1. Introduction
The family Lonchaeidae (Diptera: Tephritoidea), commonly known as lance flies, comprises a group of acalyptrate dipterans characterized by stout bodies measuring approximately 3–6 mm in length. Members of this family are typically metallic green, blue, or black in coloration, and have abundant fine setae and black halteres [1,2]. This family has a broad global distribution and currently includes 611 described species, classified into ten genera [3].
Lonchaeids exhibit a wide variety of behaviors. Most species consume decaying plant material during their larval stage; however, some develop on healthy plant tissue [4,5,6,7] or act as secondary invaders of fruits associated with other insects (usually Tephritidae) [8,9]. Furthermore, a few species exhibit predatory behavior toward other arthropods at specific developmental stages [10,11,12].
In the Neotropical region, the genera Dasiops Rondani, 1856, and Neosilba Waddill&Weems, 1978 contain the highest number of described species, some of which can be considered pests of fruit and vegetables from different plant families [13,14,15,16,17].
Neosilba batesi Curran (Diptera: Lonchaeidae) has been recorded in Colombia, Peru, Central America, Mexico, and the United States [3,8,18,19]. This species is polyphagous, typically saprophagous, and opportunistic on fruits previously infested by Anastrepha spp. (Diptera: Tephritidae) [8,20,21,22]. It takes advantage of wounds caused by prior oviposition or exit holes made by larvae emerging from the fruit. However, N. batesi can occasionally act as a primary invader, as it is sometimes the only species that emerges from fruits collected in the field [23,24].
Recently, N. batesi has been reported in association with avocado (Persea americana) fruits in some regions of Mexico [25,26]. However, these reports lack rigorous species identification and a systematic assessment of its phytosanitary status. Additionally, avocado producers in Michoacán and Jalisco have reported premature fruit drop potentially linked to this species.
Despite these circumstances, no formal study has yet confirmed the identity of N. batesi or evaluated its ecological or phytosanitary significance in avocado crops. This is relevant because Mexico is one of the world’s leading producers and exporters of avocado fruit [27], with Michoacán and Jalisco accounting for nearly 87% of national production [28]. Therefore, the objectives of this work are to: 1. confirm the taxonomic and molecular identity of N. batesi; 2. provide photographic documentation to support its recognition; and 3. generate data on its geographic distribution and relationship with avocado in Mexico.
2. Materials and Methods
2.1. Sampling and Taxonomic Identification of Insects
From November 2022 to January 2023, inspections were conducted in avocado plantations in the municipalities of Tingüindín, Ario de los Rosales, Los Reyes, and Tancítaro (Michoacán state), and Quitupan and Tuxpan (Jalisco state) (Table 1). At each site, we collected fallen Hass fruits infested with Diptera larvae, as well as tree-hanging fruits exhibiting spots or symptoms of rot. A total of 104 and 43 fruits of each type, respectively, were collected.
Fruits from each location were stored separately and transported to the Laboratory of Agrobiology at Universidad Michoacana de San Nicolás de Hidalgo in Uruapan, Michoacán, Mexico. All fruit samples from each site were placed in plastic containers (30 × 20 × 15 cm), covered with organza cloth, and held until adult emergence.
Adult specimens were preserved in 70% ethanol, then sexed, counted, and identified according to [8] and [23]. Photographs of newly emerged larvae from the fruits, pupae, adults, and male genitalia were taken with a Canon EOS Rebel T7 (Tokyo, Japan) camera fitted with a 4× microscope objective. Focus stacking was performed in Helicon Focus 8.2.2 Pro., and final image editing was performed in GIMP 2.10.34.
2.2. DNA Extraction and Sequencing
Genomic DNA was individually extracted from three morphologically identified Neosilba batesi specimens (two males and one female) using the DNeasy Blood&Tissue Kit (QIAGEN, Hilden, Germany), following the manufacturer’s protocol for insect tissues, without modifications.
A 508 bp fragment of the 5′ region of the mitochondrial cytochrome c oxidase subunit I (COI) gene was amplified using the Platinum™ Taq DNA Polymerase High Fidelity Kit (Thermo Fisher Scientific, Waltham, MA, USA). The primers used were COI-FW: 5′-TTATAATTGGDGGWTTTGGWAATTG-3′ [34] and HC02198: 5′-TAAACTTCAGGGTGACCAAAAAATCA-3′ [35]. PCR conditions consisted of an initial denaturation at 94 °C for 1 min, followed by 35 cycles of 94 °C for 15 s, 55 °C for 30 s, and 68 °C for 1 min, with a final extension at 72 °C for 5 min.
PCR products were visualized on 1% agarose gels stained with ethidium bromide and purified using the Wizard® SV Gel and PCR Clean-Up System (Promega, Madison, WI, USA). Amplicons were sequenced using the Sanger method on an Applied Biosystems™ 3130/3130xl Genetic Analyzer.
