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Description of the immature stages of the flea beetle Omophoita personata (Coleoptera: Chrysomelidae: Galerucinae: Alticini)

ABSTRACT

Alticini is the most diverse tribe of Chrysomelidae with approximately 8,000 species. Despite of its high diversity, little is known about their natural history and immature stages. Herein, we describe the immatures of Omophoita personata (Illiger, 1807) reared in laboratory from adults sampled in field. We also investigated and compared with immatures of O. octoguttata (Fabricius, 1775). Detailed morphology and chaetotaxy are presented. Larvae of O. personata have their bodies covered with tubercles and the stemmata are absent. These characteristics are shared with other Oedionychina species, reinforcing the stability of these morphological characteristics as diagnostic of this subtribe. This study provides important descriptive and comparative data that increases the knowledge about Alticini immatures.

KEY WORDS:
Larvae; larval morphology; Oedionychina; pupa description; taxonomy

INTRODUCTION

Alticini are an important component of phytophagous entomofauna in different environments. This tribe has approximately 8,000 species in about 534 genera distributed around the world (Nadein and Bezděk 2014Nadein K, Bezděk J (2014) Galerucinae Latreille, 1802. In: Leschen RAB, Beutel RG (Eds) Handbook of Zoology. Coleoptera, beetles. Morphology and systematics. Walter de Gruyter, Berlin/Boston, vol. 3, 251-259.), with 1,410 species recorded for Brazil (Linzmeier 2022Linzmeier AM (2022) Alticini in Catálogo Taxonômico da Fauna do Brasil. PNUD PNUD http://fauna.jbrj.gov.br/fauna/faunadobrasil/119960 [Accessed: 21/02/2022]
http://fauna.jbrj.gov.br/fauna/faunadobr...
). Despite all this diversity, their biology is poorly known, especially when considering the immatures stages. However, there are a few studies describing the immatures of different species (e.g., Virkki and Zambrana 1983Virkki N, Zambrana I (1983) Life history of Alagoasa bicolor (L.) in indoor rearing conditions (Coleoptera, Chrysomelidae, Alticinae). Entomologische Arbeiten aus dem Museum Georg Frey 31(32): 131-155., Casari and Duckett 1997Casari SA, Duckett CN (1997) Description of immature stages of two species of Pseudolampsis (Coleoptera: Chrysomelidae) and the establishment of a new combination in the genus. Journal of the New York Entomological Society 105: 50-64., Lee et al. 1998Lee JE, Lingafelter SW, Konstantinov AS (1998) Larval morphology of Systena blanda Melsheimer (Coleoptera: Chrysomelidae: Alticinae). Proceedings of the Entomological Society of Washington 100: 484-488., Duckett and Moyá 1999Duckett CNA, Moyá S (1999) New species of Ptocadica Harold (Coleoptera: Chrysomelidae, Alticini) from Costa Rica and Panama. The Coleopterists Bulletin 53(4): 311-319., Biondi and Nardis 1999Biondi M, Nardis G (1999) Morphological description of the first-instar larva of Derocrepis sodalis (Coleoptera, Chrysomelidae, Alticinae). Italian Journal of Zoology 66(4): 393-397., Duckett and Swigoňová 2002Duckett CN, Swigoňová Z (2002) Description of immatures stages of Alagosa januaria Bechyné (Coleoptera: Chrysomelidae). Journal New York Entomological Society 110(1): 115-126., Duckett and Casari 2002Duckett CN, Casari SA (2002) First descriptions of larval stages of Walterianella bucki Bechyné (Coleoptera: Chrysomelidae: Alticini) and notes on life history. The Coleopterists Bulletin 56(2): 170-181., Lesage and Zmudzinska-Krzesinka 2004Lesage L, Zmudzinska-Krzesinska A (2004) The immature stages of the grape flea beetles Altica chalybea Illiger and A. woodsi Isely (Coleoptera, Chrysomelidae). In: Jolivet P, Santiago-Blay JA, Schmitt M (Eds) New developments in the biology of Chrysomelidae. SPB Publishing Academic Publishing, The Hague, 503-528., Linzmeier et al. 2007Linzmeier AM, Ribeiro-Costa CS, Moura LA (2007) First descriptions of immatures for Megistops (Boheman) (Coleoptera, Chrysomelidae, Galerucinae) in a new host-plant family, with notes on life history and redescription of M. vandepolli Duvivier. Zootaxa 1615(1): 55-68., Casari and Teixeira 2011Casari SA, Teixeira EP (2011) Immatures of Syphrea uberabensis guerini Bechyné (Coleoptera, Chrysomelidae, Alticini). Revista Brasileira de Entomologia 55(1): 17-26., Hua et al. 2013Hua Y, Ge S, Beutel RG, Zhang Y, Cui J, Yang X (2013) The larvae of Altica koreana (Oglobin) and A. viridicyanea (Baly) (Coleoptera: Chrysomelidae: Galerucinae: Alticini). Zootaxa 3694(5): 461-470. https://doi.org/10.11646/zootaxa.3694.5.4
https://doi.org/10.11646/zootaxa.3694.5....
, Cabrera et al. 2016Cabrera N, Sosa A, Telesnicki M, Julien M (2016) Morphology of juvenile stages of Kuschelina bergi (Harold) with biological information (Coleoptera, Chrysomelidae, Alticini). ZooKeys 561: 51-61., Konstantinov et al. 2019Konstantinov AS, Linzmeier AM, Morais ACC, Palmer MW, Scheffer SJ, Lewis ML (2019) Discovery of the first Nearctic moss-eating flea beetle, Distigmoptera borealis Blake, 1943 (Coleoptera: Chrysomelidae: Galerucinae: Alticini). The Coleopterists Bulletin 73(3): 599-610.).

