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Home , Acanthaclisinae, Acanthaclisini, Brachynemurini, Dendroleon, Dendroleontini, Dimarini

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Evolution and success of (: , Myrmeleontidae)

Mervyn W. MANSELL

Abstract: and hold the key to the unresolved higher classification of Myrmeleontidae. Additio- Myrmeleontidae comprise the largest nal information is also forthcoming from and most widespread of Neuroptera historical biogeography. Classifications, owing to their ability to exploit a wide ran- morphological adaptations, life histories, ge of habitats including sand. A psammo- strategies and distribution pat- philous existence was facilitated by sever- terns are reviewed and discussed as a con- al larval autapomorphies in the ground- tribution to elucidating relationships with- plan of Neuroptera that pre-adapted - lions to a life in sand and ensured their in the Myrmeleontidae. evolutionary success. The progression Key words: Myrmeleontidae, higher from arboreal habitats to psammophily classification, subfamilies, evolution, bio- may reflect the phylogeny of the family geography, biology, psammophily.

Stapfia 60. zugleich Kataloge des OÖ. Landesmuseums, Neue Folge Nr. 138 (1999), 49-58

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Introduction tion that set Neuroptera on an evolutionary course and engendered a remarkable of Myrmeleontidae are a highly evolved predatory . Enigmatically, this speciali- family of Neuroptera whose larvae have adop- sation was not restrictive, but resulted in the ted a variety of predation strategies that ena- radiation of Neuroptera into an impressive ble them to exploit a wide range of habitats array of morphologically and biologically relative to other families. This versatility has diverse taxa that comprise 17 families. It also ensured their evolutionary success as the lar- provided a larval autapomorphy to underpin gest and most widespread group, rivalled only the monophyly of Neuroptera, and established by , in the neuropteroid lineage. them as a highly effective functional group of There are currently about predators. 2000 described of ant- Myrmeleontidae, in particular, have lions distributed throughout exploited the specialised larval mouthparts the world, but especially in the that pre-adapted them to a variety of habitats, arid regions of the globe culminating in psammophily (Fig. 2). This (ASPÖCK H. et al. 1980). was the key to the evolutionary success of ant- Many of these species names lions. Occupation of different habitats enab- are synonyms, but they are led these „sit and wait" ambush predators to being offset by new discoveries develop many tactics to improve their effi- that will eventually yield a far ciency as predators. These range from lurking higher number of extant taxa. camouflaged on rocks and vegetation, in tree- The family also includes some holes, concealed in dust and sand, under rock of the largest and most striking overhangs, in burrows and ultimately oi all insects, with wingspans the ability to modify the immediate sandy ranging from 30-170 mm. environment into traps. Many have ornately patterned The abundance of antlions and their vor- wings (Fig. 1) that enhance acious feeding habits are an important influ- the and cryptic ence in the regulation of populations of gro- behaviour of these great in- und-dwelling and arboreal . In arid sects. areas such as the Kalahari savannah of sou- The range of habitats and thern Africa, the biomass of large predatory predation strategies reflect larvae of (Figs 3-5) and other tribes evolutionary trends within the could well exceed that of vertebrate predators. Myrmeleontidae, from plesio- Larval and adult antlions, in turn, are a morphic arboreal habitats to sustainable food resource for and the apomorphic occupation of vertebrates in natural ecosystems. Many spe- sand. This was facilitated by a cies are restricted to specific habitats and bio- unique morphological adapta- mes, and are sensitive to ecological disturbance, Fig. 1: speciosus (LINNAEUS), Western Cape Province, South Africa. Length = 70 mm

Fig. 2: of Crambomorphus MCLACHLAN, Kalahari Savannah, South Africa, displaying the psammophilous habit. Length = 30 mm.

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Fig. 3: Larva of Golafrus oneili (PERINGUEY), Kalahari Savannah, South Africa. Length = 35 mm. (Photo: A.S. Schoeman).

Fig. 4: Larva of Crambomorphus MCLACHLAN, Kalahari Savanna, South Africa. Length = 30 mm.

