Vol. 1, no. 1, January 1985 INSECTA MUNDI 29

A Generic Review of the Acanthaclisine Based on Larvae (: MYJ;ffieleontidae) 1

A 2 3 Lionel J..i. Stange and Robert B. Miller

IRTRODUCTIOR

The tribe Acanthaclisini Navas contains 14 (Rambur), whereas Steffan (1975) provides described genera which we recognize as additional data on this species as well as valid. We have reared larvae of 8 of these on Acantbaclisis occitanica (Villers). Our (Acantbaclisis Rambur, C_troclisis Nauas, best biological data on the Acanthaclisini, FadriDa Navas, Banks, Phano­ excluding larval behavior, are based on clisis Banks, Synclisis Navas, Syngenes observations of Paranthaclisis congener Kolbe, and Navas). In addition, we (Hagen) made near Reno, Nevada. In common have studied preserved larvae from Aus- with most aurJions, P. congener Jay eggs at tralia which probably represent the genus dusk. As the female expels the eggs, she Beoclisis Navas. Th~s represents the ma- evenly coats them with sand, using the pos jority of the taxa, lacking only the small terior gonapophysis. The eggs are shallowly genera Avia Navas, Cos ina Navas, Madrasta bUlled, in cOntlast to otheI known nOn Navas, Mestressa Navas, and Stipbroneuria acanthaclisine species which lay their eggs GelS taecke:I~ Studies of these laI vae have on the surface. Some females caught just revealed structural differences, especially after dusk still had egg material on the of the mandible, which we have employed to end of their abdomens where some had been provide ident i fie at ion of these genera by broken. Their abdomens appeared empty. means of descriptions, keys, and illustra Like most species with thick abdo­ tions. Also, since no modern key exists, mens, Parantbaclisis species lay eggs in we are providing a key to the genera based batches rather than a few every day. One on adults which will prOVide some further fema Ie captured at dusk had a plump abdomen insight on the generic relationships. Ob­ and layed 20 eggs which left her abdomen seruations on the tribal differences of empty The eggs were large and oblong and Myrmeleontidae based on larvae are made hatched ~n 24 days. L~ttle or no expans~on with a preliminary key to the known tribes. of the head capsule and mandibles takes place upon hatching ~n Parantbaclisis. Tribe AcauthacJisini Navas 1912 Th~s ~s unl~ke other tribes where upon hatching the head capsule and mandibLes InclUded genera: Acanthaclisis Ram- expand w~th the mandibles hanging like limp bur, Avia Navas (= Navas), spaghetti before expansion. Other char­ Navas (-SoBra Navas, -Neboda Navas, -Neo- acteristics ale similar to larvae ill other clisis Navas, =Stenoclisis Navas, =Sograssa tribes. There are three larval instars, Navas), Costns Navas, Beocltsts Navas, and diapause, if assumed, occurs in the Hadrasta Navas, Hestressa Navas, Parantba- larva in the sand and not in the cocoon nor citsis Banks, Pbanoclisis Banks (-Hola in the egg. The cocoon is constructed of Navas), Stipbroneuria Gerstaecker (=Neriga silk beneath the sand with sand grains Navas), SyDclisis Navas, Syngenes Kolbe covering the surface. The mobile pupa digs (-Onclus Navas), Vella Navas (=Cyroplectron its way to the surface of the sand where Esben PeteISen) [Note. The genus Vellassa the adult emerges and then climbs any con­ Navas is of uncertain status]. venient object before expanding its anten- nae, wings, and abdomen. Ceneral Biology: Principi (1947) de- Description (based on larvae): Three- tailed the biology of Synclisis baetica segmented labial palpus shorter than basal width of mandible; distal palpomere l5 to 1. Contribution No. 603, Bureau of Ento 3.0 t~mes longer than w~de; sensory open~ng mology, Div. Plant Industry, Fla. Dept. Agr1c. & CosnUmer Services t ;::~;~;;s~: ;i(~~n;:~Ci~;;s),itohr;(~:~::= Gainesville, FL 2. Taxonomic Entomologist, Div. Plant When 3, basal tooth shorter than distal Industry, P. O. Box 1269, GaineSVille, one; lateral margin with or without long ~ i2~02 setae, but setae never longer than one-half 3. Cesearch ASSOCiate, P. O. Box 1092, Project City, CA 96079. greatest width of mandible; mesothoracic spiracle not borne on tubelcle, scali ab 30 INSECTA ~mNDI Vol. 1, no. 1, January 1985

