Fopius Wharton, 1987

Taxonomic History / Nomenclature
Fopius Wharton, 1987: 68-69. Type species: Rhynchosteres (Fopius) silvestrii Wharton, 1987 (original designation).

Type locality of type species: Cameroon, Nkolbisson; holotype female in US National Museum.

First described as a subgenus, then elevated to generic rank by van Achterberg and Maeto (1990).
Current status: valid genus.

Prior to 1987, most of the species now placed in the genus Fopius were placed either in Opius or Biosteres, and sometimes in Diachasma or Parasteres. Wharton (1987) initially proposed the name Fopius as a subgenus of Rhynchosteres. Van Achterberg and Maeto (1990) elevated Fopius to generic status, where it currently remains. Wharton (1999) partially revised the genus, and included a tabular key to species.

Both Fopius and Rhynchosteres will key out together in the general key that accompanies the Paroffit web site (because the sternaulus is heavily sculptured in both). From there you will be referred here to the Fopius page. For differences between the two, see the diagnosis and relationships sections.
Diagnosis and Relationships
Fopius, like Diachasmimorpha, is characterized by the small second submarginal cell, the loss of hind wing RS, and the well developed hind wing 2M and m-cu. Fopius, however, has a more heavily sculptured body (especially frons, notauli, and propleuron). The species of Rhynchosteres are similarly heavily sculptured, but usually have both clypeus and mandibles greatly modified. Additionally, most species of Fopius have the hind wing m-cu slightly recurved near the wing margin.

Fopius was originally described as a subgenus of Rhynchosteres (Wharton 1987), despite the fact that the type species of Fopius lacked the highly modified, tubular clypeus originally used to characterize Rhynchosteres. Wharton’s (1987) placement of Fopius in Rhynchosteres was based on a study of several Afrotropical species that had the midventral margin of the clypeus variously thickened and protruding, suggesting a morphocline leading to the more elaborate modifications found in Rhynchosteres sensu stricto. Van Achterberg and Maeto (1990), dissatisfied with the broadening of the concept of Rhynchosteres that this entailed, removed Fopius from Rhynchosteres and accorded it separate generic rank. The Fopius desideratus species group is particularly critical for an understanding of the relationship between Fopius and Rhynchosteres. Althought the mandible lacks the modifications found in most species of Rhynchosteres, the midventral portion of the clypeus in members of the Fopius desideratus species group is at least slightly thickened and weakly protruding. Additionally, the relative length of the first two flagellomeres in the Fopius desideratus species group is as in Rhynchosteres, the petiole is similarly structured (flatter and narrower than in more typical species of Fopius such as Fopius silvestrii), and most significantly, the ovipositor sheath shows an identical pattern of setal reduction.

Occipital carina nearly always present laterally, extending dorsally more than half height of head. Labrum completely to almost completely concealed by clypeus (Figure 1); outer surface of clypeus usually relatively flat to weakly and evenly convex, ventral margin varying from thin and evenly convex throughout to variously thickened and protruding medially, sometimes as a distinct tooth. First flagellomere shorter than second in most species. Propleuron with oblique carina immediately dorsad propleural flange, the carina bordered ventrally by crenulate to rugose sculpture. Notauli deep, complete to midpit, crenulate throughout (Figure 2). Postpectal carina present. Hind tibia dorso-posteriorly without basal carina. Fore wing (Figure 3) with second submarginal cell short; m-cu arising basad or directly in line with 2RS. Hind wing (Figure 4) with RS virtually absent in nearly all species (represented by a very weakly infumate crease in caudatus); m-cu well-developed as a distinctly pigmented crease extending nearly to wing margin in nearly all species, rarely (5%) poorly developed in some individuals, extending only half way to wing margin in caudatus. Metasoma of several species with second and rarely third terga striate; metasoma otherwise unsculptured beyond petiole. Ovipositor long, extending well beyond the tip of the abdomen.
1. Fopius silvestrii face...
2. Fopius arisanus dorsa...
3.Fopius arisanus fore wing.
4.Fopius arisanus hind wing.
Fopius is an Old World group, with nearly all of its 25 described species (as of 1999) from the tropics (Wharton 1999).

