Research Article |
Corresponding author: Gilson R.P. Moreira ( gilson.moreira@ufrgs.br ) Academic editor: Erik J. van Nieukerken
© 2018 Gabriela T. Silva, Gilson R.P. Moreira, Héctor A. Vargas, Gislene L. Gonçalves, Marina D. Mainardi, Germán San Blas, Donald Davis.
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:
Silva GT, Moreira GRP, Vargas HA, Gonçalves GL, Mainardi MD, Blas GS, Davis D (2018) Overlooked gall-inducing moths revisited, with the description of Andescecidium parrai gen. et sp. n. and Oliera saizi sp. n. from Chile (Lepidoptera, Cecidosidae). ZooKeys 795: 127-157. https://doi.org/10.3897/zookeys.795.27070
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There are still many gall systems associated with larvae of Lepidoptera in which the true gall-inducers have not been identified to species. Reports on misidentification of gall inducers have been recurrent for these galls, particularly in complex gall-systems that may include inquilines, kleptoparasites, and cecidophages, among other feeding guilds such as predators and parasitoid wasps. Here we describe and illustrate the adults, larvae, pupae and galls, based on light and scanning microscopy, of Andescecidium parrai gen. et sp. n. and Oliera saizi sp. n., two sympatric cecidosid moths that are associated with Schinus polygamus (Cav.) Cabrera (Anacardiaceae) in central Chile. Adults, immatures, and galls of the former did not conform to any known cecidosid genus. Galls of A. parrai are external, spherical, and conspicuous, being known for more than one century. However, their induction has been mistakenly associated with either unidentified Coleoptera (original description) or Oliera argentinana Brèthes (recently), a distinct cecidosid species with distribution restricted to the eastern Andes. Galls of O. saizi had been undetected, as they are inconspicuous. They occur under the bark within swollen stems, and may occur on the same plant, adjacent to those of A. parrai. We also propose a time-calibrated phylogeny using sequences from mitochondrial and nuclear loci, including specimens of the new proposed taxa. Thus in addition to clarifying the taxonomy of the Chilean cecidosid species we also tested their monophyly in comparison to congeneric species and putative specimens of all genera of Neotropical and African cecidosids.
Anacardiaceae , Cecidosid moths, insect galls, Neotropical microlepidoptera, Schinus polygamus
Cecidosidae are a small group of ancient gall-inducing and stem-mining micromoths restricted in distribution to the Southern Hemisphere. They include seven genera and 19 species that are distributed in southern South America, southern Africa, and New Zealand (for taxonomic reviews see
A similar case of such a complex system was recently found for a cecidosid-induced gall in central Chile. In fact, such a spherical, external gall was briefly described for the first time by
While searching later in the field for additional material of this new cecidosid genus, three of the authors of this study (G. R. P. Moreira, G. San Blas and H. A. Vargas) found that there was a second undescribed, cecidosid species in Chile occurring in sympatry, forming galls on swollen stems of the same S. polygamus plants, as already suggested by
Adults used in the study were reared by H.A. Vargas from galls either attached to Schinus polygamus branches or detached ones found in litter, under the canopy cover, which were maintained in small plastic vials at room temperature in the entomology laboratory of the Facultad de Ciencias Agronómicas, Universidad de Tarapacá, Arica, Chile. They were periodically inspected for emerged adults, which were then pinned and dried. Larvae and pupae were obtained by dissecting additional galls. They were fixed in Kahle-Dietrich´s fluid and preserved in 70% EtOH. For DNA analyses, additional specimens were preserved in 100% EtOH at -20 °C.
For gross morphology studies, the specimens were cleared in a 10% potassium hydroxide (KOH) solution and mounted on slides with either glycerin jelly or Canada balsam. Observations were made with the aid of a Leica M125 stereomicroscope. Structures selected to be drawn were previously photographed with an attached Sony Cyber-shot DSC-H10 digital camera. The vectorized line drawings were made with the software CorelPhotoPaint X4, using the corresponding digitalized images as a guide. At least four specimens were used for the descriptions of each life stage. Measurements were made with an attached ocular micrometer; values are presented as mean ± standard deviation unless noted otherwise.
