Membracidae

Treehoppers. Most abundant in the Neotropics. Pronotum enlarged, often with horns, keels, or spines. All species phytophagous on trees, shrubs, or herbaceous plants; some species attended by ants.

Crazy Eyes

/ Ted C. MacRae / 70 Comments

Spissistilus festinus | Stoneville, Mississippi

Spissistilus festinus (three-cornered alfalfa hopper) is one of the few truly economic pests in the otherwise bizarre and innocuous family Membracidae (treehoppers).  Its common name alludes to one of the crops it affects, but my encounters with this species are most often in soybean (I am, after all, a soybean entomologist).  Damage in this crop is caused by both adults and nymphs, whose piercing/sucking mouthparts cause girdling and breakage of the stem—often just a few inches above the soil.  This individual was seen during my travels last week in a soybean field in Stoneville, Mississippi, where numbers throughout the season were especially high this year.  Although I have seen innumerable S. festinus adults, I have never noticed their crazy, zig-zag patterned red and white eyes until I managed this closeup face shot (click on photo for best view).

This slightly cropped photo was taken with a 100mm macro lens and full extension tube set, resulting in slightly more than 2X magnification.  One of the lessons I took from was the need to pay more attention to background and value contrast.  By placing the subject a few inches in front of the dark green soybean foliage I was able to achieve a much more pleasing background than the typical black background one gets with full flash photos at high magnification.  Although both the subject and the background are green, there is still sufficient difference in shade to create contrast between them.  Light-green is one of the more difficult colors to work with when full flash is used with high shutter speeds and small apertures to maximize crispness and detail (in this case, 1/250 sec and f/16).  However, increasing ISO to 400 and lowering flash exposure compensation to -2/3 can reduce the amount of flash needed to illuminate the subject with such settings, making it easier to achieve a properly exposed and true-colored subject.

Copyright © Ted C. MacRae 2011

BugShot 2011 – Lesson 2

/ Ted C. MacRae / 11 Comments

Most of my insect photography is done up close using fast shutter speeds (to prevent motion blur) and small apertures (to maximize depth of field).  This necessitates the use of full flash – the amount of light reaching the camera sensor at f/16 and 1/250 sec is not enough to show any image at all, much less one properly exposed.  Full flash photography has its own set of challenges, but for the most part it can be used to produce excellent closeup photographs of insects, even very small ones.  One thing that has always bothered me about full flash photography, however, is the “black background” effect when photographing an insect sitting up on a plant without something else in the immediate background to reflect light.  Not that I don’t like black backgrounds—they can be used to stunning effect with the right subject.  I just don’t want it to be my only option for insects that I photograph up off the ground. Of course, it is rather a simple matter to place something in the background that is close enough to reflect light but far enough away to remain out of focus, but what I really want to be able to do sometimes is have a blue sky.  I always thought this meant natural light, with its low f stops, slow shutter speeds, and the resulting motion blur and loss of depth of field. 

Of the many things I learned today, how to include a blue sky background in a closeup photograph at f/16 was my favorite.  This is accomplished by bumping up the ISO to 400 (to make the camera sensor more receptive to light) and decreasing the shutter speed to 1/60 sec.  Keeping the f stop high maintains the depth of field, but the increased ISO and decreased shutter speed allows sufficient light from a sky background to register on the sensor.  By themselves, however, these setting will still result in an underexposed subject, which is illuminated instead by fill flash.  Despite the slower shutter speed, there is no motion blur because the “effective” shutter speed for the subject is the duration of the flash pulse rather than the camera shutter speed—it’s like combining two exposure speeds in a single photograph, one for the background and another for the subject.

The following three photographs illustrate this principle—again, they are not technically perfect photos, but rather the result of quick experimentation to understand the principles involved.  Photo 1 is from yesterday’s post and illustrates what my typical settings have always been: ISO 160, f/16, and 1/250 sec.  It’s a decent photo of the treehopper, Acutalis tartarea; however, black is perhaps the least appropriate background to choose for this black species. Until now, it would have been my only option unless I tried arranging foliage in its background.  Photos 2 and 3 are of another individual of this species that I found today (fortunately in similar orientation to the individual photographed yesterday).  In both photos I kept the flash unit set to ETTL (adjusting FEC as appropriate for the shots).  In Photo 2 I bumped up the ISO to 400 but kept the shutter speed fast (1/200 sec)—you can see some effect in that the background is not truly black, having received some light from the blue sky.  It’s not enough, however, because the shutter speed was still too fast.  In Photo 3 the ISO remained at 400 but the shutter speed was also decreased to 1/60 sec.  The shutter staying open that long allows enough light from the sky to register on the sensor and, Voila!, we have a blue sky background that creates nice value contrast with the black subject.  The subject these photos is not terribly sharp, but that is just lack of focus—not motion blur from a slow shutter (sorry, I was just practicing settings rather than going for a perfect shot). All three photos were shot with the Canon 100mm macro lens + 68 mm of extension tube (total magnification ~2X).

