Zootaxa 2230: 57–63 (2009) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2009 · Magnolia Press ISSN1175-5334(online edition) The Old-World genus Ceratothripoides (Thysanoptera: Thripidae) with a new genus for related New-World species LAURENCE A. MOUND1 & DAVID A. NICKLE2 1Honorary Research Fellow, Australian National Insect Collection, CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601 Australia. E-mail: [email protected] 2Systematic Entomology Laboratory, PSI, Agricultural Research Service, U. S. Department of Agriculture, 10300 Baltimore Avenue, Building 005, BARC-West, Beltsville, MD 20705-2350 U.S.A. E-mail: [email protected] Abstract A key is provided to five Old World species that comprise the genus Ceratothripoides Bagnall, and the species C. revelatus (Priesner) is recalled from synonymy with C. brunneus Bagnall. Five New World species previously placed in this genus are here allocated to Retanathrips Mound & Nickle gen.n., with Physothrips funestus Hood as type species. Key words: tospovirus vector, Retanathrips new genus, identification key, Ceratothripoides Introduction The genus Ceratothripoides was proposed by Bagnall (1918a) for one African species from Ghana, based on a single female specimen with distinctive antennae. Subsequent collecting indicated that the antennae of this specimen were deformed, and the species was discovered in several additional African countries. Since 1918, fourteen more species have been described in, or referred to, Ceratothripoides, although six of them have been placed into synonymy. Full nomenclatural information concerning all of these taxa is available in the web-based checklist to world Thysanoptera (Mound 2009a). However, the genus itself has remained poorly defined. Mound & Marullo (1996) discussed the five New World species that have been referred to this genus, but listed them as “Ceratothripoides”, indicating that these five were not congeneric with the Old World species. As a result these five species have remained with no formal generic assignment. Until recently, most of the species listed in Ceratothripoides have been known only to museum taxonomists. However, Murai (2000) recognized C. claratris as an important pest of tomatoes in Thailand, and Premachandra et al. (2005) recorded this species as a vector of a tospovirus on tomatoes in that country. Moreover, Halaweh & Poehling (2009) have indicated that tospovirus vector ability in this thrips is possibly controlled by a recessive allele. The identity and relationships of C. claratris are thus now of more general interest from an economic point of view. Moreover, C. brunneus has been found recently on three continents other than its native Africa. This species is well established and abundant on various crops in Peninsular Malaysia, two specimens have been examined from Puerto Rico, and a population has been studied from a greenhouse in the Netherlands (Mound & Azidah, 2009). Moreover, Nickle (2009) has recorded C. brunneus as being intercepted nine times in quarantine in North America. These records of this African species so far outside its native range, together with the studies on C. claratris as a tospovirus vector, have provided the incentive to re-examine the taxonomy and systematic relations of the species listed under this generic name. Unfortunately, the results presented here are by no means definitive. The lack of field studies in tropical countries, including valid host associations and structural variation within and between populations, have resulted in taxonomic decisions that are not necessarily reliable. Two species recognised here appear to be Accepted by C. Dalgleish: 20 Aug. 2009; published: 14 Sept. 2009 57 equally widely distributed across Africa, and are here distinguished on two apparently consistent character states. Both have been taken from various flowers, but there is no information on any host specificity. Two other species are known that appear to have overlapping ranges between the Sahel region and South East Asia, but these are distinguished only by rather weak colour differences for which the biological significance remains equivocal. The five New World species