Revista de Biología Tropical ISSN: 0034-7744 [email protected] Universidad de Costa Rica Costa Rica

Goldarazena, Arturo; Mound, Laurence A.; Strassen, Richard zur Nomenclatural problems among Thysanoptera (Insecta) of Costa Rica Revista de Biología Tropical, vol. 56, núm. 2, junio, 2008, pp. 961-968 Universidad de Costa Rica San Pedro de Montes de Oca, Costa Rica

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Nomenclatural problems among Thysanoptera (Insecta) of Costa Rica

Arturo Goldarazena1, Laurence A. Mound2 & Richard zur Strassen3 1. Neiker-Tecnalia, Instituto Vasco de Investigación y Desarrollo Agrario, Departamento de Producción y Protección Vegetal E-31080 Vitoria, España; [email protected] 2. CSIRO Entomology, GPO Box 1700 Black Mountain Canberra, ACT, Australia; [email protected] 3. Forschungsinstitut Senckenberg, Frankfurt (Main), Germany; [email protected]

Received 05-IX-2007. Corrected 12-ii-2008. Accepted 09-iii-2008.

Abstract: We present data to argue that several recent papers on the Thysanoptera of Costa Rica are affected by unsatisfactory technical procedures, including failure to recognize intraspecific structural variation. Fourteen new synonyms are recognized for Costa Rica Thysanoptera, nine generic and five specific. Rev. Biol. Trop. 56 (2): 961-968. Epub 2008 June 30.

Key words: Variation, , identification, systematic relationships.

Biological taxonomy is unique amongst These extensive, and expensive, revisionary the sciences. In most sciences an obviously studies were required because of the quarantine false hypothesis will be ignored. In contrast, significance of so many apparent pest spe- a new taxon name and the biological hypoth- cies on avocados. The 18 Scirtothrips species esis this name implies must be fully consid- described from Mexico on mango (Mangifera ered by subsequent taxonomists, so long as indica), a tree crop not native to the Americas, the name meets the minimal requirements of will require similar extensive studies. the relevant Code of Nomenclature. However, Taxonomy requires a combination of good establishing that the author of a new taxon has technical methods and critical scholarship. For published incorrect data, or drawn conclusions taxonomy this involves the examina- that are at variance with pre-existing bio- tion of well-prepared microscope slides, using logical knowledge, can involve more extensive good optical apparatus, and the preparation of research than that involved in the faulty pub- skilled illustrations and descriptions, whilst lication. Commenting on one recent example, taking into consideration published observa- Mound and zur Strassen (2001) noted the tions from previous workers, and placing all absence of biological evidence to support the this within the context of our understanding description by Johansen and Mojica-Guzman of the biology of the . The large num- (1999) of a large number of new species of ber of synonyms, more than 20% of available Scirtothrips collected mainly from two tree names worldwide, produced by workers on this crops in Mexico. Subsequently, extensive field group of insects (Gaston and Mound 1993) has studies and DNA analyses have confirmed arisen through repeated failure to recognize that only one species of Scirtothrips occurs in one or more of the above disciplines and pro- Mexico and Guatemala on Avocado (Persea cedures. This paper records similar problems americana), and examination of the holotypes that have arisen in recent years among several of five of the “new species” has resulted in taxonomic publications on Thysanoptera from their formal synonymy (Hoddle et al. 2008). Costa Rica.