Sequence chromatograms were visualized and edited manually using BioEdit v7.7.1 [36]. Multiple sequence alignment was performed using ClustalW [37], with subsequent manual adjustments to ensure positional homology. Sequence identity was confirmed via BLAST against the NCBI GenBank database (available online:
Phylogenetic analysis was conducted using MEGA11 [38]. The best-fit nucleotide substitution model was determined using a hierarchical likelihood ratio test (hLRT) and comparison of Bayesian Information Criterion (BIC) scores across 31 candidate models. The General Time Reversible model with a discrete Gamma distribution (GTR + G) was identified as the best fit (BIC = 4194.459). Phylogenetic reconstruction was performed using the Maximum Likelihood (ML) method. Initial heuristic trees were generated via Neighbor-Joining (NJ) and BioNJ algorithms based on pairwise distances, with the final topology corresponding to the highest log-likelihood value.
To root the phylogenetic tree, sequences of Silba adipata, Silba fumosa, Lonchaea iona, and Lonchaea cristula (All Diptera: Lonchaeidae) were included as outgroup taxa. These species belong to the subfamily Lonchaeinae and were selected to ensure taxonomic relevance and provide intergeneric divergence for robust phylogenetic inference.
2.3. Observation and Description of Damages in Avocado Fruits
The study was carried out in a 5 ha Hass avocado orchard in Los Reyes, Michoacán (19°35′17.20″ N, 102°23′4.82″ W; 1592 m a.s.l.). From January to April 2024, we conducted four sampling events, selecting tree-hanging fruits of various sizes that exhibited rot symptoms or necrotic epidermal spots.
Each fruit was measured for length and width, then placed individually in a plastic cup filled with vermiculite to serve as a pupation substrate. Cups were covered with organza cloth and checked daily to record and count emerged adults, which were preserved in 70% ethanol.
At every sampling site, we performed visual observations of fly behavior and photographed infested fruits to document larval presence.
3. Results
3.1. Identification and Phylogenetic Analyses
Flies of the family Lonchaeidae that emerged from avocados collected from the ground (n = 131) and trees (n = 85) in nine orchards were identified as N. batesi Curran (Figure 1), based on their chaetotaxy and the morphology of the male genitalia. Females were assigned to N. batesi because there are no taxonomic keys available for females of the genus Neosilba, and no males of other species were detected.
The larvae are musciform in shape and translucent white, with a slender and cylindrical body tapering toward the anterior end. They possess a pair of short, dark, and sclerotized posterior spiracles, characteristic of Lonchaeidae (Figure 1A). The puparia are oval, slightly broader at the front, reddish-brown in color, and exhibit remnants of the posterior spiracles on the caudal part (Figure 1B).
Adults (Figure 1C) are glossy and bluish black, measuring approximately 3–4 mm in length, with reddish eyes, black legs, a setulose lunule, plumose arista, and calypters bearing a group of long black setae, typical of the genus. In males, the epandrium is elongated, and the parameres are bilobed, with the medial lobe being slightly narrower (Figure 1D).
The COI sequences obtained from two males and one female, each comprising 470 base pairs of the partial mitochondrial COI gene, were identical to one another (GenBank accession numbers: PV243309, PV243310, PV243311; Table 1). BLAST analysis (99% query coverage) showed the highest similarity to Neosilba zadolicha (97.05%, accession KR262649.1), followed by Neosilba pendula (95.92%, accession OQ160349.1). Sequence identity with N. batesi (specimen CSCA_17X541, accession MW283302.1) from Florida was lower, at 93.86%. This relatively low similarity contrasts with the morphological identification of the specimens as N. batesi, suggesting potential taxonomic incongruence. However, COI sequences from N. batesi specimens from Veracruz, Mexico (PV504763, PV504764, PV504765), recently reported by [29] and included in this study, showed >99% identity with our sequences.
The phylogenetic analysis (Figure 2) supports the taxonomic assignment of the newly generated sequences to N. batesi, placing them within a well-supported clade corresponding to the genus Neosilba. However, these sequences formed a distinct genetic lineage, clearly separated from both the N. batesi specimen from Florida and the clade containing N. zadolicha and N. pendula. Notably, limitations in the available sequence data were evident, as each of the reference taxa was represented by a single sequence in GenBank, thereby restricting the depth of phylogenetic comparisons and the robustness of taxonomic resolution.
3.2. Distribution of Neosilba batesi in the Study Area
From November 2022 to January 2023, specimens of N. batesi emerged from the nine sites sampled in the states of Michoacan and Jalisco, Mexico. These sites ranged in elevation from 1301 to 2206 m a.s.l. (Table 2, Figure 3).
During fruit collection and transport, several larvae emerged and most reached the pupal stage; however, pupal mortality was high. Only adults of each sex were counted.
Females consistently outnumbered males, though the female-to-male ratio rarely exceeded 2:1. In addition, specimens of Drosophila melanogaster and other Drosophila spp. emerged from fallen fruits.
3.3. Neosilba batesi and Its Association with Avocado Fruits
In the avocado plantation in Los Reyes, Michoacán, 157 fruits were collected from trees across four inspections in 2024. Fruits infested by N. batesi larvae measured, on average, 5.40 cm in length and 3.90 cm in width. The average number of larvae that emerged per fruit was 9.61. Although most larvae successfully reached the pupal stage, mortality in this stage was high. The maximum dimensions recorded for infested fruits were 7.95 cm in length and 5.05 cm in width (Table 3).
