Simulium (Psaroniocompsa) jujuyense Paterson & Shannon

HERNÁNDEZ, LUIS MIGUEL, SHELLEY, ANTHONY JOHN, DE LUNA DIAS, ANTONIO PAULINO ANDRADE & MAIA-HERZOG, MARILZA, 2007, New specific synonymies and taxonomic notes on Neotropical black flies (Diptera: Simuliidae) belonging to the subgenera Chirostilbia Enderlein, Hemicnetha Enderlein, Inaequalium Coscarón & Wygodzinsky, Psaroniocompsa Enderlein and Psilopelmia Enderlein, Zootaxa 1506 (1), pp. 1-80 : 23-25

Simulium (Psaroniocompsa) jujuyense Paterson & Shannon

( Figs. 35–46 , 67–76 , 88–92 , 106–110 , 145–159 , 187–198 , 217–220 )

Simulium jujuyense Paterson & Shannon, 1927: 741 . SYNTYPES ♀ (man-biting). ARGENTINA: Zapla , Jujuy; iii.1927, (Paterson) [Syntype series lost.]

Simulium auripellitum Enderlein, 1934a: 286 . HOLOTYPE ♀, PARAGUAY: Hohenau , 250m; 12.x.1907, [Without collector’s name.] (SMT) [Examined.] New synonymy.

Psaroniocompsa mendozana Enderlein, 1936: 123 . HOLOTYPE ♂, ARGENTINA: Mendoza; [Without date.], (Jensen- Haarup, S.) — ♂ (NMHU) [Examined.] [Synonymy by Wygodzinsky, 1951: 207.]

Pliodasina guttata Enderlein, 1936: 124 . HOLOTYPE ♀, PARAGUAY: San Bernardino ; v.1910, (Fiebrig, K.) (NMHU) [Examined.] New synonymy.

Simulium bonaerense Coscarón & Wygodzinsky, 1984: 59 . HOLOTYPE ♀ (reared), Buenos Aires, Arroyo Azul, prox. a Azul y ruta 3; 26.ix.1971, (Coscarón & Wygodzinsky) (MLP) [Examined.] New synonymy.

Simulium minuanum Strieder & Coscarón 2000: 103–117 . HOLOTYPE ♀ (reared). ARGENTINA: Missiones , Azara, Arroyo Chimiray; 11.i.1999, (Strieder, M.N. & Martins, S.) (LEU) New synonymy.

Simulium jujuyense was described by Paterson & Shannon (1927) from ten man-biting females collected from the provinces of Jujuy (Zapla, Ledesma) and Tucumán in Argentina between iii–ix.1927. The location of the syntype series is unknown. All South American simuliid species described by Paterson & Shannon (1927) could not be located in any institution in Argentina or in the USNM, and they are now considered to be lost (Sixto Coscarón, pers. com. to Luis Hernández in 2005). Wygodzinsky (1951) described all life stages of S. jujuyense from reared specimens and man-biting females collected in Argentina and Ecuador, and discussed its taxonomic position with related species within Simulium . The modern concept of this species is based on material cited in his work. The general morphology of the adult thoracic pattern, genitalia and pupal gill configuration of material identified as S. jujuyense either by Wygodzinsky or Coscarón and deposited in the AMNH, MLP and BMNH is shown in Figs. 35, 36 , 67, 68 , 145–147 , 187–189 (see also Coscarón & Wygodzinsky, 1984). We are not designating a neotype at this stage for S. jujuyense because of the morphological variation found in different populations of this species, especially the pupal gill configuration, that indicates the presence of a species complex and the lack of a well-preserved, long series of reared specimens. This should be done at a later date using reared-link adults from the type locality in conjunction with molecular and cytogenetic studies.

Several species have been synonymized with S. jujuyense on which we have the following comments and we also provide explanations for four new synonyms proposed in the current work. Three others species S. spinifer Knab , S. hoffmani Vargas, and S. sicuani Smart have been recorded as synonyms of S. jujuyense by Coscarón, (1987, 1991), which is followed by Crosskey & Howard (2004), and more recently by Coscarón & Coscarón-Arias (2007). After further examination of type material these species are not here considered as synonyms of S. jujuyense and they are discussed in detail under S. spinifer .

