Rev. Biol. Trop. 50(2): 477-484, 2002 www.ucr.ac.cr www.ots.ac.cr www.ots.duke.edu INVITED PAPER

MINIREVIEW

Thysanoptera in the Neotropics

Laurence A. Mound CSIRO Entomology, GPO Box 1700, Canberra ACT, Australia. e-mail: [email protected]

Received 20-X-2001. Corrected 12-III-2002. Accepted 14-III-2002.

Abstract: It is suggested that descriptive of must be integrated into biological studies if we are to understand patterns of evolutionary and ecological diversity. Collecting and describing new taxa is easy, but understanding their position in ecosystems and how they have contributed to the origin and maintenance of bio- logical diversity is more important yet more difficult. Many authors fail to appreciate that individual thrips species are commonly highly polymorphic, both within and between sexes, with the result that 20% of species names and 30% of generic names are currently placed into synonymy. The biological significance of such poly- morphism has been little studied, but the presence of large and small males in a species is presumed to indicate some form of male/male competition for resources; this is particularly common in fungus feeding species. Amongst phytophagous species, the recognition of the host on which thrips actually breed is a prerequi- site to understanding patterns of diversity, some thrips lineages being associated with particular groups of plants whereas others exploit a diverse range of plants. Attempts to understand the diversity of thrips, including the application of cladistic methods, are severely limited by the lack of studies on the biology of individual species, although thrips exhibit a wide range of interesting biological phenomena, including various levels of sociality, gall-induction, specific pollination associations, virus transmission, and ectoparasitism.

Key words: taxonomy, morpho-species, polymorphism, collaborative studies.

Biological diversity can be considered in Thysanoptera are particularly diverse in two different ways. Taxonomists consider the the Neotropics, with considerably more than numbers and distributions of taxa, whereas 2000 species existing in this Region (Mound ecologists consider the many ways in which and Marullo 1996). These species feed on a taxa depend on each other Ð their biological variety of substrates, up to 50% being and behavioural relationships. Taxonomy is mycophagous either on spores or on hyphae, a thus seen as being limited to ‘product’, the large number feeding primarily in flowers, publication of descriptions, whereas most rather fewer feeding only on leaves, even on research biologists are concerned essentially mosses and ferns, with a few species predato- with ‘process’, how organisms live. If we are ry. In addition to this wide range of feeding to develop an understanding of the functioning habits, the biologies exhibited by these and origins of the rich biological diversity of include various levels of sociality, remarkable the Neotropics, we need to explore the com- structural polymorphisms within and between mon ground between product and process. sexes, gall-induction on leaves, specific polli- This article is a brief review of the current state nation associations, and virus transmission on of our knowledge of the order crop plants. One recent study has even demon- Thysanoptera, the thrips, considering both tax- strated ectoparasitism by a thrips species on an onomic products and biological processes, Homopteran (Izzo et al. 2002). Despite this with particular emphasis on the Neotropical range of interesting topics, the vast majority of region. studies on thrips in the Neotropics have been 478 REVISTA DE BIOLOGêA TROPICAL limited either to insecticide trials or to descrip- tions. Thus the number of taxon descriptions is tive