Field observations indicated that, in apparently healthy developing fruits on the tree, females deposit eggs at the junction of the pedicel with the fruit (Figure 4A). Occasionally, several females can be found ovipositing simultaneously on the same fruit, especially when it shows signs of rot. In fallen fruits, oviposition occurs in the exposed pulp at the pedicel attachment site (Figure 4B) or within wounds on the exocarp. Females can also lay their eggs in fruits with previous ovipositions and larvae inside. No direct oviposition through the exocarp was observed.
Eggs are whitish, elongated, and may be laid singly or in clusters (Figure 4C). Early infestation is signaled by external symptoms, such as a purple ring at the fruit’s base (Figure 4D) or purple to reddish spots that may partially (Figure 4E) or completely cover the surface of the fruit (Figure 4F). Notably, healthy avocados can grow and develop normally even when adjacent to infested ones, including those on the same peduncle (Figure 4G).
As infestation progresses, internal decay and pedicel rot lead to fruit drop, although some fruits desiccate and mummify while remaining attached (Figure 4H). Dissection of infested fruits reveals larvae of various instars feeding on the mesocarp (Figure 4I).
Larvae consume pulp at different stages of decomposition (Figure 5A,B) and may exit through wounds in the exocarp, either at the pedicel scar or at impact-induced openings. No exit holes produced by larvae through the exocarp were observed. Puparia develop either in the soil or within the fruit (Figure 5C), including mummified avocados that remain on the tree (Figure 4H).
4. Discussion
In this study, N. batesi was identified in association with avocados both on the tree and on the ground. In tree fruits, females exhibit opportunistic oviposition behavior by exploiting potential wounds or openings at the junction between the pedicel and the fruit. Conversely, on fallen fruits, they deposit their eggs in areas where the pulp is exposed, even when the fruit is in an advanced state of decomposition.
These findings are consistent with previous reports. For example, Ref. [8] documented the first association of N. batesi with fallen avocados in the state of Chiapas, Mexico. Later, Ref. [21] classified this species as a saprophagous fly associated with decaying fruits, and Ref. [22] identified N. batesi as a fruit-decomposing species in the soil, associated with Persea americana cv. Criollo, Fuerte and Hass in Michoacán, Mexico. A related association was also partially reported by [18] in FL, USA, where the species was found in wounded avocados and captured in traps. Conversely, a study on the diversity of Tephritoidea in Colombia [24] reports an association of N. batesi with avocado and other fruit species; however, the characteristics of the collected fruits are not specified.
Neosilba batesi is distributed across countries in South, Central, and North America [6], and has been reported to infest fruits of sweet orange (Citrus sinensis L.), papaya (Carica papaya L.), mango (Mangifera indica L.), Inga spp., peach palm (Guilielma gasipaes Kunth), fig (Ficus carica L.), soursop (Annona muricata L.), and other wild Annona spp. [8,20,23,29].
Several species of this genus, including N. zadolicha, Neosilba certa, N. glaberrima, N. pendula, Neosilba parva, and other Neosilba spp., have been associated with fruits of Persea americana in Brazil [40,41]. Among these, only N. glaberrima has been reported in Mexico [8,23,29]. No male specimens of N. glaberrima were detected in the present study.
The phylogenetic analysis supports the morphological identification, placing the newly obtained sequences from Michoacán within the same clade as other N. batesi from Veracruz sequences recently described by [29]. In contrast, N. batesi_CSCA 17X541, collected in Florida and submitted to GenBank in 2020 (accession number MW283302.1), clustered in a separate clade. This record lacks an associated publication and supporting taxonomic evidence, leading to the same conclusion reached by [29]: the identification of this isolate requires confirmation. Furthermore, no additional sequence data are currently available for N. batesi, N. zadolicha, or N. pendula. Therefore, further molecular data are needed to achieve a more accurate taxonomic resolution for these species.
During the study conducted in the avocado orchard in Los Reyes, Michoacán, fruits infested with N. batesi larvae (from 27 January to 18 April 2024) exhibited characteristics that would prevent their commercialization. These fruits were typically small to medium in size, falling below marketable calibers. Additionally, they appeared visually immature and contained a variable number of larvae. Notably, many pupae failed to reach adulthood, likely due to the unfavorable conditions within the containers where they were confined.
In harvested or fallen fruits, the presence of laid eggs can be detected with the naked eye at the pedicel–fruit attachment point. Additionally, the chorion may still be visible after the eggs have hatched and the larva has emerged. Infestation by N. batesi larvae in tree fruits can be visually identified early on, typically by the presence of a purple ring at the base of the fruit (at the pedicel junction), along with distinct purple to reddish discolorations that may extend to cover the entire fruit surface. Most of these fruits eventually fall to the ground, although some remain attached to the pedicel and can easily be detached.
The premature detachment of fruits can be attributed to several factors. Larval feeding activity creates galleries and induces internal decomposition, leading to structural damage that weakens the fruit’s attachment to the pedicel. Furthermore, it is probable that microorganisms are involved, enhancing the rotting process and thereby triggering the fruit’s natural abscission mechanisms. However, these hypotheses require further confirmation.
