Robert S. Copeland

A New Invasive Fruit Fly Species from the Bactrocera dorsalis (Hendel) Group Detected in East Africa

Abstract—A new fruit fly species suspected to be from the Bactrocera dorsalis (Hendel) group (originating from Asia), was detected during routine field surveys in the Coast Province of Kenya. Since most species in this group are of tremendous quarantine concern when introduced, and considering the fact that it has never before been detected or reported in continental Africa, surveys were immediately initiated covering a distance of over 3000 km across major fruit-growing and trading localities within Kenya, to determine the extent of spread of the new invasive species. We report on the detection of the flies, preliminary results of the survey, and discuss the potential effects of these flies on the horticulture industry in East Africa.Résumé—Une nouvelle mouche des fruits appartenant vraisemblablement au groupe Bactrocera dorsalis (Hendel) (originaire dTnde), a été trouvée lors d’enquêtes de terrain de routine dans la province côtière du Kenya. Depuis que la plupart des espèces de ce groupe font l’objet de mesure de quarantaine très sévère lors de leur introduction et considérant qu’elle n’a jamais été trouvée ou signalée sur le continent Africain auparavant, une enquête couvrant près de 3000 km à travers les principales localités productrices de fruits du Kenya a immédiatement été lancée afin de déterminer l’aire d’extension de cette nouvelle espèce invasive. Nous présentons des résultats sur la découverte de la mouche ainsi que sur l’enquête, et discutons des effets potentiels de cette mouche sur l’industrie horticole de l’Afrique de l’Est.

Parasitoids of medfly, Ceratitis capitata, and related tephritids in Kenyan coffee: a predominantly koinobiont assemblage

Arabica coffee was sampled from two sites in the central highlands of Kenya (Rurima, Ruiru) and one site on the western side of the Rift Valley (Koru). Three species of ceratitidine Tephritidae, Ceratitis capitata (Wiedemann), C. rosa Karsch and Trirhithrum coffeae Bezzi, were reared from sites in the central highlands, and an additional species, C. anonae Graham, was recovered from the western-most site. Ten species of parasitic Hymenoptera were reared from these tephritids. The parasitoid assemblage was dominated by koinobionts. Eight of the species are koinobiont endoparasitoids, but only one idiobiont larval ectoparasitoid was reared, and only one idiobiont pupal endoparasitoid. The effects of sampling bias on determination of parasitoid assemblage size associated with concealed hosts are discussed. The potential for use of these parasitoids in biological control is also discussed. Most of the parasitoid species recovered during this study are capable of developing on C. capitata, while several also attack C. rosa. Both flies are notorious pests of tropical and subtropical fruits.

Indigenous hosts of Ceratitis capitata (Diptera: Tephritidae) in Kenya.

Abstract To study the relationship of Mediterranean fruit fly or medfly, Ceratitis capitata (Wiedemann), to native plant hosts in an area within its original home range, fruits were sampled in diverse areas of Kenya from 1999 to 2001. Sampling effort was concentrated in and around forested areas in coastal, central highland, and western highland habitats. Medflies were reared from fruits of 55 species of plants, 51 of them indigenous; 46 of these species represent previously unknown hosts in Africa. Fruits infested by C. capitata were collected in all study sites, east and west of the Gregory Rift Valley, in xeric habitats between the coast and the central highlands, and at altitudes from sea level to 2,164 m above sea level. The conditions for year-round breeding of medfly in indigenous fruits are present at the coast, and possibly in highland areas as well. Infestation indices were comparable to those reported elsewhere from cultivated fruits. Although polyphagous in its home range, C. capitata was not distributed uniformly among species within two important host-plant families, Sapotaceae and Rubiaceae.

COLONIZATION OF FOPIUS CERATITIVORUS, A NEWLY DISCOVERED AFRICAN EGG-PUPAL PARASITOID (HYMENOPTERA: BRACONIDAE) OF CERATITIS CAPITATA (DIPTERA: TEPHRITIDAE)

Abstract Fopius ceratitivorus Wharton is a recently discovered braconid parasitoid of the Mediterranean fruit fly (= medfly), Ceratitis capitata (Wied.). Unlike other parasitoids previously used in medfly biological control, F. ceratitivorus was originally collected from medfly in its purported region of origin, east Africa. Shipments of Ceratitis spp. pupae from Kenya to a newly constructed quarantine facility in Guatemala yielded both F. ceratitivorus and its congener F. caudatus (Szèpligeti). Only the former species was successfully colonized through the use of medfly infested coffee berries. In the process of colonization it was determined that F. ceratitivorus oviposited into the eggs and recently hatched larvae of medflies and completed development in the hosts’ puparia. This is a relatively rare behavior among fruit fly parasitoids and, because tephritid eggs near the surface of fruits are particularly vulnerable to attack, one that might contribute to its success as a biological control agent.