''Typical'' insect macro settings: ISO 160, 1/250 sec, f/16

ISO increased to 400 (1/200 sec, f/16)

Shutter speed decreased to 1/60 (ISO 400, f/16)


Copyright © Ted C. MacRae 2011

BugShot 2011 – Lesson 1

/ Ted C. MacRae / 7 Comments

I’m a lucky guy! First, I’m one of the fortunate attending this weekend’s BugShot insect photography workshop. Second, this first-of-a-kind event is being held only 13 miles from my home at Shaw Nature Reserve in Gray Summit, Missouri. Third, I was “adopted” by the BugShot instructors to assist in the event. Who are the instructors? None other than John Abbott from Austin, Texas—an expert on dragonfly biology and insect action photography, Thomas Shahan from Norman, Oklahoma—master of close-up arthropod (especially jumping spider) portraiture, and Alex Wild from Champaign-Urbana, Illinois—ant photographer extraordinaire and author of the most popular insect blog on the net.  In 2009 I picked up a digital SLR camera for the first time ever—in 2011 I am rubbing shoulders, discussing exposure and lighting, and enjoying social time with three of the country’s most accomplished insect macrophotographers (and some other very cool people as well).

I have come a long way, but I still have much to learn.  Intimate understanding of lighting, exposure, and the creative use of flash still eludes me—I can do a few things well, but there is much more I can’t do at all.  Today was my first time experimenting with the effect of lighting direction, i.e. taking the flash heads off their fixed position on the front of the lens and hand-holding them in different positions.  This simple technique can have dramatic effects on the look of a photograph, as illustrated by the following two photographs.  In the first, both flash heads of my Canon MT-24EX twin flash are attached to the front of the lens (as they have been for every single flash photograph I’ve ever taken up to this point).  In the second, only the right flash head remains attached to the lens, while the left head has been detached and is being hand-held directly above the subject (in this case, the treehopper Acutalis tartarea on Solidago sp.).  Technically they are not very good photographs, but they illustrate well the dramatic differences that can be achieved by varying the position of the flash heads.  Among other things, this is a technique that I will be exploring much over the coming weeks.

Both flash heads attached to front of lens.

Right flash head attached to front of lens; left flash head held directly above subject.

Copyright © Ted C. MacRae 2011

Bichos Argentinos #9 – Membracido

/ Ted C. MacRae / 3 Comments

Enchenopa? sp. | Buenos Aires, Argentina

This treehopper that I photographed at La Reserva Ecológica Costanera Sur strongly resembles our North American species of Campylenchia due to the brown elytra and lack of any yellow markings on the pronotal crest.  However, the rounded lower margin of the frons (more apparent in the full-sized version of this photo) eliminates this genus as a possibility and suggests instead the closely related Enchenopa. 

I sent this and another photo to Andy Hamilton (Canadian National Collection of Insects, Arachnids and Nematodes) for his opinion.  Andy claims to be a hack when it comes to Neotropical Membracidae (focusing more on world Cercopidae and Holarctic Cicadellidae), but he is a much better hack than I!  In his reply, he mentions that a lot of work is still needed on tropical species and genera, and in fact none of our North American species of Enchenopa actually resemble the type-species from Brazil (Membracis monoceros).  Most of what we now consider Enchenopa will likely be referable back to the genus Membracis (type genus of the family), but where the species in the above photo will eventually fall remains anyone’s guess.

