Revising Spider Egg-predating Fly Records (Diptera: Sarcophagidae): New Insights on Prey, Taxonomy, and Distribution of Sarcophaga (Mehria) lorosa Hall in Southern South America

© 2024 Academia Sinica, Taiwan Open Access Revising Spider Egg-predating Fly Records (Diptera: Sarcophagidae): New Insights on Prey, Taxonomy, and Distribution of Sarcophaga (Mehria) lorosa Hall in Southern South America Filipe Macedo Gudin1,* , Diego Galvão de Pádua2, Pablo Ricardo Mulieri3, Benito CortésRivas4, Andrés Moreira-Muñoz5, and Rodrigo de Oliveira Araujo2,4 1Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Rua do Matão, Trav. 14, 101, Cidade Universitária, CEP 05508-090, São Paulo, São Paulo, Brazil. *Correspondence: E-mail: [email protected] (Gudin) 2Laboratorio de Entomología General y Aplicada, Centro de Investigación de Estudios Avanzados del Maule, Universidad Católica del Maule, Avenida San Miguel, 3605, Talca, Chile. E-mail: [email protected] (Pádua); [email protected] (Araujo) 3Consejo Nacional de Investigaciones Científicas y Técnicas, División Entomología, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Avenida Angel Gallardo, 470, Buenos Aires, Argentina. E-mail: [email protected] (Mulieri) 4Laboratorio de Ecología de Abejas, Departamento de Ciencias Biológicas y Químicas, Facultad de Ciencias Básicas, Universidad Católica del Maule, Avenida San Miguel, 3605, Talca, Chile. E-mail: [email protected] (Cortés-Rivas) 5Instituto de Geografía, Pontificia Universidad Católica de Valparaíso, Avenida Brasil 2241, Valparaíso, Chile. E-mail: [email protected] (Moreira-Muñoz) Received 7 July 2023 / Accepted 27 December 2023 / Published 16 April 2024 Communicated by Y. Miles Zhang Flesh flies (Diptera: Sarcophagidae) exhibit a wide range of feeding habits including necrophagy, coprophagy, kleptoparasitism, parasitism, and predation. Among them are species of Sarcophaga Meigen belonging to the subgenera Baranovisca Lopes and Mehria Enderlein that are specialized predators of spider eggs. These flies hover around spider webs and lay their larvae on the spider egg sac. While progress has been made on the taxonomy of Baranovisca and Mehria in recent decades, our knowledge about their biology, prey selection, and distribution remains limited, restricting our understanding of the evolutionary dynamics of Sarcophagidae-Araneae interactions. Here, we describe and illustrate the first record of S. (M.) lorosa Hall preying on egg sacs of Metepeira galatheae (Thorell) (Araneae: Araneidae) in Chile. The taxonomy of S. (M.) lorosa is revised, with two new junior synonyms proposed: Weyrauchimyia ruficauda Lopes and Tibana, syn. nov., and Arachnidomyia travassosi Tibana and Mello, syn. nov. Furthermore, we present an annotated catalog that comprehensively reviews the existing records of spider egg-predating Sarcophagidae, and provide an overview of the evolution of Sarcophagidae-Araneae interactions. Our catalog includes information on at least four species of Baranovisca and 10 species of Mehria that have been documented as preying on eggs from species of various spider families, such as Araneidae, Cheiracanthiidae, Clubionidae, Philodromidae, Salticidae, and Tetragnathidae. These records cover all biogeographical regions except the Afrotropical. Our results enhance our understanding of the evolution of Sarcophagidae-Araneae interactions. Key words: Araneidae, Coevolution, Egg sacs, Metepeira, Oviposition strategy Citation: Gudin FM, de Pádua DG, Mulieri PR, Cortés-Rivas B, Moreira-Muñoz A, de Oliveira Araujo R. 2024. Revising spider egg-predating fly records (Diptera: Sarcophagidae): new insights on prey, taxonomy, and distribution of Sarcophaga (Mehria) lorosa Hall in southern South America. Zool Stud 63:07. doi:10.6620/ZS.2024.63-07. Zoological Studies 63: 7 (2024) doi:10.6620/ZS.2024.63-07 1 © 2024 Academia Sinica, Taiwan BACKGROUND Flesh flies (Diptera: Sarcophagidae) are often considered as necrophagous or coprophagous flies, although there are several lineages with parasitoid or predatory habits (Pape 1996; Pape and Dahlem 2010). The subfamily