Five new species of Asphondylia (Diptera, Cecidomyiidae, Asphondyliini) from Brazilian restinga (Atlantic Forest)

Abstract

Asphondylia braziliensissp. nov.,Asphondylia fluminensissp. nov.,Asphondylia marambaiensissp. nov.,Asphondylia varroniaesp. nov., and Asphondylia xerezi sp. nov., are described based on material from Brazilian restingas. The first species induces galls on Struthanthus acuminatus (Ruiz. & Pav.) Kuijt (Loranthaceae), the second on Erythroxylum ovalifolium Peyr (Erythroxylaceae), the third on Lantana fucata Lindl. (Verbenaceae), the fourth on Varronia curassavica Jacq. (Cordiaceae), and the fifth on Heliotropium sp. (Heliotropiaceae). One host plant, Erythroxylum ovalifolium, is endemic to the Atlantic forest. The other plant species are native to Brazil. Illustrations of relevant morphological characters are provided. The new species are compared with congeneric Neotropical species. Types are deposited in the Entomological Collection of the Museu Nacional, Rio de Janeiro.

Keywords
Gall midges; Host plant; Morphology; Taxonomy.

INTRODUCTION

Asphondylia Loew, 1850 (Asphondyliina, Asphondyliini, Cecidomyiidi) is a diverse and widespread genus of Cecidomyiidae, with more than 300 species, all gall-inducers, 98 of them occur in the Neotropical region and 20 in Brazil (Gagné & Jaschhof, 2021). The genus is easily recognizable mainly by the male and pupal morphology. Males have two-toothed gonostyli and pupae exhibit upper and lower frontal horns (Gagné, 1994).

Five new species of Asphondylia are described here based on material from Brazilian restingas, a phytophysiognomy of the Atlantic forest. They induce galls on Erythroxylum ovalifolium Peyr (Erythroxylaceae), Lantana fucata Lindl. (Verbenaceae), Struthanthus acuminatus (Ruiz. & Pav.) Kuijt (Loranthaceae), Varronia curassavica Jacq. (Cordiaceae), and Heliotropium sp. (Heliotropiaceae), all plants native to Brazil. Their galls were previously reported and illustrated in the literature by Maia, 2001 and Rodrigues et al., 2014. Due to nomenclatural changes, four botanical names were updated: Lantana fucata, Struthanthus acuminatus and Heliotropium sp. - cited as Lantana lilacina Desf., Struthanthus concinnus (Mart.) Mart. and Tournefortia sp. in Rodrigues et al., 2014, respectively, and Varronia curassavica - cited as Cordia verbenacea DC. in Maia, 2001 (Miller & Gottschling, 2007). In addition, the genera Varronia and Heliotropium previously positioned in Boraginaceae are currently positioned in the families Cordiaceae and Heliotropiaceae, respectively (Miller, 2013).

According to Gagné & Jaschhof, 2021, five gall midge species have been reported on Cordiaceae, six on Erythroxylaceae, three on Heliotropiaceae, four on Loranthaceae, and 18 on Verbenaceae in the Neotropical Region.

Varronia curassavica is native to Brazil, where it occurs in five phytogeographical areas: Amazon, Atlantic Forest, Caatinga, Cerrado, and Pampa (Stapf & Silva, 2023). This plant is used in folk medicine due to its anti-inflammatory, analgesic and cicatrizing properties (Lorenzi & Matos, 2008). Heliotropium L. is a genus native to Brazil, where it is widespread, with records in all phytogeographical domains and states, being represented by 14 species (Melo, 2023). Erythroxylum ovalifolium is endemic to the Atlantic Forest, restricted to the restinga physiognomy and known only from the state of Rio de Janeiro. Struthanthus acuminatus has only been recorded in areas of the Atlantic Forest in Brazil (Caires & Dettke, 2023). Lantana fucata L., recognized as a ruderal species, is native to Tropical and Subtropical America (Silva & Lima, 2012). In Brazil, this plant is widespread, occurring from north to south, in the Atlantic Forest, Caatinga, Cerrado, and Pampas (Silva et al., 2022).

MATERIAL AND METHODS

Galls on Erythroxylum ovalifolium, Lantana fucata, Struthanthus acuminatus, Varronia curassavica and Heliotropium sp. were collected at restinga areas in the southeastern Brazil, the first in the State of Rio de Janeiro (RJ), in the following municipalities - Maricá (22°52′S, 42°54′W), Arraial do Cabo (22°55′S, 42°21′W) and Carapebus (22°23′S, 41°45′W); the second and fifth in RJ, both in the municipality of Mangaratiba (Ilha da Marambaia, Kutuca beach: (23°04′04.85″, 23°03′59.17″S, 43°59′39.41″, 43°59′33.59″W); and the fourth in the State of São Paulo, in Bertioga (23°51′S, 46°08′W) as well as in RJ, in Maricá and Carapebus. Field work was performed in different dates (see “Material Examined”).

Branches with galls were removed from the host plants and transported in plastic bags to the laboratory. Some gall samples were dissected to obtain the larvae and the remainder was kept in plastic pots covered by a fine screen to obtain adults and pupal exuviae. The specimens were first preserved in 70% ethanol and then mounted on microscope slides following the methods outlined in Gagné (1994). The genus was identified using the identification key of Gagné (1994), and the new species were proposed after comparison with literature data (host plants, gall morphology and cecidomyiid descriptions). All specimens were deposited in the Entomological Collection of the Museu Nacional/Universidade Federal do Rio de Janeiro (MNRJ).

Morphological studies and drawings were made with the aid of an optical microscope with coupled photographic camera and drawing tube. Measurements were made using a microscope slide with scale from 0.01 to 5.0 mm. Length of wings was measured from the arculus to the apex (Fig. 1A); total length of females from head vertex to posterior margin of the 8^th abdominal segment; pupal antennal horns were measured from base to apex, the length of the basal part was compared with that of the distal part (Fig. 1B). Lower facial horns are considered “aligned” when mesal and lateral horns are arranged in an almost horizontal line (Fig. 1B) and “non-aligned” when the mesal horn is clearly positioned below the lateral horns (Fig. 1C). All drawings were redrawn using Corel DRAW®. Adult morphological terminology follows Gagné (1994). The new species were compared to other Neotropical congeneric species.

Figure 1
Asphondylia spp.: (A) Asphondylia hancorniae Maia, 2023, wing, (B-C) Pupa head, frontal view: (B) Antennal horn: basal and distal part, lower facial horn, teeth aligned (seta), (C) Lower facial horn, teeth not aligned (seta).

Descriptions

Asphondylia braziliensis Maia, sp. nov. (Figs. 2A 4D)

Diagnosis: Male hypoproct rounded apically, deeply bilobed; ovipositor with needle part about 1.00-1.20 × length 7^th sternite; pupa: antennal horn 0.23-0.30 mm long, upper facial horn single and conical, three lower frontal horns not aligned, 8^th abdominal segment with 5-7 dorsal spines in the posterior row, larva: spatula with lateral teeth longer than mesal ones, mesal teeth rounded apically, three setose lateral papillae on each side of spatula.

Figure 2
Asphondylia braziliensis Maia, sp. nov., male: (A) Head, ventral view, (B) 5^th flagellomere, (C) 10^th 12^th flagellomeres, (D) Midleg, tarsal claw and empodium, lateral view, (E) Wing, (F) Terminalia, dorsolateral view.

Male: Body: 3.40-3.75 mm long (N = 3). Head (Fig. 2A): 0.40-0.50 mm long, 0.50 mm wide (N = 3), eye facets hexagonal, closely appressed; antennae: flagellomeres 1 and 2 not fused, scape obovate, setose, 0.11-0.12 mm long, 0.0 mm wide (N = 3), pedicel globose, setose, 0.05 mm long, 0.05 mm wide (N = 3), 1^st 12^th flagellomeres cylindrical, all 0.04 mm wide, circumfila longitudinally wavy, dense, anastomosing, equally spread along segments (Fig. 2B), 1^st 7^th flagellomeres 0.1-0.18 mm long (N = 3), 8^th 10^th flagellomeres 0.16-0.17 mm long (N = 3), 11^th 0.15-0.17 mm long (N = 3), 12^th flagellomere 0.12-0.13 mm long (N = 2) (Fig. 2C), proportion flagellomere neck-node 1:10; frons with 30 setae (N = 2); mouthparts: labrum long-attenuated, 0.08 mm long, 0.03 mm wide (N = 3); hypopharynx of the same shape as labrum, with long, anteriorly-directed lateral setulae, 0.10 mm long, 0.03 mm wide (N = 3); labella elongate and convex, 0.16 mm long, 0.10 mm wide (N = 1), with lateral and mesal setae; palpus 0.14 mm long: 1^st segment globoid, 0.02 mm long, 0.02 mm wide (N = 2), 2^nd segment cylindrical, 0.04 mm long, 0.02 mm wide (N = 2), 3^rd segment fusiform, 0.08 mm long, 0.02 mm wide (N = 2), all segments with setae. Thorax: scutum with two dorsocentral rows of setae, setae more abundant anteriorly and posteriorly, two groups of lateral setae more abundant anteriorly, extending from base to distal margin, scales intermixed; scutellum with scattered setae; anepimeron setose; remaining pleural sclerites bare; legs: tarsal claws curved beyond midlength, isomorphic, empodium longer than claws (Fig. 2D); wing: length 2.50 mm (N = 3) (Fig. 2E). Abdomen: trichoid sensilla not visible; 1^st 7^th tergites sclerotized, rectangular with a posterior row of setae, few scattered lateral and mesal setae and mostly covered elsewhere with scales, tergite 1 shorter than other tergites, ½ length of 2^nd tergite, 8^th tergite bare, narrow, longer sublaterally than mesally; 2^nd 8^th sternites more sclerotized than tergites, rectangular, as long as tergites, with a posterior row of setae, several mesal and lateral setae, and mostly covered elsewhere with scales; 8^th sternite with scattered setae and mostly covered elsewhere with scales. Terminalia (Fig. 2F): gonocoxite short and stout, 0.16 mm long, 0.07 mm wide (N = 2); gonostylus ovoid, 0.05 mm long, 0.04 mm wide; hypoproct deeply bilobed, rounded apically.