Alticini’s adults have a wide variety of shapes, colors and habits, and species of Oedionychina are among the largest, most colorful, and conspicuous. This subtribe is considered monophyletic and has more than 600 species in 23 genera distributed in the New World, mainly in the Neotropical Region (Ducket and Kjer 2003Duckett CN, Kjer KM (2003) Cladistic analysis of the Oedionychines of Southern Brazil (Galerucinae: Alticini) based on two molecular markers. In: Furth DG (Ed.) Special topics in leaf beetle biology. Proceedings of the Fifth International Symposium on the Chrysomelidae. Pensoft Publishers, Sofia, 117-132.). They are easily recognized by their smooth or confusedly punctuated elytra, hind legs with femur generally well developed and apical metatarsomere globose (Scherer 1983Scherer G (1983) Diagnostic key for the Neotropical alticine genera (Coleoptera: Chrysomelidae: Alticinae). Entomologische Arbeiten aus dem Museum Georg Frey 31: 1-89.). Among them Omophoita Chevrolat, 1836 has approximately 134 species distributed mainly in Central and South America (Blackwelder 1944Blackwelder RE (1944) Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America. Bulletin of the United States National Museum 185(1-6): 1-1492., Scherer 1983Scherer G (1983) Diagnostic key for the Neotropical alticine genera (Coleoptera: Chrysomelidae: Alticinae). Entomologische Arbeiten aus dem Museum Georg Frey 31: 1-89., Ducket and Kjer 2003Duckett CN, Kjer KM (2003) Cladistic analysis of the Oedionychines of Southern Brazil (Galerucinae: Alticini) based on two molecular markers. In: Furth DG (Ed.) Special topics in leaf beetle biology. Proceedings of the Fifth International Symposium on the Chrysomelidae. Pensoft Publishers, Sofia, 117-132.), of which 83 are recorded from Brazil (Linzmeier 2022Linzmeier AM (2022) Alticini in Catálogo Taxonômico da Fauna do Brasil. PNUD PNUD http://fauna.jbrj.gov.br/fauna/faunadobrasil/119960 [Accessed: 21/02/2022]
http://fauna.jbrj.gov.br/fauna/faunadobr...
).

Despite the diversity of this genus, few studies have addressed the natural history of its species or have sought to understand its role in natural environments. A few studies deal with host plants (Begossi and Benson 1988Begossi A, Benson WW (1988) Host plants and defense mechanisms in Oedionychina (Alticinae). In: Jolivet P, Petitpierre E, Hsiao TH (Eds) Biology of the Chrysomelidae. Kluwer Academic Publishers, Dordrecht, 57-71., Jolivet 1991Jolivet P (1991) Sélection trophique chez les Alticinae (Coleoptera Chrysomelidae) (suite). Bulletin Mensuel de la Société linnéenne de Lyon 60(2): 53-72.), mimicry complex (Del-Claro 1991Del-Claro K (1991) Notes on mimicry between two tropical beetles in south-eastern Brazil. Journal of Tropical Ecology 7: 407- 410.), chromosomal characterization (Almeida et al. 2009Almeida MC, Campaner C, Cella DM (2009) Cytogenetics of four Omophoita species (Coleoptera, Chrysomelidae, Alticinae): a comparative analysis using mitotic and meiotic cells submitted to the standard staining and C-banding technique. Micron 40(5): 586-596., 2010Almeida MC, Goll LG, Artoni RF, Nogaroto V, Matiello RR, Vicari MR (2010) Physical mapping of 18S rDNA cistron in species of the Omophoita genus (Coleoptera, Alticinae) using fluorescent in situ hybridization. Micron 41(7): 729-734., Mello et al. 2014Mello LRA, Tasior D, Goll LG, Artoni RF, Vicari MR, Nagaroto V, Almeida MC (2014) Physical map of repetitive DNA and karyotype evolution in three species of the genus Omophoita (Coleoptera: Alticinae). Italian Journal of Zoology 81(1): 16-24., Rosolen et al. 2018Rosolem LAM, Vicari MR, Almeida MC (2018) Accumulation of transposable elements in autosomes and giant sex chromosomes of Omophoita (Chrysomelidae: Alticinae). Cytogenetic and Genome Research 156(4): 215-222.), and just a recently published paper give some data about larval morphology and biological aspects of Omophoita octoguttata (Fabricius, 1775) (Begha et al. 2021Begha BP, Santos MH, Prado LR (2021) Redescription of Omophoita octoguttata (Coleoptera: Chrysomelidae) and its immature stages, with notes on life history. Iheringia, Série Zoologia 111: 1-9.).

