidae, Pergidae, Tenthredinidae) from southern Ecuador, with a new record for the country and some ecological data

Jean-Luc Boevé; Diego S. Marín-Armijos; Diego F. Domínguez; David R. Smith

doi:10.3897/jhr.51.9830

Research Article

Sawflies (Hymenoptera: Argidae, Pergidae, Tenthredinidae) from southern Ecuador, with a new record for the country and some ecological data

Jean-Luc Boevé^‡, Diego S. Marín-Armijos^§, Diego F. Domínguez^§, David R. Smith^|

‡ Royal Belgian Institute of Natural Sciences, Bruxelles, Belgium

§ Universidad Técnica Particular de Loja, Loja, Ecuador

| Smithsonian Institution, Washington, DC, United States of America

Corresponding author: Jean-Luc Boevé ( jean-luc.boeve@naturalsciences.be )

Academic editor: Marko Prous

This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Citation: Boevé J-L, Marín-Armijos DS, Domínguez DF, Smith DR (2016) Sawflies (Hymenoptera: Argidae, Pergidae, Tenthredinidae) from southern Ecuador, with a new record for the country and some ecological data. Journal of Hymenoptera Research 51: 55-89. https://doi.org/10.3897/jhr.51.9830

ZooBank: urn:lsid:zoobank.org:pub:062BE137-7933-4E23-AA76-061FB0949E9F

Abstract

An illustrated list of species of sawflies collected during 2014 and 2015 in southern Ecuador is given. One genus and species, Pristiphora fernandezi Smith (Tenthredinidae), is new for the country. Ecological data are provided for Lagideus podocarpus Smith, sp. n. (Pergidae) that was observed feeding on Fuchsia vulcanica L. (Onagraceae). Three new combinations are proposed: Bolivius notabilis (Konow, 1899), Plaumanniana biclinea (Konow, 1899), and Proselandria alvina (Konow, 1899) (Tenthredinidae: Selandriinae).

Keywords

Species list, páramo, host plant, Fuchsia , Lagideus , larva, defence behaviour

Introduction

Most sawflies described from South America belong to the Tenthredinidae (Smith 2003a, 2003b), Argidae (Smith 1992), and Pergidae (Smith 1990). Worldwide, the most species-rich family is the Tenthredinidae with over 5700 species (Taeger et al. 2010). The Argidae include ca. 900 species distributed throughout the world but with a greatest number and diversity occurring in the Neotropical Region (Smith 1992). They are mainly associated with angiosperms. The Pergidae include ca. 440 species, occurring in the Australasian, Nearctic, and Neotropical regions (Smith 1990; Schmidt and Smith 2006). They are the most diverse in Australia and the Neotropics.

Related to its size, Ecuador is among the countries with highest biodiversity, which is mainly due to its tropical location as well as the presence of the Andes, Amazon Basin and two major ocean streams. These combined factors lead to a high variety of distinct environments. One typical biotope for tropical highlands is called the ‘páramo’ which occurs in southern Ecuador.

Here, we list the sawflies which were collected mainly in the Podocarpus National Park (NP) that is characterized by the páramo. The collections were performed in the frame of a Global Taxonomy Initiative (GTI) project.

Methods

Sawflies were collected in southern Ecuador, mainly around Loja (e.g., Fig. 1) and during October 2014. Most sawflies were collected as adults using a net, a few being collected in yellow pan traps, while larvae were found only at one location. The specimens were stored in ethanol, but some of the larvae were reared to obtain the adults.

Figure 1.

Examples of biotopes in southern Ecuador and where sawflies were collected. a El Tiro (altitude 2815 m) b Cajanuma “Mirador” (3000 m) c Colibrí Station (2170 m) d Yangana (ca. 2500 m).

Most pictures were taken by J.-L. Boevé with the following cameras: Pentax Optio W10, Nikon Coolpix P300, and Nikon D7000. Since adult specimens are kept in ethanol, they were slightly dried to take the pictures, which were mainly intended to illustrate the habitus. Pictures were also taken by D. R. Smith (Fig. 10) and Thibaut Delsinne (Fig. 21b, e, f, g).

The sawfly specimens collected as part of the GTI project are stored in the Royal Belgian Institute of Natural Sciences, Brussels, Belgium (RBINS; J.-L. Boevé collection, with specimen reference codes starting with ‘P’), with duplicates located in the National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA (USNM), and Museo de Colecciones Biológicas, Universidad Técnica Particular de Loja, Ecuador (MUTPL). Sawfly specimens from the RBINS, Senckenberg Deutsche Entomologisches Institut, Müncheberg, Germany (SDEI), and USNM were examined. Dates are given by dd.mm.year.

Taxonomic changes presented here are by D.R. Smith. Identifications of Argidae, Pergidae, and Tenthredinidae (Nematinae) are based on Smith (1990, 1992, 2003b). Identifications of Blennocampinae and Selandriinae (Tenthredinidae) are based on Smith (2007) and unpublished manuscripts by D.R. Smith.

Results

Family Argidae

Scobina bolivari (Konow, 1899)

Fig. 2

Distribution

This species is known from Colombia, Ecuador, Peru, Bolivia, and northern Argentina (Jujuy) (Smith 1992).

Material

Los Geraneos, 04°09'S, 078°38'W, 855m, 18.10.2014, P3944.B (1 ♀), leg. A. Pauly, J.-L. Boevé.

Figure 2.

Scobina bolivari, female (P3944.B), body length 8.0 mm. a Dorsal view b ventral view.

Scobina inculta (Konow, 1906)

Fig. 3

Distribution

Known from Venezuela, Ecuador, Peru, Bolivia, and northern Argentina (Jujuy) (Smith 1992).

Material

Bombuscaro, 04°05'S, 078°57'W, 930m, 22.03.2015, P4120.A (1 ♀), leg. T. Delsinne.

Figure 3.

Scobina inculta, female (P4120.A), body length 9.5 mm. a Dorsal view b ventral view.

Scobina notaticollis (Konow, 1899)

Fig. 4

Distribution

Known from Costa Rica, Panama, Venezuela, Colombia, Ecuador, Peru, and Bolivia (Smith 1992).

Material

Vilcabamba, 04°16'S, 079°13'W, 1570m, 11.10.2014, P3891, P3892 (2 ♀), leg. A. Pauly, J.-L. Boevé.

Figure 4.

Scobina notaticollis, female (P3891), body length 8.5 mm. a Dorsal view b ventral view.

Scobina spp.

Fig. 5

Material

S. of Yangana, 04°22'S, 079°10'W, 1940m, 09.10.2014, flying, P3890 (1 ♀), leg. A. Pauly, J.-L. Boevé; nr Loja, Reserva El Madrigal, 04°02'S, 079°11'W, 2275m, 21.10.2014, on Rumex crispus (Polygonaceae), P3953.B (1 ♂), leg. J.-L. Boevé.