Fallen fruits in advanced stages of decomposition usually harbor large numbers of eggs and larvae at different instars. These larvae feed on the fruit pulp, which corroborates their saprophagous feeding habits. Based on our observations, such fallen fruits appear to serve as key reservoirs for future fly generations. Therefore, collecting and removing them from the ground could represent an effective cultural practice to reduce N. batesi populations in the field.
We hypothesize that N. batesi females prefer to oviposit on fallen, decomposing fruit rather than on apparently healthy fruit still on the tree. However, further research is required to confirm this behavioral pattern. Interestingly, no direct ovipositions through the epidermis of the fruits were observed, and some visually healthy fruits lacking larvae, wounds, or signs of disease continued to develop normally, even in the presence of N. batesi-infested fruits around on the tree and in the soil.
This observation suggests that the presence of a pre-existing pathogen or damage between the pedicel and the avocado might influence oviposition behavior on tree fruits, which is consistent with the opportunistic behavior associated with this species.
Traps baited with hydrolyzed protein or torula yeast have been reported as effective tools for capturing N. batesi in other crops [19,29,42] and may be useful for monitoring or managing this species in avocado orchards. However, comparative field studies are needed to evaluate their effectiveness under local conditions.
Given the signs of oviposition, the obvious symptoms of infestation, the small dimensions of the fruits that are typically infested and premature fruit drop caused by N. batesi, it is unlikely that infested fruits would reach the avocado export chain, which includes multiple strict phytosanitary inspections in both orchards and packing facilities.
Conceptualization, B.A.L.-S. and C.P.I.-R.; methodology, D.G.-G., O.M.-G., D.V.G.-B., M.K., B.A.L.-S. and C.P.I.-R., validation, B.A.L.-S. and C.P.I.-R.; formal analysis, M.K., B.A.L.-S. and C.P.I.-R.; investigation, D.G.-G., D.V.G.-B., B.A.L.-S. and C.P.I.-R.; resources, B.A.L.-S., O.M.-G. and D.V.G.-B.; writing—original draft preparation, B.A.L.-S. and C.P.I.-R.; writing—review and editing, B.A.L.-S. and C.P.I.-R.; visualization, B.A.L.-S. and C.P.I.-R.; supervision, B.A.L.-S. and C.P.I.-R.; project administration, B.A.L.-S.; funding acquisition, B.A.L.-S. and O.M.-G. All authors have read and agreed to the published version of the manuscript.
Not applicable.
Data is contained within the article.
We thank Hugo Armando Garcia for granting us access to the avocado orchards and for his support during the field samplings.
The authors declare no conflicts of interest.
This article has been republished with a minor correction to the readability of figure 3’s caption. This change does not affect the scientific content of the article.
Footnotes
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Figure 1 Neosilba batesi. (A) Lateral and dorsal view of larva; (B) puparium in ventral position; (C) female in lateral view; (D) male genitalia.
Figure 2 Phylogenetic tree inferred using the Maximum Likelihood method based on the General Time Reversible model [
Figure 3 Avocado orchards with the presence of Neosilba batesi (numbers 1, 2, 3, and 4 indicate the orchards mentioned in
Figure 4 Ovipositions, signs, and symptoms of Neosilba batesi larvae in avocado fruits. (A) Female ovipositing on an immature tree fruit. (B) Female ovipositing on a fallen fruit showing signs of rot and white mycelium. (C) Recently laid N. batesi eggs at the base of a fallen, rotting fruit. (D) Purple ring at the fruit’s base. (E) Infested fruit with a lateral purple spot. (F) Infested fruit is almost entirely covered by rot. (G) Infested and healthy fruits sharing the same peduncle. (H) Mummified fruit attached to the pedicel. (I) Larvae and internal rot within the fruit.
Figure 5 Decomposing avocado fruits with different developmental stages of N. batesi. (A,B) Larvae inside fruits in an advanced state of decomposition. (C) Larvae and pupae within a rotting fruit.
Accession numbers of sequences used in the phylogeny study, with corresponding organism name, submission date, country, and reference.
