Dryophthoridae of Costa Rica and Panama

Cactophagus LeConte, 1876

Diagnosis: Small to large, 7-25 mm in total body length; elongate-oval in form, uniformly black or black with red, orange or yellow markings; antennal funicle of 6 articles, scape cylindrical, apex extended to posterior margin of eye or beyond; rostrum cylindrical, curved ventrally or straight; anterior margin of pronotum straight behind eye; tarsus of 5 articles, but article 4 small, located at very base of article 3, article 3 wider than 2 and bilobed ventrally, ventral pilosity various, forming pads which cover one-third or more of ventral surface, article 5 (claw-bearing article) with apical venter smooth; legs of males and females generally similar in shape and vestiture, front coxae separated, some species with tubercle between front coxae or with metasternum markedly produced and subconical between mesocoxae; pygydium exposed beyond elytra; scutellum triangular, widest at or near base; male with aedeagus lacking lateral line.

Taxonomic notes: Separation of Cactophagus from some Rhodobaenus and Metamasius can be difficult.

Natural history: Where known, species of Cactophagus are associately mostly with arums (Araceae) and orchids (Orchidaceae). Despite the generic name, only a couple of species are associated with Cactaceae. While preliminary, based on the meager biological information available, species of Cactophagus may be restricted to Araceae and Orchidaceae. In contrast, species of Metamasius may prove restricted to Arecaceae and Bromeliadaceae.

Diversity: About 60 species of Cactophagus are known from Central and South America; 28 are known from Costa Rica and 24 from Panama, with 18 shared between the two countries.

	Key to species of Cactophagus in Costa Rica and Panama
1	Pronotum deeply laterally incised subapically immediately behind eye; in dorsal view, lateral margin with distinct, deep subapical emargination, apex distinctly tubulate; pronotum and (especially) elytra with coarse, irregular sculpture	2
	-Pronotum at most slightly laterally impressed subapically behind eye, not distinctly incised; in dorsal view, lateral margin more or less continuous to apex, at most slightly constricted subapically or rarely subemarginate; pronotum and elytra generally lacking coarse sculpture	5

2	Elytra smooth and even, lacking "wart-like" swellings, striae and intervals uniform, not distorted by swellings; elytra very slightly transversely impressed at anterior and posterior one-thirds but not deeply plicate; color black with elytral intervals 3 and 9 orange throughout most of their lengths	C. lineatus Anderson
	-Elytra with moderate to large, "wart-like" swellings which distort striae and intervals; elytra distinctly, deeply transversely plicate at anterior and posterior one-thirds; color uniformly black or elytra black with few red spots near base	3

3	Color black, elytra with distinct red spots at base of interval 3, on humerus and subapically on interval 9; Panama	C. morrisi Anderson
	-Color uniformly black; Costa Rica	4

4	Pronotum with subbasal impression weakly carinate; prosternum between coxae narrow, not tuberculate	C. verrucosus (Champion)
	-Pronotum with subbasal impression not carinate; prosternum between coxae wider, slightly tuberculate	C. gibberosus (Champion)

5	Outer apex of tibia bidentate; male femur with inner margin with distinct tooth	C. transatlanticus (Kirsch)
	-Outer apex of tibia rounded or obtusely angulate; male femur with inner margin lacking tooth, simple	6

6	Metasternum with round or conical protuberance between middle coxae	7
	-Metasternum feebly rounded or flat between middle coxae	13

7	Pronotum with three round, equidistant depressions; antennal club with pilose apical portion one-quarter or less length of entire club; elytra yellowish with black borders and spots	C. graphipterus (Champion)
	-Pronotum with one depression at middle of base; antennal club with pilose apical portion one-third or more length of entire club; elytra black, or black with red or orange bands or stripes	8

8	Metasternal protuberance rounded, not larger than bulge of coxa; mesepimeron with outer front corner turned upward or backward	C. pruinosus (Champion)
	-Metasternal protuberance conical, larger than bulge of coxa; mesepimeron with outer front corner flat	9

9	Rostrum densely, coarsely punctate laterally in basal one-half, sides in front of scrobe with hairy, elongate depression formed by coalescent punctures	C. viduus (Hustache)
	-Rostrum very finely punctate or impunctate laterally, sides in front of scrobe smooth	10

10	Rostrum with upper margin of scrobe straight, base not dilated over scrobes	C. aurofasciatus (Breme)
	-Rostrum with upper margin of scrobe with small angulation or sinuation, base dilated (at least slightly) over scrobes	11

11	Elytra black, with large black velvety patch on each elytron (feebly defined in some specimens); prosternum with round swelling in front of each coxa	C. carinipyga (Champion)
	-Elytra black, with 1 or 2 transverse orange or red bands; prosternum smooth in front of each coxa	12

