Corethrella (Corethrella) appendiculata Grabham, 1906

Published, First, 2008, The Frog-Biting Midges of the World (Corethrellidae: Diptera), Zootaxa 1804, pp. 1-456 : 139-145

Corethrella (Corethrella) appendiculata Grabham

Corethrella appendiculata Grabham 1906:343 . Type locality: Kingston, Jamaica. Lectotype ♂ designated by Belkin et al. 1970:242 (USNM). Dyar and Shannon 1924: 216. Edwards 1930:532. Lane 1942:113, 1953:81. Matheson 1944: 92. Cook 1956: 60. Stone 1968:185. Belkin et al. 1970:242. Lounibos 1983:226. McKeever 1985a, 1985b, 1986. McKeever and French 1991b:525.

Corethrella arborealis Shannon and Del Ponte 1928:102 . Type locality: Vipos, Tucuman, Argentina. Holotype ♂ (INMA). Lane 1951:333.

Corethrella confusa Lane 1939a:109 . Type locality: Inhumas , Goyaz [= Goias], Brazil. Holotype ♂ (DEFS).

DIAGNOSIS: Male adult: only extant species of Corethrella in the New World with a pale palpus with at most some slight darkening on palpal segment 5 (as in Fig. 12B), a distinct midlength wing band and with or without more basal dark scales (Fig. 64C), with a dark brown thorax (as in Fig. 53A), dark brown midfemur, base of hind tibia darkly pigmented (contrasting with pale apex of hind femur) (as in Fig. 53A), all femora with broad scales (as in Fig. 74D), and with the dorsomedial seta from base/ gonocoxite length = 0.17–0.25 (Fig. 93D). Female adult: only extant species of Corethrella in the New World with a distinct midlength wing band and with or without more basal dark scales but these never covering all of base of wing (Fig. 70I), with a dark brown thorax ( Fig. 53A), dark brown midfemur, base of hind tibia darkly pigmented (contrasting with pale apex of hind femur) ( Fig. 53A), and all femora with broad scales (Fig. 74D).

DESCRIPTION: Male adult. Descriptive statistics: see Tables 2–5. Head: Outline in anterior view laterally elongate (as in Fig. 12B). Two large setae on frons between ventromedial area of ommatida (as in Fig. 16B). Antenna pale to medium brown; pedicel with at least one distinctive, more elongate, stout, dorsal or dorsolateral seta; flagellomeres as in Fig. 23C, sensilla coeloconica distributed as in Table 1; flagellomere 13 with well-developed apical bifurcation. Palpus pale, in some with segment 5 light brown; segment 3 of nearly constant width. Thorax (as in Fig. 53A): Dark brown, pale sclerites around base of wing. Posterior portion of dorsocentral row with group of about 8 elongate setae. Prescutal suture short, not extending more than half way to dorsocentral row of setae. Anterior anepisternum divided diagonally by sinuous suture, dorsal portion about equal to ventral portion. Ventral portion of posterior anepisternum triangular, uniformly brown, with anterodorsal margin thick. Wing (Fig. 64C): Apex of R 2 basal to apex of M 1. Anterior margin with differently, discretely pigmented scales (indicating anterior margin of midlength band), with midlength band, with some darker more basal scales restricted to veins C, Sc to forming transverse band; veins (other than costa and wing margin) with well-developed scales. Halter as dark as scutellum. Legs (as in Fig. 53A): Dark brown with knees of fore-, midlegs pale, apical 0.25 of hind femur pale, base and apex of hind tibia with non-discrete dark pigmentation, pale at midlength, at least mid-, hind leg tarsomeres 2–4 with banding. Femora, in some, tibiae and in some, some tarsomeres, with broad scales (broadest on midfemur) (also some in patch of whiplike setae on posterior portion of hind tibia). Midleg with thick, subapical setae on each of at least tarsomeres 1–3. Apices of fore-, midleg fifth tarsomeres undivided, with claws slightly subapical to apical (Fig. 75F). Claw of foreleg longer than those of mid-, hind leg (Fig. 75F). Each claw without inner tooth. Anterior claws of each leg without a basal prong. Foreleg claws unequal. Midleg claws equal. Foreleg third tarsomere shorter than fourth tarsomere. Empodia slender. Abdomen (Fig. 79H): Dark brown. Genitalia (Fig. 93D): Gonocoxite dark brown basally, pale apically, gently tapering; anteromedial area with spicules similar in length to those elsewhere on gonocoxite; with well-defined dorsal row of setae, with setae 2, 3 or 2, 3, 4 thicker than others, 1 thicker than 5, with row restricted to dorsal portion of gonocoxite. With one dorsomedial stout seta, more or less of even thickness for most of length, tapering near apex. Gonostylus (partially extended) mostly straight, curved near apex to slightly but evenly curved, slender, of more or less equal thickness for entire length, expanded apically, rounded to somewhat pointed apically; one elongate, slender subbasal seta, situated anteriorly or anteroventrally; apical seta slender, short, simple. Aedeagus slender, elongate, tapering gradually to apex, pointed apically, with lateral margins fused at apex.

