Zootaxa 4802 (2): 374–382 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2020 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4802.2.10 http://zoobank.org/urn:lsid:zoobank.org:pub:3B340088-6420-40CE-8740-9E9239B8B206

The spongillafly genus Burmeister, 1839 (: ) from Brazil: Distributional, taxonomical and bionomical notes

ALICE CARVALHO ASSMAR1* & ADOLFO RICARDO CALOR1,2 1PPG Biodiversidade e Evolução, Laboratório de Entomologia Aquática—LEAq, Instituto de Biologia, Universidade Federal da Ba- hia, Rua Barão de Jeremoabo, 147, campus Ondina, Ondina, CEP 40170-115 Salvador, Bahia, Brazil. 2 �[email protected]; https://orcid.org/0000-0003-3975-3176 *Corresponding author. �[email protected]; https://orcid.org/0000-0002-3007-3679

Abstract

Sisyridae is a small family of aquatic , with 73 extant species described worldwide. In Brazil, 17 species of two genera, Climacia McLachlan, 1869 and Sisyra Burmeister, 1839, have been reported. In this paper, the family is recorded from the Caatinga biome (semiarid region), Northeastern Brazil, for the first time, including a record of the genus Sisyra from Ceará State. Additional distributional data are presented for both S. apicalis Banks, 1908 and S. panama Parfin & Gurney, 1956. Sisyra ariasi Penny, 1981 is here synonymized with S. panama, NEW SYNONYMY. Illustrations of the male genitalia of S. minuta Esben-Petersen, 1935, and bionomic notes for S. panama are provided.

Key words: aquatic insects, checklist, Neotropical region, new synonymy

Resumo

Sisyridae é uma pequena família de insetos aquáticos, com 73 espécies existentes descritas no mundo. No Brasil, 17 espécies de dois gêneros, Climacia McLachlan, 1869 e Sisyra Burmeister, 1839, são registradas. Neste artigo, a família é registrada para o bioma Caatinga (região semiárida), nordeste do Brasil, pela primeira vez, incluindo um registro do gênero Sisyra para o Ceará. Dados distributivos adicionais são apresentados para S. apicalis Banks, 1908 e S. panama Parfin & Gurney, 1956. Sisyra ariasi Penny, 1981 é aqui sinonimizada com S. panama, NOVA SINONÍMIA. Ilustrações do macho de S. minuta Esben-Petersen, 1935, e comentários sobre o comportamento de S. panama, são providenciados.

Palavras-chave: insetos aquáticos, lista de espécies, nova sinonímia, região Neotropical

Introduction

Sisyridae (spongillaflies) is an aquatic family of Neuroptera (Insecta: Neuropterida) occurring worldwide. These neuropterans are commonly known by the association of their aquatic larval stages with freshwater sponges (Po- rifera: Spongillidae), which are their main feeding source (Brown 1952; Parfin & Gurney 1956; Penny & Rafael 1982). The larvae of Sisyridae have their mandibles and maxillae modified into sucking tubes, which they insert into the hosts and drain cellular content (Brown 1952; Parfin & Gurney 1956). The diet of the adults is based on honey- dew, pollen and small invertebrates, such as mites and aphids (Pupedis 1987). The courtship is through vibratiory emissions sent through the substrate or wings movements (Brown 1952; Rupprecht 1995; Henry 2005). The imma- tures stages of the species of Sisyridae are poorly known. Hamada et al. (2014) recently provided bionomic notes on the larvae of Sisyra panama Parfin & Gurney, 1956, which, along with Banks, 1908 (Bowles 2006), are the only Neotropical species with known larvae. Currently, the family has 73 valid extant species in four genera, Climacia McLachlan, 1869, Sisyrina Banks, 1939, Sisyborina Monserrat, 1981 and Sisyra Burmeister, 1839 (Forteath et al. 2015; Oswald & Machado 2018; Oswald 2019). The latter is the most species rich genus of Sisyridae, with 48 valid species (Forteath et al. 2015;