12	Elytra with 2 red bands, one subbasal, the other at apical one-quarter; pronotum markedly sinuate at middle; scutellum distinctly emarginate anteriorly	C. duplocinctus (Champion)
	-Elytra with a single red basal band; pronotum very slightly sinuate at middle; scutellum slightly concave anteriorly	C. riesenorum Anderson

13	Antenna with pilose apical portion very small, less than one-fourth or one-fifth length of club	14
	-Antenna with pilose apical portion one-third or more length of club	16

14	Hind tibia broad, flat and "paddle-like", width at middle about one-third length	C. silron Anderson
	-Hind tibia typically slender and elongate, width at middle much less than one-third length	15

15	Pronotum with basal depression and two or more additional depressions; antennal club symmetical in profile	C. aurocinctus (Champion)
	-Pronotum with single shallow basal depression or no depression; antennal club oblique in profile, one side longer than the other.	C. strigosus (Erichson)

16	Rostrum with ventral surface with long or short hairs	17
	-Rostrum with ventral surface glabrous	18

17	Rostrum with ventral hairs sparse, tiny, basal; pronotum and elytra with velvety black areas; elytral apices truncate	C. miniatopunctatus Chevrolat (in part)
	-Rostrum with ventral hairs dense, long, in two distinct rows; pronotum and elytra without velvety black areas but with tumid orange spots; elytral apices separately rounded	C. personatus (Vaurie)

18	Elytra black, with single transverse red band at extreme base (band may be incomplete at middle or divided at sides)	19
	-Elytra without transverse red band or if present, not at base, or with other pattern of red markings	21

19	Elytral intervals at base flat or only feebly swollen	C. mesomelas (Champion)
	-Elytral intervals at base abruptly bulbous	20

20	Elytra with one red band; fourth interval bulbous at base and wider than others; pronotum black	C. sanguinolentus (Olivier)
	-Elytra with two red bands; fourth and fifth intervals bulbous at base but fourth not wider than others; pronotum with some red in most specimens	C. rubrovariegatus Bovie

21	Mesosternal process at apex distinctly narrower than one-half diameter of coxa	22
	-Mesosternal process at apex equal to or greater than one-half diameter of coxa	26

22	Elytral striae uniform, straight, not distorted by orange maculations (if maculations present); antennal scrobe with posterior edge separated from eye by no more than twice width of base of scape; inner face of front coxa lacking tubercles	23
	-Elytral striae variously redirected and distorted by swellings at orange maculations (if maculations present); antennal scrobe with posterior edge distant from eye by nearly length of antennal club or more; inner face of front coxa of male with inward-pointed tubercle	24

23	Rostrum immediately in front of eye with deep, vertical groove (formed by coalescent punctures); pygidium with dense long hairs in apical one-half, along lateral margins and along midline; form slender, pronotum longer than wide	C. lingorum Anderson
	-Rostrum immediately in front of eye with large, deep punctures; pygidium (except for apical fringe) with sparse, short hairs throughout; form more robust, pronotum about as long as wide	C. circumdatus (Champion), (in part)

24	Pronotum and basal portion of elytra coarsely, irregularly sculptured, rugulose, deeply coarsely punctate; pronotum with single transverse subbasal impression and two paramedian shallow, irregular impressions near midlength	C. dragoni Anderson, new species
	-Pronotum and elytra more or less smooth, finely evenly punctate; pronotum with only single transverse subbasal impression	25

25	Elytra each with colored subapical spot on third interval elongate and nearly extended to apex; humeral spot longer than adjacent spot	C. sinuatus (Champion)
	-Elytra each with colored subapical spot on third interval (if present), nearly round, situated far from apex; humeral spot short, not larger than others	C. ornatus (Champion)

26	Rostrum short, lower margin only three or four times longer than wide	27
	-Rostrum longer, lower margin at least five times longer than wide	28

27	Middle coxae separated by width of coxa; front coxae separated by width of antennal club; antennal club with apical pilose portion about one-third length of club	C. pulcherrimus (Champion)
	-Middle coxae separated by about one-half to two-thirds width of coxa; front coxae separated by less than width of antennal club; antennal club with apical pilose portion about one-half length of club	C. annulatus (Champion)

28	Elytra with punctures of striae large, fovea-like, deeply impressed and "pit-like", intervals and pronotum impunctate	C. fahraei (Gyllenhal)
	-Elytra with punctures of striae small to moderate, shallowly to moderately impressed, intervals finely punctate or impunctate; pronotum with punctures large, coarse and deep to small, shallow and fine	29

