Nothotrioza Burckhardt

Nothotrioza Burckhardt gen. n. Type species. Nothotrioza myrtoidis Burckhardt sp. n., by present designation. Description. Adult. Dark brown. Body large; thorax massive. Head (Fig. 3 A) deflexed from longitudinal body axis, much narrower than mesothorax, about as wide as pronotum; vertex bearing long setae; frons large with median ocellus at dorsal edge; genae convex, not forming processes, covered in long setae. Clypeus prominent, pear-shaped, laterally compressed; rostrum long. Antenna 10 -segmented, bearing a single subapical rhinarium each on segments 4, 6, 8 and 9 respectively, Pronotum much narrower than mesonotum, strongly inclined. Mesonotum covered in long conspicuous setae. Forewing (Figs 2 A–C) large hyaline; vein R+M+Cu not strictly trifurcating but with a very short, inconspicuous vein M+Cu; vein Rs long, evenly curved; bifurcation of vein M proximal of line connecting apices of veins Rs and Cu 1 a; cell m 1 much larger than cu 1; vein Cu much longer than Cu 1 b. Radular spinules present in cells m 1, m 2 and cu 1; surface spinules lacking. Hindwing shorter than forewing, hyaline. Metatibia with 6–8 (2–4 + 3–4) apical, sclerotised spurs (Fig. 3 B). Male proctiger (Figs 3 D, G) large with produced hind margin. Paramere (Figs 3 E, H) lamellar, narrowing to apex which forms sclerotised tooth. Distal portion of aedeagus (Figs 3 F, I) with hook-shaped apical dilatation. Female terminalia (Figs 4 A, B) relatively short, cuneate; proctiger with truncate apex; valvula ventralis with many teeth in apical part (Fig. 4 C). Fifth instar nymph (Figs 2 D, E) lacking sectasetae. Antenna 3 -segmented, rhinaria formula 3333. Humeral lobes absent. Tarsal arolium small, not petiolate (Fig. 4 D). Caudal plate (Figs 4 E, F) with two tubercles bearing each a tooth-like seta at hind margin. Circumanal ring (Fig. 4 E) consisting of several rows of pores. Etymology. From Greek νόθος = illegitimate, false, and Trioza, a related genus. Composition. Nothotrioza currently comprises three species, viz. N. myrtoidis and N. cattleiani, respectively, which are both associated with Psidium, as well as N. tavaresi associated with Malpighiaceae. Distribution. Brazil. Systematic relationships. White & Hodkinson (1985) subdivided the Triozidae, based on adult and nymphal characters, into the Neolithinae, Triozamiinae and Triozinae with the tribes Pauropsyllini and Triozini. Hollis & Broomfield (1989) transferred the Triozamiinae to the Homotomidae. Klimaszewski (1993) separated the Rhinopsyllidae from the Triozidae based on differences in the hind wing venation which were shown to be trivial by Burckhardt & Lauterer (1997) who synonymised the two. Li (2011) raised the Triozidae to superfamily status with four families. This classification, artificial and based on Chinese Psylloidea only, is not applicable to the world fauna and was rejected by Burckhardt & Ouvrard (2012). In the classification of White & Hodkinson (1985) Nothotrioza falls within the Neolithiinae, as Neolithus sp. sensu White & Hodkinson (1985: Fig. 157, key page 289) is a misidentification (Burckhardt 1988) referring to Nothotrioza sp. Neolithus Scott, 1882, contains the single species N. fasciatus Scott, 1882, associated with Sapium glandulosum (Euphorbiaceae). N. fasciatus has been reported from Argentina, Brazil, Paraguay and Uruguay (Hodkinson & White 1981, Burckhardt 1988). White & Hodkinson (1985) included into the Neolithiinae also tentatively Schedoneolithus, a Peruvian genus comprising a single free-living species associated with Dunalia (Solanaceae). Nothotrioza may be most closely related to Neolithus because of the large body size, the strongly deflexed head, which is much narrower than the thorax, the large frons, the strongly arched thorax, the long setae covering head and thorax, the genae not developed into conical processes, the not strictly trifurcating vein R+M+Cu of the forewing, the variable number of sclerotised apical metatibial spurs, the posteriorly lobed male proctiger, and the relatively short, cuneate female terminalia in the adults, as well as the lack of sectasetae and the circumanal ring consisting of several rows of pores in the nymphs. The two genera differ in the combination of these characters from all other triozid genera but without an analysis of all triozid genera it is difficult to judge if these characters express phylogenetic relationships. Schedoneolithus resembles the two genera in the lacking genal processes and the large frons but differs in the less strongly arched thorax, the strictly trifurcating vein R+M+Cu and the fixed number, 1 + 3, of apical metatibial spurs in the adults. The resemblence of Schedoneolithus to Neolithus is superficial and does probably not reflect close phylogenetic relationship.Published as part of Carneiro, Renê G. S., Burckhardt, Daniel & Isaias, Rosy M. S., 2013, Biology and systematics of gall-inducing triozids (Hemiptera: Psylloidea) associated with Psidium spp. (Myrtaceae), pp. 129-146 in Zootaxa 3620 (1) on pages 131-132, DOI: 10.11646/zootaxa.3620.1.6, http://zenodo.org/record/22205