374 Accepted by D. Bowles: 28 May 2020; published: 23 Jun. 2020 Oswald 2019). The genus Sisyra has a cosmopolitan distribution, excluding Antarctica, and it is most represented in the Palearctic region, with 16 species recorded (Oswald 2019). In the Neotropics, six species of Sisyra have been recorded, all of them have been reported from Brazil (Bowles 2015; Assmar & Salles 2017). Despite recent papers on adults sisyrids (Monserrat 2005; Flint 2006; Hamada et al. 2014; Bowles 2015; Ardila- Camacho & Martins 2017; Assmar & Salles 2017), the distribution of Neotropical species of Sisyra is still poorly known (Hamada et al. 2014; Bowles 2015). Among the Brazilian biomes, Sisyra previously has been recorded from the Atlantic Rainforest (S. apicalis, S. elongata Penny & Rafael, 1982 and S. panama), and the Amazon Forest (all six species), where two species occur only there, S. ariasi Penny, 1981 and S. minuta Esben-Petersen, 1935 (Assmar & Salles 2017; Machado & Martins 2019). Only one specimen of S. ariasi was ever collected, the male holotype (Penny 1981). Since its description, no other collection records of S. ariasi have been reported. The illustration of the male genitalia provided by Penny (1981) lacks important details and the description of the species is similar to that for S. panama. Recently, fieldwork conducted in the Northeast Region of Brazil resulted in the collection of several specimens of Sisyra. Due to scarce knowledge of Neotropical Sisyra, the present study aims to provide new distributional data for S. apicalis and S. panama, including the first record of the family from the Caatinga biome (semiarid region). Furthermore, S. ariasi is proposed as a junior SYNONYMY of S. panama, after comparison of the holotypes, and to present bionomical notes of S. panama adults. Finally, the male genitalia of S. minuta are here illustrated in order to complement previous illustrations.

Material and methods

Specimens analyzed are from the Museu de História Natural da Bahia (UFBA), Universidade Federal da Bahia, Salvador, Bahia, Brazil. In addition, specimens were collected during field trips in Bahia state, using white and UV lights, between 2014 and 2019, in the cities of Salvador, Jandaira and Palmeiras. The altitude in the area of the col- lections ranges from 0 to 870 m. Wings were removed, dehydrated and preserved on a slide in order to analyze venation and color pattern. The genitalia were cleared using a 10% KOH solution (Blahnik & Holzenthal 2004), heated for approximately one hour, dehydrated with 80% and 100% ethanol series, and stored in micro vials with glycerin. The specimens are preserved in 80% ethanol, except the adults observed alive, which are pinned. While alive, the specimens were kept in plastic cages and set free in closed rooms for observation. All specimens are deposited at UFBA. Photographs were taken using software Nikon NIS-Elements, attached to a Photomicroscope Nikon SMZ1500 and edited at Adobe Photoshop CC 2018. The illustrations of the dissected parts were drawn using a camera lucida attached to an optical microscope Leica DM500. Digital illustrations were vectorized in the software Adobe Illus- trator CC 2017. The photograph of the wings of Sisyra panama holotype (Fig. 15) presented a shadow underneath the wings, what resulted in a “lighter streak” in the posterior margin of the wings. Therefore, the photographs were edited in the Adobe Photoshop CC 2017 in order to smooth the shadow effects and avoid misinterpretation. Maps were drawn using the free software QGIS Desktop 3.10 (Figs 1–7). Terminology follows Aspöck & Aspöck (2008) and Breitkreuz et al. (2017) for genitalia and wings, respec- tively. The numbers attributed to the RP branches are merely to aid in identification.

