Bulletin of the British Museum (Natural History) A survey of the Ophioninae (Hymenoptera: Ichneumonidae) of tropical Mesoamerica with special reference to the fauna of Costa Rica Ian D. Gauld Entomology series Vol 57 No 1 25 August 1988 The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series. Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff of the Museum and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. Parts are published at irregular intervals as they become ready, each is complete in itself, available separately, and individually priced. Volumes contain about 300 pages and several volumes may appear within a calendar year. Subscriptions may be placed for one or more of the series on either an Annual or Per Volume basis. Prices vary according to the contents of the individual parts. Orders and enquiries should be sent to: Publications Sales, British Museum (Natural History), Cromwell Road, London SW7 5BD, England. World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.) © British Museum (Natural History), 1988 The Entomology series is produced under the editorship of the Keeper of Entomology: Laurence A. Mound Publications Manager (Entomology): W. Gerald Tremewan ISBN 0 565 06033 3 ISSN 0524-6431 Entomology series Vol 57 No 1 pp 1-309 British Museum (Natural History) Cromwell Road London SW7 5BD Issued 25 August 1988 A survey of the Ophioninae (Hymenoptera: Ichneumonidae) of tropical Mesoamerica with special reference to the fauna of Costa Rica Ian D. Gauld Department of Entomology, British Museum (Natural History), Cromwell Road, London SW7 5BD Contents PARAIGHISISWE aw etl» tans ea ta sols whe torte sade eine rae rar coe TPA Nel gis gemieate sie es GN ew wae wan y 2 DIREC GIGI DE AN. aire fa Ae cet a ee) oe emai rine a bad aa e+ Ae ee Aide ees 2 SPeneAL AIG materral StUced! +e afar nin. deceit: tare erdtls hin Ue tinge bidet hate ea bs 3 Serer OTA MiAl SCOIE: On tle SUNCLY sete csi aicts nisin sickays ais]ele 5 inva, sale eiopnin sini gic, 2a aw <0 3 Representation and distribution of material examined ......................0.. 3 MEOSIOMes Of WAtelAleXAMINeG yan cc. ides ces cae en ss wisea ese ee ns eo 4 eEUMO MOU ants tee comin te Maret, Sens eee ee CLT OMe OER. Mate eee 4 aEeelesicriteraanGettioal Characters 0: iw 'adel: Shstes 5 Ps hea theslohies Su aero 6 An historical résumé of work on the Mesoamerican Ophioninae .................. 9 IEseomCOhr REY GMINIELACN Mamet aA re cs Ph pt betel ied Rime cyaea tha tibet ear aha sca eel Ri wks Pees. « 9 Ita C ROSS A sohrhtey tush opti ta ade ishce Monat lotr n Sew oe wed HA 9 RACER CUOMO tert oivab eshte lack ambaysertey hs, «arden le ayaa Rhar¥ied kyu lar eianyenarbas «lave 10 ea aaa E rt ACN, VE (EIA PR OMUEN Sys saiscealy sisal rate minadl'n, nts Wine (a tak le «Giablnfa's > aibie slayer neidiayes 10 Sram ENED CMU O TAG) OIMIINAC: ex seas a Satya tee ci wets a aya 08s ards) andvasrwhnrg) Ameen taninioahie falls 10 TRUS EUUT Any LUSCH Wont iuearuabahtc th aetna swe vic ansh «be iade, Care “IGM a (o."eid_ by'a.a cha whsha rays. «el «CarmivQere pe velo are 10 RePereS OE CLULCUUs re rasa atte ciel cieatecaekrd.d atin 2 ; oe TERGITE 1 OF GASTER STERNITES ANTERIOR AREA SPIRACULAR AREA POSTERIOR AREA METAPLEURON & Figs 1, 2 Stylized ophionines labelled to show parts. 1, Ophion alitrunk, lateral view. 2, Enicospilus propodeum and anterior part of gaster, lateral view. Abbreviations: ATC = anterior transverse carina of propodeum; EpC = epicnemial carina; ESS = episternal scrobe; LLC = lateral longitudinal carina of propodeum; LT = laterotergite of tergite 2; N3 = metanotum; PTC = posterior transverse carina of propodeum; PTCM = posterior transverse carina of mesosternum; SAP = subalar prominence; Tg = tegula. 6 IAN D. GAULD current standard terminology is unsuitable for it. In this work the nomenclature adopted by Gauld & Mitchell (1978; 1981) is followed (Fig. 2). Various measurements and indices have been extensively used and these require explanation. The width of the face is the minimum distance between the eyes, and its height is the median vertical distance from the clypeal margin to the facial tubercle. The scutellar length and breadth are defined respectively as the length from the posterior rim of the scuto-scutellar groove to the posterior margin of the scutellum, and the distance across the posterior rim of the scuto-scutellar groove between the lateral carinae. The length of the hind trochantellus is measured mediodor- sally and the length and depth of the hind coxa are the maximum values measured in profile. The indices used are defined thus: Alar index of fore wing _ length of 1m-cu between bulla and 2m-cu Al length of 3rs-m Cubital index of fore wing length of Cul between Im-cu and Cula CI= length of Culb Frontal index of head Y maximum diameter of median ocellus distance between eyes through median ocellus Intercubital index I= length of 3rs-m ~ length of M between 2m-cu and 3rs-m Second discoidal index _ length of Cula between Culb and 2m-cu ~ length of Cul between Rs&M and 1m-cu See also Fig. 4. The geographical names used broadly follow those given in the Times Atlas of the World. Many Costa Rican localities are not shown in this atlas, but most are given on the 1 : 200 000 series of maps published by the Instituto Geografico Nacional, San José, Costa Rica. The Costa Rican localities where most collecting was undertaken are shown on Map 1. The county names given for United States localities are shown in the Rand McNally Road Atlas. Distances and altitudes are given in kilometres and metres irrespective of the units shown on original labels. Species criteria and critical characters Some contemporary ichneumonid taxonomists favour the use of subspecies for any population that differs slightly in colour or sculpture from the ‘typical’ form, but increasingly the more progressive workers are abandoning this category which is not only a nomenclatural encumbrance, but can be biologically misleading (see Gauld & Mitchell, 1978; 1981; Thorpe, 1980; Gauld, 1984a). Consequently these categories are not used in this study. Certain species are extraordinarily variable in both colour and sculpture, but usually this variation does not coincide well with geographical distribution; rather the frequency of a particular form of variation amongst individuals in a population changes from region to region. This is most marked in a number of very common species that inhabit disturbed ecosystems, such as agricultural areas, in which selective pressures are likely to favour species that are capable of attacking a variety of hosts in a variety of ephemeral niches. It is not surprising that such species OPHIONINAE OF TROPICAL MESOAMERICA 7 should exhibit great phenotypic variation, as it is well known that development in different hosts or under different microclimatic regimes can profoundly influence the morphology or colora- tion of an insect (Myers, 1977; Liu & Carver, 1982; Pungerl, 1986). Species that inhabit climatically very stable habitats, such as montane rainforests, are often morphologically and chromatically extremely uniform throughout their range. Gauld & Mitchell (1978; 1981) and Gauld (1985) have given exhaustive accounts of many of the critical characters of ophionines and this information is not repeated here. However, it is worth emphasizing that certain structural features (particularly the shape of mandibles, length of antennae, relative lengths of wing veins, form and number of the alar sclerites, form of the tarsal claws, stoutness of the gaster and pilosity of the terminal male sternites), which have been PTEROSTIGMA MARGINAL CELL Rs+2r Rs&M eal p tens T Trib BASAL CELL =e ——— = ee i a Pan Lore i Me eat Impcu Mae . eS ys hee 2nd DISCAL CELL2™\cU = _|| 1st suspiscaL CELL Cula Cul 1A 2 ~~ —— a ae a se Cu et ae R BASAL opr ae S MARGINAL 1A M a CELL < Rs u-a Cul M 1A MARGINAL CELL RAMELLUS PROXIMAL SCLERITE >= DISTAL SCLERITE oa Sag Se QUADRA ete! MARGIN OF FENESTRA Figs 3-5 Stylized ophionine wings labelled to show parts. 3, Ophion, fore and hind wings. 4, Enicospilus, distal part of fore wing. Indices used in text are measured from the points indicated by letters d-g and w-z; Al = xy/wz; Cl = ef/fh; ICI = wz/wx; SDI = fg/de. 5, Enicospilus, alar sclerites of fore wing. IAN D. GAULD UdSIIA ‘ZT [PALasaY ISA10.J UOWRY URS *[] /PUOLILD ‘(| SAALASAY IPsJIAIUOY] *6 = IL ‘1 {spues] soseparoinyy] ‘Sof URS LIS] “9 SY1eg [PUONLN BSOY PUPS ‘¢ {eyoRH [Pe OAD ‘Z VWYVNVd ‘ wae ewe ee eee en ‘yivg [PUOTLN d}skovueNny poiepap A[MIU ay} UT papnjoul MOU oP O-Z SUG “BLaqr] = J “led [PUOHRN Opeaodion ‘ZZ ‘seuinbsq *[Z :eIOq Aap OpsvssyH urg “7 ‘quedey ‘6] ‘nzeosq ap o1uojuy ues ‘gy :eqyeLiny ‘LT Sie” [LUONLN OsANINYOL ‘OT ‘Vajag eT VOULY “CT SYAe_ [LUONLN OLD o1nesg “py syeq [PUONLN SVOd UPI[OA “ET -O11090$ 9p oz® 6le@ o6pj105 B Zl ° 8le asop ung . d [euoneN lat, Bap UOUTY *g ‘faliqey ues voULy ¢ foRdR_D URIJOA UO OSA UOIDRISY “p :ISO1O ULI[OA UO eZj}LIRY] BSe>) VWNOVAVIIN ‘RI]IIAD BIULS ‘T “says SuNda]joo auruorydo yedioutad ay) yo suoneso} ayeunxoidde oy} Suimoys ‘vory eIsoD f dey NV35D0 Didlovd OPHIONINAE OF TROPICAL MESOAMERICA 9 used extensively to separate species in other regions, are also reliable for species separation in Mesoamerica. Similarly other features (such as the sculpture of the mesopleuron, metapleuron and posterior area of the propodeum) often show a considerable degree of intraspecific variation and are less reliable indicators of species-distinctness. Some Mesoamerican species, especially in the genus Enicospilus, have distinctive structural differences which have not been observed in species in other regions. For example, several Neotropical Enicospilus species have modified or incomplete posterior transverse carinae of the mesosternum (this carina is complete and unspecialized in almost all species from other regions), and a few have the second laterotergite pendant (it is folded up under the tergite in all species in all other regions). The position and completeness of the lower end of the occipital carina is also an important feature of several Neotropical species of Enicospilus and Thyreodon. A number of other critical characters of this latter genus are discussed by Porter (1984). An historical résumé of work on the Mesoamerican Ophioninae Although a few of the more widespread New World ophionines were described by classical authors (e.g. Christ, 1791; Brullé, 1846) no attempt was made to document the Mesoamerican fauna until the work of the early North American ichneumonologists E. Norton (1863) and E. T. Cresson (1865) who described a small number of north Mexican and Cuban species; in particular the latter author described a number of species of Thyreodon. Little further was published until 1886 when P. Cameron’s work in Biologia Centrali-Americana appeared. Cameron’s study was based on a fairly small collection of Mexican, Guatemalan, Costa Rican and Panamanian species made by the Godman-Salvin expedition, but it represents a first attempt at understand- ing the tropical Mesoamerican continental fauna. At the beginning of the twentieth century the European hymenopterists G. V. Szépligeti (1906) and C. Morley (1912) added a few more species, but the most comprehensive work of this period was a revision of the entire New World Ophioninae by C. W. Hooker (1912). Hooker recognized nine genera as occurring in tropical America, but two of these (Retanisia and Ophiopterus) are currently placed in other ichneumonid subfamilies. Little further descriptive work was undertaken until 1947 when R. A. Cushman revised the generic classification of the Ophioninae. He recognized eight genera, Thyreodon, Athyreodon, Rhynchophion, Ophion, Ophiogastrella, Agathophiona, Aulophion and Enicospilus as occurring in tropical America. H. Townes (1971) produced a further generic reclassification of the Ophioninae and provided keys to the world genera, but these have now been rendered partially obsolete by the work of Gauld (1979; 1985). Gauld & Lanfranco (in press) have published a synopsis of the Latin American ophionine genera together with a new generic key. At species-level, very little work has been done on the Mesoamerican ophionines after Hooker (1912) until Townes (1939) made some preliminary attempts at establishing syn- onymies. This work was continued in the important Catalogue and Reclassification of Neotropic Ichneumonidae (Townes & Townes, 1966). Except for work on Thyreodon by C. Porter (1984; 1986), there have been no recent attempts to produce keys to the diverse Mesoamerican ophionine species. The Ophioninae Diagnosis Generally large (fore wing length usually 13+ mm) slender insects with a laterally compressed gaster, long antennae and legs, and ample wings. Labrum generally exposed; mandibles bidentate. Propodeal carinae occasionally complete though usually reduced, often with posterodorsal part of propodeum modified and coarsely sculptured. Fore wing with vein 2rs-m absent, 3rs-m present, joining M well distal to 2m-cu; 1st subdiscal cell generally more sharply angled at base of Cu1a than it is at base of 1m-cu; 2nd subdiscal cell with a pigmented spurious vein extending from near end of 1A towards the hind corner of the wing; hind wing (of New World species) always with distal abscissa of Cul present. Tarsal claws pectinate. Gaster 10 IAN D. GAULD elongate; first segment more or less cylindrical anteriorly, with junctior between tergite and sternite unrecognizable; spiracles positioned near to posterior end of tergite 1; glymmae not present; ovipositor (of New World species) short, not longer than apical abdominal depth, and usually with a dorsal subapical notch. Field recognition Most of the nocturnally active ophionines (i.e. most species in the subfamily) are large, yellowish brown insects with very slender antennae and large eyes and ocelli. They are very distinctive, but inexperienced workers may confuse them with other nocturnal ichneumonids that are superficially similar. No other nocturnal Central American hymenopteran has a fore wing with venation like that of an ophionine (Fig. 3); most others (Cidaphus and most Netelia species) have both veins 2rs-m and 3rs-m present, enclosing a small cell (the areolet) (Fig. 6), but rarely if only one vein is present it is 2rs-m and this joins M opposite to or proximal to 2m-cu. Other nocturnal ichneumonids often have a distinct glymma (a pit-like structure) present in tergite 1 and have the petiolar spiracles at or before the centre of the tergite. The relatively few ophionine species that are diurnally active are predominantly black in colour with blackish or patterned wings. Although most are slender with strongly laterally compressed gasters, a few are quite stout and have short antennae. All these species may be distinguished from most other ichneumonids by the fore wing venation. Only a very few other ichneumonids have an ophionine-like venation, and these either have simple claws or entirely lack the distal abscissa of Cul in the hind wing. Classification of the group In early ichneumonid works (e.g. Morley, 1915) the name Ophioninae was applied to a large and heterogeneous group of ichneumonids with laterally compressed gasters. However, during the past 50 years this group has been subdivided into more natural taxa, and at present (Townes, 1971; Carlson, 1979; Gauld, 1985) the subfamily Ophioninae is restricted to include only those genera previously placed in the tribe Ophionini (e.g. sensu Hooker, 1912; Cushman, 1947; Townes, 1951). As currently recognized, the Ophioninae is one of the 31 subfamilies of the Ichneumonidae. Phylogenetically it is probably the sister-group of the Campopleginae (Gauld, 1985), and this pair of taxa belong to a lineage of endoparasitoids that includes the Banchinae, Ctenopelmatinae, Tersilochinae and Cremastinae There have been three major classifications