Therefore, considering the lack of information regarding immatures of this genus, is important to provide such data that can help in the understanding of its evolutionary history, as well as on the economic and ecological aspects of Alticini. Finally, this study aims to describe the external morphology and chaetotaxy of the immatures of Omophoita personata (Illiger, 1807), comparing it with other Alticini species.

MATERIAL AND METHODS

Immatures were obtained from adults sampled in field trips made in 2019 and 2020 at municipality of Realeza (-25.761349, -3.553202), state of Paraná, Brazil and reared in laboratory (26 ± 5 °C; 55-65% humidity, and natural light). The adults (couples) were maintained in transparent plastic containers (1000 ml) with a branch of the host plant - Ocimum carnosum (Spreng.) Link & Otto ex Benth. (Lamiaceae). After oviposition eggs were transferred to a new container with 1 cm of sieved and autoclaved soil (to avoid contamination by pathogens), and slightly moistened with distilled water, since this species pupate inside a cocoon in the soil. Adults and larvae were feed with the host plant and observed daily.

Immatures were killed and preserved in AGA (1-part acetic acid 100%; 1-part glycerin; 3-parts distilled water; 7-parts alcohol 100%), with specimens of each immature stages packed and labeled separately.

The external morphology of immatures and chaetotaxy were studied using stereomicroscope and microscope. Specimens were measured with a digital pachymeter. The length and width of larvae and pupa were measured in dorsal view immediately after killed. The eggs were measured before they were fixed in alcohol. The mean size is presented followed by the lowest and highest value and the number of specimens measured.

The first and third instars larvae, pupa, as well as some important taxonomic structures, were illustrated in ink using a stereomicroscope, equipped with camera lucida. The illustrations were scanned and vectorized with the Adobe Illustrator and Paint Tool Sai software. The terminology follows Costa et al. (1988Costa C, Vanin SA, Casari-Chen SA (1988) Larvas de Coleoptera do Brasil. FAPESP, São Paulo, 282 pp.) and Duckett and Swigoňová (2002Duckett CN, Swigoňová Z (2002) Description of immatures stages of Alagosa januaria Bechyné (Coleoptera: Chrysomelidae). Journal New York Entomological Society 110(1): 115-126.).

Although the material of the three larval instars were available, the description is based on the third instar larva because the first and second instars are very similar to the third. Thus, differences observed among instars are presented in the remarks section. Furthermore, as we had available immatures of O. octoguttata reared in the laboratory they were also analyzed, permitting the direct comparison between species.

Voucher specimens of both species are deposited in the Coleção Entomológica UFFS-RE, of the Universidade Federal da Fronteira Sul, Campus Realeza, Paraná, Brazil.

TAXONOMY

Omophoita personata (Illiger, 1807)

Figs 1-11

Egg (measurements in mm). Length 2.55 (2.26-2.66; N = 18), width 0.75 (0.64-0.87; N = 18). Color light brown, darker at the apex, elongated, fusiform, wider medially, narrower at the apex. Presence of micropyle at the apex. Chorion with grainy texture.

Third instar larva (measurements in mm). Body length: 12.93 (11.56-13.82; N = 4). Body width: 3.36 (3.22-3.52; N = 4).

Coloration. Head and legs black. All thoracic and abdominal segments orange in vivo, cream in AGA.