Female: Body length: 3.50-4.70 mm (N = 4). Head: 0.40-0.60 mm long, 0.50 mm wide, antennae: scape 0.12-0.13 mm long, 0.05-0.07 mm wide (N = 4), pedicel 0.05-0.07 mm long, 0.05-0.07 mm wide (N = 4), 1^st 11^th flagellomeres cylindrical, all 0.04 mm wide, circumfila comprising two longitudinal bands connected subbasally and apically by two transverse bands (Fig. 3A), flagellomeres 1 and 2 not fused, 1^st flagellomere 0.25 mm long (N = 4), 2^nd flagellomeres 0.19-0.21 mm long (N = 4), 3^rd flagellomere 0.18-0.20 mm long (N = 4), 4^th flagellomere 0.18-0.19 mm long (N = 5), 5^th and 6^th flagellomeres 0.17-0.20 mm long (N = 4), 7^th flagellomere 0.17-0.19 mm long (N = 4), 8^th flagellomere 0.16-0.18 mm long (N = 4), 9^th flagellomere 0.14-0.15 mm long (N = 4), 10^th flagellomere 0.11-0.12 mm long (N = 4), 11 flagellomere 0.06-0.08 mm long, 12^th flagellomere 0.04-0.045 mm long (Fig. 3B); mouthparts: labrum 0.11 mm long, 0.035 mm wide (N = 2), hypopharynx 0.16 mm long, 0.035 mm wide (N = 2), labellum 0.05-0.07 mm long, 0.02-0.04 mm wide at midlength (N = 4), palpus 0.19-0.23 mm long (N = 4): 1^st segment globose 0.02-0.03 mm long, 0.02-0.03 mm wide (N = 4), 2^nd segment cylindrical 0.06-0.065 mm long, 0.025-0.03 mm wide at midlength (N = 4), 3^rd segment conical 0.11-0.14 mm long and 0.02 mm wide at midlength (N = 4). Thorax: wing length: 2.90-3.00 mm (N = 3); tarsal claw more robust than in male (Fig. 3C). Abdomen (Fig. 3D): trichoid sensillae not visible, 1^st 7^th tergites as in male, 8^th tergite with posterior margin with lobes 0.15-17 mm long (N = 2), 2^nd 6^th sternites as in male, 6^th sternite 0.34 mm long (N = 1), 7^th sternite rectangular 0.54-0.63 mm long, 1.84-1.85 × length sternite 6 (N = 3), setose, mostly covered elsewhere with scales; sternite 8 not sclerotized; ovipositor: needle part 0.54-0.65 mm long (N = 3), 1.00-1.20 × length sternite 7 (N = 2). Other characters as in male.

Figure 3
Asphondylia braziliensis Maia, sp. nov. (A-C) female: (A) 3^rd flagellomere, (B) 10^th 12^th flagellomeres, (C) Hindleg, tarsal claw and empodium, lateral view, (D-F) pupa: (D) General aspect, ventral view, (E) Head, frontal view, (F) Prothoracic spiracle.

Pupa (Fig. 4A): Color: brownish. Body length: 2.80-4.20 mm (N = 8). Head (Fig. 4B): face with lateral projection; antennal horn 0.23-0.30 mm long (N = 5), conical, pointed, inner margin serrated, distal part longer than basal part; dorsal plate 0.12-0.21 mm long.0.40-0.53 mm wide (N = 6), apical seta 0.03-0.035 mm long (N = 5); upper facial horn bifid, 0.06 mm long (N = 6), conical; three lower facial horns not aligned, 0.03 mm long (N = 3); two pairs of lower facial papillae: one pair setose, the other bare; three pairs of lateral facial papillae: one pair setose, two bare; upper cephalic margin thickened laterally. Thorax: integument wrinkled (Fig. 4C); prothoracic spiracle, 0.11-0.12 mm long, longer than antennal basal width, setiform, conspicuously curved (N = 5) (Fig. 4D). Abdomen: segments 2-8 with transverse rows of crescent dorsal spines at basal half (Fig. 5A); posterior row with 12-17 spines in the 2^nd segment (N = 6), 14-22 in the 3^rd (N = 6), 14-18 in the 4^th (N = 6), 14-17 in the 5^th (N = 6), 13-16 in the 6^th (N = 6), 10-12 in the 7^th (N = 6), 5-7 in the 8^th (N = 6).

Figure 4
Asphondylia braziliensis Maia, sp. nov., pupa: (A) Thoracic integument, dorsolateral view, (B) 5^th 8^th abdominal segments, dorsal view, (C) Larva, spatula and associated papillae, ventral view, (D) Stem galls on Struthanthus acuminatus (Ruiz. & Pav.) Kuijt (Loranthaceae).

Figure 5
Asphondylia spp.: (A-B) Asphondylia parasiticolaMöhn, 1960, ventral view, redrawn from Möhn, 1960 (original drawings without scales): (A) Larva, spatula and associated papillae, (B) Pupa, head, (C) Asphondylia struthanthiRübsaamen, 1915, redrawn from Rübsaamen, 1915: Larva, spatula and associated papillae, ventral view (original drawings without scales).

Larva: Body: 2.30-2.60 mm long (N = 5); head 0.04 mm long, 0.081 mm wide (N = 2). Spatula (Fig. 5B) quadridentate, 0.24-0.25 mm long (N = 4), lateral teeth longer than mesal, 0.03 mm long, mesal teeth 0.02 mm long (N = 3); three setose lateral papillae on each side of spatula. Terminal segment 0.07-0.08 mm long (N = 3). Terminal segment with no visible papillae in the slides (terminal segment smashed).

Gall: On leaf and stem, conical, green, glabrous, complex, one-chambered on Struthanthus acuminatus (Ruiz. & Pav.) Kuijt) (Loranthaceae) (Fig. 5C).

Material examined: Holotype male, BRAZIL: Rio de Janeiro State, Mangaratiba, Ilha da Marambaia, Praia do Kutuca, 21.VI.2020, Rodrigues, A. col. (MNRJ-ENT1-69762). Paratypes: MALES - same data as holotype: 2 ♂♂ (MNRJ-ENT1-69763, MNRJ-ENT1-69764); FEMALES - same data as holotype: 2 ♀♀ (MNRJ-ENT1-69768, MNRJ-ENT1-69769); 28.VIII.2010: 2 ♀♀ (MNRJ-ENT1-69766, MNRJ-ENT1-69767); Praia Suja, 18.IX.2020: 1 ♀ (MNRJ-ENT1-69765); PUPAL EXUVIAE - same data as holotype: 2 pupal exuviae (MNRJ-ENT1-69771, MNRJ-ENT1-69778); 24.VII.2010: 8 pupal exuviae (3: MNRJ-ENT1-69770, 2: MNRJ-ENT1-69773, 3: MNRJ-ENT1-69774); 28.VIII.2010: 2 pupal exuviae (MNRJ-ENT1-69775); 18.IX.2020: 1 pupal exuviae (MNRJ-ENT1-69776); 25.II.2011: 2 pupal exuviae (MNRJ-ENT1-69772, MNRJ-ENT1-69777); PUPA - 25.II.2011: 1 pupa (MNRJ-ENT1-69772); THIRD-INSTAR LARVAE - 26.V.2010: 4 larvae (MNRJ-ENT1-69781); 24.VII.2010: 8 larvae (4: MNRJ-ENT1-69779, 4: MNRJ-ENT1-69780).

Etymology: The name “braziliensis” refers to the country where the type-material was collected.

Geographic distribution (based on gall records on the host plant): Brazil, Rio de Janeiro State, Mangaratiba municipality (Rodrigues et al., 2014).

Remarks: Only two species of Asphondylia were previously known from Struthanthus: A. parasiticolaMöhn 1960, inducer of stem and fruit galls on Struthanthus marginatus, described from El Salvador, and A. struthanthiRübsaamen 1915, inducer of fruit galls on Struthanthus sp., described from Brazil (Gagné & Jaschhof, 2021).