Figure 5.

Scobina sp. a, b Female (P3890), body length 8.0 mm c, d male (P3953.B), body length 7.5 mm a, c Dorsal views b, d ventral views.

Sericoceros gibbus (Klug, 1834)

Fig. 6

Distribution

A widespread species known from Mexico south to Guyana, Brazil, and Peru.

Material

Bombuscaro, Podocarpus NP, 04°06'S, 078°58'W, 995m, 17.10.2014, flying while crossing footpath, P3937 (1 ♀), leg. A. Pauly, J.-L. Boevé.

Remarks

Larvae have been recorded from Coccoloba manzinellensis Beurl. and C. caracasana Meisn. in Panama, and C. guanacastensis W. C. Biurger (Polygonaceae) in Costa Rica. Notes on the biology were given by Smith and Janzen (2003).

Figure 6.

Sericoceros gibbus, female (P3937), body length 9.5 mm. a Dorsal view b ventral view.

Family Pergidae

Acordulecera spp.

Remarks

Probably four species were found in this study. It is a large genus of perhaps 200 or more species that occurs from Canada to Chile. Smith (1990) listed 45 described species from the Neotropics.

Material

Valle de Quinara, 04°18'S, 079°16'W, 1550m, 09.10.2014, P3888.A, P3888.B (2 ♂), leg. A. Pauly, J.-L. Boevé; Station Colibrí, Podocarpus NP, 03°59'S, 079°06'W, 2170m, 13.10.2014, by sweeping, P3902 (1 ♂), leg. A. Pauly, J.-L. Boevé; Bombuscaro, Podocarpus NP, 04°07'S, 078°58'W, 995m, 16.10.2014, P3934 (1 ♀), leg. A. Pauly, J.-L. Boevé; nr Loja, Reserva El Madrigal, 04°02'S, 079°11'W, 2275m, 21.10.2014, near rivulet, P3956.G (1 ♀), P3956.H, P3956.I, P3956.J (3 ♂), leg. T. Delsinne, J.-L. Boevé; Cajanuma, Podocarpus NP, 04°07'S, 079°10'W, 2810m, 23.10.2014, on leaf, P3971 (1 ♀), leg. T. Delsinne, J.-L. Boevé.

Decameria varipes Cameron, 1883

Fig. 7

Distribution

This species occurs from Guatemala south to Ecuador (Smith 1990).

Material

nr Loja, Reserva El Madrigal, 04°02'S, 079°11'W, 2275m, 21.10.2014, near rivulet, P3956.A, P3956.B, P3956.C, P3956.D, P3956.E, P3956.F (6 ♂), leg. T. Delsinne, J.-L. Boevé, on leaf of Rubus sp. (Rosaceae), P3950 (1 ♂), leg. J.-L. Boevé.

Remarks

The thorax is typically partly red in most specimens; however melanic forms are occasionally found. The specimens collected here are entirely black but agree in all structural features with the red form. Colour variation is common in some Perreyiinae.

Figure 7.

Decameria varipes, male (P3956.B), body length 4.0 mm. a Dorsal view b ventral view.

Decameria sp.

Fig. 8

Material

Loja, Reinaldo Espinosa Botanical Garden, 04°02'S, 079°12'W, 2145m, 14.10.2014, P3915 (1 ♂), leg. J.-L. Boevé.

Figure 8.

Decameria sp., male (P3915), body length 5.5 mm. a Dorsal view b ventral view.

Lagideus podocarpus Smith, sp. n.

http://zoobank.org/A9FD9A79-348D-42A2-8176-AAF8A2C0DA07

Figs 9, 10, 11

Description

Female. Unknown.

Male. Length 7.0 mm. Antenna and head black; labrum and mouthparts white; mandible dark red brown. Thorax black with posterior corners of pronotum orange. Fore and midlegs light orange; fore coxa black with white stripe on outer surface and extreme apex white; mid coxa black with extreme apex white. Hind legs black with extreme apex of coxa, trochanters, and extreme base of femur white and about basal half of tibia light orange. Wings hyaline; veins and stigma black.

Head with vertex and frons shiny, very few widely scattered punctures; area adjacent to inner orbits slightly dull, micropunctured; with long golden hairs, as long as or longer than second antennomere. Antenna 1.5× head width; antennomeres 4-8 bipectinate, antennomere 3 unipectinate with inner ramus absent to very slightly indicated (Smith 2012: fig. 25); apical antennomere with rami less than half length of stem. Malar space linear. Thorax shiny; mesonotum with distinct punctures, separated by flat shiny interspaces more than puncture diameters; with long golden hair on dorsum and grey to white hair on pleurae and sterna. Hindbasitarsomere slightly longer than length of remaining tarsomeres combined; inner hind tibial spur about 0.6× length of basitarsomere. Genitalia in Fig. 10; parapenis uniformly broad, not distinctly indented on posterior margin; valviceps of penis valve with dorsal margin concave.

Figure 9.

Lagideus podocarpus, male (P3898.etc>13), body length 6.0 mm. a Dorsal view b ventral view.

Figure 10.

Male genitalia of Lagideus podocarpus. At left, genital capsule, ventral view; at right, penis valve, lateral view with dorsal margin to right.

Type material

Holotype labelled “Ad.M, ex-larva,” “ECUADOR: Cajanuma, Podocarpus NP, 04°07'S, 079°10'W, 2810 m, X.2014, on Fuchsia vulcanica, JLBoevé leg. RBINS, EtOH100%, P3898.etc>10” (RBINS). Paratypes: Same data as for holotype except P3898.etc>01 (1 ♂), P3898.etc>02 (1 ♂), P3898.etc>03 (1 ♂), P3898.etc>04 (1 ♂), P3898.etc>05 (1 ♂), P3898.etc>06 (1 ♂), P3898.etc>07 (1 ♂), P3898.etc>08 (1 ♂), P3898.etc>09 (1 ♂), P3898.etc>11 (1 ♂), P3898.etc>12 (1 ♂), P3898.etc>14 (1 ♂); these specimens are in RBINS, USNM and MUTPL.

Other material

Cajanuma, Podocarpus NP, 04°07'S, 079°10'W, 2810m, 11.10.2014, on Fuchsia vulcanica (Onagraceae), P3898 (6 larvae), 14.10.2014, on F. vulcanica, P3917 (4 larvae), P3919 (20–30 larvae), 23.10.2014, on F. vulcanica, P3968 (>30 larvae), P3969 (6 larvae), P3970 (6 larvae), P3976 (4 larvae), leg. J.-L. Boevé.

Etymology

Named for Podocarpus National Park, Ecuador, where the collections were made; a noun in apposition.