Fig. 5: Larva of Palpares annulatus STITZ, Kalahari Savannah, South Africa. Length = 25 mm. (Photo: R.G. Oberprieler). rendering them excellent bio-indicators of the attention of scientists since early times, environmental degradation. This is of particu- and are entrenched in folklore (WHEELER lar concern in South Africa where large-scale 1930, KEVAN 1992). Despite this, the global habitat fragmentation and application of systematics and phylogenetic relationships of pesticides threaten several endemic populati- Myrmeleontidae remain obscure owing to the ons such as the vulnerable Pamexis dynamic evolutionary state of the group and HAGEN (Fig. 6). Apart from their obvious bio- the consequent plethora of synonyms in the logical significance, Myrmeleontidae are ideal literature. These issues are currently being subjects for the study of behaviour, phy- addressed by several researchers, and some siology, biogeography and evolution, and is phylogenetic analyses have already been car- consequently a group that urgently warrants ried out (e.g. MANSELL 1992, STANGE 1994). study and conservation. Various authors have attempted to arrange Antlions and their habits have captured the Myrmeleontidae into subfamilies, tribes

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and subtribes, but no consensus has been rea- TlLLYARD (1916) recognized Banks' two subfa- ched, as most classifications lack a sound phy- milies and added two more tribes in studies on logenetic basis. The variety of lifestyles and the Australian fauna. EsBEN-PETERSEN (1918) biogeography of antlions may, however, reflect divided the Myrmeleontidae into two sec- evolutionary trends within the family that tions, Archaemyrmeleonida and Neomyrme- could contribute to resolving their higher clas- leonida, the former including only the Palpa- sification. The current paper consequently nni, and the latter comprising the Myrme- reviews information on supra-generic classifi- leontinae with four tribes and - cation, morphology, biology and historical tinae with six. BANKS (1927) raised the Palpa- rini and Macronemurini to subfamily status and added two more tribes, and subsequently- elevated the to subfamily level (BANKS 1943). More recent contributions have been tho- se of MARKL (1956) who classified the genera of the world into 23 tribes, adding 10 new tri- bes in the process. He did not mention subfa- milies in his treatise, but it had had the impor- tant effect of focussing attention to the higher classification of Myrmeleontidae, and provi- ding a basis for future discussions. STANGE (1961) mentioned three subfamilies, Acant- haclisinae, Macronemurinae and Dendroleon- tinae, but did not deal with or . In 1967 and 1970a, STANCE referred to four subfamilies, omitting Macro- nemurinae, and then (STANCE 1970b) synony- mized Dendroleontinae and Macronemurinae with Myrmeleontinae. HöLZEL (1969) esta- blished the tribe Isoleontini, and in 1970 he also included the tribes of Dendroleontinae in Myrmeleontinae. RlEK (1970) placed the Aus- Fig. 6: Pamexis karoo MANSELL, Karoo biogeography, particularly vicariance events, National Park, South Africa. Wing tralian Myrmeleontidae in four subfamilies, length = 34 mm as a contribution to elucidating the phylogeny Dendroleontinae, Macronemurinae, Acant- and classification of the family. haclisinae and Myrmeleontinae, but did not mention tribes. HöLZEL (1972) recognised three subfamilies, Palparinae, - tinae and Echtromyrmecinae, and STANGE (1976) listed , Palparinae and Higher classification of Myrmeleontinae. WILLMANN (I977) and Myrmeleontidae ASPÖCK H. et al. (1980) reduced the number of subfamilies to two, Palparinae and Myrme- Supra-generic classification of Myrme- leontinae, but the latter did not provide a tri- leontidae was initiated by BANKS (1899) who bal classification. NEW (1982) relegated the distinguished two groups, Myrmeleoni and family Stilbopterygidae to subfamily status, Dendroleoni. In his study of African antlions assigning Albardia VAN DER WEELE to the BANKS (1911) elevated these two groups to , and Stdbopteryx NEWMAN and the subfamilies Myrmeleoninae and Dendro- Aeropteryx RlEK to the subfamily Stilbop- leoninae, each comprising two tribes, Palpari- teryginae within Myrmeleontidae. At this sta- ni and Myrmeleonini and Dendroleoni and ge some consensus appeared to be emerging Nemoleonini respectively. regarding subfamilial classification, but none NAVAS (1912a, 1912b, 1913, 1914a, of these subfamilies was based on sound phy- 1914b, 1926) added eight more tribes, while

