Paraphotina insolita ( Rehn, 1941 )

Agudelo, Antonio A. & Rafael, José A., 2016, Review of Paraphotina (Mantodea: Photinaidae), Zoologia (e 20160055) 33 (6), pp. 1-13 : 7-13

Paraphotina insolita ( Rehn, 1941)

Figs. 6 , 10 , 14 , 18, 22, 31

Guaraunia insolitum Rehn, 1941: 2 . Type material: male holotype deposited in ANSP.

Paraphotina venezuelana Beier, 1963: 7-8 , syn. nov.; Terra, 1995:

77. Type material: male holotype (type 5632), female allotype deposited in ZSM.

Paraphotina insolita : Cerdá, 1992, partim; Terra, 1995: 77, partim;

Lombardo, 1998: 214, partim; Agudelo & Chica, 2001, 2003:

135, partim; Ehrmann, 2002; Agudelo, 2004: 49, 58, partim;

Otte & Spearman, 2005: 288, partim; Agudelo et al., 2007: 129, partim; Ehrmann & Koçak, 2009: 9, partim.

Type Locality. Venezuela (Monagas, Quiriquire ) .

Diagnosis. Length 38-40 mm. Vertex convex and slightly undulated. Juxta-ocular tubercles slightly developed ( Fig. 10 ).

Ocellar tubercle dark gray. Pseudophallus long and projected ventro-dextrally ( Fig. 28), rarely with apex curved dorsally (Figs.

18, 27). Right membranous lobe of hypophallus with moderately long setae (Fig. 22, rml).

Material examined. BRAZIL, Amazonas: Barcelos, Rio Ara-

çá-Curudurí, 00°05’50”N, 63°17’22”W, 1 male, 15-19.vi.2010,

arm., luminosa, Andreazze R, Rafael JA, Takiya D, Agudelo AA leg.

(INPA); Roraima: Rio Branco , “ Amazonas ”, 1 male, 30.viii.1924 ,

Bequaert leg., determined as Guaraunia insolitum by JWH Rehn in 1954 ( ANSP); Alto Alegre ESEC Maracá, 03°21’59”N, 61°26’04”W, 1 male 12.xii.2015 , Boldrini R, Rafael J, Xavier F leg. ( INPA) ;

COLOMBIA, Meta: San Martin, Vereda La Pascualera, 200 msnm, Figure 21-24. External genitalia, hypophallus of species of Parapho- 1 male, 17.i.2000, t. luz, Agudelo AA, Chica LM ( UDFJC); San tina (ventral view): (21) P. caatingaensis from Rio Grande do Norte, Martín, Reserva El Caduceo, 1 male, 05.ii.2011, Agudelo AA Brazil; (22) P. insolita from Meta, Colombia; (23) P. occidentalis from leg. ( INPA). Paraphotina venezuelana : VENEZUELA, Maracay: 1 male Coari, Amazonas, Brazil; (24) P. reticulata from Maranhão, Brazil). holotype, 1 female allotype; P. Volg leg., Beier det. ( ZSM). (lp) Lateral process; (rml) right membranous lobe. Scale bar: 1 mm. Distribution. Brazil (Amazonas, Roraima), Colombia

(Meta), Venezuela (Aragua, Monagas) .

Remarks. The synonymy established by CERDá (1992) holotype of P. venezuelana ( Figs. 28, 29) was found to be very between Guaraunia insolitum and P. reticulata is rejected here similar to that of P. insolita (Figs. 18, 22, see also LOMBARDO 1998: because CERDá (1992) only examined material from Venezuela figs. 4, 7, 8). In addition, the typical localities of P. venezuelana without examining the type specimen of C. reticulata , including and P. insolita are found in the northern Amazon of Venezuela its phallic complex. This erroneous synonymy was followed by ( Fig. 35 ); Therefore, the synonymy is established.

