Traumatomutilla André miscellanea: Revision of the bellica, bifurca, diabolica, and vitelligera species groups, and a new group for the new species T. pilkingtoni Bartholomay and Williams (Hymenoptera: Mutillidae: Sphaeropthalminae: Dasymutillini) Author Bartholomay, Pedro R. Author Williams, Kevin A. Author Luz, David R. Author Cambra, Roberto A. Author Oliveira, Márcio Luiz de text Insecta Mundi 2019 2019-06-28 709 709 1 37 journal article 23949 10.5281/zenodo.3674793 98c3ec1f-1711-4f1d-acb6-1702ad46ddfb 1942-1354 3674793 63A67DA8-A6A5-47E4-97F0-FFFE3D66A58A Traumatomutilla bifurca ( Klug, 1821 ) ( Fig. 26–45 ) Mutilla bifurca Klug 1821: 313 . Lectotype female (designated here), Brazil , Pará , Cametá (ZMB) Ephuta ( Ephuta ) bifurca André 1902: 58 (new combination) Mutilla bifurca Bradley 1916: 192 ( incertae sedis ) Traumatomutilla ira Casal 1969: 297 Holotype female, Brazil , Paraíba , Soledade, Juazeirinho (AMNH), examined, syn. nov. Traumatomutilla bifurca Nonveiller 1990: 75 (new combination) Diagnosis. Female. This species is separated from other members of the bifurca species group by lacking genal carina and overall setae pattern with the head completely clothed in silvery-white, dorsum of pronotum and mesonotum completely black and two sublateral oblique longitudinal patches of appressed silvery-white setae on T2. Male. Males can be diagnosed by their conspicuously striated and raised medial plate of pygidium, one pair of tooth-like projections below mid-ocellus, genital cuspis which is conspicuously slender and broadened, dorsoventrally flattened and subacute apically. Additionally, the cuspis is virtually devoid of setae with the exception of a single oblique row of short setae subapically on ventral surface. Description. Female. Body length 8.0–12.0 mm. Head . Posterior margin flat, occipital carina conspicuously swollen apicolaterally. Head width 0.75 × pronotal width. Eye almost circular; its length in frontal view 0.9 × the distance between its lower margin and mandibular condyle. Sculpture of frons and vertex concealed by dense setae; gena sparsely, coarsely and shallowly foveate-punctate; genal carina absent. Mandible unidentate apically, unarmed ventrally and dorsally. Antennal scrobe with dorsal horizontal carina conspicuously projected into a small lamella narrowly before antennal tubercle; lateral scrobal carina absent. Antennal tubercle irregularly and coarsely rugose. Flagellomere 1 2.2 × pedicel length; flagellomere 2 1.5 × pedicel length. Mesosoma . Mesosomal length 0.85 × width; pronotum as wide as mesothorax. Anterior face of propodeum well-defined, densely and finely punctate in dorsal half with longitudinal striae on ventral half. Mesosomal dorsal sculpture mostly concealed by dense setae, coarsely and densely areolate-punctate where observable. Humeral carina well developed, slightly projected apically, terminating just before low and rounded epaulet; anterolateral corners of pronotum angulate in dorsal view. Pronotal spiracles virtually flat against lateral margin of pronotum. Lateral face of pronotum sparsely and superficially punctate. Sculpture of mesopleuron and lateral face of propodeum concealed by dense setae, except at smooth impunctate dorsal fourth of metapleuron. Post spiracular area apparently undefined, concealed by dense setae. Ratios of width of humeral angles, pronotal spiracles, widest point of mesonotum, narrowest point of mesonotum and propodeum posterior to propodeal spiracles, 53:62:60:48:45. Lateral margin of mesothorax slightly constricted anterior to propodeal spiracle. Scutellar scale well-defined, very narrower than anterolateral