4802 (2): 374–382 issn 1175-5326 (print edition) https

httpsdoiorg1011646zootaxa4802210httpzoobankorgurnlsidzoobankorgpub3B340088-6420-40CE-8740-9E9239B8B206

374 Accepted by D Bowles 28 May 2020 published 23 Jun 2020

Article ZOOTAXA ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)

Zootaxa 4802 (2) 374ndash382httpswwwmapresscomjzt

Copyright copy 2020 Magnolia Press

The spongillafly genus Sisyra Burmeister 1839 (Neuroptera Sisyridae) from Brazil Distributional taxonomical and bionomical notes

ALICE CARVALHO ASSMAR1 amp ADOLFO RICARDO CALOR12

1PPG Biodiversidade e Evoluccedilatildeo Laboratoacuterio de Entomologia AquaacuteticamdashLEAq Instituto de Biologia Universidade Federal da Ba-hia Rua Baratildeo de Jeremoabo 147 campus Ondina Ondina CEP 40170-115 Salvador Bahia Brazil2acalorgmailcom httpsorcidorg0000-0003-3975-3176Corresponding author aliceassmar92gmailcom httpsorcidorg0000-0002-3007-3679

Abstract

Sisyridae is a small family of aquatic insects 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 Cearaacute State Additional distributional data are presented for both S apicalis Banks 1908 and S panama Parfin amp 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 eacute uma pequena famiacutelia de insetos aquaacuteticos com 73 espeacutecies existentes descritas no mundo No Brasil 17 espeacutecies de dois gecircneros Climacia McLachlan 1869 e Sisyra Burmeister 1839 satildeo registradas Neste artigo a famiacutelia eacute registrada para o bioma Caatinga (regiatildeo semiaacuterida) nordeste do Brasil pela primeira vez incluindo um registro do gecircnero Sisyra para o Cearaacute Dados distributivos adicionais satildeo apresentados para S apicalis Banks 1908 e S panama Parfin amp Gurney 1956 Sisyra ariasi Penny 1981 eacute aqui sinonimizada com S panama NOVA SINONIacuteMIA Ilustraccedilotildees do macho de S minuta Esben-Petersen 1935 e comentaacuterios sobre o comportamento de S panama satildeo providenciados

Palavras-chave insetos aquaacuteticos lista de espeacutecies nova sinoniacutemia regiatildeo 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 amp Gurney 1956 Penny amp 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 amp 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 amp Gurney 1956 which along with Sisyra apicalis 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 amp Machado 2018 Oswald 2019) The latter is the most species rich genus of Sisyridae with 48 valid species (Forteath et al 2015

SISYRA BuRMEISTER (NEuROPTERA SISYRIDAE) FROM BRAzIL Zootaxa 4802 (2) copy 2020 Magnolia Press middot 375

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 amp Salles 2017)

Despite recent papers on adults sisyrids (Monserrat 2005 Flint 2006 Hamada et al 2014 Bowles 2015 Ardila-Camacho amp Martins 2017 Assmar amp 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 amp 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 amp Salles 2017 Machado amp 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 Histoacuteria 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 amp 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 ldquolighter streakrdquo 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 310 (Figs 1ndash7)

Terminology follows Aspoumlck amp Aspoumlck (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 5ndash7)

Sisyra apicalis Banks 1908261 [description] Parfin amp Gurney 1956471 [revision] Penny 1981163 [record] Maes amp Flint 19944 [record] Oswald et al 2002581 [checklist] Monserrat 200510 [record] Bowles 200613 [record larval informa-tion] Flint 2006280 [record] Bowles 20153 [record] Assmar amp Salles 201786 [record]

Sisyra nocturna Navaacutes 1932155 [description] 193567 [revision] Monserrat 200510 [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 5ndash7) This also constitutes the first record of Sisyridae to Pernambuco state