Female adult. Descriptive statistics: see Tables 6–11. As for male, with following differences. Head: Coronal suture elongate, extending ventrally past ommatida (as in Fig. 16B). Antenna, light to dark brown; with flagellomeres as in Fig. 30G, sensilla coeloconica distributed as in Table 1. Clypeus ( Fig. 18R) squarish or somewhat wider than long. Mandible with small, pointed teeth. Palpus as in Fig. 34 AD; pale to light brown with palpal segment 5 darker in some. Wing (Fig. 70I). Legs: Claws of each leg equal to those of others; equal on each leg, simple (without inner teeth). Abdomen: Dark brown. Cercus dark brown.

Pupa. Described by Grabham (1906), Lane (1953), Cook (1956), McKeever and French (1991b). Thorax: Scutum, metathorax each with spherical sensory pit (as in Fig. 100A). Respiratory organ (Fig. 102I): Tubular. Abdomen (Fig. 108B): Segments 3–7 somewhat expanded laterally. Paddle only moderately elongate; apicodorsal thick spine articulating; apicoventral seta longer than thick spine.

Larva. Described by Grabham (1906), Lane (1953), Cook (1956), Borkent and McKeever (1990), McKeever and French (1991b).

Egg ( Fig. 5A). Described by McKeever and French (1991b).

DISTRIBUTION AND BIONOMICS: Corethrella appendiculata is known from Arizona and southeastern United States, throughout Central America and the Caribbean, south to northern Argentina (Fig. 118) at altitudes ranging from 0–1700 m. However, the upper elevations are represented by a female from 2km E. Santa Elena, Monteverde (1540 m) and a female from Alto de Pipe, Caracas, Miranda, Venezuela (1700 m) and these may represent different species (see below). Otherwise the highest elevation for this species is 890 m ( Brazil) with the majority of records from elsewhere under 350 m. The following give further distributional records which I have been unable to confirm: McKeever and French (1991b; United States), Belkin et al. (1970; Jamaica), Belkin and Heinemann (1975b; Bahama Islands, Cayman Islands), Belkin and Heinemann (1976b; Martinique), Heinemann and Belkin (1977b; Guatemala), Lane and Cerqueira (1958; Brazil). The single female from Piraja, Bahia, Brazil was collected by R.C. Shannon. Although there are three Piraja in the State of Bahia, Shannon and Davis (1930) described this collection site just north of Salvador (then called Bahia). The specimens from the Belkin “Mosquitoes of Middle America” project are identified in Table 12.

McKeever and French (1991b) pointed out that Nearctic specimens of C. appendiculata were restricted to the warm coastal plain in southeastern United States at elevations below 152 meters. However, further material has now become available from Mendoza Canyon, Arizona, at 946 meters. Further to this, Bradshaw and Holzapfel (1985) indicated that C. appendiculata are restricted at the north end of their range, then known in the Nearctic only from the eastern USA, by freezing temperatures. This may be so for eastern populations but the samples from Arizona indicate that at least these can withstand more severe conditions than those in the east.

The records from Gainesville and Vero Beach, Florida are from a colony maintained by L. P. Lounibos at the University of Florida. Specimens used to establish the colony were collected at both these localities.