Specimens used in scanning electron microscope (SEM) analyses were dehydrated in a Bal-tec CPD030 critical-point dryer, mounted with double-sided tape on metal stubs and coated with gold in a Bal-tec SCD050 sputter coater. They were examined and photographed in a JEOL JSM6060 scanning electron microscope at Centro de MicroscopiaEletrônica (CME) of Federal University of Rio Grande do Sul, (
The material examined is deposited in the following collections:
IADIZA Instituto Argentino de Investigaciones de las Zonas Áridas, CCT-CONICET Mendoza, Mendoza, Argentina;
IDEA Colección Entomológica de la Universidad de Tarapacá, Arica, Chile;
LMCI Laboratório de Morfologia e Comportamento de Insetos /
We assembled data from two mitochondrial loci and one nuclear protein-coding locus from 12 species representing all Neotropical lineages of Cecidosidae (Cecidonius Moreira & Gonçalves, Cecidoses, Dicranoses Kieffer & Jörgensen, Eucecidoses Brèthes and Oliera) to infer the relationship of Andescecidium gen. n. and the phylogenetic status of A. parrai sp. n. and its congeneric undescribed taxon Andescecidium sp. (Table
Total genomic DNA was purified from fresh larval tissue of Andescecidium parrai, Andescecidium sp. and two additional species of Oliera (O. saizi and Oliera sp.), using the PureLink genomic DNA kit (Life, Invitrogen, USA) following the manufacturer’s instructions. Sequences of Cecidonius pampeanus, Cecidonius sp., Cecidoses eremita Curtis, Dicranoses congregatella (Brèthes), Eucecidoses minutanus Brèthes, Oliera argentinana, Scyrotis granosa (Meyrick) were previously generated by
Approximately 2.5 Kb of gene segments were amplified by polymerase chain reaction (PCR): 1421 bp of the cytochrome oxidase subunit I (COI), 474 bp of the 16S ribosomal RNA (16S) and 395 bp of the Wingless (Wg) gene loci, with primers and conditions described in
Specimens used in this study for the phylogenetic inference in Cecidosidae.
Family | Genus | Species | Vouchers* | Pop (#) Locality | GenBank (accession number) | BOLD Systems | ||
---|---|---|---|---|---|---|---|---|
COI | 16S | Wg | COI-5P | |||||
Cecidosidae | Andescecidium | parrai | LMCI 231-5 | Til-Til/Rungue/Chile | MH750899 | – | MH750908 | MISA020-18 |
LMCI 233-5 | Cuesta Barriga/Chile | MH750900 | – | MH750906 | MISA021-18 | |||
LMCI232-6 | Cuesta La Dormida/Chile | MH750901 | – | MH750907 | MISA022-18 | |||
sp. | LMCI 163-15 | Mendoza/Argentina | MH750902 | – | – | MISA023-18 | ||
Cecidonius | pampeanus | LMCI 16-24 | Eldorado do Sul/Brazil | MH750864 | MH750886 | MH750892 | MISA024-18 | |
sp. | LMCI 14-72 | Curitiba/Brazil | MH750881 | MH750887 | MH750893 | MISA025-18 | ||
Cecidoses | eremita | LMCI 163-1 | Mendoza/Argentina | MH750879 | MH750888 | MH750895 | MISA026-18 | |
Dicranoses | congregatella | LMCI 3-1 | Canguçu/Brazil | MH750880 | MH750889 | MH750896 | MISA027-18 | |
Eucecidoses | minutanus | LMCI 163-21 | Mendoza/Argentina | MH750881 | MH750890 | MH750897 | MISA028-18 | |
Oliera | argentinana | LMCI 6-11 | Canguçu/Brazil | MH750883 | MH750891 | MH750898 | MISA029-18 | |