Copyright © Ted C. MacRae 2011

Brazil Bugs #15 – Formiga-membracídeos mutualismo

/ Ted C. MacRae / 50 Comments

Of the several insect groups that I most wanted to see and photograph during my trip to Brazil a few weeks ago, treehoppers were near the top of the list.  To say that treehoppers are diverse in the Neotropics is certainly an understatement – South America boasts an extraordinary number of bizarre and beautiful forms that still, to this day, leave evolutionary biologists scratching their heads.  The development of this amazing diversity is a relatively recent phenomenon (thinking geological scale here), as there are no known membracid fossils prior to Oligo-Miocene Dominican and Mexican amber – well after the early Cretaceous breakup of Gondwanaland split the globe into the “Old” and “New” Worlds.  With its origins apparently in South America, numerous groups continued to spring forth – each with more ridiculous pronotal modifications than the last and giving rise to the dazzling diversity of forms we see today.  Even North America got in the evolutionary act, benefiting from northern dispersal from South America’s richly developing fauna via temporary land bridges or island stepping stones that have existed at various times during the current era and giving rise to the almost exclusively Nearctic tribe Smiliini (whose species are largely associated with the continent’s eastern hardwood forests).  Only the subfamily Centrotinae, with its relatively unadorned pronotum, managed to successfully disperse to the Old World, where it remains the sole representative taxon in that hemisphere.  With a few notable exceptions, treehoppers have virtually no economic importance whatsoever, yet they enjoy relatively active study by taxonomists, evolutionists, and ecologists alike – due almost completely to the bizarreness of their forms and unique mutualistic/subsocial behaviors.

I did manage to find a few species of treehoppers during the trip (a very primitive species being featured in Answer to ID Challenge #4 – Aetalion reticulatum), and of those that I did find the nymphs in this ant-tended aggregation on a small tree in the rural outskirts of Campinas (São Paulo State) were perhaps the most striking in coloration and form.  Most were jet black, although a few exhibited fair amounts of reddish coloration, and all exhibited sharply defined white bands of wax and long erect processes on the pronotum, mesonotum, and abdomen.  I’ve seen a fair number of treehopper nymphs, but I did not recognize these as something I had seen before, and given the incomplete state of immature taxomony I feared an identification might not be possible.  Still (and I know this is probably beginning to sound like a broken record), I gave it the old college try.

I usually like to start simple and get more creative if the results aren’t satisfactory, so I went to my old friend Flickr and simply typed “Membracidae” as my search term.  Predictably, pages and pages of results appeared, and I began scanning through them to see if any contained nymphs at all resembling what I had.  After just a few pages, I encountered this photo with very similar-looking nymphs, and although no identification beyond family was indicated for the photo, I recognized the lone adult sitting with the nymphs as a member of the tribe Aconophorini – a diverse group distinguished from other treehoppers by their long, forward-projecting pronotal horn.  Luck was with me, because I happen to have a copy of the relatively recent revision of this tribe by Dietrich and Deitz (1991).  Scanning through the work, I learned that the tribe is comprised of 51 species assigned to three genera: Guayaquila (22 spp.), Calloconophora (16 spp.), and Aconophora (13 spp.).  The latter two genera can immediately be dismissed, as ant-interactions have not been recorded for any of the species in those two genera – clearly the individuals that I photographed were being tended by ants.  Further, the long, laterally directed apical processes of the pronotal horn, two pairs of abdominal spines, and other features also agree with the characters given for nymphs of the genus Guayaquila.  In looking at the species included in the genus, a drawing of a nymph that looked strikingly similar to mine was found in the species treatment for G. gracilicornis.  While that species is recorded only from Central America and northern South America, it was noted that nymphs of this species closely resemble those of the much more widely distributed G. xiphias, differing by their generally paler coloration.  My individuals are anything but pale, and reading through the description of the late-instar nymph of the latter species found every character in agreement.  A quick search of the species in Google Images was all that was needed to confirm the ID (at least to my satisfaction). 

In a study of aggregations of G. xiphias on the shrub Didymopanax vinosum (Araliaceae) in southeastern Brazil, Del-Claro and Oliveira (1999) found an astounding 21 species of associated ant species – a far greater diversity than that reported for any other ant-treehopper system.  The most frequently encountered ant species were Ectatomma edentatum, Camponotus rufipes, C. crassus, and C. renggeri, and after perusing the images of these four species at AntWeb I’m inclined to believe that the ants in these photos represent Camponotus crassus (although I am less confident of this ID than the treehoppers – corrections welcome!).  The authors noted turnover of ant species throughout the day in a significant portion of the treehopper aggregations that they observed, which they suggest probably reflects distinct humidity and temperature tolerances among the different ant species and that might serve to reduce interspecific competition among ants at treehopper aggregations.  Since treehopper predation and parasitism in the absence of ant mutualists can be severe, the development of multispecies associations by G. xiphias results in nearly “round-the-clock” protection that can greatly enhance their survival.

