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Mtodos en Ecologa y Sistemtica ISSN impreso 1659-2182 ISSN digital 1659-3049 Revisionary paper

Vol. 7(2): 16
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Characters in Thysanoptera

AXEL P. RETANA-SALAZAR* Programa Universitario de Biologa Aplicada (PUA), Centro de Investigacin en Estructuras Microscpicas (CIEMic), Ciudad de la Investigacin, Universidad de Costa Rica, 2060.

RESUMEN: Thysanoptera es un grupo de alta homoplasia como ha sido establecido en la literatura por mltiples autores. Esto ha tenido un efecto en la discusin entre los taxnomos que invierten grandes cantidades de tiempo en la discusin de la validez de los caracteres utilizados en este grupo de insectos. Esto se refleja en mltiples trabajos de taxonoma en los que se describen especies y grupos sin una fundamentacin filogentica. De igual forma decisiones tomadas en funcin de resultados filogenticos son muchas veces cuestionados ya que los mismos van en contra de una clasificacin ms cmoda. En este trabajo se presenta una revisin corta de las tendencias y problemas en los diferentes tipos de caracteres que se utilizan en Thysanoptera en la actualidad y la necesidad de tener mayor cuidado con el trato de algunos de estos caracteres en la toma de decisiones taxonmicas, en las cuales se proponen cambios en la clasificacin biolgica. PALABRAS CLAVE: Filogenia, caracteres moleculares, caracteres morfolgicos, caracteres biolgicos, caracteres teratolgicos, cladstica., Thysanoptera, Thripidae. ABSTRACT: Thysanoptera is a group of high homoplasy as has been established in the literature by many authors. This has had an effect on the debate among taxonomists investing large amounts of time discussing the validity of the characters used in this group of insects. This is reflected in many taxonomic works, described species and phylogenetic groups without a foundation. Similarly decisions based on phylogenetic results are often questioned since they go against a more comfortable classification. This paper presents a short review of trends and issues in different types of characters used in Thysanoptera today and the need for a more careful treatment of some of these characters in taxonomic decisions in which changes in the biological classification are proposed. KEY WORDS: Phylogeny, molecular characters, morphological characters, biological characters, teratological characters, cladistics., Thysanoptera, Thripidae.

INTRODUCTION Thysanoptera is a complex group especially because of its high level of homoplasy (Gauld & Mound 1982, Mound & Palmer 1983, RetanaSalazar 1998, 2000). However it is one of the groups with fewer amounts of formal phylogenetic analysis and with a very common practice of -taxonomy of the specialists that
Autor Correspondencia: apretana@gmail.com*

are more occupied in the description of new species than in the determination of the natural groups. The recognized authorities focused in the taxonomy producing several hundreds of pages per year full of subjective criteria about the importance of the characters used by each author. This practice was abandoned in the

Mtodos en Ecologa y Sistemtica ISSN impreso 1659-2182 ISSN digital 1659-3049 majority of the groups, where several authorities dont consider the publication of species description in absence of good phylogenetic analysis and revisionary works as good exercise (Retana-Salazar 1998, 2000, 2006, RetanaSalazar & Ramrez Morales 2006). Before the last 20 years only Johansen in several papers (Johansen 1982, Johansen 1983a,b, 1986a,b) approached the topic of the use of characters to rigorous analysis that show its validity in taxonomic decisions. Some other works have been strongest criticised for the choice of the characters related with no correct the hierarchical level or for the publication of some obscure cladistic results (Bhatti 1993). In several groups the comparative study of the characters in species that are close together is necessary to establish the possible change of the states of characters, but that is not enough for the determination of the evolutionary development of each character (Wiley 1981, Retana-Salazar 2006). In the last years, important papers about molecular data in Thysanoptera have appeared. One of the most important is the publication of Morris and Mound (2005). Other researchers used morphological features as principal characters, but the use of morphological characters may be dangerous if the importance of the character is not correctly evaluated as good characters in phylogenetic resolution (Bjrklund 1997, Diniz Filho 2000, RetanaSalazar & Retana-Salazar 2008). In these cases the subjective consideration of the possible evolutionary origin of the characters is a dangerous practice that shows the a priori ideas of the taxonomist more than the real evolutionary change. SYSTEMATIC PHYLOGENETICS IN THYSANOPTERA

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Mound and collaborators (1980) presented a phylogenetic paper concerning the family classification in Thysanoptera. This work has some methodological problems that prevent the repetition of these results, especially the absence of the type of analysis formally used, i.e. some details like the selected software, the polarization criterion of characters, the analysis of homoplasius and homologue characters, the statistics of the trees obtained and the support index, among other things. For this reason the works of Johansen (1982, 1986a,b) are considered the first formal phylogenetic analysis. Excellent revisionary work was performed by this author on the taxonomy, biogeography and phylogeny of Elaphrothrips, Leptothrips and Humboldthripini. In the first paper about Humboldthripini this author includes all the tribes of Thripinae. Johansen is clear in the description of characters and the detail of the methodology used for the phylogenetic reconstruction. This technique is the only weakness of this analytical work, because it does not include the use of specific software nor the analytical study of characters, but presents a good statistical approach to the distribution of characters that permits the repetition of these results. However the description of characters is the most important, with extensive character states all well-defined. That is the real strength of this work and character definition is the most important step in a good phylogenetic analysis. The work performed by Mound and collaborators (1980) as a phylogenetic analysis with absence of all the necessary technical considerations, which are indispensable for others that wish to try and reproduce those results, is an example of some critical papers that need extensive revision on the phylogenetic results performed. Several times I have tried to utilize this data matrix with different software for phylogenetic reconstruction but I havent

