HYM. RES. J. Vol. 15(2), 2006, pp. 270-276 The Genus AUotilla Schuster (Hymenoptera: Mutillidae): Phylogenetic Analysis of its Relationships, First Description of the Female and New Distribution Records DlOMEDES QUINTERO A. AND ROBERTO A. CAMBRA T. Museo de Invertebrados G.B. Fairchild, Estafeta Universitaria, Universidad de Panama, Panama, Panama; email: [email protected] [email protected] — Abstract. The morphological characteristics of the monotypic genus AUotilla Schuster are discussed; new distribution records and a morphology-based phylogenetic analysis are presented, clarifying possible relationships with other sphaeropthalmine genera. Females of the genus are described and illustrated for the first time; male genitalia are illustrated. Until now, AUotillagibbosa Schuster, 1949 Natural History, New York (AMNH); was known only from the holotype male Museum of Comparative Zoology, Har- from Cordoba, Argentina. In a list of vard University , Cambridge (MCZ); Mu- Mutillidae from Argentina, Fritz (1998) seo de Invertebrados G. B. Fairchild, Uni- included Buenos Aires as an additional versidad de Panama (MIUP); D. Broth- J. collection locality for this species, without ers's personal collection, University of providing any additional information such KwaZulu-Natal, South Africa (DJBC); and as the number of specimens examined, Institut Royal des Sciencies Naturelles de their sex, or the depository. We must Belgique (ISNB). consider the Buenos Aires record not to RESULTS be valid because it is incomplete and because we have been unable to locate any of Fritz's AUotilla specimens in his AUotilla Schuster reference collections at the AMNH, New AUotilla Schuster 1949: 89-93. Type species: York (to which they were sold by his AUotilla gibbosa Schuster, 1949, by original widow after his death) or at the Museo designation and monotypy. — de Ciencias Naturales "B. Rivadavia," Generic diameters offemale. Body cov- Buenos Aires (Roig-Alsina, pers. comm.). ered with simple setae only. Head: almost The female of the Neotropical, mono- round, narrower than mesosoma; eye tilylpuisctragteednufsor AtUhoetiflirlsat tiisme;densecrwibdeidstraibnud- gsmeanlall,canreianralyabsceinrtc;ulsacrroabanldcafrlaitna(aFibgs.en3t);; tion records and a morphology-based antenna 12-segmented; antennal tubercle phylogenetic analysis are presented. strongly projecting, with low lamellate MATERIALS AND METHODS ridge on anteromedial surface (Fig. 3); mandible slender, with preapical tooth We follow Brothers (1999) for the sub- nearly obsolete or totally absent (Fig. 3); family classification of the Mutillidae. For mandible evenly arcuate on ventral mar- SEM examination we used a JEOL model gin, without tooth or constriction; probos- JSM 5600. The AUotilla gibbosa specimens cidal fossa with carina nearly reaching examined are deposited in the follow- innermandibularbase; hypostoma without ing institutions: American Museum of tooth or tubercle; maxillary palp 6-seg- Volume 15, Number2, 2006 271 merited; labial palp 4-segmented, the sec- sized punctures, except for mostly im- ond segment almost as long as wide. punctate anterior area near lateral area of Mesosoma: broader than long, pyriform pronotum; metapleura and lateral area of (Fig. 4); dorsum strongly convex, without propodeum impunctate, smooth; setation transverse carina along posterior margin of of pronotum, mesonotum and metanotum pronotum or lateral carina; dorso-lateral similar to that of vertex; mesopleura and margins without spines; scutellar scale dorsum of propodeum with pale setae; absent; mesopleuron swollen; leg with metapleura and lateral area of propodeum apex of tarsus simple, not produced above glabrous. Metasoma: terga I and II with claws. Metasoma: first metasomal tergum small, dense punctures, sparser in apical not constricted posteriorly, sessile with areas (Fig. 5); terga