Specimens have been collected using light, Malaise and frog-call traps and by rearing, with nearly all of these reared from treeholes or cut bamboo. Larvae are often abundant and important predators in treehole communities. A few specimens were reared from bromeliads and two from the leaf axil of a terrestrial aroid ( Xanthosoma sp ); I suspect at least some of these (especially some from Costa Rica) to represent one or more separate species. Two males and three females from Mendoza Canyon, Arizona (those collected 5–6-IX- 1968) were reared from a “stream pool” but those collected 24-IX-1970 from the same canyon were reared from a treehole, similar to that of numerous C. appendiculata from elsewhere. Hinman (1934) noted a larva of this species from a moat in Louisiana but the specimen was not examined and its identity is suspect.

The serrate mandibles of the female adults and their attraction to Hyla gratiosa calls suggest that they feed on frog blood in nature but they are known to be facultatively autogenous ( McKeever and French 1991a) and have been colonized in the laboratory without access to a blood meal. Laboratory reared females will bite offered Hyla frogs ( Hartberg and McKeever 1982; Kesavaraju, unpublished, in Lounibos et al., 2008). In some Costa Rican localities, C. appendiculata is one of the most common of species attracted to taped frog calls of Hyla gratiosa . For example, at Hitoy Cerere Biological Reserve, Costa Rica, Oct. 11, 2003, females made up 38.6% of the 754 Corethrella collected that evening. However, at La Selva Research Station, Costa Rica, March 2, 2004, C. appendiculata made up only 1–5% of the total Corethrella collected (total numbers in four samples: 41, 83, 148 and 170 specimens). McKeever and French (1991b) ran a frog-call trap in numerous localities in the USA, using the same taped call of H. gratiosa , but collected adults from only one locality.

The larvae of C. appendiculata are an important component of treehole communities in Florida, influencing distribution and density of prey and competing with other predators ( Bradshaw and Holzapfel 1984, 1985; Alto et al. 2005; Griswold and Lounibos 2005 a, b, 2006). In southern Florida they are in most treeholes bearing water ( Lounibos 1983) and they are also common in treeholes in Guanacaste, Costa Rica (Chaverri, pers. comm.). In Panama, C. appendiculata are present only in smaller treeholes in the understorey and tend to occupy the same treeholes for successive generations ( Yanoviak 1999, 2001). They are likely important in treehole communities throughout their range (Fig. 118). Bradshaw and Holzapfel (1984) reported that the larval stage overwinters in northern Florida but Lounibos (1983) found pupae in every month of the year in southern Florida, although in lesser numbers in January and February. Because there appears to be no stage which can withstand dessication, the drying of treeholes in Florida reduces population levels ( Lounibos 1983) and C. appendiculata appears to be absent during the dry season in Guanacaste, Costa Rica (see “Adult Seasonality”). It is likely that this species breeds continuously in areas with continuously wet treeholes. Although affected by complete drying, water level itself appears not have an impact on population levels ( Lounibos 1983). After a sustained dry period in 1979, populations of C. appendiculata were reduced to 10% of treeholes in southern Florida but regained normal levels within 6 weeks, again reoccupying 60% of all wet treeholes.

Frank et al. (1988) recorded a few immatures in bromeliads in Miami, Florida but these specimens were not reexamined here to confirm their identity.

McKeever and French (1991a) studied the life cycle of C. appendiculata in the laboratory and found that female ovarioles grew from an average of 44.6 µm (range: 43.0–46.2) at six hours of age to 291.8 µm (range: 286.6–297.1) after 120 hours. Eggs were produced by 94% of the females 4.5–5.5 days after emergence and these hatched in 3–6 days. Larvae matured in 11–37 days and the pupal period was 3–8 days but with 95% emerging in 5–6 days. The average number of days for a complete life cycle was 33 days. Grabham (1906) reported the pupal stage to be three weeks but this result is incongruous with not only that reported by McKeever and French (1991a) but also all other species of Corethrella ; perhaps Graham (1906) was reporting the length of the total emergence period of the pupae. McKeever (1986) noted that males and females of C. appendiculata never fed on proffered 10% sucrose under laboratory conditions.