Comments

Although only the male is known, it differs from described species, and we prefer to have a name since it was reared and the life history and host plant are known. Only females are known for most species of Lagideus, and this could be the opposite sex of one of those. However, this won’t be known until the sexes are associated.

The bipectinate antennomeres 4-8 and unipectinate antennomere 3 will take this species to couplet 17c in Smith (2012) which includes L. longicus Smith and L. tapanti Smith, both known from Costa Rica. The black antenna and similarity of genitalia will put it closest to L. longicus.

Lagideus tapanti has white hairs on the head and body, antennomeres 1-3 or 4 yellowish, thorax black, and legs black with the base of the femora white; the antenna (Smith 2012: fig. 25) has the rami of the apical antennomere much longer than the stem; and the parapenis of the male genitalia are lobed posteriorly with a central, shallow posterior emargination (Smith 2012: fig. 28). Lagideus longicus is entirely black and the apical antennomere has long rami as in L. tapanti; the parapenis of the genitalia is distinctly indented posteriorly for at least a third of its medial length (Smith 2012: fig. 29); and the valviceps of the penis valve have a straight dorsal margin (Smith 2012: fig. 33).

Lagideus podocarpus can be distinguished from both these species by light orange posterior corners of the pronotum; the light orange fore and mid legs (except the partly black coxae); basal half of the hind tibia light orange; the antennae, similar to L. longicus (Smith 2012: fig. 25) except the rami of the apical antennomere are short, less than half the length of the stem; and the male genitalia (Fig. 10) which has the parapenis nearly uniformly broad, without a central posterior emargination and the valviceps of the penis valve distinctly concave dorsally.

The length varies slightly 6.0–7.5 mm. The tegulae may be black to brown or partly light orange as are posterior corners of the pronotum.

Life history

The population of L. podocarpus at Cajanuma (Fig. 11) consisted of larvae found within a circumference of ca. 100 m, on several patches of the host plant, F. vulcanica L. (Onagraceae). The larvae were attached to the underside of a leaf and they were generally aggregated in that one or a few individuals occurred per leaf. Younger larvae skeletonised the leaves and made holes in the leaf, whereas larvae of later instars attacked the leaf up to its edge. Larvae fed on both younger and older leaves. One large patch of flowering plants clearly harboured fewer larvae than the smaller and more numerous spots of non-flowering plants. The larvae being kept in rearing still appeared as healthy, despite the fact that the offered leaves (during November 2014) were less fresh and rather decaying. A couple of plant species and varieties were also offered to them to roughly establish their diet breadth. Fuchsia ‘Koralle’ was rather accepted, whereas only very few small holes were made on leaves of Fuchsia ‘Gartenmeister Bonstedt’. These two varieties are cultivars of Fuchsia triphylla that is related to F. vulcanica (Marc Reynders, personal communication). A leaf of Oenothera sp. (Onagraceae) was also tested, but not accepted.

Figure 11.

Life history traits of Lagideus podocarpus. a A patch of the host plant Fuchsia vulcanica b a plant with feeding damage, seen from above c–e larvae attached to the underside of leaves, seen from below f a larva that just moulted with exuvia visible g, h prepupal stage i cocoon j screenshot of a video clip, the thorax of the larva was touched with flexible forceps a second before k, l two screenshots of a video clip performed during a bioassay, both being spaced by 1 sec: an ant is approaching the larva (first screenshot), then touching it, which immediately provokes the defensive behaviour of the larva that directs its caudal filaments towards its aggressor (second screenshot).

Larvae of nearly all instars were found in the field, but no adults were collected by sweeping. Among the larvae kept in rearing, several individuals reached the eonymph stage (Fig. 11g–h), during the first ten days of October 2014. They formed a cocoon between fresh or rather decaying leaves (Fig. 11i), or in an offered layer of moistened sand. At the end of October, prepupae and cocoons occurred in the rearing. A total of 14 adults, all males, emerged from 20.11 to 12.12.2014.

Once disturbed, a larva of L. podocarpus reacted immediately by raising its abdomen (Fig. 11j), then slowly lowering it. This reaction could be repeated several times, under successive disturbances. After several such behaviours, the larva tended to slightly walk backwards. The prepupa reacted under disturbance by curling its body and making some slow movements. To further investigate the larval defensive behaviour, a dozen workers of the ant species Eciton burchellii Westwood (Formicidae) were collected in the field (Pueblo Viela, 15.10.2016), held without feeding, and used ca. 36 h later. The bioassay consisted of placing seven ant workers and one larva settled on a host plant leaf in an open box, the interactions being then filmed. As far as no ant contacted the larva, the latter remained merely immobile. But once contacted, the larva instantly raised its abdomen and directed the apical filaments towards the aggressor (Fig. 11k–l). Thus, these filaments seem to play a major role in defence. Their size is impressive, with a length of 13 mm for a remaining body size of 16 mm. The filaments resemble those of the larvae of Philomastix; the two genera are sister-groups (Schmidt and Walter 2014). However, the filaments protrude from the apical segment in Lagideus and the ninth one in Philomastix (Smith and Bado 2004). Moreover, it seems that they are clearly covered with spines in Lagideus only (Fig. 11; Smith and Bado 2004). It is unclear whether the defence is only physical, or includes chemical compounds. Larvae of Pergidae contain toxic peptides which are active against vertebrates and invertebrates (Boevé et al. 2014), but whether Lagideus sp. contains such compounds remains unknown.

Note that another sawfly species (Tenthredinidae) was found among the collected leaves (and stems) of F. vulcanica, but in much lower numbers (2 individuals: 11.10.2014, P3899, and 14.10.2014, P3919.C1) than Lagideus sp. (>90 in total). This plant constitutes a host plant, since a larva of the tenthredinid was placed on an intact leaf and the larva ingested significant parts of it.

Perreyia picea (Westwood, 1874)

Fig. 12

Distribution

This species is known from Colombia, Ecuador, Peru, and Brazil (Smith 1990).

Material

nr Loja, Reserva El Madrigal, 04°02'S, 079°11'W, 2270m, 21.10.2014, P3957.E (1 ♂), leg. A. Pauly, J.-L. Boevé.

Figure 12.

Perreyia picea, male (P3957.E), body length 6.5 mm. a Dorsal view b ventral view.

Family Tenthredinidae

Subfamily Blennocampinae

Metapedias sp.

Fig. 13

Distribution

This Neotropical genus occurs from Mexico to Argentina and includes about 25 species.

Material

Bombuscaro, Podocarpus NP, 04°07'S, 078°58'W, 995m, 16.10.2014, flying just above the ground, P3935 (1 ♀), leg. J.-L. Boevé; N. of Pachicutza, 04°04'S, 078°38'W, 900m, 18.10.2014, P3942.C (1 ♂), leg. A. Pauly, J.-L. Boevé; Loja, Estación Científica San Francisco, 03°58'S, 079°05'W, 1820m, 24.03.2015, P4121.B (1 ♀), leg. T. Delsinne.