saizi | LMCI 232-2 | Til-Til/Rungue/Chile | MH750904 | – | MH750910 | MISA030-18 | ||
LMCI232-4 | Cuesta La Dormida/Chile | MH750905 | – | MH750909 | MISA031-18 | |||
sp. | LMCI 163-13 | Mendoza/Argentina | MH750903 | – | – | MISA032-18 | ||
Scyrotis | granosa | LMCI 228-2 | Tsitsikamma/South Africa | MH750885 | – | – | MISA033-18 | |
pulleni |
|
Transvaal /South Africa | - | – | – | LNAUT084-14 | ||
Adelidae | Adela | septentrionella | – | – | EU884115 | – | – | – |
Incurvariidae | Incurvaria | masculella | – | – | AF150926 | – | – | – |
Heliozelidae | Antispila | ampelopsia | RMNH.INS.30326 | – | MF118352 | – | – | HELA119-15 |
Prodoxidae | Greya | enchrisa | – | – | EU884123 | – | – | – |
We used BEAST v2.02 (
The concatenated tree corroborated our hypothesis of monophyletic status for the new genus and species and their relationships within Cecidosidae with strong node support (BPP = 1) (Figure
Molecular phylogeny of Cecidosidae showing monophyletic status and relationships of Andescecidium and Oliera saizi (highlighted by green and brown squares, respectively). Bayesian time-calibrated consensus tree based on cytochrome oxidase subunit I (COI), r16S ribosomal (16S), and Wingless (Wg) genes. Representative species of other Adeloidea families (Adelidae, Incurvariidae, Heliozelidae and Prodoxidae) were used to root the tree. Numbers above branches indicate posterior probability support for the equivalent node. Blue bars indicate confidence interval (95% HDP) for each node age estimate, presented in millions of years ago (Mya).
Estimates of pairwise genetic distance (%) among Cecidosidae lineages based on concatenated mitochondrial DNA sequences using p-distance.
1. | 2. | 3. | 4. | 5. | 6. | 7. | 8. | 9. | 10. | 11. | 12. | 13. | 14. | 15. | 16. | ||
1. | Andescecidium parrai | ||||||||||||||||
2. | Andescecidium sp. | 9 | |||||||||||||||
3. | Cecidonius pampeanus | 12 | 12 | ||||||||||||||
4. | Cecidonius sp. | 14 | 15 | 7 | |||||||||||||
5. | Cecidoses eremita | 16 | 15 | 14 | 15 | ||||||||||||
6. | Dicranoses capsulifex | 19 | 18 | 19 | 21 | 21 | |||||||||||
7. | Dicranoses congregatella | 20 | 19 | 20 | 22 | 21 | 9 | ||||||||||
8. | Eucecidoses minutanus | 15 | 13 | 13 | 15 | 13 | 21 | 20 | |||||||||
9. | Oliera argentinana | 11 | 13 | 10 | 11 | 12 | 19 | 18 | 11 | ||||||||
10. | Oliera saizi | 11 | 12 | 11 | 12 | 12 | 18 | 18 | 13 | 5 | |||||||
11. | Oliera sp. | 12 | 11 | 12 | 15 | 13 | 19 | 18 | 12 | 4 | 6 | ||||||
12. | Scyrotis granosa | 21 | 20 | 20 | 23 | 22 | 22 | 23 | 23 | 21 | 20 | 19 | |||||
13. | Scyrotis pulleni | 16 | 17 | 19 | 19 | 17 | 22 | 21 | 18 | 18 | 16 | 16 | 21 | ||||
14. | Adela septentrionella | 20 | 19 | 19 | 21 | 21 | 21 | 21 | 21 | 19 | 16 | 16 | 18 | 21 | |||
15. | Incurvaria masculella | 20 | 20 | 19 | 22 | 20 | 23 | 24 | 20 | 18 | 19 | 19 | 21 | 22 | 17 | ||
16. | Antispila ampelopsia | 23 | 23 | 23 | 25 | 24 | 25 | 25 | 25 | 22 | 22 | 22 | 22 | 25 | 19 | 20 | |
17. | Greya enchrysa | 19 | 18 | 18 | 21 | 21 | 23 | 24 | 21 | 18 | 20 | 20 | 19 | 21 | 18 | 14 | 18 |
Andescecidium parrai Moreira & Vargas, sp. n.