Update 3/3/11, 9:45 a.m.:  My thanks to Chris Dietrich at the Illinois Natural History Survey, who provided me in an email exchange some clarifying comments on the origins and subsequent dispersal of the family.  The first paragraph has been slightly modified to reflect those comments.

REFERENCES:

Del-Claro, K. and P. S. Oliveira. 1999. Ant-Homoptera interactions in a Neotropicai savanna: The honeydew-producing treehopper, Guayaquila xiphias (Membracidae), and its associated ant fauna on Didymopanax vinosum (Araliaceae). Biotropica 31(1):135–144.

Dietrich, C. H. and L. L. Deitz.  1991.  Revision of the Neotropical treehopper tribe Aconophorini (Homoptera: Membracidae).  North Carolina Agricultural Research Service Technical Bulletin 293, 134 pp.

Copyright © Ted C. MacRae 2011

Bizarre, beautiful extremes

/ Ted C. MacRae / 8 Comments

No niche, it seems goes unfilled. Specialization is likely to be pushed to bizarre, beautiful extremes.–E. O. Wilson, The Diversity of Life

Wilson didn’t mention treehoppers specifically when he made the above quote, referring to the exuberance of extreme behavioral and morphological adaptations seen in the biota of the tropics, but he could have just as easily led off with them.  Treehoppers (order Hemiptera, family Membracidae) are well-known for their variety of oddly grotesque shapes resulting from a curiously inflated pronotum – presumably having evolved to resemble thorns and buds on their host plants, or the ants that vigorously defend numerous treehopper species in exchange for their sweet honeydew, or perhaps to aid in the dispersal of volatile sex pheromones (an attractive hypothesis but lacking experimental support). Despite inordinate attention in relation to their low economic importance, it remains that the pronotal modifications of many treehoppers are so bizarre that they continue to defy any logical explanation.

I must admit that, despite my passion for beetles, treehoppers were my first love.  (Well, actually anything that I could bring home from my solo wanderings in the urban woodlands and vacant lots near my childhood home and keep alive in a terrarium was my first true love, but from an academic standpoint, treehoppers were the first group to arouse my taxonomic interest as I began my transformation from child collector to serious student.) I had just begun graduate school in the Enns Entomology Museum under the late hemipterist Tom Yonke to conduct leafhopper host preference and life history studies, and although far more Cornell drawers in the museum contained Cicadellidae, it was the treehopper drawers that I found myself rifling through each afternoon after completing the day’s thesis duties. Despite their lesser number, the treehopper drawers had recently benefited from the attentions of a previous student, Dennis Kopp, whose efforts during his time at the museum concentrated on collecting treehoppers from throughout Missouri and culminated in the four-part publication, The Treehoppers of Missouri (1973-1974). I was enamored by these little beasts – specifically by their exaggerated pronotum – and started collecting them whenever I could on my forays around the state surveying for leafhoppers. They were closely enough related to leafhoppers to make them relevant to my work, only cooler – like leafhoppers on steroids! With The Treehoppers of Missouri as my bible and my desk located a half dozen footsteps from the largest treehopper collection within a several hundred mile radius, I delved into their taxonomy and, for a time, considered a career as a professional membracid taxonomist.

Fast forward nearly 30 years, and my involvement as a taxonomist is neither professional nor deals with membracids. Beetles have taken over as my focal taxon, and I conduct these studies strictly as an avocation. Still, I continue to collect treehoppers as I encounter them, and although such efforts have been largely opportunistic, I’ve managed to assemble a fairly diverse little collection of these insects as a result of my broad travels. Much of this has occurred in the New World tropics, and it is this region that is the center of diversity for the family Membracidae (fossil evidence suggests that subfamily diversification and subsequent New World radiation began during Tertiary isolation about 65 million years ago after South America separated from Africa, since only the primitive subfamily Centrotinae occurs in both the Old and the New Worlds – all other subfamilies are restricted the New World (Wood 1993)).  Every now and then, as I accumulate enough material to fill a Schmidt box, I sit down and study what I’ve collected, comparing it to my meager literature to attempt identifications.  For material I collect in eastern North America, this works fairly well, as there have been a number of publications covering different parts of this area.  Outside of this area, however, my only hope is to entice one of the few existing membracid specialists into agreeing to look at what I’ve accumulated and ask for their help in providing names, in exchange for which they will be granted retention privileges to benefit their research.