Miltogramminae contains a large clade of kleptoparasitic species (Piwczyński et al. 2017), while parasitoid and predatory larvae are frequently found in species of the subfamilies Paramacronychiinae and Sarcophaginae, attacking mostly invertebrates such as insects, snails, millipedes, earthworms, and spiders, but even producing myiasis in several species of vertebrates (Aldrich 1914; Pape 1996). For instance, the largest genus of the family, Sarcophaga Meigen, includes approximately 890 species classified in several subgenera worldwide (Buenaventura et al. 2017), some of which are adapted to parasitism or predation. Sarcophaga larvae of the subgenera Baranovisca Lopes and Mehria Enderlein are specialized predators of spider eggs; the females hover around and fly through the spider web, trying to reach the egg sac, where they lay one larva per attack (Lubin 1974; Hieber and Uetz 1990; Rayor and Uetz 1990; Hieber et al. 2002). The larva then enters the egg sac and eats the eggs, thus completing its development. Some flies of the families Bombyliidae, Chloropidae, Ephydridae, and Phoridae, and wasps of the family Ichneumonidae are also specialized parasitoids or predators of spider eggs (Sobczak et al. 2012; Fritzén and Sääksjärvi 2016; Villanueva-Bonilla et al. 2016; Gillung and Borkent 2017; Riccardi and Pádua 2021; Souza-Santiago et al. 2023). However, in some cases, the incidence of flesh flies is much higher than that of the other parasitoids/ predators (Hieber and Uetz 1990). The subgenus Baranovisca includes six species distributed in the Australasian and Oriental regions, while Mehria includes 15 species distributed in the Nearctic, Neotropical, and Palaearctic regions (Pape 1996). Although the taxonomy of these species has been improved with detailed descriptions and illustrations during the final decades of the last century (Lopes 1946 1959 1981 1985 1989; Cantrell 1981; Tibana and Mello 1992), information about their biology, prey, and distribution is restricted to a few species, which hinders a deeper understanding of the evolution of Sarcophagidae-Araneae interactions. Most records of flesh flies preying on spider eggs were recently listed (Gillung and Borkent 2017), but not all were included. Here, we record for the first time the interaction of Sarcophaga (Mehria) lorosa Hall preying on egg sacs of Metepeira galatheae (Thorell) (Araneae: Araneidae) during a desert bloom in the Copiapó and Huasco provinces from the Atacama region, Chile (Chávez et al. 2019). The taxonomy and nomenclature of S. (M.) lorosa is revised, including notes on type material and new distribution records. Furthermore, we propose two new junior synonyms: Weyrauchimyia ruficauda Lopes and Tibana, syn. nov., and Arachnidomyia travassosi Tibana and Mello, syn. nov. Finally, we present an annotated catalog that comprehensively reviews the existing records of spider egg-predating Sarcophagidae and provide an overview of the evolution of Sarcophagidae-Araneae interactions. MATERIALS AND METHODS Metepeira galatheae adults (n = 20: 5 ♂, 15 ♀) (Fig. 1A, B) and egg sacs (n = 21) (Fig. 1C) were manually collected at four localities from two provinces in the Atacama region during a desert bloom (16–17 November 2022): Copiapó Province: Sector Totoral (27°50'14"S, 71°05'05"W); and Huasco Province: Parque Nacional Llanos del Challe (28°08'55"S, 71°09'24"W), Quebrada Honda (27°59'22,30"S, 71°08'14"W), and Parque Eolico Cabo Leones (28°55'14"S, 71°26'38"W) (Fig. 2B). The collected egg sacs and spiders were deposited in Falcon tubes, closed with voile, and transported to the Laboratorio de Ecología de Abejas of the Universidad Católica del Maule (UCM), Talca, Chile. All egg sacs and spiders were kept separately in Falcon tubes at room temperature (27–30°C). Specimens of S. (M.) lorosa were identified based on examination of the holotype male deposited in the Natural History Museum (NHMUK) in London, United Kingdom. Additional material of S. (M.) lorosa was obtained from the Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” (MACN), Buenos Aires, Argentina. Morphological