| Species/Voucher Code | Estate, Country | Accession GenBank | Reference |
|---|---|---|---|
| Neosilba batesi Mx-CIQA-Mich01 | Michoacan, Mexico | PV243309 | Nucleotide sequences generated by this work |
| Neosilba batesi Mx-CIQA-Mich02 | Michoacan, Mexico | PV243310 | |
| Neosilba batesi Mx-CIQA-Mich03 | Michoacan, Mexico | PV243311 | |
| Neosilba batesi INECOL_24/21 | Veracruz, Mexico | PV504763 | [ |
| Neosilba batesi INECOL_24/22 | Veracruz, Mexico | PV504764 | [ |
| Neosilba batesi INECOL_24/30 | Veracruz, Mexico | PV504765 | [ |
| Neosilba batesi CSCA 17X541 | FL, USA | MW283302.1 | unpublished |
| Neosilba zadolicha YSUW02121210 | Brazil | KR262649.1 | [ |
| Neosilba pendula IV18-A | Brazil | OQ160349.1 | [ |
| Neosilba glaberrima INECOL_24/43 | Veracruz, Mexico | PV504766 | [ |
| Neosilba glaberrima INECOL_24/52 | Veracruz, Mexico | PV504767 | [ |
| Neosilba recurva INECOL_24/32 | Veracruz, Mexico | PV522074 | [ |
| Neosilba recurva INECOL_24/39 | Veracruz, Mexico | PV522075 | [ |
| Neosilba recurva INECOL_24/40 | Veracruz, Mexico | PV522076 | [ |
| Neosilba recurva INECOL_24/46 | Veracruz, Mexico | PV522077 | [ |
| Neosilba recurva INECOL_24/51 | Veracruz, Mexico | PV522078 | [ |
| Neosilba sp. INECOL_24/31 | Veracruz, Mexico | PV504768 | [ |
| Neosilba sp. INECOL_24/41 | Veracruz, Mexico | PV504769 | [ |
| Neosilba sp. INECOL_24/42 | Veracruz, Mexico | PV504770 | [ |
| Neosilba sp. INECOL_24/42 | Veracruz, Mexico | PV504771 | [ |
| Neosilba sp. INECOL_24/44 | Veracruz, Mexico | PV504772 | [ |
| Neosilba flavitarsis INECOL24/1M | Veracruz, Mexico | PQ834830 | [ |
| Neosilba flavitarsis INECOL24/2F | Veracruz, Mexico | PQ834831 | [ |
| Neosilba flavitarsis INECOL24/3M | Veracruz, Mexico | PQ834832 | [ |
| Silba adipata 8041568-LCO | Turkey | MK450115 | unpublished |
| Silba adipata INECOL_24/10 | Veracruz, Mexico | PV504773 | [ |
| Silba adipata INECOL_24/11 | Veracruz, Mexico | PV504774 | [ |
| Silba adipata TJMM10 | Morelos, Mexico | OM949837 | unpublished |
| Silba fumosa ZFMK-TIS-2574287 | Germany | OP831869 | [ |
| Lonchaea cristula LCR5-1 | Colombia | MZ189773 | [ |
| Lonchaea iona ZFMK-TIS-2575260 | Germany | OP831877.1 | [ |
Sites with Neosilba batesi on avocado trees Persea americana c.v. Hass and number of emerged adults per orchard of confined soil and tree fruit.
| Site | Coordinates/ | Date of Fruit Collection | (Number of Fruits)/Emerged Adults | |
|---|---|---|---|---|
| Fallen Fruits | Tree Fruits | |||
| Tingüindín, Mich. | 19°43′46″ N 102°27′7″ W/1882 | 22 December 2022 | (13)/5♂♂ 11♀♀ | (3)/1♂♂ 3♀♀ |
| Ario de los Rosales, Mich. | 19°11′10″ N 101°39′52″ W, 2206 | 10 January 2023 | (8)/4♂♂ 7♀♀ | (6)/4♂♂ 7♀♀ |
| Los Reyes 1, Mich. | 19°35′17.20″ N 102°23′4.82″ W/1592 | 10 November 2022 | (19)/10♂♂ 16♀♀ | (4)/2♂♂ 9♀♀ |
| Los Reyes 2, Mich. | 19°34′34″ N 102°19′30″ W, 2141 | 14 December 2022 | (8)/6♂♂ 10♀♀ | (5)/3♂♂ 5♀♀ |
| Los Reyes 3, Mich. | 19°39′50″ N 102°24′51″ W, 1620 | 14 December 2022 | (10)/8♂♂ 6♀♀ | (5)/5♂♂ 7♀♀ |
| Los Reyes 4, Mich | 19°38′50″ N 102°23′39″ W, 1780 | 14 December 2022 | (6)/4♂♂ 7♀♀ | (4)/6♂♂ 8♀♀ |
| Tancítaro, Mich. | 19°23′28”N 102°25′18”W, 2020 | 14 January 2023 | (14)/4♂♂ 6♀♀ | (6)/3♂♂ 5♀♀ |
| Tuxpan, Jal. | 19°32′5″ N 103°27′50″ W, 1301 | 28 December 2022 | (11)/6♂♂ 8♀♀ | (4)/3♂♂ 5♀♀ |
| Quitupan, Jal. | 19°48′2″ N 102°48′32″ W, 2040 | 29 December 2022 | (15)/5♂♂ 9♀♀ | (6)/4♂♂ 6♀♀ |
1,2,3,4 Avocado orchards indicated on the map of the study area (
Size of avocados sampled from trees and number of Neosilba batesi larvae and adults obtained across four sampling dates.
| Collect Date | Number of Fruits Collected | Average Sizes of Infested Avocado Trees (cm) | Average Number of Emerged Larvae/Fruit | Average Number of Adults | |
|---|---|---|---|---|---|
| Long | Width | ||||
| 27 January 2024 | 18 | 7.95 | 5.05 | 4.94 | 1.72 |
| 3 February 2024 | 65 | 4.57 | 3.4 | 5.07 | 1.06 |
| 5 March 2024 | 44 | 4.05 | 3.2 | 15.09 | 2.5 |
| 18 April 2024 | 30 | 5.04 | 3.93 | 13.34 | 3.8 |
1. Jaagumagi, R. Lonchaeidae. Manual of Nearctic Diptera; McAlpine, J.F.; Peterson, B.V.; Shewell, G.E.; Teskey, H.J.; Vockeroth, J.R.; Wood, D.M. Chapter 62 Research Branch, Agriculture Canada: Ottawa, ON, Canada, 1987; Volume 2, pp. 791-797.