52 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at logenetic characters, and still require apprai- modification of the malpighian tubules for sal. The tribes were also subjectively defined production and cryptonephry (HENNIC 1981). and in considerable disarray, but recently Each of these attributes further enhanced the STANCE & MILLER (1990) and STANGE (1994) ability of antlions to thrive in hot, dry sandy have attempted to rationalise them by exami- habitats. ning larval characters. They listed three subfa- The mouthparts and feeding mechanism milies, Palparinae, Myrmeleontinae and Stilb- are unique to Neuroptera. When prey is cap- opteryginae, but were not able to substantiate tured, the sharp tips of the mandibles pierce the monophyly of any of these. STANGE & the cuticle and en:vmes are infused into the MlLLER (1990) enumerated nine tribes, and STANCE (1994) added a further two. The tri- Fig. 7: Jaw structure of Palpares inclemens (WAL- bes documented by these authors are general- KER), ventral aspect, sho- ly well defined and are recognised for the fol- wing grooved mandible lowing discussions. and lacinia of maxilla. Jaw length = 8 mm.

Morphology and Feeding

Autapomorphic mouthparts, comprising the elongated mandibles and maxillary lacinia that fit together along their entire lengths to form a piercing and sucking apparatus (Fig. 7), has been the key factor in the rise of Neurop- tera as a group of specialist predators. Adapta- tions derived from this attribute include: smooth curved mandibles in larvae of Chryso- pidae, , and Psy- chopsidae; short or long straight stylets in , , , Dila- ridae, and ; and the curved toothed jaws of (Myrmeleontidae, Ascalaphidae and Nymphi- dae). The straight mandibles and maxillae are apomorphic (ASPÖCK U. 1992), and curved body through the mandibular/maxillary jaws with teeth are another derivation from canals. This kills the prey and dissolves the the simple curved-jaw condition. Such attri- soft internal tissues, the resulting fluid being butes equipped neuropteran larvae for widely drawn up through the sucking tubes into the divergent predation strategies in a variety of alimentary canal of the larva. The food conse- habitats. These range from aquatic, semi- quently never comes into contact with the aquatic, arboreal, corticolous, detritis-dwel- substrate and is not contaminated with chitin, ling, under stones, tree-holes and sheltered sand or other indigestible debris that would rock ledges and caves, to inquilines in colo- accumulate from biting and chewing mouth- nies of , and wood-boring , parts. Sand grains would also impede a che- parasites in the nests of , culminating in wing motion and this is obviated, as is the psammophily in Myrmeleontidae and Nemop- need for a mouth opening that would become teridae. A psammophilous habit does not, filled with sand. The ingestion and digestion however, infer a close relationship between of food is consequently so efficient that the these two families (MAN'SELL 1996). midgut is sealed and no solid waste, only The mouthparts engendered further auta- excess fluid, is voided during larval life. This pomorphic characteristics in larval Neurop- exempts some of the malpighian tubules from tera, including loss of labial palps, mouth ope- an excretory role, enabling them to produce ning reduced to a shallow slit, a sealed midgut, silk for cocoon construction.