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Mtodos en Ecologa y Sistemtica ISSN impreso 1659-2182 ISSN digital 1659-3049 been successful in obtaining the same results as these authors (Figs 1-3). The only alternative in obtaining these results is using constraints for the topologies depicted (Figs. 4, 5). This implies that these results are not in the set of optimal solutions and is a particular restriction for a particular result. It is relevant to note that the results obtained without the use of constraints show better statistical fit (Figs. 1-3). These results indicate that the considerations of Bhatti (1993) seem to be correct when he comments about this work: The changes they proposed in the family classification of Terebrantia did not arise from cladistic analysis. They did not reconstruct the phylogeny. They merely plotted the character states on two proposed cladograms(page 99). A similar problem is evidenced when analyzing the data matrix used for the study of the phylogenetic relationships among groups of Aeolothripidae (Marullo & Mound 1995)(Figs. 6, 7). Recent works from Morris and collaborators (1999) have several methodological problems widely discussed in phylogenetic papers. These authors use very small numbers of characters considering the number of taxa included. The characters considered are not the best at a generic and supraspecific level. Several conclusions are not well founded in these results and they considered this phylogenetic analysis as definitive in order to propose patterns of evolution. The works of Retana-Salazar (1998, 2000, 2001, 2006, 2007a,b, 2010) are classical cladistic work, using new alternatives in the phylogenetic reconstruction, particularly useful in high homoplasius groups, where the common use of parsimony may conduce to unreal results (Brooks & Wiley 1986, Retana-Salazar & Retana-Salazar 2008). All propose new hypothesis and as such are subject to proof. As required by formal science. Recently Cavallieri and Mound argued that As in all branches of
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science, conclusions in taxonomy can be no more reliable than the data on which they are based (Cavallieri & Mound 2012, page 1). However, in many of Mounds articles concerning aspects of the phylogeny of Thysanoptera, there is a lack of reliability in the method. These papers are an effort in the search of objective tools for the determination of the characters that show natural groups in Thysanoptera (Retana-Salazar & Retana-Salazar 2008). Based in his results this author demonstrates the necessity to use phylogenetic analysis to define the possible directionality of evolutionary changes and the necessity to segregate some genera into several ones but defining the natural groups using phylogenetic analysis (Retana-Salazar 2006, Retana-Salazar & Soto-Rodrguez 2007, Retana-Salazar & Nishida 2007). CLADISTICS AS A TOOL FOR TAXA DESCRIPTION The philosophy of cladistics suggests the necessity to take some axiomatic purposes for the correct reconstruction of phylogeny. Some particular groups as Frankliniella, Thrips, Hoplothrips, Elaphrothrips, Heterothrips and others, have a great number of species and the homoplasious level is high, common observation is not enough to define which characters are homologues or homoplasious. The use of a data matrix of these characters to obtain a hypothetical tree permits the phylogenetic study of the characters behaviour and its phylogenetic value. Choosing a correct out-group may be difficult when the in-group is highly homoplasious. The choice of an incorrect out-group can lead to equivocal and not congruent relationships. I suggest two practices for this delicate scenario: a) the use of several out-groups all of them with

Mtodos en Ecologa y Sistemtica ISSN impreso 1659-2182 ISSN digital 1659-3049 characters shared with the in-group (RetanaSalazar 2010), or 2) the use of a hypothetical ancestor created using characters observed in several other taxa (Retana-Salazar 2007a). In this aspect Mound and collaborators (1980) and Bhatti (1988, 1993) suggest possible data for the inference of the hypothetical ancestor. The use of multiple out-groups can be very useful in groups with few species where the diagnostic characters are well defined (Retana-Salazar 2007a). By contrast, in taxonomic groups where there is no clarity on the limits of the taxa and there is no homogeneity of the characters used in grouping, the use of multiple out-groups may not be of much help except to determine the paraphyly of the ingroup (Retana-Salazar 2010). BIOLOGICAL CHARACTERS IN THYSANOPTERA Some authors suggest (Mound & Marullo 1996) that the uses of these criteria are in several cases better than the morphological inference, because the plasticity of morphological features may lead us to mistaken associations. Lamentably these characters are diagnostic in few cases. One relevant case where biology has been the most important feature is Aulacothrips recognized as family level by Bhatti (2006) it is well supported by morphology, biology and biogeographic features. In this particular species, the only one known to be an ectoparasite of insects and whos natural hosts are the individuals of Aetalion, and some membracids is one extreme case and there is no discussion about this. Other groups are not so clear in those biological features (Alves-Silva & Del Klaro 2011). Several species of Frankliniella are closely related to Asteracea species, and only one is strictly associated with Hemerocallis, while few are strictly associated with mosses (Mojica & Johansen 1990, Mound & Marullo 1996, Retana-Salazar 1998), but it is difficult to establish that these particular groups are
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different taxa only based on their biology. Retana-Salazar (2010) shows that these biological features are only significant for the characterization of species, but not for groups of species. Biology may be highly homoplasious because biological features are very plastic, and in these cases the uses of biological features in absence of other morphological characters is not a good index for group segregation. MORPHOLOGICAL CHARACTERS AND THE
PROBLEM WITH SPECIES RECOGNITION