29	Punctures of elytral striae moderate in size, distinct; intervals finely but distinctly punctate; pronotum with distinct, moderately large deep punctures	30
	-Punctures of elytral striae small if present, not distinct; intervals impunctate; pronotum at most finely punctate	31

30	Elytra with bases of intervals 3-5 slightly swollen, interval 5 noticeably wider at base than throughout rest of length; pronotum with punctures large, coarse and deep, some coalescent; pygidium subcarinate towards apex	C. gasbarrinorum Anderson
	-Elytra with bases of intervals 3-5 flat, more or less of same width throughout lengths; pronotum with punctures moderate, individually distinct, not coalescent; pygidium evenly convex towards apex, not at all carinate	C. validirostris (Gyllenhal)

31	Pronotum with subbasal impression indistinct or absent; pronotum red or orange with black paramedian stripes; body size less than 8.0 mm	C. condylus (Vaurie)
	-Pronotum with subbasal impression distinct and moderately deep; pronotum black with lateral oblique red or orange stripe or maculation, or with no red or orange markings; body size greater than 8.0 mm	32

32	Pronotum with distinct red or orange lateral stripes that continue posteriorly onto elytra	33
	-Pronotum entirely black or black with pale red marks medially, not continuing posteriorly onto elytra	34

33	Prosternum between coxae with distinct tubercle (males) or swelling (females); front tarsus with article 3 almost fully pilose ventrally, with at most a fine median, basal glabrous line; pronotal red or orange stripes narrow, many times longer than wide; pygidium subcarinate	C. circumjectus (Champion)
	-Prosternum flat between coxae; front tarsus with article 3 with broad, triangular, basal glabrous area; pronotal red or orange stripes broad, about three times longer than wide; pygidium slightly, evenly convex	C. circumdatus (Champion), (in part)

34	Front coxae separated by slightly more than one-half width of coxa; base of elytral interval 4 swollen and directed slightly medially	C. sunatoriorum Anderson
	-Front coxae separated by at most about one-third width of coxa; base of elytral interval 4 flat or slightly convex, directed anteriorly	35

35	Male with prosternum between front coxae with distinct tubercle; pygidium with subapical swelling, apex subtruncate in lateral view; elytra with intervals 3-5 lacking series of orange dots at basal one-third	C. rectistriatus (Champion)
	-Male with prosternum flat between front coxae; pygidium at most slightly evenly convex, rounded in lateral view; elytra with row of red or orange dots on intervals 3-5 at basal one-third	C. miniatopunctatus Chevrolat, (in part)

Cactophagus annulatus (Champion)

Distribution: Costa Rica, Panama; also in South America.

Natural history: Numerous adults have been collected from epiphytic arums (Araceae) in the vicinity of Monteverde and Santa Elena. Vaurie (1967b) states that specimens have also been found on leaves of young palms.

Cactophagus aurocinctus (Champion)

Distribution: Costa Rica, Honduras, Mexico, Nicaragua and Panama.

Natural history: Vaurie (1967b) notes a single adult was collected from a Cynoches (Orchidaceae) pseudobulb.

Cactophagus aurofasciatus (Breme)

Distribution: Mexico, Panama; also in South America.

Natural history: No significant information.

Cactophagus carinipyga (Champion)

Distribution: Costa Rica, Honduras, Mexico and Nicaragua; also in South America.

Natural history: No significant information.

Cactophagus circumdatus (Champion)

Distribution: Costa Rica, Nicaragua and Panama; also in South America.

Natural history: No significant information.

Cactophagus circumjectus (Champion)

Distribution: Costa Rica and Panama; also in South America.

Natural history: Vaurie (1967b) notes a single individual associated with a palm. Adults have been collected on Araceae.

Cactophagus condylus (Vaurie)

Distribution: Costa Rica and Panama.

Natural history: No significant information.

Cactophagus dragoni Anderson

Distribution: Panama.

Natural history: Adults have been collected in montane cloud forest in the La Fortuna area of Chiriqui province. One specimen was collected in the axil of a large arum plant (Araceae).

Cactophagus duplocinctus (Champion)

Distribution: Costa Rica, Guatemala, Mexico, Nicaragua and Panama; also in South America.

Natural history: No significant information.

Cactophagus fahraei (Gyllenhal)

Distribution: Costa Rica, El Salvador, Guatemala, Honduras, Mexico and Nicaragua; also in South America.

Natural history: Vaurie (1967b) notes that this species develops in Opuntia and Cereus (Cactaceae).

Cactophagus gasbarrinorum Anderson

Distribution: Panama.