proposed for the genera of Ophioninae (Cush- man, 1947; Townes, 1971; Gauld 1985). The first author recognized three groups of genera, the Ophion, Thyreodon and Enicospilus genus-groups. Townes (1971) subsequently amalgamated Cushman’s Thyreodon and Enicospilus generic groups and recognized two tribes, the Ophionini (= the Ophion genus-group) and the Enicospilini (= the Thyreodon + Enicospilus genus- groups). Gauld (1985) pointed out that whilst Townes’s Enicospilini was holophyletic, his Ophionini was a paraphyletic assemblage, and in an attempt to produce a more representatively phylogenetic classification Gauld divided the 32 world genera of Ophioninae into five holo- phyletic genus-groups, the Ophion, Sicophion and Eremotylus genus-groups (more or less corresponding to the Ophionini sensu Townes) and the Thyreodon and Enicospilus genus- groups (= Enicospilini of Townes). The large Enicospilus genus-group was further divided by Gauld into five subgroups, the Orientospilus, Ophiogastrella, Stauropoctonus, Leptophion and Enicospilus genus-subgroups. The biology of Ophioninae Natural history Ophionines are solitary koinobiont endoparasitoids of the larvae of other holometabolous insects, that is they allow the host to develop for a period after oviposition (see also Askew & Shaw, 1986). A Nearctic species of Ophion is known to parasitize a coleopteran larva (Townes, 1971) and a European species has occasionally been recorded from a sawfly host (Thompson, OPHIONINAE OF TROPICAL MESOAMERICA (ia 1957), but virtually all other rearings are from Lepidoptera. The most usual hosts are moth larvae that feed exposed on vegetation, especially species of the families Noctuidae, Lasiocam- pidae, Lymantriidae, Saturniidae, Geometridae, Arctiidae and Sphingidae. The larvae of Rhopalocera and Microlepidoptera are seldom attacked. Samples of most ophionine species seem to comprise approximately equal numbers of both sexes, and rearing confirms a normal one to one sex ratio (Rohlfs & Mack, 1985a). Of 125 specimens of Enicospilus lebophagus reared at Santa Rosa National Park, Costa Rica in 1985, 71 were females and 54 males (D. H. Janzen, pers. comm.). This is not significantly different from a 1:1 sex ratio. Mating apparently occurs soon after emergence, but has not been observed. Most male ophionines have very uniform and unspecialized genitalia, though species of Thy- reodon often have specifically different gonosquamae (Porter, 1984), and some species of Ophiogastrella and members of the Enicospilus trilineatus species-group have ornamented, flanged aedeagi. Species-characteristic secondary sexual features are observable in some spe- cies, though the functions of these various characters have never been investigated. Males of some Enicospilus species have species-specific arrangements of hair on the posterior gastral sternites, whilst the pubescence on the tarsi of males of some Thyreodon differs among species (Porter, 1984). The females of some species of Ophiogastrella have the distal segment of the fore tarsus modified and males of all species in this genus have flattened claws (see Fig. 15). Adult ophionines occur in almost all habitats. Large numbers ot species are crepuscular or strictly nocturnal (Rohlfs & Mack, 1985b). These insects are most frequently encountered at light, but sometimes they may be observed at dusk visiting flowers. Occasionally nocturnal species can be found roosting on the underside of leaves during the day. The diurnal ophionine species are more conspicuous and may often be encountered feeding from flowers or flying amongst shrub layer vegetation. Diurnally active koinobionts are probably exposed to a greater array of potential predators than are nocturnal species (Gauld, 1987), so it is not surprising to observe that many diurnal ophionines have protective devices. Some are aposematically col- oured and resemble agressive aculeates. For example, Rhynchophion flammipennis is bluish black with orange, black-tipped wings and in flight looks extremely like some species of Pepsis (Pompilidae). Others, such as Thyreodon species, have elaborate flanges that guard the cervical membranes, and probably offer protection against asilid predators, whilst a few Thyreodon species have sharp processes on the mesoscutum which may serve to deter vertebrate predators. Adult female ophionines are not known to feed on prospective hosts (Jervis & Kidd, 1986). All the females I have dissected (12 species) are apparently pro-ovigenic, that is they emerge from the pupa with a full complement of more or less mature eggs. Most individuals have between 7 and 30 mature eggs in each of their lateral oviducts. I have seen three individuals with one or no eggs present, suggesting further oocytes do not develop during active adult life. Rohlfs & Mack (1985a) observed that adult females of Ophion flavidus commenced oviposition within 24 hours of emerging from the pupa. Oviposition is into the host larva and is accomplished rapidly by the female straddling the host and thrusting her ovipositor forwards, into the host (Vickery, 1929). The ovipositor is withdrawn almost immediately. The ophionine egg is cylin- drical, slightly curved and pale coloured. No sign of paralysis has been observed in newly parasitized hosts. Many ophionines oviposit into relatively mature larvae in their third or fourth (Vickery, 1929), or fourth or fifth instar (Moutia & Courtois, 1952; Rohlfs & Mack, 1985a), though some species attack young larvae (Price, 1975). Generally a single egg is deposited free in the haemocoel. No species of Ophioninae are known to develop gregariously. It is not known if ophionines inject viral particles or physiology-manipulating venoms into their hosts. How- ever, they may do so since species of the closely related subfamily Campopleginae are well- known to do both (Vinson & Stoltz, 1986), as are species in other subfamilies in the lineage to which the Ophioninae belongs (Stoltz et a/., 1981) The ichneumonid egg hatches to produce a caudate first instar larva (Clausen, 1940) that lives in the haemocoelic cavity of the host (Moutia & Courtois, 1952). In most cases it is not known what effect parasitization has on the host caterpillar, although parasitized hosts are sometimes, but not necessarily, smaller than healthy ones. One species of Ophion has been shown to depress host-larval food consumption (Rohlfs & Mack, 1983). 12 IAN D. GAULD An ophionine may have three or more larval instars, though this facet of their biology requires closer investigation as it is notoriously difficult to assess the number of instars that endo- parasitoids undergo (Rojas-Rousse & Benoit, 1977). Post first instar ophionine larvae are hymenopteriform in appearance (Clausen, 1940). The ophionine larva usually does not kill its host until after the host larva is fully grown, and often after it has spun a cocoon or constructed some other form of pupation retreat. The fully grown parasitoid larva eats out all the body tissue and fluids of the host, leaving only the cephalic capsule and cuticle (Vickery, 1929 and pers. obs.). Generally the host is destroyed by the parasitoid final instar larva prior to actual pupation, and the parasitoid larva then spins a thick cocoon within the host cocoon or in the host’s pupation chamber in the soil (Rohl fs & Mack, 1985a). One Palaearctic species of Ophion is unusual in that it spins a cocoon within the host pupa (Brock, 1982), whilst an American species of Enicospilus kills its host as a fully grown larva and spins a cocoon within the unbroken host larval skin (Hancock, 1926). Like many other Hymenoptera (Gauld & Bolton, 1988) ophionines apparently only have a brief pupal instar. A great deal of the period the ophionine spends within its cocoon may be passed as a final instar larva, and species which enter prolonged ‘pupal’ diapause probably do so as larvae. In north-western Europe one species of Ophion which emerges very early in spring passes the winter as a (?pharate) adult in the cocoon (Morley, 1915): Host specificity A great deal of work still needs to be undertaken on the host relationships of ophionines. It is very difficult to establish the real host range of any one species from the literature as misiden- tifications of both parasitoids and hosts abound. Host lists of the type compiled by Thompson (1957) can be misleading as they simply collect all records, but fail to re-evaluate the evidence upon which each is based (see also Askew & Shaw, 1986). All too often one finds that several congeneric sympatric species of ophionine are listed as having identical or very similar host ranges. For example, in the U.S.A. the two fairly common large ophionines Enicospilus americanus and E. glabratus were frequently confused in early literature, and they are both recorded from a similar range of saturniid, arctiid and lymantriid hosts (Hooker, 1912). Closer study has suggested that the former species only attacks saturniid larvae, whilst the latter is restricted to the larvae of arctiids and lymantriids (Gauld, 1988). The evidence presently available indicates that there is a wide range of host specificity amongst species of Ophioninae. Many will attack a variety of hosts within a given family. The Central American species Thyreodon atriventris is believed to parasitize a range of sphingid larvae, but it has never been reared from similar-sized sympatric larvae belonging to other families (Janzen, pers. comm.). Other ophionines are more restricted in their host range, such as to hosts of a single subfamily; Thyreodon santarosae is only known to parasitize ceratocam- pine saturniids (Porter, 1986 and p. 61). Other species are believed to have an even more restricted host range and some apparently are monophagous. For example, Enicospilus lebophagus has only been reared from the saturniid Rothschildia lebeau despite intensive rearing of other sympatric possible hosts (Janzen, pers. comm.; Gauld 1988). To view host range solely from a taxonomic viewpoint is undoubtedly a gross oversimplifica- tion, because a variety of other ecological parameters markedly affect host searching by parasitoids (see, Askew & Shaw, 1986; Lawton, 1986; Sato & Ohsaki, 1987). Different parasitoid species are often found in different types of habitat. For example, in Costa Rica Thyreodon rivinae and T. maculipennis may be quite common in deciduous Pacific dry forest, but they do not occur in neighbouring wet forests, where they seem to be replaced by other Thyreodon species. One of these exclusively wet forest species, T. laticinctus, is known to parasitize a species of sphingid that occurs in both wet and dry forest habitats (see p. 63), and thus this parasitoid only attacks its host over part of its host’s geographical range. Innate responses to a combination of physical and semiochemical factors (Vinson, 1981; Weseloh, 1981; Elzen et al., 1983) may lead a parasitoid to search for hosts in a very limited segment of the total environment, and within this limited niche a physically similar, or taxonomically related subset of the available species may be used as hosts. For example, Enicospilus glabratus OPHIONINAE OF TROPICAL MESOAMERICA 13 parasitizes ‘hairy’ larvae (lymantriids and arctiids) that feed exposed on the leaves of angiosperm trees, whilst E. purgatus and Ophion flavidus have been reared from a variety of noctuid species that feed as larvae on low herbaceous vegetation. A number of Central American ophionines parasitize polyphagous hosts (e.g. see Janzen, 1984b) but it is not known whether larvae are parasitized equally on all possible foodplants. Studies on other endo- parasitoids of lepidopterans indicate that the parasitoids develop with different degrees of success depending upon what food plant the host consumes (Barbosa et al. 1986; Thorpe & Barbosa, 1986). Ophionine parasitoids of saturniids in North and Central America compared: a discrepancy in host ranges As far as is known, virtually all members of one species-group of Ophioninae (the Enicospilus americanus complex) parasitize the larvae of saturniine and hemileucine saturniids. In eastern North America the host resource ‘saturniid larvae’ is divided between two polyphagous species, E. americanus which attacks a range of mostly saturniines, and E. texanus which parasitizes a variety of mostly hemileucine saturniids (Gauld, 1988). This situation is a good example of a phenomenon pointed out by Lawton (1986) — a sympatric, synchronous and phylogenetically closely related group of species share a parasitoid. However, in Mesoamerica the EF. americanus complex is represented by 27 species that are known to, or considered likely to, attack saturniids (see pp. 123-172). A number of these have been reared, and in each case a single species of Enicospilus is only known to attack one species of saturniid. For example, E. bozai parasitizes Copaxa moinieri, E. robertoi parasitizes Hylesia lineata, E. ugaldei parasitizes Automeris tridens and E. lebophagus attacks Rothschildia lebeau. No saturniid is known to be attacked by more than one of these species of Enicospilus. These data suggest that in Mesoamerica the host resource ‘saturniid larvae’ has been subdivided by a host-species specific complex of Enicospilus and thus it seems that synchronous, sympatric and closely related tropical saturniids do not share parasitoids. This is a surprising observation because, as host resources are more subdivided in tropical localities than temperate ones, one might expect tropical parasitoids to be more polyphagous than their temperate counterparts (see Janzen & Pond, 1975; Rathcke & Price, 1976). Janzen (1981) postulated an additional scenario in which he suggested that tropical koinobionts may be better at finding scarce hosts than their temperate relatives. Possibly, in the case of the E. americanus complex, evolutionary pressure for an increased ability to find rare hosts might have led to a narrowing of the ‘searching image’, resulting in a parasitoid only being able to locate a single host species, though it is difficult to imagine that such narrowing of the ‘searching image’ could alone lead to E. lebophagus only recognizing one of two very similar sympatric species of Rothschildia as a potential host. One might expect that natural selection would favour indivi- duals that could attack either saturniid species. However, insect larva are far from defenceless against koinobiont parasitoids. Many have well-developed immuno-defensive systems and are capable of encapsulating foreign bodies (such as developing parasitoids) in their haemocoel (Salt, 1975). Parasitoids overcome this defence in a variety of ways (Salt, 1968), especially by injecting substances during the oviposi- tion sequence (Vinson & Iwantsch, 1980; Guzo & Stoltz, 1987). Clearly the potential exists fora co-evolutionary race between ophionine parasitoids and their saturniid hosts, with selective pressures favouring saturniid larvae with ever more efficient immuno-defensive systems and parasitoids that have more efficient immunosuppressive venoms. Such a mechanism could have led to the evolution of extreme host-specificity, thus permitting large numbers of closely related parasitoid species to exist sympatrically and synchronously. If immunosuppressive venoms are host-specific in this species-group, one would expect that evenif E. lebophagus could be induced to oviposit in Rothschildia erycina, its progeny would be incapable of developing successfully. One relevant question that potentially could be answered by systematics is, ‘Did the host- specific tropical species arise from a northern generalist ancestor (of the E. americanus type), or have the North American generalists arisen from a more host-specific tropical species?’ A thorough cladistic study of the americanus complex could reveal the answer, but preliminary 14 IAN D. GAULD work (Gauld, unpubl.) has not proved fruitful because few synapomorphies are recognizable and excessively high homoplasy plagues the data set. It is my subjective opinion that the entire species-complex may have had a northern origin, spreading to the New World from the Old. I suggest this because species very similar to E. americanus exist in widely separated parts of the Old World (e.g. E. inflexus and E. undulatus in the Palaearctic, E. plicatus and E. grandis in the Oriental region and E. leucocotis in southern Africa), and furthermore all the structurally more primitive members of the lineage exist in the Old World (e.g. the E. cohacarus species-complex in Madagascar). E. americanus is one of the structurally least specialized New World species, and it may well be similar to the ancestor of the diverse Central American species. The Ophioninae of Mesoamerica The composition and size of the Mesoamerican fauna All of the genus-groups of Ophioninae recognized by Gauld (1985) are present in Central America, as are four of the five subgroups of the large Enicospilus genus-group (Table 1). The subfamily is represented in Mesoamerica by approximately 170 described species belonging to 12 genera, Enicospilus, Ophion, Thyreodon, Ophiogastrella, Stauropoctonus, Rhynchophion, Sicophion, Prethophion, Janzophion, Simophion, Eremotylus and Agathophiona. This is a greater generic diversity than that found in any other zoogeographical region except South America. Only two New World genera of Ophioninae are not represented in the study region, Table 1 The higher classification of New World Ophioninae OPHION genus-group Ophion Fabricius, 1798. [Widespread; also in Old World] Alophophion Cushman, 1947. [Temperate South America] Agathiophiona Westwood, 1882. [Sonoran Mexico] [Four other genera in Old World and Australia] SICOPHION genus-group Sicophion Gauld, 1979. [Tropical Central and South America] Janzophion Gauld, 1985. [Tropical Central and South America] [One other genus in Australia] EREMOTYLUS genus-group Eremotylus Foerster, 1869. [Widespread; also in Palaearctic] Trophophion Cushman, 1947. [SW. of U.S.A.] [One other genus in southern Palaearctic] THYREODON genus-group Rhynchophion Enderlein, 1912. [Tropical America, southern U.S.A.] Thyreodon Brullé, 1846. [Widespread in New World] [Three other genera in Old World] ENICOSPILUS genus-group Orientospilus subgroup Prethophion Townes, 1971. [Tropical Central and South America] Simophion Cushman, 1947. [Arid parts of Nearctic and Palaearctic regions; one possibly congeneric species from humid part of Panama] [One other genus in Old World] Ophiogastrella subgroup Ophiogastrella Brues, 1912. [Tropical Central and South America] Stauropoctonus subgroup Stauropoctonus Brauns, 1889. [Tropical Central and South America; also in Old World and Australia] [One other genus in Afrotropical region] Leptophion subgroup [Three genera in Old World tropics and Australia] Enicospilus subgroup Enicospilus Stephens, 1835. [Cosmopolitan] [One other genus in Old World; three genera in Hawaii] OPHIONINAE OF TROPICAL MESOAMERICA 15 Alophophion, which is restricted to temperate southern South America, and Trophophion, which inhabits the arid areas of California and Arizona (Cushman, 1947; Gauld & Lanfranco, in press). More than 60% of Mesoamerican species of Ophioninae, that is 116 species, belong to the genus Enicospilus. Ophion and Thyreodon are represented by about 20 species each, whilst the total of Mesoamerican species in all the other genera combined numbers only about 15. Distribution patterns within Mesoamerica It is an indisputable fact that all species of New World Ophioninae do not have the same geographical ranges, but what is perhaps more surprising is that neither do they all have totally different ranges. Rather one finds that a number of often only distantly related species have a very similar range and, for the group as a whole, there seems to be relatively few of these distribution patterns. The approximate limits of the nine preponderant ones are shown on Maps 2 and 3, and they are discussed further below. 1 PAN-AMERICAN. Only four species of Ophioninae can be said to have a more or less Pan- American distribution, that is they are widely distributed from Canada or the northern United States southwards to Argentina. These four taxa, Ophion flavidus, Enicospilus purgatus, E. dispilus (= arcuatus) and E. glabratus, are generally common insects in disturbed habitats, but they are rather less common in pristine tropical forests. All seem to be fairly polyphagous, attacking a range of lepidopterous hosts, though they may be restricted to caterpillars in one vegetational stratum. O. flavidus and E. purgatus are parasitoids of grass- and herb-feeding noctuid larvae, E. dispilus is known to attack noctuids and notodontids that feed on shrubs or low trees, whilst E. glabratus seems to attack hairy larvae feeding on trees. The widspread temperate species E. americanus apparently does not have a Pan-American distribution as I have not found it to be present in tropical America; it seems to have a disjunct distribution, being present in temperate North America and southern South America. 2 TRopIcAL AMERICAN. Several species are widespread throughout tropical South America, much of Central America, the Caribbean and into Florida. One of the commonest examples is Enicospilus trilineatus (= appendiculatus), though this is exceptional in extending quite widely into the southern United States. Other examples that have a more restricted range (generally only north to southern Florida) include E. cubensis, E. fernaldi, E. flavus and E. guatemalensis. A few other species, such as Thyreodon atriventris, Enicospilus flavoscutellatus and E. liesneri, have a very similar distribution, but are not yet known to occur in Florida. Conversely, a number of other taxa (including E. aktites, E. ulfstrandi, E. orosii and E. opleri) occur in Florida, Central America and South America, but they are not yet known from any Caribbean island. This apparent absence may simply reflect a paucity of collecting effort. All of the species that are widespread throughout the smaller Caribbean islands either have type 1 or 2 distribution patterns; all widespread Caribbean species are also present in both Central and South America, though one, E. flavus, is only recorded from one Central American locality. Most of the species showing a type 2 distribution pattern (e.g. T. atriventris, E. cubensis, E. flavoscutellatus) are rather catholic in their choice of habitat, and are found in disturbed dry forests as well as in relatively undisturbed wetter forest over a considerable altitudinal range. Others, like E. aktites, may have a preference for coastal scrub and woodland. 3 MExIcAN/NEotTroPICAL. A very large number of species of Enicospilus are widespread throughout tropical South and Central America, and extend as far north as the southern states of Mexico. They do not, however, occur on the Carribean islands or in Florida. Examples of taxa with this type of distribution include Enicospilus monticola, E. mexicanus, E. columbianus, E. exoticus, Thyreodon morosus and T. laticinctus. Virtually all of these species are more or less restricted to humid forest biomes. 4 SuB-NICARAGUAN/NEoTROPICAL. A large number of ophionines are widespread throughout tropical South America, but their range only extends north into Panama or Costa Rica, reaching to about 11°N. Taxa with such a distribution include Prethophion latus, Ophiogastrella mac- ulithorax, Enicospilus chaconi, E. tenuigena, E. laurenae and E. xanthocarpus. Several of these IAN D. GAULD 16 ‘dnois uesawy 9e1oduia} jo WI] UsYyINOs (6) ‘dnossd s1wWapus uvagqIIeD (g) :dnoss [eoido.jOaN/UENSeARSIN-Qns Jo JWT] U1dyIIOU (p) ‘dnosd [eotdoOaN/uRdIXa|Q] JO Wuny urayjiou (¢) ‘dnoss uvsuowy peoido.y Jo yun] UsdYII0U (Z) ‘sus9}{ed UONNQLsSIp Jo sy] oyeUIXxO1dde SuIMmoys voLIoWIeOSa|, Z dey Co ‘\x > tS «= TX ss Segre E ° °, \ ; . ‘ NS Le SS a \, ~s Ns Ne se “ 6 17 OPHIONINAE OF TROPICAL MESOAMERICA ‘dnoid s1wapue ursuewy jenuad (/) ‘dno13 s1wapus uesixayy (9) ‘dnoid otwapus eruemeuRg/uRory eISOD (¢) ‘susoyed UONNGLYSIpP 9wWOs Jo sj] ayewTXOIdde SuIMoYs BoLIOWIROSA ¢ dey] 7 7 A x AK « A a * > + t + + + + + 4 18 IAN D. GAULD species are associated with humid forest habitats at moderate altitudes (500-1000 m) on mountains, and the discontinuity of such habitats in the vicinity of Lake Nicaragua may have prevented them from spreading northwards. 5 Costa RIcAN/PANAMANIAN. The existence of this area as a centre of endemicity may be artefactual as it is the most intensively sampled region. However, 13 species (Ophion clio, O. uraniae, O. erato, O. flavoorbitalis, Janzophion nebosus, Sicophion fenestralis, Enicospilus georginae, E. martae, E. corcovadoi, E. erasi, E. bozai, E. mayi and Stauropoctonus bicarinatus) are only known to occur in both Costa Rica and Panama. The first seven of these are taxa that only occur at moderate to very high elevation sites. In other regions a considerable degree of endemicity has been found amongst ophionines inhabiting montane areas (Gauld & Mitchell, 1978; 1981) so perhaps a similar group of endemic species is centred on that block of mountains dominated by the Cordillera Talamanca. Seven taxa (O. euterpe, O. arribai, Simophion melanostigma, Enicospilus karrensis, E. pamelae, E. fosteriand E. hubbelli) are apparently restricted to Panama, whilst 27 taxa (Table 2) are endemic to Costa Rica. 6 MEXICAN. Twelve taxa (Thyreodon ferrugineus, T. niger, T. robur, Agathophiona fulvicornis, Janzophion saxis, Eremotylus tropicus , Enicospilus halffteri, E. sarukhani, E. gomezpompai, E. dirzoi, E. catemacoi and possibly E. masoni) are more or less endemic to Mexico. As I have not collected in Mexico, nor have I seen extensive collections from regions other than Chiapas, I can offer no further observations about any correlations between the distribution of ophionines and different major habitat types. 7 CENTRAL AMERICAN. Several common species are widely distributed throughout Central America from Mexico or the extreme south of the United States, south to Costa Rica or Panama. Some of these taxa (e.g. Thyreodon rivinae, T. maculipennis, T. apricus, Enicospilus lebophagus and E. ugaldei) seem to be associated with areas which have a pronounced dry season. Such species may range as far north as the southern periphery of the United States, but they do not seem to occur further south than Guanacaste Province, Costa Rica. Other species (such as E. donahuei, E. forsythei and E. parkeri) are associated with wetter areas and only extend north to about the Tropic of Cancer, but they extend south into central Costa Rica or Panama. 8 CARIBBEAN. Eleven species belonging to the genera Enicospilus and Thyreodon are endemic to islands in the Caribbean (Table 3). All, except the endemic Puerto Rican species E. luquillo, and T. ultor, which occurs both on Cuba and the Bahamas, are endemic to the large, geo- logically old islands of Cuba, Hispaniola and Jamaica. Seven of these taxa are restricted to just one of these islands, and only one, E. howdenorum, occurs on all three. Some other species of Enicospilus have a similar, but slightly wider, distribution. For example, E. sondrae occurs in Florida as well as on the three major Caribbean islands, and E. cressoni also occurs in Florida and Mexico. 9 TEMPERATE AMERICAN. Although no serious attempt was made to study most of the ophionines that have this distribution, in passing I noted that Enicospilus texanus and several species belonging to the genera Ophion and Eremotylus have ranges that are predominantly in the United States, but extend southwards into northern Mexico (see Townes & Townes, 1966). Many of these taxa, such as Ophion elongatus and Eremotylus subfuliginosus, seem to be associated with semi-desert areas. Comparison between the Mesoamerican fauna and that of South America Many of the ophionine genera represented in Mesoamerica are more species-rich in South America. Some, such as Ophiogastrella, Stauropoctonus and Sicophion, have their northern limits in tropical Central America, whilst others, such as Thyreodon and Enicospilus seem to decrease in species-richness as one moves north. Almost half of the species of Enicospilus present in Central America have either distribution patterns 2, 3 or 4, that is to say, they also occur widely in South America. Although the tropical South American fauna is incompletely known, an examination of the material available suggests 19 OPHIONINAE OF TROPICAL MESOAMERICA [Rory e1soD ‘seuinbsy woly uMOUyY A[UO SI WnsounpojDg ‘J “g'N] “UW OOP ‘BITIO9D PIS 18 JeIUQeY ALTIUS UT SIND90 OSTYz "W QOOT “eq[euiny 1e yeIQeY IepIUIIs UI sim990 OSTY, SE EE ee a a a a hee a ed ke ee + —_ _ _ _ —_ — + —_ — _ _ _ — + _ — _ _ — — _ 4 _ —_ —_— _ _ ~ - - ~ - - ~ _ _ _ + _ — = — — _ —_ I = _ — - - ~ - - ~ - _ —_ _ pe - —_ — _ — + _ —_ —_ _— —_ _ + _ — _ = - ~ - - ~ - - —_ — + i = — = = — — — + — — ~ - ~ - - - ~ Ww OdceEre WOSEc Ww OOST-er WW OOIT W 006-8 w006-8 WO08-9 ayIonyy P| Sseod SAIOSIY OBdP-) JOLISIp SAIOSOY ores) ep O1I9+) uBd[OA, opIsAg]UOPy uBvI[OA, Jetuqeyy ug uowley us ornelg Sado. INVLNOW isauod Gno1g ISO. ANVINO| YAMOT LAA x - + - + + + = + E + = + ~ * E x zt = - + W OO0r-€ WOEE eyoeHH =e “J&N [2 OLIaD, «= s RSOY BIS Isaxoy AUG ] sorseds IVSOADJUDS uopoaddky J ad as a4oyoisdiaq apluudydAjod auawModjau adouyvo 100909 uo1ydQ ASojijs 12410] 1zajvzuos vjjaajsvso1ydQ MUDULIIA 14ajiod 14aqnpo 108uau ya14qv3 aD[JaISAAIIWAY ea 1zauvs wuniouapsof wunApvAISa 1u0dvY9 apy19a9 av1sauadq 1040A]D 10pavjaqv snjidsonuq “BOTY BISOD 0} S1WIApus seuIUOTIYdG jo saidads Jo uoNNqUIsSIq Z IIqeL 20 IAN D. GAULD Table 3 Endemic species of Ophioninae in the Caribbean CUBA HISPANIOLA JAMAICA PuerRTO Rico BAHAMAS Enicospilus carlota + dajaboni luquillo = howdenorum masneri + ++ 141 + l | Thyreodon affinis elegans flammiger fulvescens grandis ultor +++ 144 | I++ ] l that more than 50% of this fauna is represented in Mesoamerica. Mesoamerica is thus faunistically very similar to South America. Much of Mesoamerica is geologically recently subaerial (Coney, 1982), and to a considerable extent the flora and fauna of the region is thought to comprise recent colonists from the great continental masses of South and North America (Gentry, 1982; Rich & Rich, 1983). One is tempted to suggest that species exhibiting distribution patterns 2, 3 or 4 are recent colonists that have spread into Mesoamerica from the south, the most vagile having spread through the Caribbean to Florida, and less vagile species being restricted to the Central American mainland. As one progresses north through Mesoamerica the Neotropical faunal element decreases, though it does not seem to do so smoothly. Rather, two ‘steps’ are apparent. The northernmost of these steps occurs about 20°N in southern Mexico; this is as far north as most Neotropical elements (such as E. flavoscutellatus, E. monticola and E. mexicanus) reach. Many other Neotropical species (e.g. E. chacont, E. tenuigena and E. xanthocarpus, and species of other genera such as Prethophion latus and Sicophion fenestralis) have ranges that only extend to 10-11°N in Costa Rica so the second step more or less follows the Costa Rican/Nicaraguan border. Virtually every species that is known from Nicaragua, El Salvador, Honduras, Guatemala or Belize has a range that extends into southern Mexico. A similar stepped situation has been observed for the South American herpetofauna in Mesoamerica. Savage (1982) observed that 20 taxa had the northern limit of their range in Costa Rica, 11 had the northern limit of their range in Mexico, but only six had the northern limit of their range in the intervening area. Comparison between the Mesoamerican fauna and that of North America Comparatively few species are widespread in both North and Central America, and those that are (distribution pattern 1 species) are also widespread in South America. Amongst the Ophioninae there does not seem to be any north-centred homologue of patterns 2-4. The only aggregation of species with a north- centred distribution pattern (9) comprises a very small proportion of the Meosamerican fauna, and such elements do not penetrate into tropical areas. Throughout much of North America Ophion is the dominant ophionine genus, far surpassing (both in numbers of species and individuals) all other ophionine genera combined. There are probably about 50 species of Ophion in North America (Gauld, 1985), and the genus becomes progressively less species-rich southwards throughout Central America. In tropical Meso- america most species of Ophion are restricted to cool localities at high altitudes. Enicospilus is represented in North America by 22 species (Table 4). The majority of these taxa (13) only occur on the extreme southern fringes of the United States, in areas such as Florida south of 29°N, or Hidalgo county, Texas. Most of these species are also widespread in South America (i.e they have a type 2 distribution pattern). I know of only three species of Enicospilus with a north-centred distribution — EF. americanus, whose range extends southwards OPHIONINAE OF TROPICAL MESOAMERICA 21 Table 4 Distribution of Enicospilus species in North America UNITED STATES MESOAMERICA Other Central Florida Texas California states America Caribbean aktites + = = americanus cressoni cubensis cushmani dispilus doylei fernaldi flavus glabrutus guatemalensis lebophagus neotropicus opleri orosit peigleri purgatus sarukhani sondrae texanus trilineatus ulfstrandi +r + +4 l | bt++++4+14+1 bl +t+tti¢i+¢i t+t+ti t¢tt+te+e il +¢ettett+rt+tt + + Peete tee Lett i +++ +++ 11 evkooe de foetal l++ | ++ | l+1+1+4i ‘single record from southern Louisiana. *recorded once from Panama Canal. 