The morphological characters may be very variable, but we need critical studies of the group variation for making good taxonomic decisions. Bhatti (2006) established that taxonomy as we know it is really morphotaxonomy, because we need particular morphological features for the description and comparison between species. Some particular features as the brachipterous condition of some species in Frankliniella and Thrips may lead to the recognition of natural groups with a particular morphological feature (possibly apomorphic character) that defines the group. This consideration without a formal analysis of other characters may bring us to incorrect phylogentical assumptions. In the genus Thrips it seems that this character is associated with other derived characters and the brachipterous species may form a real natural group. In Frankliniella the situation is very different and the brachipterous forms are not supported by other derived characters and the reduction of the wings seems to appear several times in the evolutionary history of the group (Retana-Salazar 2010). The number of antennal segments is analyzed by Mound and Palmer (1981) in six genus-

Mtodos en Ecologa y Sistemtica ISSN impreso 1659-2182 ISSN digital 1659-3049 groups in Thripina. This author suggested that the plesiotypic condition is 9 segments, some groups in Aptinothripina and all the genera in Pseudothripini seem to have a secondary development of this 9-segmented condition. Mound and Palmer (1981) suppose that this condition derives from the subdivision of the sixth segment. With this evidence Mound and Palmer (1981) suggest that the most common apomorphic condition is seven antennal segments. This consideration leads to the possibility of interpreting the presence of six, seven and eight as an apomorphic sequence. Recent studies in the genus-group Anaphothrips (Retana-Salazar 2007b, Retana-Salazar 2010) demonstrate that these conditions may be reversal on several occasions. They are useful in the diagnosis of genera, the studies in the Anaphothrips group evidenced that this transformation has good phylogenetic inertia with the hierarchical level of genus. A similar condition is presented by the phylogenetic analysis of the Hoodothrips genus-group (Retana-Salazar 2007a). In this particular case the fusion of segments may occur not only in the terminal segments. This feature complicates things when considering apomorphic and plesiomorphic conditions. Those are good examples of how the same characters, in genera that seem phylogenetically close, or at least included in the same family, have a very different evolutionary landscape. Mound and Palmer (1981) proposed the uses of 8 morphological characters for the segregation of genus-groups in Thripina. These authors proposed a particular polarity from these characters based on merely observational data of slides. This methodology suggested that the analysis may be erroneous; Retana-Salazar (2008) uses a data matrix with these 8 characters in the phylogenetic reconstruction of the relationships of the genus-groups proposed by Mound and Palmer (1981). In this paper
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Retana-Salazar (2008) considered those characters under a formal analysis and use for the determination of polarity recently described fossil form with a good number of specimens that permit a good observation of the major part of characters (Pealver 1998). Results of this analysis show that the values of these characters in genus-groups definition are very poor and the majority is not well associated with the phylogenetic structure of these groups. This kind of work reveals that considerations about polarity of characters in absence of formal analysis of phylogeny and based only in observation of the alpha taxonomists, may lead to grave mistakes that may be reflected in erroneous taxonomic decisions. For this reason we need to 1) establish a phylogenetic hypothesis and 2) decide based on these results which characters have a good phylogenetic fit in each case and then these will be the features used in the definition of natural groups. This fit needs to be measured using some important tools, like the measure of the phylogenetic inertia of characters (Bjrklund 1997, Diniz-Filho 2000, Retana-Salazar & Retana-Salazar 2008) and the evaluation of the less entropic topology for a same data matrix (Brooks & Wiley 1986, Retana-Salazar 2007a, Retana-Salazar & Retana-Salazar 2008). In Tubulifera the problem is the same. Mound and Marullo (1996) proposed that in the Liothrips lineage, the presence of one sensorium in antennal segment III and three sensoria in antennal segment IV are characteristics of this generic group. The phylogenetic analysis performed by Morris and collaborators (1999) using morphological features shows that this condition in antennal segments III-IV came up several times in the evolutionary history of the group. These characters have a low phylogenetic inertia and for this reason are not good in defining the lineage.