Asphondylia braziliensis differs from A. parasiticola in the following morphological characters:

• (1) larva: spatula with stalk longer in A. parasiticola than in A. braziliensis, apical teeth weakly indented in A. parasiticola and deeply indented in A. braziliensis, mesal teeth somewhat longer in A. parasiticola than in A. braziliensis, mesal teeth pointed in A. parasiticola and rounded apically in A. braziliensis, 4 setose lateral papillae on each side of spatula in A. parasiticola, 3 setose lateral papillae on each side of spatula in A. braziliensis (Fig. 5A × Fig. 4C);

• (2) pupa: antennal horns truncate in ventral view in A. parasiticola and pointed in A. braziliensis, upper facial horn bifid in A. parasiticola and simple in A. braziliensis, one lower facial horn in A. parasiticola and three in A. braziliensis (Fig. 5B × Fig. 3E);

• (3) female: the new species has body longer, all segments of the palpi shorter, scape, pedicel, 1^st, 5^th and 9^th flagellomeres shorter, 10^th flagellomeres longer, 11^th shorter or equal and 12^th equal or longer in A. braziliensis than in A. parasiticola (Table 1).

  Table 1
  Comparative length of adults and larvae of Asphondylia braziliensis Maia, sp. nov., A. parasiticolaMöhn, 1960, and A. struthanthiRübsaamen, 1915 (Diptera, Cecidomyiidae). Data on the last two species were obtained from literature.
  

  Length:	Asphondylia braziliensis, sp.nov.	Asphondylia parasiticolaMöhn, 1960	Asphondylia struthanthiRübsaamen, 1915
  Male
  Body	3.40‑3.75 mm	-	2.90‑3.00 mm
  Scape	0.110‑120µ	-	118µ
  Pedicel	50µ	-	72µ
  1st flagellomere	170‑180µ	-	256µ
  5th flagellomere	170‑180µ	-	-
  9th flagellomere	160‑170µ	-	-
  10th flagellomere	160‑70µ	-	-
  11th flagellomere	150‑170µ	-	-
  12th flagellomere	12‑130µ	-	-
  Palpus
  1st segment	20µ	-	28‑30µ
  2nd segment	40µ	-	46‑48µ
  3rd segment	80µ	-	100µ
  Female
  Body	3.50‑4.70 mm	3.70‑3.80 mm	-
  Scape	120‑130µ	184µ	-
  Pedicel	50‑70µ	75‑76µ	-
  1st flagellomere	250µ	308µ	-
  5th flagellomere	170‑200µ	212µ	-
  9th flagellomere	140‑150µ	162µ	-
  10th flagellomere	110‑120µ	108µ	-
  11th flagellomere	60‑80µ	80µ	-
  12th flagellomere	40‑45µ	40µ	-
  Palpus
  1st segment	20‑30µ	39‑40µ	-
  2nd segment	60‑65µ	90‑92µ	-
  3rd segment	110‑140µ	162‑164µ	-
  Larva
  Body	2.30‑2.60 mm	3.50 mm	2.50‑2.70 mm
  Spatula	240‑250µ	285µ	212µ

The male of A. parasiticola is unknown. Asphondylia braziliensis differs from A. struthanthi in the following morphological characters:

• (1) larva: spatula: stalk longer in A. struthanthi than in A. braziliensis, base of spatula truncate in A. struthanthi and anchor-shaped in A. braziliensis, mesal teeth divergent in A. struthanthi and parallel in A. braziliensis (Fig. 5C × Fig. 4C);

• (2) pupa: the upper frontal horn is bifid in A. struthanthi and conical in A. braziliensis (Fig. 6A × Fig. 3E), the prothoracic spiracle is cylindrical in A. struthanthi and setiform in A. braziliensis (Fig. 6B × Fig. 3F);

  

  Figure 6
  Asphondylia struthanthiRübsaamen, 1915, redrawn from Rübsaamen, 1915 (original drawings without scales): (A-B) Pupa: (A) Head, frontal view, (B) Prothoracic spiracle, (C) Male terminalia, dorsal view.
  

• (3) male: the body is shorter in A. struthanthi than in A. braziliensis, the pedicel and 1^st flagellomere, and all segments of palpi are shorter in A. braziliensis than in A. struthanthi (Table 1). The female of A. struthanthi is unknown.

Asphondylia fluminensis Maia, sp. nov. (Figs. 7A 11B)

Diagnosis: Male hypoproct rounded apically, slightly bilobed, ovipositor with needle part about 2.02-2.14 × length 7^th sternite; pupa: antennal horn 0.20-0.26 mm long, upper facial horn single and conical, three lower frontal horns aligned, 8^th abdominal segment with 7-8 dorsal spines in the posterior row, larva: spatula with lateral teeth longer than mesal ones, mesal teeth pointed apically, three setose lateral papillae on each side of spatula.

Figure 7
Asphondylia fluminensis Maia, sp. nov., male: (A) Head, ventral view, (B) 1^st 2^nd flagellomeres, (C) 5^th flagellomere, (D) 7^th 12^th flagellomeres, (E) Foreleg, tarsal claw and empodium, lateral view.

Male: Body: 2.60-3.05 mm long (N = 3). Head (Fig. 7A): 0.55 mm long, 0.55 mm 0.60 mm wide (N = 3), eye facets circular, closely appressed; antennae: flagellomeres 1 and 2 not fused (Fig. 7B), scape obovate, setose, 0.10-0.11 mm long, 0.05-0.06 mm wide (N = 4), pedicel globose, setose, 0.04-0.06 mm long, 0.05-0.06 mm wide, 1^st 9^th flagellomeres cylindrical, all 0.04-0.05 mm wide, circumfila longitudinally wavy, dense, anastomosing, equally spread along segments (Fig. 7C), 1^st flagellomere 0.21-0.22 mm long (N = 4), 2^nd flagellomere 0.17-0.19 mm long (N = 4), 3^rd 7^th flagellomeres 0.17-0.18 mm long (N = 4), 8^th 10^th flagellomeres 0.16 mm long (N = 2), 11^th flagellomere 0.15-0.16 mm long (N = 2), 12^th flagellomeres 0.15 mm long (N = 2) (Fig. 7D), proportion flagellomere node/neck: 1:16; frons with 37-40 setae (male, N = 2); mouthparts: labrum long-attenuated, 0.07-0.08 mm long, 0.03 mm wide (N = 2); hypopharynx of the same shape as labrum, 0.09-0.13 mm long, 0.03 mm wide (N = 3) with long, anteriorly-directed lateral setulae; labella elongate and convex, 0.04-0.05 mm long, 0.02-0,03 mm wide (N 3), with lateral and mesal setae; palpus 0.11-0.14 mm long: 1^st segment globoid, 0.02 mm long, 0.02 mm wide (N = 3), 2^nd segment cylindrical, 0.03-0.04 mm long, 0.02 mm wide (N = 3), 3^rd segment fusiform, 0.06-0.08 mm long, 0.02 mm wide (N = 3), all segments with setae. Thorax: scutum with two dorsocentral rows of setae, setae more abundant anteriorly and posteriorly, two groups of lateral setae more abundant anteriorly, extending from base to distal margin, scales intermixed; scutellum with scattered setae; anepimeron and anepistenum setose; remaining pleural sclerites bare; legs: tarsal claws curved beyond midlength, isomorphic, empodium longer than claws (Fig. 7E); wing: length 2.20-2.30 mm (N = 3). Abdomen (Fig. 8A): trichoid sensilla not visible; 1^st 7^th tergites sclerotized, rectangular with a posterior row of setae, few scattered lateral setae and mostly covered elsewhere with scales, 1^st tergite shorter than other tergites, 1/35 × length of 2^nd tergite, 8^th tergite band-like, bare; 2^nd 8^th sternites more sclerotized than tergites, rectangular, as long as tergites, with a posterior row of setae, several setae and midlength, few lateral setae, and mostly covered elsewhere with scales; 8^th sternite with scattered setae and mostly covered elsewhere with scales. Terminalia (Fig. 8B): gonocoxite short and stout, 0.14 mm long, 0.07-0.09 mm wide (N = 2); gonostylus ovoid, 0.05 mm long, 0.04 mm wide (N = 2); hypoproct slightly bilobed, rounded apically.

Figure 8
Asphondylia fluminensis Maia, sp. nov., male: (A) 3^rd abdominal segment-terminalia, lateral view, (B) Terminalia, dorsal view.