3except for extreme north of Mexico. through the Mexican uplands almost to the Guatemalan border (and has a disjunct representa- tion in southern South America), EF. texanus, whose range extends into northern Mexico, and E. cushmani, which is the only known endemic North American Enicospilus. Thus, to a large extent, the Enicospilus fauna of the United States can be regarded as a depauperate northern extension of the much richer Neotropical fauna. The evolutionary history of Mesoamerican ophionines The scenario suggested by the faunal elements — massive invasion of Mesoamerica from South America with some localized speciation — is probably a great oversimplification as certain species-groups which are most diverse in the Neotropics may initially have spread from the north and radiated in the south. For example, the Enicospilus americanus species-complex, the E. purgatus complex and the genus Rhynchophion all have their closest relatives in the Old World temperate regions, so possibly they colonized South America from the north and subsequently diversified in the tropics. Even Stauropoctonus, which is not now present in North America, may have spread to South America from North America at some remote time. This is suggested by the fact that the Neotropical species have obviously been derived from Old World taxa (and the group is unlikely to be old enough to predate the breakup of Gondwanaland) (see Gauld, 1985), and the most primitive Neotropical species of Stauropoctonus occurs at the northern extreme of the range of the genus in the New World (Fig. 82). Possibly all species of Enicospilus in South America may have originated from a few ancestral colonists that spread through the north from the Old World. Although the phylogeny of Enicospilus is very incompletely understood, this scenario is suggested by the following facts: a) the closest relatives of Enicospilus, that is Dicamptus and the Leptophion complex, are confined to the Old World; b) Enicospilus is represented in the Old World by at least 30 very distinctive species-groups; 22 IAN D. GAULD c) Enicospilus is represented in the New World by only five species-groups, two of which are more diverse in the Old World. The great majority of tropical American Enicospilus species seem to comprise a single species- group, the E. dispilus group (see p. 75). One group of species, the genus Janzophion, deserves special mention as it is endemic to mountain tops of Central America. Gauld (1985) hypothesized that this group and Sicophion represent ancient autochthonous South American relict groups that may have had a Gondwanic origin, as their closest relatives occur in Australia. Latitudinal gradients in species-richness Janzen (1981), analysing the distribution of species of seven ichneumonid subfamilies in North America (Acaenitinae, Banchinae, Diplazontinae, Metopiinae, Phygadeuontinae, Pimplinae and Xoridinae), found a general decrease in species-richness southwards from about 40°N. The Ophioninae, however, seems to be exceptional as it apparently shows an increasing species- richness towards the equator. Four American sites that have been reasonably well-collected are Washtenaw County, Michigan, U.S.A. (42°N), Alachua County, Florida, U.S.A. (29°N), the area around Huixtla, Chiapas, Mexico (17°N) and Guanacaste National Park, Costa Rica (11°N). Eleven species of Ophioninae have been found to occur at the Michigan site, 18 at the Floridan site, 49 in Chiapas and 69 in Guanacaste National Park. A similar latitudinal gradient was observed for the Ophioninae in Australia (Gauld, 1986), with tropical Queensland having more than twice as many species as temperate Tasmania. Endemism in Costa Rica In this study 28 species of Ophioninae have only been found to occur in Costa Rica (Table 2). Four of these (Ophion polyhymniae, O. terpsichore, O. thaliae and O. melpomene) are restricted to high altitude oak forest-dominated sites on the Cerro de la Muerte and Volcan Pods. Very similar vegetation is present at the same altitude in northern Panama, and many species (e.g. Ophion uraniae, O. clio and Enicospilus georginae) occur in this habitat in both countries. It would be quite surprising if the four apparently endemic Costa Rican Ophion species did not also occur in Panama. Similarly, Ophion arribai, only known from a high altitude oak forest in Chiriqui Province, Panama, may be expected to occur in similar habitats in Costa Rica. Political boundaries should not obscure the fact that it is the highland massif centred on the Cordillera de Talamanca which seems to be an area of endemism in Mesoamerica. Eleven species (Ophiogastrella gonzalezi, O. stilesi, O. lemairei, Thyreodon santarosae, Enicospilus gamezi, E. vilmari, E. estradarum, E. alvaroi, E. oduberi, E. hemicrescellae and E. ceciliae) have only been found to occur either in, or in sites adjacent to, the Pacific dry forest of north-western Guanacaste. This is surprising as a similar endemicity has not been observed in their lepidopterous hosts. For example, Janzen (in press) has commented that the dry forest in Santa Rosa National Park, Guanacaste, has very few endemic species of Lepidoptera. Another surprising feature of these endemics is that, with the exception of one specimen of E. alvaroi which was collected in more humid forest in Rincon de la Vieja National Park (adjacent to drier areas), none has ever been collected in wet forest sites. This is remarkable because many other (non-endemic) dry forest species frequently occur in wet forest sites. Eight species (Ophion cacaoi, O. calliope, Enicospilus gabrieli, E. chaconi, E. abelardoi, E. mengoi, E. porteri and Thyreodon species 1) occur in wet forests at altitudes between about 600 and 1100 metres. All but three of these species have been taken at more than one site. Three further species (Enicospilus brenesiae, E. haberi and E. fogdenorum) are only known to occur in cloud forests at altitudes between 1300 and 1500 metres. All of these species have only been collected in Monteverde reserve, and no Costa Rican endemics are known to occur both at Monteverde and any other site. The habitat of the remaining Costa Rican endemic species, Enicospilus baltodanorum, is unknown, but it may have been collected in lowland wet forest. A comparison of tropical ophionine faunas in Mesoamerica and South East Asia Although the species composition of the ophionine faunas of tropical Mesoamerica and South East Asia are completely different, there are interesting analogies and differences in the OPHIONINAE OF TROPICAL MESOAMERICA 23 ophionine faunas of two areas. These are of some interest because the Ophioninae seems to be the dominant (= most species-rich) group of koinobiont ichneumonids in both areas One of the most striking differences between the faunas of the two areas is the existence of a quite large group of diurnally active species (Thyreodon and Rhynchophion) in Mesoamerica, whilst only two very uncommon diurnal ophionine species (Dictyonotus spp.) are known to occur in South East Asia (Gauld & Mitchell, 1981). All three genera seem to be closely related, and are known to parasitize the larvae of sphingids, but ophionine sphingid parasitoids are more diverse and common in the New World than the Old. Relatively few species of Ophion occur in either region, and the great majority of species that do are restricted to altitudes above 2000 metres. Enicospilus is the most species-rich genus of ophionines in both tropical localities, yet the altitudinal distribution of the species differs strikingly between the regions. In the Central American states of Costa Rica and Panama the majority of Enicospilus species are encountered from altitudes ranging from sea-level to 1600 metres. Rather few species have been collected above 2000 metres and none at all have been taken above 3000 metres. In Borneo and New Guinea (which are typical examples of the South East Asian tropics) relatively few species of Enicospilus are encountered at low elevation sites and the greatest species-richness occurs between 1500 and 2500 metres. In New Guinea several species even occur above 3000 metres, though at this altitude species-richness is very low. Table 5 presents a comparison of the Enicospilus species occurring in four well-collected Mesoamerican sites (Barro Colorado Island and Guadalupe Arriba in Panama, Santa Rosa National Park and Monteverde Reserve in Costa Rica) and four sites studied by the author at similar altitudes in Borneo and New Guinea. The lowland sites (<400 m) in Borneo are very much less species-rich than the cloud forest site (Gunong Pagon/Bukit Retak in Brunei), whilst in Mesoamerica the lowland sites are similar to or have a greater species-richness than the cloud forest site. Whilst lowland sites in South East Asia are less species-rich than upper montane forest sites (2300 m), the lowland sites in Mesoamerica are very much more species-rich than upper montane Mesoamerican sites. In the Old World tropics, many species of Enicospilus have a much more restricted atitudinal range than their Mesoamerican congeners. For example, in Brunei 21 of the 34 species occurring between 1500 and 1700 metres did not occur at neighbouring low altitude sites (Gauld, 1986b), whilst not one of the 19 species collected at 2300 metres on Mt Kaindi, Papua New Guinea, has been collected at 1000 metres on the same mountain (Gauld & Mitchell, 1981). In Costa Rica 20 of the 38 species collected at Monteverde (1300-1400 m) also occur in Santa Rosa National Park (300 m) whilst all the species occurring at Guadalupe Arriba, Panama (2200 m) have also been collected at Monteverde (Costa Rica). Table 5 Comparison of species-richness of Enicospilus at different altitudes in Central America and South East Asia CENTRAL AMERICAN SITE No. spp. SouTH East ASIAN SITE* No. spp. Barro Colorado Is., 38 Coastal Brunei, 0-100 m 8 Panama, 150 m Santa Rosa Nat. Pk, 50 Brunei dipterocarp forest, 19 Costa Rica, 300 m 100-300 m Monteverde Res., Costa 38 Pagon/Retak, Brunei, 1500-1700 m 34 Rica, 1300-1500 m Guadalupe Arriba, 3 Mt Kaindi summit, 19 Panama, 2200 m Papua New Guinea, 2300 m *sources Gauld & Mitchell, 1981; Gauld, 1986. 24 OPHION Fabricius arribai sp. n. cacaoi sp. n. calliope sp. n. clio sp. n. erato sp. n. euterpe sp. n. flavidus Brullé ancyloneura Cameron biangularis Taschenberg concolor Szépligeti diversus Szépligeti politior Morley flavoorbitalis Cameron melpomene sp. n. polyhymniae sp. n. terpsichore sp. n. thaliae sp. n. uraniae sp. n. AGATHOPHIONA Westwood fulvicornis Westwood SICOPHION Gauld fenestralis sp. n. JANZOPHION Gauld nebosus Gauld Saxis sp. n. EREMOTYLUS Foerster tropicus sp. n. RHYNCHOPHION Enderlein flammipennis (Ashmead) THYREODON Brullé apricus Porter atriventris (Cresson) grenadensis Ashmead ornatus (Szépligeti) rufothorax Cameron thoracicus (Ashmead) erythrocera Cameron laticinctus Cresson maculipennis Cresson morosus Smith rivinae Porter santarosae Porter species 1 PRETHOPHION Townes latus Townes SIMOPHION Cushman melanostigma (Cameron) comb. n. OPHIOGASTRELLA Brues gonzalezi sp. n. lemairei sp. n. maculithorax Brues stilesi sp. n. STAUROPOCTONUS Brauns bicarinatus (Cushman) ENICOSPILUS Stephens undulatus species-group IAN D. GAULD Nomenclatural summary abelardoi sp. n. aktites Gauld alvaroi sp n. americanus (Christ) cecropiae (Sanborn) druryi (Kriechbaumer) rugosus (Brullé) bozai sp. n. brevis (Morley) cameronii (Dalla Torre) curvinervis (Cameron) renovatus (Morley) chaconi sp. n. chiriquensis (Cameron) calcator (Morley) syn. n. clarkorum sp. n. cushmani Gauld enigmus sp. n. gamezi sp. n. glabratus (Say) angulatus (Hooker) syn. n. arctiae (Ashmead) cubitalis (Morley) excubitalis Walkley gomezpompai sp. n. halffteri sp. n. hallwachsae sp. n. lebophagus Gauld major (Morley) mayi sp. n. mexicanus (Cresson) peigleri Gauld quintanai sp. n. robertoi sp. n. sarukhani sp. n. scuintlei sp. n. tenuigena (Kriechbaumer) texanus (Ashmead) ugaldei sp. n. umanai sp. n. venezuelanus (Szépligeti) ramidulus species-group doylei sp. n. neotropicus Hooker purgatus (Say) lateralis (Brullé) flaviceps (Brullé) volubilis (Holmgren) trilineatus species-group carlota sp. n. cubensis (Norton) dajaboni sp. n. dirzoi sp. n. lupemejia sp. n. porteri sp. n. trilineatus (Brullé) striatus (Brullé) OPHIONINAE OF TROPICAL MESOAMERICA sphacelatus (Erichson) nigricauda (Taschenberg) vecors (Tosquinet) appendiculatum (Felt) syn. n. brasiliensis (Szépligeti) maculiceps Cameron brevinervis (Morley) syn. n. columbianus species-group columbianus (Enderlein) martae sp. n. dispilus species-group baltodanorum sp. n. georginae sp. n. guatemalensis (Cameron) guindoni sp. n. haberi sp. n. hacha sp. n. hemicrescellae sp. n. howdenorum sp. n. hubbelli sp. n. Jesicae sp. n. karrensis sp. n. kelloggae sp. n. kleini sp. n. barbarae sp. n. lacsa sp. n. bima sp. n. laurenae sp. n. brenesiae sp. n. leoni sp. n. burgosi sp. n. carri sp. n. catemacoi sp. n. ceciliae sp. n. cepillo sp. n. colini sp. n. corcovadoi sp. n. cornifuscus nom. n. fuscicornis (Cameron) cressoni Hooker devriesi sp. n. dispilus (Szépligeti) arcuatus (Felt) syn. n. liesneri sp. n. lovejoyi sp n. luisi sp. n. luquillo sp. n. maculipennis (Cameron) parvifasciatus Cameron syn. n. madrigalae sp. n. marini sp. n. maritzai sp. n. masneri sp. n. masoni sp. n. mengoi sp. n. monticola (Cameron) 25 donahuei sp. n. antomelas (Enderlein) syn. n. duckworthi sp. n. elegans (Szépligeti) syn. n. echeverri sp. n. fuscipennis (Szépligeti) syn. n. erasi sp. n. oduberi sp. n. estradarum sp. n. opleri sp. n. exoticus (Morley) orosii sp. n. bicolor (Szépligeti) syn. n. pamelae sp. n. dichromus Townes & Townes syn. n. parkeri sp. n. fernaldi Hooker persimilis (Szépligeti) flavoscutellatus (Brullé) pescadori sp. n. thoracicus (Cresson) randalli sp. n. trimaculatus (Taschenberg) sanchezi sp. n. trispilus (Szépligeti) syn. n. simoni sp. n. attritus (Enderlein) syn. n. sondrae sp. n. flavus (Fabricius) stevensi sp. n. flavarius (Thunberg) teodorae sp. n. concolor (Cresson) ulfstrandi sp. n. guyanensis Cameron vegai sp. n. fogdenorum sp. n. vilmari sp. n. forsythei sp. n. woldai sp. n. fosteri sp. n. xanthocarpus (Szépligeti) gabrieli sp. n. flavosignatus (Enderlein) syn. n. galilea sp. n. xanthostigma (Szépligeti) gallegosi sp. n. 26 IAN D. GAULD 12 = 14 Figs 6-14 6, central part of fore wing, Netelia sp. 7-9, fore wings; (7) Janzophion nebosus; (8) Pre- thophion latus; (9) Sicophion fenestralis. 