Mtodos en Ecologa y Sistemtica ISSN impreso 1659-2182 ISSN digital 1659-3049 However this kind of characters is good for the segregation of generic taxa, particularly the number of antennal sensoria in antennal segment IV in the Karnyothrips lineage (Retana-Salazar & Soto-Rodrguez 2007). Recently Minaei and collaborators (2007) redefined the genus Noeheegeria, and in this paper, without any particular phylogenetic arrangement the authors considered that this feature (antennal sensoria), is indeed valid for the segregation of genera. These authors analyzed the presence of this feature in a group of species that shows this particular character, but its presence is not enough for separating if we do not know whether these characters are plesiotypic or apotypic, because apormorphous characters could support all segregation. This biological problem to define which characters are apomorphous and which of them are plesiotypic, as well as the determination of homoplasius characters, is only resolved by phylogenetic analysis. These problems with morphological characters are biological hurdles usually appointed in literature. Moreover, there are other kinds of problems with morphology in Thysanoptera. The preparation procedure can affect morphology. For example, Hoyers medium creates undulating setae and produce distortion of the colour and the original shape of the specimen. This is due to it not being viscous at all and the cover slide then stretches the sample, destroying the original shape of the specimen. The macerating substance affects coloration and the clarity with which some characters are observed. Excess pressure on the cover slide stretches the sample producing distortion of the morphological features, for example wider abdomen and head bases, together with longer mouth parts and displaced and longer setae (Retana-Salazar & Mound 1994). Retana-Salazar & Mound (1994) proposed that variation within a species can be greater than
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variation among several species, but this affirmation needs further studies because highly variables species such as Fr. occidentalis and Fr. intonsa may be a species complex. Mound (2005) suggests that phenotypic plasticity is a great source of morphological variation within species, but this asseveration needs several controlled experiments to prove it and not just the personal guess of a taxonomist. MORPHOLOGICAL TRAITS OF THE IMMATURE
STAGE

The characters of the immature stages could be useful determining the species as in the case of Selenothrips rubrocinctus where the species name refers to the main feature of the larva II, the presence of a red band at the middle of body. In other cases, such as Taeniothrips inconsequens where has been reported the presence of a well-developed toothed comb in the terminal segments of the abdomen of the larva II, which is characteristic of the species (Lacasa and Llorens 1996)(Fig. 8). The presence of distinctive characters in some species has caused to put aside the integral description of the larva II, thereby losing a large amount of information that can be of value in studies of phylogeny. The characters of the immature can be useful in phylogeny and taxonomy, making it necessary to develop studies of the ontogeny of the characters, as has been done in groups like Culicidae (Harbach 1991). In Thysanoptera have been studied little morphological characters of the different stages of development. There is no clear characterization of the genera at the immature level but if there is numerous descriptions of the larvae of some species of thrips. A first step in systematic research into the characters of larval forms in Thysanoptera is to determine the general features that define the different genera and taxonomical value of these characters.

Mtodos en Ecologa y Sistemtica ISSN impreso 1659-2182 ISSN digital 1659-3049 An interesting example is the study of the immature of Frankliniella insularis where the adult female is characterized by the absence of medial comb on the posterior margin of sternum VIII (Fig. 9a), among other features, but the observation of larvae II shows that this comb is not present in abdominal segment VIII but a similar but small and complete similar structure is in abdominal tergum IX in this stage (Fig. 9b). This can be useful in the polarization of this character. The structure of the segments of the antenna can be a good Terebrantia taxonomic character. In Thripidae segments III-IV of immatures are very developed and segments V-VII are forming a stylus (Fig. 10a, b). In Aeolothripidae antennal segments are evident since very early as well developed units of similar structure to that of adults but with fewer segments (Fig. 11a). In addition, in Aeolothrips intermedius has ascertained the presence of a double comb in the caudal segments of the abdomen (Fig. 11b), while in adults there is no evidence of such structures. Preliminary data has given rise to start at the University of Costa Rica a project to describe systematically the characters of the larvae of the principal species of thrips. These data will begin studies on the importance of ontogeny in phylogenetic characters polarization. TERATOLOGICAL CHARACTERS These characters are abnormalities recorded by taxonomists after the revision of a long series of individuals. Such variation is highly diverse, in most animals it may be identified because the individuals involved either deviate markedly from type as to be recognized as freaks or because the abnormalities involved are asymmetrical (Mayr 1969). Teratological characters may not be used in taxonomic decisions, however recently Goldarazena and
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collaborators (2008), used a particular character in a few individuals to synonymize a recently described species. This type of mistake may be dangerous because some journals publish these equivocal papers. Asiain and Mrquez (2009) show that several specimens in some particular Coleoptera groups, are highly asymmetrically modified and for this reason asymmetries are not good characters for taxonomic work. Asymmetry is a good indicator that this character is not possible to be valuated to propose taxonomic changes (Mayr 1969). Cappe de Baillon (1927) and Balazuc (1948) studied and classified the teratological variation. Recent studies in highly polluted areas of Europe has shown that some sensitive species like Frankliniella intonsa may be drastically affected promoting an increase in the rate of these abnormalities and the majority of these monstrosities are associated with antennal deformations (Vasiliu-Oromulu et al. 2008). However, the abnormalities reported as a result of environmental issues, reflected in embryological deformations in species such as Frankliniella intonsa, are also found in specimens collected in undisturbed systems in tropical regions. This is the case of some species of Aeolothrips and Liothrips collected in Costa Rica and in some specimens of Erotidothrips mirabilis collected in Asian national parks. It is necessary, as has been done in other insects groups, to begin producing literature that reports and illustrates these morphological changes which may lead to taxonomic errors in Thysanoptera. MOLECULAR DATA IN THYSANOPTERA In several cases this kind of data are considered the panacea for all the taxonomic and phylogenetic problems. But the real landscape is very different. In some aspects molecular data are more objective than traditional characters, but in other aspects these molecular characters