Head (Fig. 1). Hypognathous, rounded in frontal view, well sclerotized; slightly inserted into prothorax; epicranial suture Y-shaped; coronal suture short; frontal sutures pale, reaching antennal sockets. Endocarina long, robust, extending from base of frontal sutures to clypeus, externally represented by a slight depression, internally ridge-shaped. Each epicranial plate with one pair of filiform setae medially (eps1-2) and two pairs of filiform setae near antenna (eps3-6). Frons with three pairs of filiform setae (frs1-3), one pair near antenna (frs1), one pair medially (frs2), and one medially, near the frontoclypeal suture (frs3). Antennae small, glabrous, two-segmented; antennal sockets membranous, well-developed; base of first antennomere well sclerotized, wider than long; second antennomere cylindrical with conical apex. Stemmata absent. Clypeus transverse, sclerotized basally, membranous apically mainly in the median region. Labrum sclerotized, transverse, slightly wider than clypeus, with four pairs of filiform setae: two pairs of long setae, one laterally (lbs1) and one medially (lbs2), and two pairs of short setae (lbs3-4) near the anterior margin; anterior angles rounded, anterior margin notched medially. Mandibles (Fig. 2) symmetrical, palmate, strongly sclerotized, with five teeth: teeth 1 and 2 the longest, similar in length, teeth 3 and 4 slightly serrated, teeth 4 and 5 short; external face with two prominent filiform setae, one near tooth 1 (mds1) and other dorsomedially (mds2); condyle present, well developed; penicillus with a row of 14 short and thick setae. Maxilla (Fig. 3): cardo narrow, transverse, with one filiform setae (cas1) near outer margin; stipes elongated, wider basally, sclerotized, apically membranous, with two filiform setae (sts1-2); palpiger with two setae ventrally (pas1); mala pronounced, basally sclerotized, with a projection palpiform containing two segments, surrounded by six thick setae, followed by a line of flattened setae on the internal region; maxillary palp three-segmented: first palpomere glabrous, second palpomere with one ventral seta (mps1) and one ventrolateral seta (mps2); third palpomere with one ventral filiform microseta (mps3). Labium: postmentum greater than the prementum, weakly sclerotized, with two pairs of long filiform setae, one in the anterior region (mes1) and one in the posterolateral region (mes2), and one pair of short filiform setae anterolaterally (mms1); prementum basally sclerotized, with one pair of filiform setae near the base (prs1), three pairs of short setae (prs2-4) between the labial palps and two pairs of sensilla between the labial palps; labial palpi two-segmented. Hypopharynx densely covered with short and wide setae. Epipharynx (Fig. 4) with four pairs of thick and short setae (epis1-4) in the anterior angles and a dense set of thin setae in anterior margin, medially.

Figures 1-4
Head and mouthparts of third instar larva of Omophoita personata: (1) head, frontal view; (2) mandible, ventral view; (3) maxillae and labium, ventral view; (4) epipharynx, ventral view. Scale bars: 1 = 0.3 mm; 2-4 = 0.25 mm.

Prothorax (Fig. 5). Slightly smaller than following thoracic segments; pronotum with 20 short tubercles, each bearing one filiform setae, distributed as follows: two rows in the pronotum, one row on anterior margin with 12 tubercles (apt1-6) followed by one row with four tubercles (ppt1-2) (Fig. 5), one lateral pair (lpt) and one anteroventrolateral pair (alt); prosternum with median Y-inverted-shaped sclerotization, containing two pairs of short setae (vpt1-2) medially (Fig. 7).

Figures 5-6
Larvae of Omophoita personata: (5) third instar larva, dorsal view; (6) first instar larva, dorsal view. Scale bars: 5 = 2.25 mm; 6 = 0.75 mm.

Figures 7-9
Third instar larva of Omophoita personata: (7) head, thorax and abdomen, ventral view; (8) leg, ventral view; (9) abdominal segments VIII-X, ventral view. Scale bars: 7 = 1.2 mm; 8 = 0.45 mm; 9 = 0.5 mm.

Mesothorax and metathorax (Figs 5, 7). With one pair of dorsolateral digitiform tubercles (mdt), bearing two filiform setae (from here called bisetose tubercles) and 10 short tubercles, bearing one filiform setae (from here called unisetose tubercles), distributed as follows: two transverse dorsal rows, the anterior row with one pair of tubercles (amt1) and the posterior row with two pairs of tubercles (pmt1-2) (Fig. 5), one anterolateral pair (lmt) and one ventral pair (vmt1) (Fig. 7); presence of one ventral bisetose tubercle (vmt2) (Fig. 7), anteriorlly to the pair of unisetose tubercles. Mesothorax with one pair of annuliform spiracles above the anterolateral tubercle.

Legs (Fig. 8). Partially sclerotized; coxa partially sclerotized, with trapezoidal shape, with ten filiform setae, distributed as follows: three long setae (cs1-3) in the posterior margin of the sclerotized region, one long setae (cs4) and one short seta (cms1) on the ventral inner margin, three short setae (cms2-4) in the anterior margin of the sclerotized region and two short setae (cms5-6) on the dorsal outer margin; trochanter with four filiform setae (ts1-4) ventrally; apex of femur with six filiform setae (fs1-6), the ventral seta (fs1) is the longest; tibia with seven filiform setae (tis1-7): five medially (tis1-5) and two short setae (tis6-7) at the apex. Tarsungulus elongated, almost straight, with one filiform seta (tas1) basally.