idd-treehoppersMost recently, I was able to convince Illinois Natural History Survey entomologist Chris Dietrich to take a look at the material I had accumulated during the past ten years or so, which included many specimens from Mexico and a smattering from other world areas, including South Africa. Chris did his doctoral work at North Carolina State University under “Mr. Membracid” himself, Lewis Deitz, and has since been conducting evolutionary and phylogenetic studies on Membracidae and the related Cicadomorpha. I recently received this material back from Chris (photo above), the majority of which he had been able to identify to species – only a few specimens in the more problematic genera were left with a generic ID.

Oaxaca)

Campylocentrus sp. (Mexico: Oaxaca)

Oaxaca)

Hyphinoe obliqua (Mexico: Oaxaca)

Puebla)

Poppea setosa (Mexico: Puebla)

Oaxaca)

Umbonia reclinata (Mexico: Oaxaca)

Puebla)

Umbonia crassicornis male (Mexico: Puebla)

umbonia_crassicornis_female

Umbonia crassicornis female (Mexico: Puebla)

The selection of photos here show a sampling of some of the more interesting forms contained within this batch of newly identified material – all of which hail from southern Mexico. Campylocentrus sp. is an example of the primitive subfamily Centrotinae, distinguished among most membracid subfamilies by the exposed scutellum (not covered by the expanded pronotum).  Hyphinoe obliqua is an example of the largely Neotropical subfamily Darninae, while Poppea setosa represents one of the more bizarre ant-mimicking species of the subfamily Smiliinae.  Umbonia is a diverse genus in the subfamily Membracinae, occurring from the southern U.S. south into South America. Umbonia crassicornis is one of the most commonly encountered species in this genus, with the photos here showing the high degree of sexual dimorphism it exhibits.  As membracids go, these species are quite large (10 mm in length from frons to wing apex for Campylocentrus sp. and P. setosa, a slightly larger 11-13 mm for the others); however, the many smaller species in this family are no less extraordinarily ornamented.  I’ve also included a photo (below) of one of the drawers from the main collection after incorporating the newly identified material – this drawer represents about half of my treehopper collection, with the largely Nearctic tribe Smiliini and the primitive family Aetalionidae contained in another drawer. In all, the material contained one new subfamily, six new tribes, 13 new genera¹, and 30 new species for my collection. For those with an appetite for brutally technical text, a checklist of the species identified, arranged in my best attempt at their current higher classification, is appended below (any treehopper specialist who happens upon this should feel free to set the record straight on any errors). For each species, the country of origin (and state for U.S. specimens) is indicated along with the number of specimens, and higher taxa new to my collection are indicated with an asterisk(*). Don’t worry, I didn’t type this up just to post it here – it’s a cut/paste job from my newly updated collection inventory for Membracoidea. Happy reading!

¹Wildly off topic, and perhaps of interest only to me, but two of the genera represented in the material are homonyms of plant genera: Oxyrhachis is also a Madagascan genus of Poaceae, and Campylocentrus is a Neotropical genus of Orchidaceae. Scientific names of plants and animals are governed by separate ruling bodies (ICBN and ICZN, respectively), neither of which specifically prohibit (but do recommend against) creating inter-code homonyms. The number of such homonyms is surprisingly high – almost 9,000 generic names have been used in both zoology and botany (13% of the total in botany) (source).  Fortunately, there is only one known case of plant/animal homonymy fr BOTH genus- and species-level names – Pieris napi japonica for a subspecies of the gray-veined white butterfly (Pieridae) and Pieris japonica for the popular ornamental plant Japanese andromeda (Ericaceae).

treehopper_drawer
REFERENCES:

Kopp, D. D. and T. R. Yonke. 1973-1974. The treehoppers of Missouri: Parts 1-4. Journal of the Kansas Entomological Society 46(1):42-64; 46(3):375-421; 46(3):375-421; 47(1):80-130.

