Between 2009 and 2011, specimens of delphacids (adults and nymphs) were collected by Hernández and Cabrera Walsh (2014) feeding on L. g. subsp. hexapetala. Samples were identified as P. paraguayensis, collection field trips were resumed in 2019 in areas where L. g. subsp. hexapetala is most commonly distributed (from Buenos Aires to the northeastern part of Argentina). Adults and nymphs of P. paraguayensis were collected both, by aspirating individuals from the leaves and stems of field host plants, and laboratory colonies. Specimens were preserved in 70% EtOH for morphological studies. Live adults and immature specimens from the lower Delta were used to describe colour patterns (Table 1).

During summer months (December to March), general culture consisted of outdoor pools (1×1.5 m) with L. g. subsp. hexapetala plants brought from the field and enclosed in square meshed cages. In cooler months (May to August) or if there is no availability of greenhouse conditions, P. paraguayensis can also be reared in small containers inside rearing chambers at FuEDEI facilities. We were able to maintain colonies in two types of containers with different water availability and size. One to three L cylindrical plastic containers with polyester gauze or nylon covers and enough water to irrigate the plant roots (Fig. 1A). Here, 10–20 cm plants can be placed vertically but insects are harder to observe or retrieve. The second device consisted of 20×15×5 cm rectangular plastic containers with pierced plastic lids and polyester gauze. Stems were placed horizontally and inside water picks for irrigation (Fig. 1B). Excess condensation on the walls is not recommended.

Figure 1. 

Different rearing and test devices for Pissonotus paraguayensis A device for aquatic plants utilised for rearing and host range tests B device for rearing and tests with plants in water-pics C small petri dishes with L. g. susp. hexapetala utilised in life cycle study D hanging plastic cylinders utilised in host range test for terrestrial plants.

Identification of P. paraguayensis was based on the original description and photographs of the holotype from the Arizona State University, Lois B. O’Brien Collection (ASU: ASULOB). Female (brachypters and macropters) specimens, captured in the same place and host plant were structurally matched with available males, then used for dissections, images and complementary descriptions are deposited in the Museo de La Plata entomological collection (MLP). Immature stages: description was based on 24-hour-old nymphs from laboratory colonies. Specimens were submitted to 95% ethyl ether to preserve colour, cleared in cold 10% KOH solution and fixed in Faure liquid for microscopic examination and illustration. Eggs: obtained by dissecting oviposition scars. First instar: described in detail, focused on major differences are highlighted for older stages and also between presumptive brachypterous and macropterous forms of the fifth instar. Measurements: range and median, given in mm, taken from anaesthetised specimens. Dimensions were expressed as L.: total body length (from the tip of vertex to the tip of abdomen), W.: maximum body width (across the widest part of the mesothorax), t.l.: thoracic length (from the anterior margin of the pronotum to the posterior margin of the metanotum along the midline). Other measurements are relative. Morphological terminology and measurements of adults follow Bartlett and Deitz (2000), but the wing venation and female genitalia refer to Bourgoin et al. (2015) and Asche (1985) respectively. Carination and arrangement of pits of nymphs generally follows Vilbaste (1968) updated after by Yang and Yeh (1994). External morphology was observed with a Leica EZ5 stereomicroscope and photographs were taken with an adapted RRID 18 HD digital camera and a Canon EOS 90D. Illustrations were imported into Adobe Photoshop 7.0 for labelling and plate composition.