Abdomen (Figs 5, 6, 9). Segments I-VII with one pair of dorsolateral digitiform tubercles (adt1), unisetose, with filiform setae (Figs 5, 7) and 12 short tubercles, with filiform setae, distributed as follows: dorsally one anteromedially bisetose tubercle (aat1), followed by a row of two pairs of unisetose tubercles (pat1-2) (Fig. 5); laterally one bisetose pair of tubercles (lat1); ventrally five bisetose tubercles, one of them medially (vat1) and the others in a posterior transverse row (vat2-3) (Fig. 7). Segment VIII quadrangular, with one pair of dorsolateral digitiform tubercles (adt2), unisetose, directed posteriorly (Figs 5, 9); three bisetose dorsal tubercles with one short anterodorsal tubercle (aat2) and one pair of more developed tubercles (pat3) near the posterior margin (in these tubercles the basal seta is shorter) (Fig. 5); one pair of short lateral bisetose tubercles (lat2) (Fig. 9); ventral tubercles as in the previous segments (Fig. 9). Segment IX (Fig. 9) with 14 filiform setae, four dorsal pairs (abs1-4), near the posterior margin and three ventral pairs (vas1-3) arranged diagonally (Fig. 9). Segment X not visible in dorsal view; it emerges from segment IX, forming the anal pseudopod, fusiform (Fig. 9). Annuliform spiracles present in segments I-VIII, above the lateral tubercles.

Pupa (measurements in mm). Body length: 7.33 (6.17-8.27; N = 26). Body width: 3.73 (3.13-4.38; N = 26).

Coloration. Yellow in vivo, pale yellow in AGA.

Head (Fig. 10). Rounded, not visible dorsally, with three pairs of filiform setae (hs1-3) as follows: one interantennal pair on frons, near the median line (hs1) and two pairs (hs2-3) at the vertex; mouthparts developed and visible; labrum with a pair of microsetae (lms1) in the anterior angles; antennae developed, with 11 antennomeres.

Prothorax (Fig. 10,11). With eight pairs of setae, four pairs anteriorly (aps1-4), two pairs medially (mps1-2) and two pairs (pps1-2) near the posterior margin.

Figures 10-11
Pupa of Omophoita personata: (10) ventral view; (11) dorsal view. Scale bars: 1.5 mm.

Mesothorax and metathorax. Mesonotum and metanotum (Fig. 11) with two pairs of setae (ms1-2) medially. Mesothorax with one pair of annuliform spiracles. Fore and mid-legs (Figs 10, 11) with three filiform setae at the apex of femur (pfs1-3; msfs1-3); hind legs with two setae at apex of femur (mtfs1-2).

Abdomen (Figs 10, 11). Segments I-VII with three pairs of setae inserted in small tubercles (abs1-3), evenly spaced, with two dorsal pairs and one lateral pair. Segment VIII with two pairs of short setae (abms1-2), thinner than those of the previous segments. Segment IX bearing one pair of urogomphi with apex directed posteriorly; base of each urogomphus with one microseta (abms3). Segments I-V with one pair of annulliform spiracles.

Remarks. The first-instar larva (Fig. 6) of O. personata has differences from advanced larval instars. Beyond the size (body length: 3.62 mm, 3.24-3.99 mm, N = 7; body width: 0.76 mm, 0.72-0.89 mm, N = 7), the following differences were observed, with characteristics of second and third instars shown between parenthesis: head visible in dorsal view (not visible), pronotum with darker coloration (lighter); prosternum with one pair of median setae (two pairs); mesonotum and metanotum with one pair of egg bursters (egg burster absent), one short bisetose tubercle (met1) in the anterior dorsal row (one pair of short unisetose tubercles in the anterior dorsal row); mesosternum and metasternum with one pair of setae posterior to the bisetose tubercle (one pair of unisetose tubercles posterior to the bisetose tubercle); abdominal segments I-VII with one dorsomedially bisetose tubercle (aat1), followed by one pair of unisetose tubercles (pat1) (one dorsal anteromedially bisetose tubercle, followed by a row of two pairs of unisetose tubercles); segment VIII with one bisetose anteromedially tubercle (aat2) and two posterior unisetose tubercles (pat2) (posterior tubercles bisetose). The second larval instar of O. personata was almost identical to the third instar, with differences in the body size (length: 6.91 mm, 6.00-7.82 mm, width: 1.70 mm, 1.60-1.80 mm; N = 2).