Sisyra apicalis Banks, 1908 (Figs 1, 3, 5–7)

Sisyra apicalis Banks, 1908:261 [description]; Parfin & Gurney 1956:471 [revision]; Penny 1981:163 [record]; Maes & Flint 1994:4 [record]; Oswald et al. 2002:581 [checklist]; Monserrat 2005:10 [record]; Bowles 2006:13 [record, larval informa- tion]; Flint 2006:280 [record]; Bowles 2015:3 [record]; Assmar & Salles 2017:86 [record]. Sisyra nocturna Navás 1932:155 [description]; 1935:67 [revision]; Monserrat 2005:10 [junior synonym of S. apicalis]

Remarks. This species is widely distributed, being recorded from the southern United States to Brazil (Oswald 2019). In Brazil, this is the first record of S. apicalis from Bahia and Pernambuco states, in the Northeast Region of the country (Figs 1, 3, 5–7). This also constitutes the first record of Sisyridae to Pernambuco state.

SISYRA Burmeister (Neuroptera: Sisyridae) from Brazil Zootaxa 4802 (2) © 2020 Magnolia Press · 375 The remarkable characteristic of S. apicalis is the straight distally projected male gonocoxite 9, with rounded apex and curving internally in dorsal view (Assmar & Salles 2017). Material examined. BRAZIL: Bahia, Jandaira, Reserva Copern, 11°36’51.9” S 37°38’46.9” W, 22 m a.s.l., 11.x.2016, light pan trap (UV and white light), Kiszewiski; Silva, F.; Dias, E. & Campos, R., 1 male (UFBA, N0035); Salvador, Parque São Bartolomeu, 12°53’45.6” S 38°28’11.31” W, 22.x.2019, light pan trap, Miranda, M. & Pereira, R., 1 male (UFBA, N0052). Minas Gerais: Aimorés, AE UNE Elieser Batista [Aimorés hydroelectric power plant, Eliezer Batista], Pedra Lorena, 19°26’44” S 53°06’21” W, 15.i.2011, CDU trap, França, D., 1 fe- male (UFBA, N0039). Pernambuco: Tamandaré, REBIO Saltinho, riacho Saltinho, 8°43’47.5” S 35°10’31.4” W, 20.iv.2018, white light pan trap, Pereira-Silva, R., Lima, R.P. & Gomes, D.W.S., 1 male (UFBA, N0034); same data, except 19.iv.2018, 3 m.a.s.l, 2 males (UFBA, N0036, N0042); Cabo Santo Agostinho, Refúgio de Vida Silvestre Gurjaú, Sede, waterfall, 8°21’30” S 34°56’30” W, 01–02.x.2019, white and UV light pan trap, Pereira-Silva, R., 1 male (UFBA, N0081).

Sisyra minuta Esben-Petersen, 1935 (Figs 1–2, 8–11)

Sisyra minuta Esben-Petersen, 1935:152 [description]; Parfin & Gurney 1956:478 [revision]; Penny 1981:164 [record]; Flint 2006:281 [record].

Remarks. Esben-Petersen (1935) described this species based on a male specimen from “Taderinha”, Brazil, which according to Parfin & Gurney (1956) is a misspelling to “Taperinha” (Amazonas, Brazil). Later, Penny (1981) and Flint (2006) recorded S. minuta from Manaus city, Amazonas state, Brazil, with specimens from Adolpho Ducke Reserve (Fig. 2). Esben-Petersen (1935) described the species as small, with yellowish brown forewing and hyaline hind wing (Figs 8–9), without providing a description of the male genitalia. Parfin & Gurney (1956) and Penny (1981) illus- trated and described the male genitalia of the holotype; however, important structures were not highlighted. There- fore, additional illustrations of this species are here presented in order to complement previous illustrations of the male genitalia (Figs 10–11). The remarkable characteristic of S. minuta is the short male gonocoxite 9, with the apex sharply pointed dorso- ventrally; “beak-like” as stated by Penny (1981). The specimen analyzed in this paper has differences in the male genitalia from what is described and illustrated in the literature. These differences could be attributed to the angle of the genitalia during the illustration or drawing styles. The primary differences are in the gonostyle 11, which has al- most same size than the apex, with a lateral “finger-like” joint to the gonocoxite 9 (Fig. 11). In addition, the sharply pointed apex of gonocoxite 9 has a second projection on the inner region, which bears a distal seta (Fig. 10). Parfin & Gurney (1956) illustrated the male genitalia of S. minuta showing a small gonostyle 11, without lateral joint, and the sharply pointed apex of the gonocoxite 9, without an inner projection. Penny (1981) did not identify the inner small projection of the gonocoxite 9, but highlighted three sharply pointed structures, which could be interpreted as the gonostyle 11, the gonocoxite 9 apex and the lateral joint of the gonostyle 11 with gonocoxite 9, which could appear as a sharp projection depending on the view. Material examined. BRAZIL: Amazonas, Manaus, Reserva Adolpho Ducke, 3°5’0” S 60°0’0” W, 14– 18.xi.2009, Malaise trap, Mendonça, M.M., 1 male (UFBA, N0040).