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 56 (2): 961-968, June 2008 961 Nakaharathrips Retana-Salazar, 2007 evidence. The conclusion seems to be derived from a mathematical paper (Retana-Salazar and This was erected with Anaphothrips Retana-Salazar, 2004) in which it is claimed sudanensis Trybom as type species. However, that a difference such as the number of anten- as indicated in the world list of Thysanoptera nal segments that can be scored must indicate (Mound 2005), this species, in the form of one that different taxa are involved. If this logic of its 10 recognized synonyms, Neophysopus were to be accepted, then it would be essential medioflavus Schmutz, is the type species to explain why “segment VI fully divided + of the genus Neophysopus Schmutz. Thus segment VI with a partial division” constitutes Nakaharathrips is clearly a synonym of an autapomorphy distinct from “segment VI Neophysopus because they share the same type not divided”. However, the available biologi- species. According to the generic key and the cal evidence is contrary to this phenetic claim, data matrix published with Nakaharathrips, with the number of antennal segments varying the species A. sudanensis is uniform in colour. in some species between individuals within This is incorrect, because this is one of the most populations, and some individuals having a remarkably bicoloured species in the Thripidae, different number of segments on the left and with the median abdominal segments almost right antennae (Palmer 1992). Moreover, there white but the head and thorax and the terminal are several genera of Thripidae in which differ- abdominal segments dark brown (see website, ent species have a different number of antennal PADIL). This bicoloured condition has been segments (Sakimura and O’Neill 1979, zur emphasized by many previous authors, as indi- Strassen 2003: 259, 269). Given the many cated by some of the synonyms of sudanensis, character states, including biological attributes such as citricinctus, bicolor, and flavicinctus (Nakahara 1995) that are shared among the 52 as well as medioflavus. All of this informa- species currently treated in Anaphothrips, there tion is available on the web (Mound 2005), as is little justification in transferring a few of well as in the formal taxonomic literature. A them to a different genus. The claim that having further problem in the published data matrix eight segments is the plesiomorphic condition is that both Nakaharathrips and Anaphothrips and nine segments the apomorphic condition is are scored as having ocelli reduced, whereas also questionable, because the presence of nine macropterae of both A. obscurus and A. sudan- antennal segments is characteristic of most bas- ensis have well-developed ocelli (see PADIL). al-clade Thysanoptera (Mound et al. 1980). We In Thysanoptera, development of ocelli is conclude that Neophysopus (=Nakaharathrips) closely correlated with wing length; the use of should continue to be treated as a synonym of these in a phylogenetic analysis as independent Anaphothrips. characters is therefore questionable. The systematic position of Neophysopus Nicolemma Retana-Salazar, 2007 (=Nakaharathrips) has been undisputed for many years, either as a subgenus or as a This genus was erected for a single new synonym of Anaphothrips (Priesner 1949, species, N. garitai Retana-Salazar, although the Stannard 1968). The decision (Retana-Salazar type specimens of this species had previously 2007) to segregate into one genus those spe- been studied and identified by Mound and cies that have antennal segments VII and VIII Marullo (1996) as Aurantothrips orchidaceus fully distinct from VI, together with those (Bagnall). The new genus and species was that have segment VI partially divided by an characterized on two structural features: anten- incomplete oblique suture, and to place into a nal segments VII and VIII fused to produce a second genus those species in which segment 7-segmented antenna, and a spherical structure VI is completely undivided, is not supported present in the abdomen that was interpreted as by any additional morphological or biological a spermatheca. However, fusion of the terminal

962 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 56 (2): 961-968, June 2008 two antennal segments is not unique to the claim that if differences can be observed and few specimens listed from Costa Rica. Similar scored mathematically then these differences fusion has been observed on either the left or must be interpreted as indicating that different right antenna of three females within a long taxa are involved, regardless of any biological series of Aurantothrips orchidaceus collected data. However, any such a mathematical sys- from orchids in Colombia, and one male of tem will be biased if only selected data is this species from Britain has both antennae considered. 7-segmented (specimens in the Natural History Museum, London). Thus the first of the char- Quirosiella Retana-Salazar, 2000 acter states used to distinguish the new genus is merely intraspecific variation. Similarly, the This genus was erected for two species, spherical structure in the abdomen that is illus- a new species, Q. sotoi from Costa Rica, as trated (Retana-Salazar 2007) is clearly visible type species and Pseudothrips longiceps Hood in many specimens of Aurantothrips orchida- based on a single female from Brazil. The ceus, including the Lectotype female. We there- generic definition was not accompanied by any fore conclude that Nicolemma is a synonym of comparison with other genera, being distin- Aurantothrips Bhatti, and that N. garitai is a guished only in a key to five genera by tergite synonym of A. orchidaceus (Bagnall). VII lacking a marginal comb of microtrichia. This state was contrasted with species that are Aurantothrips Bhatti, 1978 known to exhibit a variable number of marginal microtrichia laterally on this tergite (Stannard, This genus was erected (Bhatti 1978: 90) 1951, Palmer and Mound 1985, Mound and for a single Neotropical species from orchids, Marullo 1996). In view of this variation, the Anaphothrips orchidaceus Bagnall, described genus Quirosiella is here regarded as a syn- originally from greenhouse specimens in onym of Psectrothrips Hood, to which genus Europe. Sakimura (1967) examined 24 females P. longiceps was transferred by Palmer and and nine males and concluded, on the basis of Mound (1986). weak differences in colour and chaetotaxy, that A cladogram provided by Retana-Salazar two subspecies could be distinguished. Mound (2000) involved 11 of the 17 nominal spe- and Marullo (1996) reviewed the evidence cies in the genera Psydrothrips Palmer and produced by Sakimura, but reported further Mound, Psectrothrips Hood (= Lacandonithrips variation in colour and structure in specimens Johansen) and Pseudothrips Hinds, but no data from Colombia and Brazil that cast doubt on matrix was provided to enable the suggested the existence of two biological entities. The relationships to be assessed, although there subsequent decision (Retana-Salazar 2007) that was a table indicating the characters used. the two subspecies are distinct species was not Moreover, no indication was given of which of supported by any evidence of having studied the 11 species the author did, or did not, person- any specimens. Accompanying the decision ally study; for several species the data evident- were two short paragraphs copied (but trans- ly came from published literature. One species lated) from Mound and Marullo (1996) that not included in the cladogram for which speci- detailed the differences between the specimens mens were available was a Costa Rican species available to Sakimura. However, the subse- named after its collector, Pseudothrips retanae quent paragraph by the same authors, discuss- Mound & Marullo. The cladogram for the 11 ing the variation in and between samples species placed the two species of Quirosiella from Colombia and Brazil, was neither trans- as sister to two species of Pseudothrips, and lated nor discussed. Again, the logic seems to this pair of genera as sister to five species of be derived from Retana-Salazar and Retana- Psectrothrips. This pattern of relationships Salazar (2004), in which the authors appear to is particularly surprising because the males