Wood, T. K. 1999. Diversity in the New World Membracidae. Annual Review of Entomology 38:409-435.
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Superfamily MEMBRACOIDEA
Family MEMBRACIDAE
Subfamily CENTROTINAE

 *Tribe BOOCERINI
*Campylocentrus curvidens (Fairmaire) [Mexico] – 4
Campylocentrus sp. [Mexico] – 1

*Tribe GARGARINI
*Umfilianus declivus Distant [South Africa] – 3

*Tribe OXYRHACHINI
*Oxyrhachis latipes (Buckton) [South Africa] – 1

Tribe PLATYCENTRINI
Platycentrus acuticornis Stål [Mexico] – 11
Platycentrus obtusicornis Stål [Mexico] – 3
Platycentrus brevicornis Van Duzee [USA: California] – 7
Tylocentrus reticulatus Van Duzee [Mexico] – 4

*Tribe TERENTIINI
*Stalobelus sp. [South Africa] – 1

*Subfamily HETERONOTINAE

*Tribe HETERONOTINI
*Dysyncritus sp. [Argentina] – 1

Subfamily MEMBRACINAE

Tribe ACONOPHORINI
Aconophora sp. female [Mexico] – 1
*Guayaquila xiphias (Fabricius) [Argentina] – 7

Tribe HOPLOPHORIONINI
Platycotis vittata (Fabricius) [USA: Arizona, California] – 3
Umbonia crassicornis (Amyot & Serville) [Mexico] – 73
Umbonia reclinata (Germar) [Mexico] – 8

Tribe MEMBRACINI
Enchenopa binotata complex [Mexico] – 1
Enchenopa sp. [Argentina] – 6

Subfamily DARNINAE

Tribe DARNINI
Stictopelta nova Goding [Mexico] – 9
Stictopelta marmorata Goding [USA: Texas] – 1
Stictopelta pulchella Ball [Mexico] – 11
Stictopelta varians Fowler [Mexico] – 3
Stictopelta sp. [USA: Arizona, California] – 5
Stictopelta sp. [Mexico] – 5
Stictopelta spp. [Argentina] – 6
*Sundarion apicalis (Germar) [Argentina] – 2

*Tribe HYPHINOINI
*Hyphenoe obliqua (Walker) [Mexico] – 1

Subfamily SMILIINAE

Tribe AMASTRINI
Vanduzeea triguttata (Burmeister) [USA: Arizona] – 2

Tribe CERESINI
Ceresa nigripectus Remes-Lenicov [Argentina] – 3
Ceresa piramidatis Remes-Lenicov [Argentina] – 4
Ceresa ustulata Fairmaire [Argentina] – 1
Ceresa sp. female [Argentina] – 1
Poppea setosa Fowler [Mexico] – 11
Tortistilus sp. [USA: California] – 1

Tribe POLYGLYPTINI
*Bilimekia styliformis Fowler [Mexico] – 3
Polyglypta costata Burmeister [Mexico] – 18

Tribe SMILIINI
Cyrtolobus acutus Van Duzee [USA: New Mexico] – 1
Cyrtolobus fuscipennis Van Duzee [USA: North Carolina] – 1
Cyrtolobus pallidifrontis Emmons [USA: North Carolina] – 1
Cyrtolobus vanduzei Goding [USA: California] – 4
Cyrtolobus sp. [USA: Arizona] – 2
*Evashmeadea carinata Stål [USA: Arizona] – 4
*Grandolobus grandis (Van Duzee) [USA: Arizona] – 1
Ophiderma sp. [Mexico] – 1
Palonica portola Ball [USA: California] – 4
Telamona decora Ball [USA: Missouri] – 4
Telamona sp. [USA: Texas] – 1
*Telamonanthe rileyi Goding [USA: Texas] – 2
*Telonaca alta Funkhouser [USA: Texas] – 1
Xantholobus sp. [Mexico] – 1

Subfamily STEGASPINAE

Tribe MICROCENTRINI
Microcentrus perditus (Amyot & Serville) [USA: Texas] – 1
Microcentrus proximus (Fowler) [Mexico] – 1

Family AETALIONIDAE
Subfamily AETALIONINAE

Aetalion nervosopunctatum nervosopunctatum Signoret [Mexico] – 9
Aetalion nervosopunctatum minor Fowler [USA: Arizona] – 2
Aetalion reticulatum (Linnaeus) [Argentina, Uruguay] – 26

Copyright © Ted C. MacRae 2009

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