III and IV with small, tergum 2 (Fig. 5); tergum II evenly convex, sparse punctures; tergum V mostly without rows of longitudinal carinae; ter- smooth, except basal lateral area with gum II with felt lines; sternum II without a few punctures; tergum VI with scale-like felt lines; tergum VI with surface totally surface sculpture basally, scales diminish- sculptured and evenlymerging with restof ing in size toward apex, gradually turning tergum, pygidial area poorly defined by into granules (Fig. 6); sterna I and VI postero-lateral carina, only visible under smooth; sterna II and III with small, high magnification (>30x) (Fig. 6). somewhat sparse punctures; sterna IV and V mostly smooth, except apex with Allotilla gibbosa Schuster, 1949 small, dense punctures; tergum I mostly (Figs 3-8) with pale setae, a few dark setae at apex; Allotilla gibbosa Schuster, 1949: 93-95, Holotype tergum II with dark setae, except lateral male, Argentina: Cordoba, col. W. M. Davis, area and apex with pale setae; terga III and Harvard University, MCZ, type 30516, exam- IV mostly with pale setae; tergum V mostly ined. glabrous, with a few pale and dark setae — laterally; tergum VI glabrous; sterna I to V Description offemale. Integumental col- with pale setae; sternum VI—glabrous. or: head, mesosoma, all metasomal sterna, Additional male characters. The external terga I, III and IV, reddish brown; antennae male genitalia and the penis valve (Figs 7- and legs yellow-red; mandible reddish 8) are illustrated here for the first time brown except apical third blackish; tergum (paramere, cuspis and digitus were de- II reddish brown except posteriorly with scribed previously but not illustrated). The truwpoteldatemraeld,iatlrlayn;svetresreg,a bVlacakndspoVtIs bilnatcekr.- vcoulsspeilslaapheaxs d(Fiisgt.in7c)t.ivTehelopnegnissetavealoven htahse Head: vertex and gena with sparse, medi- a subapical tooth, more distant from the um-sized punctures one or more diameters apical tooth than in males of Protophotopsis apart (Fig. 3); punctures of frons denser, (seeFigs 11-14,CambraandQuintero 1997). less than one diameter apart; vertex and — frons with long, sparse, erect and semi- Material examined (56 males, 5 females). All erect, dark setae; gena, clypeus laterally males were captured with Malaise traps (B. asentdae.hMypeossotsoommaa:wpirthonolotnugm, asnpdarsmee,sopnaol-e GDaerlcseitnenecocloll.l).).anPdARfAemGaUleAsY:witBhoqpuitefralolntDraeppsar(tT-. tmeutmanwiotthumpuwnictthurterasnsavserosne rveetritceuxlat(Feibg.an4)d; imsetnrta:ciPona,rq2u3e9 Nmac(i2o1na1l2'TeS,ni6e1nt3e9'EnWc)iso1,6A-1d9miSne-p 2003, 20 males [MIUP, DJBC]; same loc, 20-24 (Fig. 4); dorsum of propodeum mostly Mar 2004, 6 males [MIUP]; same loc, 23-26 Sep densely micropunctate-rugose (Fig. 4) ex- 2004, 3 males [MIUP]; Siracua, 275 m (21 02' S, cept a narrow impunctate area near meta- 61 45' W) 20-22 Sep 2003, 21 males [MIUP, notum; mesopleuron with dense, medium- MCZ, AMNH, ISNB, DJBC]; Estancia Maria 272 Journalof Hymenoptera Research Vicenta, 235-244 m (20 55' S, 61 23' W) 26-30 sessile, evenly merging with second; and Sep 2004, 5 males [MIUP]; TransChaco, Mister seven Sphaeropthalmina (Nanotopsis, Pro- Long, (20°35' S, 62 02' W), 17Sep 2003, 1 female tophotopsis, Reedomutilla, Scaptodactyla, Li- [MIUP]; Parque Nacional Teniente Enciso, maytilla, Suareztilla and Limaytilla), females TransChaco, 23-25 Sep 2004, 3 females [MIUP, with head nearly round, narrower than the ISBN, DTBC]; same data but 24-25 Sep 2003, 1 mesosoma, a—nd genal carina absent. female [MCZ]. Presidente Hayes mDepartment: Characters. Twenty-three binary and Reserva Tinfunke, La Verde, 146 (23 56' S, multistate characters of adult male (M) 69°29' W) 29 Nov-1 Dec 2003, 1 male [MIUP]. — and female (F) external morphology and Variations. Female frons dark reddish male genitalia were coded for analysis; all