sent; dolichasters absent on head; abdomi­ and "uncati" (tibial spurs evenly curved). nal sternite VIII without submedian teeth; These 2 groups appear to be fairly natural stern~te IX w~thout enlarged pa~r of d~g­ d~vis~ons but since the names are not de­ ging setae j tergite IX without median lobe rived from included genera they are no.ina bear~ng med~an sc1erotized process. nuda. Perhaps when the larval stages of Larval Behavior: None of the larvae Avia, Cosina, Madrasta, Mestre8sa, and studied construct pltfall traps. Two dis­ Stipbroneuria are known, a logical sub­ tinct types of locomot~on are present. division of the tribe can be made. The two Backward movement only is found in the most highly evolved genera based on larval genera Phanoclisis and Vella. Forward and structure and behavior are Centrocllsls and backward movement are found in Acantha­ Paranthaclisis. Both appear to be select ~liaia, Ceatre~1181a, Fadriaa. Bee~llala. feedelS In the labolatolY and show re Paranthacliais. 8yngenea. and 8ynclisis. ductions in the mandibular teeth to 2 (Cen­ The latter three genera are fast runners Lroclisis) or I (Parantbaclisia). Also. and burrowers. Centroclisis larvae are both ~enera have highly reduced antennae slow creepers and diggers. Acauthac11s1a, Beoclisia, and Fadriua larvae are fast burrowers but seldom move forward rapid ly, at least under laboratory conditions. The behavior. Both genera are fast runners and larvae usually inhabit open tracts of sand burrowers. apparently favoring coastal where considerable sand depth is required beach dunes, having relatively long man- for temperature regulation, protection of dibles and with sternite IX relatively the.large cocoon. escape and concealment pointed and without peg-like setae on ster- from predators, as well as space for nite VIII (Figs. 8. 9). Finally. Phano­ hunting prey. Most species are incessant cl1s1s appears to be relatively plesio- feeders (except notably Centroclisis and morphic in many adult characters (distal Parauthaclisis) and require a fairly high palpomere, pretarsal claws) but its larva sand SUI face tempel atule to pupate. The moves only backward. a behav~ora1 trait cocoon is constructed in a single day and shared by the unrelated Vella. the period from construe t ion 0 f the cocoon Measurements: Measurements (in mm) of to the emergence of the adult is 54~3 days the head capsule were made from the ventral in all of the genera reared so far except surface. The longitudinal measurement was Fadriaa (35 days). ...11 of the genera made along the m~dline from the base of the studied. except Centroclisis, feed when the head capsule to the level of the mandible sand temperature is warm, but avoid extreme base. The width was measuled at the widest temperatures above or below safe levels by part of the head capsule. The mandible burrowing deeply in the sand It is common length was measuled (dOl sal side) from the to see them active on the surface in mid- base to the distal side of the distal morning. late afternoon, and very warm tooth. The width was measured at the nights. Centroclisis is the except ion as widest part of the mandible. The average the species studied is strictly nocturnal measurements are given with the number of Discus8ion: This tribe is well de­ specimens examined in parenthesis. fined by both larval and adult characters Illustrations' The photographs are and shows limited structural diversity in all of the 3rd ins tar larva except for compar~son to most tr~bes of antl~ons. Figure 8 (8yncUsis baetica) which is based Some group characters are present which on a 2nd instar larva. Figure]6 of Acan- suggest at least 4 subgroups: Group 1 ­ t:haclisis sp. shows a deformed middle tooth Centroclisis and Paranthaclisis; Group 2 - of the right mandible. The left mandible Syngenes and Synclisis, GIOUp 3 Acantha of Figure 16 and both mandibles of Figure clisis, Fadrina. and Phauoclisis; Group 4 - 20 show the normal condition of the man- Beoclisis and Vella. Navas (1912) divided dibular teeth. The photographs were taken the tribe into 2 groups; "uneinati" (adult by Robert B. M~l1er. tibial spurs bent at nearly right angle) Vol. 1, no. 1, January 1985 INSECTA MUNDI 31

Key to Genera of Acanthaclisini

LARVAE

1. Sternite VIII without short, blunt, peg-like setae although stout but point- ed dIggIng setae plesent (Figs. 8 11) 2 Sternite VIII with numerous short, peg-like setae (apex truncate) (Figs. 3 7) ....•...... •...... •...•.•...... 5 2. Ventral head capsule densely setose, except along midline (Fig. 21); Western Hemisphere ••••••••••••••••••••••••••••••••••••••••••••••••••••••••••• Vella Ventral surface of head capsule glabrous except laterally (Fig. 25) •••••• 3 3. Mandihle with prominent setae on interior margin of mandible between tooth 1 and mandibular base; sternite IX broadly rounded (Fig. 11); Australia ...... Beoclisis Mandible without setae between tooth 1 and mandibular base;sternite IX pointed (Figs. 12, 13) 4 4. Anterior margin of clypeal-labrum produced as a rounded lob (Fig. 13); Ethiopian •••••.••••••••.••••••••••.••••••.•••••••••••••••••••••••• Syngenes Anterior margin of clypeal-labrum emarginate {Fig. l2);Palearctic ...... Synclisis 5. Mandible with 1 or 2 teeth and without setae on the interior margin of mandible (Figs. 18, 19) ••••••••••••••••••••••••••..••.•••..•••••••••••••• 6 Mandl. lIle with :3 teeth and setae present on Inteliol malgin of mandib1e(Figs. 14-16) •••••••••••••••••••••..•••••••••.•••••••••••••••••••••••••••••••••• 7 6. Mandible with 1 tooth (Fig. 19); Nearctic ParanthaclisiB Mandible with 2 teeth (Fig. 18); Old World •••••••••••••••••••• Centroclisis 7. Tooth 3 of mandible 10ngel than middle tooth by at least 1.5 times(Fig. 14); Ethiopian •••••••••••••••••••••••••••••••••••••••••••••••••••••••••• Fadrina Tooth 3 of mandible shol ter than middle tooth (Figs. 15-16) S 8. Mandible with tooth 2 noticeably closer to tooth 3 than tooth 1 (Fig. 16); distance between mandibular teeth 1 and 3 greater than greatest width of mandible; ventral head capsule with thinly scattered setae;larva able to run forward ••••••••••••••••••••••••••••••••••••• ••• ••• ACautbaclisis Mandible with tooth 2 slightly closer to tooth 1 than 3 (Fig. 15); distance hetween mandihu1ar teeth equal to or slightly less than greatest widtb of mandih1e; ventral head capsule glabrous; larva moves backward on1y;Etbiopian ••.••..•••••••••••.••••••••••••••••••••••••••••••••••••••••••••• Phanoclisis