In general, the main differences from the first to second instar are the loss of egg bursters, replaced by two pairs of unisetose tubercles and the addition of tubercles and setae in the dorsal region of abdominal segments I-VIII.

Analyzing the immatures of O. octoguttata no differences were found in external morphology and chaetotaxy in comparison to O. personata. Only differences in size were recognized as follows (measurements in mm): egg length: 2.40 (N = 1); width: 0.72 (N = 1). First instar body length: 3.75 (3.10-5.12; N = 4); body width: 0.82 (0.69-0.95; N = 4). Second instar body length: 7.62 (N = 1); body width: 1.54 (N = 1). Third instar body length: 12.73 (12.23-13.97; N = 4); body width: 4.26 (3.85-4.51; N = 4). Pupae body length: 7.15 (6.08-8.21; N = 5), body width: 3.90 (3.54-4.31; N = 5).

Material examined. Adults (10 males, 16 females): BRAZIL, Paraná state: 3 males, 3 females, Realeza municipality -25.761349, -3.553202, 27.I.2020, Rech, T., col./Omophoita personata, Linzmeier det. 2020; 2 males, 3 females, same data but 10.III.2020; 3 males, 2 females, same data but 13.III.2020; 1 male, 1 female, same data but 10.I.2020/Omophoita octoguttata; 1 male, 7 females, same data but 15.V.2019, Copini, C., col./Omophoita octoguttata.

Immatures (17 eggs, 10 first instar larvae, 2 second instar larvae, 6 third instar larvae, 30 pupae): All immatures were obtained from adults from same locality already mentioned and reared in laboratory. Omophoita personata: 16 eggs; 6 first instar larvae; 2 second instar larvae; 3 third instar larvae; 25 pupae. Omophoita octoguttata: 1 egg; 4 first instar larvae; 3 third instar larvae; 5 pupae. All immatures were preserved inside the same container, identified as “Material voucher 001”.

DISCUSSION

The immatures of Omophoita personata here described and the immatures of Omophoita octoguttata also analyzed are very similar and the larvae cannot be differentiated based on their external morphology and chaetotaxy. Larvae of these species have thoracic and abdominal segments with prominent tubercles and no stemmata. According to Duckett and Swigoňová (2002Duckett CN, Swigoňová Z (2002) Description of immatures stages of Alagosa januaria Bechyné (Coleoptera: Chrysomelidae). Journal New York Entomological Society 110(1): 115-126.), such characteristics can be used to distinguish larvae of the Oedionychina subtribe from the other Alticini, since they are found in different species as Walterianella bucki Bechyné, 1956, Alagoasa parana Samuelson, 1985, Alagoasa januaria Bechyné, 1955 and Kuschelina gibbitarsa (Say, 1824). The absence of stemmata is pointed out as a characteristic of miner or subterranean species and would also be a characteristic of individuals in the process of recolonizing arboreal environments (Reid 1995Reid CAM (1995) A cladistic analysis of subfamilial relationships in the Chrysomelidae sensu lato (Chrysomeloidea). In: Pakaluk J, Ślipinski SA, Crowson RA (Eds) Biology, phylogeny, and classification of Coleoptera: papers celebrating the 80th birthday of Roy A. Crowson. Muzeum I Instytut Zoologii PAN, Warzawa, 559-631., Duckett and Swigoňová 2002Duckett CN, Swigoňová Z (2002) Description of immatures stages of Alagosa januaria Bechyné (Coleoptera: Chrysomelidae). Journal New York Entomological Society 110(1): 115-126.). Although O. personata and O. octoguttata feed externally, larvae were found spending much of their time under the soil or protected under leaves, which could, somehow, be related to the absence of stemmata.

The eggs of O. personata are also similar to those of other Oedionychina species, both in their shape and by the fact that they are laid in clusters, with variations only in size.