ASSMAR amp CALOR376 middot Zootaxa 4802 (2) copy 2020 Magnolia Press

The remarkable characteristic of S apicalis is the straight distally projected male gonocoxite 9 with rounded apex and curving internally in dorsal view (Assmar amp Salles 2017)

Material examined BRAzIL Bahia Jandaira Reserva Copern 11deg36rsquo519rdquo S 37deg38rsquo469rdquo W 22 m asl 11x2016 light pan trap (uV and white light) Kiszewiski Silva F Dias E amp Campos R 1 male (uFBA N0035) Salvador Parque Satildeo Bartolomeu 12deg53rsquo456rdquo S 38deg28rsquo1131rdquo W 22x2019 light pan trap Miranda M amp Pereira R 1 male (uFBA N0052) Minas Gerais Aimoreacutes AE uNE Elieser Batista [Aimoreacutes hydroelectric power plant Eliezer Batista] Pedra Lorena 19deg26rsquo44rdquo S 53deg06rsquo21rdquo W 15i2011 CDu trap Franccedila D 1 fe-male (uFBA N0039) Pernambuco Tamandareacute REBIO Saltinho riacho Saltinho 8deg43rsquo475rdquo S 35deg10rsquo314rdquo W 20iv2018 white light pan trap Pereira-Silva R Lima RP amp Gomes DWS 1 male (uFBA N0034) same data except 19iv2018 3 masl 2 males (uFBA N0036 N0042) Cabo Santo Agostinho Refuacutegio de Vida Silvestre Gurjauacute Sede waterfall 8deg21rsquo30rdquo S 34deg56rsquo30rdquo W 01ndash02x2019 white and uV light pan trap Pereira-Silva R 1 male (uFBA N0081)

Sisyra minuta Esben-Petersen 1935(Figs 1ndash2 8ndash11)

Sisyra minuta Esben-Petersen 1935152 [description] Parfin amp Gurney 1956478 [revision] Penny 1981164 [record] Flint 2006281 [record]

Remarks Esben-Petersen (1935) described this species based on a male specimen from ldquoTaderinhardquo Brazil which according to Parfin amp Gurney (1956) is a misspelling to ldquoTaperinhardquo (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 8ndash9) without providing a description of the male genitalia Parfin amp 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 10ndash11)

The remarkable characteristic of S minuta is the short male gonocoxite 9 with the apex sharply pointed dorso-ventrally ldquobeak-likerdquo 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 ldquofinger-likerdquo 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 amp 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 3deg5rsquo0rdquo S 60deg0rsquo0rdquo W 14ndash18xi2009 Malaise trap Mendonccedila MM 1 male (uFBA N0040)

Sisyra panama Parfin amp Gurney 1956(Figs 1 3ndash4 6 12ndash20)

Sisyra panama Parfin amp Gurney 1956474 [description] Penny 1981166 [record] Flint 2006281 [record] Hamada et al 2014281 [bionomic notes and record] Bowles 20154 [record] Assmar amp Salles 201787 [record]

Sisyra ariasi Penny 1981168 [description] NEW SYNONYMY

Remarks Both holotypes of S ariasi (Figs 12ndash14) and S panama (Figs 15ndash16) 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)


Penny (1981) affirmed that S panama and S ariasi have two distinctive crossveins between ldquoR3 and R4+5rdquo (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 17ndash18) Besides S panama has only one crossvein between RP2 and RP3 according to Parfin amp Gurney (1956)

Regarding the male genitalia both illustrations and descriptions of these two species are not detailed Parfin amp Gurney (1956) described the male gonocoxite 9 of S panama as ldquostoutrdquo ldquowith long setae and distal inner teethrdquo and ldquocaliper-likerdquo In addition the authors described the basal portion of the ldquoparamererdquo (gonostyle 11) as triangular The illustration shows the postero-dorsal margin elongated in relation to the ventral one (Parfin amp Gurney 1956) In contrast according to Penny (1981) S panama male gonocoxite 9 is ldquoshort quadrangular with several small teeth at the apexrdquo lacking the ldquoventral thumb-like loberdquo 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 19ndash20) Sisyra panama has a ventral ldquothumb-like loberdquo which is the gonostyle 11 called ldquoparameresrdquo by Parfin amp 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 amp Gurney 1956 Description and illustration of the male genitalia of S panama are provided here (Figs 19ndash20)