ADULTS

1, Tibial spnrs bent at nearly right angle. often with flange 2 Tibl.al spurs gently curved •••••••••••••••••••••••••••••••.••••••••••••••• 8 2 Rindfemllr witbout elongate sensory hair; costal area of forewing usually biareolate, upper series of cellules not much smaller than lower series ••• 3 Hindfemur with elongate sensory hair; costal alea simple, ol,if biareo1ate, then upper series of cellules much smaller than lower series ••••••••••••• 4 3. Midfemur with 1 elongate sensory hair; male ectoploct with long postventral lobe; terga V and VI without silvery pubescence; Ethiopian, Middle East •••• ••••••••••••••••.•••••••••••••••••••••••••••••••••••••••••. •• Syngenes Ko1he Midfemur with 2 elongate sensory hairs; male ectoproct short; terga V and VI

Forewin: cos ta 1 area predominate ly biareolate; ocu lar rim without setae

with long setae (much longer than width of scape); Old World .. •••••••••••••••••••••••••••••••••••••.••• • •••..•..... Centroclisis Navas Forefemur and midfemur with only 1 elongate sensory hal.r; ocular rim with very short setae (much shorter than width of scape); Nearctic . ••••••• •• •• ••••••••.•.•••. •• •••••••••••••• •• ••••••••• •• Paranthaclisis Banks 32 INSECTA MUNDI Vol. 1, no. 1, January 1985

1 Paranthaclisis sp. 2 Centroclisis punctulata

3 Centroclisis punctulata

6. Pretarssl claws slender,longer than tibial spurs; pronotum longer than wide; distal palpomere of labium with small, oval-shaped sensory opening;Ethiopian ...... •.•... . Ph4D.oclis18 Banks Pretarsal claws smaller than tibial SpuTs;pronotum as wide or wider than long; distal palpomere of labium with an elongate slit-like sensoryopenlng ...... 7 7. Tibial spur produced 8S flange just before IIbend"; hindfemur not abruptly swollen near base; male ectoproct with postventral lobe more than 3 times longer than wideiPalearctic ••••••.•.••••••••••••••.••• Acantbaclisia Rambur Tibial spur not expanded just before "bend" but much wider somewhat before this point; hindfemur abruptly swollen near base; male ectoproct with post- ventral lobe much broader than long; Ethiopian Fadrlna Navas 8. Hindfemur with elongate sensory hair 9 Hindfemur without elongate sensory hair 10 Vol. 1, no. 1, January 1985 INSECTA ~ruNDI 33

9. Hindwing posterior area at medial fork at least 1.5 times higher than great­ est presectoral height; hindwing presectoral area without accessory longi- tudinal vein; hindwing without large dark spots; Western Hemisphere •••••••••••••••••••••••••••••••••••••••••••••••••••••••••••••• •• Vella Navas Hindwing posterior area at medial fork no higher than greatest presectoral height,hindwing presectoral area with short accessory longitudinal vein near middle next to radial vein;hindwing with large dark spot near stigma and center of wing at about 2/3 distance to apex, India, Burma (Note. One species described from Africa but mislabeling is suspected) •••••••••••••••• ••••••••••••••••••••••••••••••••••••••••••••••••• Stiphroaeuria Gers taecker 10. Radial sector arises much closer to cubital fork in forewing than to medial fork in hindldng; distal palpomere of labium relatively short with a long oval-shaped sensory opening; Australia •••••••••••••••••••••••• Cos ina Navas Radial sector in about same relative position to cubital fork Uoreldng) and medial fork (hindwing) in both wings; distal palpomere of labium elongate with a long slit-like sensory opening 11 11 Forewing costa] area narrow and single-celled at basal 1/2 sometimes be- coming biareolate distally although not changing much 1n width before stigma ...... ] 2 Forewing costal area broad and 2-celled from near base •••••••••••••••••• 13 12. Eye very large, greatest ocular width nearly twice that of interocular distance; male tergite VI posteriorly with dense black or white setae in marked contrasttoprecedingtergites; Ethiopian AV1a Navas Eye only moderately enlarged, less than 1.5 times interocular distance; male tergite VI without unusual vestiture; Australian MeBtreBBa Navas 13. Hindwing with hypostigmatic cell less than 4 time longer than high; apex of hindwing with strong concavity beyond cell distal to hypostigmatic cell; Philippines •••••••••••••••••••••••••••••••••••••••••• Hadrasta Navas Hinowing with hypostigmatie cell more than 7 times longer than high, hind wing without concavity in apical field; Australia, ?Indonesia •••••••••••••• • ••••••••••••••••••••••••••••••••••••••••••••••••••••••••••• Beoclisis Navas