Figure 13.

Metapedias sp. a, b Female (P4121.B), body length 6.0 mm c, d male (P3942.C), body length 6.5 mm. a, c Dorsal views b, d ventral views.

Waldheimia atripennis (Fabricius, 1804)

Fig. 14

Distribution

This is a relatively common species occurring from Costa Rica to Brazil and Argentina.

Material

Bombuscaro, road to Podocarpus NP, 04°06'S, 078°58'W, 975m, 08.10.2014, P3883 (1 ♂), leg. A. Pauly, J.-L. Boevé; Bombuscaro, Podocarpus NP, 04°06'S, 078°58'W, 995m, 16.10.2014, P3936.D (1 ♂), leg. A. Pauly, J.-L. Boevé; Los Geraneos, 04°09'S, 078°38'W, 855m, 18.10.2014, P3944.A (1 ♀), leg. A. Pauly, J.-L. Boevé, P3941 (1 ♂), leg. J.-L. Boevé; Miasi, 04°15'S, 078°20'W, 875m, 19.10.2014, P3945 (1 ♂), leg. A. Pauly, J.-L. Boevé, on/around leaf of Anthurium sp. (Araceae), P3946.A, P3946.B, P3946.C (3 ♂), leg. J.-L. Boevé; Romerillos, nr Podocarpus NP, 04°09'S, 078°56'W, 1100m, 22.10.2014, P3967.O (1 ♀), P3967.U, P3967.W (2 ♂), leg. A. Pauly, J.-L. Boevé.

Figure 14.

Waldheimia atripennis, male (P3967.U), body length 7.0 mm. a Dorsal view b ventral view.

Waldheimia erebus (W.F. Kirby, 1882)

Fig. 15

Distribution

This species is known from Guyana, Brazil, Colombia, Ecuador, and Peru.

Material

Bombuscaro, road to Podocarpus NP, 04°06'S, 078°58'W, 955–975m, 08.10.2014, flying around plants of Sticherus sp. (Gleicheniaceae), P3866, P3867, P3868, P3869 (4 ♂), leg. J.-L. Boevé, P3880, P3881, P3882, (3 ♂), leg. A. Pauly, J.-L. Boevé; Pueblo Viela, 04°38'S, 079°08'W, 1060m, 15.10.2014, on leaf of Anthurium sp., P3926.B (1 ♀), P3926.A, P3926.C, P3926.D, P3926.E, P3926.F, P3926.H (6 ♂), leg. A. Pauly, J.-L. Boevé; Bombuscaro, Podocarpus NP, 04°06'S, 078°58'W, 995m, 16.10.2014, P3936.B, P3936.C, P3936.E (3 ♂), leg. A. Pauly, J.-L. Boevé; Romerillos, nr Podocarpus NP, 04°09'S, 078°56'W, 1100m, 22.10.2014, P3967.V (1 ♀), leg. A. Pauly, J.-L. Boevé, P3962.B, P3962.C, P3962.D, P3962.E (4 ♂), leg. J.-L. Boevé, P3967.Q, P3967.R, P3967.S, P3967.T, P3967.X, P3967.Y (6 ♂), leg. A. Pauly, J.-L. Boevé, flying, P3959, P3961, P3963.I, P3963.J, P3963.K, P3963.L, P3963.M, P3963.N, P3963.O (9 ♂), on leaf of Hedychium coronarium (Zingiberaceae), P3960.C, P3960.D (2 ♂), on leaf of Anthurium sp., P3958.F, P3958.G (2 ♂), leg. J.-L. Boevé; Bombuscaro, 04°05'S, 078°57'W, 930m, 22.03.2015, P4120.D, P4120.E, P4120.G (3 ♂), leg. T. Delsinne; Zamora, Copalinga, 04°05'S, 078°57'W, 1000m, 21.07.2015, P4125 (1 ♂), leg. T. Delsinne.

Figure 15.

Waldheimia erebus. a, b Female (P3926.B), body length 7.5 mm c, d male (P3967.R), body length 8.0 mm. a, c Dorsal views b, d ventral views.

Waldheimia pellucida Konow, 1904

Fig. 16

Distribution

Known from Colombia, Ecuador, Peru, and Bolivia.

Material

Bombuscaro, road to Podocarpus NP, 04°06'S, 078°58'W, 955m, 08.10.2014, flying, P3865 (1 ♂), leg. J.-L. Boevé, P3875, P3876, P3877 (3 ♂), leg. A. Pauly, J.-L. Boevé; Pueblo Viela, 04°38'S, 079°08'W, 1060m, 15.10.2014, on leaf of Anthurium sp., P3928.A (1 ♀), leg. A. Pauly, J.-L. Boevé; N. of Pachicutza, 04°04'S, 078°38'W, 900m, 18.10.2014, P3942.B (1 ♂), leg. A. Pauly, J.-L. Boevé; Miasi, 04°15'S, 078°20'W, 875m, 19.10.2014, on/around leaf of Anthurium sp., P3946.E, P3946.F, P3946.G (3 ♂), leg. J.-L. Boevé; Romerillos, nr Podocarpus NP, 04°06'S, 078°57'W, 995m, 22.10.2014, on leaf of Anthurium sp., P3958.D (1 ♂), leg. J.-L. Boevé.

Figure 16.

Waldheimia pellucida, male (P3958.D), body length 6.5 mm. a Dorsal view b ventral view.

Waldheimia sp.

Fig. 17

Material

nr Loja, Reserva El Madrigal, 04°02'S, 079°11'W, 2270m, 21.10.2014, P3957.A (1 ♂), leg. A. Pauly, J.-L. Boevé.

Figure 17.

Waldheimia sp., male (P3957.A), 9.5 mm. a Dorsal view b ventral view.

Subfamily Nematinae

Pristiphora fernandezi D.R. Smith, 2003

Fig. 18

Distribution

Pristiphora is the only native genus of Nematinae in the Neotropics. Nine species were recorded by Smith (2003), from Mexico to Brazil. Pristiphora fernandezi was described from Colombia, and the specimen from the Podocarpus NP is the first record of both the species and genus from Ecuador.

Material

Station Colibrí, Podocarpus NP, 03°59'S, 079°06'W, 2170m, 17–20.10.2014, with yellow pan trap, P3948 (1 ♀), leg. A. Pauly, J.-L. Boevé.

Figure 18.

Pristiphora fernandezi, female (P3948), body length 8.0 mm. a Dorsal view b ventral view.

Subfamily Selandriinae

Neotropical Selandriinae are poorly studied. Identifications below are based on a key to genera in Smith (2007). Correct combinations of many of the described species have not been published, including three species collected in this study. To apply the correct name to them, the new combinations are presented here.