that are here referred to a new genus are based on particularly few specimens, with no knowledge of intra- and interpopulation variation. The restricted objective here is to clarify the nomenclature, illustrate character states that appear to facilitate identifications, and to indicate problems that require field and laboratory studies on these insects within the tropics of Africa and the Americas. Ceratothripoides Bagnall Ceratothripoides Bagnall, 1918a: 201. Type-species C. brunneus Bagnall, by monotypy. Diagnosis: Antennae 8-segmented, segments III–IV with forked sensorium and many microtrichia, segment I with pair of dorso-apical setae. Head with three pairs of ocellar setae; two setae comprising pair I usually arising one in front of the other (Fig. 7) (side-by-side in funtumiae); ocellar setae pair III arising within ocellar triangle; compound eyes with no pigmented facets (in cameroni and claratris with 4 or 5 weakly pigmented facets); maxillary palps 3-segmented. Pronotum with two pairs of posteroangular setae, 3–4 pairs of posteromarginal setae. Mesonotum with median setae close to posterior margin, anterior campaniform sensilla present or absent. Metanotum reticulate, median setae at anterior margin (Figs 1, 5), campaniform sensilla present or absent. Prosternal ferna entire, basantra without setae; mesothoracic sternopleural sutures complete, mesofurcal spinula present, metafurca spinula absent (except funtumiae). Forewing first vein with about 7 setae basally, 2 setae distally; second vein with complete row; discal seta present or absent on clavus. Tergites without ctenidia, median setae (S1) far apart, VI–VII with setae S3 much smaller than S4; tergite VIII posterior margin with complete comb of microtrichia (Figs 2, 6); X with dorsal split. Sternites III–VI with 3 pairs of marginal setae, VII with S1 and S2 arising well in front of posterior margin. Male sternites III–VII with 2 or more rows of small pore plates (Fig. 4); tergite IX with median setae slender or slightly stout (Fig. 10). Generic relationships. Many of the characters listed above, particularly the presence of a pair of dorso- apical setae on the first antennal segment, are shared with the species of Pezothrips, also with the legume- flower associated species of the genera Megalurothrips, Odontothrips and Odontothripiella. Nearly all of these species are endemic to the Old World. In contrast to the legume-flower thrips, the species of Pezothrips and Ceratothripoides have numerous small pore plates (Fig. 4) on the sternites of the males (Mound, 2009b). The character states of the first antennal segment and the male sternal pore plates also occur in species of the Trichromothrips genus-group (Masumoto & Okajima, 2005), but males in that group usually have paired drepanae on abdominal tergite IX. The systematic relationships of the nine species currently listed (Mound, 2009a) in Pezothrips continue to be problematical. The type species, P. frontalis (Uzel), has no comb on tergite VIII, whereas this is present laterally in P. kellyanus (Bagnall), but complete and long in P. dianthi (Priesner). Eight of these nine species are reputed to have setae S1 on sternite VII closer together than their length, although this is not true of P. kellyanus. However, all nine have setae S2 on sternite VII arising at the margin, in contrast to the condition in the species of Ceratothripoides. Key to species 1. Forewings brown or deeply shaded; compound eyes with no pigmented facets..........................................................2 -. Forewings uniformly pale with no dark shading; compound eyes with 4 (or 5?) weakly pigmented facets...............4 58 · Zootaxa 2230 © 2009 Magnolia Press MOUND & NICKLE 2. Pronotal posteromedian pair of setae almost as long as metanotal median setae (Fig. 8); metafurcal spinula present; male tergite IX medially with two pairs of stout setae (Fig. 10).................................................................. funtumiae -. Pronotal posteromedian setae scarcely 0.5 as long as metanotal setae (Fig. 1); metafurcal spinula absent; male tergite IX with median setae slender .......................................................................................................................................3 3. Mesonotum and metanotum with no campaniform sensilla (Fig. 1); forewing clavus with 6 or 7 marginal setae but no discal seta (Fig. 1); tergite VIII campaniform sensilla usually anterior to median pairs of setae (Fig. 2)................. ........................................................................................................................................................................brunneus -. Mesonotum and metanotum with pair of campaniform sensilla (Fig. 5); forewing clavus with 5 or 6 marginal setae and one discal seta (Fig. 5); tergite VIII campaniform sensilla posterior to median pairs of setae (Fig. 6)....revelatus 4. Legs extensively shaded brown; antennal segment V brown .........................................................................claratris -. Legs yellow; antennal segment V yellow ......................................................................................................cameroni Ceratothripoides brunneus Bagnall Ceratothripoides brunneus Bagnall, 1918a: 201 Physothrips marshalli Bagnall, 1918b: 66 Physothrips ventralis Hood, 1918: 116 This species was described originally from a single female with deformed antennae. This female was collected at Aburi, Ghana, 15.xi.1915, in Cola shoots and buds. The syntypes of marshalli were collected at this same locality, 30.x.1915, from flowers of Hibiscus sinensis. The original specimens of these two species have been re-examined and clearly represent the same species. Hood described ventralis from “numerous specimens” in various flowers from Kamerun and Nigeria (Ibadan). Pitkin (1978) selected as Lectotype of ventralis the female specimen that Hood had labeled as Holotype, and this was collected in Cameroon, 23 November 1915, from flowers of monkshood. This specimen lacks metanotal campaniform sensilla and the forewing clavus lacks a discal seta, but on tergite VIII the campaniform sensilla are posterior to the median setae. The first two character states indicate that, based on this Lectotype, the species ventralis is correctly considered a synonym of brunneus. The paralectotypes of ventralis from Cameroon in the USNM are also identifiable as brunneus, and of the 24 paralectotypes of ventralis in the USNM from Nigeria that are mentioned by Pitkin (1978), 23 are also identifiable as brunneus. However, one paralectotype from Nigeria, collected on Melia azederach, is here identified as revelatus, based on the two character states indicated in the key above, and a second paralectotype with identical data has also been studied (in BMNH) that also represents revelatus. In the Senckenberg Museum Frankfurt there are two female and one male paratypes of ventralis that were collected in Cameroon, and these represent brunneus. This species can be distinguished from other members of this genus by means of the characters in the key above, but the position of the paired campaniform sensilla on tergite VIII is not constant even in females, and in males these sensilla are often close to the posterior margin. Specimens of brunneus have been examined from the following countries: Ghana, Sierra Leone, Ivory Coast, Nigeria, Cameroon, Ethiopia, Uganda, Congo, Angola, and South Africa (in BMNH & SMF); Netherlands (in greenhouse) (in SMF); Puerto Rico and Peninsular Malaysia (in ANIC). This was one of the most common Thripidae collected during a recent survey of crop thrips in Peninsular Malaysia (Mound & Azidah, 2009). Two males have been studied (in ANIC) that were collected on Citrus at Maricao, Puerto Rico, “4.10.07”, and submitted for identification by Prof. Irma Cabrera. Ceratothripoides cameroni (Priesner) Taeniothrips cameroni Priesner, 1934: 28 Bhatti (1990) indicated that C. cameroni has antennal segment V yellow or shaded and legs yellow, whereas C. claratris has antennal V brown, and legs extensively brown. This statement concerning C. cameroni is true OLD-WORLD GENUS CERATOTHRIPOIDES Zootaxa 2230 © 2009 Magnolia Press · 59 for the original specimens from Sudan, but other specimens collected subsequently from that country have the antennae variable in