taxonomy. not a good measure of our knowledge of bio- logical diversity, particularly when, as in some groups, up to 50% of names fall as synonyms, OBJECTIVES OF TAXONOMY and high synonymy rates remain a modern methodological, not merely an historical, arte- Descriptive taxonomy is essentially sub- fact (Gaston and Mound 1993). jective, museum taxonomists rarely testing the In contrast, an apparently similar ques- veracity of the hypotheses they erect, these tion, “Why are there so many species of being the new species that they describe insects?” resonates through most biological (Gaston and Mound 1993). As a result, many disciplines. It involves comparative studies biology students avoid taxonomy, even when between species and the ways in which they recognizing that biological observations need share available resources. It involves species to be viewed in a comparative, evolutionary, turnover between localities (Bartlett et al. framework. This has led to an increasing lack 1999), habitats and seasons, and thus requires of taxonomic specialists over the last 30 years, information on how species disperse, and how and this in turn limits the range of biodiversity they exploit ecosystems that are in a constant questions that can be asked. For example, in state of flux. This one question thus subsumes tropical forests, neither the ecology of tree so many other questions that are fundamental flowers, nor the dynamics of leaf-litter sys- to understanding how biological diversity has tems, can be studied in depth, due to the lack arisen, and how we will conserve it. The ques- of taxonomic expertise for many of the insect tion “Why?” is about process and opens up groups involved. Molecular technology is new avenues of thought; the question “How broadening the interest in taxonomy, such tools many?” is about product and, by itself, is of being seen to provide more objective methods more political than scientific interest. for examining relationships between taxa. But the lack of defined objectives for descriptive taxonomy remains a problem, both for attract- THRIPS SPECIES-LEVEL ing new students and for funding the long-term TAXONOMY AND BIOLOGY employment of specialists. The lack of clarity in taxonomic objec- Almost 5 500 species of Thysanoptera are tives is curious (Mound 1998). The question currently considered valid worldwide, in nine “How many species of insects are there?” is families and almost 750 genera. Judging from deceptively attractive, and has received con- material in the major museum collections siderable attention in recent years. Suggestions there are probably about 10 000 thrips species have been made that funding be sought to in the world. The total number of described describe all of the world’s taxa (Anonymous species from Central and South America is 1994). Even if we assume that a precise answer about 1 600 (Mound and Marullo 1996), and to this question is possible, the use that could more than 700 species of thrips have been be made of that answer to generate further sci- described from Brazil. This latter figure sug- entific questions is not clear. Probably as many gests that the thrips fauna of Brazil is quite as 50% of named insect species are based on well known, in that a good proportion of the single samples, even single individuals, with species have been given names. But few of little known either of their biology or their these species can be recognized from the litera- variation. Moreover, probably 5% of named ture, and little is known about how most of insect species are known only from unrecog- them live, beyond that some are probably fun- nizable, although not necessarily old, descrip- gus-feeding, or grass-feeding or flower-feeding. INTERNATIONAL JOURNAL OF TROPICAL BIOLOGY AND CONSERVATION 479