10, 11, scutellum and propodeum, dorsal; (10) Enicospilus sp.; (11) Ophiogastrella sp. (arrows indicate anterior area of propodeum). 12, Stauropoctonus sp., hind trochantellar segments. 13, 14, hind wings; (13) Sicophion fenestralis; (14) Enicospilus sp. (arrows indicate junction of Cul and cu-a). 1 N - Nn oo Ko) OPHIONINAE OF TROPICAL MESOAMERICA 27 Key to genera occurring in Mesoamerica Premecibestinaderalywentirely absent i0 2. ci. iw ssie eke Mecsas tw Teich we eeees eee e eee. 2, Occipital carina present, usually complete on upper part of head, but sometimes narrowly aM NMENE UMLEARRRRICTER CHE SAIN | Fi as fe i estos Zvial AA Sion iglace ge hyetols « Vin hieMwe nen lene © 4 Mid and hind trochantelli with distal margin produced into an acute, curved tooth (Fig. 12); Mirmicleles twisted abOUt OO”... «meetin crhe ws Vln eles tye ope aietcie STAUROPOCTONUS (p. 72) Mid and hind trochantelli simple; mandibles not twisted, or twisted less than 20°........... 3 Fore wing with vein Rs+2r slender, almost straight; 2nd discal cell slender, longer than Ist subdiscal cell (Fig 8); epicnemial carina absent laterally. ............ PRETHOPHION (p. 64) Fore wing with vein Rs+2r basally broadened, curved; 2nd discal cell stout, shorter than Ist subdiscal cell (Fig. 7); epicnemial carina present laterally.............. JANZOPHION (p. 49) Labium with glossae greatly lengthened so as to form an elongate tube which reaches back to level of hind coxae (Fig. 18); female with subgenital plate greatly enlarged, longer than tergite 3 Bne-o lacks Mexican INSCGtSi. - oeyerpregseee = <= on oe es «Mya ae w= AGATHOPHIONA (p. 48) Mouthparts not or only slightly lengthened, the glossae at most reaching to about the epicnemial carina (Figs 16, 17); subgenital plate of female usually not enlarged and shorter than tergite 3. 5 Hind wing with abscissa of Cul between M and cu-a at most 0.6 times as long as cu-a, generally Stlogter, So:junction Of Gril andicy-a is close to M(iig. 13) Joe saci sees ive ce sseeees-- 6 Hind wing with abscissa of Cul between M and cu-a at least 0.8 times as long as cu-a, generally longer so junction of Cul and cu-a is either intermediate between M and IA (see Figs 45, 46), Cra CROSe COMLA (EI. a etme enn. AEE «= < + cs s+ « OnE oir... 5. 8 Fore wing with Rs centrally dipped (Fig. 9); pterostigma very broad; discosubmarginal cell AME KUOLIVAOLOAGIY OLAMLOUS: sorte pe Rat 6c gcse visi w Wc eee st aise che cree oO SICOPHION (p. 51) Fore wing with Rs centrally almost straight; pterostigma very narrow; discosubmarginal cell MEANS ta SMa MADLOUS AGA AMMEMIONY | screenees sss cceee ns cose sees we eee 7 Hind part of alitrunk, in profile, with propodeum greatly enlarged and metapleuron short and deep (Fig. 19); tergite 2 in profile long, generally longer than tergite 3, and with laterotergite pendant; mouthparts not particularly long, usually more or less concealed (Fig. 16). THYREODON (p. 54) Hind part of alitrunk, in profile, with propodeum not particularly enlarged, metapleuron longer than high (Fig 20); tergite 2 in profile much smaller than tergite 3, with laterotergite folded under; maxillae and labium elongate, projecting ventrally by a distance about equal to length CIMLOR IE TMPAGE (NG WM acon eter ache ace acute a MKS forse eS bles, 6 pe, n, tee a eve RHYNCHOPHION (p. 53) Fore wing with Rs+2r slightly sinuous, often thickened for more than half its length, and with a hairless fenestra in the discosubmarginal cell adjacent to it and extending about 0.5 of its length or more; fenestra often bearing pigmented sclerites in membrane (Fig. 21). ENICOSPILUS (most) (p. 74) Fore wing with Rs+2r straight or basally abruptly curved, slender or basally thickened; discosubmarginal cell only glabrous in anterior corner (Figs 22, 23), never with pigmented SG OMUCs MEV) OMEN DAN Guat Meee nae ewe ka Sous kas eke ed ieusias uleusiiee: agen e cr aue.eke, ours 9 Fore tibial spur, when viewed from behind, with a membranous flange along the comb (Fig. 25); feLeite 2usually withithymora close tovantenOmend..0..+-0-+-0--00++502--+4ee-+-. 0-0 10 Fore tibial spur, when viewed from behind, without a membranous flange (Fig. 24); tergite 2 with thyridia separated from anterior margin by more than their own diameter .......... 11 Fore wing with Rs+2r basally thickened and abruptly curved (Fig. 22); clypeus with margin flat, often blunt, never subapically impressed so margin is very thin; lm-cu arcuate or sinuous, with at most a weak trace of a vein stub................ 0.0 eee ee eee EREMOTYLUS (p. 52) Fore wing with Rs+2r at most slightly broadened basally, never abruptly curved (Fig. 23); clypeus with margin subapically impressed, thin; 1m-cu steeply curved or centrally elbowed usually withyallong vein stubipresemty. (ewan oe «ets ee eye el tierarcltercl. eles OPHION (p. 29) 28 IAN D. GAULD Figs 15-25 15, Ophiogastrella gonzalezi, male hind tarsal claw. 16-18, head, front view; (16) Thyreodon atriventris; (17) Rhynchophion flammipennis; (18) Agathophiona fulvicornis. 19, 20, propodeum and anterior segments of gaster, lateral view; (19) Thyreodon rivinae; (20) Rhynchophion flammipennis. 21-23, fore wing; (21) Enicospilus sp.; (22) Eremotylus tropicus; (23) Ophion flavidus. 24, 25, fore tibial spur, seen from front; (24) Enicospilus sp.; (25) Ophion sp. OPHIONINAE OF TROPICAL MESOAMERICA 29 11 Mandible moderately to strongly tapered distally, generally twisted 15° or more (Figs 157, 158); posterior transverse carina of mesosternum usually present, rarely interrupted centrally; propodeum with anterior groove shallow and broad, so anterior surface is long (Fig. 10). ENICOSPILUS (few) (p. 74) — Mandible weakly and evenly tapered distally, not twisted; posterior transverse carina of mesosternum absent except as lateral vestiges; propodeum with anterior groove short I Re SB ORE Mee ny ae Oe Ce ee ty ae 12 12 Epicnemial carina distinct laterally on mesopleuron; males with pectinal comb all round distal Geen cn (lavrened claws (Fig. 05) o... 2... e er dew cence weeds OPHIOGASTRELLA (p. 66) — Epicnemial carina only present as a trace medioventrally, laterally entirely absent; males with pectinal comb simple, not extending around distal apices of claws........ SIMOPHION (p. 65) The OPHION genus-group OPHION Fabricius Ophion Fabricius, 1798: 210, 235. Type-species: Jchneumon luteus L., by subsequent designation, Curtis, 1836: 600. Paniscus Schrank, 1802: 316. Type-species: Jchneumon luteus L., by monotypy. [Psylonychia Szépligeti, 1905: 21. Nomen nudum. | Stenophthalmus Szépligeti, 1905: 23. Type-species: Stenophthalmus algiricus Szépligeti, by subsequent designation, Viereck, 1914: 137. [Homonym of Stenophthalmus Becker, 1903.] Pachyprotoma Kohl, 1906: 223. Type-species: Ophion (Pachyprotoma) capitatus Kohl, by monotypy. Australophion Morley, 1912: 4, 30. Type-species: Ophion peregrinus Smith, by monotypy. Neophion Morley, 1912: 4, 30. Type-species: Neophion crassus Morley, by subsequent designation, Viereck, 1914: 100. Apatophion Shestakov, 1926: 262. Type-species: Apatophion mirsa Shestakovy, by original designation. Platophion Hellén, 1926: 13. Type-species: Ophion areolaris Brauns, by subsequent designation, Cush- man, 1947: 475. Potophion Cushman, 1947: 476. Type-species: Potophion caudatus Cushman, by original designation. [Psylonychia Cushman, 1947: 476. Unavailable name, proposed in synonymy. | Apomesus Townes, 1971: 54. Type-species: Apomesus longiceps Townes, by original designation. Mecetron Townes, 1971: 60. Type-species: Stenophthalmus choaspese Uchida, by original designation. Diacnosis. Mainly orange or brownish yellow species with hyaline or slightly yellowish wings; fore wing length 10-27 mm. Mandibles stout, not twisted; maxillae and labium unspecialized; clypeus apically impressed, truncate to weakly convex; occipital carina complete (in Neotropical species). Notauli usually present anteriorly; mesopleural furrow curving upwards from episternal scrobe to subalar prominence; posterior transverse carina of mesosternum only represented by vestiges laterally. Fore wing with Rs+2r more or less straight, not angled basally; discosubmarginal cell with a glabrous area in anterior corner; pterostigma moderately broad; 2nd discal cell large, with 1m-cu rather angulate centrally and frequently with a well-developed vein stub; hind wing with first abscissa of Rs weakly to strongly bowed; distal abscissa of Cul variously positioned, but usually about equidistant between M and 1A. Fore tibial spur with a membranous flange behind comb. Gaster moderately slender; second segment of gaster with laterotergite folded under; tergite 2 longer than tergite 3. REMARKS. Ophion is a large cosmopolitan genus that is most species-rich in the Holarctic realm. In the United States and in Mexico north of about 20°N Ophion species comprise a majority of the ophionine fauna, and a number of these species, such as O. elongatus Hooker, have ranges that extend southwards through the mountains into central and southern Mexico. It is not possible to work out these northern Central American species in isolation from the much richer Nearctic fauna, and such a study is beyond the scope of the present work. However, a number of Ophion species occur on the mountains in the southern part of Central America, and as this fauna seems to be distinct from the more northern fauna, a preliminary taxonomic treatment is presented below. Townes & Townes (1966) list nine Ophion species as occurring in the Neotropical region, of which three, O. flavidus Brullé (= ancyloneura Cameron), O. flavoorbitalis Cameron and O. melanostigma Cameron, are recorded from Central America. I have examined the holotype of O. melanostigma and conclude that it is not a species of Ophion as it lacks a membranous flange on the fore tibial spur and has no umbo. These features suggest it belongs to the Enicospilus genus-group, and I have assigned it tentatively to the genus 30 IAN D. GAULD Figs 26-34 Ophion species. 26, O. calliope, discosubmarginal cell. 27, 28, distal hamuli; (27) O. arribai; (28) O. melpomene. 29-34, central part of fore wing; (29) O. erato; (30) O. cacaoi; (31) O. melpomene; (32) O. arribai; (33) O. flavoorbitalis; (34) O. thaliae. OPHIONINAE OF TROPICAL MESOAMERICA 31 Simophion (see p. 65). I have found both of the remaining two species in southern Central America, together with 11 endemic new species which are described below. In southern Central America the great majority of Ophion species are restricted to medium or very high altitude sites. Eight of the 13 species have only been collected above 2100 m, and one species is restricted to sites above 2700 m. Only four rather uncommon species have been collected in lowland or lower montane (1500 m or below) forested sites, and a single species, O. flavidus, is widely distributed and associated with disturbed habitats. At high altitudes Ophion species comprise a large proportion of the ophionine fauna. For example, the only ophionines known to occur above 3,000 m in Costa Rica are two Ophion species, whilst at Guadalupe Arriba (2200 m) in Panama, four of the nine ophionine species are Ophion. Key to species of Ophion occurring in southern Mesoamerica 1 Discosubmarginal cell of fore wing narrowly glabrous adjacent to Rs&M (Fig. 26); propodeum with anterior transverse carina absent or present only as a central vestige, and with posterior transverse carina strong laterally, sometimes more or less complete .................... 2 — Discosubmarginal cell of fore wing proximally fairly evenly hirsute, without a distinct glabrous band adjacent to Rs&M (Figs 29-34); propodeum often with both anterior and posterior transverse carinae strong, or with the anterior one stronger and more complete than the iemeemer one, or with no'carinae really discermible ...0 0... occ ccc ne weenie se ceens 3 2 Head in dorsal view with genae strongly inflated behind eyes (Fig. 35); lower tooth of mandible slightly longer than the upper and slightly angled downwards (Fig. 38); tarsi only weakly depressed, the hind leg with the penultimate tarsal segment about 2.8 times as long as broad; flagellum yellowish brown; female with subgenital plate more or less unspecialized (Fig. 40) euterpe sp. n. (p. 34) — Head in dorsal view with genae rounded behind eyes (Fig. 36); lower tooth of mandible slightly shorter than the upper, and not angled downwards (Fig. 37); tarsi very strongly depressed, ventrally flat, the hind leg with the penultimate tarsal segment about 1.4 times as long as apically broad; flagellum black; female with subgenital plate very large and plough-share shaped, projecting beyond apex of gaster (Fig. 39).................... calliope sp. n. (p. 35) 3 Head rather long, its width across the eyes 1.2 times length of head from vertex to clypeal apex (Fig. 42); hind leg with trochantellus dorsally longer than broad (Fig. 43); a predominantly blackish brown or very dark reddish brown species................ polyhymniae sp. n. (p. 36) — Head short and broad with width across the eyes 1.3—1.5 times length of head from vertex to clypeal apex (Fig. 41); hind leg with trochantellus dorsally shorter than broad (Fig. 44); pale Vellonish Drown! tO DrOWMISM'OLAN GO SPECIES is kaw ek ee ee eee ce weet et seercades 4 4 Hind wing with marginal cell proximally very long, with the junction of Rs and rs-m near centre so distal abscissa of Rs appears to be very short (Fig. 45). A small pallid yellowish brown species frequently encountered in disturbed habitats below LELULOPaTET Wepre Pee RT, CN ge Maree: 278 PER ae Ae AME se, ig alee ne oie slecen flavidus Brullé (p. 37) — Hind wing with marginal cell proximally short, with the junction of Rs and rs-m well proximal to centre so distal abscissa of Rs appears to be quite long (Fig. 46)....................005- 5 5 Very large insects, fore wing length 25+ mm; mid tibial spurs subequal in length, the longer 1.1— 1.2 times length of the shorter. Propodeum with posterior transverse carina close to anterior one so that area superomedia is subquadrate; marginal cell of hind wing proximally broadly glabrous. ..... clio sp. n. (p. 39) — Small to moderately large insects, fore wing length <24 mm; mid tibial spurs unequal, the femeer 15— 126 timewtenrtn Of TNE SMOFED a6 coe c se cea ne kee e dec ceewsceseasoes 6 6 Posterior transverse carina of propodeum close to anterior one, with area superomedia more or less quadrate; fore wing with ramellus very short or absent (Fig. 30) and hind wing with marginal cell proximally almost uniformly hirsute ...................... cacaoi sp. n. (p. 40) — Posterior transverse carina of propodeum well behind anterior one, so that area superomedia, if discernible, is much longer than broad; fore wing with ramellus moderately to very long (Figs 29, 31-34), if shorter than abscissa of Cul between Im-cu and Cula then hind wing with Matpinal cell proximally glabrous in part (Fig. 53) 0... ee eet 7 32 IAN D GAULD Figs 35-49 Ophion species. 35, 36, head, dorsal; (35) O. euterpe; (36) O. calliope. 37, 38, mandible; (37) O. calliope; (38) O. euterpe. 39, 40, posterior end of gaster of female, lateral view; (39) O. calliope; (40) O. euterpe. 41, 42, head, front view; (41) O. flavidus; (42) O. polyhymniae. 43-44, hind trochantellar segments; (43) O. polyhymniae; (44) O. flavidus. 45, 46, distal part of hind wing; (45) O. flavidus; (46) O. thaliae. 