Andescecidium gen. n. resembles Cecidonius in having pupae bearing strong spines on the anterior margins of the abdominal terga and larvae with two stemmata, which make them unique among cecidosids. However, Andescecidium shows several adult, larval, and gall features which in conjunction differentiate it from the latter. Adults are much larger, bear mandibular rudiments and have reduced three-segmented maxillary palpi, which are respectively absent and with two in the small Cecidonius. The ovipositor in Andescecidium is reduced in size, and associated with an inconspicuous dorsal crest in the oviscapt cone that is used to lay eggs on superficial external buds. The ovipositor of Cecidonius is very long, having the oviscapt cone dorsally anchored by a conspicuous crest, thus allowing oviposition deep into the stem. Contrary to the larvae of Cecidonius, which are unique among cecidosids in having setae of much longer size on the thorax, those of Andescecidium show these structures uniform on thorax and abdomen. The spherical galls of Andescecidium are associated with stem buds, growing on the external surface from the beginning; they are pedunculate and have fully developed walls. Those of Cecidonius grow initially under the bark, erupting through the stem surface later in ontogeny, and with their bases remaining open when mature, clogged with feces.
(Figs
Male. Forewing length ca. 12.1 mm (n = 1). Head (Figure
Female. Forewing length ca. 11.5 mm (n = 2). Similar to male in general, except that sensilla are less abundant and smaller on the antennae, and abdominal sternum VII with caudal margin more sclerotized, bearing a dense ring of stout, elongate setae (Figure
Andescecidium parrai adult morphology under light microscopy (LMCI 218-03). A head, anterior view B metathoracic furcasternum, posterior (closed arrow points to left furcal apophysis) C fore-, median- and hindlegs, from left to right, respectively D female genitalia, dorsal (open and closed arrows point to the anterior ends of left anterior and posterior right apophyses, respectively) E distal portion of female genitalia in detail, lateral (enlarged area marked by rectangle in D; closed arrow points to the ovipositor tip; asterisk identifies oviscapt seen by transparency; bracket indicates distal margin of sternum eight). Scale bars: 0.3 mm (A, B), 1 mm (C), 0.2 mm (D, E).
The genus name is derived from a composition between the Portuguese Andes and Cecidia (a gall; from the Greek kekídion). Thus, the epithet refers to the Andes Mountains where the galls of Andescecidium were first found.
As discussed for the monotypic genus.
As discussed for the monotypic genus.
Chile: Holotype ♂, Cuesta Barriga roadside, Padre Hurtado, Metropolitan Region, emerged June 2013, ex gall on Schinus polygamus collected 22.V. 2013, HA Vargas & GRP Moreira leg. (
Immatures used for morphological description, fixed in Kahle-Dietrich’s fluid, preserved in 70% EtOH: 33°31'24"S, 70°54'35"W, Cuesta Barriga roadside, Padre Hurtado, Metropolitan Region, HA Vargas & GRP Moreira leg., 13 larvae (22.V.2013, LMCI 218-3, 8, 9, 10, 12; 25.XI.2013, LMCI 231-5, 6), 5 pupae (22.V.2013, LMCI 218-3, 5, 11, 14), 3 galls (22.V.2013, LMCI 218-5, 11, 13); 3°3'42"S, 71°0'35"W, roadside on Cuesta La Dormida, border Til-Til/Valparaiso, HA Vargas & GRP Moreira leg., 4 larvae (28.XI.2013, LMCI 233-6), 5 galls (22.V.2013, LMCI 216- 1 to 5, 11); 33°00'31"S, 70°53'52"W, 712m, roadside near Til-Til, Rungue, Metropolitan Region, HA Vargas & GRP Moreira leg., 6 larvae (26.XI.2013, LMCI 232-6, 7), 4 galls (26.XI.2013, LMCI 232-8), G San Blas leg., 12.X.2011, 3 larvae (donated to LMCI 163-14), 46 larvae (IADIZA), 5 galls (IADIZA), 7 pupae (extracted from dried galls on 3.V.2012, IADIZA), G San Blas leg., 15.III.2013, 3 dried pupae; 35°36'00"S, 71°07'17.02"W, 426m, Ruta J-65, 25km from Curicó, Maule Region, G San Blas leg., 12.X.2011, 16 larvae; 36°39'30"S, 72°16'41"W, 43m, roadside in Cruce Nebuco, Bio-Bio Region, HA Vargas, LE Parra & GRP Moreira leg., 2 galls (30.V.2013, LMCI 223-1) ; 35°31'53"S, 71°18'4"W and 35°31'29"S, 71°18'33"W, P. Sta. Edilia, San Clemente, Maule Region, LE Parra & GT Silva leg., 25.XI.2016, 2 larvae donated to LMCI (330-1, 2).