Problems in descriptive taxonomy Mexican biodiversity, particularly the inter-rela- tionships of this important tree crop with the Describing large numbers of species is not native flora and fauna. In this instance, each simple. For example, more than 150 species ‘new species’ was defined on trivial structural are recognized in the Frankliniella features that are known to be highly variable (Nakahara 1997), 120 of these being known within several pest species of Scirtothrips. In from the Neotropics. However, in the collec- indicating that such taxonomic conclusions are tions of the Natural History Museum, London, unreliable, Mound and Strassen (2001) suggest- and the US National Museum, Washington, ed that taxonomic decisions are sometimes too there are several thousand microscope slides of important to other biologists to be left solely to unidentified Frankliniella species from the descriptive taxonomists. South American Cordillera. Most of these Some correlation presumably exists unidentified Frankliniella cannot be sorted sat- between floristic diversity and the diversity of isfactorily even to putative ‘morpho-species’, -feeding thrips. But host plant exploita- because we are unable to distinguish securely tion by thrips ranges from strict monophagy to between patterns of intra- and inter-population extensive polyphagy, often within genera. variation. For many, even males and females Thus two species of Echinothrips are known to cannot be associated satisfactorily, due to be strict monophages, on Tsuga and on colour differences between sexes and structur- Selaginella, whereas a third member of the al differences between large and small individ- genus is a pest on a wide range of plants in uals. It is not unusual to find adults of two or greenhouses (Collins 1998). In the genera more species of Frankliniella in the same set Frankliniella and Scirtothrips, a few species of flowers. To establish the intraspecific varia- are similarly monophagous, although both tion, and the true host plant relationships of a genera include some of the most extreme reasonable number of species to provide a sat- polyphagous pest thrips. This unpredictable isfactory base-line, requires extensive field- pattern of host exploitation clearly increases work. Such field studies need to take into the difficulty of distinguishing intra- from account the behaviour of wind dispersed thrips inter-specific variation in museum specimens that can be found resting, sometimes in large in the absence of careful field studies. numbers and in mixed species-assemblages, on Problems of species recognition occur in plants that are not used for larval development most large genera. Fungus-feeding thrips in the (Mound and Marullo 1996: 17). genus Hoplothrips show remarkable polymor- A facile response to such real biological phism, both within and between sexes. Males problems is to describe as ‘new species’ each of and females can look so different from each the structural variants that can be observed, even other that they may be placed in different gen- in the absence of any correlated biological data. era, and extensive population samples are One recent example of considerable economic needed to establish ranges of intraspecific vari- importance concerns the leaf-feeding species of ation. Moreover, any attempt to apply one of the genus Scirtothrips. In Mexico, Johansen and the 35 species names that are available in the Mojica-Guzman (1999) recorded 21 species of genus from Central and South America Scirtothrips from Mangifera, and described 18 requires access to the collections of J.D. Hood of these as ‘new species’. However, Mangifera at the U.S. National Museum, Washington, is not native to the Americas, hence each of because no identification keys have been pub- these thrips presumably has a native host plant lished. Thus there are serious scientific, as well within Mexico from which it invades cultivated as technical, problems for anyone in the mango trees. The mere description of ‘new Neotropics wishing to understand thrips taxon- species’ tells us nothing about the functioning of omy and biology. 480 REVISTA DE BIOLOGêA TROPICAL