Simulium auripellitum was described by Enderlein (1934a) based on a female holotype collected by Schrottky biting man at Hohenau in Paraguay (see Material Examined). The original description covers too few characters for identification today, but Coscarón & Wygodzinsky (1984) provided further details following examination of the type material and other specimens from Brazil and Paraguay. In his description of S. jujuyense Wygodzinsky (1951) stated that S. auripellitum is very similar to this species, but refrained from any synonymy due to some differences in leg coloration, smaller body length in auripellitum and the fact that Enderlein placed this species in Simulium s.s. in which females do not have claws on the legs. Shelley et al. (2000) comprehensively reviewed the taxonomy of S. auripellitum and suggested that the minor morphological differences between S. jujuyense , S. bonaerense and S. auripellitum indicated possible synonymy with S. jujuyense as the senior synonym. We have re-examined the holotype of S. auripellitum , which is in the SMT. The specimen is in good condition and has been pinned through the left side of the scutum. We have taken digital images of the holotype prior to dissection of the head, legs [only three partly damaged, the others are missing], wings, abdomen and genitalia, which are now on a slide; the thorax remains pinned. We have also compared this material with numerous linked-reared specimens identified as S. auripellitum in the BMNH and IOC collections. The thoracic pattern of the holotype of S. auripellitum ( Fig. 37, 38 ) and the morphology of the nudiocular area and cibarium ( Figs. 89 , 107 ), wing venation, legs coloration, and genitalia ( Figs. 148–150 ), agree with the general morphology of specimens identified as S. auripellitum from Brazil deposited in the IOC and BMNH ( Figs. 35–40 ) (see also Shelley et al., 2000). The general morphology of this material falls within the variation found in S. jujuyense ( Figs. 35–32 , 69–70 ). In addition, the configuration of the pupal gill filaments also lies within the variation found in S. jujuyense ( Wygodzinsky, 1951; see also Fig. 131 in Shelley et al., 2000). Therefore, we regard both species as conspecific.

We confirm the synonymy of Psaroniocompsa mendozana with S. jujuyense proposed by Wygodzinsky in 1951. Enderlein (1936) described P. mendozana based on a single male collected in Mendoza, Argentina. We have examined the holotype, which is deposited in the NMHU. The specimen is in relatively good condition with the left wing, abdomen and genitalia on a slide (Material Examined). The thoracic pattern ( Figs. 71, 72 ) and the morphology of the genitalia ( Figs. 190, 191 ) fall within the variation of S. jujuyense .

We consider S. guttatum (Enderlein) to be a new synonym of S. jujuyense for the following reasons. Pliodasina guttata was superficially described by Enderlein in 1936 based on a single female collected in San Bernardino, Paraguay by Fiebrig. Vargas & Díaz Nájera (1953) then examined this specimen and stated that it was very close to S. incrustatum , but refrained from synonymising it with the latter species. In 1962 Stone also examined this specimen, agreed with Vargas & Díaz Nájera’s (1953) statement and placed Pliodasina as a synonym of Notolepria Enderlein. Stone followed this paper in 1963 with an annotated list of generic group names in the family Simuliidae . He concluded that Vargas & Diaz Nájera (1953) incorrectly assigned S. incrustatum Lutz to the subgenus Notolepria , and considered both S. incrustatum and S. guttatum as belonging to the subgenus Simulium , thereby making Pliodasina Enderlein a junior synonym of Simulium . This subgeneric synonymy has been followed by Coscarón (1987, 1991) and Crosskey & Howard (2004). Coscarón & Wygodzinsky (1984), and Coscarón (1987, 1991) considered S. guttatum (Enderlein) as a species inquirenda in the subgenus Psaroniocompsa Enderlein. However, Crosskey & Howard (2004) cited it as a valid species in the incrustatum species group of the subgenus Psaroniocompsa . We have examined the holotype of S. guttatum , which is housed in the NMHU. We have taken digital images of the thorax prior to dissection to the head and four legs, which are on a second slide. One wing, one front and hind leg and the genitalia have been previously dissected and are on two slides; the thorax and part of the abdomen remain pinned (see Material Examined). The pinned female has few setae left on the thorax and so the setal pattern is not evident [this pattern has been used to separate S. auripellitum from S. incrustatum by Shelley et al., 1997; 2000]. Consequently, comparison was made between the distribution of setal scars of S. guttatum and that of a female S. auripellitum [here synonymized with S. jujuyense ] in which part of the thorax was devoid of setae and a female topotype of S. incrustatum with setae missing. Scar distribution indicated that in S. guttatum the setae were arranged randomly as in S. auripellitum and not in groups in longitudinally diverging lines as in S. incrustatum . The female thoracic pattern ( Figs. 41, 42 ), head morphology (nudiocular area and cibarium) ( Figs. 90 , 108 ), wing venation, leg coloration and genitalia ( Figs. 151–153 ) of S. guttatum all fall within the variation found in S. jujuyense . Therefore, we remove S. guttatum from its synonymy with S. incrustatum and place it as a synonym of S. jujuyense .