Mtodos en Ecologa y Sistemtica ISSN impreso 1659-2182 ISSN digital 1659-3049 are inherently subjective (Lee 2004). The considerations of some specialists that state molecular data as being the solution to our problems only denote little knowledge about this topic. Frequently the molecular data are not consistent with the resolution. Same sets of data show very different solutions and the question is what is the truth? In reality, all the answers are true because the inconsistency of the resolution of different sets of data is an effect of different methodologies for the study of the data matrix (Kjer et al. 2006, Klass 2007). In other cases the molecular data concludes with resolutions without consistency with the biology of the group (Mound & Morris 2007) Mound and Morris (2007) used molecular data to reconstruct the phylogeny of the Thysanoptera. The only clear result that these authors obtained was the monophyly of Thysanoptera. The only arrangements with support for natural groups definition was Tubulifera and Terebrantia, the same two groups defined by Haliday in 1836. The family groups are not well supported and the classical families are not consistent natural groups. These results are totally inconsistent with the biological and morphological features. Several times molecular studies have the same importance than a study in the Amazonia to conclude that this is a forest. On the other hand, the molecular data needs good support in morphological features associated with the phylogenetic results, otherwise these results are inconsistent. Crespi and collaborators (2004) with molecular data matrix conclude that in some groups as the clade formed by (Advenathrips+Vicinothrips)(Koptothrips(Triad othrips(Crespithrips(Glaridothrips+Xaniothrips ))))) in the basal node of Koptothrips the enlarged forelegs character appears, which may
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diagnose this group, but this character is not constant in the species of this group. Studies conducted in other arthropod groups have concluded that molecular data may indicate the presence of genetic units with a high rate of genetic divergence which could indicate the presence of undescribed species. However, it is unclear what level of divergence is needed for allowing the delimitation of two lineages (Camacho et al. 2011). CONCLUSIONS Thysanoptera is one of the highest homoplasious groups such as Hymenoptera and Hemiptera (Nieto 1999, Gauld & Mound 1982, Mound & Palmer 1983). Mound supposes that the operational difficulty with systematic studies on Phlaeothripinae is that many taxa at generic level and above are defined by loss of apomorphous characters. Gauld & Mound (1982) conclude that genera in groups of insects with high levels of homoplasy need to be defined polythetically, which is through the presence or absence of one or more unique characters. About this point Mayr (1969) proposed three statements that need to be satisfied for a polythetic taxon: a) each species possesses a large number of the total number of characters of the taxon, b) each character is possessed by a large number of the species and c) no characters are possessed by every species of the aggregate but is missing in the species of all the other taxa. The polythetic taxa created by Mound as Holopothrips does not follow the anterior statements and these genera defined in absence of real characters are a big black box where the biology and morphology needs to be reviewed for a proper taxonomy of these species (RetanaSalazar & Nishida 2007).

Mtodos en Ecologa y Sistemtica ISSN impreso 1659-2182 ISSN digital 1659-3049 In the last years some authors considered that genitalia is not highly informative in thrips taxonomy, and only some structures highly variables as pseudovirga are more or less useful (Mound & Minaei 2007). This consideration is opposite to other data about morphological variation derived from analysis of long series of specimens and the analysed together with the geographic variation (Retana-Salazar 2006, 2009). These opposite results are indicators of the complexity of thrips characters. Despite the considerations regarding the observations of some taxonomists, there is a big amount of literature that demonstrates the utility of the male genitalia in taxonomic works (CrdobaAguilar 2000). Molecular data in Thysanoptera is not good for taxonomic decisions and makes for poor characters in order to define biological groups (Mound & Morris 2007). Thysanoptera like other groups, has several problems if molecular data are considered as the best solution to taxonomic and phylogenetic problems. Molecular data is not the solution to taxonomic issues and the objectivity in some aspects of this kind of data is accompanied by the subjectivity in other considerations (Lee 2004). The ability of molecular data to determine species by genetic similarity, remains valid if the species of the group are well known, at the same time, the possibility of using molecular data to identify new species appears to be an unreliable technique as well, if the data-bases are not large enough or if taxonomic confirmations of their work are lacking (Meyer & Paulay 2005). Teratological studies are necessary in Thysanoptera because some erroneous taxonomic decisions have been proposed using this kind of characters. In some other groups of insects these particular characters have been studied for the taxonomic significance of these (Asiain & Mrquez 2009). Goldarazena and collaborators (2008) synonymyze the species
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Nicolemma garitai with Aurantothrips orchidaceus based on teratological characters and these decisions are not well founded and lead to grave mistakes in taxonomic work. These kinds of publications in sections used in journals for discussion of particular areas of conflict with few arguments destroy good taxonomic proposals for the sole comfort of taxonomists. We need a natural classification of Thysanoptera, but this group is highly homoplasius, the alternative is to carry out phylogenetic studies that help us in the definitions of higher taxa and the study of the phylogenetic inertia associated to characters that show us which features are good for the definition of natural groups that may be considered as taxa. Several times specialists consider that the characters that are good in a particular group may be extrapolated to other taxa. In highly homoplasius groups this a priori consideration may conduce to consider as valid artificial groups with homoplasius characters used for their diagnosis. Mound and Palmer (1981) published an interesting paper where they analyzed eight morphological characters in 50 genera of Thripini, they conclude that these features have a high taxonomic value and propose the polarization of these characters based only on the mere observation of specimens and in absence of formal analysis. Based on these characters these authors propose some genus groups Frankliniella genus group, Thrips genus group, Taeniothrips genus group, Mycterothrips genus group and other. Recent studies based in polarity of characters using fossil data and a new methodology to study the fossil insertion in phylogenetic trees show that these characters are not good for defining genus groups and are not informative. Only two characters are useful