Same data as above, fixed and preserved in 100% EtOH at -20 °C for DNA extraction: n = 2, LMCI 231-5; n = 6, LMCI 233-5, 6.
Genitalia preparation on slide of male by LE Parra, UCCC, Parque Nacional La Campana, Sector Ocoa, Hijuelas, Valparaíso Region, L. Sáiz leg., March-April 1993, with label “Cecidoses argentinana (Brèthes), Det. LE Parra”; added label “Andescecidium parrai Moreira & Vargas, Det. GRP Moreira; 9.V.2018”, 1♂ (LMCI 218-03-04) and 1♀ (LMCI 218-03-03), cleared in KOH 10%, preserved in 70% EtOH, same data as the holotype.
Named in honor of Prof. Dr. Luis E. Parra, from the Universidad of Concepción, who for the first time characterized such a gall-inducing moth, for his great contribution to the development of Lepidopterology in Chile.
Last instar larva (Figs
Thorax (T) and abdomen (A) subcylindrical, creamy-white and covered with microtrichia (Figure
Morphology of Andescecidium parrai last larval instar under scanning electron microscopy. A, B head, lateral (A) and antero-dorsal (B) views C stemmata, lateral D antenna, lateral E labium, lateral F maxilla, lateral G thorax, lateral H first abdominal spiracle, lateral I detail of second abdominal segment, showing aligned setae (arrow indicates spiracle), lateral J callus of second abdominal tergum, latero-dorsal K last abdominal segments, postero-lateral L ventral lobe of tenth abdominal segment (enlarged area marked with rectangle in K). Scale bars: 150 µm (A, B), 30 µm (C, L), 20 µm (D, E), 50 µm (F, I), 300 µm (G), 10 µm (H), 100 µm (J, K).
Pupa (Figs
Andescecidium parrai pupal morphology with scanning electron microscopy. A, C head and prothorax, under ventral and lateral views, respectively B frontal process of head (= gall cutter), ventral D clypeus and labrum, ventral E abdominal terga, lateral F spines and setae of second abdominal tergum in detail, lateral G abdominal pleura and sterna, latero-dorsal H spiracle of third abdominal segment, dorsal I tergal spines of ninth abdominal segment, postero-lateral. Scale bars: 500 µm (A, C, E), 150 µm (B, D, F), 300 µm (G), 30 µm (H), 70 µm (I).
The large, short pedunculated galls of A. parrai develop externally from the beginning, on stem buds of S. polygamus terminal and sub-terminal branches (Figure
Little is known about biology of A. parrai galls. Our field observations suggest that the species is univoltine, galls starting to develop during late spring and summer, when most leaf buds are found on S. polygamus plants (
Natural history of Andescecidium parrai on Schinus polygamus. A host-plant habitat at Cuesta La Dormida, intersection Til-Til/Valparaíso, beginning of road to Rungue, Chile, 33°3'42"S, 71°0'35"W (seta on the horizon points to the El Altar mountain, Metropolitan region) B young, developing galls, lateral C mature gall, lateral, with pedunculus (indicated by arrow) in detail in the upper right corner D, E old, empty galls under lateral and posterior views, respectively (setae point to apical open). Scale bars: 4 mm (B, C, E), 3 mm (D).