carinae; anterolateral carinae unequal, disconnected from each other and scutellar scale; scabrous intervals on scutellar area apparently indistinct, concealed by dense setae if present. Posterior face of propodeum longer than dorsal face. Metasoma . Ratios of width of T1, width of T2 and length of T2, 45:97:104. T2 with maximum width posterior to midlength. Sculpture of T1–6, except pygidium, mostly concealed by dense setae, except laterally on T2 sparsely and coarsely foveate-punctate. S1 with longitudinal sharp carina, higher posteriorly; S2–5 densely and coarsely foveate-punctate, more finely and densely so in S3–5; anteromedial crest-fold vestigially present on S2; subapical slope on S2 vestigial, observable only laterally. Pygidium ovate, defined by carinae along its entire length, with well defined longitudinal wavy costae; interstice apparently smooth; apical margin with minute transverse striae. Male. (Hitherto undescribed) Body length 8.0–12.0 mm. Head. posterolateral angles rounded, vertex conspicuously swollen posteromedially; head width 0.9 × pronotal width. Eyes almost rounded, slightly longitudinally ovate; ocelli small; OOD 4.8 × DLO, IOD 1.5 × DLO. Occipital carina distinct, strongly convex. Front, vertex and gena sculpture concealed by setae; front with a pair of tooth-like projections below median ocellus. Gena ecarinate. Antennal scrobe broadly concave, with prominent transverse dorsal carina extending from internal margin of eye to antennal tubercle, 2/3 of antennal tubercle height in lateral view. Clypeus weakly convex, densely setose, setae concealing sculpture, apical margin straight. Scape bicarinate. Flagellomere 1 1.8 × pedicel length; flagellomere 2 1.3 × pedicel length. Mandible obliquely tridentate apically, inner tooth larger than middle tooth; lacking dorsal or ventral projections; maxilla and labium elongate; maxillary palp 6-segmented, third and fourth segments slightly flattened and apically expanded, other segments almost cylindrical. Labial palp 4-segmented, second and third segments slightly flattened and apically expanded, other segments almost cylindrical. Mesosoma. Epaulets evident, slightly prominent, rounded, distad from humeral carina by more than its own length. Anterior face of pronotum densely micropunctate laterally and with a distinct, impunctate, shining and slightly concave area medially; lateral face of pronotum finely and densely punctate; dorsal face of pronotum densely setose, sculpture concealed. Meso and metapleuron sculpture concealed by setae; mesopleuron with acute tubercle on dorsal half; Tegula convex, glabrous except with long appressed setae anterolaterally. Mesonotum and scutellum densely punctured; notaulus absent, parapsis indistinguishable; scutellum slightly convex; axilla produced posterolaterally as truncate tooth, contiguously punctate but smooth posteriorly. Metanotum sculpture concealed by dense appressed setae. Propodeum strongly convex, entirely densely reticulate, except lateral face near metapleuron smooth. Metasoma . T1 sub-petiolate, 0.5 × as wide as T2, sculpture concealed by dense silvery-white setae. S1 with weakly pronounced ridge-like carina. T2 0.75 × as long as wide, observable sculpture with dense coarse punctures, sculpture concealed by dense silvery-white setae elsewhere. S2 sparsely and coarsely punctate, more sparsely so medially; with posteromedial, longitudinal, acutely sub-ovate pit filled with dense silvery-white setae. T3–5 with sculpture concealed by dense silvery-white setae. T6 densely and coarsely punctate. S3–4 sculpture concealed by dense silvery-white setae. S5–7 densely and