The first instar larvae of O. personata and O. octoguttata are also similar to those of Alagoasa bicolor (Linnaeus, 1767), A. januaria, W. bucki and Kuschelina bergi (Harold, 1881) (Virkki and Zambrana 1983Virkki N, Zambrana I (1983) Life history of Alagoasa bicolor (L.) in indoor rearing conditions (Coleoptera, Chrysomelidae, Alticinae). Entomologische Arbeiten aus dem Museum Georg Frey 31(32): 131-155., Duckett and Swigoňová 2002Duckett CN, Swigoňová Z (2002) Description of immatures stages of Alagosa januaria Bechyné (Coleoptera: Chrysomelidae). Journal New York Entomological Society 110(1): 115-126., Duckett and Casari 2002Duckett CN, Casari SA (2002) First descriptions of larval stages of Walterianella bucki Bechyné (Coleoptera: Chrysomelidae: Alticini) and notes on life history. The Coleopterists Bulletin 56(2): 170-181., Cabrera et al. 2016Cabrera N, Sosa A, Telesnicki M, Julien M (2016) Morphology of juvenile stages of Kuschelina bergi (Harold) with biological information (Coleoptera, Chrysomelidae, Alticini). ZooKeys 561: 51-61.). Omophoita personata, O. octoguttata, A. januaria and K. bergi share egg bursters on meso and metanotum. This feature is absent in W. bucki, and for A. bicolor there is no information about this structure (Virkki and Zambrana 1983Virkki N, Zambrana I (1983) Life history of Alagoasa bicolor (L.) in indoor rearing conditions (Coleoptera, Chrysomelidae, Alticinae). Entomologische Arbeiten aus dem Museum Georg Frey 31(32): 131-155.). The first instar of the two Omophoita’ species can be differed from A. bicolor by the filiform setae in all tubercles (in A. bicolor the setae of the main tubercles are club-shaped) and absence of prothoracic and anal shields (prothoracic and anal shields present in all A. bicolor larval instars); from A. januaria by the head setae filiform (in A. januaria setae of frons and vertex are spatulate); from W. bucki by the digitiform dorsolateral tubercles (in W. bucki the tubercles are large, lobe-like) and presence of filiform setae in all tubercles (in W. bucki the setae are club-shaped).

Regarding the third instar larvae, the two Omophoita’ species are very similar to A. januaria in morphology and chaetotaxy. However, in the abdominal segment VIII, that have the same morphology, a difference was found in terminology: the unisetose dorsolateral digitiform tubercles of O. personata were considered as two dorsal unisetose tubercles by Duckett and Swigoňová (2002Duckett CN, Swigoňová Z (2002) Description of immatures stages of Alagosa januaria Bechyné (Coleoptera: Chrysomelidae). Journal New York Entomological Society 110(1): 115-126.). Furthermore, in the description of segments IX and X of O. personata, the segment X was considered as being formed by the anal pseudopod. In A. januaria, Duckett and Swigoňová (2002Duckett CN, Swigoňová Z (2002) Description of immatures stages of Alagosa januaria Bechyné (Coleoptera: Chrysomelidae). Journal New York Entomological Society 110(1): 115-126.) describe the segment X as being formed by the anal pseudopod and another part with four pairs of setae arranged in a longitudinal row. This row was considered part of segment IX in O. personata as observed in the description of W. bucki (Duckett and Casari 2002Duckett CN, Casari SA (2002) First descriptions of larval stages of Walterianella bucki Bechyné (Coleoptera: Chrysomelidae: Alticini) and notes on life history. The Coleopterists Bulletin 56(2): 170-181.) and has three pair of setae.

Comparing O. personata and K. bergi some differences were found such as: 16 unisetose tubercles on pronotum arranged in two rows, the anterior with 12 unisetose tubercles and the posterior with four unisetose tubercles (15 unisetose tubercles on pronotum, nine anterior and six posterior unisetose tubercles in K. bergi), meso and metasternum with a bisetose median anterior tubercle and two unisetose posterior tubercles (two bisetose median tubercles in meso and metasternum in K. bergi), and mandibles with five teeth, one pair of filiform setae and penicillus formed by a row of 14 short and thick setae (mandibles with four teeth, one seta and penicillus formed by two thick setae in K. bergi). More conspicuous differences were observed in relation to Walterianella bucki. Omophoita personata has frons with filiform setae (in W. bucki the setae on frons are stout), filiform setae on pronotum (in W. bucki the pronotal setae are club-like), segment IX with seven pairs of setae arranged closer to each (in W. bucki the pairs of setae are arranged more sparsely), dorsolateral tubercles digitiform (in W. bucki tubercles are much more developed.

The pupae of O. personata and A. januaria have similar chaetotaxy of the head and pronotum (mesonotum, metanotum and legs not described by Duckett and Swigoňová (2002Duckett CN, Swigoňová Z (2002) Description of immatures stages of Alagosa januaria Bechyné (Coleoptera: Chrysomelidae). Journal New York Entomological Society 110(1): 115-126.), but were illustrated), and those of O. personata can be differentiate by one pair of setae on anterior angles of labrum (absent in A. januaria), segment VIII with two pairs of setae (glabrous in A. januaria), apex of urogomphi directed posteriorly with one pair of microsetae (apex directed inward to the center, glabrous in A. januaria). To K. bergi and W. bucki the pupa’s description was not included in the studies (Duckett and Casari 2002Duckett CN, Casari SA (2002) First descriptions of larval stages of Walterianella bucki Bechyné (Coleoptera: Chrysomelidae: Alticini) and notes on life history. The Coleopterists Bulletin 56(2): 170-181., Cabrera et al. 2016Cabrera N, Sosa A, Telesnicki M, Julien M (2016) Morphology of juvenile stages of Kuschelina bergi (Harold) with biological information (Coleoptera, Chrysomelidae, Alticini). ZooKeys 561: 51-61.), and to Alagoasa bicolorVirkki and Zambrana (1983Virkki N, Zambrana I (1983) Life history of Alagoasa bicolor (L.) in indoor rearing conditions (Coleoptera, Chrysomelidae, Alticinae). Entomologische Arbeiten aus dem Museum Georg Frey 31(32): 131-155.) shows the development, but also does not describe the morphology.