Biology and systematics of gall-inducing triozids (Hemiptera: Psylloidea) associated with Psidium spp. (Myrtaceae)

Carneiro, Renê G. S., Burckhardt, Daniel, Isaias, Rosy M. S. (2013): Biology and systematics of gall-inducing triozids (Hemiptera: Psylloidea) associated with Psidium spp. (Myrtaceae). Zootaxa 3620 (1): 129-146, DOI: http://dx.doi.org/10.11646/zootaxa.3620.1.

Nothotrioza myrtoidis

Biology of N. myrtoidis Psidium myrtoides is an evergreen plant which has little demarcation of leaf flushing, maturation and senescence phenophases (Fig. 5 A). Mature leaves are found throughout the year in all individuals of the population. A variable number of individuals presents senescence and their leaves fall along the year. In October 2009, all the analysed individuals showed leaf senescence, coinciding with the more pronounced phenological activity of budburst in this population. Leaf flushing decreased but persisted in a smaller portion of the population throughout the year. The induction of the galls by Nothotrioza myrtoidis began in October and lasted until early December. Gall growth and development started in November and lasted until September of the following year. In early August, some galls reached maturation, which lasted until November when the peak of senescent galls was registered (Fig. 5 B). In October of 2009 and 2010, during leaf flushing, two events of induction were observed, reaching infestation rates ??of 30.3 % and 17.2 %, respectively. The second instar nymphs were found in 90 % of the galls from November 2009 to February 2010. In March they were reduced to about 80 %, and reached the lowest percentage in July. The highest percentage of occurrence of the third instar was registered in April (approximately 80 %), while the fourth and fifth instars from May to June and from July to September, respectively. From August to October, adults were found inside the galls, and by the end of this period, the life cycle of the insects restarted (Fig. 6). The first instar nymphs were not counted because they were located on the leaf surfaces and became usually detached during handling and fixation of the leaves. There was a positive correlation between the volume of the galls and the nymph developmental stages (r = 0.89), which formed five groups (Table 2). The development of N. myrtoidis passes through five nymphal stages, taking one year to complete its whole life cycle. Females lay their eggs strictly on the margins of young leaves of P. m y r t o i d e s. The first instar nymphs (Fig. 7 A) emerge and migrate to the limb, where they settle and begin to feed. Inside the galls, the nymphs pass through successive developmental stages until the fifth instar (Figs 7 B–E), which is the predecessor of the adults. These are distinguished from the nymphs mainly by the presence of functional wings (Fig. 7 F). Females can be distinguished from males in the morphology of the terminalia. Females possess a short, cuneate ovipositor (Fig. 7 F, detail on the left) whereas males are characterised by the tubular proctiger and the subglobular subgenital plate with the papameres and aedeagus (Fig. 7 F, detail on the right). Nymphs of Nothotrioza myrtoidis were parasitized by an undescribed species of Galeopsomyia (Hymenoptera: Eulophidae) (Fig. 7 G, arrow). When pupating (Fig. 7 H) the parasitoid ends its feeding activities, having caused the death of the gall inducer. Galeopsomyia sp. uses the gall as shelter during the development period until the adult stage (Fig. 7 I), when it actively digs an escape tunnel, reaching the external environment (Fig. 7 I, detail). The gall inducers presented 15.7 % of parasitism and 29.8 % of mortality, along the insect’s life cycle. A relationship between the red colour of the gall and the non-parasitized condition of the gall inducer was found (χ 2 = 10.67; p = 0.0048) (Fig. 8).Published as part of Carneiro, Renê G. S., Burckhardt, Daniel & Isaias, Rosy M. S., 2013, Biology and systematics of gall-inducing triozids (Hemiptera: Psylloidea) associated with Psidium spp. (Myrtaceae), pp. 129-146 in Zootaxa 3620 (1) on page 138, DOI: 10.11646/zootaxa.3620.1.6, http://zenodo.org/record/22205

Nothotrioza cattleiani Burckhardt, sp. n.