Female: Body length: 2.80-2.90 mm (N = 2). Head: 0.05 mm long, 0.04 mm long (N = 2), antennae: scape 0.11 mm long, 0.05 mm wide (N = 1), pedicel 0.05-0.07 mm long, 0.06-0.07 mm wide (N = 2), 1^st 11^th flagellomeres cylindrical, circumfila comprising two longitudinal bands connected subbasally and apically by two transverse bands (Fig. 9A), 1^st flagellomere 0.23-0.28 mm long, 0.05 mm wide (N = 2), 2^nd flagellomere 0.28 mm long. 0.05 mm wide (N = 1), 3^rd flagellomere 0.16-0.18 mm long, 0.04-0.05 mm wide (N = 2), 4^th 5^th flagellomeres 0.15-0.18 mm long, 0.04-0.05 mm wide (N = 2), 6^th flagellomere 0.15-0.17 mm long, 0.04-0.05 mm wide (N = 2), 7^th flagellomere 0.17 mm long, 0.04-0.05 mm wide (N = 1), 8^th flagellomere 0.14 mm long, 0.04 mm wide (N = 1), 9^th flagellomere 0.10 mm long, 0.04 mm wide (N = 1), 10^th flagellomere 0.08 mm long, 0.04 mm wide (N = 1), 11^th flagellomere 0.05 mm long, 0.04 mm wide (N = 1), 12^th flagellomere globose, 0.04 mm long, 0.04 mm wide (N = 1); proportion flagellomere neck-node 1:15; frons with 26 setae; mouthparts: labrum 0.05-0.05 mm long, 0.04 mm wide (N = 2), hypopharynx 0.10 mm long, 0.04 mm wide (N = 2), labellum 0.04-0.045 mm long, 0.03 mm wide at midlength (N = 2), palpus 0.14 mm long (N = 1): 1^st segment globose 0.02 mm long, 0.02 mm wide (N = 1), 2^nd segment cylindrical 0.05 mm long, 0.02 mm wide at midlength (N = 1), 3^rd segment fusiform 0.07 mm long, 0.01 mm wide at midlength (N = 1). Thorax: wing length: 2.30-2.40 mm (N = 2) (Fig. 9B); tarsal claws curved beyond midlength, isomorphic, empodium longer than claws (Fig. 9C). Abdomen (Fig. 9D): trichoid sensillae not visible, 1^st 7^th tergites as in male, 8^th tergite with posterior margin with lobes 0.08-0.09 mm long (N = 2), 2^nd 6^th sternites as in male, 6^th sternite 0.23 mm long (N = 2), 7^th sternite 0.42 mm long, 1.82 × length of sternite 6 (N = 2), setose, mostly covered elsewhere with scales; sternite 8 not sclerotized; ovipositor: needle part 0,85-0.90 mm long, 2.02-2.14 × length sternite 7 (N = 2). Other characters as in male.

Figure 9
Asphondylia fluminensis Maia, sp. nov., female: (A) 5^th flagellomere, (B) Wing, (C) Foreleg, tarsal claw and empodium, lateral view, (D) Abdomen, lateral view.

Pupa (Fig. 10A): Color: brownish. Body length: 2.70-3.15 mm (N = 10). Head (Fig. 10B): dorsal plate 0.15-0.17 mm long, 0.40-0.46 mm wide (N = 7), antennal horn 0.20-0.26 mm long (N = 10), conical, pointed, inner margin serrated, distal part shorter than basal part; apical seta 0.04 mm long (N = 7), antennal width 0.06-0.07 mm; one upper facial horn conical, 0.02 mm long (N = 5); three lower facial horns aligned, 0.01 mm long (N = 5); two pairs of lower facial papillae: one pair setose, the other bare; three pairs of lateral facial papillae: one pair setose, two bare; upper cephalic margin thickened laterally, face with lateral projection. Thorax: prothoracic spiracle 0.10-0.11 mm long (N = 6), 1.3 × antennal basal width, setiform, not curved (N = 6) (Fig. 10C), integument wrinkled (Fig. 10D). Abdomen: segments 2-8 with transverse rows of crescent dorsal spines at basal half (Fig. 10E); posterior row with 19-27 spines in the 2^nd segment, 17-25 in the 3^rd, 18-24 in the 4^th, 15-22 in the 5^th, 15-23 in the 6^th, 12-19 in the 7^th and 7-8 in the 8^th.

Figure 10
Asphondylia fluminensis Maia, sp. nov., pupa: (A) General aspect, ventral view, (B) Head, frontal view, (C) Prothoracic spiracle, (D) Thoracic integument, dorsal view, (E) 7^th abdominal segment, dorsal view.

Larva: Body: 2.10 mm long (N = 1); head 0.05 mm long, 0.8 mm wide (N = 2). Spatula (Fig. 11A) quadridentate, 0.21 mm long (N = 1), lateral teeth longer than mesal, 0.035 mm long, mesal teeth 0.002-0.03 mm long (N = 2); three setose lateral papillae on each side of spatula. Terminal segment 0.06 mm long, four pairs of terminal papillae (three setose, one shorter than the others, and one corniform) (N = 1).

Figure 11
Asphondylia spp.: (A) A. fluminensis Maia, sp. nov., larva, spatula and associated papillae, ventral view, (B) Flower bud galls on Erythroxylum ovalifolium Peyr. (Erythroxylaceae), (C) A. erythroxylisMöhn, 1959, larva, spatula, ventral view, redrawn from Möhn, 1959 (original drawing without scale).

Gall: On flower bud, globoid, green, glabrous, one-chambered on Erythroxylum ovalifolium Peyr (Erythroxylaceae) (Fig. 11B).

Material examined: Holotype male, BRAZIL: Rio de Janeiro State, Maricá, APA de Maricá, 11.VIII.1990, V. Maia col. (MNRJ-ENT1-69782). Paratypes: MALES - same data as holotype: 1 ♂ (MNRJ-ENT1-69783); 01.IX.1997: 2 ♂♂ (MNRJ-ENT1-69786, MNRJ-ENT1-69787); Arraial do Cabo, 11.VIII.1990: 2 ♂♂ (MNRJ-ENT1-69784, MNRJ-ENT1-69785); FEMALES - same data as holotype: 3 ♀♀ (MNRJ-ENT1-69788: 1, MNRJ-ENT1-69789: 2); PUPAL-EXUVIAE - same data as holotype: 12 pupal exuviae (1: MNRJ-ENT1-69791, 4: MNRJ-ENT1-69793, 4: MNRJ-ENT1-69795, 3: MNRJ-ENT1-69796); 01.IX.1997: 2 pupal exuviae (MNRJ-ENT1-69797); 07.VIII.1998: 3 pupal exuviae (MNRJ-ENT1-69790); 28.IX.1990: 1 pupal exuviae (MNRJ-ENT1-69794); Carapebus, 29.VII.1998: 4 pupal exuviae (MNRJ-ENT1-69792); THIRD-INSTAR LARVAE - APA de Maricá, 18.V.2021: 1 larva (MNRJ-ENT1-69798); 08.IX.1988: 2 larvae (MNRJ-ENT1-69799).

Additional material: Rio de Janeiro state: Maricá, APA de Maricá, MALES - same data as holotype: (MNRJ-ENT1-69800, MNRJ-ENT1-69801); PUPAL-EXUVIAE - same data as holotype: 11 pupal exuviae (1: MNRJ-ENT1-69803, 6: MNRJ-ENT1-69806, 4: MNRJ-ENT1-69809); Carapebus, 29.VII.1998, 6 pupal exuviae (MNRJ-ENT1-69802); 26.IX.1998: 5 pupal exuviae (3: MNRJ-ENT1-69804, 2: MNRJ-ENT1-69805); APA de Maricá, 07.VIII.1988: 3 pupal exuviae (MNRJ-ENT1-69807); 01.IX.1997: 1 pupal exuvia (MNRJ-ENT1-69808).

Etymology: The name “fluminensis” means “of Rio de Janeiro State”.

Geographic distribution (based on gall records on the host plant): Brazil, Rio de Janeiro State, Rio de Janeiro (Grumari), Maricá, Arraial do Cabo and Carapebus (Maia, 2013).

Remarks:Asphondylia erythroxylisMöhn 1959 is the only previously described congeneric species on Erythroxylum P. Browne. It induces fruit galls on E. mexicanum and is known from El Salvador (Gagné & Jaschhof, 2021). The new species differs from A. erythroxylis in the following morphological characters: (1) larva: the mesal teeth of A. fluminensis is clearly deeper than those of A. erythroxylis (Fig. 11C); (2) female: the body of A. fluminensis is longer and the last three flagellomeres are shorter in the new species than in A. erythroxylis. Furthermore, the last three flagellomeres are subequal in length in the new species, while in A. erythroxylis they are progressively shorter toward apex (Table 2). The male and pupa of A. erythroxylis are unknown.

Table 2
Comparative length of females and larvae of Asphondylia fluminensis Maia, sp. nov. and A. erythroxylisMöhn, 1959 (Diptera, Cecidomyiidae). Data on A. erythroxylis were obtained from literature.

Length:	Asphondylia fluminensis, sp. nov.	Asphondylia erythroxylis Möhn, 1959
Female
Body	2.80‑2.90 mm	2.20 mm
Scape	110µ	118µ
Pedicel	50‑70µ	58µ
1st flagellomere	230‑280µ	207µ
5th flagellomere	150‑180µ	156‑157µ
9th flagellomere	100µ	100µ
10th flagellomere	50µ	70µ
11th flagellomere	40µ	48‑49µ
12th flagellomere	40µ	36µ
Palpus
1st segment	20µ	23‑24µ
2nd segment	50µ	42µ
3rd segment	70µ	70‑72µ
Larva
Body	2.10 mm	-
Spatula	210µ	293µ

Asphondylia marambaiensis Maia, sp. nov. (Figs. 12A 14F)

Diagnosis: Male hypoproct rounded apically, deeply bilobed, ovipositor with needle part about 1.13-1.16 × length 7^th sternite; pupa: antennal horn 0.23-0.29 mm long, upper facial horn single and conical, three lower frontal horns aligned, 8^th abdominal segment with 6-9 dorsal spines in the posterior row.

Figure 12
Asphondylia marambaiensis Maia, sp. nov., male: (A) 5^th flagellomere, (B) Palpus, (C) Midleg, tarsal claw and empodium, lateral view, (D) 5^th abdominal segment-terminalia, ventral view, (E) Terminalia, dorsal view.