colour, even with segments IV–V light brown, and legs have extensive light brown shadings. Specimens examined from Senegal (in BMNH) are similar to these, and are thus slightly darker than the type specimens from Sudan. Specimens from Nigeria and Saudi Arabia (in BMNH) have rather darker hind femora, and antennal segment V is also brown. In contrast, all available specimens from India and Thailand have brown hind femora and brown antennal segment V, and these are identified as C. claratris. Whether these slight differences in colour represent two distinct species, or whether they are symptoms of intraspecific variation, as suggested by zur Strassen (1975), remains unclear. The palest specimens, those that would be associated with the name C. cameroni, appear to come from the more arid areas, whereas the darker specimens, that would be associated with the name C. claratris, come generally from more humid areas. As indicated below, the two names are here retained because C. claratris is used for a form that is of economic significance, but the biological reality of these two colour forms requires further field studies. Ceratothripoides claratris (Shumsher) Taeniothrips claratris Shumsher, 1946: 178 Mycterothrips moultoni Seshadri & Ananthakrishnan, 1954: 213 Ceratothrips reticulatus Reyes, 1994: 183 [synonymised by Bhatti, 2003: 8] The significance of this species is discussed above under C. cameroni, of which it is possibly a variant. The two remain separated here primarily because the name C. claratris has become widely used in Asia in economic entomological literature (Premachandra et al., 2005). The name claratris has been used for specimens from various localities across India and Thailand, and also from Luzon in the Philippines (Bhatti, 1990, 2003). Ceratothripoides funtumiae (Bagnall) Physothrips funtumiae Bagnall, 1913; 292 This species remains known only from the original specimens from Uganda (in BMNH, SMF and USNM), together with 15 females from southern Nigeria (in BMNH). In contrast to the other species placed in this genus, the head bears ocellar setae pair I in the normal position for Thripidae, side-by-side; the other four species have these two setae placed irregularly or even one in front of the other (Fig. 7). Moreover, the metanotal reticulation is almost equiangular or even linear rather than transverse, and paired campaniform sensilla are present (Fig. 8). Campaniform sensilla are also present near the anterior margin of the mesonotum, and the metafurca bears a spinula. The forewing clavus is similar to that of revelatus, with five marginal setae and one discal seta. Antennal segments III–IV are brown with the apex contrastingly yellow (much as in Pezothripskellyanus), and antennal segment VIII is twice as long as VII. In females, the campaniform sensilla on tergite VIII are anterior to the median setae. In males, tergite IX bears two pairs of moderately stout setae medially (Fig. 10), and sternites III–VII bear a transverse pore plate at the anterior and three irregular rows of about 20 small pore plates behind this. Of the specimens of this species from Nigeria, four were collected with C. brunneus at Ibadan in February, 1964, on “lily flower”. Ceratothripoides revelatus (Priesner) stat. rev. Taeniothrips revelatus Priesner, 1937: 169 60 · Zootaxa 2230 © 2009 Magnolia Press MOUND & NICKLE FIGURES 1–10. Ceratothripoides species. C. brunneus 1–4: (1) Pronotum, mesonotum, metanotum and forewing clavus. (2) Tergite VII–IX. (3) Tergites VII–IX of male. (4) sternal pore plates of male. C. revelatus 5–7: (5) Mesonotum, metanotum and forewing clavus. (6) tergites VII–VIII. (7) Head. C. funestus 8–10: (8) Pronotum, mesonotum and metanotum. (9) Tergites VII–VIII. (10) Tergites VIII–IX of male. Although the original description of revelatus mentions an “allotype”, neither holotype nor type locality was specified. Priesner based the species, in part, on specimens identified by Karny (1925) as T. ventralis, from Kampala, Uganda, but listed further specimens from Njala, Sierre Leone, and Rutshuru, Belgian Congo. There is a long series of syntypes of revelatus in BMNH from Kampala, Uganda, 15.i.1921, bearing