Adiaclema sp.

Fig. 19

Remarks

A large genus of about 30 species which occur from Mexico to Argentina.

Material

Romerillos, nr Podocarpus NP, 04°09'S, 078°57'W, 1135m, 22.10.2014, flying near rivulet, P3964.B (1 ♀), leg. J.-L. Boevé.

Figure 19.

Adiaclema sp., female (P3964.B), body length 9.0 mm. a Dorsal view b ventral view.

Andeana farcta (Konow, 1900)

Figs 20, 21a–c

Distribution

This species is known from Venezuela, Colombia, Ecuador, and Peru.

Material

Yangana, 04°25'S, 079°09'W, 2450m, 09.10.2014, P3889 (1 ♂), leg. D. Domínguez, J.-L. Boevé; Cajanuma, Podocarpus NP, 04°07'S, 079°11'W, 2700m, 11.10.2014, by sweeping herbs, P3893, P3894, P3895, P3896 (4 ♂), leg. J.-L. Boevé, P3897 (1 ♂), leg. Carlos R. Carreira, J.-L. Boevé; N. of Santiago, 03°48'S, 079°16'W, 2550m, 12.10.2014, under a leaf, P3900.A (1 ♂), leg. J.-L. Boevé, P3900.B (1 ♂), leg. A. Pauly, J.-L. Boevé; Cajanuma, Podocarpus NP, 04°07'S, 079°10'W, 2750m, 14.10.2014, P3921 (1 ♂), leg. A. Pauly, J.-L. Boevé; Loja, Reinaldo Espinosa Botanical Garden, 04°02'S, 079°12'W, 2145m, 14.10.2014, by sweeping grass, P3911 (1 ♂), leg. J.-L. Boevé; S. of Yangana, 04°25'S, 079°09'W, 2515m, 15.10.2014, P3932 (1 ♂), leg. A. Pauly, J.-L. Boevé; Valladolid, along river, 04°33'S, 079°08'S, 1605m, by sweeping grass, P3929 (1 ♀), leg. J.-L. Boevé; Vilcabamba (in the car), P3933 (1 ♂), leg. J.-L. Boevé; nr Loja, Reserva El Madrigal, 04°02'S, 079°11'W, 2270m, 21.10.2014, P3955.A (1 ♂), leg. T. Delsinne, J.-L. Boevé, P3957.B, P3957.C (2 ♂), leg. A. Pauly, J.-L. Boevé, P3954.A, P3954.B, P3954.C (3 ♂), leg. J.-L. Boevé, on Sida sp. (Malvaceae), P3952.A, P3952.B, P3952.C (3 ♂), leg. J.-L. Boevé; Loja, La Argelia, 04°01'S, 079°12'W, 2130m, 25.10.2014, landing on the ground, P3977 (1 ♀), leg. A. Pauly, J.-L. Boevé; nr Loja, Reserva El Madrigal, 04°02'S, 079°11'W, 2270m, 27.03.2015, one specimen on Bidens pilosa (Asteraceae), P4123.A, P4123.B (2 ♂), leg. T. Delsinne.

Figure 20.

Andeana farcta. a, b Female (P3929), body length 10.0 mm c, d male (P3894), body length 8.5 mm. a, c Dorsal views b, d ventral views.

Figure 21.

Pictures of sawfly adults taken in the field. a, c Andeana farcta (P3952 or P3954) b A. farcta (Reserva El Madrigal, 27.03.2015) d Proselandria sp. (P3905).

Pictures of sawfly adults taken in the field. e, f Neonapeptamena sp. (P3973.B) g Stromboceridea albilabris (Reserva El Madrigal, 27.03.2015) h Bolivius notabilis (P3966).

Andeana sp.

Fig. 22

Remarks

A genus of about 15 species in the Neotropics.

Material

N. of Pachicutza, 04°04'S, 078°38'W, 900m, 18.10.2014, P3940.D (1 ♀), leg. J.-L. Boevé; Loja, Estación Científica San Francisco, 03°58'S, 079°05'W, 1820m, 26.03.2015, P4122 (1 ♀), leg. T. Delsinne.

Figure 22.

Andeana sp., female (P3940.D), body length 8.5 mm. a Dorsal view b ventral view.

Aneugmenus sp.

Fig. 23

Remarks

Smith (2005) recorded two species in northern South America, Venezuela, Colombia, and Ecuador, the southern extent of this Holarctic genus. One species, A. merida Smith, was reared from bracken fern, Pteridium aquilinum (L.) in Venezuela. The specimen collected here is a male and could be either of the two species; the male is not known for A. colombia Smith.

Material

nr Loja, Reserva El Madrigal, 04°02'S, 079°11'W, 2280m, 21.10.2014, by sweeping on Pteridium sp. (Dennstaedtiaceae), P3949 (1 ♂), leg. J.-L. Boevé.

Figure 23.

Aneugmenus sp., male (P3949), body length 6.0 mm. a Dorsal view b ventral view.

Bolivius notabilis (Konow, 1899), comb. n.

Figs 21h, 24

Stromboceros notabilis Konow 1899: 294; Konow 1903: 148 (female); Konow 1905: 98; Oehlke and Wudowenz 1984: 399; Taeger et al. 2010: 534.

Remarks

The male holotype cited by Oehlke and Wudowenz (1984) at SDEI is here considered a lectotype designation. Konow did not give the number of specimens available, but it was the only one found at SDEI. It is labelled “Callanga, Cuczo, Peru,”, “Coll. Konow,” “TYPE”, “Stromboeros notabilis Knw., Peru.”.

Distribution

This species is now known from Ecuador, Peru, and Bolivia.

Material

Bombuscaro, road to Podocarpus NP, 04°06'S, 078°58'W, 975m, 08.10.2014, P3873, P3874, P3878 (3 ♂), leg. A. Pauly, J.-L. Boevé; Bombuscaro, Podocarpus NP, 04°06'S, 078°58'W, 995m, 16.10.2014, P3936.A (1 ♂), leg. A. Pauly, J.-L. Boevé; N. of Pachicutza, 04°04'S, 078°38'W, 900m, 18.10.2014, P3940.C (1 ♀), leg. J.-L. Boevé, P3940.B (1 ♂), leg. J.-L. Boevé, P3942.A (1 ♂), leg. A. Pauly, J.-L. Boevé; Romerillos, nr Podocarpus NP, 04°09'S, 078°56'W, 1100m, 22.10.2014, P3962.A (1 ♂), leg. J.-L. Boevé, P3967.B, P3967.C, P3967.D, P3967.E, P3967.F, P3967.G, P3967.H, P3967.K, P3967.L, P3967.P (10 ♂), leg. A. Pauly, J.-L. Boevé, flying, P3963.A (1 ♀), P3963.B, P3963.C, P3963.D, P3963.E, P3963.F, P3963.G, P3963.H (7 ♂), flying near rivulet, P3964.A (1 ♀), P3965, P3966 (2 ♂), leg. J.-L. Boevé; Bombuscaro, 04°05'S, 078°57'W, 930m, 22.03.2015, P4120.C (1 ♀), leg. T. Delsinne.