Chinajariyawong et al. provide an extensive list of opiine species and their tephritid hosts collected in Thailand and Malaysia during surveys conducted between 1986 and 1994 (Chinajariyawong et al. 2000). The specimens were primarily collected from various Bactrocera spp. and include the following Fopus species: arisanus (Sonan), deeralensis (Fullaway), persulcatus (Silvestri), skinneri (Fullaway), and vandenboschi (Fullaway) with arisanus and vandenboschi being the most commonly collected species of Fopius.

No referenced distribution records have been added to the database for this OTU.
Identification of Species and Subspecies
There are no keys available that cover all of the species of Fopius that have been reared from fruit-infesting Tephritidae. However, keys to most of these species can be found in Wharton and Gilstrap (1983) and Fischer (1987), with both works treating the species under the generic name Biosteres. Wharton (1999) also presented a tabular summary of morphological characteristics for all the known species. Only one species, Fopius okekai Kimani-Njogu and Wharton, 2002, has been described since then.

Fopius persulcatus species group

The Fopius persulcatus species group is characterized largely by striate sculpture on the second metasomal tergum. The frons is also densely covered with large punctures, these sometimes coalescing to form rugulose or rugulo-striate sculpture. This species group consists of Fopius alternatae (Tobias) and Fopius myolejae (Tobias) from Russia; and the Indo-Pacific species Fopius arisanus (Sonan), Fopius carpomyiae (Silvestri), Fopius persulcatus (Silvestri), Fopius skinneri (Fullaway), and Fopius vandenboschi (Fullaway). The species are very similar to one another, but differ primarily in coloration, length of ovipositor, and configuration of the ovipositor tip. Following their successful introduction to Hawaii during the biological control program against oriental fruit fly (Clausen et al. 1965), Fopius arisanus and Fopius vandenboschi were intensively studied, and much is now known about their biology (with most of the early literature on arisanus published under the name Opius oophilus Fullaway). Other than the original host records, however, little is known about the other species. Data on persulcatus are particularly problematic because of widespread confusion regarding the identity of this species during the Hawaiian oriental fruit fly program, and the subsequent description of several subspecies (Fischer 1963). Though the best known species are parasitoids of tephritids in the dacine genus Bactrocera Macquart in their native range, other species have been reared from trypetine tephritids in the genera Carpomya Costa, Myoleja Rondani, and Rhagoletis Loew. Based on the similarities in the shape of the ovipositor tip, these species preferentially attack either the egg or early instar larva of their host (though all eventually emerge from the puparium). This biology, however, has only been confirmed for arisanus (attacking eggs) and vandenboschi (attacking primarily first instars). The species of this group are known from Pakistan and India east through Indonesia and north through Taiwan, Japan, and far eastern Russia.

Fopius desideratus species group.

Members of this species group that have been reared from fruit-infesting tephritids include Fopius bevisi (Brues), Fopius desideratus (Bridwell), Fopius niger (Szépligeti), Fopius ottotomoanus (Fullaway) and Fopius okekai Kimani-Njogu and Wharton. As noted above, this group is characterized by the reduced setae on the ovipositor sheath, slight thickening medially along the ventral margin of the clypeus, and a longer, somewhat flatter petiole than is found in members of the Fopius silvestrii group (discussed below). The punctation on the frons tends to be weaker, with transverse striae usually clearly present medially. The species of the Fopius desideratus species group are very similar to one another, differing primarily in color. Members of this species group have been collected from Cameroon and Nigeria east to Kenya and Tanzania as well as South Africa and Madagascar, and undoubtedly occur throughout subsaharan Africa. Most but not all of the specimens reared to date have been from Dacus infesting Cucurbitaceae. The few remaining records were from ceratitidine tephritids on various hosts. All members of this species group have large, subapical nodes on the ovipositor, suggesting that they may attack late instar larvae of their hosts.

Fopius silvestrii species group.

This group is currently defined largely by the absence of features that define the two species groups already mentioned: the second metasomal tergum is unsculptured and the setae on the ovipositor sheath are not reduced. Both Fopius silvestrii (Wharton) and Fopius ceratitivorus Wharton have been reared from medfly, Ceratitis capitata, infesting coffee, and Fopius silvestrii has also been reared from Dacus bivittatus (Bigot) infesting squash (Steck et al. 1986, Wharton 1987). The two other members of this species group have not been reared. Members of the Fopius silvestrii species group have much the same distribution pattern as those of the Fopius desideratus species group, and are differentiated from one another largely by color (Fopius silvestrii is dark and Fopius ceratitivorus is pale) and ovipositor length. Fopius ceratitivorus is known only from Kenya and extreme northeastern South Africa; Fopius silvestrii has been collected in West Africa and western Kenya.