Acanthaclisls Rambur 1842 widely spaced teeth on the mandibles, and a (Figs. 4, 16, 20) middle tooth longer than the distal tooth, Reference: Steffan (1975), Brauer (855), which distinguishes it from other known Hagen (1873, 1887), Redtenbacher (1884), genera. It is unique in having thinly- Willmann (1977) scattered setae on the ventral head cap Description: Head capsule (l) 4.9 mm sule. long, 3.8 wide; mandible 1.5 long, .8 wide, Behavior. Large larvae were found in anterior margin of clypeal-labrum produced Egypt in deep sand around the base of a as a rounded lobe; antenna longer than palm tree. They moye bael,ward and forward basal width of mandible; distal palpomere rapidly. However, in pursuit of prey they of labium about 2 times longer than broad, appear to move only backliard. sensory opening small, not bulging; man- Larvae Studied' Acautbaclisis sp. d1ble strongly broadened at tooth I, dis­ EGYPT:Ismailaia, June 6, 1984, R. B. Miller tance between teeth ] and 3 greater than and]. A. Stange (3 SC, 5 MC). greatest width of mandible; tooth 2 longest and c loser to tooth 3 than to 1; usua 11 y 2 Centroc]iSfS Navas 1909 setae between teeth 1 and 2, 1 seta between (ngs. 2, 3, 18, 22) teeth 2 and 3; setae on exterior margin of (=Sogra Navas 1912, =Heboda Navas 191], mandible extending somewhat beyond tooth 2, =Heoclisis Navas 1914, =Sograssa Navas longest setae about 1/3 greatest mand1bular 1924, *'Stenoclisis Navas 1932) width; ventral head capsule with several we 11 separated setae (2nd and 3rdins tars) , Description: Head capsule (1) 3.4 mm or glabrous (1st instar); abdomen with long long, 3.0 wide; mandible 1.1 long, .55 dark setae especially laterally on ster wide, anterior margin of clypeal-labrom not nites I-VI; sternite VIII and IX with many produced, widely emarginate; distal palpo­ peg-like setae, sternite IX broadly mere of labium about 2 times longer than rounded. wide; sensory opening moderately large, This genus has a combination of peg bulging; antenna ahout as long as basal like setae on sternites VIII and IX, three width of mandible; mandibles thick with 2 34 INSECTA MUNDI Vol. 1, no. 1, January 1985

4 Acanthaclisis sp. 5 Fadrina sp.

6 Paranthaclisis 7 Phanoclisis longicollis

8 Synclisis baetica 1 mm. 9 Syngenes longicornis strong teeth and no setae on along interior The 2-toothed mandible without setae margin; teeth of about equal size; exterior on the inner margin 1s unique among the margin of mandible with short setae ending known genera of the tribe. at tooth 2; mandible of uniform width to Behavior: The larva of the species distal tooth; abdomen with numerous long studied mOves both backward and forward but black setae on sternites I-VII; sternltes quite slowly. The mandibles are the most VIII and IX with numerous short, blunt peg­ powerful observed in the Acanthaclisini like setae; sternite IX broadly rounded. which allows the larva -to feed on harder- Vol. 1, no. 1, January 1985 INSECTA ~roNDI 35 bodied prey than most other types of setae; sternite IX bluntly rounded. ant1ions known to us. The mandibles are The larvae of Fadrin8 represent an bui It for more powerful grasping of prey. extreme 3-toothed mandible type where the This suggests why no setae are present on teeth are very close together and near the the inner margin of the mandibles, as these base of a strongly broadened mandible (Fig. would be broken if present. Other acantha- 14). clisine larvae studied drag prey sWiftly Behavior: Larvae move backward or through the sand killing them relatively forward, but are reluctant to run forward. slowly as they are eaten. They rely on Larvae were found living in open sand. speed and strength rather than fast-acting T,arvae pursue prey by digging rapidly back- toxicity. However, although the Centro­ ward and then whipping the head backward to cl1sis larva studied was very powerful it grasp prey when they are beneath it They was slow moving. Observations revealed never ran forward after prey. Two species that very strong tox1n was 1njected into of tsdrina were reared. In one species, prey since large prey were paralyzed in a the anterior margin of the clypeal-labrum few seconds and were 11fe1ess within 10 1S S inuate, the antennae are equal in seconds. This is important to the larva length to the basal width of the mandible, because it lacks sufficient speed to pro· and tooth 3 is tW1ce as long as tooth 2. tect itself against its prey by rapidly In the other species, the anterior margin digging through the soil. The larva was of the clypeal-labrum is produced as a not an incessant feeder as are other genera broadly-rounded lobe, the antennae are discussed except Parsnthaclisis. It would twice as long as the basal width of the rest for a few days after a sizeable meal mandible, and tooth 3 is 1.5 times longer without digging near the surface. Also, it than tooth 2. proved to be strictly nocturnal, and would Larvae Studied. SOUTH AFRICA. Cape dig deeply if exposed to light. It cap­ Province, 4 km. N. Clanwilliam, January 30, tured prey by slowly digging baclEward and 1983, R. B. Hiller, L. A. Stange (l se), whipping its head backward at prey. The Ba ines Kloof (3 MC, 1 SC). time passed from cocoon construction to adult emergence was 56 days. This appears Beoclisis Navas 1923 to be the largest acanthaclisine genus in (Figs. 11, 17, 25) terms of described species (34) and also the most Widely distributed, from South References. Adams (936), Mathew (1950) Africa north to Morocco and east to Malaya. Description: Head capsule (1) 4.12 mm Since the 1srvse 1 i up deep under the sand long, 331 wide; mandible 1.9 long, .66 during the day they should be searched for wide; anterior margin of clypeal-lahrum at night. weakly s1nuate, slightly emarginate at Larvae Studied: Centroclisis punctu- middle; antenna much longer than bas~l lata Navas. Tunisia: Tozeur, May 17, 1~84, w1dth of mandible; distal palpomere of R. B. Miller, L. A. Stange (l MC, 3rd labium about 2 to 2.5 times longer than instar skin). wide; mandible broadened at level of tooth I, distance between teeth 1 and 3 longer FadriDa Na~as 1912 than greatest mandibular width; tooth 3 (Figs. 5, 14, 26) slightly longer than 2 which is slightly longer than 1; tooth intervals vary (2 DescriptioBI Head capsule (l) 3.0 mm species seen), tooth 2 closer to tooth 3 long, 2.3 wide; mandible 1.0 long, .5 (sp. A, Fig. 17) or to tooth 1 (sp. B); wicle; anterior margin of clypeal-Iabrum setae on exterior margin of mandible extend variable, sinuate, or produced as very from near base to level of tooth 3, longest broadly-rounded lobe; distal palpomere of setae about 1/4 greatest mandibular width, labium about 2 times longer than wide; ventral surface of head capsule glabrous; sensory opening not bulging; mandible "ery abdomen with long, black, hair-like setae broad at level of tooth 1 and beyond to 3; on sternites I-VII; sternites VIII and IX tooth 3 at least 1.5 times longer tban without short, peg-like setae, but short, tooth 2; usua lly 1 seta between teeth; pointed setae present especially along setae on exterior margin of mandible rela- midline; sternite IX bluntly rounded tively long, some almost 1/3 greatest man- We have studied only 2 preserved lar- dibu1ar w1dth; ventral head capsule vae from AustraI1a Wh1Ch represent 2 dif­ glabrous; abdomen with numerous long, black ferent species It is possible that at setae on sternites I-VII; sternites VIII least I of these could be Cosin8 Navas. and IX with numerous short, blunt, peg-like The large size of both larvae precludes 36 INSECTA MUNDI Vol. 1, no. 1, January 1985