Folding wings like a cockroach: A review of transverse wing folding ensign wasps (Hymenoptera: Evaniidae: Afrevania and Trissevania)

We revise two relatively rare ensign wasp genera, whose species are restricted to Sub-Saharan Africa: Afrevania and Trissevania. Afrevania longipetiolata sp. nov., Trissevania heatherae sp. nov., T. hugoi sp. nov., T. mrimaensis sp. nov. and T. slideri sp. nov. are described, males and females of T. anemotis and Afrevania leroyi are redescribed, and an identification key for Trissevaniini is provided. We argue that Trissevania mrimaensis sp. nov. and T. heatherae sp. nov. populations are vulnerable, given their limited distributions and threats from mining activities in Kenya. We hypothesize that these taxa together comprise a monophyletic lineage, Trissevaniini, tr. nov., the members of which share the ability to fold their fore wings along two intersecting fold lines. Although wing folding of this type has been described for the hind wing of some insects four-plane wing folding of the fore wing has never been documented. The wing folding mechanism and the pattern of wing folds of Trissevaniini is shared only with some cockroach species (Blattodea). It is an interesting coincidence that all evaniids are predators of cockroach eggs. The major wing fold lines of Trissevaniini likely are not homologous to any known longitudinal anatomical structures on the wings of other Evaniidae. Members of the new tribe share the presence of a coupling mechanism between the fore wing and the mesosoma that is composed of a setal patch on the mesosoma and the retinaculum of the fore wing. While the setal patch is an evolutionary novelty, the retinaculum, which originally evolved to facilitate fore and hind wing coupling in Hymenoptera, exemplifies morphological exaptation. We also refine and clarify the Semantic Phenotype approach used in previous taxonomic revisions and explore the consequences of merging new with existing data. The way that semantic statements are formulated can evolve in parallel, alongside improvements to the ontologies themselves.

Host Records for Fruit-Feeding Afrotropical Tortricidae (Lepidoptera)

We present host records for the following Afrotropical Tortricidae reared from fruit in Kenya : Idiothauma nr africanum Walsingham, Paraccra mimesa Razowski, Apotoforma nr uncifera Razowski, Eugnosta percnoptila (Meyrick), Phtheochroa aarviki Razowski & Brown, Actihema hemiacta (Meyrick), Choristoneura dinota (Meyrick), Choristoneura occidentalis (Walsingham), Procrica nr ophiograpta (Meyrick), Metamesia nr episema (Diakonoff), Epichoristodes acerbella (Walker), Cryptaspasma phycitinana Aarvik, Cryptaspasma subtilis Diakonoff complex, Cryptaspasma caryothia (Meyrick) complex, Cryptaspasma n. sp., Lobesia vanillana (Joannis), Lobesia semosa Diakonoff, Eccopsis nebulana Walsingham, Eccopsis praecedens Walsingham, Afrothreutes madoffei Aarvik, Afroploce nr karsholti Aarvik, Metendothenia balanacma (Meyrick), Endothenia ator Razowski & Brown, Anthozela chrysoxantha Meyrick, Anthozela psychotriae Razowski & Brown, Concinocordis wilsonarum Razowski & Brown, Crocidosema plebejana Zeller, Crocidosema lantana (Busck), Gypsonoma scolopiae Razowski & Brown, Cosmetra nereidopa (Meyrick), Cosmetra sp. 1, Cosmetra sp. 2, Cosmetra sp. 3, Cosmetra podocarpivora Razowski & Brown, Cosmetra taitana Razowski & Brown, Thaumatotibia leucotreta (Meyrick), Thaumatotibia batrachopa (Meyrick), Thaumatotibia salaciae Razowski & Brown, Cryptophlebia semilunana (Saalmüller), Cryptophlebia sp. 1, Cryptophlebia peltastica (Meyrick), Cydia connara Razowski & Brown, Cydia sennae Razowski & Brown, Cydia nr choleropa (Meyrick), Fulcrifera crotalariae Razowski & Brown, Fulcrifera nr periculosa (Meyrick), Fulcrifera sp. 1, Fulcrifera sp. 2, Dracontogena continentalis Karisch, Dracontogena solii Aarvik & Karisch, Grapholita limbata Diakonoff, Grapholita mesosocia (Meyrick), Grapholita nr mesosocia (Meyrick), Selania exornata (Diakonoff), Eucosmocydia monitrix (Meyrick), Stenentoma sorindeiae Razowski & Brown, and Thylacogaster garcinivora Razowski & Brown. Nearly 75 % of all our reared tortricids are members of Grapholitini, a tribe that includes numerous pests of fruit worldwide. Approximately 77 % of the reared species are recorded from only one or two different plants species (frequently in the same plant genus). Plant families that support the greatest numbers of tortricid fruit-feeders are Rubiaceae (13 tortricid species), Ochnaceae (11 tortricid species), Sapindaceae (10 tortricid species), Rosaceae (8 tortricid species), Connaraceae (7 tortricid species) and Fabaceae (7 tortricid species). We also review previously reported food plants and provide a few new records for four foliage- and/or fruit-feeding tortricids from the Afrotopical Region, e.g. Goniotorna erratica (Diakonoff), Taiteccopsis taitana Razowski, Cydia choleropa (Meyrick) and Leguminivora ptychora (Meyrick). Identifications are based on morphology and DNA barcodes (COI).