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 56 (2): 961-968, June 2008 963 of Pseudothrips species exhibit a remarkable Vargasia character that occurs in only six other genera Retana-Salazar & Soto-Rodriguez, 2007 of the worldwide family Thripidae, each of these being from South America (Mound and This genus was distinguished from Marullo 1996, De Borbón 2008). These genera Karnyothrips Watson, with Hindsiana mela- include Psydrothrips but not Psectrothrips, leuca Bagnall as type species (although quoted and the character is the presence of a glandular incorrectly as Karnyothrips melaleucus). The opening on the antecostal region of sternite III, authors of this new generic name indicated that whereas the normal condition in this family is they had studied specimens of K. melaleucus for a transverse porose area medially on several and K. ochropezus. However, they claimed to sternites posterior to the antecostal ridge (zur include a total of 24 species in this new genus, Strassen 2003, Tyagi et al. 2008). although they did not provide the names of any The species Q. sotoi was compared to of these species nor any evidence of having Psectrothrips longiceps (Hood), a species pre- studied any of them. Vargasia was distinguished viously recorded from the same locality, Las on the basis that the two species studied have Alturas in Costa Rica (Mound and Marullo only three sensoria on antennal segment IV, 1996: 179). The new species was distinguished and this was contrasted with Karnyothrips fla- because the metanotal reticulations were con- vipes, the type species of Karnyothrips, which sidered to lack internal markings, and the intero- the authors claimed to have four sensoria on cellar setae were considered shorter. However, this segment. However, specimens of both K. the metanotal internal markings are visible flavipes and K. melaleucus have been stud- under phase contrast in two female paratypes ied in the collections of the Natural History of Q. sotoi that have been studied (deposited Museum, London, and the Australian National in the Senckenberg Museum, Frankfurt, and Collection, Canberra, and the number the Natural History Museum, London), and of sensoria on segment IV varies from three to these markings are also clear in the previously four in both species, within and between popu- collected specimens from Las Alturas, Costa lations (Okajima 2006: 397). Both species are Rica. Unfortunately, the head of the paratype known to have a similar biology as predators deposited in the Senckenberg Museum is tilted of scale insects, and they are essentially similar forward such that the length of the ocellar setae in structure. Since there is neither morphologi- cannot be measured accurately, in contrast to cal nor biological evidence for supporting the the Hood holotype and the specimens slide new genus, Vargasia is here considered a syn- mounted in London. In view of the variation onym of Karnyothrips. In describing this new Q. sotoi and P. longiceps are here considered to genus, the authors also returned two species to represent the same species. Apterygothrips from Karnyothrips, but present- The generic name Linneothrips Johansen ed no evidence of examining either species, nor and Mojica (1996) should be considered a yet of examining any species of Apterygothrips synonym of Pseudothrips Hinds. Contrary to for reference. Retana (2000: 57), who sent the same species for description to both Mexico and Europe, the Willeia paper by Johansen and Mojica although listed Retana-Salazar & Soto-Rodriguez, 2007 as 1995 was not published until 30.viii.1996, as confirmed by the editor of the journal; the This new genus was erected for two spe- publication data of the Mound and Marullo cies, with Zygothrips americanus Hood as type book was 4.iii.1996. species (but quoted incorrectly as Karnyothrips