, mm. 10 Vella fallax 11 Heoclisis Sp.

~', . ,:.~ , " . .:.~ .~;:.: . .-... . -.. ~.:-­ ~'.. .~ ~ . ..~ _.~ . ~ -'h. • " 2 mm.

12 Synclisis baetica 13 Syngenes longicornis them from being Mestre.sa Navas , the third Paraathacl181a Banks 1907 kno\o'n genus from Australia. (Flg•. 1, 6, 19, 23) Behavior: Mathew (1950) stated that these larvae only move backward. Adams Referencesl Hagen (1887), Stange (1936) remarked that SOme specimens ""ere 0970, 1980) found at the base of trees. Mansell (per­ De8cription: Head capsule (3 hagen!) sonsl communication) has reared at least 2.8 mm long, 2.7 wide; mandible .9 long, one species found under rock overhangs in .5 wide; anterior margin of clypeal-labrum Austra 1 ia. He observed that the larvae can not produced medially, roundly emargtnatej walk forwards with great agility, but very distal palpomere of labium about 1.5 times seldom seem to do this, preferring to move longer than wide, sensory opening large and backwards through the sand when pursuing bulging; antenna with about 11 segments and prey and leave conspicuous tracks. shorter than basal width of mandible; man­ Larvae Studied: AUSTRALIA: Queensland, dible only slightly broadened at level of near Hughender, September 7, 1972, R. E. tooth; mandible with I tooth; mandible Woodruff (1 FSCA); 3 miles S. Woodstock, without setae except for short setae on September 6,1972, R. E. Woodruff (1 FSCA). exterior margin from base to about level of Vol. 1, no. 1, January 1985 INSECTA ~ruNDI 37 tooth; ventral surface of head capsule broadened at level of tooth I, distance glabrous; abdomen with short hair-like between 3 teeth equal to or slightly less setae on sternites I-VII; sternites VIII than greatest width of mandible; mandibular and IX with numerous peg-like digging tooth 2 longer than tooth 3, tooth 2 some setae; sternlte IX bluntly rounded. what closer to I than to 3; usually 1 seta The one-toothed mandible is unique between teeth; setae on exterior margin of among known genera of the trlbe. also, the mandible about as long as one-half mandibu­ reductlon of the antenna lS shared only by lar Width; ventral surface of head capsule Centroclisis and one species of Fadrina. glabrous; sternites VIII and IX with numer­ Behavior: These larvae are found in ous peg-like digging setae; sternite IX sand dune areas and flat areas of deep bluntly rounded. sand, and move both forward and backward Behavior: The se larvae exh i 6i t on 1y rapidly. Colonies from Nevada and Cali­ backward movement and were found in sand fornia fed with difficulty but P. hageni dune habitats. They dig very rapidly after (Banks) from Baja California and Parantba- prey. cltsts sp. from Florida fed on lepidop Larvae Studied: Phanoclis18 100gi­ terous larvae and were reared. Field ob- collis (Rambur). TUNISIA: 30 km. NW Gafsa, servations in Nevada showed P. nevadensis Qued el Kebid, May 15, 1~84, R. '8. Miller, to be feeding on lepidopterous larvae and L. A. Stange (4 Me, scL EGYPT:Abu Rawash beet Ie larvae of the family Coccine 11 idae. (near Caito), Nay 30, 1984, R. B. foliller, Larvae dig rapidly backward after prey and L. A. Stange (2 MC, 2 SC). grab it by whipping their head backward a fter they are underneath it They never Synelisis Na~a8 1919 pursue prey by running forward. None of (Figs. 8, 12) the species feed constantly, at least in the laboratory, but those from Nevada have References: Redtenbacher (1884), Principi prolonged periods of diapause corresponding (l9lj7), Steffan (1975) to those perlods of the year when their Description: Head capsule (3) 4.7 mm habitat has constant sub-freezing tempera- long, 3.5 wide; mandible 2.0 long, .6 l~ide; tures. anterior margin of clypea I-labrum not pro­ Larvae Studied: Parantbaclisis ba&eni duced med ia lly, slight ly emarginate media1- (Banks). MEXICO:Baja California, Chapa la ly; distal palpomere of labium about 2 Dry Lake, June 20, 1~83, R. ll. Miller, L. times longer than wide, sensory opening A. Stange (4 SC, FSCA); CALIFORNIA:Inyo moderately large and bulging; antenna with County, Stovepipe Wells, Death Valley, about 13 segments, longer than basal width January 3, 1965, F. Parker, L. A. Stange, of mandible; mandible weakly broadened at (20 SC, FSCA), San Bernardino County, Ke1 level of tooth 1, width at thlS point less so, Au gus t 15, 1 97 3, R. B. Mille r (l MC) • than length between base and tooth 1; man- Parantbaclisis sp. FLORIDA:Gulf dible typlcally wlth 1 seta between teeth I County, St. Joseph's Peninsula, November I, and 2, 2 setae between teeth 2 and 3, smal] 1978, L. A. Stange H FSCA, 1 Me). setae on exterior margin of mandlble from Parantbaclisis congener (Hagen). near base to about tooth 2; mandible with 3 NEVADA. Store, County, 27 miles E. Reno, teeth, tooth 3 longer than 2 WhlCh lS August 9, 1980 (11 nc, SC, FSCA, from longer than 1; tooth 2 closer to I than to eggs); Washoe County, 4 miles N. Nixon, 3; ventral surface of head capsule glab­ January 4, 1981, R. ll. Miller (4 MC, SC). rous, distal palpomere of labium about 2 Parantbaclisis nevadensis Banks. times longer than wide; abdomen with fine, NEVADA' Washoe County, Pyramid Lake, June pale, hair like setae on sternites I VII, 10, 1980, R. B. Miller (l MC). sternites VIII without short, peg-like setae; sternite IX pointed (Fig. 8). Phanoclisis Banks 1913 The shape of the last sternites (Fi gs 7, 15, 24) (broadly pointed) is distinctive as is the shape of the mandible (evenly tapered from Description: Head capsu Ie (I) 3 9 mm tooth 1 to 3), showing the most similarity long, 3.2 wide; mandible 1.5 long, .8 wide; to Syngenes. However, the anterior margins anter ior margin of clypeal-labrum produced of the clypeal-Iabrum of the 2 genera are as a rounded lobe, slightly indented quite different. medially, distal palpomere of laolus about BebaVior: These larvae are very fast 3 times longer than wide, sensory opening runners and fast burrowers, preferring open small, not bulging, antenna longer than sand dune areas. They wl.ll run forward or basal width of mandible; mandible great ly dig backward after pre~ 38 INSECTA MUNDI Vol. 1, no. 1, January 1985 ;J'