Karny’s handwriting and identification as ventralis. Moreover, one female “paratype” of ventralis in BMNH actually represents revelatus and has the following data: S. Nigeria, Ibadan, Melia azederach fls, 14.i.1915 (Jobbins- OLD-WORLD GENUS CERATOTHRIPOIDES Zootaxa 2230 © 2009 Magnolia Press · 61 Pomeroy). In this collection there is also one female and one male of revelatus, labeled: Ibadan, Moor Plantation, 11.v.1964, in lemon flowers. Although long considered a synonym of brunneus, this species can be distinguished by means of the indicated character states. In addition to the material listed above, the following specimens in the BMNH collections are here identified as revelatus: Ghana, Tafo, iv–v.1971, 14 females, 4 males (labelled brunneus) from cocoa flowers; Sao Tome, 15.vii.1974, 5 females 1 male from cocoa flowers; Kenya, Nairobi National Museum, 6.ix.1986, 25 females, 6 males from Whitefieldia elongata flowers. The males in the series from Nairobi are unusual in that they are almost yellow rather than brown in colour. The countries from which this species is now recorded are: Ghana, Sierra Leone, Nigeria, San Tome, Angola, Congo, Mozambique, Kenya and Uganda. Retanathrips gen. n. Mound & Nickle Type-species Physothrips funestus Hood, 1915 Members of this new genus apparently share the character states of Ceratothripoides listed above, judging from the few available specimens and the published descriptions. These New World species differ from the species of Ceratothripoides as follows: tergite VIII with posteromarginal comb absent at least medially; sternite VII setae S2 close to posterior margin; ocellar setae pair I arranged transversely. The five species listed by Mound & Marullo (1996) as “Ceratothripoides” are here re-assigned to this new genus. However, since these species are known from so very few specimens, their structure and identity requires further study on freshly collected material. Species included. Retanathrips chilticus (Johansen, 1986: 354) comb.n. Described in Ceratothripoides, this species is based on a single female, collected from Datura arborea flowers, 26.vii.1981, in Hidalgo, Mexico. The original illustration indicated that the comb on the posterior margin of tergite VIII is very weak medially. Retanathrips funestus (Hood, 1915: 24) comb.n. Described in Physothrips from Texas, this species is considered a senior synonym of Taeniothrips martellorum Medina Gaud from Puerto Rico (Mound & Marullo, 1996). Specimens have also been examined from Mexico, Jamaica, Trinidad and Costa Rica. Retanathrips inconstantis (Johansen & Mojica, 1986: 372) comb.n. Described in Wegenerithrips, this species is based on two females, collected from mosses, 24.x.1982, in Veracruz, Mexico. The description indicated that there is no posteromarginal comb on tergite VIII, and the setal row on the forewing first vein is almost complete. Retanathrips lagoenacollus (Moulton, 1933: 130) comb.n. Described in Taeniothrips, this species is known only from two females and one male collected at ”St. Theresa, Brazil”, 27 & 29.vi.1928. Retanathrips silvestris (Hood, 1935: 83) comb.n. Described in Taeniothrips from Panama, this species is recorded from Costa Rica and Trinidad. However, as indicated by Mound & Marullo (1996) the specimens under this name are possibly merely large individuals of R. funestus. Acknowledgements The continued support of several colleagues is gratefully acknowledged: Richard zur Strassen, Senckenberg Museum, Frankfurt (SMF); Jon Martin, Natural History Museum, London (BMNH); Sueo Nakahara, U.S. Department of Agriculture, Beltsville, Maryland, USA (USNM). We are grateful to Dr Masami Masumoto of Yokohama Plant Quarantine Station, for many valuable comments on classification problems among Thripinae, and to Prof. Irma Cabrera of Puerto Rico for sending specimens for identification. Gary L. Miller and Matt Buffington kindly provided comments on a draft. 