Figure 24.

Bolivius notabilis. a, b Female (P3940.C), body length 6.5 mm c, d male (P3940.B), body length 7.0 mm. a, c Dorsal views b, d ventral views.

Dochmioglene sp.

Fig. 25

Remarks

A genus of about 10 species in Central and South America.

Material

Loja, Estación Científica San Francisco, 03°58'S, 079°05'W, 1820m, 24.03.2015, P4121.A (1 ♀), leg. T. Delsinne.

Figure 25.

Dochmioglene sp., female (P4121.A), body length 8.0 mm. a Dorsal view b ventral view.

Inea sp.

Fig. 26

Remarks

A genus of about 20 species in Mexico, Central America, and northern South America.

Material

Bombuscaro, Podocarpus NP, 04°06'S, 078°58'W, 955m, 08.10.2014, flying, P3870, P3871, P3872 (3 ♂), leg. J.-L. Boevé; Pueblo Viela, 04°38'S, 079°08'W, 1060m, 15.10.2014, on leaf of Anthurium sp., P3926.G (1 ♂), leg. A. Pauly, J.-L. Boevé; Bombuscaro, Podocarpus NP, 04°06'S, 078°58'W, 995m, 16.10.2014, P3936.F (1 ♂), leg. A. Pauly, J.-L. Boevé; Los Geraneos, 04°09'S, 078°38'W, 855m, 18.10.2014, P3944.C (1 ♀), leg. A. Pauly, J.-L. Boevé, P3943.A, P3943.B (2 ♂), leg. J.-L. Boevé; Bombuscaro, 04°05'S, 078°57'W, 930m, 22.03.2015, P4120.F (1 ♂), leg. T. Delsinne; Romerillos, nr Podocarpus NP, 04°06'S, 078°57'W, 995m, 22.10.2014, on leaf of Anthurium sp., P3958.E (1 ♀), leg. J.-L. Boevé.

Figure 26.

Inea sp., male (P3943.A), body length 5.5 mm. a Dorsal view b ventral view.

Neoanapeptamena sp.

Figs 21e–f, 27

Remarks

This genus includes about three species and occurs only in Colombia, Ecuador, and Peru.

Material

Cajanuma “Mirador”, Podocarpus NP, 04°07'S, 079°11'W, 3000m, 14.10.2014, on leaf of Macrocarpaea arborescens (Gentianaceae), P3918 (1 ♂), leg. J.-L. Boevé, 23.10.2014, P3973.A, P3973.B (2 ♀), leg. T. Delsinne, J.-L. Boevé; Cajanuma, Podocarpus NP, 04°07'S, 079°10'W, 2750m, 14.10.2014, P3920 (1 ♀), leg. A. Pauly, J.-L. Boevé, 2810m, 23.10.2014, P3972.A, P3972.B (2 ♀), leg. T. Delsinne, J.-L. Boevé.

Figure 27.

Neoanapeptamena sp., female (P3973.B), body length 9.0 mm. a Dorsal view b ventral view.

Plaumanniana biclinea (Konow, 1899), comb. n.

Fig. 28

Stromboceros biclinius Konow 1899: 293; Konow 1905: 97; Oehlke and Wudowenz 1984: 369; Taeger et al. 2010: 532.

Strongylogaster biclinius : Enderlein 1920: 361.

Remarks

Two males and four females at SDEI are labelled “Callanga, Cuczo, Peru,” “Coll. Konow,” “TYPUS”, and one male has the label “Stromboceros biclinius Konow, Peru.” The male with Konow's, determination label is hereby designated lectotype. The other specimens are paralectotypes.

This species is now known from Colombia, Ecuador, and Peru.

Material

Pueblo Viela, 04°38'S, 079°08'W, 1060m, 15.10.2014, on leaf of Anthurium sp., P3927.A, P3927.B, P3927.C, P3927.D, P3927.E, P3927.F, P3927.G, P3927.H (8 ♂), leg. A. Pauly, J.-L. Boevé; Romerillos, nr Podocarpus NP, 04°06'S, 078°57'W, 995m, 22.10.2014, on leaf of Anthurium sp., P3958.C (1 ♀), leg. J.-L. Boevé.

Figure 28.

Plaumanniana biclinea. a, b Female (P3958.C), body length 9.0 mm c, d male (P3927.B), body length 8.5 mm. a, c Dorsal views b, d ventral views.

Plaumanniana sp.

Fig. 29

Remarks

About 20 species are included in this genus which occurs form Central America to Argentina.

Material

Miasi, 04°15'S, 078°20'W, 875m, 19.10.2014, flying under leaf of Anthurium sp., P3946.H (1 ♀), leg. T. Delsinne, J.-L. Boevé; Bombuscaro, 04°05'S, 078°57'W, 930m, 22.03.2015, P4120.B (1 ♀), leg. T. Delsinne; Romerillos, nr Podocarpus NP, 04°09'S, 078°56'W, 1100m, 22.10.2014, P3967.A (1 ♀), leg. A. Pauly, J.-L. Boevé, P3967.I, P3967.J, P3967.M, P3967.N (4 ♂), leg. A. Pauly, J.-L. Boevé, on leaf of Hedychium coronarium, P3960.A, P3960.B (2 ♂), leg. J.-L. Boevé, on leaf of Anthurium sp., P3958.A, P3958.B (2 ♂), leg. J.-L. Boevé.

Figure 29.

Plaumanniana sp. a, b Female (P3946.H), body length 9.0 mm c, d male (P3960.A), body length 7.5 mm. a, c Dorsal views b, d ventral views.

Proselandria alvina (Konow, 1899), comb. n.

Fig. 30

Stromboceros alvinus Konow, 1899: 292; Oehlke and Wudowenz 1984: 366; Taeger et al. 2010: 532.

Remarks

Konow described both sexes of this species from “Peru (Callanga, Cuczo)”. Two males and one female labelled as types are at SDEI. The lectotype, here designated, is a male labelled “Callanga, Cuczo, Peru,” “coll. Konow,” “TYPE”, “Stromboceros alvinus Knw, Peru.” The female and other male have the same data but lack a determination label. The female is a paralectotype, but the male belongs to the genus Bolivius. D. R. Smith did not designate a lectotype prior to Oehlke and Wudowenz (1984) who stated one was designated by D. R. Smith in 1976.

Distribution

This species is now known from Colombia, Ecuador, and Peru.