964 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 56 (2): 961-968, June 2008 americanus). The authors did not indicate that synonym of Karnyothrips. The authors failed they had examined any specimens of either to include their new name in their abstract, but species; apparently relying on data in the key they did include a nomen nudum “Aguilaria” to species provided by Mound and Marullo as a new genus; presumably one name replaced (1996). The genus was distinguished from the other during manuscript preparation. Karnyothrips on two character states. First, it was claimed that antennal segment IV bears Jironiella only three sensoria rather than four, but as Retana-Salazar & Soto-Rodriguez, 2007 indicated above this is variable in the type species of Karnyothrips, and moreover K. This new genus was erected for a single americanus has been examined at the Natural new species, J. saidi. The species was based History Museum, London and actually bears on specimens collected from Cyperaceae in only two sensoria on this segment. Second, K. Costa Rica, and the authors claimed that it americanus has the anteromarginal setae on the could be distinguished from Haplothrips spe- pronotum well developed instead of reduced, cies through the absence in the head of a although the length of this pair of setae is maxillary bridge. With the exception of this variable among other species of Karnyothrips. character, the description agrees closely with Given this variation, the known similarities in specimens of Haplothrips graminis Hood, a biology, and the absence of any further support- widespread species in Costa Rica and Colombia ing morphological evidence, the generic name on Cyperaceae and Poaceae. Subsequently, one Willeia is here regarded as a further synonym paratype female of J. saidi in the collections of of Karnyothrips. the Natural History Museum, London, has been examined under phase contrast illumination, Dioclesianothrips and remnants of a maxillary bridge are clearly Retana-Salazar & Soto-Rodriguez, 2007 visible. In H. graminis the maxillary stylets are particularly wide apart, and the maxillary This new genus was erected for a single bridge is therefore unusually long and slender. species, Haplothrips (Xylaplothrips) sono- This structure is thus very easily displaced rensis Stannard (but quoted incorrectly as during slide preparation; certainly it is not Karnyothrips sonorensis). The authors stated visible in all of the slides of this species in col- that they had examined specimens of this lections both in London and in Canberra. As North American species in the collection of a result, J. saidi is here placed into synonymy A.P. Retana-Salazar, but did not indicate how with the common Central American species, these specimens were identified nor the locality H. graminis, and Jironiella becomes a further from which they were collected. If the speci- synonym of the worldwide genus Haplothrips mens came from Costa Rica then this would Amyot and Serville. be the first published record from Central America. The genus was distinguished from Gynaikothrips garitacambroneroi Karnyothrips and also from Willeia because Retana-Salazar, 2006 antennal segment IV bears only two sensoria. However, contrary to the statement given by the This species was described from 13 authors, antennal segment IV of K. americanus females and two males, collected in Costa also bears only two sensoria, and the form of Rica. The plant species from which these speci- the other antennal segments in this species and mens were collected is not clearly stated in the in K. sonorensis is very similar, as indicated by original description, but judging from Garito- Stannard (1968: 431). There is thus no support Cambronero and Lizano-Fallas (2006) it was for placing this one species in a separate genus, Ficus benjamina. The thrips was distinguished and Dioclesianothrips is therefore considered a from Gynaikothrips uzeli (Zimmerman) by the

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 56 (2): 961-968, June 2008 965 greater length of the pronotal posteroangular illustrations of the male “genitalia” are based setae. However, the measurements given fall on the unexpanded epiphallus, an inflatable sac within the range of variation found in samples whose shape varies with the extent of inflation. of G. uzeli from various parts of South East The actual pseudovirga was not illustrated, Asia including the Maldive Islands. Moreover, and although the pseudovirga can be useful in G. uzeli seems to be increasing in distribution recognizing some Haplothripini (Mound and in the Caribbean region and southern States of Minaei 2007) its structure has not been demon- the USA on F. benjamina, presumably through strated to be useful for distinguishing species the activities of the horticultural trade (Boyd in the majority of leaf-feeding . and Held 2006). One female paratype of G. The number and length of setae on the apex of garitacambroneroi, deposited in the Natural the parameres is not stable between individual History Museum, London, has been examined, males of Gynaikothrips species examined from and as a result the species is here considered various parts of Asia and Australia. a synonym of the Asian species G. uzeli. The

NOMENCLATURAL SUMMARY

Anaphothrips Uzel, 1895: 142 Nakaharathrips Retana-Salazar, 2007: 329. syn.n.