LOMBARDO (1998), who revised the holotype of G. insolitum but considered it a representative of C. reticulata . The review of the Paraphotina occidentalis Lombardo, 1998 phallic complex of the type specimen of C. reticulata (Figs. 20, Figs. 7 , 11 , 15 , 19, 23, 32

24) and comparison with LOMBARDO’ s (1998) figures of the phallic complex of the Guaraunia insolitum holotype showed that these Paraphotina occidentalis Lombardo, 1998: 213 ; Agudelo & Chica, 2001: 34; Ehrmann, 2002: 268; Otte & Spearman, 2005: 287; species are not synonymous. The specimens of P. reticulata present pseudophallus project dextro-dorsally, while in P. insolita it Agudelo et al., 2007: 129; Ehrmann & Koçak, 2009: 9. Type is projected to dextro-ventrally. Aside from this, specimens of P. material: male holotype, male paratype deposited in ANSP. insolita lack lateral process on the hypophallus, while P. reticu- Diagnosis. Length 45-50 mm. Robust appearance for the lata possesses a well-developed and strongly sclerotized lateral genus ( Fig. 7 ). Ocellar tubercle dark brown. Arabesques on proprocess, projecting to the right (dorsal view). This aspect was notum, forefemora and foretibiae well-marked. Pseudophallus corroborated by P. insolita material from northern Brazil, Amazo- long and projected to the right (Fig. 19, psph). Hypophallus nas and Roraima, and the Colombian and Venezuelan “Llanos”, with numerous and minute setae on the lateral membranous also when compared with material of P. reticulata from the east lobe (Fig. 23, rml).

of the Amazon. The morphology of the phallic complex of the Type locality. Peru (Huallaga, Aguaytia) .

7/13 9/13 Paraphotina reticulata Saussure, 1871 a: 196 ; Giglio-Tos, 1915: 29) revealed that they are not conspecific, thus the synonymy

72; 1927: 330; Beier, 1930: 28; 1935: 124; Cerdá, 1992: 13, between these species is rejected here.

partim; Terra, 1995: 77; Ehrmann, 2002: 268, partim; Otte &

Spearman, 2005: 287; Agudelo et. al., 2007: 129; Ehrmann DISCUSSION

& Koçak, 2009: 9.

Diagnosis. Length 36-45 mm. Ocellar tubercle dark. Within a morphological context, the definition of most Pseudophallus long and projected to the right (Fig. 20, psph). Photinaidae genera has been generally well supported by easily Hypophallus with a strongly sclerotized lateral process, right recognizable diagnostic features. Except in cases like Metriomanmembranous lobe with moderately long setae (Fig. 24, rml). tis, Photina and Photinella , the genera of this family appear to

Type Locality. Brazil, Pará. be acceptably defined, but their phylogenetic relationships are

Material examined. Cardioptera reticulata : BRAZIL, Pará, not necessarily as clear. After the review of Orthoderella by ROY & holotype male ( reticulata Saussure “Natal”) ( MNHG). Additional STIEWE (2011), the distinction between Paraphotina and Orthodematerial. BRAZIL. Amazonas: Barcelos, Rio Araçá , comunidade Ba- rella has become more evident. The behavior of Paraphotina cuquara, 00°09’17”N, 63°10’35”W, 2 males, 06.vi.2010, suspensa species is reminiscent of the observations described by BRUNNER dossel 25 m; Alvim E, Rafael JA, Agudelo AA leg. ( INPA); Novo & GANDOLFO (1990) for females of Orthoderella ornata (per. obs.). Airão, M.E. Baixo rio Jaú, 01°56’28”S, 61°26’45”W, 1 male, 22- However, Orthoderella is distinguished from Paraphotina by a 23.vii.1995, luz mista, luz negra, lençol, Motta C, Xavier F leg. long head with a nearly straight vertex, and by well-developed ( INPA- 0027036); Maranhão: Caxias, Reserva Ecológica Inhamum, juxta-ocular tubercles. The pronotum in Orthoderella is slender, 2 males, 01-03.xii.2005, Limeira F et. al., leg. ( CZMA); same date its arabesque brands less conspicuous, and the lateral margins but Reserva Ecológica Inhamum, Povoado Coités, 1 male, 22- of the prozona are nearly parallel. Wings in female Orthoderella 24.x.2006, Souza SO, Silva JJ leg. ( INPA); same data but 1 male, are noticeably shorter than the abdomen, exposing its distal 19-21.i.2007, Souza SO, Silva JJ leg.( INPA); same data but 1 male, half when at rest. In contrast, females of Paraphotina generally 08-11.viii.2008, Limeria F, leg. ( INPA); same municipality but expose only the distal third of the abdomen. The genera Ortho- Morro do Alecrim, Campus da UEMA), 1 male, 26-30.iii.2009, r. derella and Paraphotina are distinguished from other Photinaidae entomológica, Barbosa EA, Neto MB leg. ( INPA); Carolina, povoa- genera by the arabesque marks on their pronotum, forefemora do Canto Grande, Balneario Urupuxete, 3 males, 23-27.ii.2009, and foretibiae.