Male: Body length: 3.50-3.65 mm long (N = 2). Head: 0.50 mm long, 0.50 mm wide (N = 2), eye facets circular, closely appressed; antennae: flagellomeres 1 and 2 not fused, scape obovate, setose, 0.12 mm long, 0.07 mm wide (N = 2), pedicel globose, setose, 0.07 mm long, 0.06 mm wide (N = 2), 1^st 12^th flagellomeres cylindrical, all 0.05 mm wide, circumfila longitudinally wavy, dense, anastomosing, equally spread along segments (Fig. 12A), 1^st flagellomere 0.25-0.26 mm long (N = 2), 2^nd flagellomere 0.22-0.23 mm long (N = 2), 3^rd 4^th flagellomeres 0.20-0.21 mm long (N = 2), 5^th 6^th flagellomeres 0.20-0.22 mm long (N = 2), 7^th flagellomere 0.20-0.23 mm long (N = 2), 8^th flagellomere 0.20-0.22 mm long (N = 2), 9^th flagellomeres 0.20-0.21 mm long (N = 2), 10^th flagellomere 0.19 mm long (N = 2), 11^th flagellomere 0.18-0.19 mm long (N = 2), 12^th flagellomere 0.17 mm long (N = 2); proportion flagellomere neck-node 1:10; frons with 25 setae (N = 1); mouthparts: labrum, hypopharynx and labella kneaded; palpus (Fig. 12B) 0.18-0.19 mm long: 1^st segment globoid, 0.02 mm long, 0.02 mm wide (N = 2), 2^nd segment cylindrical, 0.05 mm long, 0.02 mm wide, 3^rd segment fusiform, 0.11-0.12 mm long, 0.02 mm wide (N = 2), all segments with setae. Thorax: scutum with two dorsocentral rows of setae, setae more abundant anteriorly, two groups of lateral setae more abundant anteriorly, extending from base to distal margin, scales intermixed; scutellum with scattered setae; anepimeron setose; remaining pleural sclerites bare; legs: tarsal claws curved beyond midlength, isomorphic, empodium as long as claws (Fig. 12C); wing: length 2.70 mm (N = 1). Abdomen (Fig. 12D): trichoid sensilla not visible; 1^st 7^th tergites sclerotized, rectangular with a posterior row of setae, few scattered lateral setae and mostly covered elsewhere with scales, 1^st tergite shorter than other tergites, ⅓ × length of 2^nd tergite, 8^th tergite band-like, bare; 2^nd 8^th sternites more sclerotized than tergites, rectangular, as long as tergites, with a posterior row of setae, several setae and midlength, few lateral setae, and mostly covered elsewhere with scales; 8^th sternite with scattered setae and mostly covered elsewhere with scales. Terminalia (Fig. 12E): gonocoxite short and stout, 0.14-0.16 mm long, 0.07-0.08 mm wide (N = 2); gonostylus ovoid, 0.05 mm long, 0.05 mm wide; hypoproct deeply bilobed, rounded apically.

Female: Body length: 4.00-4.50 mm (N = 3). Head (Fig. 13A): 0.45-0.55 mm long, 0.50-0.55 mm wide (N = 3), facets circular, antennae: scape 0.13-0.14 mm long, 0.05-0.08 mm wide (N = 3), pedicel 0.07-0.08 mm long, 0.07-0.08 mm wide (N = 3), 1^st 10^th flagellomeres cylindrical, all 0.05 mm wide, circumfila comprising two longitudinal bands connected subbasally and apically by two transverse bands (Fig. 13B), 1^st flagellomere 0.28-0.31 mm long (N = 3), 2^nd flagellomere 0.22-0.25 mm long (N = 3), 3^rd flagellomere 0.21-0.23 mm long (N = 3), 4^th flagellomere 0.21-0.24 mm long (N = 3), 5^th flagellomere 0.22-0.24 mm long (N = 3), 6^th flagellomere 0.21-0.24 mm long (N = 3), 7^th flagellomere 0.22-0.23 mm long (N = 3), 8^th flagellomere 0.20-0.22 mm long, 9^th flagellomere 0.16-0.18 mm long, 10^th flagellomere 0.10-0.13 mm long, 11-12^th flagellomeres 0.06-0.07 mm long (Fig. 13C), proportion flagellomere neck-node 1:11; mouthparts: labrum 0.08-0.10 mm long, 0.035 mm wide (N = 1), hypopharynx 0.14 mm long, 0.035 mm wide (N = 1), labellum 0.06-0.07 mm long, 0.03 mm wide at midlength (N = 2), palpus 0.24 mm long (N = 3): 1^st segment globose 0.03 mm long and 0.03-0.04 mm wide (N = 3), 2^nd segment cylindrical 0.05 mm long, 0.03 mm wide at midlength (N = 3), 3^rd segment ellipsoidal, 0.16 mm long, 0.025-0.03 mm wide at midlength (N = 3). Thorax: wing length: 2.80-3.10 mm (N = 3) (Fig. 13D). Abdomen (Fig. 13E): trichoid sensillae not visible, 1^st 7^th tergites as in male, 8^th tergite with posterior margin with lobes 0.12-0.14 mm long (N = 3), 2), sternites 2^nd 6^th sternites as in male, 6^th sternite 0.35-0.39 mm long (N = 2), 7^th sternite 0.56-0.59 mm long, 1.51-1.60 × length of 6^th sternite, setose, mostly covered elsewhere with scales; sternite 8 not sclerotized; ovipositor: needle part 1.13-1.16 mm long, 1.16-2.02 × length sternite 7 (N = 2). Other characters as in male.

Figure 13
Asphondylia marambaiensis Maia, sp. nov., female: (A) Head, ventral view, (B) 5^th flagellomere, (C) 10^th 12^th flagellomeres, (D) Wing, (E) Abdomen, lateral view.

Pupa (Fig. 14A): Color: brownish. Body length: 3.50-4.00 mm (N = 4). Head (Fig. 14B): dorsal plate 0.43 mm long, 0.17 mm wide (N = 2), antennal horn 0.23-0.29 mm long (N = 3), conical, rounded apically, inner margin serrated, distal part shorter than basal part; apical seta 0.025 mm long (N = 1); antennal width 0.08 mm (N = 3), one upper facial horn conical, 0.04 mm long (N = 3); three lower facial horns aligned, 0.03 mm long (N = 3); two pairs of lower facial papillae: one pair setose, the other bare; three pairs of lateral facial papillae: one pair setose, two bare; upper cephalic margin thickened laterally, face with lateral projection. Thorax: prothoracic spiracle 0.11-0.13 mm long (N = 4), 1.37-1.62 × antennal width, setiform, conspicuously curved (N = 2) (Fig. 14C), integument wrinkled (Fig. 14D). Abdomen (Fig. 14E): segments 2-8 with transverse rows of crescent dorsal spines at basal half; posterior row with 11-15 spines in the 2^nd segment, 13-16 in the 3^rd, 12-16 in the 4^th, 11-17 in the 5^th, 10-17 in the 6^th, 9-12 in the 7^th and 6-9 in the 8^th.

Figure 14
Asphondylia marambaiensis Maia, sp. nov., (A-E) pupa: (A) General aspect, ventral view, (B) Cephalic horns, ventral view, (C) Prothoracic spiracle, (D) Thoracic integument, dorsal view, (E) Abdomen, dorsal view, (F) Leaf galls on Lantana fucata Lindl. (Verbenaceae).

Larva: unknown.

Gall: on leaf, globose, green, hairy, complex, one-chambered on Lantana fucata Lindl. (Verbenaceae) (Fig. 14F).

Material examined: Holotype male, BRAZIL: Rio de Janeiro State, Mangaratiba, Ilha da Marambaia, Praia do Kutuca, 21.VI.2010, Rodrigues, A. col. (MNRJ-ENT1-69810). Paratypes: same data as holotype: 1 ♂ (MNRJ-ENT1-69811); 3 ♀♀ (MNRJ-ENT1-69812, MNRJ-ENT1-69813, MNRJ-ENT1-69814), and 5 pupal exuviae (2: MNRJ-ENT1-69815, 3: MNRJ ENT1-69816).

Etymology: The name “marambaiensis” refers to the type-locality.

Geographic distribution (based on gall records on the host plant): Brazil, Rio de Janeiro State, Mangaratiba (Ilha da Marambaia) (Rodrigues et al., 2014), Arraial do Cabo (Carvalho-Fernandes et al., 2016); Espírito Santo State, Santa Teresa (Maia et al., 2014); and Minas Gerais State, Serra do Cipó (Coelho et al., 2009).

Remarks: In the Neotropical Region, Asphondylia camaraeMöhn 1959, known from El Salvador and Colombia, is the only congeneric species associated with the genus Lantana. It induces galls on unopened flowers of Lantana camara and Lantana urticifolia (Verbenaceae) (Gagné & Jaschhof, 2021). Asphondylia marambaiensis differs from A. camarae in the following morphological characters: (1) male: body, antennomeres and 3^rd segment of palpi shorter in A. camarae than in Asphondylia marambaiensis; 1^st and 2^nd segments of palpi longer in A. camarae than in Asphondylia marambaiensis; (2) female: body, pedicel, and 1^st, 5^th, 9^th and 12^th flagellomeres shorter in A. camarae than in Asphondylia marambaiensis, scape longer or equal (Table 3). The pupa of A. camarae is unknown.