terminology follows Cumming and Wood (2017). Fly vouchers were deposited in UCM (5 ♂ and 1 ♀), and spider vouchers in UCM (13 ♀) and UNAP (5 ♂ and 2 ♀). Metepeira galatheae specimens were identified by Dr. Andres Taucare-Rios de la Universidad Arturo Prat (UNAP), Iquique, Chile. The catalog follows a format similar to that presented by Guimarães (1977) and Arnaud (1978). Sarcophagidae species are listed alphabetically according to their respective biogeographical regions, with valid names and junior synonyms accompanied by authorship. Published records of egg predation are given with the name of the fly followed by respective spider prey, author, year of publication, page number, locality, and notes on the record. Comments and emendations regarding taxonomy, nomenclature or reliability of the record are included in square brackets. Secondary page 2 of 14Zoological Studies 63: 7 (2024) © 2024 Academia Sinica, Taiwan Fig. 1. Record of Sarcophaga (Mehria) lorosa Hall (Diptera: Sarcophagidae) in Metepeira galatheae (Thorell) (Araneae: Araneidae). A–C, Female, male, and egg sac of M. galatheae, respectively; D, Puparium of S. (M.) lorosa. page 3 of 14Zoological Studies 63: 7 (2024) © 2024 Academia Sinica, Taiwan references citing only previous records were not included. The classification of Sarcophagidae follows Pape (1996), and the classification of spiders follows Gloor et al. (2017). Digital images were taken using a Leica MC170 HD digital camera attached to a Leica S8AP0 stereomicroscope with an LED illumination dome (Kawada and Buffington 2016). Multiple layers were stacked using the software Helicon Focus A8.1.0.0. Distributional data of S. (M.) lorosa were obtained directly from the labels and plotted on a map using SimpleMappr (Shorthouse 2010). When necessary, Fig. 2. A, Distribution map of Sarcophaga (Mehria) lorosa Hall, highlighting the type localities of the junior synonyms Arachnidomyia travassosi Tibana and Mello, syn. nov., and Weyrauchimyia ruficauda Lopes and Tibana, syn. nov.; B, Study areas in the Atacama region, Chile, during a desert bloom. Map created with SimpleMappr (https://www.simplemappr.net/). page 4 of 14Zoological Studies 63: 7 (2024) © 2024 Academia Sinica, Taiwan coordinates were estimated using Global Gazetteer Version 2.3. RESULTS Records of Sarcophaga (Mehria) lorosa in egg sacs of Metepeira galatheae A total of 21 egg sacs of M. galatheae were collected and reared. Six specimens of S. (M.) lorosa emerged from two egg sacs (Fig. 4): four flies (4 ♂) emerged between 27–28 November 2022 from one egg sac collected on 16 November 2022, and two flies (1 ♂ and 1 ♀) emerged on 30 November 2022 from one egg sac collected on 16 November 2022. We verified that no flies emerged from the remaining 19 egg sacs and manually opened them to search for fly puparia. Nineteen hatched puparia were found in seven egg sacs (Fig. 1D): Two egg sacs with one puparium, three egg sacs with two puparia, one egg sac with five puparia, and one egg sac with six puparia. One egg sac had a puparium and a dead specimen of S. (M.) lorosa. Therefore, approximately 42% of the egg sacs had been attacked by S. (M.) lorosa. TAXONOMY Sarcophagidae Macquart, 1834 Subfamily Sarcophaginae Macquart, 1835 Genus Sarcophaga Meigen, 1824 Subgenus Mehria Enderlein, 1928 Sarcophaga (Mehria) lorosa Hall, 1937 (Figs. 2A, 3, 4) Sarcophaga lorosa Hall, 1937: 367. Holotype male (NHMUK), examined (Fig. 3A, C, E). Type locality: Chile, Santiago. Weyrauchimyia ruficauda Lopes and Tibana, 1982: 142. Holotype male (MNRJ, lost). Type locality: Chile, Arica y Parinacota [as Tarapacá], Lluta. [Junior secondary homonym of Sarcophaga ruficauda Zetterstedt, 1838.]