In treeholes in Florida, later instar larvae of C. appendiculata appear to have a negative impact on the number of Aedes triseriatus (Say) . Early instar larva do not prey on mosquito larvae ( Lounibos 1983). Kesavaraju et al. (2007) found moving mosquito larvae were more susceptible to predation by larvae of C. appendiculata than were those at the water surface and that earlier instar culicid larvae were also more vulnerable. Grabham (1906) reported larvae in Jamaica fed on small Crustacea. Larvae spend most of their time anchored to substrate by their siphons and only occasionally came to the surface to breathe. At least under laboratory conditions, larvae of Toxorhynchites rutilus may prey on larvae of C. appendiculata but it is uncertain whether they do so in nature ( Lounibos 1985).

Lounibos et al. (2008) showed that mature fourth instar larvae of C. appendiculata in Florida kill but do not consume surplus mosquito larvae, especially 1–2 days before pupation. Prey were preferentially killed by grabbing and damaging the thorax of Aedes albopictus . The significance of this behavior is uncertain.

A lethal iridescent virus in Louisiana, originally found in larvae of C. brakeley i (possibly C. condita ) was successfully transmitted to larvae of C. appendiculata under laboratory condition ( Chapman et al. 1971). Larvae are susceptible to the very high concentrations of the nematode Reesimermis nielseni in the laboratory but it is unknown whether they have an impact in nature ( Ignoffo et al. 1973).

TAXONOMIC DISCUSSION: Males and females were associated through the shared presence of a common pigmentation pattern and were collected together and reared from the same habitat from a number of locations. Nevertheless, it is very likely that C. appendiculata includes more than one species. There was striking variation in wing pigmentation (especially degree of pigmentation basal to midlength band) and length, male palpal segment 3 length/width, female palpal segment 3/5, degree of pigmentation of female palpus, female number of anepimeral setae, and pupal morphology (chaetotaxy, pigmentation). However, I was unable to find consistent differences that would characterize any further species under this name.

There are certainly two species represented under the name C. appendiculata in the Osa Peninsula of southwestern Costa Rica that can be distinguished on the basis of their pupal stage. One is pale brown with, at most, slightly darker pigmentation on the abdominal tergites (either general light pigmentation or with lateral patches) and abdomen tergites 2–6 without elongate lateral setae. The other has dark brown pigmentation on the thorax and tergites 1–2, a broad medial dark brown band on tergites 3–7, pale brown segment 8, genital sacs and paddles, and abdominal segments 4–7 each with a strong lateral seta. I was unable to adequately distinguish the adults and therefore have not named the second species.

A male from 2.1 km W Teapa, Tabasco, Mexico, 100 m, 14-VII-1970 (USNM; Belkin “Mosquitoes of Middle America” project MEX 558-112; reared from bromeliad) was very similar to C. appendiculata but the gonocoxite was entirely dark brown; it is almost certainly an unnamed species but the gonocoxites were partially shriveled and in poor condition and I therefore have not named it. Another male, from just east of Ma Retraite, Paramaribo, Suriname, 5 m, 1-IV-1967 (USNM; Belkin “Mosquitoes of Middle America” project SUR 210-20; reared from bromeliad) likely has uniformly dark gonocoxites but the genitalia had been stained and could not be confidently interpreted; a female from the same locality appeared indistinguishable from those of other C. appendiculata but was excluded on the basis that it is likely conspecific with the male. The fact that the previous two specimens were reared from bromeliads, while nearly all C. appendiculata have been reared from treeholes or cut bamboo also suggests that they represent unnamed species.

A female from 2 km E. St. Elena, Costa Rica had two sensilla coeloconica on each of flagellomeres 10–12 and a female from Alto de Pipe, Caracas, Miranda, Venezuela, also reared from a bromeliad, had two sensilla coeloconica on each of flagellomeres 9 and 10 and had more lightly pigmented halters and these may also represent unnamed species. The latter two specimens were collected at the highest elevation of any C. appendiculata . The specimen from Venezuela, recognized by Belkin as “ Corethrella sp. 16", is likely teneral (weak tarsomeres) and is perhaps merely an unusual specimen of C. appendiculata . A female from French Guiana (FG 170) was identified by Belkin as “ Corethrella sp. 12" and I have identified it here as C. appendiculata . The scutellum was, unlike other C. appendiculata , significantly lighter than the scutum and it may indeed represent an unnamed species. A male adult C. bromelicola from Martinopolis, São Paulo, Brazil identified by Lane (1942) was actually a member of C. appendiculata .