2. MacGowan, I.; Rotheray, G.E. Lonchaeidae (Lance flies). Manual of Afrotropical Diptera. Volume 3. Brachycera—Cyclorrhapha, excluding Calyptratae; Kirk-Spriggs, A.H.; Sinclair, B.J. South African National Biodiversity Institute: Pretoria, South Africa, 2021; pp. 1587-1596.
3. MacGowan, I. World Catalogue of the family Lonchaeidae (Diptera, Cyclorrhapha, Acalyptratae). Zootaxa; 2023; 5307, pp. 1-96. [DOI: https://dx.doi.org/10.11646/zootaxa.5307.1.1] [PubMed: https://www.ncbi.nlm.nih.gov/pubmed/37518661]
4. Norrbom, A.L.; Korytkowski, C.A. Lonchaeidae (Lance flies). Manual of Central American Diptera; Brown, B.V.; Borkent, A.; Cumming, J.M.; Wood, D.M.; Woodley, N.E.; Zumbado, M.A. NRC Research Press: Ottawa, ON, Canada, 2010; Volume 2, pp. 857-863.
5. Strikis, P.C.; De Deus, E.G.; Silva, R.A.; Pereira, J.D.B.; Jesus, C.R.; Massaro-Júnior, A.L. Conhecimento sobre Lonchaeidae na Amazônia brasileira. Moscas-das-Frutas na Amazónica Brasileira: Diversidade, Hospedeiros e Inimigos Naturais; Silva, R.A.; Lemos, W.P.; Zucchi, R.A. Embrapa: Macapá, Brazil, 2011; pp. 206-215.
6. MacGowan, I.; Okamoto, T. New species of Lonchaeidae (Diptera: Schizophora) from Japan and a re-evaluation of genus Setisquamalonchaea Morge. Entomol. Sci.; 2013; 16, pp. 196-202. [DOI: https://dx.doi.org/10.1111/ens.12003]
7. Juárez Maya, M.; Morales Galván, Ó.; Valdez Carrasco, J.; Illescas Salvador, D.; Gutiérrez Ruelas, J.; Illescas Riquelme, C.P. Morphology of immature stages of the black fig fly Silba adipata (Diptera, Lonchaeidae). Biodivers. Data J.; 2024; 12, e137798. [DOI: https://dx.doi.org/10.3897/BDJ.12.e137798] [PubMed: https://www.ncbi.nlm.nih.gov/pubmed/39619294]
8. McAlpine, J.F.; Steyskal, G.C. A revision of Neosilba McAlpine with a key to the world genera of Lonchaeidae (Diptera). Can. Entomol.; 1982; 114, pp. 105-137. [DOI: https://dx.doi.org/10.4039/Ent114105-2]
9. Strikis, P.C.; Prado, A.P. Neosilba (Tephritoidea: Lonchaeidae) species reared from coffee in Brazil, with description of a new species. Proceedings of the 7th International Symposium on Fruit Flies of Economic Importance: From Basic to Applied Knowledge; Salvador, Brazil, 10–15 September 2006; pp. 187-193.