Research potential (Mound and Marullo 1996). Such typological taxonomy contributes little to our understanding More positively, the problems indicated of biodiversity. above can viewed as offering outstanding These problems raise once again the ques- opportunities for anyone with serious interests tion of research objectives. Much taxonomic in biosystematics. The polymorphism and descriptive work is essentially haphazard, taxa structural allometry that is so common in the being described as they become available, often species of Hoplothrips, as well as several other for no better reason than that an author wants genera of fungus feeding thrips, is known to be named specimens in a museum collection. In associated with competition for resources, contrast, description of new taxa is best carried including male/male combat (Crespi 1986a, b). out when this extends knowledge in some spe- Fungus-feeding thrips could provide an excel- cific way, be this of structural variation, host lent tool for investigating various ecological plant relationships, or geographical range; also, and behavioural phenomena in the Neotropics. it is necessary at times to provide a valid name Kiester and Strates (1984) described remark- for a species on which particular observations able sub-social behaviour in one large species are being made by other biologists. A contrast- that lives on the trunks of Gustavia trees in ing approach to the sequential description of Panama feeding on the fungal spores of a new taxa, as a discipline isolated from the rest lichen species. The adults of this thrips can be of biology, is a research programme in Australia observed leading out parties of immatures to focussed on thrips associated with Acacia trees particular feeding sites each morning, appar- (Crespi and Mound 1997). This programme ently along pheromone trails, and then leading involves behavioural and host relationship stud- them back again in the evening to a home ies, as well as morpho- and molecular-taxo- crevice in the bark. nomic studies (Mound and Morris 2001, Morris Even greater opportunities for innovative et al. 2002). In Costa Rica, the ALAS biological studies are provided by the species- ( of La Selva) Project is another rich genus Elaphrothrips. At times, three or approach that incorporates taxonomy within the more species of this genus can be found on a broader objectives of field biology. Research single tree, feeding on fungal spores in bunches programmes with similar broad objectives of dead leaves. But it is not known if two or could be devised on Neotropical thrips, such as more species share the same dead leaf, or if they the diversity of fungus-feeding species on dead feed on spores of the same fungal species. That leaves and twigs, or the inhabitants of the flow- is, nothing is known of how such thrips compete ers of particular plant families in which particu- for available resources. All Elaphrothrips lar groups of thrips are known to be common, species exhibit variation in male size and body such as and Heterothripidae, or armature, from which it can be concluded that montane Asteraceae and the Frankliniella min- they indulge in male/male competition, but uta group. nothing is known for neotropical species of A major area for research, still largely interactions between either con-specific or con- ignored, is the relationship between thrips and generic individuals occupying the same habitat. flowers, whether as pollinators or as pollen The only recent paper on fungus-feeding thrips predators. In neotropical forests an intriguing in the Neotropics (Johansen and Retana 1999), problem is the large number of thrips adults described four new species on a total of seven that can be found in the fallen flowers of specimens, all taken in traps. The authors thus Bignoniaceae trees, these flowers falling daily provided no supporting biological evidence for in large numbers from the canopy to the forest their ‘new species’, and the character states used floor. Presumably the adult thrips fly back into to differentiate the species are known to be vari- the tree canopy to oviposit later in the day, able within populations of related species because their larvae are not usually found in INTERNATIONAL JOURNAL OF TROPICAL BIOLOGY AND CONSERVATION 481 such fallen flowers. However, no life history and genus levels (Mound and Marullo 1996). studies have been attempted on these common Relationships between the 250 genera of thrips, nor is their significance in floral biolo- are particularly difficult to gy understood. evaluate. The formal suprageneric classifica- The significance of thrips as pollinators tions that have been proposed (Priesner 1960), has been largely ignored in the Neotropics, although commonly quoted, are a poor reflec- although recent studies have demonstrated that tion of relationships. In contrast, the three Frankliniella diversa is probably a host specif- informal ‘lineages’ indicated by Stannard ic pollinator of Castilla elastica (Moraceae) (1957), and further developed by Mound and (Sakai 2001), and Del Claro et al. (1997) Marullo (1996), are operationally ineffective. examined the co-existence of ants and a Similarly amongst the , although species of in the flowers of many authors quote Tribal names, these Tribes Peixotoa tomentosa (Malpighiaceae). Thrips and Sub-tribes are not defined satisfactorily. are not easy insects to work with, but two recent studies on the inter-dependence of a Problems in thrips systematics plant species and a host-specific thrips have demonstrated how rewarding such work can be; Intra-specific polymorphism has been Macaranga flowers in south east Asia (Moog et mentioned above as an operational problem for al. 2002), and Macrozamia cycads in Australia species recognition, but such variation also (Terry 2001). Another interaction between causes problems in establishing generic classi- thrips and plants that has been little studied in fications. For example, species of Liothrips the Neotropics is gall-induction (Mound and usually have a single pair of setae on the head Kranz 1997), although this habit is known in behind