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 (Cearaacute state) and Caeteacute-Accediluacute 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 ldquoBrejo de Altituderdquo and it is floristically similar to the Amazonia biome (Takiya et al 2016 Queiroz et al 2017) Caeteacute-Accediluacute 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 19ndash20) 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 ldquothumbrdquo 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 ldquoback and forthrdquo 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 amp 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


FIGuRES 1ndash7 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) Cearaacute 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


Material examined BRAzIL Bahia Palmeiras Caeteacute-Accediluacute Vale do Capatildeo rio do Batista 30vindash03vii2018 Pennsylvania white light trap Calor Assmar Gomes Miranda amp Tavares 1 female (uFBA N0033) Salvador 19BC coacuterrego Cascatildeo 12deg58rsquo16rdquo S 38deg30rsquo39rdquo W 19viii2014 white sheet Gomes V Ferreira E Alves Lima 1 female (uFBA N0038) same data except Alphaville I Residencial Itaparica L Tavaresrsquos house 12deg56rsquo39rdquo S 38deg23rsquo507rdquo W 01viii2019 Tavares L 1 male (uFBA N0044) same data except 02x2019 Tavares L 1 female (uFBA N0048) Cearaacute ubajara Serra do Ibiapaba Parque Nacional de ubajara Murimbeca stream 03deg49rsquo189rdquo S 40deg54rsquo166rdquo W 868 m 28x2011 white light pan trap Gomes V amp Duarte T 1 male (uFBA N0041)

FIGuRES 8ndash11 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 Caeteacute-Accediluacute Palmeiras (Bahia state) and Serra do Ibiapaba ubajara (Cearaacute 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)


FIGuRES 12ndash20 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 amp Gurney 1956 (15) holotype forewing and (16) hind wing with label (17) forewing of Sisyra panama Parfin amp 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


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 Cearaacute states) It is the first record of the family from Cearaacute 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 Fundaccedilatildeo de Amparo agrave Pesquisa do Estado da Bahia grant Ndeg BOL03162017) and Improvement Coor-dination of Higher Education Personnel (CAPES Coordenaccedilatildeo de Aperfeiccediloamento de Pessoal de Niacutevel 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

References

Aspoumlck u amp Aspoumlck H (2008) Phylogenetic relevance of the genital sclerites of Neuropterida (Insecta Holometabola) Sys-tematic Entomology 33 97ndash127

httpsdoiorg101111j1365-3113200700396xArdila-Camacho A amp Martins CC (2017) First record of spongillaflies (Neuroptera Sisyridae) from Colombia Zootaxa

4276 (1) 129ndash133 httpsdoiorg1011646zootaxa427618Assmar AC amp Salles FF (2017) Taxonomic and distributional notes on Spongilla-flies (Neuroptera Sisyridae) from South-

eastern Brazil with first interactive key to the species of the country Zootaxa 4273 80ndash92 httpsdoiorg1011646zootaxa427316Banks N (1908) Neuropteroid insects notes and descriptions Transactions of the American Entomological Society 34 255ndash

267Banks N (1939) New genera and species of neuropteroid insects Bulletin of the Museum of Comparative Zoology 85 439ndash

504Blahnik RJ amp Holzenthal RW (2004) Collection and curation of Trichoptera with an emphasis on pinned material Necto-

psyche Neotropical Trichoptera Newsletter 1 8ndash20Bowles DE (2006) Spongillaflies (Neuroptera Sisyridae) of North America with a key to the larvae and adults Zootaxa 1357

