Agrilus lucindae Hespenheide, 2010

Hespenheide, Henry A., 2010, New Agrilus Curtis (Coleoptera: Buprestidae) from México and Costa Rica mimicking parasitic wasps, Zootaxa 2545, pp. 39-46 : 43-45

Agrilus lucindae Hespenheide , New Species

( Figs. 8, 9 )

Description. Holotype female: Slender, in cross-section flattened above, convex below, 5.85 mm long, 1.45 mm wide; head, tibiae and beneath black; pronotum with midline and lateral margins black, otherwise dark reddish-purple; elytra with anterior1/2 metallic red, 1/4 beyond middle black, and posterior 1/4 golden; femora and basal 4 antennomeres green; complex pattern of setae: pale, recumbent, and relatively inconspicuous on lower 1/2 of front, along midline and margins of pronotum and on pronotum beneath, on elytra in oval spots at basal 1/4 just interior to suture, and in broad, somewhat golden oblique band at middle from suture to lateral margins; setae longer and denser in broad transverse band at apical 1/4 of elytra, in a broad oblique patch from middle coxae to upper posterior angles of metasternum, on posterior coxae, on posterior 1/3 of dorsal portion of abdominal ventrite 1, and on lateral portions of abdominal ventrite 3.

Head with front convex, somewhat swollen and bulbous with slight bluish reflections above eyes, with moderate depression along midline, broader and deeper above antennal insertions, surface transversely rugose with slight golden reflections between eyes; epistoma with transverse ridge between antennal insertions, width 1/3 of distance between inner margins of eyes, ventral margin shallowly emarginate, eyes small, oval; antennae long, slender, serrate from antennomere 5, antennomere 5 triangular, 2/5 wide as long, segments 6– 11 sharply triangular.

Pronotum slightly narrower than elytra at posterior margin, with sides weakly expanded outward nearly to apex, then narrowing slightly; when viewed from side marginal and submarginal carinae slightly undulate, narrowly separated for anterior 5/6; from above, anterior margin produced as narrowly rounded-angulate lobe; basal margin shallowly angulately emarginate at middle of each elytron, slightly emarginate before scutellum; disk convex in cross section, with distinct depression along midline and distinct narrow depressions at posterior angles and along lateral margins; prehumeral carinae absent; disc distinctly rugose. Scutellum rounded in front, acuminate behind with strong raised transverse carina.

Elytra subequal in width at humeri and at apical 2/3, lateral margins slightly emarginate between, then narrowing to broadly and obliquely rounded-truncate apices; disk weakly costate on basal 1/2, nearly flat on apical 1/2, each elytron with small, strong oval depression at base; surface coarsely imbricate on basal 1/2, more weakly so on apical 1/2; dorsal portions of ventrites 1–2 visible from above.

Prosternum with sides of prosternal process narrowing between coxae to acute apex; prosternal lobe nearly transverse. Posterior coxae with posterior margins nearly straight, upper exterior angles obtuse. Abdomen with suture between ventrites 1 and 2 obsolete, dorsal portions of ventrites 1–5 narrow, slightly broader at posterior portion of ventrite 1. Hind legs with first metatarsal segment equal in length to next three combined, tarsal claws cleft with inner portion shorter and blunter on all tarsi.

Type material. Holotype: Costa Rica: Prov. Heredia, F. La Selva, 3 km S Pto. Viejo, 10o 26’ N 84o 01’ W, 11-iii-1980, H.A. Hespenheide, [at] Justicia ( INBC). Paratype: Prov. Cartago, Guayabo Nat. Mont., ~ 1100 m, 0 9o 58’14”N; 83o41’24”W, 3.6–7.2008, Cate, Barries, Nagy ( INBC); Heredia Pr., La Selva Biol. Sta., 3 km S Pto. Viejo, 10 o 26’ N 84o 01’ W, 18-iv-1988, H.A. Hespenheide, ( CHAH).

Host. This species was collected as an adult on Justicia aurea Schltdl. (Acanthaceae) .

Etymology. This species is named in honor of Lucinda McDade who has studied Justicia aurea ( McDade & Kinsman 1980) and other Acanthaceae at La Selva and elsewhere and was lead editor of a volume reviewing research at La Selva ( McDade et al. 1994).

Discussion. In addition to general differences in size and color, Agrilus braconoides has the marginal and submarginal carinae of the pronotum abruptly and narrowly separated only for the anterior 1/2; a coppery-red metasternum and reddish reflections on the abdomen; a golden epistoma; and more elongate antennae with outer antennomeres narrowly rounded. A. lucindae differs in having the marginal and submarginal carinae of the pronotum narrowly separated for anterior 5/6 and gradually convergent toward the base; metasternum, abdomen; and epistoma black; and somewhat shorter antennae with the outer antennomeres sharply acute. The paratypes measure 6.90 and 7.00 mm in length.

As noted above under A. bellamyi , the mimetic Agrilus described here belong to two very different species groups, the Mexican A. braconicoloratus and A. bellamyi and the Costa Rican A. lucindae and A. braconoides . Their superficial resemblance to each other is another example of what I have termed mimetic homoplasy ( Hespenheide 2005). The relationships of these four species with non-mimetic Agrilus remain to be determined.