coarsely punctate, except apical third of S7 impunctate and asetose; apical margin of S7 with weakly acute medial projection. Pygidium with raised striated medial plate surrounded by lateral carinae. Wings. Forewing with elongate sclerotized pterostigma, stigma length 2x its height; marginal cell broadly truncate apically; three submarginal cells, 3r-m ending before M. Legs. Mid and hind tibiae each without strong spines dorsally, distinct apical secretory pore on inner surface near base of inner spur; spurs straight on margins, not serrate. Genitalia . Parapenial lobe weakly digit-like apically. Paramere free length 1.15 × the free length of cuspis and 3.5 × the free length of digitus; virtually straight in dorsal view and weakly and gradually upcurved in lateral view, virtually asetose. Cuspis free length 3 × the free length of digitus. slightly sinuose throughout and slightly expanded apically in dorsal view, virtually straight throughout and sharp apically in lateral view; with a single subapical row of short setae ventrally. Digitus with numerous short setae basally on dorsal surface. Penis valve with strongly concave on internal surface, closely bidentate apically, apical distance between teeth 0.05 × length of valve; with few short setae on apical margin and subapically on external surface. Coloration and variations. Female. Body and appendages black except mandibles partially reddishbrown. Tibial spurs pale-yellow. Head setae predominantly silvery-white with dorsal half of gena and posterior margin of vertex having black setae. Mesosomal setae predominantly silvery-white except dorsum of pronotum, mesonotum, and dorsum of propodeum medially, with black setae. Legs setae predominantly silvery-white except tarsi setae partially black to reddish-black. Metasomal setae predominantly white, except T1 medially, T2 medially and sublaterally, T3 medially, and T4–5 sublaterally with black setae varying in density. No significant color variations were found in the females examined. Male. Body and appendages black. Tibial spurs yellowish-white. Wings hyaline brown, apical third infuscated brown on forewing; veins brown, hindwing completely hyaline. Head setae silvery-white. Mesosomal setae predominantly silvery-white, except mesonotum, tegulae, and scutellum with black setae. Legs setae silvery-white; tibial spurs white. Metasomal setae predominantly silvery-white, except posterior half of T2, T5 medially, T6–7, and S7 with brownish-black to black setae varying in density. No significant color variations were observed for any of the males examined. Material examined. 1733 ♀ , 71♂ . Type material. Lectotype of Traumatomutilla bifurca , ♀ , BRAZIL , Bahia , Sieber & Gomes ( ZMB ). Holotype of Traumatomutilla ira , ♀ , BRAZIL , Paraíba , Soledade, Juazeirinho, vi.1956 , A.G.A. Silva ( AMNH ). Other material. BRAZIL : Pará : Algodoal, Praia da Princesa, 1♀ , 22.viii.1981 , Edmar L. Oliveira ( MPEG ); Cachimbo , 1♀ , 19.viii.1978 , M.J.G. Hopkins & H.C. Hopkins ( MPEG ) ; Conceição do Araguaia , 1♀ , vii.1959 , M. Alvarenga ( MPEG ) ; Mangabeira , Mocajuba , 3♀ , vii.1953 , O. Rego ( AMNH ) ; Maranhão : Barreirinha , Parque Nacional Lençóis Maranhenses , 02°39’80’’S 42°49’88’’W, 1♀ , 07.v.2016 , J.A. Rafael & F.L. Oliveira ( INPA ) ; Barra do Corda , 1♀ , 07.ii.1955 , Expedição do Departamento de Zoologia ( MPEG ) ; Imperatriz , 1♀ , 13.vi.1978 , M.F. Torres ( MPEG ) ; Mirador , Parque Estadual do Mirador, Base da Geraldina , 12♀ , 28.viii–03.ix.2008 , F. Limeira-de-Oliveira ( CZMA ) ; Riachão , Fazenda Altos , 9♀ , 18–22.viii.2009 , F. Limeira-de-Oliveira ( CZMA ) ; Caxias , Reserva Ecológica