Material

Pueblo Viela, 04°38'S, 079°08'W, 1060m, 15.10.2014, on leaf of Anthurium sp., P3928.C (1 ♂), leg. A. Pauly, J.-L. Boevé; Bombuscaro, Podocarpus NP, 04°06'S, 078°58'W, 995m, 17.10.2014, P3938 (1 ♀), leg. A. Pauly, J.-L. Boevé; N. of Pachicutza, 04°04'S, 078°38'W, 900m, 18.10.2014, P3940.A (1 ♂), leg. J.-L. Boevé.

Figure 30.

Proselandria alvina. a, b Female (P3938), body length 5.5 mm c, d male (P3928.C), body length 5.5 mm. a, c Dorsal views b, d ventral views.

Proselandria carminea (Jörgensen, 1913)

Fig. 31

Remarks

This species occurs only in western South America from Colombia to northern Argentina.

Material

nr Loja, Reserva El Madrigal, 04°02'S, 079°11'W, 2270m, 27.03.2015, P4123.O, P4123.P (2 ♂), leg. T. Delsinne.

Figure 31.

Proselandria carminea, male (P4123.O), body length 5.5 mm, lateral view.

Proselandria spp.

Figs 21d, 32

Remarks

About 20 species are known, from Mexico to Argentina.

Material

Loja, Reinaldo Espinosa Botanical Garden, 04°02'S, 079°12'W, 2145m, 14.10.2014, P3903 (1 ♀), leg. J.-L. Boevé, P3904, P3909, P3910 (3 ♂), leg. J.-L. Boevé, P3916 (1 ♂), leg. A. Pauly, J.-L. Boevé, flying, P3908 (1 ♂), by sweeping grass, P3912 (1 ♀), on leaf of Ismene longipetala (Amaryllidaceae), P3905, P3907 (2 ♀), P3906 (1 ♂), leg. J.-L. Boevé; Pueblo Viela, 04°38'S, 079°08'W, 1060m, 15.10.2014, on leaf of Anthurium sp., P3928.B (1 ♀), leg. A. Pauly, J.-L. Boevé; Miasi, 04°15'S, 078°20'W, 875m, 19.10.2014, on/around leaf of Anthurium sp., P3946.D (1 ♀), leg. J.-L. Boevé.

Figure 32.

Proselandria sp. a, b Female (P3928.B), body length 6.0 mm c, d female (P3903), body length 6.0 mm e, f male (P3904), body length 6.0 mm. a, e Dorsal views b, d, f ventral views, c lateral view.

Stromboceridea albilabris (Konow, 1885)

Figs 21g, 33

Distribution

Stromboceridea occurs from southwestern United States through western South America to northern Argentina. This species is known from Colombia, Ecuador, Peru and Bolivia.

Material

Bosque “El Sayo”, 03°49'S, 079°17'W, 2760m, 12.10.2014, under leaf of bamboo (Poaceae), P3901 (1 ♂), leg. J.-L. Boevé; Cajanuma, Podocarpus NP, 04°07'S, 079°10'W, 2750m, 14.10.2014, P3922, P3923, P3924 (3 ♂), 2810m, 23.10.2014, P3975.A, P3975.B, P3975.C, P3975.D, P3975.E, P3975.F (6 ♂), leg. A. Pauly, J.-L. Boevé; Loja, Reinaldo Espinosa Botanical Garden, 04°02'S, 079°12'W, 2145m, 14.10.2014, by sweeping grass, P3914 (1 ♀), leg. J.-L. Boevé; S. of Yangana, 04°32'S, 079°08'W, 1880m, 15.10.2014, on leaf of Anthurium sp., P3930.A, P3930.B, P3930.C, P3930.D, P3930.E (5 ♂), leg. J.-L. Boevé, P3931.A, P3931.B, P3931.C, P3931.D (4 ♂), leg. A. Pauly, J.-L. Boevé; nr Loja, Reserva El Madrigal, 04°02'S, 079°11'W, 2270m, 21.10.2014, P3957.D (1 ♀), leg. A. Pauly, J.-L. Boevé, P3955.B (1 ♂), leg. T. Delsinne, J.-L. Boevé, on Rumex crispus, P3953.A (1 ♂), leg. J.-L. Boevé; Cajanuma “Mirador”, Podocarpus NP, 04°07'S, 079°10'W, 2810m, 23.10.2014, P3974 (1 ♂), leg. T. Delsinne, J.-L. Boevé; nr Loja, Reserva El Madrigal, 04°02'S, 079°11'W, 2270m, 27.03.2015, P4123.E (1 ♀), leg. T. Delsinne, P4123.C, P4123.D, P4123.I, P4123.J, P4123.K, P4123.L, P4123.M, P4123.N (8 ♂), leg. T. Delsinne.

Figure 33.

Stromboceridea albilabris. a, b Female (P3957.D), body length 9.5 mm c, d male (P3922), body length 7.5 mm. a, c Dorsal views b, d ventral views.

Stromboceridea sp.

Fig. 34

Remarks

About 20 species are known.

Material

Loja, Reinaldo Espinosa Botanical Garden, 04°02'S, 079°12'W, 2145m, 14.10.2014, by sweeping grass, P3913 (1 ♀), leg. J.-L. Boevé; nr Loja, Reserva El Madrigal, 04°02'S, 079°11'W, 2280m, 21.10.2014, near rivulet, P3951 (1 ♂), leg. J.-L. Boevé; nr Loja, Reserva El Madrigal, 04°02'S, 079°11'W, 2270m, 27.03.2015, P4123.F, P4123.G, P4123.H (3 ♀), leg. T. Delsinne, P4123.Q, P4123.R (2 ♂), leg. T. Delsinne.

Figure 34.

Stromboceridea sp., male (P3951), body length 5.5 mm. a Dorsal view b ventral view.

Tioloma nigrita Strand, 1911

Fig. 35

Distribution

Known only from Ecuador.

Material

El Tiro, road Loja - Zamora, 03°59'S, 079°09'W, 2815m, 08.10.2014, P3884 (1 ♀), leg. J.-L. Boevé, P3886 (1 ♀), leg. A. Pauly, J.-L. Boevé, P3885, P3887 (2 ♂), leg. A. Pauly, J.-L. Boevé; El Tiro, road Loja - Zamora, 03°59'S, 079°09'W, 2815m, 17.10.2014, with yellow pan trap, P3939 (1 ♀), 17–20.10.2014, with yellow pan trap, P3947 (1 ♂), leg. A. Pauly, J.-L. Boevé.

Figure 35.

Tioloma nigrita. a, b Female (P3939), body length 8.5 mm c, d male (P3947), body length 7.0 mm. a, c Dorsal views b, d ventral views.