armadilha luminosa, Limeira F et al., leg. ( CZMA). same data but The four species of Paraphotina are superficially very simi- 2 males, 23-27.ii.2009, armadilha luminosa. Limeira F et al., leg. lar, which renders their identification somewhat difficult to ( INPA); same data but Fazenda Cincorá, 3 males, 17-22.x.2009, naked eye. Paraphotina insolita and P. reticulata , in particular, are armadilha luminosa, Limeira F et al., leg. ( CZMA); same data difficult to distinguish morphologically. The fast identification but 1 male, 17-22.x.2009, armadilha luminosa, Limeira F et of P. caatingaensis comb. nov. and P. occidentalis , is generally al., leg. ( INPA); Bom Jardim, REBIO, Res. Biol. Gurupí, 2 males, less problematic, because their sizes are at opposite ends of the 15-19.iv.2010, Limeira F et al. leg. ( INPA); Parnarama, Povoado Paraphotina range. Thus, it becomes necessary to resort to a more Brejo de São Felix, 1 male, 03-06.vi.2010, Souza SS, Lino TR leg. detailed observation, including the dissection of the genitalia. ( INPA); Pará: 1 male, holotype ( reticulata “Natal”) MHS ( MHNG); While the phylogenetic relationships within Photinaidae Santo Antonio do Tauá, 2 males, 5-15.x.1979, Michel Boulard are consecutively being analyzed, we are just beginning to leg., genital preparation n° 3488 ( MNHN); Belém, Campus de understand the life history and systematics of the concerned Pesquisa do Museu Paraense Emílio Goeldi, 1 female, 10.iii.2014, genera. Some Photinaidae genera such as Metriomantis and Photi- Favacho C. leg. ( MPEG); Piaui: Parque Nacional de Sete Cidades, na are in urgent need of revision. Ecological and behavioral data 04.09909ºS, 41.70952ºW, 2 males, 15-25.iv.2012, Somavilla A, for Paraphotina as well as for the large majority of Photinaidae Oliveira leg. ( INPA). genera are rare. Behavioral data were studied in Orthoderella by

Distribution. Brazil (Amazonas, Maranhão, Pará, Piaui). BRUNNER & GANDOLFO (1990) and in Cardioptera brachyptera by

Remarks. This species had been described with doubtful TERRA (1992, 1996a, b). Taxonomic approaches are the basis for provenance. The holotype ( Fig. 1 ) of C. reticulate bears a label subsequent evolutionary analysis.

giving the typus locality as Natal ( South Africa). However, SAU- Biogeographical aspects. The distribution of Neotropical SSURE (1871a) added a note to the description commenting that species of Mantodea seem to show some patterns similar to the specimen may have come from Pará ( Brazil). This comment those of several others groups studied methodically in the bioappeared again in his succeeding work ( SAUSSURE 1871b). We geographical context, e.g., Diptera (AMORIM & PIRES 1996), fitting confirmed the State of Pará as type locality due to the conspe- within subregions and areas of endemism proposed by NIHEI & cific specimens examined are principally from Maranhão and CARVALHO (2007). These relationships of the continental northern Pará. CERDá (1992) established P. venezuelana as synonym of P. Neotropical region and its biogeographical components were reticulata . The comparison between phallic complexes of the briefly described by RAFAEL & CUMMING (2009), while the biogeoholotypes of P. reticulata and P. venezuelana (Figs. 20, 24, 28, graphical patterns were discussed by AMORIM (2009). In recent reviews of Neotropical Mantodea ( LOMBARDO & IPPOLITO 2004, ROY rica y algunos aspectos bioecológicos ( Insecta: Mantodea ). & EHRMANN 2009, ROY & STIEWE 2011), the distribution maps of Revista Colombiana de Entomologia 29: 127-136.

species largely coincided with these classifications ( Fig. 35 ). All AGUDELO AA, LOMBARDO F, JANTSCH LJ (2007) Checklist of the Neospecies of Paraphotina occur in South America. Three species, P. tropical mantids ( Insecta, Dictyoptera, Mantodea ). Biota insolita , P. occidentalis , and P. reticulata , occur in Northwestern Colombiana 8: 105-158.