do Inhamum , 5♀ , 02–11.ix.2005 , M.M.B.G.J. Júnior ( CZMA ) ; Mirador , Parque Estadual do Mirador, Base da Geraldina , 5♀ , 22–26.viii.2006 , F. Limeira-de-Oliveira ( CZMA ) ; Mirador , Parque Estadual do Mirador, Base da Geraldina , 5♀ , 26.viii.2006 , F. Limeira-de-Oliveira ( CZMA ) ; Mirador , Parque Estadual do Mirador , Base dos Cágados , 06°46’37’’S 45°06’34’’W , 3♀ , 26.xi–03.xii.2011 , F. Limeira-de-Oliveira ( CZMA ) ; Caxias , Reserva Ecológica do Inhamum , 2♀ , 17.ix.2005 , F. Limeira-de-Oliveira ( CZMA ) ; Caxias , Ecológica do Inhamum , 2♀ , 30.viii.2005 , F. Limeira-de-Oliveira ( CZMA ) ; Mirador , Parque Estadual do Mirador , Base dos Cágados , 06°46’37’’S 45°06’34’’W , 1♀ , 06.viii.2011 , F. Limeira-de-Oliveira ( CZMA ) ; Mirador , Parque Estadual do Mirador , Base dos Cágados , 06°46’37’’S 45°06’34’’W , 1♀ , 27.ix–02.x.2011 , F. Limeira-de-Oliveira ( CZMA ) ; Mirador , Parque Estadual do Mirador , Base dos Cágados , 06°46’37’’S 45°06’34’’W , 1♀ , 01–05.vi.2011 , F. Limeira-de-Oliveira ( CZMA ) ; Mirador , Parque Estadual do Mirador, Base da Geraldina , 1♀ , 22viii–03.ix.2008 , F. Limeira-de-Oliveira ( CZMA ) ; Caxias , Reserva Ecológica do Inhamum , 1♀ , 25.viii.2005 , F. Limeira-de-Oliveira ( CZMA ) ; Mirador , Parque Estadual do Mirador , Base da Geraldina , 1♀ , 21–25.vi.2008 , F. Limeira-de-Oliveira , J.A. Rafael e P.A.M. Moraes ( CZMA ) ; Mirador , Parque Estadual do Mirador , Base da Geraldina , 06°37’25’’S 45°52’08’’W , 4♂ , 30.vii–06. viii.2011 , F. Limeira-de-Oliveira & D.W.A. Marques ( CZMA ) ; Mirador , Parque Estadual do Mirador, Base da Geraldina , 1♂ , 21–26.viii.2006 , F. Limeira-de-Oliveira ( CZMA ) ; Mirador , Parque Estadual do Mirador , Povoado Pindaíba ( Mel ), 06°39’44’’S 45°01’37’’W , 1♂ , 01–05.vi.2007 , F. Limeira-de-Oliveira , M.M. Abreu & J.S. Pinto-Júnio ( CZMA ) ; Mirador , Parque Estadual do Mirador , Posto Avançado do Mel , 06°43’48’’S 45°00’22’’W , 1♀ , 18–25.iii.2012 , F. Limeira-de-Oliveira & D.W.A. Marques ( CZMA ) ; Mirador , Parque Estadual do Mirador , Base dos Cágados , 06°46’37’’S 45°06’34’’W , 1♂ , 26.xi–03.xii.2011 , F. Limeira-de-Oliveira & D.W.A. Marques ( CZMA ) ; São Luis , Vila Maranhao , 1♀ , 29.ix.1992 , R . Cambra ( MIUP ) ; São Luis , Vinhais , 1♀ , 23.ix.1992 , D. Quintero ( MIUP ) ; Piauí : Parque Nacional de Sete Cidades , 2♀ , 08.ii.2013 , M.L. Oliveira ( INPA ) ; 1♀ , 10.ii.2013 , P.C. Grossi ( INPA ) ; BR135 , 35 km south of Bom Jesus , 2♀ , 01.ii.2001 , J.P. Pitts ( EMUS ) ; Parque Nacional de Sete Cidades , 2♂ , 07–13.ii.2013 , A. Somavilla ( INPA ) ; Teresina , 2♀ m, 19.vii.1993 , A. Casimiro ( MIUP ) ; Ceará , M. Calmon [Miguel Calmon], 1♂ , 18.4.1909 , Ducke ( MNHN ) ; Rio Grande do Norte : Natal , Lagoa Nova , 1♀ , 24.iv.2011 , S.M.N. T . Melo ( UFRN ) ; Natal , Campus CB UFRN, 1♀ , 03.iii.2008 , I. V . Almeida ( UFRN ) ; Natal , Campus CB UFRN, 1♀ , 25.xi.2008 , M.A. Silveira ( UFRN ) ; Natal , Parque das Dunas , 1♀ , 22.v.2008 , M.G.C. Bezerra ( UFRN ) ; Nísia Floresta , Praia de Barreta , 1♀ , 04.ii.2011 , R . M.B. Silva ( UFRN ) ; Nísia Floresta , Estrada de acesso à FLONA , 1♀ , 15.x.2005 , I. Moreno ( UFRN ) ; Baixa Verde , 1♀ , Mann ( USNM ) ; Goiás , Parque Nacional Chapada dos Veadeiros , 9♂ , 2005 ( UFES ) ; nr. Araguaina , 1♀ 10–12.ix.1963 ( BMNH ) ; Paraíba , Juazeirinho , Soledade , 4♀ , ix.1956 , C. R . Gonçalves ( DGMC ) ; Juazeirinho , Soledade , 4♀ , ix.1956 , A.G.A. Silva ( FSCA ) ; Juazeirinho , Soledade , 2♀ , vi.1956 , C. R . Gonçalves ( EMUS ) ; Souza , 1♀ , 29.xi.1938 , A. Silva ( FSCA ) ; 1240 ♀ , vi.1956 , A.G.A. Silva ( DZUP ) Santa Luzia , 1♀ , vi.1956 , A.G.A. Silva ( FSCA ) ; Santa