14 Fadrina sp. 2 mm. 15 Phanoclisis longicollis

16 Acanthaclisis sp. 17 Heoclisis sp.

18 Centroclisis punctulata 19 Paranthaclisis hageni Larvae Studiedl Synclisis baetlea Syngeneo Kolbe 1897 (Rambur). SPAIN:San Lucar de Barramcda, (6 -OneIu8 Navas 1912) SC); TUNISIA:Bizerte, May 28, 1984, R. B. (Figs. 9, 13) ~1111er, L. A. Stange 00 SC, FSCA, 10 Mcl; EGYPT:Hannovil Beach (SW of Alexandria), Description: Head capsule (3) 3.5 mm June 7, 1984, R. B. Miller, L. A. Stange (4 long, 2.8 widej mandible 1.60 long, .5 MC, SC). wide; anterior margin of clypeal-labrum Vol. 1, no. 1, January 1985 INSECTA BUNnI 39 produced as a rounded lobe; antenna longer less than between base of mandible and than basal width of mandible; distal palpo- tooth; 3 mandibular teeth evenly spaced, mere of labium about 2 times longer than tooth 2 and 3 are nearly equal in length, wide, sensory opening small, not bulging; longer than tooth 1; ventral surface of mandible moderately broadened at level of head capsule densely setose; sternite VIII tooth 1 (Fig 13); mandible witb distal and IX densely setose, many long truncate tooth much longer than middle tooth, tooth setae, no peg-l1ke setae, but short, point­ 1 and 2 much closer togetber than tooth 2 ed setae on sternite VJlI; sternite IX and 3; distance between teeth somewhat bluntly rounded. longer (about 1.5 times) than &reatest The densely pubescent ventral surface mandibular Width; usually 2 setae between of the head capsule is a diagnostic char- teeth I and 2 and between teeth 2 and 3; acter 1n the tr1be. setae on exterior margin of mandible at Behavior: We have found the larvae in least 1/3 length of mandible, extending to dunes (V. assi.i1is) and in fairly deep tooth 3; abdomen with only 5 pale setae on tracts of sand (V. americana and V. fal- steInites I VII; steInite VIII with weakly lax). These laIliae only UlOlle backwaId produced digging setae, no peg-like setae; through the sand leaving conspicuous trails sternite IX with many small peg like setae, on the surface. V. americana has been sternite IX pointed. observed feeding on Hyrmeleon larvae in Syngenes larvae ale ditinguished fIom Florida. other acanthaclisine larvae by the median Larvae Studied: Vella americana lobe of the clypeal-labrum in combination (Drury). FLORIDA:Highland County,S miles with the lack of the peg like digging setae S. Sebring, March 12, 1980, R. B. Miller, on sternite VIII. Also, the mandible is L. A. Stange (3 FSCA, SC); Archbo Id Bio­ relatively slender with tooth 1 and 2 logical Station, March 13, 1980, R. B. closer together than 2 and 3. Tooth 3 is Miller, L. A. Stange (2 SC, 2 MC); Okaloosa the longest in contrast to most genera in Couoty, 2 miles N. Holt, October IS, 1983, the tribe. The 9th s terni te is pointed L. A. Stange (2 SC). (Fig 9), most similar to the condition of Vell8 8ssimi 1 i s Banks PERIJ·I.a I.iber- Synclisis. tad, Laredo Sand Hills, July I, 1974, L. A. Bebayi or- We observed large numbers of Stange (JO SC); July 16, 1982, R. B Hi 1- larvae in sand dune areas of coastal South ler, L. A. Stange (2 SC, 4 MC). Africa. lhey can run fast forward and Vella faltax fallax (Rambur). MEXI- burrow quickly. They use both of the capa­ CO:Baja Califoria Sur, Juncalito Beach, May b1lit1es 1n capturing prey. In South 1983, R. B. M111er, L. A. Stange (2 SC, 2 Africa, their prey consisted mostly of MC); Tamaulipas, San Antonio, June 26, large sand-burrowing Lep1doptera larvae. 