47, O. uraniae, posterior part of gaster of male, lateral view. 48, 49, epicnemium and lower part of anterior alitrunk, anterolateral view; (48) O. flavoobitalis; (49) O. uraniae. OPHIONINAE OF TROPICAL MESOAMERICA 33 Figs 50-53 Ophion species. 50, 51, propodeum, lateral view; (50) O. thaliae; (51) O. flavoorbitalis. 52, 10 11 53, marginal cell of hind wing; (52) O. erato; (53) O. terpsichore. Lower corner of epicnemial carina, just lateral to a very strongly concave area, acute (Fig. 49); male with apex of aedeagus produced into a small acute protuberance (Fig. 47); mesopleuron reddish brown, with three pallid marks on upper part ................ uraniae sp.n. (p. 41) Lower corner of epicnemial carina, just lateral to concave part, generally rounded, rarely angled at about 90° (Fig. 48); male with apex of aedeagus rounded; mesopleuron variously coloured, never dark reddish brown with three dorsal pale areas................0..000- 8 Hind wing with proximal 0.3 of marginal cell densely hirsute centrally (Figs 52, 53)......... 9 Hind wing with proximal 0.3 of marginal cell broadly glabrous, at most with isolated hairs aS ie 4 Sa elas Pe a i SEU Ps Se A oe ee ne een 12 Alitrunk laterally yellowish, with mesoscutum and mesosternum marked with dark brown; fore REI PAUUICL TARMICLIUS, VEL Vs GED (ENP eG) ata eialecs aye of galelnie 4 Asie clase «a ere © ix ejaye'sse o'er srs.e aia/e eh 6.6 10 Alitrunk laterally orange or reddish brown, of similar colour to mesoscutum and mesosternum; fore wine with ramellus of moderate length (Figs 31,32). ........ cc esse ec sece cn ereeees 11 Central flagellar segments slender, 2.1—2.3 times as long as broad; marginal cell of hind wing only glabrous along anterior margin (Fig. 52); mesopleuron highly polished, with indistinct microreticulation; gaster with anterior segments pale yellowish orange, segments 5+ TIUSCALG He dM, BEC area SEAS PUAN BTRIATR leg AIR Cs rave lerayeneral elo ace6, 8 sxe Spans eratosp.n. (p. 43) Central flagellar stout, about 1.5 times as long as broad; marginal cell of hind wing glabrous all around proximal margin (Fig. 53); mesopleuron weakly polished, conspicuously granulate; gaster with anterior segments dark brown, the posterior segments brownish orange. terpsichore sp. n. (p. 44) Fore wing with ICI = 0.40-0.44; CI>0.75; Rs+2r basally only slightly broadened, joining pterostigma well away from its base (Fig. 31); hind wing with all distal hamuli subtending similar angle to vein R1 (Fig. 28); scutellum reddish brown......... melpomene sp. n. (p.'44)) Fore wing with ICI = 0.74-0.83; CI<0.60; Rs+2r basally abruptly thickened, joining pterostigma close to base (Fig. 32); hind wing with the most proximal 3 or more of the distal hamuli almost parallel to R1, subtending a far smaller angle than the most distal of the hamuli chigt27); see leigwihvitishr Ah A bP eden lel sleek eet). oe arribai sp. n. (p. 45) 34 IAN D. GAULD 12 Fore wing with first subdiscal cell barely narrowed distally (Fig. 34); lateral longitudinal carina of propodeum only present as an anterior vestige, which is curved up to reach margin of propodeal spiracle (Fig. 50); hind wing with distal abscissa of Cul joining cu-a closer to 1A than: towMites ceii. as Se cee Oe tae oak aees.s ec oe Mes eee ate ahd eos thaliae sp. n. (p. 46) — Fore wing with first subdiscal cell proximally very broad and strongly distally narrowed (Fig. 33); lateral carina of propodeum complete (Fig. 51); hind wing with distal abscissa of Cul joining’ cz-a'closer tovMithanito TACs 25.2 seeee- oe ae flavoorbitalis Cameron (p. 47) Ophion euterpe sp. n. (Figs 35, 38, 40) Description. Mandibles stout, weakly narrowed apically, with upper tooth slightly broader and shorter than the lower tooth which is slightly bent downwards apically; outer mandibular surface coarsely punctate, with microreticulation between punctures, and with basal part of mandible broadly concave; distal segment of maxillary palp slender; malar space 0.2 times as long as basal mandibular width. Head in front view with greatest width across eyes 1.25 times length of head from vertex to clypeal margin; orbits ventrally slightly divergent; clypeus in profile weakly convex, with isolated punctures, apically with margin strongly impressed. Lower face centrally convex, polished and sparsely punctate. Head in dorsal view with genae grossly inflated behind eyes; posterior ocellus close to but not touching eye; occipital carina mediodorsally convex, ventrally weak but curved to join hypostomal carina well above base mandible. Antenna quite short and stout, with 50 flagellar segments; 20th segment 1.7 times as long as broad. Mesoscutum polished and closely punctate, in profile abruptly rounded; notauli vestigial, only dis- tinguishable as slight impressions near anterior end. Mesopleuron polished, centrally finely and quite closely punctate; lower corner of epicnemial carina sharply produced after concavity, and with upper end inclined towards anterior margin of pleuron, but evanescent. Scutellum in profile weakly convex, not laterally carinate. Metapleuron moderately convex, indistinctly punctate. Propodeum in profile abruptly rounded; anterior transverse carina present only as a central vestige, posterior transverse carina inter- rupted centrally, laterally quite strong; lateromedian longitudinal carinae vestigial; lateral longitudinal carina present anteriorly, joined to spiracular margin. Fore wing length 14 mm; CI = 0.90; ICI = 0.54; SDI = 1.16; cu-a opposite to the base Rs&M; discosubmarginal cell very sparsely hirsute, with moderately large glabrous area anteriorly, and with a glabrous area along Rs&M; Rs+2r joining pterostigma near centre; Ist subdiscal cell sparsely hirsute; 1m-cu centrally angled, ramellus slightly shorter than abscissa of 1m-cu between its base and bulla. Hind wing with 8 hamuli on R1, the distal ones more tightly curved than the proximal ones; Ist abscissa of Rs proximally abruptly curved through about 50°; marginal cell proximally quite short, with junction between Rs and M proximal to centre, and with distal abscissa of Rs quite long; proximal 0.3 of marginal cell more or less glabrous with a few scattered hairs centrally; distal abscissa of Cul joining cu-a more or less intermediate between M and 1A. Fore leg with tibia quite strongly with scattered spines on outer surface; fore tibial spur with an unusually fine microtrichial comb. Mid leg with longer tibial spur 1.5 times length of the shorter. Hind leg with coxa in profile 1.6 times as long as deep; trochantellus dorsally 0.2 times as long as broad; hind tarsus slender, the 4th segment 2.8 times as long as broad; claws of female long and weakly curved, with long pectinae. Gaster stout; tergite 2 in profile 2.6 times as long as posteriorly deep; thyridia obovate and separated from anterior margin of tergite by about 0.5 times its own length. Female with subgenital plate long but unspecialized; ovipositor moderately long and straight, its sheath slender. Male unknown. Colour generally orange; interocellar area yellowish orange; flagellum orange; pterostigma orange; wings distinctly yellowish. REMARKS. Ophion euterpe is immediately distinguishable from all other southern Mesoamerican species of Ophion by its grossly swollen temples. The stout gaster, enlarged subgenital plate, the proximally glabrous discosubmarginal cell, slightly flattened distal tarsal segments and weak anterior propodeal transverse carina are features found both in this species, and in an even more modified form, in O. calliope suggesting the two may be related BIOLOGICAL INFORMATION. Only a single specimen of O. euterpe has been collected and it was taken at light in lowland rainforest on Barro Colorado Island, Panama. It is the only species of Ophion that has been collected in lowland rainforest. Nothing is known of the biology of this insect. MATERIAL EXAMINED Holotype 9, Panama: Barro Colorado Island, v.1978 (Wolda) (RNH). OPHIONINAE OF TROPICAL MESOAMERICA 35 Ophion calliope sp. n. (Figs 26, 36, 37, 39) DEscrIPTION. Mandibles stout, very weakly narrowed apically, with upper tooth similar to and about as long as the lower tooth; outer mandibular surface punctate, with microreticulation between punctures, and with basal part of mandible broadly concave; distal segment of maxillary palp slightly incrassate; malar space ().1 times as long as basal mandibular width. Head in front view with greatest width across eyes 1.4 times length of head from vertex to clypeal margin; orbits ventrally slightly convergent; clypeus in profile weakly convex, with isolated punctures and with margin strongly impressed. Lower face centrally convex, polished and punctate. Head in dorsal view with genae rounded behind eyes; posterior ocellus contiguous with eye; occipital carina mediodorsally rather weak, convex, ventrally curved to join hypostomal carina almost at mandible. Antenna short and stout, with 47-51 flagellar segments; 20th segment 1.5—1.6 times as long as broad. Mesoscutum polished and finely punctate, in profile abruptly rounded; notauli weakly impressed only near anterior end. Mesopleuron polished, centrally finely and quite sparsely punctate; lower corner of epicnemial carina bluntly rounded after concavity, and with upper end inclined towards anterior margin of pleuron, and joining it just above level of lower corner of pronotum. Scutellum in profile evenly convex, laterally carinate for 0.3—0.5 of its length. Metapleuron convex, finely punctate. Propodeum in profile abruptly rounded; anterior transverse carina indistinct, posterior transverse carina interrupted centrally, laterally strongly raised into crests; lateromedian longitudinal carinae weak in posterior part; lateral longitudinal carina absent, not joined to spiracular margin by a short carina. Fore wing length 19-21 mm; CI = 0.65-0.70; ICI = 0.99-1.03; SDI = 1.19-1.30; cu-a more or less opposite to the base Rs&M; discosubmarginal cell sparsely hirsute, with moderately large glabrous area anteriorly and with narrow glabrous area along Rs&M; Rs+2r joining pterostigma close to base; Ist subdiscal cell sparsely hirsute; 1m-cu centrally angled, ramellus generally longer than abscissa of lm-cu between its base and bulla. Hind wing with 9-10 hamuli on R1, the distal ones more tightly curved than the proximal ones; 1st abscissa of Rs proximally abruptly curved through about 70°; marginal cell proximally short, with junction between Rs and M far proximal to centre, and with distal abscissa of Rs long; proximal 0.3 of marginal cell with a central hirsute area; distal abscissa of Cu1 joining cu-a almost intermediate between M and 1A. Fore leg with tibia slightly flattened, with numerous spines on outer surface. Mid leg with longer tibial spur 1.2-1.3 times length of the shorter. Hind leg with coxa in profile 1.6-2.0 times as long as deep; trochantellus dorsally 0.1—-0.2 times as long as broad; all tarsi ventrally flattened, especially the distal two segments; hind tarsus with the 4th segment 1.3—1.5 times as long as broad; claws of female long and weakly curved, with long close pectinae, those of male similar but with pectinae finer and close together. Gaster stout; tergite 2 in profile about 3 times as long as posteriorly deep; thyridia elliptical and separated from anterior margin of tergite by about 0.30.5 times its own length. Female with subgenital plate large, in profile plough-share shaped, ventrally convex and projecting beyond apex of gaster; posterior margin medially incised; ovipositor short and decurved, its apex slender; ovipositor sheath stout. Male with subgenital plate transverse, finely hirsute; gonosquama short, apically obliquely truncate and dorsally with a weak subapical notch; aedeagus simple, but with weak rounded lateral keels. Colour uniformly orange-brown with mesoscutal vittae weakly infuscate, and head yellowish; interocel- lar area yellow; flagellum black, scape and pedicel orange; pterostigma orange; wings slightly yellowish. REMARKS. This is one of the most distinctive American species of Ophion on account of the highly modified subgenital plate and the stout and strongly flattened distal tarsal segments. No other southern Meso- american species has a black flagellum like this insect. The relationships of O. calliope are not clear, but slightly modified tarsi and a slightly enlarged subgenital plate are found in O. euterpe and the two taxa may be related. This hypothesis of relationship is strengthened by the fact that both species also have a narrow glabrous band along the proximal periphery of the discosubmarginal cell, and have more or less lost the anterior transverse carina of the propodeum. BIOLOGICAL INFORMATION. In Costa Rica Ophion calliope occurs in lower montane humid forests at altitudes between 700 and 1400 m. This is the only species of Ophion known to occur in intact forests at this altitude, though other species occur in disturbed areas at a similar altitude. Nothing is known of the biology of this insect though the form of the ovipositional apparatus suggests it might be attacking a young hairy or spined host. This is suggested by the fact that the subgenital plate forms a sort of shield that only allows the extreme apex of the ovipositor to project. The apex of the gaster is thus protected, and the ovipositor cannot penetrate far into the host. 36 IAN D. GAULD MATERIAL EXAMINED Holotype 2, Costa Rica: Alajuela Prov.: San Ramon Reserve, Rio San Lorencito, 800 m, xi.1986 (Chacon) (BMNH) Paratypes. Costa Rica: Alajuela Prov.: 1 2, San Ramon Reserve, Rio San Lorencito, 800 m, xi.1986 (Chacon) (BMNH); Puntarenas Prov.: 7 0’, 13 9, Finca Las Cruces, 6 km S of San Vito de Java, 1400 m, x.1986 (Eger) (BMNH, CNC, MNCR, TC); San José Prov: 1 9, Braulio Carrillo National Park, Estacion Carrillo, 700 m, v.1985 (Chacon) (BMNH). Ophion polyhymniae sp. n. (Figs 42, 43) DescriPTIon. Mandibles stout, weakly narrowed apically, with upper tooth slightly broader and longer than the lower tooth, with distal edge slightly concave; outer mandibular surface coriaceous proximally, distally polished and sparsely punctate and with basal part of mandible with a small concave area; distal segment of maxillary palp slender; malar space 0.4—0.5 times as long as basal mandibular width. Head in front view with greatest width across eyes 1.1—1.2 times length of head from vertex to clypeal margin; orbits ventrally slightly convergent; clypeus in profile weakly convex, with weak isolated punctures, apically with margin strongly impressed. Lower face centrally weakly convex, weakly polished, with fine sparse punctures and with the area between them microreticulate/coriaceous. Head in dorsal view with genae rounded behind eyes; posterior ocellus separated from eye by about 0.1 times its maximum diameter; occipital carina mediodorsally arched upwards almost to a point, ventrally obsolescent, not clearly joining hypostomal carina. Antenna quite long and slender, with 51-55 flagellar segments; 20th segment 1.7-2.0 times as long as broad. Mesoscutum polished, coriaceous, in profile evenly rounded; notauli vestigial, only distinguishable as slight impressions near anterior end. Mesopleuron polished, centrally finely and very closely punctate; lower corner of epicnemial carina about a right angle after broad concavity, and with upper end inclined towards anterior margin of pleuron, which it reaches just above level of lower corner of pronotum. Scutellum in profile fairly weakly convex, not laterally carinate. Metapleuron weakly convex, finely punctate, with intervening area coriaceous. Propodeum in profile evenly rounded; anterior transverse carina complete, posterior transverse carina broadly incomplete centrally, laterally quite strong; laterome- dian longitudinal carinae absent; lateral longitudinal carina present anteriorly, joined to spiracular margin. Fore wing length 17-19 mm; CI = 0.50-0.57; ICI = 0.51-0.62; SDI = 