Mtodos en Ecologa y Sistemtica ISSN impreso 1659-2182 ISSN digital 1659-3049 for phylogenetic structure and both are plesiotypic and consequently not good for determination of natural groups (Retana-Salazar 2008). That is an example of the problems carried on with classification systems based on taxonomic observation of characters in absence of formal phylogenetic analysis. Pealver and Nel (2010) have recently published an excellent paper about the transcendent importance of the fossil evidence in determination of the contribution of morphological characters and based in the evidence and a new and better conducted phylogenetic analysis conclude that the family Stenurothripidae needs to be considered a good taxon and an acceptable name better than Adiheterothripidae. More evidence is needed to support this new hypothesis but at least this proposal is solidly founded in data and formal analysis. Previous studies have spurious phylogenetic analysis where the repetition of the analysis shows a different result to those that have been published (Mound & Marullo 1998).

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trees obtained. Recently Retana-Salazar & Retana-Salazar (2008) published a book where the uses of entropy in the study of biological systems are proposed, particularly the evolutionary systems and the determination of new taxa. ACKNOWLEDGEMENTS To Ral Ramrez-Morales and Stephen Smith for the English revision of this paper and Alexander Rodrguez-Arrieta for his comments to this paper, and also The project Descripcin y ultraestructura de los thrips (Thysanoptera: Insecta) de Mesoamrica 810-A6-239 and the project "Estudio morfolgico y gentico de los estados inmaduros de thrips (Thysanoptera: Insecta) de relevancia econmica en Hispanoamrica", N810-Bl-224. The collections of the British Museum of Natural History, London, England, Senckenberg Museum, Frankfurt, Germany, Museo Geominero de Madrid, Espaa, Collection de Thysanoptera du Muse d'Histoire Naturelle de Paris, National Collection of South Africa, Pretorya, South Africa, IBUNAM, Coyoacan, Mexico DF, Universidad Autnoma de la Habana, Cuba, Centro Nacional de Sanidad Vegetal de Cuba, Universidad Autnoma Agraria Antonio Narro, Coahuila, Mexico, Universidad Autnoma de Nayarit, Nayarit, Mexico, INBio, Santo Domingo de Heredia, Costa Rica. Alcides Snchez-Monge for outstanding production of SEM images. REFERENCES
Alves-Silva, E., Del Klaro, K. 2011. Ectoparasitism and phoresy in Thysanoptera: the case of Aulacothrips dictyotus (Heterothripidae) in the Neotropical savanna. Journal of Natural History 113. Balazuc J. 1948. La teratologie des Coleopteres et experiences de transplantation sur Tenebrio

The only real tool to define hypothetical natural groups, based on the presence of apomorphous characters, is the phylogenetic reconstruction and the analysis of the phylogenetic inertia of the characters, that permits the consideration of which characters are better to be used in taxonomic work. There arent any good indicators for phylogenetic inertia of characters in qualitative data sets, but some authors have proposed the use of the retention index for this evaluation (Bjrklund 1997, Diniz-Filho 2000). Several times, with high complex groups like Thysanoptera, the parsimony analysis is not a good tool because the results of the reconstruction are a false clade, for this reason Brooks & Wiley (1986) suggest the uses of the entropy concept in phylogenetic reconstruction as a way of evaluation of the
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molitor L. Memoires de Meseum National dHistoire Naturelle 25:1-293. Bhatti JS. 1993. Phylogenetic relationships among Thysanoptera (Insecta) with particular reference to the families of the Order Tubulifera. Zoology (Journal of Pure and Applied Zoology) 4:93-130. Bhatti JS. 2006. The classification of Terebrantia (Insecta) into families. Oriental Insects 40:339375. Bjrklund M. 1997. Are comparative methods always necessary? Oikos 80:607-612. Brooks DR, Wiley EO. 1986. Evolution as Entropy: Toward a Unified Theory of Biology, Chicago Univ. Press. Camacho, A.I., Dorda, B.A., Rey, I. 2011. Identifying cryptic speciation across groundwater populations: first sequences of Bathynellidae (Crustacea, Syncarida). Graellsia 67(1):7-12. Cappe de Baillon P. 1927. Recherches sur al teratologie des insectes. Encyclopedie dEntomologie 8:5-291. Cavalleri, A., Mound, L.A. 2012. Toward the identification of Frankliniella species in Brazil (Thysanoptera, Thripidae). Zootaxa 3270:1-30. Crdoba-Aguilar A. 2000. Evolucin y diversidad de la morfologa de los genitales masculinos en insectos. Folia. Entomol. Mex. 110:95-111. Diniz-Filho JAF. 2000. Mtodos Filogenticos Comparativos. Holos Editora. Brasil. Gauld ID, Mound LA. 1982. Homoplasy and the delination of holophyletic genera in some insect groups. Systematic Entomology 7:73-86. Goldarazena A, Mound LA, zur Strassen R. 2008. Nomenclatural problems among Thysanoptera (Insecta) of Costa Rica. Revista de Biologa Tropical (Forum) 56(2):961-968.
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Harback, RE. 1991. Ontogeny of the larval stage of Sabethes chloropterus, with special reference to setal development and phylogenetic implications for the family Culicidae (Diptera). Mosquito Systematics 23(1):10-24 Johansen RM. 1982. Algunos aspectos de la biologa, ecologa, conducta y distribucin de geogrfica del gnero Torvothrips (Insecta: Thysanoptera). Anales del Instituto de Biologa. Universidad Nacional de Mxico 52:205-222. Johansen RM. 1983a. El gnero Elaphrothrips Buffa, 1909 (Thysanoptera: Phlaeothripidae) en el continente Americano; su sistemtica, evolucin, biogeografa y biologa. Monografa del Instituto de Biologa, Univeridad Nacional de Mxico 1:1267. Johansen RM. 1983b. Nuevos thrips (Insecta: Thysanoptera; Terebrantia, Thripidae, Thripinae) de La Sierra Madre Oriental y del Eje Volcnico Transversal de Mxico. Anales del Isntituto de Biologa, Universidad Nacional de Mxico 53:91132. Johansen RM. 1986a. Revisin de la tribu Humboldthripini Johansen, 1983 (Insecta; Thysanoptera; Thripinae). Anales del Isntituto de Biologa, Universidad Nacional de Mxico 56:697744. Johansen RM. 1986b. Nuevos conceptos taxonmicos y filogenticos del gnero Elaphrothrips Buffa, 1909 (Thysanoptera: Phlaeothripidae) del continente americano y descripcin de dos especies nuevas. Anales del Instituto de Biologa, Universidad Nacional de Mxico 56:745-868. Lacasa Plasencia A. y Llorens Climent, J.M. 1996. Thrips y su control biolgico. Divulgacin Tcnica 18, Consejera del Medio Ambiente, Agricultura y Agua, Murcia, Espaa 312pp. Lee M. 2004. The molecularisation of Taxonomy (Review). Invertebrate Systematics 18:1-6.