1ndash19Bowles DE (2015) New distributional records for Neotropical spongillaflies (Neuroptera Sisyridae) Insecta Mundi 0400

1ndash7Breitkreuz LC Winterton SL amp Engel MS (2017) Wing tracheation in Chrysopidae and other Neuropterida (Insecta) a

resolution of the confusion about vein fusion American Museum Novitates 2017 (3890) 1ndash45 httpsdoiorg10120638901Brown HP (1952) Life history of Climacia areolaris (Hagen) a neuropterous lsquoparasitersquo of fresh water sponges American

Midland Naturalist 47 130ndash160 httpsdoiorg1023072421701Burmeister HCC (1839) Handbuch der Entomologie zweiter Band Berlin Theodor Christian Friedrich Enslin1839 757ndash

1050Devetak D (1998) Detection of substrate vibration in Neuropteroidea a review Acta Zoologica Fennica 209 87ndash94Devetak D amp Amon T (1997) Substrate vibration sensitivity of the leg Scolopidial organs in the green lacewing Chrysoperla

carnea Journal of Insect Physiology 43 433ndash437 httpsdoiorg101016S0022-1910(96)00121-7


Esben-Petersen P (1935) Two new species of Neuroptera Konowia 14 151ndash153Flint OS Jr (2006) New species and records of Neotropical Sisyridae with special reference to Sisyra (Insecta Neuroptera)

Proceedings of the Biological Society of Washington 119 279ndash286 httpsdoiorg1029880006-324X(2006)119[279NSARON]20CO2Forteath GNR Purser J amp Osborn AW (2015) A new species of Sisyra Burmeister 1839 (Neuroptera Sisyridae) from Four

Springs Lake and Wadleyrsquos Dam Northern Tasmania Austral Entomology 54 217ndash220 httpsdoiorg101111aen12118Hamada N Pes AMO amp Fusari LM (2014) First record of Sisyridae (Neuroptera) in Rio de Janeiro state Brazil with

bionomic notes on Sisyra panama Florida Entomologist 97 281ndash284 httpsdoiorg1016530240970140Henry CS (2005) Acoustic communication in Neuropterid insects In Drosopoulos S amp Claridge MF (Eds) Insect Sounds

and Communication Physiology Behaviour Ecology and Evolution CRC press and Taylor and Francis Group Boca Raton pp 153ndash166

httpsdoiorg1012019781420039337ch10Machado RJP amp Martins CC (2019) Sisyridae in Cataacutelogo Taxonocircmico da Fauna do Brasil PNuD Available from http

faunajbrjgovbrfaunafaunadobrasil11051 (accessed 4 October 2019)Maes JM amp Flint OS Jr (1994) Dilaridae Berothidae y Sisyridae (Planipennia) tres familias nuevas para la fauna de Nica-

ragua Revista Nicaraguumlense de Entomologia 29 3ndash5McLachlan R (1869) New species of Hemerobiina with synonymic notes (first series) Entomologistrsquos Monthly Magazine 6

21ndash27Monserrat VJ (1981) Sobre los Sisiacuteridos de la Regioacuten Oriental (Neuroptera Planipennia Sisyridae) Revista Espantildeola de

Entomologia 57 165ndash186Monserrat VJ (2005) Nuevos datos sobre algunas pequentildeas familias de neuroacutepteros (Insecta Neuroptera Nevrorthidae Os-

mylidae Sisyridae Dilaridae) Heteropterus Revista de Entomologia 5 1ndash26Navaacutes L (1932) Deacutecadas de insectos nuevos Deacutecada 21 Broteacuteria (Ciecircncias Naturais) 28 109ndash119Navaacutes L (1935) Monografiacutea de la familia de los Sisiacuteridos (Insectos Neuroacutepteros) Memorias de la [Real] Academia de Ciencias