It is important to mention that comparing larvae and pupa of O. octoguttata with the description presented by Begha et al. (2021Begha BP, Santos MH, Prado LR (2021) Redescription of Omophoita octoguttata (Coleoptera: Chrysomelidae) and its immature stages, with notes on life history. Iheringia, Série Zoologia 111: 1-9.), we were able to give complementary information since more detailed data about first and second instar larvae, mouthparts morphology, and pupa chaetotaxy were added. Moreover, some inconsistences related with morphological misinterpretations were also observed. Among them are the position of thoracic spiracles in mesothorax (not in prothorax as reported by the authors) as found in the vast majority of the Coleoptera (Costa et al. 2006Costa C, Ide S, Simonka CE (2006) Insetos imaturos - metamorfose e identificação. Editora Holos, Ribeirão Preto, 249 pp.), number of antennomeres and palpomeres, two and three respectively (not three and four, respectively), abdominal segments I-VII with two dorsolateral digitiform tubercles and 12 short tubercles (not eight tubercles), 16 filiform setae on pronotum (not ten setae), number of seta on each tubercle, since some are bisetose and other are unisetose (not all tubercles bisetose), and legs chaetotaxy.

Descriptions of immatures of Alticini species (Table 1) that do not belong to the Oedionychina, such as Pseudolampsis darwini (Scherer, 1964), Pseudolampsis guttata (LeConte, 1884), Megistops vandepolli Duviver, 1889 and Distigmoptera borealis Blake, 1943, have more conspicuous morphological differences compared to O. personata. Among them are the absence of prominent tubercles, the presence of sclerites throughout the body, presence of stemmata and distinct chaetotaxy pattern. These characteristics may be related to the environmental niches they occupy and their eating habits (external feeders of Angiosperms, leaf miners, moss inhabiting, etc) reflecting the evolutionary history of these groups, since such species are positioned in different clades within Alticini (Duckett and Kjer 2003Duckett CN, Kjer KM (2003) Cladistic analysis of the Oedionychines of Southern Brazil (Galerucinae: Alticini) based on two molecular markers. In: Furth DG (Ed.) Special topics in leaf beetle biology. Proceedings of the Fifth International Symposium on the Chrysomelidae. Pensoft Publishers, Sofia, 117-132.). As for larvae, more conspicuous differences are observed between the pupae of O. personata, Pseudolampsis darwini and P. gutatta, which differ in chaetotaxy as well as in the presence of urogomphi (absent is Pseudolampsis’ species). The urogomphi are found in O. personata, O. octoguttata and Megistops vandepolli which differs from the Omophoita’ species by the urogomphi facing inward, legs glabrous and body covered with more setae.

Table 1
Morphological characteristics of larval first and third instar, and pupa of Alticini species.

Not all species addressed in this study have the pupa described. The lack of information and incomplete or superficial descriptions may be due to the lack of specimens in pupal stage, the difficulty of finding them in nature, conservation of the material and the disregard of pupae’ characteristics for species characterizations.

Overall, the immatures of O. personata and O. octoguttata, as well as the other immatures of Oedionychina, have very similar morphological characteristics, reinforcing the morphological stability of immatures of this group. These immatures characteristics along with those of adults, help to better characterize Oedionychina and could help clarify relationships between the groups.

ACKNOWLEDGMENTS

The authors wish to thank the reviewers for their valuable comments and contributions. TR thanks to Coordenação de Aperfeiçoamento de Pessoal de Nível Superior for the doctoral scholarship (process 88882.457426/2019-01). We also thank to H. Doerrier from HD translations for the language translation. Any proficiency inadequacies in the final English version are responsibility of the authors.

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ADDITIONAL NOTES

Edited by

Editorial responsibility

Ângelo Parise Pinto

Data availability

Data citations

Linzmeier AM (2022) Alticini in Catálogo Taxonômico da Fauna do Brasil. PNUD PNUD http://fauna.jbrj.gov.br/fauna/faunadobrasil/119960 [Accessed: 21/02/2022]

Publication Dates

  • Publication in this collection
    02 Sept 2022
  • Date of issue
    2022

History

  • Received
    13 Sept 2021
  • Accepted
    03 June 2022
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