Table 3
Comparative length of adults of Asphondylia marambaiensis Maia, sp. nov. and A. camaraeMöhn 1959 (Diptera, Cecidomyiidae). Data on A. camarae were obtained from literature.

Length:	Asphondylia marambaiensis Maia, sp. nov.	Asphondylia camarae Möhn, 1959
Male
Body	3.50‑3.65 mm	2.80 mm
Scape	120µ	120µ
Pedicel	60‑70µ	54µ
1st flagellomere	250‑260µ	201µ
5th flagellomere	200‑220µ	155µ
9th flagellomere	200‑210µ	136µ
10th flagellomere	190µ	127µ
11th flagellomere	180‑190µ	126µ
12th flagellomere	170µ	124µ
Palpus
1st segment	22µ	32‑33µ
2nd segment	50µ	72‑73µ
3rd segment	110‑120µ	103µ
Female
Body	4.00‑4.50 mm	3.00 mm
Scape	130‑140µ	142µ
Pedicel	70‑80µ	62µ
1st flagellomere	280‑310µ	278µ
5th flagellomere	220‑250µ	210µ
9th flagellomere	160‑180µ	144µ
10th flagellomere	100‑130µ	100µ
11th flagellomere	60‑70µ	69µ
12th flagellomere	60‑70µ	42µ
Palpus
1st segment	30µ	44µ
2nd segment	50µ	90µ
3rd segment	160µ	153µ

Asphondylia varroniae Maia, sp. nov. (Figs. 15A 18C)

Diagnosis: Male hypoproct simple, pointed apically ovipositor with needle part about 1.89-200 × length 7^th sternite; pupa: antennal horn 0.24-0.27 mm long, upper facial horn single and conical, three lower frontal horns aligned, 8^th abdominal segment with 10-11 dorsal spines in the posterior row, larva: spatula with lateral and mesal teeth subequal in length, mesal teeth rounded apically, three setose lateral papillae on each side of spatula.

Figure 15
Asphondylia varroniae Maia, sp. nov., male: (A) Head, ventral view, (B) 5^th flagellomere, (C) Wing, (D) Foreg, tarsal claw and empodium, lateral view, (E) Abdomen, lateral view, (F) Terminalia, dorsal view.

Male: Body length: 2.70-2.80 mm (N = 3). Head (Fig. 15A): 0.40 mm long, 0.40 mm wide (N = 2), eye facets hexagonal, closely appressed; antennae: flagellomeres 1 and 2 not fused, scape obovate, setose, 0.10-0.11 mm long, 0,05 mm wide (N = 3), pedicel globose, setose, 0.05-0.06 mm long, 0.05-0.06 mm wide (N = 3), 1^st 10^th flagellomeres cylindrical (11^th 12^th flagellomeres missing), circumfila longitudinally sinuous, equally spread along segments (Fig. 15B), 1^st flagellomere 0.16-0.20 mm long, 0.05-0.06 mm wide (N = 4), 2^nd flagellomere 0.19-0.20 mm long, 0.04 mm wide (N = 3), 3^rd flagellomere 0.17-0.18 mm long, 0.04-0.05 mm wide (N = 3), 4^th flagellomere 0.15-0.17 mm long, 0.04 mm wide (N = 3), 5^th flagellomere 0.16-0.19 mm long, 0.04 mm wide (N = 3), 6^th flagellomere 0.16-0.17 mm long, 0.04 mm wide (N = 2), 7^th flagellomere 0.16 mm long, 0.04 mm wide (N = 2), 8^th flagellomere 0.15-0.16 mm long, 0.04 mm wide (N = 2), 9^th flagellomere 0.15 mm long, 0.04 mm wide (N = 2), 10^th flagellomere 0.14 mm long, 0.04 mm wide (N = 1), proportion flagellomere neck-node: 1:10; frons corrugated; mouthparts: labrum long-attenuate 0.07 mm long, 0.03 mm wide; hypopharynx of the same shape as labrum, 0.10 mm long, 0.03 mm wide (N = 1) with long, anteriorly-directed lateral setulae; labella elongate and convex, with lateral and mesal setae (labella length and width not measured as they are smashed); palpus 0.12-0.14 mm (N = 2), 1^st segment globoid, 0.02 mm long, 0.2 mm wide (N = 2), 2^nd segment cylindrical, 0.04-0.05 mm long, 0.02-0.025 mm wide (N = 2), 3^rd segment cylindrical. 0.06-0.07 mm long, 0.025 mm wide (N = 2), all segments with setae. Thorax: scutum with two dorsocentral rows of setae, setae more abundant anteriorly, and one irregular lateral row of setae on each side, scales intermixed; scutellum with scattered setae; anepimeron and anepisternum setose; remaining pleural sclerites bare; legs: tarsal claws curved beyond midlength, isomorphic, empodium as long as claws (Fig. 15C); wing (Fig. 15D): length 1.90-2.50 mm (N = 4). Abdomen (Fig. 15E): trichoid sensillae not visible; 1^st 7^th tergites sclerotized, rectangular with a posterior row of setae, few lateral setae, and mostly covered elsewhere with scales, 1^st tergite shorter than other tergites, ½ × length of 2^nd tergite, tergite8^th band-like, bare; 2^nd 8^th sternites more sclerotized than tergites, rectangular; 2^nd 6^th sternites with a posterior row of setae, several setae at midlength, few lateral setae, and mostly covered elsewhere with scales; 7^th sternites with a posterior row of setae, several mesal setae, few lateral setae, 8^th sternite entirely covered with setae, more abundant posteriorly, and mostly covered elsewhere with scales. Terminalia (Fig. 15F): gonocoxite short and stout, 0.12-0.13 mm long (N = 3), gonostylus spherical, 0.04 mm long, 0.04 mm wide (N = 1), hypoproct simple, pointed apically.

Female: Body length: 2.50-2.70 mm (N = 3). Head: 0.40 mm long, 0.40 mm wide (N = 1), antennae: scape 0.10 mm long, 0.06 mm wide (N = 2), pedicel 0.05 mm long, 0.05 mm wide (N = 2), 1^st 11^th flagellomeres cylindrical, 12^th spheroid, all 0.04 mm wide, circumfila comprising two longitudinal bands connected subbasally and apically by two transverse bands (Fig. 16A), 1^st flagellomere 0.21-0.23 mm long (N = 3), 2^nd flagellomere 0.17-0.21 mm (N = 3), 3^rd flagellomere 0.16-0.18 mm (N = 3), 4^th flagellomere 0.16-0.17 mm (N = 3), 5^th flagellomere 0.14-0.17 mm long (N = 3), 6^th flagellomere 0.14-0.16 mm (N = 3), 7^th flagellomere 0.13-0.14 mm long (N = 3), 8^th flagellomeres 0.11 mm (N = 3); 9^th flagellomere 0.09-0.10 mm (N = 3), 10^th flagellomere 0.07-0.08 mm long (N = 3), 11^th flagellomere 0.06 mm long (N = 3), 12^th flagellomere 0.04 mm long (N = 3) (Fig. 16B); proportion flagellomere neck-node: 1:13; mouthparts: labrum 0.07 mm long, 0.04 mm wide, hypopharynx 0.11 mm long, 0.04 mm wide, labellum 0.05 mm long, 0.03 mm wide at midlength (N = 3), palpus 0.15 mm long (N = 1): 1^st segment globoid 0.03 mm long and 0.02 mm wide (N = 1), 2^nd segment cylindrical 0.05 mm long, 0.015 mm wide at midlength (N = 1), 3^rd segment fusiform, 0.07 mm long and 0.015 mm wide at midlength (N = 1). Thorax: wing length: 1.90-2.00 mm (N = 2); legs: isomorphic, tarsal claw more robust than in male (Fig. 16C). Abdomen (Fig. 16D): trichoid sensillae not visible, 1^st 7^th tergites sclerotized, rectangular with a posterior row of setae, few lateral setae and mostly covered elsewhere with scales, 8^th tergite with distal margin with lobes 0.09 mm long (N = 1), 2^nd 6^th sternites as in male, 6^th sternite 0.20-0.21 mm long (N = 3); 7^th sternite 0.40-0.45 mm long, 2.00-2.14 × length sternite 6 (N = 3), setose and mostly covered elsewhere with scales; sternite 8 not sclerotized; ovipositor: needle part 0.80-0.85 mm long, 1.89-2.00 × length sternite 7 (N = 3). Other characters as in male.

Figure 16
Asphondylia varroniae Maia, sp. nov., female: (A) 5^th flagellomere, (B) 9^th 12^th flagellomeres, (C) Foreg, tarsal claw and empodium, lateral view, (D) Abdomen, lateral view.