Exactas Fisico-Quimicas y Naturales de Zaragoza 4 1ndash87Oswald JD (2019) Neuropterida species of the World Available from httplacewingtamueduSpeciesCatalogMain (ac-

cessed 4 October 2019)Oswald JD Contreras-Ramos A amp Penny ND (2002) Neuroptera (Neuropterida) In Bousquets J amp Morone JJ (Eds)

Biodiversidad Taxonomiacutea y Biogeografiacutea de Artroacutepodos de Meacutexico hacia una siacutentesis de su conocimiento Volume 3 universidad Nacional Autoacutenoma de Meacutexico Distrito Federal Mexico pp 559ndash581

Oswald JD amp Machado RJP (2018) Biodiversity of the Neuropterida (Insecta Neuroptera Megaloptera and Raphidiop-tera) Insect Biodiversity science and society 2 627ndash672

httpsdoiorg1010029781118945582ch21Parfin SI amp Gurney AB (1956) The spongilla-flies with special reference to those of the western hemisphere (Sisyridae

Neuroptera) Proceedings of the United States National Museum 105 421ndash529 httpsdoiorg105479si00963801105-3360421Penny ND (1981) Neuroptera of the Amazon Basin Part 1 Sisyridae Acta Amazonica 11 157ndash169 httpsdoiorg1015901809-43921981111157Penny ND amp Rafael JA (1982) Two new species of Sisyridae (Neuroptera) from the Amazon basin Neuroptera International

II 2 53ndash58Pupedis RJ (1987) Foraging behavior and food of Adult Spongila-flies (Neuroptera Sisyridae) Annals of the Entomological

Society of America 80 758ndash760 httpsdoiorg101093aesa806758Queiroz LP de Cardoso D Fernandes MF amp Moro MF (2017) Diversity and evolution of flowering plants of the Caatinga

Domain In Silva JMC Leal IR amp Tabarelli M (Eds) Caatinga The Largest Tropical Dry Forest Region in South America Springer New York pp 23ndash63

httpsdoiorg101007978-3-319-68339-3_2Rupprecht R (1995) Anmerkungen zum Paarungsverhalten von Sisyra Galathea 2 15ndash17Rupprecht R (1997) Wie Schwammfliegen Vibrationen erzeugen Galathea 3 11ndash13Takiya DM Santos APM Pinto AcircP Henriques-Oliveira AL Carvalho ADL Sampaio BHL Clarkson B Moreira

FFF Avelino-Capistrano F Gonccedilalves IC Cordeiro IDRS Cacircmara JT Barbosa JF Souza WRM amp Rafael JR (2016) Aquatic Insects from the Caatinga checklists and diversity assessments of ubajara (Cearaacute State) and Sete Ci-dades (Piauiacute State) National Parks Northeastern Brazil Biodiversity Data Journal 4 (e8354) 1ndash195

httpsdoiorg103897BDJ4e8354


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 amp Salles 2017)

Despite recent papers on adults sisyrids (Monserrat 2005 Flint 2006 Hamada et al 2014 Bowles 2015 Ardila-Camacho amp Martins 2017 Assmar amp 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 amp 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 amp Salles 2017 Machado amp 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 Histoacuteria 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 amp 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 ldquolighter streakrdquo 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 310 (Figs 1ndash7)

Terminology follows Aspoumlck amp Aspoumlck (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 5ndash7)

Sisyra apicalis Banks 1908261 [description] Parfin amp Gurney 1956471 [revision] Penny 1981163 [record] Maes amp Flint 19944 [record] Oswald et al 2002581 [checklist] Monserrat 200510 [record] Bowles 200613 [record larval informa-tion] Flint 2006280 [record] Bowles 20153 [record] Assmar amp Salles 201786 [record]

Sisyra nocturna Navaacutes 1932155 [description] 193567 [revision] Monserrat 200510 [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 5ndash7) This also constitutes the first record of Sisyridae to Pernambuco state




























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Acknowledgment


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4802 (2): 374–382 issn 1175-5326 (print edition) https

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