DNA was extracted from whole bodies of adults using Qiagen DNeasy Blood & Tissue Kit. A fragment of 658 bp of the cytochrome c oxidase I (COI) gene was amplified using universal barcoding primers LCO1490 and HCO2198 (Folmer et al. 1994). PCR amplification was done in a 25 μL volume using the following thermocycling protocol: 2 min at 95 °C; 5 cycles of 40 secs at 94 °C, 40 secs at 45 °C, 1 min at 72 °C; 35 cycles of 40 secs at 94 °C, 40 secs at 51 °C, 1 min at 72 °C; 5 min at 72 °C; held at 4 °C. PCR products were checked in 1% agarose gels and purified by adding 0.5 μL (10 u) Exonuclease I (Exo I) and 1 μL (1 u) Shrimp Alkaline Phosphatase (SAP). Samples were incubated at 37 °C for 15 min and reaction was stopped by heating the mixture at 85 °C for 15 min. Both strands of each fragment were sequenced using Sanger technology in Macrogen Inc. sequencing service (South Korea) with the same primers used for PCR amplification. Sequence quality check, primer trimming, and alignment were performed on CodonCode Aligner version 10.0.2. Sequences were deposited in GenBank after running a BLAST search to check for possible contaminations.

Tests were performed inside rearing chambers (25 °C, 12 hrs. light/12 hrs. dark). Mature couples (n = 10) were collected from the colony and placed individually in rectangular plastic containers (20×15 cm) with fine pierced lids. Inside, a 10-cm long stem of L. g. subsp. hexapetala with a water pic was exposed for 24 hrs to each couple (Fig. 1B). Only brachypterous females were used for this test since macropterous females might have different reproductive behaviour. After the exposure, the number of oviposition scars per stem were recorded and plants were stored again in the rearing chamber until nymph emergence. Development time of eggs and the number of emerged nymphs per stem after 24 hrs of exposure were recorded. Also, laboratory-reared plants were dissected to quantify the number of eggs per oviposition scars.

To follow the developmental time of the immature stages, newly emerged nymphs (n = 50) were randomly selected from the previously mentioned containers and placed individually in Petri dishes with 5-cm long stems and wet tissue paper (Fig. 1C). Nymphs were observed every 1–2 days and developmental time for each instar was recorded by checking for exuviae. All means are reported with ±SD.

To estimate the host range, no-choice tests were performed in a greenhouse with controlled temperatures (26 °C day/24 °C night) and natural light conditions (summer). Because the planthopper is a sap-feeding insect, whole plants were utilised. Depending on the plant species life form, two different devices were used. If the plant was terrestrial or did not resist/withstand stems being cut to be enclosed in small containers, hanging acetate cylinders with nylon mesh on both ends were placed containing ~ 15 cm of a stem (Fig. 1D). Each cylinder was placed on independent plants. If plants were aquatic, then a single 15-cm long stem was placed inside a 3-lt plastic container with water and polyester gauze as a lid (Fig. 1A). For each plant species, between three and five repetitions were carried out, depending on plant availability. Inside each device, one male and one brachypterous female of P. paraguayensis were released to test survival and oviposition. For survival, the number of days elapsed from the beginning of the experiment until the death event of both males and females, were recorded. Two types of feeding controls were set up: one with the insects on the known host plant, L. g. subsp. hexapetala as a positive control, and another with insects only with wet tissue paper as a negative control with no food. The experiment was terminated at 52 days. For the test plant list, nine Ludwigia species were considered, six from the Jussiaea section, to which the main host plant belongs (L. grandiflora (Michx.) Greuter & Burdet subsp. grandiflora, L. g. subsp. hexapetala, L. peploides (Kunth) P. H. Raven subsp. peploides, L. p. subsp. montevidensis (Spreng.) P. H. Raven, Ludwigia peploides (Kunth) P. H. Raven subsp. glabrescens (O. Kuntze) P. H. Raven and Ludwigia hookeri (Micheli) H. Hara; two from the Macrocarpon section, Ludwigia bonariensis (Micheli) Hara and Ludwigia neograndiflora (Munz) Hara; and one from the Myrtocarpus section, Ludwigia serícea (Cambess.) Hara. Two other species, one closely related species from another genus of Onagraceae, Oenothera affinis Cambess. and one that is not closely related but commonly co-exist in the natural habitat, Myriophyllum aquaticum (Vell.) Verdc were also evaluated.