brown to black; tergum IV varying from were treated as unweighted and unor- totally reddishbrown toblack or the lateral dered. No autapomorphies were used. areas black with reddish brown in the The character matrix used is given in middle. Males from Paraguay are identical Table 1. The following characters were to the holotype, except that the propodeal employed for cladistic analysis: lateral area is rugose on the holotype, but Head: punctate with smooth areas to rugose- — punctate or totally rugose in specimens Head shape (F): small, almost—round, from Paraguay. We consider this variation not broader than mesosoma; 1 trans- to be size-related: male rugosity increases versely subquadrate, large, distinctly with body length. In addition, larger males broader than mes—osoma. have the notauli nearly obsolete (same as 2. Head (M, F): wit—hout large conical the holotype), but notauli are absent in projection ventrally; 1 with large conical smaller males. Total length, females: 3.5- projection ventrally.— — 5 mm; males: —4-7 mm. 3. Scrobal carina (F)—: presen—t; 1 absent. Distribution. Paraguay and Argentina. 4. Genal carina (F): absent; 1 present. AUotillagibbosa was previously known only 5. Mandible basal vent—ral margin (M, F): - from the holotype from Cordoba, Argen- with constric—tion; 1 with broad lamellate projection; 2 almost straight. tina. Antennal tub—ercle (F): -without lamellate PHYLOGENETIC ANALYSIS projection; 1 with lamellate projection on — anteromedial surface. T/7A77. To test the subtribal position of 7. Antenna (F): 0-12-segmented; 1-13-seg- AUotilla (currently included in the subtribe mented. — — Pseudomethocina, Brothers 1975), and to 8. Ocelli (M): small (diurnally active); 1 we recognize its phylogenetic affinities, large (nocturnally active). selected as the outgroup the following two genera: Timulla (Mutillini) and Dasy- Mesosoma: lAamberirsic(aD)a;syalsabirnignrio,upgetnaxuas, wnoetseplreecsteendtthien Dorsum of mesosoma (F): —longer than wafionltdlhofmuwlailniygnwliy1n7SgoeSudpthhmaaAelmerseo:rpitthceanalnPmsideinusitdroigmbeeuntteihrooan-,s bbStrr—hhooaaanapdde,leoronsfgo.mtmheeastnoismloeosnmga;as(F1)—b:rod—iasdtsiunabcsrtelocyrtabsnrlgoiugalhdaterlr;y cina (Euspinolia, Tallium, Atillmu, Calomu- 1 violin- shaped, strongl—y constricted at tilla, Horcomutilla, Pseudomethoca, Hoplo- the propodeal spiracles; 2 p—yriform. crates, Pappognatha, Hoplomutilla and AUo- 3. Axill—a of mesonotum (M): not expand- tilla), females with head transversely sub- ed; 1 expanded posterolateral^ as a rect- quadrate, broader than the mesosoma, angular or acute pro—truberance.— genal carina present (except Euspinolia 4. Scutellar scale—(F): prese—nt; 1 absent. and Tallium), first metasomal segment 5. Notauli (M): present; 1 absent. Volume 15, Number 2, 2006 273 Table 1. Data matrix for the 23 characters used in the phylogenetic analysis Characters 274 Journalof Hymenoptera Research sent autapomorphies in the present analy- sis: posterolateral tubercles on vertex, Atlllum pTrreasumeanttomiuntislolameanfdemCaelpheasloomfutiDlalsay;mumtailnlda-, HIPHosrecuHoodmpoullmoiecltlrhaaotcesa ibles covered with dense, short pubes- Calomutilla Hoplomutilla cence, an autapomorphy of Pappognatha; Pappognatha humeral angle of pronotum produced as H^Eusplnolla a hook-like tooth, an autapomorphy of —Talhum Nanotopsls Gurisita females (males unknown); posteri- Protophotopsis or coxa with tooth apically on inner Allotilla margin, an autapomorphy of Vianatilla Us-[TScaptodactyla females; tibia 2 with one apical spur, an -XystromuLtlimlayllallla autapomorphy of Acanthophotopsis males; -Reedomutilla males apterous or brachypterous, Morsyma -Dasylabrls -Suareztilla and (apterous), Myrmilloides Stethophotopsis — (brachypterous), Dasymutilla (rarely bra- Atlllum Psoudomcthoca chypterous); tergum II with arcuate trans- I' Horcomutllla