Sisyra panama Parfin & Gurney, 1956 (Figs 1, 3–4, 6, 12–20)

Sisyra panama Parfin & Gurney, 1956:474 [description]; Penny 1981:166 [record]; Flint 2006:281 [record]; Hamada et al. 2014:281 [bionomic notes and record]; Bowles 2015:4 [record]; Assmar & Salles 2017:87 [record]. Sisyra ariasi Penny 1981:168 [description]. NEW SYNONYMY.

Remarks. Both holotypes of S. ariasi (Figs 12–14) and S. panama (Figs 15–16) were examined and no significant differences were found between the two in the lateral view. Other views are not possible to analyze because the male holotype of S. ariasi is fixed on slide and the genitalia is not cleared (Fig. 12).

376 · Zootaxa 4802 (2) © 2020 Magnolia Press Assmar & CALOR Penny (1981) affirmed that S. panama and S. ariasi have two distinctive crossveins between “R3 and R4+5” (RP2 and RP3) (Fig. 13). However, the additional crossvein found between these veins in the right forewing of S. ariasi holotype is a modification exclusive to this wing, and this additional crossvein is not present in the left forewing of the same specimen (Fig. 12). A male specimen of S. panama, collected in Salvador, Bahia, presented this same modification in the left forewing and in the hind wing (Figs 17–18). Besides, S. panama has only one crossvein between RP2 and RP3, according to Parfin & Gurney (1956). Regarding the male genitalia, both illustrations and descriptions of these two species are not detailed. Parfin & Gurney (1956) described the male gonocoxite 9 of S. panama as “stout”, “with long setae and distal inner teeth” and “caliper-like”. In addition, the authors described the basal portion of the “paramere” (gonostyle 11) as triangular. The illustration shows the postero-dorsal margin elongated in relation to the ventral one (Parfin & Gurney 1956). In contrast, according to Penny (1981) S. panama male gonocoxite 9 is “short, quadrangular, with several small teeth at the apex”, lacking the “ventral thumb-like lobe”, present in S. ariasi. The last cited characteristic would be the only difference between the two species in the male genitalia (Penny 1981). Comparing the male genitalia, there are no significant differences between S. ariasi and S. panama (Figs 12, 19–20). Sisyra panama has a ventral “thumb-like lobe”, which is the gonostyle 11, called “parameres” by Parfin & Gurney (1956). In addition, the apex of the male gonocoxite 9 of both species are truncate, although Penny (1981) illustrated the gonocoxite 9 of S. ariasi with the apex rounded compared to slightly rhomboidal in S. panama (Figs 12, 20). It is possible that the apparent differences are due to the angle at which the illustrations were made. As can be seen in Penny (1981), the genitalia are drawn in an oblique position relative to S. panama. Due to the absence of distinctive characters between the two species, Sisyra ariasi Penny, 1981, NEW SYNONYMY, is now attributed to S. panama Parfin & Gurney, 1956. Description and illustration of the male genitalia of S. panama are provided here (Figs 19–20). This is the first record of Sisyra panama and the family Sisyridae from Caatinga biome in Brazil. The specimens of S. panama were collected in two highland areas, Serra de Ibiapaba, Ubajara (Ceará state) and Caeté-Açú, Palmei- ras (Bahia state), where orographic rainfall is common, thus enabling the existence of wet forest islands surrounded by Caatinga vegetation (Queiroz et al. 2017). Serra de Ibiapaba is characterized as a “Brejo de Altitude”, and it is floristically similar to the Amazonia biome (Takiya et al. 2016; Queiroz et al. 2017). Caeté-Açú is a district of Palmeiras city, located in the Chapada Diamantina Complex, which