Pupa: Color: brownish. Body length: 2.80-3.30 mm (N = 5). Head (Fig. 17A): dorsal plate 0.40-0.42 mm long, 0.15-0.20 mm wide (n = 4), antennal width 0.07-0.08 mm (N = 5), antennal horn 0.24-0.28 mm long (N = 4), conical, pointed, inner margin serrated, distal part shorter than basal part; apical seta 0.03 mm long (N = 2); one upper facial horn conical, 0.02-0.03 mm long (N = 4); three lower facial horns aligned, 0.015 mm long (N = 4); two pairs of lower facial papillae: one pair setose, the other bare; three pairs of lateral facial papillae: one pair setose, two bare; upper cephalic margin thickened laterally, face with lateral projection. Thorax: integument wrinkled (Fig. 17B), prothoracic spiracle 0.08-0.09 mm long (N = 5), setiform, slightly curved, as long as antennal width (Fig. 17C). Abdomen (Fig. 17D): segments 2-8 with transverse rows of crescent dorsal spines; posterior row with 21-27 spines in the 2^nd segment, 25-32 in the 3^rd, 25-30 in the 4^th, 25-29 in the 5^th, 21-30 in the 6^th, 16-19 in the 7^th and 10-11 in the 8^th (N = 6).

Figure 17
Asphondylia varroniae Maia, sp. nov., pupa: (A) Head, frontal view, (B) Prothoracic spiracle, (C) Thoracic integument, dorsal view, (D) 4^th 7^th abdominal segments, dorsal view.

Larva (Fig. 18A): Body length: 1.90-2.40 mm (N = 2); head 0.06 mm long, 0.07 mm wide (N = 1). Spatula (Fig. 18B) quadridentate, 0.22-0.24 mm long (N = 2), lateral and mesal teeth subequal in length, 0.025 mm long, mesal teeth rounded apically, lateral teeth pointed apically; three setose lateral papillae on each side of spatula, terminal segment 0.18 mm long (N = 1), four pairs of terminal papillae (three setose, one shorter than the others, and one corniform).

Figure 18
Asphondylia spp.: (A-C) Asphondylia varroniae Maia, sp. nov.: (A) General aspect, (B) Spatula and associated papillae, ventral view, (C) Flower bud galls on Varronia curassavica Jacq. (Cordiaceae), (D) Asphondylia cordiaeMöhn, 1959, redrawn from Möhn, 1959, spatula and associated papillae, ventral view (original drawing without scale).

Gall: On flower bud, green, globoid, with white trichomes, one-chambered on Varronia curassavica Jacq. (Boraginaceae) (Fig. 18C). The host plant was reported as Cordia verbenacea D.C. (synonym of Varronia curassavica) and the gall-inducer misidentified as Asphondylia cordiaeMöhn 1959 in Maia, 2001.

Material examined: Holotype male, BRAZIL, São Paulo, Bertioga, Jardim São Lourenço, 25.IV.2005, V. Maia col. (MNRJ ENT1-69817). Paratypes: MALES - Fazenda Pinto, 26.IV.2005: 2 ♂♂ (MNRJ ENT1-69818, MNRJ ENT1-69820); Itaguaré, 24.VI.2005: 1 ♂ (MNRJ ENT1-69819); FEMALES - Itaguaré, 22.II.2005: 1 ♀ (MNRJ ENT1-69821); 30.VII.2004: 1 ♀ (MNRJ ENT1-69823); Fazenda Pinto, 26.IV.2005: 1 ♀ (MNRJ ENT1-69822); PUPAL EXUVIAE - Fazenda Pinto, 26.IV.2005: 1 pupal exuvia (MNRJ ENT1-69820); Itaguaré, 22.II.2005: 1 pupal exuvia (MNRJ ENT1-69821), Jardim São Lourenço, 15.V.2005: 1 pupal exuviae (MNRJ ENT1-69825); Itaguaré, 24.VI.2004: 3 pupal exuviae (MNRJ ENT1-69826); 16.V.2005: 1 pupal exuvia (MNRJ ENT1-69827); 30.VII.2004: 2 pupal exuviae (MNRJ ENT1-69828); Rio de Janeiro, APA de Maricá, 03.VII.1998: 1 pupal exuvia (MNRJ ENT1-69824); Carapebus, 26.IX.1998: 1 pupal exuvia (MNRJ ENT1-69829); THIRD INSTAR LARVAE - APA de Maricá, 05.VI.1998: 1 larva (MNRJ ENT1-69830); Carapebus, 26.IX.1998: 1 larva (MNRJ ENT1-69829).

Additional material: Rio de Janeiro, APA de Maricá, VII.1998: 1 ♀ (MNRJ ENT1-69831); Bertioga, Jardim São Lourenço, 25.IV.2005: 2 pupal exuviae (MNRJ ENT1-69832).

Etymology: The name “varroniae” is the genitive of the plant genus.

Geographic distribution (based on gall records on the host plant): Brazil: Minas Gerais, Lagoa Santa (Cerrado); Espírito Santo, Anchieta-Piúma (Atlantic Forest) Rio de Janeiro State, Rio de Janeiro, Maricá, Reserva Biológica União, Saquarema, Araruama, Arraial do Cabo, Quissamã, Carapebus, Campos de Goitacazes, São João da Barra (Atlantic Forest); São Paulo State, Bertioga, Ubatuba (Atlantic Forest); Santa Catarina, Babitonga (Atlantic Forest); Rio Grande do Sul, Porto Alegre (Atlantic Forest) (Maia, 2021).

Remarks: There is only one previously known species of Asphondylia on Cordiaceae, Asphondylia cordiaeMöhn, 1959 on Cordia dentata Poir. described from El Salvador. The new species differs from Asphondylia cordiae in the following characters: (1) larva: body longer, spatula with mesal teeth wider, stalk thinner and anterior end longer in the new species than in A. cordiae (Fig. 18B × Fig. 18D); (2) adults: body and female flagellomeres longer in the new species than in A. cordiae (Table 4); (3) male: hypoproct simple only in A. varroniae. The pupa of A. cordiae is unknown.

Table 4
Comparative length of adults and larva of Asphondylia cordiaeMöhn, 1959 and A. varroniae Maia, sp. nov. (Diptera, Cecidomyiidae). Data on the first species were obtained from literature.

Length:	Asphondylia cordiae Möhn, 1959	Asphondylia varroniae Maia, sp. nov.
Male
Body	2.10 mm	2.70‑2.80 mm
Scape	100µ	100‑110µ
Pedicel	46‑48µ	50‑60µ
1st flagellomere	195µ	160‑200µ
5th flagellomere	158‑160µ	160‑190µ
9th flagellomere	-	150µ
10th flagellomere	-	140µ
11th flagellomere	-	-
12th flagellomere	-	-
Palpus
1st segment	19‑20µ	20µ
2nd segment	28‑30µ	40‑50µ
3rd segment	72µ	60‑70µ
Female
Body	2.20‑2.30 mm	2.50‑2.70 mm
Scape	112‑114µ	100µ
Pedicel	48‑50µ	50µ
1st flagellomere	198µ	210‑230µ
5th flagellomere	130µ	140‑170µ
9th flagellomere	86µ	90‑100µ
10th flagellomere	63µ	70‑80µ
11th flagellomere	50‑52µ	60µ
12th flagellomere	32‑34µ	40µ
Palpus
1st segment	19‑20µ	30µ
2nd segment	48‑52µ	50µ
3rd segment	70‑72µ	70µ
Larva
Body	1.70‑1.80 mm	1.90‑2.40 mm
Spatula	198µ	220‑240µ

Asphondylia xerezi Maia, sp. nov. (Figs. 19A 21C)

Diagnosis: Male hypoproct rounded apically, deeply bilobed, ovipositor with needle part about 2.45 × length 7^th sternite; pupa: antennal horn 0.20-0.25 mm long, upper facial horn single and conical, three lower frontal horns not aligned, 8^th abdominal segment with 7-10 dorsal spines in the posterior row, larva: spatula with lateral and mesal teeth subequal in length, mesal teeth rounded apically, three setose lateral papillae on each side of spatula.

Figure 19
Asphondylia xerezi Maia, sp. nov.: (A) male terminalia, lateral view, (B-G) female, (B) head, ventral view, (C) 5^th flagellomere, (D) 10^th 12^th flagellomeres, (E) Wing, (F) Hindleg, tarsal claw and empodium, lateral view, (G) Ovipositor, ventral view.

Male: Body: 3.20 mm long (N = 1). Head: 0.45 mm long, 0.45 mm wide (N = 1), eye facets circular, closely appressed; antennae: flagellomeres 1 and 2 not fused, scape obovate, setose, 0.06 mm long, 0.06 mm wide (N = 1), pedicel globose, setose, 0.05 mm long, 0.06 mm wide (N = 1), 1^st 7^th flagellomeres cylindrical, all 0.04 mm wide (8^th to 12^th flagellomeres missing), circumfila longitudinally wavy, dense, anastomosing, equally spread along segments, 1^st flagellomeres 0.22 mm long (N = 1), 2^nd 7^th flagellomeres 0.17-0.18 mm long (N = 1), proportion flagellomere neck-node 1:09; frons smashed; mouthparts smashed. Thorax: scutum with two dorsocentral rows of setae, setae more abundant anteriorly, two groups of lateral setae more abundant anteriorly, extending from base to distal margin, scales intermixed; scutellum with scattered setae; anepimeron setose; remaining pleural sclerites bare; legs: tarsal claws curved beyond midlength, empodium as long as claws; wing smashed. Abdomen: trichoid sensilla not visible; 1^st 7^th tergites sclerotized, rectangular with a posterior row of setae, few scattered lateral setae and mostly covered elsewhere with scales, 8^th tergite tergite8^th band-like, bare; 2^nd 8^th sternites sclerotized, rectangular, narrower than tergites, with a posterior row of setae, several setae and midlength, few lateral setae, and mostly covered elsewhere with scales; 8^th sternite with setae at ⅔ distal and mostly covered elsewhere with scales. Terminalia (Fig. 19A): gonocoxite short and stout, 0.15 mm long, 0.07 mm wide (N = 1); gonostylus ovoid, 0.06 mm long, 0.05 mm wide (N = 1); hypoproct deeply bilobed, rounded apically.