A typical member of Oliera, having two-segmented labial palpi, pupa with five-pointed frontal process and galls enclosed within swollen terminal branches of Schinus plants (for a generic review, see
Male. Forewing length varying from 5.5 to 6.3 mm (n = 3). Head: Vertex and frons covered with smooth, narrow, brownish gray scales. Compound eyes black. Antennae filiform, about half the length of forewing, with brownish gray scales dorsally, ventral half with short sensilla. Labrum semicircular, short. Mandibles poorly developed, as small stubs. Pilifers absent. Maxillae with galeae reduced to small lobes; maxillary palpi tri-segmented (ratios of segments from base ~1.0:0.9:0.6). Labial palpi tri-segmented (ratios of segments from base ~1.0:0.8:0.7). Maxillary and labial palpi with brownish gray scales. Thorax: Mostly with brownish gray scales. Wing venation (Figure
Oliera saizi adult morphology, under light microscopy (paratypes). A fore and hindwing venation, dorsal view B schematic representation of female oviscapt cone, lateral (asterisk indicates the distal end of internal dorsal crest; closed and open arrows point to the anterior ends of anterior and posterior left apophyses, respectively) C ovipositor tip in detail, lateral (enlarged area marked by rectangle in B) D male genitalia, ventral (phallus and juxta omitted) E distal section of right valva pectinifer in detail (enlarged area marked by rectangle in D) F phallus, ventral G juxta, ventral. Scale bars: 1 mm (A), 200 µm (B), 10 µm (C), 100 µm (D, F, G), 30 µm (E).
Female. Forewing length varying from 5.2 to 5.9mm (n = 7). Similar to male in general, but with abdominal sternumVII showing a ring of stout setae on caudal margin. Genitalia (Figure
Chile: Holotype ♂, Cuesta Barriga roadside, Padre Hurtado, Metropolitan Region, emerged December 2013, ex gall on Schinus polygamus collected 25 November 2013, HA Vargas & GRP Moreira leg. (
Immature specimens used for descriptions: 33°31'24"S, 70°54'35"W, Cuesta Barriga roadside, Padre Hurtado, Metropolitan Region, HA Vargas & GRP Moreira leg., 6 larvae (25.XI.2013, LMCI 231-3, 4); 33°3'42"S, 71°0'35"W, roadside on Cuesta La Dormida, near border Til-Til/Valparaiso, Metropolitan Region, HA Vargas & GRP Moreira leg., 8 larvae (20.V.2013, LMCI 216-14), 2 pupae (28.XI.2013, LMCI 233-3); 33°00'31"S, 70°53'52"W, roadside near Til-Til, Rungue, Metropolitan Region, HA Vargas & GRP Moreira leg., 4 larvae (26.XI.2013, LMCI 232-4); 36°48'22"S, 71°44'36"W, roadside near Recinto, Bio-Bio Region, HA Vargas, LE Parra & GR. Moreira leg., 30.V.2013, 12 larvae (LMCI 222-2 to 6) and 2 pupal exuviae (LMCI 222-1).
Data as above, fixed and preserved in 100% EtOH at -20 °C for DNA extraction (n = 2, LMCI 232-4; n = 2, LMCI 233-2).
Puente El Yeso, Cajón del Maipo, Metropolitan Region, R. Charling C. leg., 4 males, with one genitalia (DRD 16536) and one wing (DRD 29925) preparations; 2 females, with one wing and genitalia preparation (DRD 16537),
Named in honor of Prof. Dr. Francisco Sáiz, from the Universidad Católica de Valparaiso, who first noticed the existence of these galls, for his great contribution to the development of Cecidology in Chile.
Last instar larva (Figs
Morphology of Oliera saizi last larval instar under scanning electron microscopy. A head, antero-lateral view B labrum and clypeus, anterior C antenna, lateral D maxilla, anterior E labium, ventral F detail of maxilla (indicated by asterisk) and mandibula (pointed by arrow) bases, showing protuberances G mesothoracic leg rudiment, lateral H callus of third abdominal tergum, latero-dorsal I last abdominal segments, lateral. Scale bars: 250 µm (A), 40 µm (B), 20 µm (C, E, F), 30 µm (D), 60 µm (G), 100 µm (H), 200 µm (I).