Natural history: The only known specimens were collected near 1300m in the vicinty of the La Fortuna Dam in Chiriqui province. The habitat along the Continental Divide Trail and surrounding areas is a wet cloud forest.

Cactophagus gibberosus (Champion)

Distribution: Costa Rica.

Natural history: No significant information.

Cactophagus graphipterus (Champion)

Distribution: Costa Rica, Guatemala, Mexico and Panama; also in South America and introduced into the United States.

Natural history: Vaurie (1967b) reports this species is associated with orchids. Specimens have been collected from Cattleya, Cycnoches, Lycaste, Odontoglossum, Oncidium, and other species having large, soft bulbs. Larval feeding and pupation takes place in the bulbs. Adults feed on leaves and other parts of the plants.

Cactophagus lineatus Anderson

Distribution: Costa Rica.

Natural history: No significant information.

Cactophagus lingorum Anderson

Distribution: Costa Rica.

Natural history: Adult specimens were collected at 700-950 m in wet premontane evergreen forest, and at 1520 m, likely in upper montane evergreen forest or perhaps cloud forest. A single specimen was beaten from the roots and stems of epiphytic Araceae. A tentatively conspecific specimen from La Amistad Pacifico was collected at 2400 m.

Cactophagus mesomelas (Champion)

Distribution: Costa Rica, Mexico, Nicaragua and Panama; also in South America.

Natural history: No significant information.

Cactophagus miniatopunctatus Chevrolat

Distribution: Belize, Costa Rica, Guatemala, Mexico, Honduras and Nicaragua; also in South America.

Natural history: Vaurie (1967b) notes that adults have been collected on flowers of Araceae and on palms. Some specimens intercepted in quarantine were associated with pineapples and bananas.

Cactophagus morrisi Anderson

Distribution: Panama.

Natural history: Specimens were collected along the Continental Divide Trail in the vicinity of La Fortuna in Chiriqui province. The habitat along the trail is a wet cloud forest.

Cactophagus ornatus (Champion)

Distribution: Costa Rica, Nicaragua and Panama; also in South America.

Natural history: No significant information.

Cactophagus personatus (Vaurie)

Distribution: Costa Rica and Panama.

Natural history: No significant information.

Cactophagus pruinosus (Champion)

Distribution: Costa Rica and Panama; also in South America.

Natural history: No significant information.

Cactophagus pulcherrimus (Chevrolat)

Distribution: Costa Rica, Mexico and Panama.

Natural history: Vaurie (1967b) notes a single specimen found inside a stem of Anthurium (Araceae) and adults have been found copulating on unidentified Araceae.

Cactophagus rectistriatus (Champion)

Distribution: Costa Rica, Nicaragua and Panama; also in South America.

Natural history: No significant information.

Cactophagus riesenorum Anderson

Distribution: Costa Rica.

Natural history: No significant information.

Cactophagus rubrovariegatus Bovie

Distribution: Costa Rica and Panama; also in South America.

Natural history: No significant information.

Cactophagus sanguinolentus (Olivier)

Distribution: Belize, Costa Rica, Guatemala, Honduras, Mexico, Nicaragua and Panama; also in South America.

Natural history: Vaurie (1967b) notes an incidental capture of a specimen on a palm leaf.

Cactophagus silron Anderson

Distribution: Costa Rica.

Natural history: No significant information.

Cactophagus sinuatus (Champion)

Distribution: Costa Rica and Panama.

Natural history: No significant information.

Cactophagus strigosus (Erichson)

Distribution: Panama; also in South America.

Natural history: No significant information.

Cactophagus sunatoriorum Anderson

Distribution: Panama.

Natural history: Three of the four known specimens were collected in wet cloud forest at 1300m on the Continental Divide trail in the La Fortuna area of Chiriqui province.

Cactophagus transatlanticus (Kirsch)

Distribution: Costa Rica; also in South America.

Natural history: Vaurie (1967b) records some specimens collected from bromeliads.

Cactophagus validirostris (Gyllenhal)

Distribution: Costa Rica, Guatemala, Honduras, Mexico and Panama; also in South America.

Natural history: Vaurie (1967b) reports specimens associated with bananas and Cinchona (Rubiaceae) as well as the collection of adults from Philodendron pertusum and P. hastatum (Araceae).

Cactophagus verrucosus (Champion)

Distribution: Costa Rica.

Natural history: Numerous adults have been collected from the Monteverde area on the stems and roots of various Araceae.

Cactophagus viduus (Hustache)

Distribution: Costa Rica, Nicaragua and Panama; also in South America.

Natural history: Vaurie (1967b) reports that a single specimen was reared from a stem of Philodendron.