10. Herard, F.; Mercadier, G. Natural enemies of Tomicus piniperda and Ips acuminatus (Coleoptera: Scolytidae) on Pinus sylvestris near Orléans, France: Temporal occurrence and relative abundance, and notes on eight predatory species. Entomophaga; 1996; 41, pp. 183-210. [DOI: https://dx.doi.org/10.1007/BF02764245]
11. Wermelinger, B. Development and distribution of predators and parasitoids during two consecutive years of an Ips typographus (Coleoptera, Scolytidae) infestation. J. Appl. Entomol.; 2002; 126, pp. 521-527. [DOI: https://dx.doi.org/10.1046/j.1439-0418.2002.00707.x]
12. Okamoto, T.; MacGowan, I.; Su, Z.H. Predation on the pollinating fig wasp of Ficus erecta by larvae of Silba sp. (Diptera: Lonchaeidae). Entomol. Sci.; 2012; 15, pp. 288-293. [DOI: https://dx.doi.org/10.1111/j.1479-8298.2012.00522.x]
13. Strikis, P.C.; Prado, A.P. A new species of genus Neosilba (Diptera: Lonchaeidae). Zootaxa; 2005; 828, pp. 1-4. [DOI: https://dx.doi.org/10.11646/zootaxa.828.1.1]
14. Nicácio, J.; Uchôa, M.A. Diversity of frugivorous flies (Diptera: Tephritidae and Lonchaeidae) and their relationship whit host plants (Angiospermae) in environments of south Pantanal region, Brazil. Fla. Entomol.; 2011; 94, pp. 443-466. [DOI: https://dx.doi.org/10.1653/024.094.0309]
15. Lemos, L.N.; Adaime, R.; Costa Neto, S.V.; Deus, E.G.; Jesus-Barros, C.R.; Strikis, P.C. New findings on Lonchaeidae (Diptera: Tephritoidea) in the Brazilian Amazon. Fla. Entomol.; 2015; 98, pp. 1227-1237. [DOI: https://dx.doi.org/10.1653/024.098.0433]
16. Sousa, E.M.; Louzeiro, L.R.F.; Strikis, P.C.; Souza-Filho, M.F.; Raga, A. Host plants and distribution records of lance flies (Diptera: Lonchaeidae) in São Paulo State, Brazil. EntomoBrasilis; 2021; 14, e942. [DOI: https://dx.doi.org/10.12741/ebrasilis.v14.e942]
17. Martins, D.S.; Fornazier, M.L.; Uramoto, K.; Guimarães, J.A.; Ferreira, P.S.F.; Ventura, J.A.; Guarçoni, R.C.; Culik, M.P.; Zanuncio Junior, J.S.; Fornazier, M.J. Frugivorous flies (Diptera: Tephritidae; Lonchaeidae) associated with guava tree: Species diversity, parasitoids and population fluctuation in the Espírito Santo state, Brazil. Pesqui. Agropecu. Trop.; 2024; 54, e77329. [DOI: https://dx.doi.org/10.1590/1983-40632024v5477329]
18. Ahlmark, K.; Steck, G.J. A new U.S. record for a secondary fruit infester Neosilba batesi (Curran) (Diptera: Lonchaeidae). Insecta Mundi; 1997; 11, 116.
19. Imbachi, K.; Quintero, E.; Manrique, M.; Kondo, T. Evaluación de tres proteínas hidrolizadas para la captura de adultos de la mosca del botón floral de la pitaya amarilla, Dasiops saltans Townsend (Diptera: Lonchaeidae). Rev. Corp. Cienc. Tecnol. Agropecu.; 2012; 13, pp. 159-166. [DOI: https://dx.doi.org/10.21930/rcta.vol13_num2_art:251]
20. Curran, C.H. New American Diptera No. 534. Am. Mus. Novit.; 1932; 534, pp. 1-15. Available online: https://www.biodiversitylibrary.org/bibliography/91295 (accessed on 14 June 2025).
21. Hernández-Ortiz, V. Diptera. Catálogo de Insectos y Ácaros Plaga de los Cultivos Agrícolas de México; Deloya-López, A.C.; Valenzuela-González, J.E. Sociedad Mexicana de Entomología A.C.: Xalapa, México, 1999; pp. 69-82.
22. Aluja, M.; Díaz-Fleischer, F.; Arredondo, J. Non host status of commercial Persea americana “Hass” to Anastrepha ludens, Anastrepha obliqua, Anastrepha serpentina and Anastrepha striata (Diptera: Tephritidae) in México. J. Econ. Entomol.; 2004; 97, pp. 293-309. [DOI: https://dx.doi.org/10.1093/jee/97.2.293] [PubMed: https://www.ncbi.nlm.nih.gov/pubmed/15154448]
23. Illescas-Riquelme, C.P.; González-Hernández, H.; Valdez-Carrasco, J.M.; Llanderal-Cázares, C.; Ruiz-Montiel, C. Lonchaeidae (Diptera: Tephritoidea) associated with the genus Annona in Mexico. Southwest. Entomol.; 2015; 40, pp. 121-130. [DOI: https://dx.doi.org/10.3958/059.040.0110]
24. Herrera, A.M.; Canal, N.A.; Agudelo-Martínez, J.C.; Pérez-Buitrago, N. Diversidad y ecología de Tephritoidea (Insecta: Diptera) en el norte de la Orinoquía colombiana. Rev. Biol. Colomb.; 2022; 70, pp. 423-436. [DOI: https://dx.doi.org/10.15517/rev.biol.trop.2022.48808]
25. APEAM. Neosilba batesi. Origen de la Especie y Cuidados Para Evitar su Incidencia en Los Huertos de Aguacate. Unidad de Investigación y Desarrollo de la APEAM. 2024; Available online: https://fliphtml5.com/dpfeq/morn/neosilba_batesi/ (accessed on 28 February 2025).
26. Reynoso, L.F. Llega Plaga Desde Colombia a Huertas de Aguacate en Uruapan: Investigador. Quadrantin Michoacán. 2024; Available online: https://www.quadratin.com.mx/principal/llega-plaga-desde-colombia-a-huertas-de-aguacate-en-uruapan-investigador/ (accessed on 30 February 2024).
27. Cruz-López, D.F.; Caamal-Cauich, I.; Pat-Fernández, V.G.; Gómez-Gómez, A.A.; Espinoza-Torres, L.E. Posicionamiento internacional del aguacate (Persea americana L.) producido en México. Rev. Mex. Agronegocios; 2020; 47, pp. 561-571.