Amazonia. Paraphotina insolita is found in North Amazonia (N AGUDELO AA, RIVERA J (2015) Some taxonomic and nomenclatural Am) in Orinoco savannas ( Colombia, Venezuela) and Brazil changes in American Mantodea ( Insecta, Dictyoptera) – Part (Roraima and Amazonas, Barcelos). Paraphotina occidentalis oc- I. Zootaxa 3936: 335-356. doi: 10.11646/zootaxa.3936.3.2 curs mainly in Southwestern Amazonia (SW Am) with records AMORIM DS (2009) Neotropical Diptera Diversity: richness, patin the municipalities of Bujari (Acre), Novo Airão, Tefé, Auta- terns, and perspectives, p. 71-97. In: PAPE T, BICKEL D, MEIER zes, and Beruri (Amazonas), but also with peripheral records in R (Eds.) Diptera Diversity: Status, Challenges and Tools. North Amazonia (Presidente Figueiredo). Paraphotina reticulata Leiden, Brill , 459p.

has been more widely recorded from North Amazonia and AMORIM DS, PIRES MR (1996) Neotropical biogeography and a Southweastern Amazonia extending until the Atlantic Forest , in method for maximum biodiversity estimation, p. 183-219. Maranhão. Paraphotina caatingaensis has been recorded only in In : BICUDO CE, MENEZES NA (Eds.) Biodiversity in Brazil: A the Brazilian domains of Cerrado and Caatinga, from Maranhão first approach. São Paulo, CNPq, vi+ 326p.

to Minas Gerais. No records from Southeast Amazonia (SE Am) BEIER M (1930) Ergebnisse einer zoologischen Sammelreise nach are known to date, but according to the pattern recorded for Brasilien, insbesondere in das Amazonasgebiet, ausgeführt the known species, it is possible that an unknown species of von Dr. H. Zerny. Annalen des Naturhistorischen Museums Paraphotina inhabits this zone. Many questions remain before in Wien 44: 27-32.

we will understand the relationships among Photinaidae genera, BEIER M (1935) Mantodea . Subfamilie: Mantinae . In: WYTSMAN P especially biogeographical aspects and their closer relationship (Ed.) Genera Insectorum. Bruxelles, Fasc. 203, 146p.

with Neotropical Acanthopidae and Liturgusidae more than BEIER M (1942) Neue und seltene Mantodeen aus deutschen Mantidae groups. Museen. Annalen des Naturhistorischen Museums in Wien 52: 126-154.

ACKNOWLEDGMENTS BEIER M (1963) Neue und bemerkenswerte Mantiden verschiedener Herkunft. Stuttgarter Beiträge zur Naturkunde, Serie This research was financed with resources from the A, 106: 1-11.

Conselho Nacional de Desenvolvimento Técnico e Científico BEIER M (1964) Blattopteroidea-Mantodea, p. 850-970. In: BRONN (CNPq) through the doctoral scholarship to AAA, (process HG (Ed.) Bronn’s Klassen und Ordnungen des Tierreichs, 156567/2010-5), and Fundação de Amparo à Pesquisa do Esta- Insecta-Arthropoda, Part III. Leipzig, Akademische Verlagsdo do Amazonas FAPEAM/PAPAC (286/2013). We are indebted gesellschaft Geest & Portig K.-G.

to Peter Schwendinger ( MHNG), Dieter Doczkal ( ZSM), Klaus BEIER M (1968) Mantodea (Fangheuschrecken) , p. 1-47. In: Schönitzer ( ZSM) and their respective institutions for granting HELMCKE JG, STARCK D, WERMUTH H (Eds.) Handbuch der Zooaccess to type specimens and the entomological collections. logie. Berlin, Walter de Gruyter & Co, vol. 4 .

To Francisco Limeira ( CZMA), Orlando Tobias ( MPEG) and BEIER M (1970) Dictyoptera (Blattoidea et Mantoidea), p. 38-41. Ricardo Andreasse ( UFRN) for providing access to material of In: TUXEN SL (Ed.) Taxonomist’s glossary of genitalia in their respective institutions. We express our gratitude to Kai insects. Copenhagen, Munksgaard, 359p.