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Marullo R, Mound LA. 1995. Su una classificazione sopra-generica della famiglia Aeolothripidae (Thysanoptera). Atti del XVII Congresso Nazionale Italiano di Entomologia, Udine: 8790. Mayr E. 1969. Principles of Systematic Zoology. McGraw-Hill Inc. New York. p 26. Meyer CP, Paulay G. 2005. DNA barcoding: Error rates based on comprehensive sampling. PLoS Biol 3(12): e422. Minaei K, Azemayeshfard P, Mound LA. 2007. The southern Paleartic genus Noeheegeria(Thysanoptera:Phlaeothripidae): redefinition and key to species. Tijdschrift loor Entomologie 150:5-64. Mojica-Guzmn A, Johansen RM. 1990. Estduio sucesional de tisanpteros (Insecta), habitants de lquenes y musgos, en cinco localidades de la Sierra Madre Oriental, Estado de Hidalgo, Mxico. Anales del Isntituto de Biologa, Universidad Nacional de Mxico 61:197-256. Morris DC, Mound LA, Scwarz MP, Crespi BJ. 1999. Morphological phylogentics of Australina gall-inducing thrips and their allies: the evolution of host-plant affiliations, domicile use and social behaviour. Systematic Entomology 24:289-299. Mound LA. 2005. Fighting, fligth and fecundity: behavioural determinants of Thysanoptera structural diversity.pp 81-105 in Ananthakrishnan, TN & Whitman D. (eds). Insects and phenotypic plasticity. Science Publishers Inc. Enfield, NH, USA. Mound LA, Palmer JM. 1983. The generic and tribal classification of spore-feeding Thysanoptera (Phlaeothripidae: Idolothripinae) 46(1):1-174. Mound LA, Heming BS, Palmer JM. 1980. Phylogenetic relationships between the families of recent Thysanoptera (Insecta). Zoologycal Journal of the Linnean Society 69:111-141.

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Mound LA, Palmer JM. 1981. Phylogenetic relationships between some genera of Thripidae (Thysanoptera). Entomologica Scandinava Suppl. 15:153-170. Mound LA, Marullo R. 1998. Two new basal-clade Thysanoptera from California with Old World affinities. Journal of the New Yoek Entomological Society 106:81-94. Mound LA, Minaei K. 2007. Australian thrips of the Haplothrips lineage (Insecta: Thysanoptera). Journal of Natural History 41(45-48):2919-2978. Mound LA, Morris DC. 2007. The Insect Order Thysanoptera: Classification versus Systematics. Zootaxa 1668:395-411. Pealver-Moll E. 1998. Estudio taxonmico y paleoecolgico de los insectos del Mioceno de Rubielos de Mora (Teruel). Instituto de Estudios Turolenses, Excma. Diputacin Provincial de Teruel, Espaa. 177pp. Pealver-Moll E. & Nel P. 2010. Hispanothrips from early Cretaceous Spanish amber, a new genus of the resurrected family Stenurothripidae (Insecta: Thysanoptera). Ann. Soc. Entomol de France (n.s) 46(1-2):138-147. Retana-Salazar AP.1998a. Reestablecimiento de los gneros Frankliniella, Exophthalmothrips y Bolbothrips (Thysanoptera: Thripidae) Revista de Biologa Tropical 46(2):385-396. Retana-Salazar AP. 1998b. Una visin filogentica de Frankliniella (Thysanoptera: Thripidae). Revista de Biologa Tropical 46(2):397-406. Retana-Salazar AP. 2000. Revisin y Filogenia del grupo genrico Pseudothrips (Thysanoptera: Thripidae). BRENESIA 54:51-62. Retana-Salazar AP. 2006. Filogenia y revision taxonmica de la superfamilia Pediculoidea n.stat. (Anoplura: Insecta). BRENESIA 66:15-24.