15181, R. B. Miller, L A. Stange (I FSCA, I Larvae Studied: Syngenes longicornis MC). . (Rambud. SOUTH AFIUCA:Natal, Warner Vella fallax haHlensis Smith. FLORI­ Beach, 10 miles S. of Durban, January 1983, DA:Monroe County, Bahia Honda, October 17, R. B. UilieI, L. A. Stange (Io SC, FSCA, 11 1982 (I se, 9 nc). MC); Cape Province, Mossel Bay, January 26, 1983, R. B. UHleI, L. A. Stange (I se). Higher Category Larval Characters Vella Navas 1913 (Figs. 10, 2l) To help identify larvae of the tribe (=Gyroplectron Esben-Petersen 1928) Acanthaclisini we are providing a key to the subfamilies and tribes of Myrmeleonti- References: Hagen 0873, 1887), Stange dae. However, many genera (about 75%) are (1970, 1980) unknown in the larval stage as are several Description: Head capsule (4) 4.2 mm tribes (Maulini, Pseudimarini, Porrerini) long, 3.5 wide ({allax) or 4 7 long, 3.8 and many slJbtribes (Dimarellina, Peri- wide (americana); mandible 1.1 long, .6 clystina, Voltorina, Acanthoplectrina, wide lfallax) or 19 long, 7 wide (ameri- etc) are unknown to date in tbe larva 1 cana); anterior margin of clypeal-labrum stage. Therefore, the following key will (Eg. 10) nearly straight, weak Iy indented undollbted Iy not fit al [ the genera, at middle; antenna much longer than basal especially highly specialized genera, and width of mand1ble; distal palpomere about 3 should be cons1dered as a preliminary key. times longer than wide, sensory area mod- It is obvious to us that the Dendroleontini erately long, bulging; mand1 ble weakly and Nemo1eontin1 (including G1enurini, broadened at tooth 1, width at this point Creoleontin1, Formicoleont1ni, etc.) are 40 INSECTA MUNDI Vol. 1, no. 1, January 1985

21 Vella fallax 22 Centroclisis punctulata

20 Acanthaclisis sp.

24 Phanoclisis longicollis 23 Paranthaclisis hageni

2 mm.

25 Heoclisis sp. 26 Fadrina sp. Vol. 1, no. 1, January 1985 INSECTA MUNDI 41 closely related, and therefore some genera tioo of the terminal abdominal segments may overlap in structure. The Brachy­ (highly modified digging setae, presence or nemur!n! includes the tribes Cepini, Myrme­ absence of submedial teeth on sternite caelurlni and Nesoleontini according to our VIII), the shape of the mandible, the de­ larval studies, but some bizarre larvae are velopment of the labial palpuB, and the apparent in this tribe

Key to Subfamilies and Tribes of Myrmeleontidae LARVAE (Porrerini unknown)

1. Tergite IX with median lobe bearing median 8clerotized process or sternite IX with highly modified pair (or more) of digging setae (blade-like to broadly triangular); sternite VIII with pair of submedial teeth near poster- ior margin ••••...... •.•.•••...••..•..•.•••...••..•.••••••••••....••...••• 2 Tergite IX without median lobe bearing sclerotized process; sternite IX without highly modified digging setae; sternite VIII with or without submed­ ial teeth near posterior margin ••••.•••••••••••• Myrmeleontinae •.••.••• 3 2. Tergite IX with median lobe bearing median sclerotized process; sternite IX without highly modified digging spines; sternite VIII with inconspicuous submedial teeth; mandible with 3 teeth; Australia ....••.•• Stilbopteryginae Tergite IX with median lobe without sclerotized median process; sternite IX with pair (or more) of submedian digging setae greatly enlarged as blade­ like to broadly triangular processes (except some Dimare.); sternite VIII with well developed submedian teeth near posterior margin;mandible with 2 to 5 teeth; Widespread except Australia and North America ••••.••••• Palpar1nae 3. Labial palpus shorter than basal width of mandible;mesothoracic spiracle not borne on tubercle; head without dolichssters; backward movement only (many

genera) or backward and forward •••••••••.••.••••.•••••••••••••••••• o. 0 • • 4 Labial palpus longer than basal width of mandible or mesothoracic spiracle borne on tubercle; head often with dolichasters; backward and forward move-