Nothotrioza cattleiani Burckhardt sp. n. (Figs 2 A, D; 3 C, G–I; 4 B, C, F) Neotrioza tavaresi sensu Butignol & Pedrosa-Macedo, 2003, nec Crawford, 1925. Material examined. Holotype male, Brazil: Paraná, Piraquara, 5.x. 2003, Psidium cattleianum (C. A. Butignol) (MZSP, dry mounted). Paratypes. Brazil: 21 males, 28 females, 35 nymphs, same data as holotype (BMNH, ELEF, MHNG, MZSP, NHMB, dry and slide mounted, and in 70 % ethanol); 3 males, 3 females, 3 nymphs, same data but 12.ix. 1997 (NHMB, dry and slide mounted); 2 males, 3 females, 3 nymphs, same data but 12.ix. 1997 (ELEF, NHMB, dry mounted and in 70 % ethanol). Description. Adult. Antennal segment 10 (Fig. 3 C) with longer terminal seta about 3 times as long as and shorter terminal seta slightly longer than segment 10. Forewing (Fig. 2 A) with vein Rs strongly curved; cell cu 1 relatively large and high, distance between apices of veins M 3 + 4 and Cu 1 a around 0.6 times as long as that between apices of veins Cu 1 a and Cu 1 b, and distance between apices of vein Cu 1 a and Cu 1 b about 2.6 times as long as length of vein Cu 1 b; field of radular spinules in cell m 2 distingly shorter than those in cells m 1 and cu 1. Male proctiger (Fig. 3 G) with narrowly rounded posterior margin, more or less evenly narrowing from widest point towards the base. Paramere, in profile, with convex, slightly undulate hind margin (Fig. 3 H). Distal portion of aedeagus with small apical hook (Fig. 3 I). Female proctiger (Fig. 4 B) with blunt tip apically. Fifth instar nymph (Fig. 2 D). Caudal plate (Fig. 4 F) with V-shaped hind margin. Measurements and ratios in Table 1. Etymology. Named after its host species Psidium cattleianum. Distribution. Brazil: Paraná (Butignol & Pedrosa-Macedo, 2003, as Neotrioza tavaresi). Host plant. Psidium cattleianum (Myrtaceae). Gall. Rounded, unilocular gall on the adaxial side of the leaves (Butignol & Pedrosa-Macedo, 2003).Published as part of Carneiro, Renê G. S., Burckhardt, Daniel & Isaias, Rosy M. S., 2013, Biology and systematics of gall-inducing triozids (Hemiptera: Psylloidea) associated with Psidium spp. (Myrtaceae), pp. 129-146 in Zootaxa 3620 (1) on pages 133-134, DOI: 10.11646/zootaxa.3620.1.6, http://zenodo.org/record/22205

Reacquisition of New Meristematic Sites Determines the Development of a New Organ, the Cecidomyiidae Gall on Copaifera langsdorffii Desf. (Fabaceae)

The development of gall shapes has been attributed to the feeding behavior of the galling insects and how the host tissues react to galling stimuli, which ultimately culminate in a variable set of structural responses. A superhost of galling herbivores, Copaifera langsdorffii, hosts a bizarre “horn-shaped” leaflet gall morphotype induced by an unidentified species of Diptera: Cecidomyiidae. By studying the development of this gall morphotype under the anatomical and physiological perspectives, we demonstrate the symptoms of the Cecidomyiidae manipulation over plant tissues, toward the cell redifferentiation and tissue neoformation. The most prominent feature of this gall is the shifting in shape from growth and development phase toward maturation, which imply in metabolites accumulation detected by histochemical tests in meristem-like group of cells within gall structure. We hypothesize that the development of complex galls, such as the horn-shaped demands the reacquisition of cell meristematic competence. Also, as mature galls are green, their photosynthetic activity should be sufficient for their oxygenation, thus compensating the low gas diffusion through the compacted gall parenchyma. We currently conclude that the galling Cecidomyiidae triggers the establishment of new sites of meristematic tissues, which are ultimately responsible for shifting from the young conical to the mature horn-shaped gall morphotype. Accordingly, the conservative photosynthesis activity in gall site maintains tissue homeostasis by avoiding hypoxia and hipercarbia in the highly compacted gall tissues