McKeever (1985b) noted that some male C. appendiculata have two subbasal setae on the gonostylus but all the specimens I examined had only one. McKeever (1985a, b) provided detailed studies of the internal reproductive systems of female and male C. appendiculata , respectively.

Some of the female C. appendiculata from 5 km NE Tárcoles, Costa Rica, 2-IX-1993 studied here were sequenced by Pawlowski et al. (1996) in their molecular analysis of the Culicomorpha (as Corethrella sp. ).

The lectotype of C. appendiculata and the holotypes of C. arborealis and C. confusa were originally on pins but were placed on microscope slides for this study. The male genitalia of C. confusa had been previously slide-mounted. Grabham (1906) described C. appendiculata on the basis of males, females, pupae and larvae but did not indicate how many specimens of each were in the type series. Belkin et al. (1970) noted the presence of 10 cotypes in the series from which they designated a lectotype and I have also studied these. Of the remaining paralectotypes, one male was slide-mounted, four males and two females were left on pins, and two pinned specimens were in such poor condition that their gender was uncertain. It is not known where the pupal and larval paralectotypes are located. I have added paralectotype labels to this series of specimens.

McKeever (1986) provided a detailed description of the male and female adult mouthparts, including excellent SEM photomicrographs and McKeever (1985a, 1985b) described the male and female reproductive systems. McKeever (1986) described the male and female mouthparts of C. appendiculata . The dorsal apotome of the pupa was described by Reinert (1999). Hartberg and McKeever (1982) described the 6 diploid chromosomes (2n = 6) of C. appendiculata .

The first instar larva of C. appendiculata has a prothoracic proleg represented by a transverse row of hooks ( Borkent and McKeever 1990). Their function is unknown but likely serve to help larvae reach water when eggs are thrown above waterline in treeholes.

Most of the specimens from the Belkin “Mosquitoes of Middle America” project included below likely have associated larval and/or pupal exuviae but I have not searched for these in the collection of the USNM.

The holotype of C. confusa did not have any locality label but Lane (1939a) reported the specimen to have come from Inhamas, Goyaz, Brazil. Edwards (1930) reported two specimens in the BMNH from Grenada but I did not examine these and their identification, therefore, is uncertain.