Luzia , 1♀ , vii.1956 , A.G.A. Silva ( EMUS ) ; Tocantins : Itaguatins , Ilha São Domingos , 1♀ , 20.vii.1989 , F.F. Ramos ( MPEG ) ; Jalapão , 1♀ , 30.x.2005 , C.A. Nogueira ( CEUFT ) ; Lagoa da Confusão, Campo dos Ipês , 1♀ , 24.ix.2005 , C.A. Nogueira ( CEUFT ) ; Dianopolis , 1♀ , 16.i.1962 , J. Bechyne ( MIUP ) ; Mato Grosso : Utiariti , Rio Papagaios , 1♀ , 27.x.1966 , Lenko & Pereira ( MZUSP ) ; 1♀ , 29.x.1966 , Lenko & Pereira ( MZUSP ) ; 1♀ , x.1966 , Lenko & Pereira ( MZUSP ) ; 1♀ , 22–31.x.1966 , Lenko & Pereira ( MZUSP ) ; 1♀ , 01–12.xi.1966 , Lenko & Pereira ( MZUSP ) ; Chapada , 2♀ , x. ( CMNH ), 1♀ , iv. ( CMNH ), 1♀ , v. ( CMNH ) ; 1♀ , vi. ( CMNH ) ; Alto Xingú , P. Leonardo [Posto Indígena Leonardo Villas-Bôas, 12°11’50”S 53°22’46”W ], 7♀ , 1963, R . Arlé ( MPEG ) ; Cuiabá , Coxipó , 2♀ , 10.x.1988 , Ulisses M. Bezerra ( UFMT ) ; Cuiabá , Salgadeira , 1♀ , 28.v.1990 , Luzia ( UFMT ) ; Cuiabá , 2♀ , 05.x.1988 , Laura M. Rodrigues ( UFMT ) ; Chapada dos Guimarães , Água Fria , 1♀ , 10.v.1984 , Sebastião Marcolino ( UFMT ) ; Nova Mutum , 1♀ , 21.viii.1991 , Humberto Fonseca ( UFMT ) ; Paranatinga , Fazenda Agrochapada , Córrego Santiago , 3♀ , 18.viii.1990 , Wellington Péche ( UFMT ) ; Santa Teresinha , near Barra do Tapirapé , 2♀ , 14.i.1963 , B. Malkin ( CASC ) ; Santa Teresinha , near Barra do Tapirapé , 2♀ , 26.viii.1957 , B. Malkin ( CASC ) ; Capitão Vasconcelos , Rio Tuatuari , Xingú Basin , 1♀ , 31.vii.1957 , B. Malkin ( CASC ) ; Barra do Tapirapé , 1♀ , 11.xii.1962 , B. Malkin ( CASC ) ; Barra do Corda , 2♀ , 11.vii.1955 , Exp. Dep. Zool. ( CASC ) ; Mato Grosso do Sul , Chapada [dos Guimarães], 1♀ , i.1960 , Canuto Amann. ( MZUSP ) ; Bahia : [Chapada Diamantina], Vila do Ventura , Morro do Chapéu , 1♀ , 26.i.2011 , T . Mahlmann ( INPA ) ; Camaçari , Barra do Jacuípe , 3♀ , 21.ii.2010 , A. Köhler ( CESC ) ; 4 km northwest of Lençóis , 1♂ , 22.x.1999 , Freddy ( UEFS ) ; Lençóis , 1♂ , 29.iv.1999 , Freddy ( UEFS ) ; Guarajuba , Camacari , 4♀ , 4.i.1992 , J. Delabie ( MIUP ), 2♀ , 24.vi.1993 ( MIUP ) ; Sambaiba , 1♀ , 20.x.1994 , Nascimento ( MIUP ) ; Minas Gerais : Rosário Oeste , 1♀ , xi.1970 , Dirings ( MZUSP ) ; 1♀ , xi.1971 , Dirings ( MZUSP ) ; 2♀ , Dirings ( MZUSP ). An additional 50♂ and 352♀ from various localities in Brazil and different collections were also examined . Distribution. Brazil . Host. Unknown. Remarks. The lectotype of Mutilla bifurca is herein officially designated because, although the specimen was labelled as a lectotype by Mickel ( Fig. 31 ), the taxonomic act was never published. The taxonomic history of T. bifurca is uncertain regarding at what moment it was moved to Traumatomutilla . André (1902) moved the species from Mutilla Linnaeus to Ephuta Say s.l. which contained most species that would eventually be included in Dasymutilla Ashmead. Bradley (1916) revised Ephuta s.s. and provided a list of species north of Mexico whilst Mickel (1928) published the first comprehensive treatment of Dasymutilla . Unfortunately, neither author provided information on the status of the remaining species that once comprised Ephuta s.l. Casal (1969) was actually the first author to mention the combination Traumatomutilla bifurca without, however, providing any remarks about the species or stating that it was a new combination. Several specimens in different collections have identification labels from Mickel as Traumatomutilla bifurca (pers. com.) which is why we hypothesize that Casal assumed the species had already been moved into Traumatomutilla by Mickel even without any official publishing of the taxonomic act. Casal (1969) originally separated