Survival data of females was analysed through Kaplan-Meier survival curves using the ‘survfit’ and ‘ggsurvplot’ functions from Survival package (ver. 3.4-0) of RStudio statistical program (ver. 2023.06.1). A non-parametric Log-rank statistical test was used to assess statistical differences in overall survival. To better understand the influence of each test plant in P. paraguayensis survival, the Cox regression model was employed to estimate the hazard ratio (HR) and P values based on χ² test, and 95% confidence intervals (95%-CI). HR is the rate of occurrence of an event during a given time interval, where HR > 1 means that exposure to the factor increases the rate of occurrence of the event, and HR < 1 decreases the rate. If the HR = 1 we say that the factor behaves as the reference (Kleinbaum and Klein 2012). Forest Plot for Cox proportional hazards model was created using the ‘coxph´ and ‘ggforest’ functions of R packages survival and survminer (0.4.9). P < 0.05 defined significance. All analyses were conducted with R 4.3.1 (R Core Team 2022).

Adult

This species was described in detail by the author from a brachypterous male from Paraguay; the brachypterous female was briefly discussed, referring only to its body size with some observations on the variations in colour patterns among specimens from different geographic locations (Bartlett and Deitz 2000).

Male (Fig. 2). Although the brachypterous male has been fully described and illustrated by the author, we added representative images of that morph in order to facilitate its association and recognition, given the dimorphic sexual coloration of this species.

Figure 2. 

Pissonotus paraguayensis. Habitus. Brachypter male A dorsal view B ventral view.

Female (Figs 3, 4). Note: As Bartlett and Deitz (2000) noted, females of Pissonotus have been little used in taxonomy probably due to the scarce availability of specimens reliably associated with males. Illustrations and complementary comments not included in the original description, particularly on variation in colour between or within conspecific populations and descriptions of some structural features of the female genitalia are given for the first time.

Figure 3. 

Pissonotus paraguayensis. Habitus. Brachypter female A dorsal view B ventral view. Macropter female C dorsal view D ventral view.

Figure 4. 

Pissonotus paraguayensis female A abdomen, genital region, ventral view B pregenital sternite C gonapophysis IX D apex of gonapophysis IX E apex of hind leg (post tibial spur and tarsi) of adults.

Brachypter. Body length: 2.91–3.16 (3.01, n = 5); tegmina length: 0.80–0.90 (0.82, n = 4) (Fig. 3A, B).

Colouration : In dorsal view body generally dark brown, paler on vertex and along midline of abdomen and widening above genital and anal segments. Vertex and upper frons are mostly brownish with carinae and anterior margin paler. Tegmina dark brown with a distinguishable distal transverse narrow white band, sometimes medially incomplete; veins concolourous with venation obscure becoming obsolete toward apex (not reticulate). In ventral view, head with a distinctive transverse whitish band on epistomal margin of frons, which extends laterally along the genae to the base of the pre- and mesocoxae; blackish brown area over postclypeus and basal ½ of coxae contrasting with the yellowish colouration of anteclypeus, rostrum, thoracic sternites and legs. Apex of rostrum, longitudinal anterior highlights on antennomeres I and II and dorsal margins of femora and tibiae, and the apex of last tarsal segments and ungues, black. Abdomen paler along midventral line from sternum V to the caudal region.

Structure : Antennal foveae elevated, placed close to ventral margin of the compound eye. Tegmina quite longer as broad (6:1), distally rounded on leading margin and apically truncate, not exceeding tergite II. Calcar slender, moderately foliaceous; shorter than basitarsus (0.8:1), usually with a row of 13–15 black tipped teeth on ventral margin with 2–4 smaller teeth irregularly spaced external to the row (Fig. 4E).

Macropter. Total length (including wings): 3.66–3.70 (3.68, n = 4). Body length (excluding wings): 3.00 (3.00, n = 4), fore-wings length: 2.90 (n = 4). (Fig. 3C, D).

Colouration : Similar to brachypterous form in colour pattern of head, antennae, sternothorax, and legs. General body colouration uniformly darker; with distinctive yellowish areas: dorsally, restricted on caudal apex of mesonotum, base of tegula and anal segment, and ventrally, on the posterior margin of abdominal sternites, pregenital sternite, inner margins of gonocoxa VIII and gonapophysis VIII. Wings semi-hyaline, fore-wing veins light brown, proximal portion of remigium and clavus and some longitudinal and transverse veins, quite dark.