Mimicry of braconids by Agrilus . I previously proposed ( Hespenheide 1989) that Agrilus braconicoloratus might be mimicking braconids, including species in the genus Atanycolus ( Figure 11 ), which are parasitoids of wood-boring beetles ( Quicke & Sharkey 1989), and models may include other Neotropical braconid genera as well as wasps in the family Ichneumonidae ( Gauld 1991) . Resemblance of other beetles and reduviid bugs to braconids and ichneumonids has been noted long ago as indicated by such specific epithets as “ braconiformis ” or “ braconoides ” and “ ichneumoniformis ” ( Isthmiade ichneumoniformis Bates, 1870 , in Clarke 2009; Epirhyssa braconoides Porter, 1978 ). The likelihood that braconids are distasteful has been discussed by Quicke (1986), although Gauld (1991) implies that ichneumonids may not be distasteful and would therefore be Batesian mimics of braconids. Leathers and Sharkey (2003) described mimicry of Central American braconids of the genus Alabagrus Enderlein by reduviids (also mentioned in Sharkey 1988). Although members of Alabagrus are parasitoids of Lepidoptera , other similarly colored braconids (probably in the genus Atanycolus ; Sharkey, personal communication) have been seen on tree falls in Panama and Costa Rica (reported in Opitz 2008) and are presumably parasitoids of wood-boring insects or their predators.

The following is a preliminary list of Central American species or genera that I or others have observed to share a braconid-like color pattern at La Selva Biological Station and/or in Panamá:

Coleoptera

Cleridae

Enoclerus spp.—Hespenheide collection

Epiphlaeus fundurufus Opitz—in association with a braconid and Enoclerus sp. ( Opitz 2008)

Megaphloeus marginipes (Chevr.) —Hespenheide collection Cerambycidae

Cosmisoma plumicornis (Drury) —Hespenheide collection

Isthmiade perpulchra Linsley (1961) Curculionidae - Baridinae

Pardisomus cleroides (Champion) ( Prena 2003) Mordellidae

undetermined species—Hespenheide collection Diptera

Tipulidae (all in Hespenheide collection, Byers, in litt.)

Hexatoma (Eriocera) braconoides (Enderlein)

Hexatoma (Eriocera) sp.

Stigmatomera amazonica Westwood

Stigmatomera (Stigmatomera) sp. Heteroptera

Reduviidae

Hiranetis nr. braconiformis (Burmeister) ( Leathers and Sharkey 2003)

5 additional undetermined species— Hespenheide collection Hymenoptera

Ichneumonidae

Acrotophus sp. —Hespenheide collection Chrysididae

Cleptidea cf. panamensis Kimsey (1986)

Mimicry complexes that include braconids or ichneumonids and cerambycid beetles of the genus Isthmiade are also found in South America ( Clarke 2009), as well as Centrocerum exornatum (Newman) (see http://www.sel.barc.usda.gov/ Coleoptera /elaphid/Centroce.htm). Poulton (1935) reported the reduviid Hiranetis cingulatus Stål as a braconid mimic, and Elkins (1969) gives the names of several other genera of putative mimetic reduviids ( Xystonyttus , Neotropiconyttus , and Graptocleptes ). Mimicry complexes that include braconids are not restricted to the Neotropics; Quicke (1986) documents similar relationships between African braconids and cerambycids. Other taxa bearing the specific name “ braconoides ” include the Asian cercopid Callitettix braconoides (Walker) (Heteroptera; http://www.catalogueoflife.org/annual-checklist/ 2010/details/species/id/7487773), the Antillean trichopteran Chimarra braconoides Walker ( Philopotamidae, Flint 1998 ; http://insectdatabases.oeb.harvard.edu/Caribbean/Mantisweb/FMPro?-DB=Image.drd&- Lay=web&-Format=images_dr.htm&Species_ID=CMNH288385&-Find), and the South American mirids Monalonion braconoides Walker (= M. annulipes Signoret ; http://research.amnh.org/pbi/catalog/ names.php?b_id=3314) and Rayieria braconoides (Walker) (Heteroptera; Carvalho 1981).

Members of other hymenopteran families— e.g., mutillids and sawflies—share very similar patterns and are almost certainly Mullerian co-models by possessing strong stings or distastefulness. The functional roles of other taxa participating in these mimicry complexes—whether Mullerian co-models or Batesian mimicsremain to be determined; for example, the chrysidid Cleptidea panamensis is in the C. mutilloides (Ducke) species-group and may be a parasitoid of sawflies ( Kimsey 1986). A number of clerid beetles commonly share the color patterns discussed here (for example, see the image of the North American Thanasimus dubius (Fabr.) at http://www.forestryimages.org/browse/detail.cfm?imgnum=0013010), and beetles from other families have specific names such as “ cleroides ” and their role may be especially interesting. Although conoderine weevils participate in a number of mimicry complexes ( Hespenheide 1995), they don’t appear to participate in this one, presumably because they lack the elongate, narrow bodies of the models.

Figures

FIGURE 11. Atanycolus sp. on log, Madden Forest, Panamá Province, Panamá; photograph by author.

FIGURES 1 – 9. Wasp-mimicking Agrilus: A. braconicoloratus Hespenheide, figures 1 – 2, dorsal and lateral views, figure 3, male genitalia; A. bellamyi New Species, figures 4 – 5, dorsal and lateral views; A. braconoides New Species, figures 6 – 7, dorsal and lateral views; and A. lucindae New Species, figures 8 – 9, dorsal and lateral views. Scale bars = 1 mm.