has phytophysiognomies similar to the Atlantic rainforest and it contains the only Rupestrian Grassland vegetation of the Caatinga (Queiroz et al. 2017). Male genitalia (Figs 19–20). Ectoproct and sternite 9 transversal, with numerous long dentigerous-based setae and simple setae. Ectoproct larger dorsally, rounded margins. Sternite 9 long, quadrangular, posterior margin longer than anterior laterally. Gonocoxite 9 stout, quadrangular, truncate apex with inner distal teeth dorsally. Numer- ous long, large dentigerous-based setae on gonocoxite 9 posterior margin, mostly postero-ventrally, simple setae throughout. Gonostyle 11 triangular in lateral view, resembling a ventral “thumb”, apex acute, curving to midline. Bionomic notes. Two adults were collected during different periods and observed alive, in order to analyze their behavior. A male and a female were observed, the first was kept alive for three days, and the last for approxi- mately 14 days, being fed with a mix of water and honey, placed on cotton. Both individuals landed on the cotton to feed on the solution. It was observed that the specimens mostly walked, instead of flying in the enclosure. While walking, they oc- casionally stopped, vibrated their maxillary palps, made a “back and forth” movement with the body, touched their abdomen on the surface or got really close to it, and then starts walking again (behavior also reported to S. nigra (Retzius, 1783) by Rupprecht (1995)). This vibration and abdominal movements have been reported for some Neu- ropteran families, including Sisyridae, as a conspecific communication and courtship behavior (Henry 2005). The specimen emits the vibration that travel through the substrate and is received by the other nearby individuals through the subgenual organs in the legs (Devetak & Amon 1997; Devetak 1998). In Sisyra, the use of the mandibles and the abdomen to produce vibration have been reported by Rupprecht (1995; 1997) and Henry (2005), respectively. Here, it was observed the maxillary palp was the body part that vibrated. Related to the flight behavior, both specimens started flight from a vertical landing position by dropping itself from the surface and then flying. When it was in a horizontal position, a spiral flight was performed, until it reached a certain height. In both cases, their flight was without apparent pattern.

SISYRA Burmeister (Neuroptera: Sisyridae) from Brazil Zootaxa 4802 (2) © 2020 Magnolia Press · 377 Figures 1–7. Maps of the new distributional records of Sisyra Burmeister, 1839 species (1) in Brazil. Emphasis in the (2) Amazonas state; (3) Northeast region of Brazil, with the first records to states of (4) Ceará and (5) Pernambuco, and additional records to (6) Bahia state; and (7) Minas Gerais state. The half-green half-red hexagon means superposition of species records.

378 · Zootaxa 4802 (2) © 2020 Magnolia Press Assmar & CALOR Material examined. BRAZIL: Bahia, Palmeiras, Caeté-Açú, Vale do Capão, rio do Batista, 30.vi–03.vii.2018, Pennsylvania white light trap, Calor, Assmar, Gomes, Miranda & Tavares, 1 female (UFBA, N0033); Salvador, 19BC, córrego Cascão, 12°58’16” S 38°30’39” W, 19.viii.2014, white sheet, Gomes, V.; Ferreira, E; Alves; Lima, 1 female (UFBA, N0038); same data, except Alphaville I, Residencial Itaparica, L. Tavares’s house, 12°56’39” S 38°23’50.7” W, 01.viii.2019, Tavares, L., 1 male (UFBA, N0044); same data, except 02.x.2019, Tavares, L. 1 female (UFBA, N0048). Ceará, Ubajara, Serra do Ibiapaba, Parque Nacional de Ubajara, Murimbeca stream, 03°49’18.9” S 40°54’16.6” W, 868 m, 28.x.2011, white light pan trap, Gomes, V & Duarte, T., 1 male (UFBA, N0041).