The diversity of Chironomidae (Diptera) associated with Hydrilla verticillata (Alismatales: Hydrocharitaceae) and other aquatic macrophytes in Lake Tanganyika, Burundi.

ABSTRACT To search for potential biological control agents of the aquatic weed, Hydrilla verticillata (L.f.) Royle, emerging chironomid adults were collected from aquatic macrophytes sampled between 2007 and 2009 from near shore sites in Lake Tanganyika, Burundi. Initial surveys identified H. verticillata populations at all sampled locations between Bujumbura and Nyanza Lac. Twenty-six (26) species of Chironomidae emerged from collections of four plant species; Hydrilla, Ceratophyllum demersum variety apiculatum (Cham.) Asch., Potamogeton schweinfurthii A.Benn., and Vallisneria spiralis f. aethiopica (Fenzl) T.Durand and Schinz. Twenty-four of the chironomid species were new country records, but none of them represented undescribed species. Dicrotendipes fusconotatus (Kieffer) dominated the chironomid community, comprising 82% of 32,090 reared adults. The six most common species contributed over 96% of the total midge fauna. Most species were uncommon or rare; nine species were represented by 10 or fewer specimens. A species accumulation curve for the 25 chironomid species reared from Hydrilla suggested that our sampling completely describes the community associated with this plant in northern Lake Tanganyika. Quantitative &bgr;-diversity values indicated that chironomid communities of the two Hydrocharitaceae species, Hydrilla and Vallisneria, were most similar to each other, even though they have very different growth forms. Chironomids also emerged in greater numbers from the two Hydrocharitaceae than from the other plants. No chironomid species, including Polypedilum wittei Freeman and Polypedilum dewulfi Goetghebuer, two species formerly considered for possible biological control of Hydrilla, were specific to that plant. Polypedilum species emerged from all sampled aquatic macrophytes. No chironomid-caused damage was seen on Hydrilla. African Chironomidae do not appear to be suitable candidates for biological control of Hydrilla.

FRUIT FLIES (DIPTERA, TEPHRITIDAE) OF KAKAMEGA FOREST, KENYA

ABSTRACT A list of the Tephritidae from Kakamega Forest and it environs was compiled from published records, Malaise trap, baited trap, and sweep-net collections, rearings from flowers and fruits, and examination of museum specimens. One hundred and thirty-five species are recorded, and collection information is provided for all of them. Twenty-five of these species are undescribed, or were so at the time our study began. Frugivorous Tephritidae were reared from 134 of 378 (35%) fruit collections made within the forest. Fruits of 17 of 47 plant families (36%), and 37 of the 123 plant species (30%) yielded Tephritidae. The plant families of greatest importance in the life histories of frugivorous species were the Sapotaceae, Rubiaceae, Flacourtiaceae, Rosaceae and Solanaceae, while the Asteraceae provided hosts for most of the flower-feeding species. The affinities of the Kakamega forest tephritid fauna with those of other regions, in particular the main central and western African rain forests and the East African coastal forests, are discussed.