men t ••••.....••••....•..•..•. 0 • • • ••• • • • • •••••••••• • • ••••••••••••••••••• •• 5 4. Mandible with some setae on exterior margin as long or longer than greatest mandibular width; sternite VIII with pair of inconspicuous submedian teeth near posterior margin; make pitfall traps ••••••••••••••....• Myrmeleontini Mandible with longest setae on exterior margin less than one-half greatest mandibular width; sternite VIII without teeth on subapical margin; pitfall traps absent ••••••••••••••••••.•••••.••••••••••••.••.•.•••• Acanthaclisini 5. Mandible with distal tooth equal to or shorter than middle tooth. distal tooth often set at different angle (acute projecting anteriorly); usually middle tooth closer to distal tooth than to basal tooth; sternite VIII often with pair of inconspicuous submedial teeth near posterior margin (absent in Mophie. Scotoleon); scoli usually absent on abdomen and much reduced on tho­ rax (known exceptions are Cnopboleon and Jaffuel~) (includes Gepini. Neso- leontini. and Myrmecaelurini) ••• 0.... Brachynemurini Mandible with distal tooth longer than middle tooth (ClenuTU8 with 2 teeth). teeth more or less parallel and usually equidistant; sternite VIII with or without submedian teeth; scoli often developed on thorax. less common on ab-

domen •••••••••••••• 0 • o ••••••••••••••• 0 ••••••••••••• 0 ••••• o •••••••••••••• 0 6 6. Meeoscutum usually with tuft of setae at middle; 9th abdominal segment near­

ly as long a9 wide •••• 0.0 ••••••••••••••••••••••••••••••••••• Dendroleontini Mes09cutum without tuft of setae at middle; 9th abdominal segment at least 2 times wider than long ••••.•••••••••••••.••••.•••••••..•...•• Nemoleontini 42 INSECTA MUNDI Vol. 1, no. 1, January 1985

Cenera Studied (Larval Stage) 8lBLIOGVJ'HY

(Approximate number of genera Adams, N. B. 1936. Ant-lions. Australian in parentheses) Mus. Mag. 6,22-25. Brauer, F. M. 1855. Beitrage zur Kennt­ Palparioae (14), niss der Verwandlung der Neuroptera. Palparini (9): No.a, Palpares, SteDares Ueber das Vorkommerund und die Leben­ Dimarin! (includes Echthromyrmicini) swelse der AcantbacllBie occitAnica 0), Di.area Villers. Verh. Zool. Bot. Wien. 5:1­ Maulin! (1): None 10, 1 plate. Pseudimarini (l): None Hagen, H. 1873. Die Larven von Myr.e­ St1lbopterygiDae (2), St1lbopteryx leon. Stettin. ent. Ztg. 34,249-295. MyrmeleontiDae (138), Hagen. H. 1887. Stray notes on Myrme­ Acanthaclisini (14): Acantbaclis18, Cen­ leonidae. Part 2. Canadian Entomol. troct!s!., Fadr:l.Da, Heoc1181a, Phano­ 19,147-156. clials, Paranthaclisl., Syncl!.!., Mathew. W. 1950. Notes on the ant-lions Syngenes, Vella of the subfamily Myrmeleontinae. W. Brachynemurlni (includes Cepini, 160­ Australian Nat. (Perth) 2:64-66. leonttni, Myrmecaelurini, Nesoleon­ Navas, L. 1912. Notas Sobre :Mirme- tini) Abatoleon, Ameromyi., Brachyne- leonidos. Broteria (Ser. ZooI.) .uruB, Cbaetoleoo, euet., Furgel- 10,29-97. la, Cepu8, Cnopholeon, Jaffuelia, Principi, M. M. 1947. Contributi all0 Lemole.u8, MenkeleoD, Myr.ecaeluruB, Studio dei "Neurotteri" Itali­ Mcphie, Scotoleon, Soltel' ani. VI. Synclisis baetica Ramb. Dendroleontini (31): Banki8u8, Dendro­ Boll. Ist. Ent. U. 8010gna 16,234­ leon, Tricholeon 253. Myrmeleontini (5): Myraeleon, Weeliu8 Redtenbacher, J. 1884. Ubersicht der Nemoleontini (includes Distoleontini Myrmeleoniden-Larven. Denkschr. Formicaleontini, Protoplectrini, Creo­ Akad. Wiss. W1en 48,335-368, Figs. Ieontini, Obini, Nyutini, Glenurini. 1-118. DimarellinU (64): Creoleon~ Di.to­ Stange. L. A. 1970. Revision of the leon, Elachyleon~ Bremoleon~ Cymnoc­ ant-lion tribe Brachynemurini of neaia~ Clenurue, Nava801eon~ Neuro­ North America. Uni v. Calif. Publ. leon~ Obue~ Peam.oleon Entomn!. 55,1-192. Stange, L. A. 1980. The ant-lions of ACKNOWLEDGMENTS Florida. II. Genera based on larvae. Florida Dept. Agric. & We thank Dr. Mervyn Mansell (Pretoria, Consumer Serv., Div. Plant Indus­ South Africa) for critical comments. Lar­ try. Entomol. Circular 22:1-4. vae used in this study are deposited in the Steffan, J. R. 1975. Les Larves de following collections: Florida State Col­ Fourmilions de Is Faune de France. Ann. Soc. ent. Fr. (N.S.) lection of , Gainesville. FL (FSCA); Robert B. Miller Collection, Pro­ 11(2) ,383-410. ject City. CA (MC); Lionel A. Stange Col­ Willmann, R. 1977. Die Myrmeleontidae lection. Gainesille, FL (SC). Acknow­ der Dodekanes/Agais. Zool. Jb. ledgment is made to the National Geographic Syst. 104,98-136, Fig. 1-34, Table. Society for 3 grant to Stange (1977) to study Neuroptera in South America.