MATERIAL EXAMINED: Lectotype, adult male on microscope slide, labeled “ Lectotype ♂ Corethrella appendiculata Grabham, 1906 by B., H.+ P, 1970”, “Kingston, Jam”, “M Grabham Collector”, “Cotype No. 9960 U.S. N.M.” ( USNM). Holotype, adult male on two microscope slide, first one with most of body labeled, “ Corethrella confusa Ln. 39”, “ Holotype ”, “4333”, “1.080”, “ Corethrella appendiculata Ln. , 41 Gbm, 06”; second one, with genitalia labeled. “ Corethrella confusa Ln. 39, 4333, ♂ term., No. 1.080, Lamina 279, Divisao 1, Gaveta 136, HOLOTYPE - genitalia Corethrella confusa Lane ”, ( DEFS) . Holotype, adult male on microscope slide, labeled, “ Corethrella arborealis Shannon & Del Ponte ”, “ Holotypus ”, “Type”, “389”, “#174 Ex treehole, Vipos, 4.III.27, E. Del Ponte ”, “ Corethrella appendiculata Det. A. Borkent ” ( INMA) . 5 ♂, 2 ♀, 2 unknown gender paralectotypes of C. appendiculata labeled as for holotype (1 ♂ on slide; remainder on pins; USNM) . Other material: 2 ♂, 3 ♀, Arizona, Pima Co., Mendoza Canyon , 5-IX-1968 ( USNM) ; 2 ♀, from previous locality but 6-IX-1969 ( USNM) ; 11 ♀, from previous locality but 24-IX-1970 ( USNM) ; 1 ♀, Macon , Georgia, United States, X-1944 ( DEFS) ; 3 ♂, 2 ♀, Sulphur , Louisiana, United States, 4- IV-1968 ( CNCI) ; 1 ♀, Moss Bluff , Louisiana, United States, 12-X-1965 ( CNCI) ; 3 ♂, 3 ♀, 4 pupal exuviae, 2 fourth instar larvae, from either Gainesville and/or Vero Beach, Florida, from colony mainained by L.P. Lounibos ( CNCI) ; 3 ♂, 1 ♀, 10 km W Ciudad Obregon, Sonora , Mexico, 20 m, 4-VI-1971 ( USNM) ; 1 ♂, 3 ♀, 7.4 km N Comalcalco , Tabasco, Mexico, 10 m, 12-VII-1970 ( USNM) ; 1♂, Reserva Ria Celestún , Celestún , Km 7.5 Carretera Celestún-Kinchil, Yucatán, Mexico, 0–15 m, 8–9-IV-1997 ( IDRE) ; 2 ♂, 1 ♀, 34 km S. Chilpancingo, Guerrero, Mexico, 900? m, 7-VIII-1966 ( USNM) ; 2 ♂, 1 ♀, 15 km from Paso de San Juan , Ver- acruz, Mexico, 100 m, 9-VII-1970 ( USNM) ; 1 ♂, Tamazunchale , San Luis Potosi, Mexico, 200 m, 21-VII- 1965 ( USNM) ; 1 ♀, across Rio Moctezuma from Tamazunchale, San Luis Potosi, Mexico, 200 m, 20-VII- 1965 ( USNM) ; 1 ♀, Tapachula , Chiapas, Mexico, 20-IX-1941 ( USNM) ; 1 ♂, about 1.5 km NW Bluefields, Zelaya , Nicaragua, 5 m, 248- XI-1971 ( USNM) ; 2 ♂, 2 ♀, each with pupal exuviae, 4 pupae, Santa Cruz ACT Bosque Nacional Diria, Guanacaste, Costa Rica, 131 m, 2-XII-2000 ( INBC) ; 1 ♂, OTS field station, Palo Verde , Puerto Humo, Guanacaste, Costa Rica, 15 m, 20-VIII-1971 ( USNM) ; 1 ♀, OTS field station, about 5 km NE of headquarters, Palo Verde , Puerto Humo, Guanacaste, Costa Rica, 10 m, 18-VIII-1971 ( USNM) ; 2 ♂, Estación Los Negritos , Sector Las Torres , Palo Verde National Park, Bagaces, ACT, Guanacaste, Costa Rica, 60 m, 21-IX-2000 ( INBC) ; 1 ♀ and pupal exuviae, 1 pupa, from previous locality but 40 m ( INBC) ; 1 ♀, Cerro Noguera , Palo Verde National Park, Bagaces, ACT, Guanacaste, Costa Rica, 180 m, 22-IX-2000 ( INBC) ; 1 ♀, pupal exuviae, Sendero Heliconias , Volcán Arenal