the eyes, and males have one pair of several species of the genus Holopothrips (see setae on the ninth abdominal tergite much Mound and Marullo 1996: 290). shorter and stouter than the other setae. In con- trast, several Neotropical species, otherwise similar to Liothrips in structure and biology, THRIPS SYSTEMATICS have been placed in a genus Pseudophilothrips AND RADIATION because they have two pairs of long setae on the head, and the males have all the setae on Systematics involves the creation of a the ninth tergite elongate. In studying the biol- hierarchical classification that reflects the pre- ogy of one host-specific species that lives on sumed phylogeny of a group. One indication of Didymopanax, Del Claro and Mound (1996) the weak state of thrips systematics is the recognized that both of these character states unusually high number of monobasic genera. vary within populations. This is not the only Worldwide in the , 20 of the 36 gen- related species in which these character states era of , and 100 of the 200 vary, thus suggesting that Pseudophilothrips genera of Thripinae, each includes only a sin- is, at best, merely a paraphyletic subset of gle species. Similarly, of the 400 genera of Neotropical species within the worldwide more than 200 each includes genus Liothrips. Again, the absence of data on only one species. Such a classification is of the host plants of so many of these presumably limited use to other biologists, because it gives host-specific species severely limits any con- no indication of the phylogenetic relationships sideration of the patterns of variation and radi- between species. ation in this genus. The ineffectiveness of a classification The genus Holopothrips provides an comprising so many monotypic genera is par- instructive example of problems in the genus ticularly important because of the lack of any level classification of Neotropical phylogenetic structure between the sub-family Phlaeothripinae. Mound and Marullo (1996) 482 REVISTA DE BIOLOGêA TROPICAL recognized 31 species in this genus, but placed morphology. In one of the few applications of five genera as synonyms. However, not all the cladistics to thrips in the Neotropics, Retana species exhibit the full suite of character states (1998) prepared a data matrix for 24 character that define this genus, and the genus is thus states of 25 species of the genus Frankliniella. defined polythetically (Gauld and Mound But in claiming that particular character states 1982). The alternative is to recognize more were apomorphic for particular groups of than 10, mainly monotypic, genera within this species, this author made little allowance for lineage and thus obscure any relationships homoplasy. For example, several members of between the species. From the biodiversity this genus that breed solely on grasses have the viewpoint, the most interesting question posed head slightly prolonged in front of the eyes. by this suite of closely related species is their Although this can be interpreted as an apomor- level of host specificity in exploiting the avail- phy for an “F. tenuicornis group”, it is equally able flora. In the absence of good field studies, likely to be a convergent adaptation to living this remains unexplored, despite the light that on grasses, given that it is so common in grass it could shed on phylogenetic relationships thrips of other genera of Thripidae. Similarly, within this uniquely Neotropical lineage. any attempt to define a genus Exophthalmothrips from Frankliniella on the New methods basis that some of the facets of the compound eyes are enlarged must specify precisely which Molecular tools are giving us new meth- ommatidia involved. Clear definition of char- ods of investigating evolutionary relationships, acter states is the basis of sound cladistics, and these techniques sometimes indicate because subtle distinctions in how such states remarkable structural variation within lineag- are defined can lead to very different conclu- es. For example, Bhatti (1992) erected a mono- sions (Mound et al. 2001). Without clear defi- typic family for a remarkable Australian genus nitions of how character states are evaluated Xaniothrips. However, not only is this consid- and scored, mathematically sophisticated ered an unsatisfactory assessment of the mor- cladistic analyses (Retana and Soto-Rodríguez phological evidence (Mound and Morris 2001) remain of limited use in analysing evo- 1999), new molecular data (Morris et al. 2002) lutionary patterns. indicate that Xaniothrips is closely related to Koptothrips, a genus of very different looking species. These two genera comprise kleptopar- FUTURE STUDIES asitic thrips whose differing methods of invad- ing the domiciles of their host species have led, The Neotropics offer biologists a richly presumably, to a great divergence in their body endowed laboratory for investigating evolu- structure. Such a lack of congruence between tionary products and processes. The twin aims morphological and molecular data poses a of taxonomy Ð the recognition and naming of problem for traditional descriptive taxonomy, species, and the creation of predictive system- although parsimony applied to the full data set atic classifications Ð provide a service to the can provide a practical guide. rest of science by facilitating interdisciplinary Parsimony, in the form of cladistics, has communication about organisms. Through its been little used in studies on thrips classifica- emphasis on evolutionary relationships, taxon- tion, primarily because of operational difficul- omy contributes directly to that ultimate objec- ties associated with homoplasy in structural tive of all biological studies Ð understanding character states (Gauld and Mound 1982). the origins and maintenance of biological Cladistics is certainly an important tool with diversity. Viewed in this way, taxonomy and which to handle variation in a rational manner. systematics have much broader objectives than But cladistic methods must be based on sound the mere description and cataloguing of taxa. INTERNATIONAL JOURNAL OF TROPICAL BIOLOGY AND CONSERVATION 483