Another species very similar to S. jujuyense is S. bonaerense described by Coscarón & Wygodzinsky in 1984 from reared adults collected in Argentina (Rio Negro and Buenos Aires). The authors stated in the original description that all the specimens previously identified by Coscarón & Wygodzinsky (1960) and Coscarón (1968) as S. jujuyense belong to S. bonaerense . They were only able to separate S. bonaerense from S. jujuyense and S. auripellitum by the primary branches of the gill bifurcating more basally and its relatively larger size. They further separated S. bonaerense from S. auripellitum by the more intense greenish yellow pubescence on its thorax. However, Shelley et al. (2000) suggested that the minor morphological differences between S. jujuyense , S. bonaerense and S. auripellitum indicate a possible synonymy with S. jujuyense as the senior synonym. We have examined the holotype of S. bonaerense , which is deposited in the MLP, and several females and males (as paratypes) deposited in the MLP and BMNH collections. The holotype is in good condition and is glued by the left side of the thorax to a card point together with its pupal pelt. The thoracic pattern ( Figs. 43, 44 ), female nudiocular area ( Fig. 91 ), cibarium ( Fig. 108 ), and the morphology of the female ( Figs. 154–156 ) and male genitalia ( Figs. 193–195 ) are very similar to that found in S. jujuyense . In all the specimens examined the pupal gill filaments always bifurcate more basally ( Fig. 217 ) than in S. jujuyense and the filaments appear to be stouter and shorter. However, variation in this pattern occurs. We have examined one specimen from Buenos Aires, identified by S. Coscarón in 1979 as S. bonaerense , in which the dorsal primary branch bifurcates more apically than the bifurcations on the other two primary branches ( Figs. 218, 219 ). This represents a variation between the typical S. bonaerense configuration and the highly variable one of S. jujuyense and it is probable that more intermediate variations will be found if larger numbers of specimens are examined. Similarly, pupal gill configuration was seen to vary considerably in species of the amazonicum group when large numbers of specimens were available for examination ( Shelley et al., 2006). Therefore, we synonymize S. bonaerense with S. jujuyense .

Simulium minuanum was described by Strieder & Coscarón (2000) from numerous adults, pupae and larvae collected in Argentina and Brazil. Strieder & Coscarón (2000) recognized the close similarities between S. minuanum and S. auripellitum . They produced a table in which they compared S. minuanum with other related species ( S. angrense , S. auripellitum , S. jujuyense and S. bonaerense ). The only salient character that distinguished S. minuanum from the other species being the absence of a sub-basal tooth on the claw, which is present in S. angrense and S. auripellitum , but reduced in S. jujuyense and S. bonaerense . The holotype is said to be have been deposited in LEU, but we have been unable to obtain this material for study. However, we have examined numerous paratypes and specimens with no type status in the MLP and BMNH collections (Material Examined). All specimens appear to have been recovered from alcohol and some of the thoraxes have collapsed. Nevertheless, the thoracic pattern ( Figs. 45, 46 ), female head (nudiocular area and cibarium ( Figs. 92 , 110 ) and the morphology of the female ( Figs.157–159 ) and male genitalia ( Figs. 196–198 ) and the pupal gill configuration ( Fig. 220 ) fall within the variation found in S. jujuyense and we consider both species as conspecific. We regard the presence or absence of a tooth on the claw as varying intraspecifically in several Neotropical species and hence do not accept this character as interspecific.

The female of S. jujuyense is similar to S. incrustatum from which it can only be distinguished by the different setation and thoracic pattern (see Shelley et al., 2000). In Brazil it is a widely distributed species in smaller rivers in southern and central states (Material Examined; Coscarón, 1987, 1991). It has also been recorded from Argentina (type locality), Bolivia, Colombia, Ecuador, Paraguay and Uruguay (see Material Examined; Coscarón, 1987, 1991; Coscarón and Wygodzinsky, 1984; Crosskey & Howard, 2004). In Argentina this species breeds in small streams and rivers attached to submerged vegetation together with S. wolffhuegeli (Enderlein) and S. rubiginosum (Enderlein) ( Coscarón & Wygodzinsky, 1984; Coscarón, 1991) and the females are anthropophilic. In Brazil, streams in forested areas are favoured and only small numbers of specimens are ever found in rivers or biting man. Wygodzinsky (1951) stated that in Tucumán province, Argentina, the females of S. jujuyense are highly anthropophilic and very aggressive. In the region of Aconquija this author carried out experimental infection with the nematode Mansonella ozzardi (Manson) , but no development of the worm was recorded. The record of S. jujuyense for Peru ( Crosskey & Howard, 2004) stems from the synonymy of S. spinifer with this species by Coscarón (1987, 1991). However, S. spinifer is considered here as a valid species in Simulium s.l. [see S. spinifer ]. Consequently, Peru has been omitted form the distribution record of this species.