Thorax (T) and abdomen (A) creamy white, mostly covered with rounded microtrichia; prothoracic shield weakly defined by a pair of slightly differentiate areas, with reduced microtrichia. A1–7 with well-developed callus (Figure
Pupa (Figs
Oliera saizi pupal morphology using scanning electron microscopy. A, C head and prothorax, under ventral and lateral views, respectively B frontal processes of head (= gall cutter), ventral D spiracle of second abdominal segment, lateral E abdominal terga VI and VII, lateral F detail with supraspiracular setae and spines of abdominal tergum VI, lateral G detail with subspiracular setae of abdominal segment five, lateral H, J last abdominal segments, under lateral and dorsal views, respectively I single tergal spine on abdominal segment nine in detail, lateral. Scale bars: 250 µm (A), 100 µm (B, I), 350 µm (C), 30 µm (D), 200 µm (E, H, J), 150 µm (F), 50 µm (G).
Oliera saizi develops spindle-shaped galls enclosed within swollen stems of S. polygamus subterminal branches (Figure
Little is also known about the biology of O. saizi. Similar to what has been found for O. argentinana (
Natural history of Oliera saizi on Schinus polygamus. A host-plant habitat at Cuesta Barriga, near Santiago city, Chile, 33°31'24"S, 70°54'35"W (asterisk locates the plant; open arrow indicates commune of Padre Hurtado in the valley, Metropolitan region) B apical branches showing swollen stem with galls under bark C pupal exuvium protruded from the gall exit hole, just after the adult emergence (enlarged area pointed by arrow in (B) D intact empty gall, shown by detaching the bark (indicated by arrow) E empty galls, with decaying gall-wall still remaining, lateral F old gall signs appearing as small craters on surface of dried branch. Scale bars: 5 mm (B), 2 mm (C), 3 mm (D, E), 15 mm (F).
Galls of both A. parrai and O. saizi have been found only on branches of Schinus polygamus (Cav.) Cabrera (Anacardiaceae), a small tree with single and glabrous leaves and slender spiny branches (
Geographic distribution of Andescecidium parrai (green circles) and Oliera saizi (blue circles) in central Chile. Regions are indicated by Roman numbers except for the Metropolitan (RM). Localities are indicated by cardinal numbers: 1 Parque Nacional La Campana 2 Cuesta La Dormida 3 Til-Til/Hungue 4 Cuesta Barriga 5 Cajón del Maipo 6 Curicó 7 San Clemente 8 Cruce Nebuco, and 9 Recinto; for a complete description of corresponding localities, see list of specimens examined.
This study illustrates further how important are intensive studies including the immature stages to achieve correct identification of true gall-inducing in Lepidoptera. It confirms preliminary findings of
Andescecidium parrai and O. saizi resulted as unique lineages in the present study from both morphological and molecular analyses. The former appeared as one of the most derived lineages to be evolved within the extant cecidosids. The genetic distance of A. parrai is ca. 10% from the nearest related species, an additional undescribed cecidosid existing in Mendoza, Argentina, which was included in the present study for comparison. Oliera saizi diverged ca. 6%, from an undescribed Oliera species collected at the same latitude in Argentina, and ca. 5% from the type species O. argentinana, sampled in southern Brazil. Thus molecular data suggest that although described as monotypic, there is at least one more species belonging to Andescecidium, and another to Oliera. In fact, these species share morphological characteristics with congenerics regarding the gall and larval stage, but unfortunately they have not been reared yet to the pupal and adult stages, upon which their descriptions are pending.
The present study also provides support to previous statements in the literature in the sense that inclusion of immature stages and galls is essential for elucidation of the taxonomy of Cecidosidae (
A complete scenario of relationships within Cecidosidae, inferred in the Bayesian calibrated-time tree, retrieved high posterior probabilities for all nodes. It was somewhat unexpected to obtain a fully resolved genus-level tree, based on only two to three markers (see
In this study we recorded for the first time two new species of cecidosids for Chile that belong to two relatively recent genera (Andescecidium and Oliera) that apparently have representatives distributed on both the west and east sides of the Andes. Based on a broader survey exploring several southern South America biomes,
We thank the staff members of Centro de Microscopia Eletrȏnica and Thales O Freitas (