Structure : Similar to the brachypterous form, with mesothorax more robust. Wings fully developed, longer than abdomen, fore-wings extending ~ 1/3 of their length beyond the tip; hind-wings shorter. Fore-wing with a simple venation pattern, with scattered small seta-bearing tubercles irregularly distributed on the veins. Remigium veins mainly non-bifurcated, nodal cells poorly defined, sometimes absent, with four long postnodal cells distally open, CuP apically weakened; pre and post nodal cells elongated; clavus in angle, postclaval margin forming an obtuse angle (100–120°).

Genitalia (Fig. 4A–D): Ovipositor sclerotised and relatively long and narrow, shortly surpassing the posterior margin of anal segment. Pregenital sternite subtriangular, elongate, longer in middle line than its wide at its widest part (1.5:1), anterior margin strongly narrowed “like a neck” and lightly rounded at apex (Fig. 4A, B). Gonocoxa VIII, more or less lanceolate, regularly expanded at base; as long as 1/3 the length of the gonapophyses VIII (Gy VIII) which is uniformly slender and elongated. Gonapophyses IX slender, slightly curved, wider at basal 1/3, distal 1/3 with minute teeth on dorsal margin (Fig. 4C, D); dorsal gonoplac (Gp) or third valvula uniformly wide. Anal style moderately short.

Host Plant. In the field, adults and nymphs were observed feeding on Ludwigia grandiflora subsp. hexapetala.

Recorded natural enemies. None.

Distribution. South Central South America: Brazil, Paraguay and apparently Bolivia (Bartlett and Deitz 2000). Argentina: Santa Fé, Entre Ríos and Buenos Aires.

Pissonotus paraguayensis was found in wetlands of central Argentina that belong to the Del Plata Basin, expanding its geographic range to central-east of Argentina and now making Buenos Aires province its southernmost limit of distribution in America (Table 1).

DNA barcode. The two sequenced specimens (1♀b, 1♂b) from Brazo Largo, Entre Ríos (33°51'53.4"S, 58°52'59.4"W) had the same 658 bp COI haplotype, which was deposited in GenBank under accession number OR523788.

Material examined. Argentina: 6♀b, 1♀m, Buenos Aires, Otamendi, 34°13'04"S, 58°55'52"W, s/ L. g. subsp. hexapetala, 21-01-2019, Faltlhauser col.; 10♂, same place and host, 10-03-2019, Hernández col.; 15 ♂, same place and host, 07-06-2019, Hernández col.; 1 ♀m, 8-10-2008, (named 1 ♀ Delphacidae), Cabrera Walsh col.; 15♂, 3♀b, laboratory reared (FuEDEI), 7-06-2019, Hernández; 5♀m, 2♀b, 1♂b, Buenos Aires, Magdalena, 35°3'48.9"S, 57°33'14.9"W, s/ L. g. subsp. hexapetala, 3-11-2021, Faltlhauser col.

Remarks. Although the colour pattern is useful to recognise members of the genus, some variations were observed among brachypterous specimens of both sexes. General colouration varies from black to blackish-orange, particularly the distal transverse white band of tegmina that may be complete, medially incomplete, or exceptionally absent, and the dorsal surface of the abdomen with more diffuse colouration towards the caudal region. Nevertheless, the distinctive pattern colouration described on the head and thorax is rather uniform in both sexes and morphs. The drumming organ in males shows a dimorphic distinctive yellowish colouration. Macropterous forms show more homogeneous colouration among specimens.

Immature stages. Fig. 5.

Figure 5. 

Immature stages in dorsal view of Pissonotus paraguayensis A first instar C second instar E third instar G fourth instar I fifth instar (presumptive brachypter form). Apical portion (from tibiae) of hind leg B first instar D second instar F third instar H fourth instar J fifth instar (presumptive brachypter form). Scale bars: 0.5 mm (A); 1mm (C, E, G. I).