T. ira from T. bifurca by subtle, subjective and not easily observable characteristics, namely the sculpture of T2 above the lateral felt line and relative size of the setal patterns of T2, the limits of which are subjected to the interpretation of the observer. Neither difference was evident or even observable in any of the specimens analyzed, including reference material identified by Casal himself after 1969. Sparser setae laterally on T2 is common in several species of every species group of Traumatomutilla (e.g. T. latevittata ( Cresson, 1902 ) , T. angustata André, 1906 ). Regarding the setal patterns of T2, we observed that specimens from more southern regions ( Mato Grosso do Sul and Tocantins cerrado areas) have the oblique stripes much shorter and wider, ending before the midline of T 2 in some cases. This contrasts with specimens from north and northeastern cerrado or Caatinga areas which have long and slender setal stripes on T2. The lectotype falls between these two forms, with the setal stripes on T2 evidently longer than southern specimens and wider than northeastern specimens. Most specimens identified by Casal as T. ira are from Caatinga regions in northeastern Brazil , whilst specimens from Cerrado were usually identified as T. bifurca . We observed no structural or color characters to support separation between Caatinga and Cerrado specimens into distinct species. We have seen more than 120 specimens from Cerrado, Caatinga and Amazon areas that cannot be differentiated based on the characteristics proposed by Casal (1969) . Consequently, we propose T. ira Casal, 1969 as a junior synonym of T. bifurca . The males herein associated with T. bifurca were repeatedly and consistently collected in large numbers and in multiple areas where females of T. bifurca were commonly collected. Although T. bifurca overlaps in distribution with T. oxira in Paraíba state , these males can’t be considered as candidates for association with T. oxira because this species is apparently restricted to the locality of Juazeirinho and males and females of T. bifurca have been collected together in multiple other localities. Figures 26–27. Traumatomutilla bifurca ( Klug, 1821 ) , ♀, lectotype, line 2mm. 26) Dorsal habitus. 27) Lateral habitus. Figures 28–31. Traumatomutilla bifurca , ♀, lectotype. 28) Head, frontal view. 29) T6 (pygidium), posterior view. 30) T2, lateral view. 31) Lectotype labels. Figures 32–33. Traumatomutilla bifurca , ♂, line 2mm. 32) Dorsal habitus. 33) Lateral habitus. Figures 34–37. Traumatomutilla bifurca , ♂. 34) Head, frontal view. 35) Head, dorsolateral view (frontal tubercles in detail). 36) T6 (pygidium), posterior view. 37) T6 (pygidium), posterolateral view. In addition to the peculiar color characteristics of the bifurca species group, the ecarinate gena on the females, frontal tubercles and raised pygidium on the males set this group apart from most Traumatomutilla . Although male characteristics of T. bifurca are apparently unique and exclusive within the Dasymutillini , a large number of Dasymutilla females also have ecarinate gena and many species of Dasymutilla do not have integumental markings on T2, something that only occurs in the bifurca group within Traumatomutilla . This, however, does not indicate that T. bifurca could be transferred to Dasymutilla in the near future. This conclusion is supported by the fact that the phylogeny of Williams (2012) found evidence that Traumatomutilla and Dasymutilla are reciprocally monophyletic and that species of the latter are predominantly North or Central American ( Williams and Pitts 2013 ; Cambra et al. 2018 ).