DNA Barcodes of Microlepidoptera Reared from Native Fruit in Kenya

This paper provides metadata for DNA barcode (COI) data in GenBank for a collection of small moths (microlepidoptera except Blastobasidae and Tortricidae) reared from native fruits throughout Kenya. This paper aims to make DNA barcode data available to document ongoing research, to contribute to the International Barcode of Life (iBOL; www.ibol.org) and Kenya Barcode of Life projects, and to encourage enhancement in identifications, in line with the concept of DNA barcode data release papers and the Fort Lauderdale principles for genetic data (Schindel et al. 2011). Many of these records represent undescribed species, and we have purposefully refrained from assigning new names until the relevant taxa can be studied in sufficient detail. Under the Fort Lauderdale principles, we ask others to refrain from assigning new species names to these records outside of the context of proper systematic study. Data for 251 sequences representing 114 barcode clusters (putative species) have been released on GenBank (accession numbers GU695820GU695823, GU695866, HQ947262, JF847884-JF847887, JN284900,KF643008KF643239, KF808331-KF808337) including the standard fields for the BARCODE data standard (Benson et al. 2012) andmore data, including images and host plants, are available on BOLD (www.boldsystems. org; Ratnasingham and Hebert 2007, 2013), in a dataset accessible using a DOI (dx.doi.org/10.5883/DS-KFML1). From 1999 to 2004, Copeland organized an insect-rearing program from native fruit collected at sites throughout Kenya (Copeland et al. 2009). Although the study was focused on fruit flies (Tephritidae) and their parasitoids, the collections also yielded many Lepidoptera. This is the third in a series of papers focused on the Lepidoptera (Adamski et al. 2010, Razowski and Brown 2012), resulting from ongoing collaboration among the International Centre of Insect Physiology and Ecology (ICIPE), the Smithsonian Institution’s National Museum of Natural History (USNM), the National Museums of Kenya (NMK), and International Barcode of Life project based at the University of Guelph. This is also a contribution to a series of papers documenting DNA barcodes for moths from Kenya (Martins et al. 2013, Miller et al. 2013).

The African Burrowing Mayfly, Povilla adusta (Ephemeroptera: Polymitarcyidae), Damages Hydrilla verticillata (Alismatales: Hydrocharitaceae) in Lake Tanganyika

ABSTRACT The submersed aquatic plant, Hydrilla verticillata (L.f.) Royle (hydrilla) is a serious invasive weed in the southern USA, but in Central African lakes, it is not considered a problem. Previous surveys in Lake Tanganyika identified 2 species of chironomid midges in the genus Polypedilum Kieffer that putatively caused tip-mining damage to hydrilla. We returned to Lake Tanganyika to further investigate the midges and to explore for other natural enemies of hydrilla. High proportions of stems collected at 2 locations south of Bujumbura were found to have bored apical meristems. No insects were observed in the burrows, but large numbers of nymphs of the African burrowing mayfly, Povilla adusta Navas, were dislodged when hydrilla stems were shaken under water. Because P. adusta is known to burrow in living aquatic plants, wood and several inorganic substrates, we hypothesized that P. adusta nymphs caused the boring damage to hydrilla. To test this hypothesis, undamaged hydrilla was exposed to P. adusta nymphs in a series of laboratory experiments. In all experiments combined, 4% of hydrilla stems exposed to P. adusta were damaged, whereas no stems in containers without P. adusta were damaged. Damage was characterized as lateral or vertical boring in apical meristems and, less frequently, in stems. To quantify the level of damage in the field, stems were collected from 2 locations on 6 occasions. The proportions of stems with bored apical meristems ranged from 5–52% and 13–57% of stems were missing apical meristems. Although Povila adusta caused substantial damage to hydrilla in Lake Tanganyika, it should not be considered for biological control because it is not a herbivore, and is known to burrow in a wide variety of substrates. The Polypedilum spp. midges earlier reported to bore into hydrilla apical meristems were not abundant in the field and not found associated with boring damage. The damage earlier attributed to the midges was likely due mostly or entirely to P. adusta. These midges should not be pursued further as candidate biological control agents of hydrilla.