However, to achieve such objectives taxonom- Bhatti, J.S. 1992. The Order Tubulifera (Insecta): Its char- ic studies must involve good field biology in acters and classification into families. Zool. J. Pure order to recognize structural and biological Appl. Zool. 3: 127-162. variation, and where possible must be support- Collins, D.W. 1998. Recent interceptions of Echinothrips ed by suitable molecular studies to distinguish americanus Morgan (Thysanoptera, Thripidae) species and to recognize relationships. Such an imported into England. Entomol. Month. Mag. 134: approach requires that taxonomists are viewed, 1-4. organized and funded as collaborators within multidisciplinary research projects, rather than Crespi, B. 1986a. Territoriality and fighting in a colonial thrips, Hoplothrips pedicularius, and sexual dimor- as isolated individuals working to their own, phism in Thysanoptera. Ecol. Entomol. 11: 119-130. frequently ill-defined, agendas. Crespi, B. 1986b. Size assessment and alternative fighting tactics in Elaphrothrips tuberculatus (Insecta: RESUMEN Thysanoptera). Anim. Beha. 34: 1324-1335.

Se ha sugerido que la taxonomía descriptiva de los ti- Crespi, B. & L.A. Mound. 1997. Ecology and evolution of sanopteros (trips) debe integrarse dentro de los estudios bio- social behaviour among Australian gall thrips and lógicos si queremos ser capaces de entender los patrones de their allies. pp. 166-180. In J. Choe & B.J. Crespi. diversidad evolutiva y ecológica. Recolectar y describir (eds.). Evolution of Social Behaviour in Insects and nuevos datos es fácil, pero entender su posición en los eco- Arachnids. Cambridge University. sistemas y como ellos contribuyen al origen y mantenimien- to de la diversidad biológica es más importante y aún más Del Claro, K. & L.A. Mound. 1996. Phenology and difícil. Muchos autores han fallado al apreciar que es común description of a new species of Liothrips que las especies individuales de trips son altamente polimór- (Thysanoptera; Phlaeothripidae) from Didymopanax ficas, tanto dentro como entre sexos, con el resultado de que (Araliaceae) in Brazilian cerrado. Rev. Biol. Trop. el 20% de los nombres específicos y el 30% de los genéri- 44: 193-197. cos son actualmente considerados como sinonimias. El sig- nificado biológico de tal polimorfimo ha sido poco estudia- Del Claro, K., R. Marullo & L.A. Mound. 1997. A new do, pero se presume que la presencia de machos grandes y Brazilian species of Heterothrips (Insecta, pequeños en una especie indica alguna forma de competen- Thysanoptera), coexisting with ants in the flowers of cia entre machos por los recursos; lo cual es muy común en Peixotoa tomentosa (Malpighiaceae). J. Nat. Hist. las especies que se alimentan de hongos. Dentro de las es- 31: 1307-1312. pecies fitófagas, el reconocimiento de la planta hospedera sobre la cual los trips realmente se crían es un prerequisito Gaston, K.J. & L.A. Mound. 1993. Taxonomy, hypothesis- para entender los patrones de diversidad, algunos linajes de testing and the biodiversity crisis. Proc. Roy. Soc., tisanopteros están asociados con grupos particulares de London B251: 139-142. plantas mientras que otros utilizan un diverso ámbito de plantas. Los intentos de entender la diversidad de los trips, Gauld, I.D. & L.A. Mound. 1982. Homoplasy and the incluyendo la aplicación de métodos cladistas, están limita- delineation of holophyletic genera in some insect dos severamente por la carencia de estudios sobre la biolo- groups. Syst. Entomol. 7: 73-86. gía de especies individuales, aunque los trips exhiben un amplio ámbito de fenómenos interesantes biológicos, que Izzo, T.J., S.M.J. Pinent & L.A. Mound. 2002. Aulacoth- incluye varios niveles de comportamiento social, inducción rips dictyotus (Heterothripidae), the first ectoparasi- de irritación, asociaciones específicas de polinización, trans- tic thrips (Thysanoptera). Florida Entomol. 85 (1). misión de virus y ectoparasitismo. Johansen, R.M. & A. Mojica-Guzman. 1999. The genus Scirtothrips Shull, 1909 (Thysanoptera: Thripidae, REFERENCES Sericothripini), in Mexico. Folia Entomol. Mexicana 104: 23-108. Anonymous. 1994. Charting the Biosphere. Systematics Agenda 2000. Technical Report. Johansen, R.M. & A.P. Retana Salazar. 1999. Especies nuevas de Thysanoptera (Insecta: Phlaeothripidae) Bartlett, R., J. Pickering, I.D. Gauld, & D. Windsor. 1999. de Costa Rica. Brenesia 51: 59-67. Estimating global biodiversity: tropical beetles and wasps send different signals. Ecol. Entomol., 24: Kiester, A.R. & E. Strates. 1984. Social behaviour in a 118-121. thrips from Panama. J. Natur. Hist. 18: 303-314. 484 REVISTA DE BIOLOGêA TROPICAL