Aurantothrips Bhatti, 1978: 90 Nicolemma Retana-Salazar, 2007: 324. syn.n.

Aurantothrips orchidaceus (Bagnall); Bagnall, 1909: 33 Nicolemma garitai Retana-Salazar, 2007: 324. syn.n.

Psectrothrips Hood, 1937: 262 Quirosiella Retana-Salazar, 2000: 55. syn.n.

Psectrothrips longiceps (Hood); Hood, 1954: 211 Quirosiella sotoi Retana-Salazar, 2000: 54. syn.n.

Pseudothrips Hinds, 1902: 146 Linneothrips Johansen & Mojica, 1996: 59. syn.n.

Pseudothrips retanae Mound & Marullo, 1996: 181 Linneothrips bicolor Johansen & Mojica, 1996: 62. syn.n.

Gynaikothrips uzeli (Zimmermann); Zimmermann, 1900: 12 Gynaikothrips garitacambroneroi Retana-Salazar, 2006: 6. syn.n.

Haplothrips Amyot & Serville, 1843: 640. Jironiella Retana-Salazar & Soto-Rodriguez, 2007: 632. syn.n.

Haplothrips graminis Hood, 1921: 69 Jironiella saidi Retana-Salazar & Soto-Rodriguez, 2007: 633. syn.n.

966 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 56 (2): 961-968, June 2008 Karnyothrips Watson, 1923: 23 Vargasia Retana-Salazar & Soto-Rodriguez, 2007: 631. syn.n. Willeia Retana-Salazar & Soto-Rodriguez, 2007: 632. syn.n. Dioclesianothrips Retana-Salazar & Soto-Rodriguez, 2007: 632. syn.n. “Aguilaria” Retana-Salazar & Soto-Rodriguez, 2007: 635. syn.n.

RESUMEN Thysanoptera: Thripidae). Folia entomologica Mexicana 93 [1996]: 39-70. [published 30.viii.1996] Presentamos datos para apoyar nuestro argumento Johansen, R.M. & Mojica-Guzman, A. 1999. The genus de que varios artículos recientes sobre los Thysanoptera Scirtothrips Shull, 1909 (Thysanoptera: Thripidae, de Costa Rica se han visto afectados por procedimientos Sericothripini), in Mexico. Fol. Ent. Mexicana 104: técnicos insatisfactorios, incluyendo el no reconocer la 23-108. variación estructural intraespecífica. Presentamos nueve sinonimias en los tisanópteros de Costa Rica: nueve a nivel Mound, L.A., Heming, B.S. & Palmer, J.M. 1980. de género y cinco a nivel de especie. Phylogenetic relationships between the families of recent Thysanoptera. Zool. J. Linn. Soc. London. 69: Palabras clave: variación, taxonomía, identificación, rela- 111-141. ciones sistemáticas. Mound, L.A. & Marullo, R. 1996. The Thrips of Central and South America: An Introduction. Mem. Entomol. International 6: 1-488. [published 4.iii.1996] REFERENCES Mound, L.A. & Minaei, K. 2007. Australian insects of the Bhatti, J.S. 1978. Systematics of Anaphothrips Uzel 1895 Haplothrips lineage (Thysanoptera – Phlaeothripinae). sensu latu and some related genera. Senck. Biol. 59: J. Nat. Hist. 41: 2919-2978. 85-114. Mound, L.A. & zur Strassen, R. 2001. The genus Boyd, D.W. & Held, D.W. 2006. Androthrips ramachand- Scirtothrips (Thysanoptera: Thripidae) in Mexico: a rai (Thysanoptera: ): an introduced critique of the review by Johansen & Mojica-Guzmán thrips in the United States. Fla Ent. 89: 455-458. (1998). Fol. Ent. Mexicana 40: 133-142.

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