Altitudinal distribution

Sawflies were collected in the field at altitudes ranging from ca. 900 to 3000 m (Fig. 36). Some trends can be deduced from this figure, although with caution because, first, for several species only one or a couple of specimens could be collected, and, second, no systematic (e.g., trapping) methodology was used. At all collection sites, however, no specimens were discarded a priori. Thus, conclusions drawn from this figure may become pertinent by considering relatively common species, and/or when these were collected at several sites far from each other.

Figure 36.

Distribution of collected sawfly species in function of the altitude. The histogram shows mean values with standard deviations; numbers of individuals are given between square brackets.

Among species with more than eight exemplars, Inea sp., the three identified Waldheimia species, the two Plaumanniana species, and Bolivius notabilis occurred at an altitude of ca. 1000 m. In contrast, Stromboceridea albilabris and Andeana farcta were collected above 2000 m. The variation in specific altitudes is rather low (Fig. 36) and most of these nine species were collected at different sites, which suggests that each species is indeed restricted to a given altitude. Three other species, Lagideus podocarpus, Tioloma nigrita, and Neoanapeptamena sp., were collectßed at an altitude of 2500–3000 m, the two latter taxa being known to occur at high altitudes (Malaise 1963, Strand 1913).

The ‘altitudinal slots’ of the sawfly species are probably linked to those of their host plants and/or to abiotic factors. However, such ecological data are still missing for most South American sawflies, and the present identification and description of L. podocarpus is just one example of a species that has remained unknown, and which are living at a high altitude. It emphasizes the importance of preserving a diversity of biotopes. In the case of a country such as Ecuador, the páramo clearly belongs to such valuable natural environments characterized by both a high altitude and very high biological value.

Acknowledgments

Thibaut Delsinne is warmly thanked for the identification of ants, and for providing a significant amount of specimens and several pictures. We thank Alain Pauly (RBINS, Brussels) who also provided us with specimens, Hugo Omar Cabrera (Herbario UTPL) for the identification of plants, Ximena Cueva and Carlos Ruiz Carreira (MUTPL) for attending some field trips, and M. Reynders (Botanical Garden Meise, Belgium) for providing Fuchsia plants, as well as the subject editor Marko Prous, and the reviewers Stefan Schmidt and Andrew Liston for comments on the manuscript. D. R. Smith thanks A. Taeger (SDEI) for allowing study of Konow's type material. The GTI project was managed in collaboration with C. R. Carreira and David Donoso (MUTPL) and funded by the Belgian Directorate-General for Development Cooperation. Permits were obtained from the Ministerio del Ambiente Loja, Ecuador.

References

Boevé J-L, Rozenberg R, Shinohara A, Schmidt S (2014) Toxic peptides occur frequently in pergid and argid sawfly larvae. PLoS ONE 9(8): e105301. doi: 10.1371/journal.pone.0105301

Enderlein G (1920) Symphytologica II. Zur Kenntnis der Tenthrediniden. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 9 [1919]: 347–374.

Konow FW (1899) Neue südamerikanische Stromboceros-Arten (Fam. Tenthredinidae). Wiener Entomologische Zeitung 18(10): 289–299.

Konow FW (1903) Neue Tenthrediniden. (Hym.) (Blennocampides, Selandriades und 1 Varietät von Allantus fasciatus Scop.). Zeitschrift für systematische Hymenopterologie und Dipterologie 3(3): 145–153.

Konow FW (1905) Hymenoptera Fam. Tenthredinidae. In: Wytsman P (Ed.) Genera Insectorum, Fasc. 29. Bruxelles, 176 pp.

Malaise R (1963) HymenopteraTenthredinoidea, subfamily Selandriinae. Key to the genera of the world. Entomologisk Tidskrift 84(3-4): 159–215.

Oehlke J, Wudowenz J (1984) Katalog der in den Sammlungen der Abteilung Taxonomie der Insekten des Institutes für Pflanzenschutzforschung, Bereich Eberswalde (ehemals Deutsches Entomologisches Institut), aufbewahrten Typen - XXII (Hymenoptera: Symphyta). Beiträge zur Entomologie 34(2): 363–420.

Schmidt S, Smith DR (2006) An annotated systematic world catalogue of the Pergidae (Hymenoptera). Contributions of the American Entomological Institute 34(3): 1–207.

Schmidt S, Walter GH (2014) Young clades in an old family: Major evolutionary transitions and diversification of the eucalypt-feeding pergid sawflies in Australia (Insecta, Hymenoptera, Pergidae). Molecular Phylogenetics and Evolution 74: 111–121. doi: 10.1016/j.ympev.2014.02.002

Smith DR (1990) A synopsis of the sawflies (Hymenoptera, Symphyta) of America south of the United States: Pergidae. Revista Brasiliera de Entomologia 34(1): 7–200.

Smith DR (1992) A synopsis of the sawflies (Hymenoptera: Symphyta) of America south of the United States: Argidae. Memoirs of the American Entomological SocietyNo. 39, 201 pp.

Smith DR (2003a) A synopsis of the sawflies (Hymenoptera: Symphyta) of America South of the United States: Tenthredinidae (Allantinae). Journal of Hymenoptera Research 12(1): 148–192.

Smith DR (2003b) A synopsis of the sawflies (Hymenoptera: Symphyta) of America South of the United States: Tenthredinidae (Nematinae, Heterarthrinae, Tenthredininae). Transactions of the American Entomological Society 129(1): 1–45.

Smith DR (2005) Two new fern-feeding sawflies of the genus Aneugmenus Hartig (Hymenoptera: Tenthredinidae) from South America. Proceedings of the Entomological Society of Washington 107(2): 273–278.

Smith DR (2007) Capítulo 18, Familia Tenthredinidae. In: Fernandez F, Sharkey MJ (Eds) Introducción a los Hymenoptera de la región Neotropical. Sociedad Colombiana de Entomologia y Universidad Nacional de Colombia, Bogota D. C., 243–252.

Smith DR (2012) The South American genus Lagideus Konow (Hymenoptera: Pergidae: Syzygoniinae), a supplement. Zootaxa 3413: 1–18.

Smith DR, Bado SG (2004) First food plant record for Lagideus Konow (Hymenoptera: Pergidae), a new species feeding on Fuchsia and Ludwigia (Onagraceae) in Argentina. Journal of Hymenoptera Research 13(1): 120–124.

Smith DR, Janzen DH (2006) Food plants and life histories of sawflies of the family Argidae (Hymenoptera) in Costa Rica, with descriptions of two new species. Journal of Hymenoptera Research 12: 193–208.

Strand E (1913) Hymenoptères. Tenthrédinides, Pompilides, Crabronides, Apides. Mission du service géographique de l’armée pour la mesure d’un arc de méridien équatorial en Amérique du Sud (1899-1906) 10(2): 13–32.

Taeger A, Blank SM, Liston AD (2010) World Catalog of Symphyta (Hymenoptera). Zootaxa 2580: 1–1064.