verse band of dense, curled setae and i—Calomutilla slight integumental ridge at anterior margin of band, in most Dimorphomu- tilla females; female with felt line on sternum II, autapomorphy of Patquiatilla; sternum II with anteromedian seta-filled pit, in some males of Dasymutilla and Traumatomutilla; tegula elongated to or beyond the level of transscutal arti- culation, autapomorphy for Timulla males; eye inner margin deeply and abruptly notched, autapomorphy for Timulla males. Figs 1-2. Preferred minimum-length cladograms of A heuristic search of trees derived from cfhoaurracttreeresdadtearivperdesefnrtoemd hienurTiastbilce 1anaulsyisnigs roaftcthheet parsimony analysis was carried out using (WINCLADA) and mult* (NONA). Tree length = 62; NONA version 2.0 using WinClada version consistency index = 0.43; retention index = 0.66. 1.00.08 (Nixon 2002), resulting in four Synapomorphies are showby the blackcircles. cladograms. We preferred two of these minimal-length cladograms (Figs 1-2), see the mesosoma, and because these were the Results and Discussion. The following only female and male Sphaeropthalminae options were used: maximum trees to keep collected from the same geographic area, = 1000; number of replications (mult*N) = Paraguay's Chaco, that we recognized as = 1000; starting trees per rep (hold/) 100; belonging to a genus whose female was not random seed = 1000; unconstrained known. search; search strategy of multiple TBR + The relationships between American TBR (mult* max*). mutillid genera (none is known to be RESULTS AND DISCUSSION present outside of America, except for Timulla) and those of Africa and Australia The female ofAllolilla gibbosa was recog- are almost totally unknown, although nized based on morphological similarities different but closely related genera are to the known male, mainly the distinctive involved. Vicariant biogeographical hy- and peculiar inflation and broadening of potheses and their common ancestral geo- Volume 15, Number 2, 2006 275 Figs 7-8. Allotilla gibbosa, male genitalia. 7. Para- meres dorsal view. 8. Penis valve, lateral view. Abbreviations: P = paramere; ST = subapical tooth; VS = volsellar setae. Figs 3-4. Allotilia gibbosa, female. 3. Head, dorsal view. 4. Mesosoma, postero-dorsal view. Abrevia- graphic areas suggest close evolutionary tions: LP = lamellate projection of antennal tubercle, relationships with South American genera. M = metanorum, P = pronotum, Pr = propodeum, S Some genera widely distributed in the = scape. Americas, such as Sphaeropthalma and Traumatomutilla, cannot be included in the present cladistic analysis because they are •• .,-: Wm——— as yet poorly defined. > The heuristic analysis resulted in four cladograms, the two preferred minimal- * length cladograms (Figs 1-2) postulate a sister relationship ofAllotilla with Scapto- dactyla and Linun/1ilia. The other two cladograms postulate a less parsimonious, complex relationship for the selected South Amerian taxa and, following Occam's razor, were not accepted: Allotilla + [(Scap- + + + todach/la Limaytilla) (((Xystromutilla ((Reedomutilla + (Dasylabris + Suareztilla)))]. These preliminary cladograms suggest than Allotilla is more closely related to genera in the subtribe Sphaeropthalmina than to those in the Pseudomethocina. At present, we do not know of any unique morphological characters to separate these two subtribes. The biology of Allotilla gibbosa is un- known, but the black integument, moder- Figs 5-6. Allotilla gibbosa, female metasoma, dorsal ately sized eyes, and very small ocelli of view. 5. Tergum 1 and basal part of tergum 2. 