Figures

PLATE 3. FIGURES 33–44. Colour patterns of the female thorax. For each species first figure is with anterior illumination and second figure with posterior illumination. 33, 34: Psaroniocompsa opalinifrons holotype; 35, 36: S. jujuyense (Argentina); 37, 38: S. auripellitum holotype; 39, 40: S. auripellitum (Brazil); 41, 42: Pliodasina guttata holotype; 43, 44: S. bonaerense holotype.

PLATE 4. FIGURES 45–58. Colour patterns of the female thorax. For each species first figure is with anterior illumination and second figure with posterior illumination. 45, 46: S. minuanum paratype; 47, 48: S. costaricensis holotype; 49, 50: S. stellatum holotype; 51, 52: S. spinifer holotype; 53, 54: S. spinifer paratype; 55, 56: S. hoffmani lectotype; 57, 58: S. sicuani lectotype.

PLATE 5. FIGURES 59–78. Colour patterns of the male thorax. For each species first figure is with anterior illumination and second figure with posterior illumination. 59, 60: S. acarayense paratype; 61, 62: S. clarki paratype; 63, 64: S. nahimi paratype (as allotype); 65, 66: S. subnigrum (Brazil); 67, 68: S. jujuyense (Argentina); 69, 70: S. auripellitum (Brazil, with silver cunae]; 71, 72: Psaroniocompsa mendozoana holotype; 73, 74: S. bonaerense paratype; 75, 76: S. minuanum paratype; 77, 78: S. sicuani paralectotype.

PLATE 6. FIGURES 79–96. Female nudiocular area. 79: Thyrsopelma striginotum holotype; 80: S. pillosum holotype; 81: S. dekeyseri paratype; 82: S. acarayense paratype; 83: Trichodagmia manicata lectotype; 84: Thyrsopelma argentata holotype; 85: S. clarki paratype; 86: S. subnigrum (Brazil); 87: Psaroniocompsa opalinifrons holotype; 88: S. jujuyense (Argentina); 89: S. auripellitum holotype; 90: Pliodasina guttata holotype; 91: S. bonaerense paratype; 92: S. minuanum paratype; 93: S. costaricensis holotye; 94: S. spinifer holotype; 95: S. hoffmani lectotype; 96: S. sicuani lectotype.

PLATE 7. FIGURES 97–114. Female cibarium. 97: Thyrsopelma striginotum holotype; 98: S. pillosum holotype; 99: S. dekeyseri paratype; 100: S. acarayense paratype; 101: Trichodagmia manicata lectotype; 102: Thyrsopelma argentata holotype; 103: S. clarki paratype; 104: S. subnigrum (Brazil); 105: Psaroniocompsa opalinifrons holotype; 106: S. jujuyense (Argentina); 107: S. auripellitum holotype; 108: Pliodasina guttata holotype; 109: S. bonaerense paratype; 110: S. minuanum paratype; 111: S. costaricensis holotye; 112: S. spinifer holotype; 113: S. hoffmani lectotype; 114: S. sicuani lectotype.

PLATE 9. FIGURES 127–141. Female genitalia (cercus and pararprocts, eighth sternite and gonopophyses). 127–129: Trichodagmia manicata lectotype. 130–132: Thyrsopelma argentata holotype; 133–135: S. clarki paratype; 136–138: S. subnigrum (Brazil). 139–141: S. subnigrum (specimens previously identified as S. nahimi by L.M.Hernández).Arrow indicates a sclerotized artefact on the left lateral arm of the genital fork that resembles an anterior process.

PLATE 10. FIGURES 142–156. Female genitalia (cercus and paraproct, eighth sternite and gonopophyses (except for S. costaricensis).142–144: Psaroniocompsa opalinifrons holotype; 145–147: S. jujuyense (Argentina); 148–150: S. auripellitum holotype; 151–153: Pliodasina guttata holotype; 154–156: S. bonaerense paratype.

PLATE 11. FIGURES 157–170. Female genitalia (cercus and pararprocts, eighth sternite and gonopophyses). 157– 159: S. minuanum paratype; 160–161: S. costaricensis holotye; 162–164: S spinifer holotype; 165–167: S. hoffmani lectotype; 168–170: S. sicuani lectotype.

PLATE 13. FIGURES 183–201. Male genitalia (ventral plate, gonocoxite and gonostyle, paramere). 183–185: S. subnigrum (Brazil), 185: Specimen identified as S. nahimi by L.M.Hernández; 187–189: S. jujuyense (Argentina); 190–192: Psaroniocompsa mendozana holotype; 193–195: S. bonaerense paratype; 196–198: S. minuanum paratype; 199–201: S. sicuani paralectotype.

PLATE 15. FIGURES 217–220. Pupal gill filaments. 217: S. jujuyense (Argentina); 218: S. bonaerense paratype; 219: S. bonaerense (gill variation, Argentina); 220: S. minuanum paratype. Arrow indicate the more apical bifurcation of the dorsal primary branch.