Description: Eggs. (Fig. 7A) (n = 10). Length: 0.86, width: 0.23.

Eggs milky white, ellipsoidal. Chorion translucent.

First Instar. (Fig. 5A, B) (n = 5). L:1.04 (0.93–1.07) W: 0.33 (0.29–0.36); t.l: 0.35 (0.25–0.41).

Uniform whitish body; with distinctive red eyes; uniform greyish antennal segments.

Form elongate, subcylindrical, dorsoventrally slightly flattened. Vertex subquadrate, with lateral margins often divergent in front and behind the eyes; length sub-equal to width (1.2:1); fastigium (lateral view) rounded; lateral and submedian carinae of vertex extending onto frons. Frons with lateral margin slightly convex and carinate, ~ 2× longer than wide in midline, regularly wider towards the frontoclypeal suture; lateral margins carinate and paralleled by submedian carina which is prominent at base and regularly evanescent towards apex; each laterofrons at level below the eyes 1.5× wider than interfrons; with 13 sensorial pits on each side: nine pits on area between submedian and lateral carinae (six visible in ventral view, three in dorsal view); dorsally four pits between lateral carina and eyes. Antennae three-segmented; scape, short, slightly wider than long, antennal pedicel sub-cylindrical ~ 2–2.5× the length of scape, without sensory pits; base of flagellum bulbous ~ 0.5× the pedicel length. Rostrum reaching the posterior margin of metacoxae, apical segment 0.6× longer than subapical.

Thoracic nota divided into three pairs of plates by longitudinal mid-dorsal line. Pronotal plates subtrapezoidal, lateral carinae divergent and slightly convex towards posterior margin. Each plate with seven pits extending from near middorsal line posterolaterally to lateral margin (two pits in line next to lateral carinae on posterior ½ of segment and five pits extending laterally from the external side of carina along the posterior border of plate, one proximal to it and four most lateral not visible in dorsal view). Mesonotum as long as metanotum. Mesonotal plate with four pits, two median on disk, and two next to lateral margin. Pro, meso, and metatarsi with two tarsomeres; apical tarsomeres subconical, curved, with a pair of apical claws. Metatibiae without lateral spines, with apical transverse row of four stiff spines; tiny spur < 2× the length of longest apical spines of the metatibia and ~ 1/4 length of metatarsomere 1. Metatarsomere 1 shorter than metatarsomere 2 (0.5:1) with four apical spines.

Abdomen with nine apparent segments, widest across segment 5. Tergite 5 with one pit and tergites 6–8 each with three pits on either side of the midline. Segment 9 surrounds anus with three pits on each side, two subapical and one dorsal median.

Second Instar. (Fig. 5C, D) (n = 5). L: 1.57 (1.54–161); W: 0.61 (0.61–0,73); t.l.: 0.48 (0.44–0.52).

Pale yellowish body, with distinctive black markings on the head and legs: a stripe on apex of vertex and upper frons extended at lower level of the eyes, along the frontal surface of antennal segments I and II, and several marks limited to: upper border of antennal fovea, apex of rostrum, outside apices of femoro-tibial joints, base of basitarsus and apices of legs.

Antennal foveae elevated, placed under the compound eyes, close to its lower margin; antennal pedicel sub-cylindrical ~ 3× the length of scape, with three small pits: two on apical margin and the third subapical on posterior surface; flagellum whip-like distally, bulbous at base, ~ 0.25× the pedicel length. Rostrum extends almost to bases of metacoxae, apical segment 0.2× shorter than subapical. Pro, meso, and metanotum subequal, pronotum slightly shorter (0.7:1); posterior margin of mesonotum straight at middle, metanotum deeply excavated. Wing-pads undeveloped. Metatibiae with two small lateral spines (one near base and one in basal ½), and five stiff spines on the apical transversal row; spur ~ 1/4 length of metatarsomere 1, with two similar sized teeth, one on lateral aspect added to the apical. Metatarsomere 1 as long as metatarsomere 2, with apical transversal row of five spines.