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The major constraint to life in hot, dry, SELL 1987). Many occupy sheltered dusty led- sandy conditions is desiccation, but the excre- ges, including Bankisus NAVÄS (MANSELL tory system of antlions prevents unnecessary unpubl.), Tricholeon ESBEN-PETERSEN (MAN- moisture loss through faeces, and the crypto- SELL 1988a) and some NAVÄS in nephridial malpighian tubes regulate the volu- southern Africa, and BANKS (STAN- me of fluid excreted. A silken cocoon, incor- GE &. MILLER 1990) in the New World. Larvae porating sand grains, is spun on the substrate of a few genera, Cymothales (C. spectabilis where the larva lived, or in sand. The cocoon ESBEN-PETERSEN, C. eccemros (WALKER)) provides a tough impenetrable shelter that (MANSELL unpubl.) and Elicura and Eremoleon protects the from parasites, predators, (STANGE & MILLER 1990, STANGE 1994) live excess soil moisture (rain) and desiccation, both in tree holes and on rock ledges. Bankisus and the sand grains provide effective camouf- larvae are found on rock ledges and on tree lage. Behavioural traits such as diurnal rhyth- roots under sandy overhangs (MANSELL unpu- ms that regulate movement through the sand, bl.). One of the most enigmatic larvae is that and larval coloration (Fig. 3) also protect of Navasoleon boliviana BANKS, from Bolivia, psammophilous antlions from excessive heat that lives upside down on bare rock on the and desiccation. These attributes have enab- ceilings of caves and has adapted its pupal led the great majority of antlions to exploit emergence procedure accordingly (MiLLER &. the limitless sandy environments of the world, STANGE 1985). This habit resembles that of Proctolyra TjEDER (Ascalaphidae) whose lar- leading to their radiation under the protective vae also live upside down but under stones in mantle of sand. the Karoo biome of South Africa (MANSELL unpubl.), and may emphasise the close relati- onship between Ascalaphidae and Myrme- Habitats of Myrmeleontidae leontidae. Eremoleon nigribasis Banks has pro- gressed further to living in animal burrows The progression of Myrmeleontidae can (MILLER 1990), a habit that could have been be traced through examples from a diversity of exploited by those adapted to living in cave lifestyles. These include: arboreal, living expo- mouths. This diversity of habitats reflects, sed on vegetation or in tree holes; exposed on even within a single genus, the versatility of rock surfaces; in fine dust and detritus under certain groups of antlions, but could also indi- rock overhangs and small caves; detritus in cate that they are less specialized than the animal burrows; freeliving in sand and psam- psammophilous species, by not being confined mophilous pit-builders. Larvae of Neulatus to one restricted habitat. They also retain NAVÄS live in Puya (Bromeliaceae) plants in several plesiomorphic traits such as abdominal Chile, while Jaffuelia NAVÄS live camouflaged scoli and slender jaws. on boulders (STANGE 1994). The larvae of these two genera resemble Ascalaphidae and Arboreal and freeliving larvae conform to may represent the archetypal mode of existen- the general groundplan of Neuroptera mani- ce of antlions. The close relationship of fest in other large families including Ascala- Ascalaphidae with Myrmeleontidae has fre- phidae, Chrysopidae, Hemerobiidae, Coniop- quently been emphasized (STANGE & MILLER terygidae, and, many other ins- 1990, STANGE 1994). Gnopholeontini larvae ects. In most insect orders true psammophily live fully exposed on rocks or tree bark and (freeliving, completely immersed in sand) is a rely on camouflage for protection (MILLER rare phenomenon that could be considered a 1990), similar to the modus operandi of derived condition. Beyond the Neuroptera, it Ascalaphidae. From there it would have been occurs mainly in Diptera and Coleoptera, a simple advance to living in tree holes and both highly evolved holometabolous orders. then to dust and detritus on rock ledges under The majority of tribes, Acanthaclisini, protective overhangs and detritus in animal , Palparidini, Palparini, Stilbop- burrows. Several genera inhabit tree holes, Eli- terygini, and most Myrmecae- cum NAVÄS (STANGE 1994), Dendroleon lurini and are psammophiles. BRAUER and Cymothales GERSTÄCKER (MAN- This could be considered an autapomorphy for