Schütte ( ZMH) and César Favacho ( MPEG) who kindly provided BRUNNER D, GANDOLFO D (1990) Morphological adaptations to an important photos and information for this work. unusual defensive strategy in the mantid Orthoderella ornata ( Insecta: Mantidae ). Journal of Zoology 222: 129-136.

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dea), p. 43-60. In: FERNáNDEZ CF, ANDRADE MG, AMAT G (Eds.) EHRMANN R (2002) Mantodea-Gottesanbeterinnen der Welt. Insectos de Colombia. Bogotá, Universidad Nacional de Münster , Natur und Tier Verlag, 519p.

Colombia, Facultad de Ciencias, vol. 3, 604p. EHRMANN R, KOÇAK AÖ (2009) The Neotropical Mantids ( Insecta:Dic- AGUDELO AA, CHICA LM (2001) Presencia del género Paraphotina tyoptera: Mantodea ) (Ehrmann 30.v.2009). Cesa News 49: 1-18. Giglio-Tos, 1915 ( Mantodea : Vatidae: Photininae ) en Co- FRANÇOIS A, ROY R (2015) Le genre Microphotina Beier, 1935 : deux lombia. Boletín Científico del Museo de Historia Natural espèces, ou une seule? ( Mantodea , Photinaidae ). Bulletin de Universidad de Caldas 5: 33-37. la Société entomologique de France 120: 389-396.

AGUDELO AA, CHICA LM (2003) Mántidos de la Orinoquía Colom- GIGLIO-TOS E (1915) Mantidi esotici. Generi e specie nuove. biana: contribución al conocimiento de su diversidad gené- Bullettino della Società Entomologica Italiana 46: 31-108.

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HEBARD M (1928) The group Luzarae of the subfamily Phalangop- ROY R (2002) Révision du genre Néotropical Macromantis Saussinae (Orthoptera, Gryllidae). Transactions of the American sure, 1871 (Dictyoptera, Mantidae). Bulletin de la Société Entomological Society 54: 1 -56. Entomologique de France 107: 403-418.

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Phasmidae). London, British Museum, Natural History, 501p. ROY R, EHRMANN R (2009) Révision du genre Zoolea Audinet-Ser- LOMBARDO F (1998) A new species of Paraphotina Giglio-Tos, ville (Mantodea, Mantidae, Vatinae). Revue Française d’En- 1915 from Peru (Insecta: Mantodea). Journal of Orthoptera tomologie 31: 1-22.

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Serville, 1831 (Mantodea, Mantidae, Acanthopinae) with SAUSSURE H DE (1871 b) Synopsis des Mantides Américains, p. comments on their Phylogeny. Annals of the Entomological 1-186. In: Memories pour servir a L’Histoire Naturelle du Society of America 97: 1076-1102. Mexique, des Antilles et États-Unis 4ª. Genève, George MENEZES E, BRAVO F (2013) Uma nova espécie de Orthoderella H, 186p.

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of Polietina Schnabl & Dziedzicki (Diptera: Muscidae): Neo- SHEVTSOVA E, HANSSON C, JANZEN DH, KJAERANDSEN J (2011) Stable tropical areas relationships and Amazonia as composite area. structural color patterns displayed on transparent insect Systematic Entomology 32: 477 -501. doi: 10.1111/j.1365- wings. Proceedings of the National Academy of Sciences 3113.2006.00376.x 108: 668-673. doi: 10.1073/pnas.1017393108

OTTE D, SPEARMAN LA (2005) Mantida species file, catalog of the SVENSON GJ, BRANHAM MA (2007) Case 3402: Photinini LeConte, mantids of the world. Philadelphia, Association of Insect 1881 (Insecta: Coleoptera) and Photininae Giglio-Tos, 1915 Diversity, 489p. (Insecta: Mantodea): proposed resolution of homonymy RAFAEL J, CUMMING J (2009) Revision of the genus Macrostomus between family-group names. Bulletin of Zoological No- Wiedemann (Diptera: Empididae: Empidinae). I. The ferru- menclature 64: 1-9.

gineus species-group. Zootaxa 2064: 39-56. SVENSON G J, WHITING MF (2009) Reconstructing the origins of REHN JWH (1941) A new genus of Mellierid Mantid from Vene- praying mantises (Dictyoptera, Mantodea):the roles of Gondzuela Orthoptera -Mantidae. Notulae Naturae, Academy wanan vicariance and morphological convergence. Cladistics Natural Science of Philadephia 70: 1-4. 25: 468-514. doi: 10.1111/j.1096-0031.2009.00263.x

RIVERA J (2010) A historical review of praying mantid taxonomy TERRA PS (1992) Zelo materno em Cardioptera brachyptera (Manto- and systematics in the Neotropical Region: State of know- dea, Vatidae, Photininae). Revista Brasileira de Entomologia ledge and recent advances (Insecta: Mantodea). Zootaxa 36: 493-503.