National Park, Alajuela, Costa Rica, 550-620 m, 10-XII-1999 ( INBC) ; 1 ♀, 2 km E Santa Elena, Monteverde , Costa Rica, 1540 m, 26-VIII-1993 ( CNCI) ; 1 ♀, Rio Esparta , 1 km S Esparta on road to Artieda, Puntarenas, Costa Rica, 110 m, 28-XII-1971 ( USNM) ; 2 ♂, 1 ♀, 1 km S Pitahaya , Puntarenas, Puntarenas, Costa Rica, 1 m, 29-XII-1971 ( USNM) ; 1 ♂, larval, pupal exuviae, Estación Quebrada Bonita , Carara National Park, Costa Rica, 10 m, 3-XII-2002 ( INBC) ; 1 ♀, from previous locality but 26-VII-1993 ( CNCI) ; 5 ♀, Tárcoles , Costa Rica, 11-XI-1993 (4 ♀, CNCI; 1 ♀, IDRE) ; 16 ♀, 5 km NE Tárcoles , Costa Rica, 2-IX-1993 ( CNCI) ; 12 ♀, from previous locality but 17-VIII-1993 ( CNCI) ; 1 ♀, from previous locality but 20-VII-1993 ( CNCI) ; 7 ♀, Manuel Antonio National Park , Costa Rica, 17-XI- 1993 ( CNCI) ; 1 ♀, pupal exuviae, 200 m N Casa Marenco, Sector Los Patos , Corcovado National Park, Puntarenas, Costa Rica, 78 m, 22-II-2000 ( INBC) ; 1 ♂ with pupal exuviae, 1 ♀, Sendero Los Patos-Sirena , Corcovado National Park, Puntarenas, Costa Rica, 78 m, 16-V-2000 ( INBC) ; 1 ♂, larval, pupal exuviae, El Neque, Jimenez , Golfito, Costa Rica, 3 m, 22-VII-2002 ( INBC) ; 1 ♀, pupal exuviae, 0.3 km S del puente sobre el Rio Tamales, Jiménez , Golfito, Costa Rica, 10 m, 14-I-2003 ( INBC) ; 1 ♀, pupal exuviae, Estación El Bonito, Piedras Blancas National Park , Golfito, Costa Rica, 100 m, 21-VII-2001 ( INBC) ; 1 ♂, pupal exuviae, Estación La Leona , Falda SE del Cerro Rubio, Corcovado National Park, Puntarenas, Costa Rica, 200 m, 19-VII-2002 ( INBC) ; 1 ♂, pupal exuviae, 400 m W. entrada al Parque , Rivera rio Rincón , Sector Los Patos, Corcovado National Park, Puntarenas, Costa Rica, 86 m, 23-II-2000 ( INBC) ; 1 ♂, 1 ♀ with pupal exuviae, 1 ♀ with larval and pupal exuviae, Estación Isla del Caño , Corcovado National Park, Puntarenas, Costa Rica, 90 m, 28-VIII- 2000 ( INBC) ; 1 ♂, pupal exuviae, as for previous locality but 29-VIII-2000 ( INBC) ; 1 ♂, pupal exuviae, 200 m N. Casa Marenca, Corcovado National Park , Puntarenas, Costa Rica, 78 m, 22-II-2000 ( INBC) ; 1 ♂ with pupal exuviae, 1 ♀ with larval and pupal exuviae, Estación Sirena , Corcovado National Park, Puntarenas, Costa Rica, 3 m, 30-XI-2001 ( INBC) ; 1 ♂, 3 ♀, 5 km NW Rincón, Osa Peninsula , Costa Rica, 10-VIII-2001 (1 ♂, IDRE; 3 ♀, CNCI) ; 1 ♂, 5 km W. Rincón, Sierpe Centro Juvenil Tropical , Aguabuena, Costa Rica, 80 m, 10- VIII-2001 ( INBC) ; 4 ♂ (1 with associated larval and pupal exuviae), El Naranjal, Silvestre Golfito R.N., Puntarenas, Costa Rica, 80 m, 22-VII-2001 ( INBC) ; 5 ♀, Orilla de E Agujas, Golfo Dulce R.F., Puntarenas, Costa Rica, 250-350 m, 30-VI-1-VII-2000 ( INBC) ; 2 ♀, nr administration building (9°58.52'N, 83°27.15'W), Barbilla National Park , 500 m, 10-II-2006 ( CNCI) ; 1 ♂, Finca La Selva , Puerto Viejo, Heredia, Costa Rica, 100 m, 8-VIII-1971 ( USNM) ; 7 ♀, La Selva Biological Station , Puerto Viejo de la Sarapiqui, Heredia, Costa Rica, 40 m, 1-III-2004 ( CNCI) ; 6 ♀, as