54 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at this section of Myrmeleontidae that supports that occurs throughout the world - on all con- the status of a subfamily Myrmeleontinae, tinents and islands that support Myrmeleont- with another other major comprised of idae. There are a number of closely related pit- non-psammophilous forms in the subfamily building genera that possess features indica- Dendroleontinae (MANSELL 1996). Morpho- ting a close relationship with this genus. The logical characters, including loss of scoli, distributions of Myrmeleon and its allies sug- development of fossorial appendages and setae gest that psammophily and pit-construction on abdominal segment 8, increased tooth evolved very early in the myrmeleontid linea- number in most Palparini, and the ability to ge, before the separation of the continents move rapidly backwards through sand (Fig. 2), (MANSELL 1996). The tribe Acanthaclisini is are commensurate with a sand-dwelling exi- not as widespread as Myrmeleon, but is stence and could be considered autapomor- nonetheless represented on all continents by phies as additional support for the Myrme- different genera. The tribe is comprised enti- leontinae. This subfamily has advanced furt- rely of psammophiles and is also an ancient her in being able to modify the immediate lineage, as indicated by the distributions of its sandy environment into pitfall traps to impro- constituent genera. Psammophily is not mani- ve their predatory skills. A pit-building habit, fest in , a tribe that occurs although a derived feature, is extremely throughout the world, although individuals ancient, but it further enhanced the success of are rare. This tribe is also an ancient lineage antlions, in their psammophilous habit (MAN- (STANGE & MILLER 1990, MANSELL 1996) whose members may manifest an early stage in SELL 1996). It is limited to the Myrmeleontini the transition from arboreal habits to psam- and a few genera in other tribes such as Myr- mophily. Dendroleontini have undergone a mecaelurini. There are several variations to great radiation in Australia, a continent that the pit-building adaptation that has been car- has long been separated, and a detailed study ried to extremes by the Australian antlion of this phylogenetically significant fauna is Callistoleon manselli NEW, 1993 (MANSELL urgently needed. The possibility that Stilbop- 1988b). Further characters, unsegmented tar- terygini, a relictual tribe confined to Austra- si, sensory setae for detection of vibrations, a lia, were precursors to psammophilous antli- highly flexible cervical region and eyes borne ons cannot be ignored. The larva of Stilbop- on protuberances (secondarily reduced in ceryx linearis NAVÄS from South Australia Myrmeleon and other pit-builders), are general resembles those of Ascalaphidae, including myrmeleontoid features that also pre-adapted well developed scoli, yet it has a psammo- myrmeleontid larvae for psammophily. philous habit (MCFARLAND 1968), and other The Dendroleontinae are not as clearly myrmeleontid traits are also manifest. Unfor- supported, although the long slender legs and tunately, only one larva of Stilbopterygini is opposable pretarsal claws in adults could be currently known, and it has not been compre- considered, along with slender form of the lar- hensively described. Myrmecaelurini appear val jaws. A closer study of the rich Australian to be confined to the Old World and are fauna would certainly reveal additional cha- psammophilies, with several pit-builders racters to clarify the status of this subfamily. (MANSELL 1996). Dimarini and Brachynemu- rini are psammophiles confined to the Ameri- cas, while Gnopholeontini and Lemolemini live mainly on rocks or are arboreal and are Biogeography confined to the New World. Nemoleontini occur throughout the world and manifest a The 12 tribes of Myrmeleontidae, as deli- wide range of larval habits. At present little mited by STANGE & MILLER (1990) and STAN- information can be gained from consideration GE (1994) manifest distinctive lifestyles that of this tribe, as it is certainly polyphyletic. were reviewed by MANSELL (1996), and distri- STANGE & MILLER (1990) include Palparidini butions that could provide further phylogene- in Palparini, but on adults characters the three tic information. The tribe Myrmeleontini species of Palparidius PERINGUEY, all endemic includes Myrmeleon LINNAEUS, the only genus © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at