2638: 44-64. doi: 10.11646/%25x TERRA PS (1995) Revisão sistemática dos gêneros de Louva-A-Deus RIVERA J, SVENSON G (2016) The Neotropical ‘polymorphic ear- da região Neotropical ( Mantodea ). Revista Brasileira de less praying mantises’ – Part I: molecular phylogeny and Entomologia 39: 13-94.

revised higher-level systematics ( Insecta: Mantodea , Ac- TERRA PS (1996a) Comportamento sexual de Cardioptera brachypanthopoidea). Systematic Entomology 41: 607-649. doi: tera ( Mantodea ). Revista Brasileira de Entomologia 40: 3- 7. 10.1111 /syen.12178 TERRA PS (1996b) Zelo materno em Photina amplipennis Stål ( Mantodea , Vatidae). Revista Brasileira de Entomologia 40: 9-10.

Submitted: 22 March, 2016 WESTWOOD JO (1889) Revisio Insectorum Familiae Mantida-

Received in revised form: 20 May 2016

rum, speciebus novis aut minus cognitis descriptis et

Accepted: 14 June 2016

delineatis. London, Gurney & Jackson, 55p.

Editorial responsibility: Ângelo Parise Pinto WIELAND F (2013) The phylogenetic systems of Mantodea ( Insecta:

Dictyoptera). Species, Phylogeny and Evolution 3: 3-222. YAGER DD, SVENSON GJ (2008) Patterns of praying mantis audi-

tory system evolution based on morphological, molecular, Author Contributions: AAA conducted the taxonomic review;

neurophysiological and behavioural data. Biological Jour- AAA and JAR analyzed the data and wrote the paper.

nal of the Linnean Society 94: 541-568. doi: 10.1111/j. Competing Interests: The authors have declared that no

1096-0031.2009.00263.x competing interests exist.

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Figures

Figure 35. Map of the geographic records for species of Paraphotina with areas of endemism proposed by AMORIM (2009). Note that geographical records of P. reticulata include transition areas for each of the other three species. (AF) Atlantic Forest; (N Am) North Amazon; (SE Am) South East Amazon; (SW Am) South West Amazon.

Figures 5-8. Males of Paraphotina (dorsal habitus): (5) P. caatingaensis from Bahia, Brazil; (6) P. insolita from Meta, Colombia; (7) P. occidentalis from Coari, Amazonas, Brazil; (8) P. reticulata from Maranhão, Brazil. Scale bar: 10 mm.

Figures 9-12. Head of males of Paraphotina (frontal view): (9) P. caatingaensis from Bahia, Brazil; (10) P. insolita from Meta, Colombia; (11) P. occidentalis from Coari, Amazonas, Brazil; (12) P. reticulata from Maranhão, Brazil. Scale bar: 1 mm.

Figures 13-16. Pronota of males of Paraphotina (dorsal view): (13) P. caatingaensis from Bahia, Brazil; (14) P. insolita from Meta, Colombia; (15) P. occidentalis from Coari, Amazonas, Brazil; (16) P. reticulata from Maranhão, Brazil. Scale bar: 1 mm.

Figures 1-4. Types of Paraphotina (dorsal habitus): (1) Paraphotina reticulata (originally Cardioptera reticulata Saussure, 1871) holotype male deposited in MHNG; (2-3) Paraphotina venezuelana, (2) holotype male, (3) allotype deposited in ZSM; (4) Paraphotina caatingaensis comb. nov. (originally Orthoderella caatingaensis Menezes & Bravo, 2013) holotype male deposited in MZFS. Arrow highlights the basal spot on fore coxa. Scale bar: 10 mm. Photo of P. caatingaensis courtesy of E. Menezes (MZFS).