Figures 8–11. Sisyra minuta Esben-Petersen, 1935 (8) forewing; (9) hind wing; and male genitalia (10) dorsal view and (11) lateral view. Abbreviations: C= costal; Sc= subcostal; RA= radial anterior; RP= radial posterior; MA= medial anterior; MP= me- dial posterior; CuA= cubital anterior; CuP= cubital posterior; A= anal; Ect= ectoproct; gx 9= gonocoxite 9; gx 11= gonocoxite 11; gs 11= gonostyle 11; st 9= sternite 9.

Conclusion

This is the first record of Sisyridae from the Caatinga Biome of Brazil, thus expanding the distributional range of the family in the country. The areas where some specimens of Sisyra panama were collected, Caeté-Açú, Palmeiras (Bahia state) and Serra do Ibiapaba, Ubajara (Ceará state), possess unique climate and vegetational dynamics, with phytophysiognomies of the Cerrado (tropical savanna ecoregion) and Atlantic Rainforest, although it is embedded within the Caatinga Domain (Takiya et al. 2016; Queiroz et al. 2017).

SISYRA Burmeister (Neuroptera: Sisyridae) from Brazil Zootaxa 4802 (2) © 2020 Magnolia Press · 379 Figures 12–20. Specimens of Sisyra Burmeister, 1839. Sisyra ariasi Penny, 1981 male holotype (12) mounted on slide, (13) forewing and (14) hind wing with label, both mounted on slide with broken base. Sisyra panama Parfin & Gurney, 1956 (15) holotype forewing and (16) hind wing with label; (17) forewing of Sisyra panama Parfin & Gurney, 1956 with additional crossvein and (18) hind wing of the same specimen, collected in Salvador city, Bahia state, Brazil. Male genitalia of S. panama in (19) dorsal view and (20) lateral view.

380 · Zootaxa 4802 (2) © 2020 Magnolia Press Assmar & CALOR The number of Sisyra species recorded from Brazil is now five. In the Northeast of Brazil, the number of spe- cies is increased from one to three species, including S. apicalis (Bahia and Pernambuco states), S. elongata (Bahia state) and S. panama (Bahia and Ceará states). It is the first record of the family from Ceará state (S. panama) and Pernambuco state (S. apicalis), along with new records from Bahia state. The Sisyridae fauna of Brazil as well as all of the Neotropics is still poorly known and we reinforce the need for additional research in order to understand if the gaps in the family distribution are natural or due to lack of study. Identification of adult sisyrids through analysis of their wings is important, but using male genitalia is crucial in order to avoid misidentification.

Acknowledgment

We are grateful to the researchers and curators Dr. Torsten Dikow and Dr. Oliver Flint Jr. (in memoriam), for all their support before and during the visit to Smithsonian Institution, Washington, DC, USA. In addition, thanks to the Smithsonian Institution for the grants awarded in their Fellowship Program. We thank Dr. Marcio Luiz de Oliveira (INPA), for the loan of specimens examined in this study and for support during the visit to the institution. In addi- tion, we thank Dr. Caleb Martins (USP) for his help in the acquisition of photographs of some species we analyzed, as well as in the careful review of this manuscript. Thanks to Foundation for Research Support of the State of Bahia (FAPESB, Fundação de Amparo à Pesquisa do Estado da Bahia, grant N° BOL0316/2017) and Improvement Coor- dination of Higher Education Personnel (CAPES, Coordenação de Aperfeiçoamento de Pessoal de Nível Superior, Postgraduate Support Program, PROAP) for all financial support. Finally, we thank to Msc. Leon Tavares, who col- lected the living specimens observed in this study and Msc. Rafael Pereira for samples collected.

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