to southern Africa, are closer to the Dimarini. Acknowledgement comprises three Afrotropical genera whose larvae remain unknown. The tribe Pal- I thank Prof. H. Aspöck, University of parini is clearly monophyletic and has radia- Vienna for inviting me to participate in this ted since the breakup of Gondwanaland project, and for his valuable comments on (MANSELL 1992, 1996). The protective cover- the manuscript. ing of sand has enabled palparines to evolve into some of the largest and most efficient psammophilous predators in the Insecta. The Zusammenfassung resultant large adults, in turn, had to adapt by developing cryptic coloration, resulting in the Die Myrmeleontiden stellen die größte magnificient wing patterns of all Palparini und am weitesten verbreitete Familie der that enhance the camouflage of these great Neuroptera dar; dies steht im Zusammen- insects which are certainly a recent myrme- leontoid innovation. hang damit, daß sie ein großes Spektrum von Habitaten - einschließlich Sand - The idea of two of Myrmeleontidae, besiedelt haben. Die Adaption an den Sand represented by the subfamilies Myrmeleon- wurde durch eine Reihe larvaler Autopor- tinae and Dendroleontinae is reiterated. mophien im Grundplan der Neuroptera Psammophily has been the key to success in ermöglicht und war letztlich das Erfolgsre- the majority of antlions, and can be used to zept in der Evolution der Myrmeleontiden. distinguish the two groups. Myrmeleontinae Der Übergang von arboralen Habitaten are well supported by apomorphic features, but this is not as clear in Dendroleontinae where zum Psammophilie spiegelt sich in der Phy- the larvae of many taxa are still unknown. logenie der Familie wider und ist mögli- The two clades were already well established cherweise der Schlüssel für das Verständ- before the breakup of the continents, as reflec- nis bisher ungelöster Fragen der Klassifika- ted by the distributions of Myrmeleontini, tion der Familie. Wichtige Information ist Acanthaclisini and Dendroleontini. These are zudem von biogeographischen Analysen the only tribes that occur in all the areas occu- von Verbreitungsmustern von Myrme- pied by antlions. Other tribes may have appea- leontiden zu erwarten. Konzepte der Klas- red after the commencement of continental sifikation, morphologische Anpassungen, separation, as manifest by the radiation of Pal- Entwicklungszyklen, Beutefangstrategien parini on the African continent. The current und Verbreitungsbilder von Myrmeleont- distribution of Palparini, and their vicariance iden werden besprochen und im Lichte der biogeogaphy were discussed by MANSELL Aufklärung von Verwandtschaftsbeziehun- (1996). gen diskutiert.

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HÖLZEL H. (1972): Die Neuropteren Vorderasiens. IV. NAVAS L. (1913): Bemerkungen über die Neuropteren Myrmeleonidae. — Beiträge zur Naturkundli- der Zoologischen Staatssammlung in München. chen Forschung in Südwestdeutschland 1: 3- V. — Mitteilungen Münchener Entomologi- 103. schen Gesellschaft 4: 9-15. KEVAN D.K.MCE. (1992): Antlion ante Linne: Myp\iJ\- NAVAS L. (1914a): Insectes Nevropteres, Planipennia KOXCOV to Myrmeleon (Insecta: Neuroptera: Myr- et . In: Voyage de Ch. Alluaud et R. meleonidae). In: CANARD M., ASPÖCK H. & M.W. Jeannel en Afrique Orientale (1911-1912). 1-52. MANSELL (Eds), Current Research in Neuroptero- — Librairie Albert Schultz, Paris. logy. — Proceedings of the Fourth International Symposium on Neuropterology. Bagneres-de- NAVAS L. (1914b): Neuröpterosnuevos o poco conoci- Luchon, France, 1991. 203-232. Toulouse. dos (Segunda Serie) Memorias de la real Acade- mia de Ciencias y Artes de Barcelona 11: 105- MANSELL M.W. (1987): The ant-lions of southern Afri- 119. ca (Neuroptera: Myrmeleontidae): genus Cymo- thales GERSTAECKER, including extralimital spe- NAVÄS L. (1926): Nevropteres d'Egypte et de Palesti- cies. — Systematic 12: 181-219. ne. II. Familie des Myrmeleonides. — Bulletin de la Societe Royale Entomologique d'Egypte MANSELL M.W. (1988a): The Myrmeleontidae (Neur- 1926: 26-62. optera) of southern Africa: genus Tricholeon ESBEN-PETERSEN. — Neuroptera International 5: NEW T.R. (1982): A reappraisal of the status of the 45-55. Stilbopterygidae (Neuroptera: Myrmeleon-

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Author's address: Dr Mervyn W. MANSELL ARC-Plant Protection Research Institute Private Bag X134 Pretoria 0001 South Africa E-mail [email protected] Fax 27 012 325 6998

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