62 · Zootaxa 2230 © 2009 Magnolia Press MOUND & NICKLE References Bagnall, R.S. (1913) Brief descriptions of new Thysanoptera. I. Annals and Magazine of Natural History, (8)12, 290– 299. Bagnall, R.S. (1918a) Brief descriptions of new Thysanoptera IX. Annals and Magazine of Natural History, (9)1, 201– 221. Bagnall, R.S. (1918b) On the rubber thrips (Physothrips funtumiae Bagn.) and its allies. Bulletin of entomological Research, 9, 65–70. Bhatti, J.S. (1990) Catalogue of insects of the Order Terebrantia from the Indian Subregion. Zoology (Journal of Pure and Applied Zoology), 2, 205–352. Bhatti, J.S. (2003) The genera Tenothrips and Ewartithrips (Terebrantia: Thripidae) and pigmented facets of eye in some Terebrantia. Thysanoptera 2003, 1–10 Halaweh, N. & Poehling, H.-M. (2009) Inheritance of vector competence by the thrips Ceratothripoides claratris (Shumsher) (Thysanoptera: Thripidae). Journal of Applied Entomology, 133, 386–393. Hood, J.D. (1915) Descriptions of new American Thysanoptera. Insecutor inscitiae menstruus, 3, 1–40. Hood, J.D. (1918) A new Physothrips from western Africa (Thysanoptera). Insecutor inscitiae menstruus, 6, 116. Hood, J.D. (1935) Ten new Thysanoptera from Panama. Proceedings of the Biological Society of Washington, 48, 83– 106. Johansen, R.M. (1986) Una especie nueva Mexicana de Ceratothripoides Bagnall (Insect; Thysanoptera; Thripidae) de la Sierra Madre Oriental. Anales del Instituto de Biologia. Universidad Nacional de Mexico, 56, 353–358. Johansen, R.M. & Mojica, A.M. (1986) Estudio de revision del genero Wegenerithrips Johansen, 1983 (Insecta; Thysanoptera; Thripidae) de trips briofagos. Anales del Instituto de Biologia. Universidad Nacional de Mexico, 56, 365–382. Karny, H. (1925) On some tropical Thysanoptera. Bulletin of Entomological Research, 16, 125–142. Masumoto, M. & Okajima, S. (2005) Trichromothrips Priesner (Thysanoptera, Thripidae) of Japan and Taiwan, with descriptions of four new species and a review of the Trichromothrips group of genera. Zootaxa, 1082, 1–27. Moulton, D. (1933) The Thysanoptera of South America II. Revista de Entomologia, 3, 96–133. Mound, L.A. (2009a) Thysanoptera (Thrips) of the World – a checklist. http://www.ento.csiro.au/thysanoptera/ worldthrips.html Mound, L.A. (2009b) Sternal pore plates (glandular areas) of male Thripidae (Thysanoptera). Zootaxa, 2129, 29–46. Mound, L.A. & Azidah, A.A. (2009) Species of the genus Thrips (Thysanoptera) from Peninsular Malaysia, with a checklist of recorded Thripidae Zootaxa, 2023, 55–68. Mound, L.A. & Marullo, R. (1996) The Thrips of Central and South America: An Introduction. Memoirs on Entomology, International, 6, 1–488. Murai, T. (2000) Damage to tomato by Ceratothripoides claratris (Shumsher) (Thysanoptera: Thripidae) in central Thailand and a note on its parasitoid, Goetheana shakespearei Girault (Hymenoptera: Eulophidae). Applied Entomology and Zoology, 35, 505–507. Nickle, D.A. (2009) Commonly intercepted trips at U.S. ports-of-entry from Africa, Europe, and the Mediterranean. IV. Miscellaneous thripine genera excluding Frankliniella, Iridothrips, and Thrips (Thysanoptera: Thripidae). Proceedings of the Entomological Society of Washington, 111, 215–238. Pitkin, B.R. (1978) Lectotype designations of certain species of thrips described by J.D. Hood and notes on his collection (Thysanoptera). Proceedings of the Entomological Society of Washington, 80, 264–295. Premachandra, W.T.S.D., Borgemeister, C., Sétamou, M, Achilles, T. & Poehling, H-M. (2005) Spatio-Temporal Distribution of Ceratothripoides claratris (Thysanoptera: Thripidae) on Tomatoes in Thailand. Environmental Entomology, 34, 883–890. Priesner, H. (1934) Three new Thysanoptera from the Sudan. Bulletin de la Société Royal Entomologique d'Ègypte, 18, 28–32. Priesner, H. (1937) Neue Thysanopteren aus Zentral-Afrika. Revue Zoologie et Botanique africaine, 30, 169–180. Reyes, C. (1994) Thysanoptera (Hexapoda) of the Philippine Islands. Raffles Bulletin of Zoology, 42, 107–507. Seshadri, A. & Ananthakrishnan, T.N. (1954) Some new Indian Thysanoptera - 1. Indian Journal of Entomology, 16, 210–226. Shumsher, S. (1946) Studies on the systematics of Indian Terebrantia. Indian Journal of Entomology, (1945) 7, 147–188. Zur Strassen, R. (1975) Thysanopterologische Notizen (3) (Insecta: Thysanoptera). Senckenbergiana biologica, 56, 75– 88. OLD-WORLD GENUS CERATOTHRIPOIDES Zootaxa 2230 © 2009 Magnolia Press · 63