6. the males suggest that they are diurnal. Tergum 6. The following morphological characters of 276 Journalof Hymenoptera Research AUotilla females indicate that they spend Spichiger et al. (2004), investigating the most of their lives underground: small, geographical zonation in South America of flattened eyes, relatively short legs with 32 common tree species encountered in a fore tarsal rake (used to excavate soil), Paraguay, found that the xeromorphic and a mostly reddish-brown integument. forests of the Chaco area act as an edaphic Extensive visual samplings carried out barrier to many species that are centered in during daylight hours in Teniente Enciso northern Argentina. The genus AUotilla is National Park did not yield any AUotilla reported only from northern Argentina females from the ground or on the sparse and Paraguay. vegetation; females were collected only If additional sphaeropthalmine genera with pitfall traps. These capture data lend eventually are included in the data matrix support to the postulated underground, presented here, the larger data matrix burrowing habits of the females. Probably might provide a better resolution in the theyparasitize small, underground-nesting phylogenetic analysis. aculeates. Females of the here-recognized ACKNOWLEDGMENTS closely related sphaeropthalmine genera, Scaptodactyla and Limaytilla, sister genera Our sincere thanks to Thibaut Delsinne, ISBN, for to AUotilla, have a similar morphological sorting and sending us all the mutillids from his habitus, suggesting that they have similar pitfall traps from Paraguay. We are grateful to hypogeal lives and burrowing activities. Philip D. Perkins, MCZ, for the loan of the holo- A morphology-based phylogenetic anal- type of A. gibbosa; Bolivar R. Garcete B., Museo ysis of 18 mostly South American mutillid lNeacctiionngalmuHitsitlolriida Nsapteucriamlendse; PAanrnaegutatye fAoirelclool,- genera and one from outside America Smithsonian Tropical Research Institute, and Denis (Dasylabris) permits us to construct a hypo- J. Brothers, University of KwaZulu-Natal, South thetical scenario of biogeographic diver- Africa for reading the manuscript and for provid- gences. The vicariant event that divided ing us with very helpful suggestions to improve its the population of the common ancestor of ScoEntMenwtosr;k.Alcides Muhoz, University of Panama, for the taxa presented in Table 1 was the uplifting of the Andean high mountain LITERATURE CITED range. This uplifting event was followed by ecological divergence of the two An- Brothers, D. J. 1999. Phytogeny and evolution of dean regions: the elongated West costal wdeaas,psV,esapntosidaenadabnedesA(pHoyidmeean)o.ptZeoroal,ogiCcharySscirdiopit-a region, draining into the Pacific Ocean, 28: 233-249. isolated from a more extensive and ecolog- Cambra. T. R. A. and D. Quintero, A. 1997. A revi- ically diverse Eastern region. The ancestor sion of Protopmotopsis Schuster (Hymenoptera: population of the present-day Euspinolia, Mutillidae). journal of Hymenoptera Research 6: 263-272. vaAinnadnaelsmtaoxssoltnop,eexswc.lausTshiveieslocylaotCmehdmioloennanatnahcneedstPaoPcreirfuio-cf Fritz,JA.rMt.J.roA.paon1dd9o9s8S..AMruCgtoeisnlctlaiirndooasne,.,PEpedd.isc4.i4o5nB-ei4so5d1Siuviren.:rsMiodrardondee, both Tallium and Limaytilla + Scaptodactyla Nixon, K. C. 2002. Winclada version 1.00.08. Published + AUotilla was relegated to the region east by the author, Ithaca, NY. ocfhatnhegeAsndberso.uTghhetclaibmoautticbayndtvheegetAantdioenaanl SchusgRteregari,pohnR..oMfII.I.t1hA9e49K.MeuCytoinlttloriidSbauuebtfiaoomnfisltitheoeswaRNreedportaersoMeponintcoea-dl uplift, particularly the desertification of and the descriptions of several New Genera the Chaco region, was the driving selective (Hymenoptera). Entomologica Americana (n.s.) 29: force that caused the diversification and 59-140. C evolutionary split between Tallium and the Spichiger, R., Calenge, and B. Bise. 2004. Geo- sister genera of AUotilla + Scaptodactyla + sgpreacpiheiscalchazroancatteriiosnticinofthetheNeoPtarroapgiucasy-oPfartarneae Limaytilla. Basin. Journal ofBiogeograpln/ 31: 1489-1501.