for previous locality but 2-III-2004 ( CNCI) ; 1 ♂, 1 ♀, each with pupal and larval exuviae, 1 ♂, 1 ♀, each with pupal exuviae, Sendero a la Catarata, Hitoy Cerere Biological Reserve, Costa Rica, 250 m, 3-XI-2000 ( INBC) ; 17 ♀, Hitoy Cerere Biological Reserve , nr. administration building (9°40.30'N, 83°01.20'W), 100 m, 11-X-2003 ( CNCI) ; 1 ♀, 3 km E. Cahuita , Costa Rica, 30-X-1993 ( CNCI) ; 1 ♂, pupal exuviae, Estación Puerto Vargas , Cahuita National Park, Prov. Limón, Costa Rica, 3 m, 19-VI-2004 ( CNCI) ; 3 ♀, Reserva Chimuri , Cahuita, Costa Rica, 15 m, 7-VI-2000 ( CNCI) ; 1 ♂, Barro Colorado , Panama, 10-I-1957 ( USNM) ; 1 ♂, as previous locality but 31-V-1996 ( CNCI) ; 1 ♂, 2.1 km S Margarita, Colon, Canal Zone , Panama, 5 m, 22-XI-1965 ( USNM) ; 1 ♂, 1 ♀, Gatun , Canal Zone, Panama, 20-X-1926 ( USNM) ; 1 ♂, Finca Engombe , Santo Domingo, Distrito Nacional, Dominican Republic, 20 m, 20-VIII-1971 ( USNM) ; 1 ♀, as previous locality but 26-VIII-1971 ( USNM) ; 1 ♂, as previous locality but 1-IX-1971 ( USNM) ; 2 ♂, 1 ♀, Antrim Valley , Massacre, St. Paul, Dominica, 250 m, 7-VI-1965 ( USNM) ; 1 ♂, 1 ♀, Bise Quarry , Castries, Castries, St. Lucia, 30 m, 17-VII-1964 ( USNM) ; 1 ♀, Grande Anse , Dauphin, St. Lucia, 50 m, 20-VII-1964 ( USNM) ; 1 ♂, Three Rivers , Colonairie, Charlotte, St. Vincent, 200 m, 13-XII-1963 ( USNM) ; 1 ♀, Freeland , Mesopatamia, St. George, St. Vincent, 150 m, 23-XI-1963 ( USNM) ; 2 ♀, Tucker Valley , Trinidad and Tobago, 1 m, 23-X-1957 ( USNM) ; 1 ♂, St. George , Verdant Vale, 200 m, Trinidad and Tobago, 29-I-1966 ( USNM) ; 1 ♂, 1 ♀, St. George , Guanapo Valley, “AripoGuanapo Trace 9 milepost “, Trinidad and Tobago, 150 m, 16-IV- 1964 ( USNM) ; 1 ♂, 1 ♀, St. George , about 3.5 km E of Mason Hall, Caledonia, Trinidad and Tobago, 250 m, 26-XI-1965 ( USNM) ; 1 ♀, Nariva , “Archers Estate “, Trinidad and Tobago, 50 m, 5-XI-1964 ( USNM) ; 1 ♀, Nariva , “Charuma Forest “, Trinidad and Tobago, 50–150 m, 8-X-1964 ( USNM) ; 1 ♂, 1 ♀, MAC Cacao Dispersion Center , Ocumare de la Costa, Aragua, Venezuela, 100 m, 12-VII-1969 ( USNM) ; 1 ♀, Alto de Pipe , Caracas, Miranda, Venezuela, 1700 m, 26-XI-1967 ( USNM) ; 1 ♂, Guamitas , Maracay, Aragua, Venezuela, 740 m, 15-VII-1969 ( USNM) ; 1 ♀, Maracay , Venezuela, 27-VI-1927 ( USNM) ; 1 ♀, 5 km SE Cayenne, Guyane , French Guiana, 5 m, 31-I–1-II-1965 ( USNM) ; 1 ♀, 45 km S Sinnamary, Inini , French Guiana, 20 m, 21- III-1967 ( USNM) ; 3 ♂, 1 ♀, Bahia, Brazil (1 ♂, 1 ♀, USNM; 2 ♂, BMNH) ; 2 ♂, 6 ♀, Bahia , Brazil, III-1929 ( USNM) ; 1 ♀, as previous locality but XII-1928 ( USNM) ; 1 ♀, Piraja , Bahia, Brazil, 7-II-1929 ( USNM) ; 1 ♂, Martinopolis , São Paulo, Brazil, IX-1940 ( USNM) ; 1 ♂, 1 ♀, Boraceia Field Station , Salesopolis, São Paulo, Brazil, 890 m, 26-I-1975 ( USNM) ; 1 ♀, as previous locality but 850 m, 22-II-1975 ( USNM) .

DERIVATION OF SPECIFIC EPITHET: The name appendiculata (appendage) may refer to the larval posteroventral hooks referred to by Grabham (1906) as “appendages”.