Female: Body length: 3.20 mm (N = 1). Head (Fig. 19B): 0.40 mm long, 0.35 mm wide, antennae: scape 0.09 mm long, 0.05 mm wide, pedicel 0.05 mm long, 0.05 mm wide, 1^st 11^th flagellomeres cylindrical, all 0.04 mm wide, circumfila comprising two longitudinal bands connected subbasally and apically by two transverse bands (Fig. 19C), flagellomeres 1 and 2 not fused, 1^st flagellomere 0.24 mm long, 2^nd 6^th flagellomeres 0.17-0.18 mm long, 7^th flagellomere 0.15 mm long (N = 4), 8^th flagellomere 0.14 mm long (N = 1), 9^th flagellomere 0.09 mm long (N = 1), 10^th flagellomere 0.07 mm long, 11 flagellomere 0.06 mm long, 12^th flagellomere 0.04 mm long (Fig. 19D); proportion flagellomere neck-node 1:15; mouthparts: labrum 0.07 mm long, 0.04 mm wide (N = 1), hypopharynx 0.11 mm long, 0.04 mm wide (N = 1), labellum 0.07 mm long, 0.04 mm wide at midlength, with 5 pairs of mesal setae (N = 1); palpus 0.22 mm long (N = 1): 1^st segment globose 0.02 mm long, 0.02 mm wide (N = 2), 2^nd segment cylindrical 0.05 mm long, 0.02 mm wide at midlength (N = 2), 3^rd segment claviform 0.15 mm long and 0.02-0.03 mm wide at midlength (N = 2). Thorax: wing length: 2.35 mm (N = 1) (Fig. 19E); tarsal claws more sclerotized and robust than in male (Fig. 19F). Abdomen: trichoid sensillae not visible, 1^st 7^th tergites as in male, 8^th tergite with posterior margin with lobes 0.07-0.08 mm long (N = 2), 2^nd 6^th sternites as in male, 6^th sternite 0.17-020 mm long (N = 2), 7^th sternite 0.35-45 mm long (N = 1), 2.05-2.25 × length sternite 6 (N = 2), setose (except basally), mostly covered elsewhere with scales; sternite 8 not sclerotized; ovipositor (Fig. 19G): needle part 0.86 mm long (N = 1), 2.45x length sternite 7 (N = 1). Other characters as in male.

Pupa (Fig. 20A): Color: brownish. Body length: 2.90-3.20 mm (N = 3). Head (Fig. 20B): dorsal plate 0.38-0.40 mm long, 0.18-0.20 mm wide (N = 2); antennal horn 0.20-0.25 mm long (N = 3), conical, pointed, distal part longer than basal part, inner margin serrated; dorsal plate 0.20 mm long, 0.39-0.40 mm wide (N = 3), apical seta 0.05 mm long (N = 3); one upper facial horn conical, 0.07-0.11 mm long (N = 3); three lower facial horns not aligned, 0.04 mm long (N = 2); two pairs of lower facial papillae: one pair setose, the other bare; three pairs of lateral facial papillae: one pair setose, two bare; upper cephalic margin thickened laterally; face with pronounced lateral projection. Thorax: prothoracic spiracle 0.10 mm long (N = 2), as long as antennal basal width, setiform, slightly curved (N = 5) (Fig. 20C), integument wrinkled (Fig. 20D). Abdomen: segments 2-8 with transverse rows of crescent dorsal spines at basal half; posterior row with 26-30 spines in the 2^nd segment (N = 3), 24-31 in the 3^rd (N = 3), 24-29 in the 4^th (N = 3), 20-28 in the 5^th (N = 3), 20-22 in the 6^th (N = 3), 13-18 in the 7^th (N = 3), 7-10 in the 8^th (N = 3).

Figure 20
Asphondylia xerezi Maia, sp. nov., pupa: (A) General aspect, ventral view, (B) Head, frontal view, (C) Prothoracic spiracle, (D) Thoracic integument, dorsal view.

Larva (Fig. 21A): Body: 2.20 mm long (N = 1); head retracted. Spatula (Fig. 21B) quadridentate, 0.24 mm long (n = 1), lateral and mesal teeth subequal in length (0.02 mm long) (N = 1), lateral teeth more pointed than mesal; three setose lateral papillae on each side of spatula. Terminal segment smashed.

Figure 21
Asphondylia spp.: (A-C) Asphondylia xerezi Maia, sp. nov., (A) General aspect, ventral view, (B) Spatula, ventral view, (C) Fruit gall on Heliotropium sp. (Heliotropiaceae), (D) Asphondylia tournefortiaeRübsaamen, 1915, larva, spatula and associated papillae, ventral view, redrawn from Rübsaamen, 1915 ((original drawing without scale).

Gall: on fruit, globoid, green, glabrous, multichambered on Heliotropium sp. (Heliotropiaceae) (Fig. 21C).

Material examined: Holotype male, BRAZIL, Rio de Janeiro, Mangaratiba, Ilha da Marambaia, Praia Grande, 16.XII.2009, A.R. Rodrigues leg. (MNRJ ENT1-69833). Paratypes: FEMALES - Praia do Kutuca, 18.VII.2010: 2 ♀♀ (MNRJ ENT1-69834, MNRJ ENT1-69835); PUPAL EXUVIAE - same data as holotype, 1 pupal exuvia (MNRJ ENT1-69837); Praia do Kutuca, 18.VII.2020: 2 pupal exuviae (MNRJ ENT1-69838); PUPA - 1 pupa (MNRJ ENT1-69836); THIRD INSTAR LARVA - Praia do Kutuca, 25.II.2011: 1 larva (MNRJ ENT1-69839).

Etymology: The species is named in honor of Dr. Roberto Xerez (Universidade Federal Rural do Rio de Janeiro) responsible for collection permission in the locality-type (area of the Brazilian Navy).

Geographic distribution: Brazil, Rio de Janeiro State, Mangaratiba municipality (Rodrigues et al., 2014).

Remarks: There is only one previously known species of Asphondylia on Heliotropiacae, A. tournefortiaeRübsaamen 1915 on Heliotropium angustiflorum (Ruiz & Pav.) Govaerts (reported as Tournefortia angustiflora Ruiz & Pav.) and Myriopus volubilis Small (reported as T. volubilis L) from Brazil.

Adults of Asphondylia xerezi Maia, sp. nov. have longer body and shorter scape, pedicel and flagellomeres (Table 5) than in A. tournefortiae. The larval spatula of A. xerezi is longer than in A. tournefortiaeRübsaamen, 1915. In addition, mesal and lateral teeth are subequal in length the new species, while in A. tournefortiae mesal teeth are shorter than lateral ones (Fig. 21B × Fig. 21D).

Table 5
Comparative length of adults and larva of A. tournefortiaeRübsaamen 1915 and A. xerezi Maia, sp. nov. (Diptera, Cecidomyiidae). Data on the first species were obtained from literature.

Length:	Asphondylia tournefortiae Rübsaamen, 1915	Asphondylia xerezi Maia, sp. nov.
Male
Body	2.90‑3.00 mm	3.20 mm
Scape	118µ	60µ
Pedicel	72µ	50µ
1st flagellomere	256µ	220µ
5th flagellomere	216µ	170‑180µ
9th flagellomere	218µ	-
10th flagellomere	-	-
11th flagellomere	-	-
12th flagellomere	-	-
Palpus
1st segment	28‑30µ	-
2nd segment	46‑48µ	-
3rd segment	100µ	-
Female
Body	-	3.20 mm
Scape	-	90µ
Pedicel	-	50µ
1st flagellomere	-	240µ
5th flagellomere	-	170‑180µ
9th flagellomere	-	90µ
10th flagellomere	-	70µ
11th flagellomere	-	60µ
12th flagellomere	-	40µ
Palpus
1st segment	-	20µ
2nd segment	-	50µ
3rd segment	-	150µ
Larva
Body	-	2.20 mm
Spatula	212µ	240µ

• https://zoobank.org/C7DC7A88-F5D8-4356-9E9C-85994FFDE18F

ACKNOWLEDGMENTS:

I would like to thank Dr. Alene Ramos Rodrigues for collecting specimens of Asphondylia marambaiensis and A. xerezi, the Brazilian Navy for collection permission and infrastructure, and the Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro for financial support.

• The author is grateful to Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ) for financial support (number process: E 26/210.300/2021).

• Published with the financial support of the “Programa de Apoio às Publicações Científicas Periódicas da Universidade de São Paulo”

REFERENCES

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Edited by

Edited by:

Carlos José Einicker Lamas

Publication Dates

• Publication in this collection
  18 Oct 2024
• Date of issue
  2024

History

• Received
  11 Sept 2023
• Accepted
  28 Nov 2023
• Published
  05 Feb 2024