Other species groups

The remaining three species, Fopius caudatus (Szépligeti), Fopius deeralensis (Fullaway), and Fopius schlingeri Wharton, do not readily cluster into distinctive species groups. Fopius caudatus is readily separated from all other species of Fopius by the distinctive band of setae and punctures on the frons extending between the eye and ocelli. Fopius caudatus resembles members of the Fopius desideratus species group in the morphology of the clypeus and petiole, but has a distinctly different ovipositor (strongly narrowed, suggesting oviposition in the host egg) as well as several features unusual for members of the genus Fopius (Wharton 1997). The commonly collected Fopius caudatus also superficially resembles Fopius silvestrii, and both have been reared from ceratitidines in coffee samples. Fopius caudatus has thus far been reared exclusively from ceratitidines (Steck et al. 1986, Wharton et al. 2000), and is known from tropical regions of both eastern and western Africa.

Both Fopius deeralensis and Fopius schlingeri are from Queensland, where they have been reared from various species of Bactrocera in a variety of host fruits (Clausen et al. 1965, Wharton 1997). Fopius schlingeri, for example, attacks at least four species of Bactrocera developing in the fruit of at least three plant families. Fopius deeralensis differs from Fopius schlingeri in having the ventral margin of the clypeus distinctly pointed midventrally, a distinct subapical ridge on the ovipositor, and the second metasomal tergum is usually weakly striate, at least basally. There are slight differences among species of Fopius in the exposure of the labrum and Fopius schlingeri provides a good example of a species with a partially exposed labrum. The ovipositor in Fopius schlingeri is virtually identical in form to that of Fopius arisanus, strongly suggesting a biology similar to the latter in which the parasitoid oviposits into the egg of its host. Field observations kindly supplied by Greg Quimio during his PhD research at the University of Queensland appear to confirm this. These two Australian species differ in only minor ways from members of the Fopius persulcatus group, and may need to be included there after further study.

1. Fopius arisanus metas...
2.Fopius arisanus fore wing.
3.Fopius arisanus hind wing.
4.Fopius arisanus lateral habitus.
5. Fopius arisanus b...
6. Fopius arisanus showi...
7.Fopius caudatus habitus
8.Fopius ceratitivorus habitus
9.Fopius silvestrii habitus.
10. Fopius silvestrii face...
11.Fopius vandenboschi dark form.
12.Fopius vandenboschi pale form.
Biology / Hosts
All species of Fopius reared as of 2000 are parasitoids of Tephritidae. A few generalizations can be made about hosts and distribution patterns, even though our current knowledge is somewhat limited. Native hosts in all of the species groups of Fopius treated here include members of the subtribes Ceratitidina and/or Dacina, both in the tephritid tribe Dacini, subfamily Trypetinae (Norrbom et al. 1998). Except where they have been introduced for biological control, members of the persulcatus species group occur outside the range of fruit-infesting Ceratitidina, and several of them have been reared from members of the tribe Trypetini in addition to members of the Dacina. Rearing records from within their native ranges (Clausen et al. 1965, Steck et al. 1986) suggest that the 17 species of Fopius for which we have host records are restricted to fruit-infesting tephritids, but that there are different levels of specificity. Some are currently known only from a single host, others (e. g. Fopius caudatus on ceratitidines) have only been reared from a narrow group of hosts, and several have been reared from both Dacini and Trypetini. Where introduced outside their native range for biological control, Fopius arisanus and Fopius vandenboschi have been able to attack other fruit-infesting tephritids (Clausen et al. 1965, Wharton et al. 1981). Although some of the species of Diachasmimorpha introduced to Hawaii to control fruit-infesting Tephritidae occasionally attack gall-making (but not flower-infesting) Tephritidae, Fopius arisanus and Fopius vandenboschi do not (Duan et al. 1996).