Third instar. (Fig. 5E, F) (n = 4). L: 1.78 (1.62–1.81); W: 0.89 (0.85–0.93)); t. l.: 0.56 (0.50–0.62).

Similar pattern colour to second instar, with darker markings.

Antennal pedicel sub-cylindrical, ~ 3× the length of scape, with four to five sensory pits; flagellum with bulbous portion ~ 0.1 the pedicel length. Rostrum overlapping the bases of metacoxae, apical segment 0.3× shorter than subapical.

Mesonotal plate with the two discal pits on both sides of the lateral carinae. Fore-wing pads short, each covering 1/2 of metanotal segment laterally, with three pits, a pair on costal area and another near middle. Metanotum plates median length as long as mesonotum, shallowly excavated on posterior margin. Metatrochanter subcylindrical, with row of ten or eleven interlocking flattened folds on posteromedial aspect. Metatibial spur 3/4 the length of metatarsomere 1, with one apical tooth and two to three teeth on lateral margin. Metatarsomere 1 with apical transverse row of five spines; apical tarsomere with an arrange of fine sensory hairs along it.

Fourth instar. (Fig. 5G, H) (n = 10). L: 1.80 (1.69–1.89); W: 0.87 (0.73–1.00); t. l: 0.55 (0.51–0.56).

Colour pattern similar to former instar. Most specimens with conspicuous brownish marking along dorsal portions of thoracic nota continuing onto wings pads and abdomen.

Antennal pedicel sub-cylindrical ~ 2× the length of scape, with approx. five or six pits on the apical ½ (Fig. 5B). Rostrum extending to bases of metacoxae, apical segment slightly shorter than subapical.

Fore-wing pads short, each covering ~ 1/2 of hind wing pad, laterally with three pits, one on costal area and two next to subcostal carina. Metanotum median length sub-equal to mesonotum; hind-wing pad oval, reaching the base of the first abdominal segment. Metatibial spur almost as long as the metatarsomere 1 in middle, with one apical tooth and three or four on lateral margin. Metatarsi with two tarsomeres: tarsomere 1 with apical transverse row of six spines, tarsomere 2 subconical, similar to apical tarsomeres of other legs, slightly shorter than length of tarsomere 1, partially subdivided in middle, with three short ventrolateral spines in middle of plantar surface.

Fifth instar. Presumptive brachypterous form (Fig. 5I, J) (n = 10) L: 2.27 (2.18–2.36); W: 0.99 (0.96–1.01); t. l: 0.65 (0.62–0.70). Presumptive macropterous form (Fig. 6A, B) (n = 2). L: 2.3; W: 1; t.l; 0.9.

Figure 6. 

Fifth instar (presumptive macropterous form) Pissonotus paraguayensis A dorsal view B ventral view.

Colour pattern similar to former instars but darker. Dorsally blackish mottled with light to dark brown and cream on head, thorax and abdomen, with notorious yellowish pits; ventrally, head with postclypeus also darkened at base. Obvious black mark on legs: on base of coxae, outside apices of femoro-tibial joint, longitudinal stripes on dorsal margins of femora and tibiae, basal tarsomere of pro and mesotarsi and the apex of last tarsal segments and claws.

Form elongate, sub-cylindrical, widest across mesothoracic wing pads. Head lightly protruding beyond the anterior margin of eyes, ~1/4 of the eyes length. Vertex subquadrate, length slightly longer than wide (1:07), anterior margin straight, posterior convex, basal compartments shallowly concave; lateral and submedian carinae prominent, continuing onto frons from the level of middle of the eyes to clypeal margin. Frons sub-rectangular; widest at middle, width. ~ 0.9× the length; carinate lateral margins slightly convex, these lateral carinae extending from vertex to clypeal border and paralleled by a pair of straighter submedian carinae regularly evanescent towards clypeal border. Clypeus narrowing distally, consisting of subconical basal postclypeus not carinated along its length, and cylindrical distal anteclypeus. Antennae with scape cylindrical, length subequal to width; pedicel subcylindrical, ~ 3× longer than scape, with nine or ten pits on the apical ½. Rostrum surpassing mesocoxae, apical segment slightly longer than apical (1.2:1).