28. SIAP. Aguacate Mexicano. Servicio de Información Agroalimentaria y Pesquera. 2024; Available online: https://www.gob.mx/cms/uploads/attachment/file/885638/Brochure_aguacate_2024.pdf (accessed on 14 June 2025).
29. Lasa, R.; Navarro-de-la-Fuente, L.; MacGowan, I.; Williams, T. A Complex of Lance Flies (Diptera: Lonchaeidae) Infesting Figs in Veracruz, Mexico, with the Description of a New Species. Insects; 2025; 16, 458. [DOI: https://dx.doi.org/10.3390/insects16050458] [PubMed: https://www.ncbi.nlm.nih.gov/pubmed/40429171]
30. Han, H.-Y.; Ro, K.-E. Molecular phylogeny of the superfamily Tephritoidea (Insecta: Diptera) reanalysed based on expanded taxon sampling and sequence data. J. Zool. Syst. Evol. Res.; 2016; 54, pp. 276-288. [DOI: https://dx.doi.org/10.1111/jzs.12139]
31. Talamonte de Oliveira, T.C.; Pereira Gomes, L.R.; Egan, S.P.; Brandão-Dias, P.F.P.; Riverón, A.Z.; Del Bianco Faria, L. Diversity of Diptera and their parasitoids associated with Inga vera Willd. (1806) (Fabaceae) with new host record in Minas Gerais, Brazil. Rev. Agrogeoambient.; 2024; 16, e20241918. [DOI: https://dx.doi.org/10.18406/2316-1817v16nunico20241918]
32. Reimann, A.; Rulik, B. The Lonchaeidae (Diptera) of the GBOL project, with the description of a new Priscoearomyia species. Contrib. Entomol.; 2024; 74, pp. 165-179. [DOI: https://dx.doi.org/10.3897/contrib.entomol.74.e127094]
33. Balseiro Teheran, F.J.; Uribe Soto, S.I. Códigos de barra de ADN para identificación de especies del género Lonchaea fallen 1820 (Diptera: Lonchaeidae) de Antioquia. Rev. Fac. Cienc.; 2021; 10, pp. 51-66. [DOI: https://dx.doi.org/10.15446/rev.fac.cienc.v10n2.94053]
34. Prosser, S.W.; deWaard, J.R.; Miller, S.E.; Hebert, P.D. DNA barcodes from century-old type specimens using next-generation sequencing. Mol. Ecol. Resour.; 2016; 16, pp. 487-497. [DOI: https://dx.doi.org/10.1111/1755-0998.12474] [PubMed: https://www.ncbi.nlm.nih.gov/pubmed/26426290]
35. Folmer, O.; Black, M.; Hoeh, W.; Lutz, R.; Vrijenhoek, R. DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Mol. Mar. Biol. Biotechnol.; 1994; 3, pp. 294-299. [PubMed: https://www.ncbi.nlm.nih.gov/pubmed/7881515]
36. Hall, T. BioEdit Sequence Alignment Editor. GitHub Pages. 2021; Available online: https://thalljiscience.github.io/ (accessed on 14 June 2025).
37. Thompson, J.D.; Higgins, D.G.; Gibson, T.J. CLUSTAL W: Improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Res.; 1994; 22, pp. 4673-4680. [DOI: https://dx.doi.org/10.1093/nar/22.22.4673] [PubMed: https://www.ncbi.nlm.nih.gov/pubmed/7984417]
38. Tamura, K.; Stecher, G.; Kumar, S. MEGA 11: Molecular Evolutionary Genetics Analysis Version 11. Mol. Biol. Evol.; 2021; 38, pp. 3022-3029. [DOI: https://dx.doi.org/10.1093/molbev/msab120] [PubMed: https://www.ncbi.nlm.nih.gov/pubmed/33892491]
39. Nei, M.; Kumar, S. Molecular Evolution and Phylogenetics; Oxford University Press: New York, NY, USA, 2000.
40. Uchôa, M.A.; Oliveira, I.; Molina, R.M.S.; Zucchi, R.A. Species diversity of frugivorous flies (Diptera: Tephritoidea) from hosts in the cerrado of the state of Mato Grosso do Sul, Brazil. Neotrop. Entomol.; 2002; 31, pp. 515-524. [DOI: https://dx.doi.org/10.1590/S1519-566X2002000400002]
41. Raga, A.; Souza-Filho, M.F.; Strikis, P.C.; Montes, S.M.N.M. Lance fly (Diptera: Lonchaeidae) host plants in the State of São Paulo, Southeast Brazil. Entomotropica; 2015; 30, pp. 57-68.
42. López-Bautista, E.; García-Sánchez, A.N.; Sierra-Gómez, U.A.; Tejeda-Reyes, M.A.; Illescas-Riquelme, C.P. Efecto nulo de trampas y cebos en la captura de la mosca negra del higo, Silba adipata. Avances en Agricultura Sostenible y Cambio Climático; Soto-Ortiz, R.; Avilés-Marín, S.M.; Brígido-Morales, J.G.; Escobosa-García, M.I. Astra: Jalisco, México, 2024; pp. 441-447.
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