Unless you are already on this page, considerably more information on species groups and use in biological control can be found on the Fopius page (and associated links) of the Paroffit website page:

Biology and Behavior
Oviposition in the egg stage of the host is an intriguing biological feature, and one that has been of considerable interest to biological control workers. This phenomenon has been well documented in Fopius arisanus (Sonan), (most of the biological information on this species was published under the species name oophilus Fullaway), perhaps the most significant parasitoid introduced to Hawaii for tephritid biological control (Clausen et al. 1965). Evidence that this biological trait is found in several other species of Fopius is gradually accumulating (see discussion by Waterhouse 1993). Based solely on ovipositor morphology, it is likely that the following species share this trait with arisanus: Fopius carpomyiae (Silvestri), Fopius persulcatus (Silvestri), Fopius schlingeri Wharton and Fopius caudatus (Szépligeti) (Wharton 1999). More recently, several studies have demonstrated that both Fopius caudatus and Fopius ceratitivorus can successfully oviposit in host eggs (Wharton et al. 2000, Lopez et al. 2003, Wang et al. 2004, and Bokonon-Ganta et al. 2005).

There are no specimens currently determined for this OTU, or those specimens determined for this OTU are not yet mappable.

This page was assembled by Bob Wharton and Danielle Restuccia. It is part of a review of the genera of World Opiinae, conducted at Texas A&M University. We are particularly grateful to Xanthe Shirley, Andrew Ly, Patricia Mullins, Trent Hawkins, Lauren Ward, Cheryl Hyde, Karl Roeder, and Andrea Walker, who did nearly all of the imaging (together with Danielle) for this project. Matt Yoder and Istvan Miko provided guidance on databasing issues associated with our use of mx and HAO respectively. This project would not have been possible without the kindness of many curators at museums throughout the world who gave generously of their time to Bob Wharton and his students. In particular, I thank Henry Townes (deceased) and David Wahl (American Entomological Institute, Gainesville), Gordon Nishida (Bernice P. Bishop Museum, Honolulu), Norm Penny, and Bob Zuparko (California Academy of Sciences, San Francisco), Bill Mason (deceased), Mike Sharkey, Andrew Bennett, and Henri Goulet (Canadian National Collection, Ottawa), Paul Dessart (deceased) (Institut Royal des Sciences Naturelles de Belgique, Brussels), Marc De Meyer (Koninklijk Museum voor Midden-Afrika, Tervuren), Axel Bachmann (Museo Argentino de Ciencias Natureles, Buenos Aires), Eberhard Koenigsmann (deceased) and Frank Koch (Museum fuer Naturkunde der Humboldt-Universitaet, Berlin), J. Casevitz Weulersse and Claire Villemant (Museum National d’Historie Naturelle, Paris), James O’Connor (National Museum of Ireland, Dublin), Jenö Papp (National Museum of Natural History, Budapest), Kees van Achterberg (National Museum of Natural History, Leiden), Max Fischer, Herb Zettel, and Dominique Zimmermann (Naturhistorisches Museum, Wien), Per Persson and Lars-Åke Janzon (Naturhistoriska Riksmuseet, Stockholm), Ermenegildo Tremblay (Silvestri Collection, Portici), Erasmus Haeselbarth (Staatliche Naturwissenschaftliche Sammlungen Bayerns, Munich), Tom Huddleston and Gavin Broad (The Natural History Museum, London), Paul Marsh and Robert Kula (USDA Systematic Research Laboratory and US National Museum of Natural History, Washington, D. C.), Vladimir Tobias (deceased) and Sergey Belokobylskij (Zoological Institute, Academy of Sciences, St. Petersburg), and Roy Danielsson (Zoological Institute, Department of Systematics, Lund) for facilitating loans and general assistance associated with examination of holotypes and other material in their care. This work was supported largely by NSF/PEET DEB 0328922 and 0949027, with REU supplements 0723663, 1026618, 1213790, and 1313933 (to Wharton). Page last updated July, 2015. The material on this page is freely available, but should be acknowledged if used elsewhere.

This material is based upon work supported by the National Science Foundation under Grant Numbers DEB 9300517, DEB (PEET) 9712543, DEB (PEET) 0328922 with REU supplements 0723663 and 1026618 and DEB 0949027 with REU supplements 1213790 and 1313933. Any opinions, findings, and conclusions or recommendations expressed in this material are those of the author(s) and do not necessarily reflect the views of the National Science Foundation.