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Retana-Salazar AP. 2007a. El grupo genrico Hoodothrips (Terebrantia: Heliothripidae). Acta Zoolgica Lilloana 51(1):15-38. Retana-Salazar AP. 2007b. Los tisanpteros del grupo genrico Anaphothrips (Thysanoptera: Thripidae), conenfasis en Amrica Central. Revista de Biologa Tropical 55(1):321-333. Retana-Salazar AP. 2010. El grupo genrico Frankliniella: el significado filogentico de sus principales caracteres morfolgicos (Thysanoptera: Thripidae; Thripini). Mtodos en Ecologa y Sistemtica 5(3):1-22. Retana-Salazar AP, Mound LA. 1994. Thrips of the Frankliniella minuta group (Insecta: Thysanoptera) in Costa Ricen Asteracea flowers. Revista de Biologa Tropical 42:639-648. Retana-Salazar AP, Ramrez-Morales R. 2006. Establecimiento de un nuevo gnero de piojos (Phthiraptera: Pediculidae) asociado al hombre (Primates: Hominidae). BRENESIA 65:61-70. Retana-Salazar AP, Nishida K. 2007. First gallinducing thrips on Elaphoglossum ferns: A new genus and species of thrips Jersonithrips galligenus from Costa Rica (Insecta, Thysanoptera, Phlaeothripidae). Senckenbergiana biologica 87(2):143-148. Retana-Salazar AP, Soto-Rodrguez GA. 2007. Revisin taxonmica del grupo HaplothripsKarnyothrips (Thysanoptera: Phlaeothripidae). Revista de Biologa Tropical 55(2):627-635. Retana-Salazar AP. 2008. Un enfoque cladista para la insercin de fsiles en la filogenia. Revista Geolgica de Amrica Central. 39:93-106. Retana-Salazar AP, Retana-Salazar SA. 2008. Entropa Biolgica. Especies y Filogenia. Editorial del Instituto Centroamericano para la Investigacin Biolgica y la Conservacin (ECIBRC). San Jos, Costa Rica 167p.

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Vasiliu-Oromulu L, Jenser G, Brbuceanu D. 2008. Frankliniella intonsa (Trybom, 1895) a very sensitive bioindicator for air pollution. Acta Phytopatologica et Entomologica Hungarica, 43(1): 401-408. Wiley EO. 1981. Phylogenetics. The Theory and Practice of Phylogenetic Systematics. New York: Wiley-Interscience.

Received: 10 de Diciembre 2011 Reviewed: 03 de Febrero 2012 Accepted: 23 de Agosto 2012.

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Figure 1. Topology obtained by applying a heuristic analysis of the data matrix presented by Mound et al. 1980. Statistics L=144, CI=0.94, R=0,65, analysis performed with PAUP 3.1.1.

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Figure 2. Topology obtained by applying a heuristic analysis of the data matrix presented by Mound et al. 1980. Statistics L=144, CI=0.94, R=0,65, analysis performed with PAUP 3.1.1.

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Figure 3. Best topology obtained by applying an exhaustive analysis of the data matrix presented by Mound et al. 1980. Statistics L=144, CI=0.94, R=0,65, analysis performed with PAUP 3.1.1.

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Figure 4. First topology proposed by Mound et al. 1980

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Figure 5. Second topology proposed by Mound et al. 1980

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Figure 6. Phylogenetic tree of groups of the Aeolothripidae published by Marullo and Mound (1995).

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Figure 7. Topology obtained by exhaustive analysis of the data matrix proposed by Marullo and Mound (1995). PAUP 3.1.1 was used.

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Figure 8. Taeniothrips inconsequens immature stage II abdominal segment IX, dorsal view. SEM microphotography, 300X, Hitachi 570S (scale: 5,5cm100m)

Figure 9. Frankliniella insularis. A) Adult female comb on VIII tergum, abdominal segments. SEM microphotography, 600X, Hitachi 570S (scale: 5,7cm50m). B) Immature stage II comb on IX tergum, abdominal segments. SEM microphotography, 600X, Hitachi 570S (scale: 5,7cm50m)

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Figure 10. Thripidae antennal segments of immature stage II. A) Head and antennal segments I-II (Taeniothrips inconsequens, SEM microphotography, 500X, Hitachi 570S, scale: 5,7cm60m). B) Antennal segments III-VII (Taeniothrips inconsequens, SEM microphotography, 700X, Hitachi 570S, scale: 5,7cm43m).

Figure 11. Aeolothrips intermedius. A) Antenna, immature stage II (SEM microphotography, 400X, Hitachi 570S, scale: 5,7cm75m). B) Abdominal terga VIII-X, immature stage II (SEM microphotography, 400X, Hitachi 570S, scale: 5,5cm75m).

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