Pronotal plates sub rectangular; anterior margin follows posterior margin of head, posterior border almost straight; each plate with straight short posterolaterally directed carina originating on anterior margin in median 1/3 and terminating near middle of plate, carina bordered along inner margin by row of six pits extending posterolaterally to lateral border of plate (three pits in line next to lateral carinae and three near posterior margin; one lateralmost pits not visible in dorsal view). Mesonotal median length ~ 1.5× that of pronotum; each plate bearing an elongate lobate wingpad covering lateral ½ of metanotal wingpad; with posterolaterally directed carina originating on anterior margin in median 1⁄4 and terminating on posterior margin, with five pits: two on notum, one on each side of carina, and three laterad (two on wingpad). Metanotal median length ~ 0.75× that of mesonotum; each plate bearing an elongate lobate wingpad extending to tergite 2; with weak longitudinal carina originating on anterior margin in median terminating near posterior margin; one pit on wingpad, just lateral to carina. When developing to macropterous adults the fore-wing pads extend up to the apices of the hind pads, reaching tergite 5.

Pro- and mesocoxae elongate, posteromedially directed; metacoxae fused to sternum. Metatrochanter subcylindrical, with a row of 12–14 flattened folds on the posteromedial aspect which interlocks with those on the adjoining trochanter. Metatibiae with subtriangular, slender, moderately foliaceous spur, a little shorter than metatarsomere 1, with a row of seven-nine teeth on lateral margin and one apical tooth. Metatarsomere 1, longer than tarsomere 2 plus 3 (1:05), with apical transverse row of seven black-tipped spines on plantar surface; tarsomere 2 cylindrical, with apical transverse row of four black-tipped spines on plantar surface; tarsomere 3 sub conical, similar to apical tarsomere of other legs.

Abdomen with nine apparent segments, slightly flattened dorsoventrally, widest across segment 5. Segment 9 surrounds anus with three evident pits on each side, two subapical and one dorsal median, in both sexes.

Specimens examined. Argentina: Otamendi, Buenos Aires (34°03'48.2"S, 58°49'19.59"W), 10 nymphs V, 10 nymphs IV, 10 nymphs III, 10 nymphs II and 10 nymphs I, 07-06-2019, on L. g. subsp. hexapetala, Faltlhauser col.; 2 nymphs III, 1 nymph I; 21-I-19, on L. g. subsp. hexapetala, Faltlhauser col.

The authors have declared that no competing interests exist.

No ethical statement was reported.

La Plata National University (UNLP) (Grant N° 730); Fundación para el Estudio de Especies Invasivas (FuEDEI); Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET).

Conceptualization: AMMRL, AJS, ACF, MCH. Data curation: AMMRL, NAS, ACF. Formal analysis: AMMRL, AJS, ACF. Funding acquisition: AMMRL, AJS. Investigation: MCH, ACF, AJS, AMMRL, AF. Methodology: ACF, MCH, AMMRL. Project administration: ACF, AMMRL, AJS. Resources: AJS, AMMRL. Software: ACF, NAS, AF. Supervision: MCH, AJS, AMMRL. Validation: ACF, MCH. Visualization: ACF, AMMRL. Writing - original draft: ACCF, MCH, AJS, AMMRL. Writing - review and editing: NAS, AJS, AMMRL, MCH, AF, ACF.

Ana M. Marino de Remes Lenicov https://orcid.org/0000-0001-8678-5408

Ana C. Faltlhauser https://orcid.org/0000-0003-3121-3650

Alvaro Foieri https://orcid.org/0000-0001-5623-3629

Nicolas A. Salinas https://orcid.org/0000-0002-1055-4475

M. Cristina Hernández https://orcid.org/0000-0003-4382-773X

Alejandro J. Sosa https://orcid.org/0000-0002-1680-8712

All of the data that support the findings of this study are available in the main text or Supplementary Information.