. Syn. nov. Arachnidomyia travassosi Tibana and Mello, 1992: 293. Holotype male (CNC), examined (Fig. 3B, D, F). Type locality: Chile, O’Higgins, La Leonera. Syn. nov. Material examined: Argentina: 1 ♂ (MACN): Neuquén, Parque Nacional Lanín, Ñorquinco, S 39°09,073 W 71°15,475, trampa Malaise, 9.i.2013, Olea, Mulieri and Patitucci leg.; Chile: 5 ♂, 1 ♀ (UCM): Huasco Province, Parque Nacional Llanos del Challe, 28°08'55"S, 71°09'24"W, reared from Metepeira galatheae egg sacs, 27–30.xi.2022, D.G. Pádua et al. leg. Diagnosis: Distinguished from the other New World species of subgenus Mehria by the following morphological features: (1) face with deep goldenyellow pruinosity, (2) wing vein R1 with setae, (3) abdominal tergite 5 reddish or yellow, and (4) genital segments (syntergosternite 7+8 and epandrium) reddish. Phallic morphology most similar to S. (M.) guyanensis Lopes and S. (M.) lindae Lopes, but separable from S. (M.) lindae by the median stylus of similar length to the lateral styli and its vesica shorter than width of distiphallus; and separable from S. (M.) guyanensis by the shape of juxta slightly folded dorsoapically, not rounded (see Lopes 1946: 125, fig. 15). Males (Fig. 4A, C, E, G, H): Body length: 7–8 mm. Wing length: 5.5–6 mm (n = 6). Head (Fig. 4E): Parafacial, fronto-orbital plate, and postocular orbits with silvery gray pruinosity; frontal vitta blackish; face with deep golden-yellow pruinosity. Facial ridge setose on lower half; frontoorbital plate and parafacial with row of setulae close to eye, parafacial with additional row of setulae in lower half. Frons 0.15–0.18 head width at level of ocellar triangle. Antenna black; first flagellomere approximately twice pedicel length; arista short plumose on proximal half. Frontal setae 10–12, well-developed, row reaching level of apex of pedicel; rows of frontal setae parallel for most of their length, diverging at the level of antennal insertion. Two reclinate orbital setae; proclinate orbital setae absent; ocellar setae developed and proclinate; outer vertical seta undifferentiated from postocular setae. Gena with silver pruinosity, covered with black setulae; genal groove blackish; postgena with silver pruinosity, covered with black setulae. Palpus, prementum, and labella blackish. Thorax (Fig. 4A, C): Black with silverygray pruinosity. Acrostichal setae 2+1; dorsocentral setae 2+3; intra-alar setae 1+2; supra-alar setae 2+3; postpronotal setae 3–4; notopleural setae 4. Postalar wall setulose. Postalar callus with 2 setae. Proepisternum bare. Katepisternal setae 3. Scutellum with a pair of basal and subapical setae; apical setae present; discal setae absent. Wing: Tegula black; basicosta yellowish; veins brown. Costal spine not developed; third costal sector setulose ventrally; vein R1 with setae dorsally; vein R4+5 with setulae dorsally from base to crossvein r-m. Cell r4+5 open at wing margin. Legs: Black, except for brown tarsi. Mid femur without ctenidium; mid tibia with 1 median anterior seta; posterior femur with anterodorsal row, and anteroventral row with stronger setae on median surface; posterior tibia with three dorsal setae, one anterior row, and three ventral setae. Tarsal claws long, subequal to length of tarsomere 5. Abdomen (Fig. 4A, C): Syntergite 1+2 to tergite 4 black with silvery-gray pruinosity; tergite 5 reddish. page 5 of 14Zoological Studies 63: 7 (2024) © 2024 Academia Sinica, Taiwan Tergites 2 and 3 without median marginal setae; tergite 4 with complete row of marginal setae; tergite 5 with row of marginal setae. Sternite 5 slightly reddish and V-shaped, with fringe of setae along the median margin of each arm. Terminalia (Fig. 4G, H): Syntergosternite 7+8 and epandrium reddish. Syntergosternite 7+8 with three pairs of setae. Cercus apically curved and pointed. Fig. 3. Holotype male of Sarcophaga lorosa Hall (NHMUK) and holotype male of Arachnidomyia travassosi Tibana and Mello (CNC), respectively. A, B: Lateral habitus and labels; C, D: Dorsal habitus; E, F: Head in frontal view. page 6 of 14Zoological Studies 63: 7 (2024) © 2024 Academia Sinica, Taiwan Fig. 4. Sarcophaga (Mehria) lorosa Hall specimens reared from egg sacs of Metepeira galatheae (Thorell). A, C, E, G, H: Lateral habitus, dorsal habitus, head in frontal view, terminalia in posterior view, and terminalia in lateral view of male, respectively; B, D, F: Lateral habitus, dorsal habitus, and head in frontal view of female, respectively. page 7 of 14 Zoological