Moog, U., B. Fiala, W. Federle & U. Maschwitz. 2002. Mound L.A. & D.C. Morris. 2001. Domicile constructing Thrips pollination of the dioecious ant plant phlaeothripine Thysanoptera from Acacia phyllodes Macaranga hullettii (Euphorbiaceae) in Southeast in Australia. Syst. Entomol. 26: October 2001. Asia. Amer. J. Bot. [in press] Mound L.A. & R. zur Strassen. 2001. The genus Morris, D.C., M.P. Schwarz, S.J.B. Cooper & L.A. Scirtothrips (Thysanoptera: Thripidae) in Mexico: a Mound. 2002. Phylogenetics of Australian Acacia critique of the review by Johansen & Mojica- thrips: the evolution of behaviour and ecology. Mol. Guzmán (1998). Folia Entomol. Mex. 40(1): 133- Phylog. Evolut. (in press). 142.

Mound, L.A. 1977. Species diversity and the systematics of Nakahara, S. 1997. Annotated list of the Frankliniella some New World leaf-litter Thysanoptera (Phlaeothri- species of the world (Thysanoptera: Thripidae). pinae; Glyptothripini). Syst. Entomol. 2: 225-244. Contributions on Entomol. Int. 2: 355-389.

Mound, L.A. 1998. Insect taxonomy in species-rich coun- Priesner, H. 1960. Das System der Tubulifera (Thysa- tries - The way forward? Ann. Soc. Entomol. Brasil noptera). Anzeiger der math.-naturwiss. Klasse der 27: 1-8. österreischen Akad. Wiss. 1960: 283-296.

Mound, L.A. & B. Kranz. 1997. Thysanoptera and Plant Retana, A.P. 1998. Una vision filogenetica de Galls: Towards a Research Programme. pp. 11-24. In Frankliniella (Thysanoptera: Thripidae). Rev. Biol. A. Raman. (ed.). Ecology and Evolution of Plant- Trop. 4: 397-406. feeding Insects in Natural and Man-made Environments. National Institute of Ecology, New Retana, A.P. & G.A. Soto-Rodriguez. 2002. Filogenia del Delhi; Backhuys Publishers, Leiden, Netherlands. Genero neotropical Zeugmatothrips (Thysanoptera: Phlaeothripidae). Gayana 65(2): 119-128. Mound L.A. & R. Marullo. 1996. The Thrips of Central and South America: An Introduction. Mem. Sakai, S. 2001. Thrips pollination of androdioecious Entomol., International 6: 1-488. Castilla elastica (Moraceae) in a seasonal tropical forest. Amer. J. Bot. 88: 1527. Mound L.A., R. Marullo & J.W.H. Trueman. 2001. The greenhouse thrips, haemorrhoidalis, and Stannard, L.J. 1957. The phylogeny and classification of its generic relationships within the sub-family the North American genera of the sub-order Panchaetothripinae (Thysanoptera; Thripidae). J. Tubulifera (Thysanoptera). Illinois Biol. Monogr. Insect Syst. Evolut. 32: 1-12. 25: 1-200.

Mound L.A. & D.C. Morris. 1999. Abdominal armature in Xa- Terry, I. 2001. Thrips and weevils as dual, specialist pol- niothrips species (Thysanoptera; Phlaeothripidae), klep- linators of the Australian Cycad Macrozamia com- toparasites of domicile-producing thrips on Australian munis (Zamiaceae). Int. J. Plant Sci. 162: 1293- Acacia trees. Australian J. Entomol. 38: 179-188. 1305.