Studies 63: 7 (2024) © 2024 Academia Sinica, Taiwan Pregonite with rounded and well-sclerotized apex, postgonite with short setulae. Phallus with vesica bifid, short; lateral stylus well-sclerotized; juxta rounded apically, slightly folded dorsoapically. Female (Fig. 4B, D, F): Differs from male as follows: Body length: 8 mm. Wing length: 6.5 mm (n = 1). Frons 0.27 head width at level of ocellar triangle. Frontal setae 8; reclinate setae 2, with posterior seta lateroclinate; proclinate orbital setae 2; inner vertical setae parallel; outer vertical seta about 2/3 inner vertical seta. Tarsal claws shorter than tarsomere 5. Distribution: Argentina (Neuquén) and Chile (Arica y Parinacota, Atacama, O’Higgins, Región Metropolitana de Santiago) (Fig. 2A). Remarks: We propose W. ruficauda and A. travassosi as junior synonyms of S. lorosa Hall. Both nominal species share the above enumerated external characters of color of face pruinosity, wing vein R1, abdominal tergite 5, and genitalia with S. lorosa. They also share a short, bifid vesica, and juxta slightly folded dorsoapically, which are here considered to provide a conspecific match. Pape (1996) transferred W. ruficauda to Sarcophaga and proposed A. travassosi as its synonym, keeping S. (M.) travassosi (Tibana and Mello) as the valid name because of the secondary homonymy between S. (M.) ruficauda (Lopes and Tibana) and Sarcophaga ruficauda Zetterstedt. Unfortunately, the holotype male of W. ruficauda was lost in the fire that consumed most of the MNRJ entomological collection (Cunha 2018; Escobar 2018). Further clarification is required regarding the type locality of A. travassosi. Pape (1996) recorded the city of La Leonera in the Biobío region as the type locality. However, the holotype label explicitly states the city La Leonera in the O’Higgins region (Fig. 3B), which is more than 400 km north of La Leonera in the Biobío region. The exact locality can be verified by consulting the works of the Chilean entomologist Luis Enrique Peña, the collector of the holotype. Peña (1974) listed the type locality of several insects collected by him and included two localities in O’Higgins: La Leonera and Cerro Poqui. Cerro Poqui is a hill located at the border of the regions O’Higgins and Región Metropolitana de Santiago, which shows that Peña used O’Higgins as a reference to the O’Higgins region. Thus, there is no reason to consider that Peña was not accurate when recording La Leonera in the O’Higgins region. The golden-yellow pruinosity on the face of S. (M.) lorosa is not commonly seen in Sarcophagidae and is not present in Nearctic and Palaearctic species of Mehria. In the Neotropical region, only S. (M.) insularis Lopes has the face and frontal vitta slightly goldenyellow (Lopes 1946), but lacks setae on vein R1. Based on the specimens examined here, the distribution of S. (M.) lorosa seems to be restricted to southwestern South America, specifically on the western side of the Andes Mountains (Fig. 2A). Annotated catalog of records of spider eggpredating Sarcophagidae Four species of the subgenus Baranovisca have been recorded as egg predators from at least four species of Araneidae; and 10 species of the subgenus Mehria have been recorded from at least 13 species of the families Araneidae, Cheiracanthiidae, Clubionidae, Philodromidae, Salticidae, and Tetragnathidae (Table 1). Records are from all biogeographical regions except the Afrotropical. Australasian Region Sarcophaga (Baranovisca) arachnivora Lopes From egg sacs of Ordgarius magnificus (Rainbow). Recorded by Lopes (1985: 51, as B. arachnivora) as a parasite in egg sacs of Dicrostichus magnificus Rainbow [= O. magnificus (Rainbow)]) in Hornsby Heights, New South Wales, Australia. Sarcophaga (Baranovisca) cyrtophorae Cantrell From egg sacs of Cyrtophora moluccensis (Doleschall). Recorded by Lubin (1974: 329, as an unidentified Sarcophagidae [identified later by Yefremova and Lubin (2020)]) as preying on eggs of C. moluccensis (Doleschall) at Wau Ecology Institute, Wau, Morobe, Papua New Guinea; by Cantrell (1980: 42, as Parasarcophaga reposita Lopes [misidentification, see Cantrell (1986)]) as preying on eggs of C. moluccensis in Brisbane, Queensland, Australia; by Cantrell (1981: 29, as P. reposita Lopes [misidentification, see Cantrell (1986)]) as preying on eggs of C. moluccensis in Queensland, Australia, and Papua New Guinea [locality not specified]; by Cantrell (1986: 3, as Parasarcophaga cyrtophorae) as preying on eggs of C. moluccensis in Brisbane, Queensland, Australia; and by Yefremova and Lubin (2020: 3) as preying on eggs of C. moluccensis at Wau Ecology Institute, Wau, Morobe, Papua New Guinea. Sarcophaga (Baranovisca) reposita Lopes From egg sacs of an unidentified spider. Recorded by Lopes (1959: 65, as Parasarcophaga (Rosellea) reposita) as preying on eggs of an page 8 of 14Zoological Studies 63: 7 (2024) © 2024 Academia Sinica, Taiwan unidentified spider in Sidney, New South Wales, Australia. Nearctic Region Sarcophaga (Mehria) davidsonii Coquillett From egg sacs of Argiope argentata (Fabricius) and Phidippus octopunctatus (Peckham and Peckham). Recorded by Coquillett (1892: 24) as preying on eggs of Phidippus opifex (McCook) [= Attus opifex McCook = Attus octopunctatus Peckham and Peckham = Phidippus octopunctatus (Peckham and Peckham)] in Los Angeles, Los Angeles County, California, United States of America; and by Davidson (1894: 269) as preying on eggs of A. argentata (Fabricius) in Santa Catalina Island and Redondo Beach, Los Angeles County, California, United States of America. Sarcophaga (Mehria) hinei Aldrich From egg sacs of Larinioides cornutus (Clerck), L. sclopetarius (Clerck), and Philodromus cespitum (Walckenaer). Recorded by Auten (1925: 244) as preying on eggs of Aranea frondosa Comstock [as Walckenaer, error] [= Araneus cornutus Clerck = L. cornutus (Clerck)], Epeira sclopetaria (Clerck) [= Araneus sclopetarius = L. sclopetarius], and Philodromus canadensis Emerton [= Aranea cespitum Walckenaer = P. cespitum (Walckenaer)]) in Put-In-Bay region, Ottawa County, Ohio, United States of America. Remarks: Gillung and Borkent (2017) listed these records for S. (M.) sexpunctata (Fabricius) as a senior synonym of S. (M.) hinei, following the classification in Systema Dipterorum (Evenhuis and Pape 2022). However, this synonymy has never been published and both nominal species are considered as valid in Pape (1996). They also included P. aureolus (Clerck) as prey of S. (M.) hinei as a record made by Auten (1925), but there is no record of this spider species in the original article. Sarcophaga (Mehria) subaenescens Aldrich From egg sacs of an unidentified spider. Table 1. List of spider egg-predating Sarcophagidae and their respective prey according to biogeographical regions. Details about records, localities, and respective references can be found in the catalog in the main text Sarcophagidae species Araneae species Australasian Region Sarcophaga (Baranovisca) arachnivora Araneidae: Ordgarius magnificus Sarcophaga (Baranovisca) cyrtophorae Araneidae: Cyrtophora moluccensis Sarcophaga (Baranovisca) reposita Unidentified spider Nearctic Region Sarcophaga (Mehria) davidsonii Araneidae: Argiope argentata Salticidae: Phidippus octopunctatus Sarcophaga (Mehria) hinei Araneidae: Larinioides cornutus Araneidae: Larinioides sclopetarius Philodromidae: Philodromus cespitum Sarcophaga (Mehria) subaenescens Unidentified spider Neotropical Region Sarcophaga (Mehria) guyanensis Araneidae: Argiope sp. Sarcophaga (Mehria) insularis Unidentified spider Sarcophaga (Mehria) lindae Araneidae: Metepeira atascadero Araneidae: Metepeira incrassata Sarcophaga (Mehria) lorosa Araneidae: Metepeira galatheae Sarcophaga (Mehria) silbergliedi Araneidae: Argiope argentata Tetragnathidae: Tetragnatha sp. Sarcophaga (Mehria) sp. Araneidae: Argiope trifasciata Oriental Region Sarcophaga (Baranovisca) banksi Araneidae: Argiope catenulata Araneidae: Argiope pulchella Palaearctic Region Sarcophaga (Mehria) sexpunctata Araneidae: Larinioides cornutus Clubionidae: Clubiona sp. Cheiracanthiidae: Cheiracanthium punctorium page 9 of 14Zoological Studies 63: 7 (2024)