1.17—1.20; cu-a subopposite to the base Rs&M,; discosubmarginal cell evenly hirsute, with moderately large glabrous area anteriorly, and without a glabrous area along Rs&M; Rs+2r joining pterostigma close to centre; Ist subdiscal cell fairly evenly hirsute; 1mm-cu centrally angled, ramellus distinctly shorter than abscissa of 1m-cu between its base and bulla. Hind wing with 9-11 hamuli on R1, the distal ones more tightly curved than the proximal ones; 1st abscissa of Rs proximally abruptly curved through about 60°; marginal cell proximally short, with junction between Rs and M far proximal to centre, and with distal abscissa of Rs long; proximal 0.3 of marginal cell glabrous peripherally but centrally hirsute; distal abscissa of Cu1 usually joining cu-a closer to 1A than to M, but sometimes intermediate between the veins. Fore leg with tibia subcylindrical without obvious spines on outer surface. Mid leg with longer tibial spur 1.4 times length of the shorter. Hind leg with coxa in profile 2.0—2.1 times as long as deep; trochantellus dorsally 1.0—1.1 times as long as broad; hind tarsus slender, the 4th segment 2.3—2.7 times as long as broad; claws of female long and weakly curved, with long pectinae, those of male similar but with pectinae slightly shorter and closer together. Gaster slender; tergite 2 in profile more than 4 times as long as posteriorly deep; thyridia elliptical and separated from anterior margin of tergite by about 0.5 times its own length. Female with subgenital plate moderately long, unspecialized; ovipositor moderately long, straight and apically slender, its sheath slender. Male with subgenital plate transverse, unspecialized; apex of aedeagus rounded. Colour generally blackish brown or dark reddish brown, orbits and sometimes vertex indistinctly suffused with whitish yellow; flagellum orange brown, proximally infuscate; mesepimeron orange-brown; legs reddish brown; flagellum orange; pterostigma orange-brown; wings slightly infumate. VARIATION. There is some variation in propodeal carination, and although most individuals have the anterior transverse carina complete it may be weak or even virtually absent in some specimens. Many other Ophion species show great variation in propodeal carination and the range of variation exhibited here is not considered exceptional REMARKS. Ophion polyhymniae is immediately distinguishable from all other southern Mesoamerican species of Ophion by its dark coloration, its rather elongate head and wide malar space. The rather fine OPHIONINAE OF TROPICAL MESOAMERICA 37 microreticulate sculpture observable over much of the alitrunk of O. polyhymniae is also found in O. melpomene. However, on the basis of this character I hesitate to suggest that these two species are closely related because similar sculpture is present on species from high altitudes in other regions, including the Himalayas and New Guinea. This suggests that this sculptural character may, in some way be an adaptation to high altitude existence, and perhaps a poor indicator of phylogenetic affinity BIOLOGICAL INFORMATION. Ophion polyhymniae occurs in montane habitats on the Cerro de la Muerte in Costa Rica where individuals have been collected quite frequently (considering how often the habitat has been sampled) between March and June in Quercus forests at altitudes of 2700-3200 m. O. polyhymniae flies at dusk and in the early part of the night; after about 2100 h individuals seem to be very lethargic. The hosts of this insect are not known. MATERIAL EXAMINED Holotype CO’, Costa Rica: Cartago Prov.:;Cerro de la Muerte, 1 km NE. Cerro Asuncion, 3100 m, iii. 1985 (Janzen & Hallwachs) (BMNH) Paratypes. Costa Rica: Cartago Prov: 1 0’, Cerro de la Muerte, 2700 m, vi.1986 (Gauld) (BMNH); San José Prov.: 3 0’, Cerro de la Muerte, 2 km E. Cerro Asuncion, 3140 m, iv.1984 (Janzen, Hallwachs & Gauld) (BMNH); 1 o’, 2 2, Cerro de la Muerte, 3200 m, iii-iv.1985 (Masner & Goulet) (CNC). Ophion flavidus Brullé (Figs 23, 41, 44, 45) Ophion flavidus Brullé, 1846 : 143. Lectotype 9, BRAZIL (MNHN), designated by Townes & Townes (1966: 168) [examined]. Ophion biangularis Taschenberg, 1875: 432. Holotype 2, BRAZIL (Halle). [Synonymized by Townes & Townes, 1966: 168.] Ophion ancyloneura Cameron, 1886: 294. Holotype 9, GUATEMALA (BMNH) [examined]. [Syn- onymized by Townes & Townes, 1966: 168.] Ophion diversus Szépligeti, 1906: 131. Lectotype 9, PARAGUAY (TM), designated by Townes & Townes (1966: 168) [Synonymized by Townes & Townes, 1966: 168. ] Ophion concolor Szépligeti, 1906: 132. Lectotype Oo’, ARGENTINA (TM), designated by Townes & Townes (1966: 169) [Synonymized by Townes & Townes, 1966: 169. ] Ophion politior Morley, 1912: 56. Lectotype 9, BRAZIL (BMNH), designated by Townes & Townes (1966: 169) [examined]. [Synonymized by Townes & Townes, 1966: 169.] Description. Mandibles moderately stout, weakly narrowed apically, more or less equally bidentate; outer mandibular surface punctate, coriaceous between punctures, and with basal part of mandible broadly concave; distal segment of maxillary palp slender; malar space 0.1 times as long as basal mandibular width. Head in front view with greatest width across eyes 1.3—1.4 times length of head from vertex to clypeal margin; orbits ventrally slightly convergent; clypeus in profile rather weakly convex, with isolated punc- tures with margin quite strongly impressed. Lower face centrally weakly convex, polished and quite densely punctate. Head in dorsal view with genae rounded behind eyes; posterior ocellus very close to but not contiguous with eye; occipital carina mediodorsally convex, ventrally curved to join hypostomal carina well above base mandible. Antenna quite long and stout, with 53-59 flagellar segments; 20th segment 1.7— 1.6 times as long as broad. Mesoscutum polished, finely punctate, in profile evenly rounded; notauli fairly weakly impressed near anterior end, but readily discernible as rugose area. Mesopleuron polished, centrally usually closely punctate, sometimes with traces of microreticulation present between punctures, rarely with punctures small and intervening area smooth and highly polished; lower corner of epicnemial carina bluntly rounded after concavity, and with upper end inclined towards anterior margin of pleuron, which it generally reaches just above lower corner of pronotum. Scutellum in profile weakly convex, laterally carinate on anterior 0.1-0.3. Metapleuron quite strongly convex, punctate. Propodeum in profile abruptly rounded; anterior transverse carina almost invariably complete, posterior transverse carina complete or centrally obsoles- cent; lateromedian longitudinal carinae generally strong and reaching to anterior transverse carina, thus enclosing distinct areae superomedia and petiolaris, or weaker and only enclosing the area petiolaris; lateral longitudinal carina complete, joined to spiracular margin. Fore wing length 10-14 mm; CI = 0.44-0.60; ICI = 0.50-0.63; SDI = 1.15—1.29; cu-amore or less opposite to the base Rs&M; discosubmarginal cell sparsely hirsute, with moderately large glabrous area anteriorly, without a glabrous area along Rs&M; Rs+2r joining pterostigma close to centre; 1st subdiscal cell sparsely hirsute; 1m-cu centrally angled, ramellus shorter than abscissa of 1m-cu between its base and bulla. Hind 38 IAN D. GAULD wing with 5-8 hamuli on R1, the distal ones more tightly curved than the proximal ones; 1st abscissa of Rs proximally abruptly curved through about 85°; marginal cell proximally long, with junction between Rs and M near to centre, and with distal abscissa of Rs quite short; proximal 0.3 of marginal cell centrally closely hirsute, periphery glabrous; distal abscissa of Cu1 joining cu-a very much closer to 1A than to M. Fore leg with tibia subcylindrical, with isolated spines on outer surface. Mid leg with longer tibial spur 1.4-1.5 times length of the shorter. Hind leg with coxa in profile 1.8—2.0 times as long as deep; trochantellus dorsally 0.40.6 times as long as broad; hind tarsus slender, the 4th segment 2.62.9 times as long as broad; claws of female long and weakly curved, with long close pectinae; claws of male similar but with pectinae shorter and closer together. Gaster quite slender; tergite 2 in profile about 4 times as long as posteriorly deep; thyridia elliptical and separated from anterior margin of tergite by about 0.5 times its own length. Female with subgenital plate large, unspecialized, triangular in profile; ovipositor moderately long, straight and slender, its sheath slender. Male with subgenital plate transverse, unspecialized; aedeagus apically slender, rather abruptly angled Colour generally pallid yellowish; interocellar area yellow; flagellum orange; pterostigma orange; wings very slightly yellowish. VARIATION. The most obvious variation occurs in the extent of the propodeal carination. Generally the anterior and posterior transverse carinae are very strong and usually the anterior one is laterally abruptly curved back to join the posterior one; the lateromedian longitudinal carinae are generally complete to the anterior carina so the areae petiolaris, superomedia and dentipara are delineated. However, any or all of these carinae may be weak or indistinct (though the anterior one is never absent if the posterior one is present), so at the most extreme no areae are enclosed. Although most specimens are pallid in colour, isolated individuals may be darker brownish orange and I have seen one individual which has dark mesoscutal vittae. RemARKS. Ophion flavidus can be most easily recognized by the proximally elongated marginal cell in the hind wing. No other Mesoamerican species has this cell so lengthened. The rather pallid colour of most specimens is also quite distinctive as most other Ophion species are darker reddish brown BIOLOGICAL INFORMATION. Ophion flavidus is one of the commonest and most widely distributed American species of Ophioninae. It has been recorded from New York State south to Florida and Texas and west to Kansas in the United States (Carlson, 1979), and it occurs throughout the Caribbean (Wolcott, 1923; Gowdey, 1926) and Latin America south to Argentina (Costa Lima, 1962; Townes & Townes, 1966; de Santis & Esquivel, 1966). In Central America it is frequently encountered in the drier lowlands on the Pacific side of the continent, and O. flavidus is the only species of Ophion known to occur in Santa Rosa National Park, Costa Rica, where it is quite common at the start of the wet season. The cumulative seasonal data (lights and malaise traps) for 1984-6 are: Jick oi AM, de Ande dO Mee Ale rg abbqyare vie O. flavidus also seems to occur commonly in agricultural and even suburban areas at higher elevations up to about 1300 m, such as at Monteverde or San Antonio de Escazti in Costa Rica. At the former site this species appears to be less seasonal than it does in the lowlands and I have seen isolated specimens collected during all months of the year. A large number of individuals were found in association with the grass-eating noctuid Mocis repanda (F.) on Volcan Cacao, though none was actually reared. I have also seen a number of individuals from higher altitudes, such as Guadalupe Arriba in Panama (2200 m), but I have only seen a single specimen that has been collected above 2400 m. At Guadalupe Arriba O. flavidus is comparatively rarely collected and the cumulative seasonal data from two years’ collecting with a light trap are: Jo ReveM cAnieibbheSeoldioed) cS acONreD S..0=-aolet) clr see eisnertugsee2 O. flavidus is a common endoparasitoid of the larvae of a variety of species of Noctuidae that feed on herbaceous vegetation in disturbed and agricultural habitats and include some notorious pests. It is thus an economically important insect. For example, in Alabama, O. flavidus is one of the most common larval parasitoids of the Fall Armyworm, Spodoptera frugiperda (Smith), and investigations are currently being undertaken with a view to maximizing its potential for biological control purposes (Rohlfs & Mack, 1985b). There are also reliable records of this species parasitizing Agrotis ipsilon (Hufnagel), Heliothis zea OPHIONINAE OF TROPICAL MESOAMERICA 39 (Boddie), Peridroma saucia (Hiibner), Pseudaletia unipuncta (Haworth) and Spodoptera eridania (Cramer) (Costa Lima, 1962; Carlson, 1979). The biology of O. flavidus has recently been studied by Rohlfs & Mack (1983; 1985a & b), and the details given by Vickery (1927) about O. bilineatus Say may actually refer to this species. MATERIAL EXAMINED Lectotype 2 (Ophion flavidus Brullé), Brazil: ‘Ouest’ (MNHN). Holotype 2 (Ophion ancyloneura Cameron) Guatemala: Capetillo (BMNH). Lectotype 2 (Ophion politior), Brazil: Rio Grande (BMNH); paralectotypes 2 2, same locality as lectotype. Costa Rica: Alajuela Prov.: 1 9, Finca San Gabriel, 3 km W. Dos Rios, 850 m, vi.1986 (Gauld) (BMNH); 1 2, 8.2 km N Vara Blanca, iv.1984 (Janzen & Hallwachs) (BMNH); Guanacaste Prov: 2 0’, 29 Q, Casa Mengo, SW. side Volcan Cacao, 1000 m, vi-x.1987 (Janzen & Hallwachs) (BMNH); 1 9, Finca Biesnan, 11 km E Quebrada Grande, 500 m, vi.1985 (Gauld) (BMNH); 1 0',5 km NE. Quebrada Grande, 600 m, vi.1986 (Gauld) (BMNH); 1 ©’, Rincon de la Vieja National Park, 900 m, iii.1984 (Janzen, Hallwachs & Gauld) (BMNH); 22 o’, 29 2, Santa Rosa National Park, 300 m, months as above 1984-7 (Janzen, Hallwachs & Gauld) (BMNH; MNCR); Puntarenas Prov.: 13 0’, 11 2, Monteverde, 1300 m, i-xii. 1985-6 (Haber) (BMNH): San José Prov: 1 0’, Cerro de la Muerte, San Gerardo de Dota, 2430 m, xii. 1981 (Janzen & Hallwachs) (BMNH); 1 0’, 1 9, San Antonio de Escazu, 1300 m, v-vi.1981 (Eberhard) (UM). Guatemala: 1 co’, 10km W. Amatitlan, 1200 m, vi.1974 (O’Brien) (BMNH). Dominican Republic: 1 CO’, Santiago Prov., La Cumbre, 1000 m, iv.1978 (Woodruff) (BMNH). Nicaragua: 2 O' , Grenada, 3km W. of Nandaime, vii.1974 (O’Brien) (FSCA); 2 0’, 1 2, Leon, vii.1985 (BMNH, ULN). Panama: 3 CO’, 13 9, Chiriqui, Guadalupe Arriba, 2200 m, months as above, 1984/5 (Wolda) (BMNH). U.S.A.: Florida: 2 0’, 1 ©, Alachua Co., Gainesville, x.1971, iii, x.1972 (Mead) (FSCA); 1 9, same locality, iii.1980 (Stange) (FSCA); 2 o’, 2 2, Hillsborough Co., Tampa, iii.1987 (Eger) (BMNH); 1 2, Marion Co., 14 km SSW. Ocala, x.1975 (Wiley) (FSCA). Ophion clio sp. n. Description. Mandibles stout, very weakly narrowed apically, with upper tooth slightly broader and slightly longer than the lower tooth, with its blade slightly indented apically; outer mandibular surface punctate, with microreticulation between punctures, and with basal part of mandible broadly concave; distal segment of maxillary palp slender; malar space 0.1 times as long as basal mandibular width. Head in front view with greatest width across eyes 1.4—1.5 times length of head from vertex to clypeal margin; orbits ventrally slightly convergent; clypeus in profile moderately convex, with isolated punctures, apically slightly coriaceous and with margin strongly impressed. Lower face centrally convex, polished and sparsely punctate. Head in dorsal view with genae constricted behind eyes; posterior ocellus contiguous with eye; occipital carina mediodorsally convex at centre, slightly produced upwards, ventrally curved to join hypostomal carina well above base mandible. Antenna quite long and stout, with 60-63 flagellar segments; 20th segment 1.7—1.8 times as long as broad. Mesoscutum weakly polished and indistinctly punctate, in profile evenly rounded; notauli strongly impressed near anterior end. Mesopleuron polished, centrally finely and quite sparsely punctate; lower corner of epicnemial carina bluntly rounded after concavity, and with upper end inclined towards anterior margin of pleuron, but with upper end evanescent. Scutellum in profile weakly convex, not laterally carinate. Metapleuron moderately convex, punctate. Propodeum in profile abruptly rounded; anterior transverse carina more or less complete, posterior transverse carina more or less complete, laterally strongly raised into crests; lateromedian longitudinal carinae weak but more or less complete; lateral longitudinal carina complete, joined to spiracular margin. Fore wing length 25—27 mm; CI = 0.63-0.70; ICI = 0.77-0.86; SDI = 1.24—-1.26; cu-amore or less opposite to the base Rs&M; discosubmarginal cell sparsely hirsute, with moderately large glabrous area anteriorly, without a glabrous area along Rs&M; Rs+2r joining pterostigma slightly proximal to centre; 1st subdiscal cell very sparsely hirsute; 1m-cu centrally angled, ramellus much shorter than abscissa of 1m-cu between its base and bulla. Hind wing with 9-10 hamuli on R1, the distal ones more tightly curved than the proximal ones; Ist abscissa of Rs proximally abruptly curved through about 70[; marginal cell proximally short, with junction between Rs and M far proximal to centre, and with distal abscissa of Rs long; proximal 0.3 of marginal cell very sparsely pubescent centrally, becoming glabrous; distal abscissa of Cul joining cu-a slightly closer to 1A than to M. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.1—-1.2 times length of the shorter. Hind leg with coxa in profile 1.9-2.0 times as long as deep; trochantellus dorsally 0.4—0.6 times as long as broad; hind tarsus slender, the 4th segment 2.6—2.9 times as 40 IAN D. GAULD long as broad; claws of female long and weakly curved, with long close pectinae; claws of male similar but with pectinae shorter and closer together. Gaster quite slender; tergite 2 in profile about 4 times as long as posteriorly deep; thyridia elliptical and separated from anterior margin of tergite by about 0.5 times its own length. Female with subgenital plate unspecialized; ovipositor moderately long and straight, its sheath slender. Male with subgenital plate transverse, unspecialized; aedeagus apically simple, rounded. Colour generally yellowish with darker orange on face centrally, mesoscutum and mesosternum; gaster with segments pale yellow centrally, with dorsal, ventral and posterior margins slightly infuscate so gaster appears mottled; interocellar area yellow; flagellum orange; pterostigma orange; wings very slightly yellowish. VARIATION. The marginal cell of the hind wing of the male is entirely glabrous proximally. Remarks. Ophion clio is the largest southern Mesoamerican species of the genus. It is most easily recognized by the combination of this large size, its coloration, and the sculpture of the propodeum — the posterior transverse carina is strongly raised laterally, and is close to the anterior carina so the weakly discernible area superomedia is subquadrate. O. clio is partially sympatric and synchronous with another large species, O. uraniae, but the two differ in colour, shape of the aedeagus, and sculpture of the propodeum. BIOLOGICAL INFORMATION. Ophion clio occupies sites at altitudes ranging from 2200 to above 3100 m on the Cerro de la Muerte, Costa Rica where I have collected it in Quercus forests. At Guadalupe Arriba, Panama, it is uncommon and two years light-trapping yielded only a single specimen. Nothing is known of the biology of this insect. MATERIAL EXAMINED Holotype 2, Costa Rica: San José Prov.: Cerro de la Muerte, 2 km E. Cerro Asuncion, 3140.m, iv. 1984 (Janzen, Hallwachs & Gauld) (BMNH) Paratypes. Costa Rica: San José Prov: 1 0’, Cerro de la Muerte, San Gerardo de Dota, 2430 m, xii.1981 (Janzen & Hallwachs) (BMNH); 1 @, Cerro de la Muerte, 2 km E. Cerro Asuncion, 3140 m, iv.1984 (Janzen, Hallwachs & Gauld) (BMNH). Panama: 1 9, Chiriqui, Guadalupe Arriba, v.1984 (Wolda) (BMNH). Ophion cacaoi sp. n. (Fig. 30) Description. Mandibles stout, very weakly narrowed apically, with upper tooth slightly longer and stouter than the lower tooth; outer mandibular surface punctate, quite polished, with microreticulation between punctures only discernible basally, and with basal part of mandible broadly concave; distal segment of maxillary palp slender; malar space 0.1 times as long as basal mandibular width. Head in front view with greatest width across eyes 1.4-1.5 times length of head from vertex to clypeal margin; orbits ventrally slightly convergent; clypeus in profile moderately convex, with isolated punctures, polished and with margin strongly impressed. Lower face centrally convex, polished and closely punctate. Head in dorsal view with genae rounded behind eyes; posterior ocellus contiguous with eye; occipital carina mediodor- sally convex, at centre slightly produced upwards, ventrally curved to join hypostomal carina well above base mandible. Antenna quite long and fairly stout, with 58-60 flagellar segments; 20th segment 1.5—1.7 times as long as broad. Mesoscutum weakly polished with fine weak close punctures, and with intervening area slightly cor- iaceous; mesoscutum in profile evenly rounded; notauli strongly impressed before anterior end, extreme anterior end weak. Mesopleuron polished, centrally finely punctate, with intervening area microreticulate; lower corner of epicnemial carina slightly acutely angled after strong concavity, and with upper end abruptly turned towards anterior margin of pleuron, but usually evanescent, though sometimes reaching pleural margin just above level of lower corner of pronotum. Scutellum in profile weakly convex, not laterally carinate. Metapleuron strongly convex, finely punctate but with fine coriaceous sculpture on areas between punctures. Propodeum in profile rather evenly declivous, weakly polished; anterior transverse carina more or less complete, posterior transverse carina more or less complete, laterally raised into crests; lateromedian longitudinal carinae usually complete to anterior transverse carina, enclosing a subquadrate area superomedia; lateral longitudinal carina complete, joined to spiracular margin by a raised ridge. Fore wing length 17-18 mm; CI = 0.36-0.50; ICI = 0.84-1.00; SDI = 1.01—1.05; cu-a slightly proximal to the base Rs&M,; discosubmarginal cell sparsely hirsute, with moderately small glabrous area anteriorly, without a glabrous area along Rs&M; Rs+2r joining pterostigma slightly proximal to its centre; 1st OPHIONINAE OF TROPICAL MESOAMERICA 41 subdiscal cell sparsely but fairly uniformly hirsute; 1m-cu centrally rounded, weakly angled, ramellus extremely short, at most about 3 times as long as broad, very much shorter than abscissa of 1m-cu between its base and bulla. Hind wing with 6—7 hamuli on R1, the distal ones more tightly curved than the proximal ones; 1st abscissa of Rs proximally fairly evenly curved through about 70°; marginal cell proximally short, with junction between Rs and M far proximal to centre, and with distal abscissa of Rs long; proximal 0.3 of marginal cell fairly evenly hirsute, not peripherally glabrous; distal abscissa of Cu1 joining cu-a more of less equidistantly between M and 1A. Fore leg with tibia slightly flattened, with isolated spines on outer surface. Mid leg with longer tibial spur 1.4-1.5 times length of the shorter. Hind leg with coxa in profile 1.9-2.2 times as long as deep; trochantellus dorsally 0.10.2 times as long as broad; hind tarsus slender, the 4th segment 2.7—2.8 times as long as broad; claws of female long and weakly curved, with long close pectinae; claws of male similar but with pectinae shorter and closer together. Gaster moderately slender; tergite 2 in profile about 3 times as long as posteriorly deep; thyridia elliptical and separated from anterior margin of tergite by about 0.5 times its own length. Female with subgenital plate unspecialized; ovipositor moderately long and straight, its sheath slender. Male with subgenital plate transverse, unspecialized; aedeagus apically curved, culminating in a short acute process. Colour generally reddish brown, with orbits, vertex and interocellar area yellowish, mesoscutal margin, scutellum and two marks on upper part of mesopleuron yellowish; flagellum orange-brown; pterostigma orange; wings quite strongly yellowish. REMARKS. Ophion cacaoi is structurally rather similar to O. uraniae. Both species have similarly modified epicnemial carinae and apically acute aedeagi, suggesting they are a sister-species pair. However, they differ consistently in a number of features. O. cacaoi has a very short ramellus, a more or less evenly hirsute marginal cell in the hind wing and the area superomedia is distinct and subquadrate to transverse; O. uraniae has a longer ramellus, a proximally narrowly glabrous marginal cell in the hind wing and, if its area superomedia is discernible, then it is elongate. O. cacaoi also has a larger ICI than uraniae, a shorter and stouter hind trochantellus, stouter antennae, and has two rather than three pale marks on the upper part of the mesopleuron. BIOLOGICAL INFORMATION. Ophion cacaoi is only known to occur in forests on the upper slopes of Volcan Cacao in north-western Guanacaste, Costa Rica. This mountain, together with Volcan Orosi, provides a rather isolated island of moderate altitude forest at the northern end of the Cordillera Guanacaste. Possibly this species arose from a geographically isolated population of O. uraniae, or the ancestral population of both was split by a vicariance event. Nothing is known of the biology of O. cacaoi. MATERIAL EXAMINED Holotype ©’, Costa Rica: Guanacaste Prov., Casa Mengo on SW. side of Volcan Cacao, 1000 m, vi.1987 (Janzen) (BMNH). Paratypes. 3 CO’, 2 2, same locality as holotype, vi-vii.1987 (Janzen) (BMNH). Ophion uraniae sp. n. (Figs 47, 49) Description. Mandibles stout, very weakly narrowed apically, with upper tooth slightly longer than the lower tooth; outer mandibular surface punctate, quite polished, with microreticulation between punctures only discernible basally, and with basal part of mandible broadly concave; distal segment of maxillary palp slender; malar space 0.1 times as long as basal mandibular width. Head in front view with greatest width across eyes 1.3—1.5 times length of head from vertex to clypeal margin; orbits ventrally slightly convergent; clypeus in profile moderately convex, with isolated punctures, polished and with margin strongly impressed. Lower face centrally convex, polished and quite closely punctate. Head in dorsal view with genae constricted behind eyes; posterior ocellus contiguous with eye; occipital carina mediodorsally convex, at centre slightly produced upwards, ventrally curved to join hypostomal carina well above base mandible. Antenna quite long and fairly slender, with 54-57 flagellar segments; 20th segment 1.8-2.5 times as long as broad. Mesoscutum weakly polished with fine weak close punctures, and with intervening area slightly cor- iaceous; mesoscutum in profile evenly rounded; notauli moderately impressed near anterior end. Meso- pleuron polished, centrally finely punctate, with intervening area microreticulate; lower corner of epicnemial carina acutely angled after concavity, and with upper end abruptly turned towards anterior margin of pleuron, but usually evanescent, though sometimes reaching pleural margin just above level of lower corner of pronotum. Scutellum in profile weakly convex, not laterally carinate. Metapleuron strongly convex, finely punctate. Propodeum in profile rather evenly declivous, strongly polished; anterior 42 IAN D. GAULD transverse carina more or less complete, posterior transverse carina more or less complete, laterally raised into crests; lateromedian longitudinal carinae usually complete to posterior transverse carina, less fre- quently present to anterior transverse carina in which case it encloses an elongate area superomedia; lateral longitudinal carina complete, joined to a rather remote spiracular margin by a raised ridge. Fore wing length 16-21 mm; CI = 0.44-0.59; ICI = 0.44-0.63; SDI = 1.19-1.27; cu-a slightly proximal to the base Rs&M; discosubmarginal cell sparsely hirsute, with moderately small glabrous area anteriorly, without a glabrous area along Rs&M; Rs+2r joining pterostigma slightly proximal to its centre; 1st subdiscal cell sparsely hirsute; 1m-cu centrally angled, ramellus much shorter than abscissa of 1m-cu between its base and bulla. Hind wing with 6-8 hamuli on R1, the distal ones more tightly curved than the proximal ones; Ist abscissa of Rs proximally abruptly curved through about 80°; marginal cell proximally short, with junction between Rs and M far proximal to centre, and with distal abscissa of Rs long; proximal 0.3 of marginal cell centrally hirsute, peripherally glabrous; distal abscissa of Cu1 joining cu-a more or less equidistantly between M and 1A. Fore leg with tibia subcylindrical, with isolated spines on outer surface. Mia leg with longer tibial spur 1.3—1.4 times length of the shorter. Hind leg with coxa in profile 1.9-2.2 times as long as deep; trochantellus dorsally 0.40.5 times as long as broad; hind tarsus slender, the 4th segment 2.7—2.9 times as long as broad; claws of female long and weakly curved, with long close pectinae; claws of male similar but with pectinae shorter and closer together. Gaster quite slender; tergite 2 in profile about 4 times as long as posteriorly deep; thyridia elliptical and separated from anterior margin of tergite by about 0.5 times its own length. Female with subgenital plate unspecialized; ovipositor moderately long and straight, its sheath slender. Male with subgenital plate transverse, unspecialized; aedeagus apically culminating in a short acute process. Colour generally reddish brown, with orbits, vertex and interocellar area yellowish, mesoscutal margin, scutellum and three marks on upper part of mesopleuron yellow; flagellum orange; pterostigma orange; wings quite strongly yellowish. VARIATION. One specimen from Volcan Poas lacks the paler markings on the alitrunk. There is a tendency for many individuals to have the lateromedian longitudinal carinae of the propodeum fused to form a single median longitudinal carina REMARKS. Ophion uraniae is most easily recognized by the angulate lower corner of the epicnemial carina. Males are particularly distinctive on account of the apically acute aedeagus, and the colour pattern is generally quite distinctive. It resembles O. cacaoi in these features but the two species may be separated by the characters given in the key (and see also remarks under O. cacaoi). O. uraniae occurs at higher altitude than cacaoi, and the two species are geographically separated, the former occurs in Central Costa Rica and south into northern Panama, whilst the latter is only known from the extreme northern end of the Cordillera Guanacaste. O. uraniae has been collected in large numbers with O. thaliae, but the latter is smaller, has the epicnemial carina rounded ventrally and lacks any trace of a lateral longitudinal carina. BIOLOGICAL INFORMATION. Ophion uraniae occurs in central Costa Rica, and thence southwards at moder- ately high elevations (2200-2500 m) into northern Panama. It can occur in large numbers; 34 individuals were taken in Costa Rica during a single night at one site. It also seems to be active throughout the year as two years collecting at Guadalupe Arriba, Panama yielded the following seasonal distribution: Jit iieioAr OMI adie AGES OR NIND APO eh SANS] eae ore Nothing is known of the hosts of this insect. MATERIAL EXAMINED Holotype 2, Costa Rica: San José Prov: Cerro de la Muerte, San Gerardo de Dota, 2430 m, xii.1981 (Janzen & Hallwachs) (BMNH) Paratypes. Costa Rica: Alajuela Prov.: 1 0’, 1 2, Volcan Pods National Park, 2350 m, vii. 1982 (Janzen & Hallwachs) (BMNH); 2 0’, 1 9, same locality, xii. 1982 (Janzen & Hallwachs) (BMNH): San José Prov: 17 oO’, 17 2, Cerro de la Muerte, San Gerardo de Dota, 2430 m, xii.1981 (Janzen & Hallwachs) (BMNH, CNC, MNCR, TC, USNM). Panama: 34 ©’, 26 9, Chiriqui, Guadalupe Arriba, 2200 m, months as above, 1984-5 (Wolda) (BMNH) OPHIONINAE OF TROPICAL MESOAMERICA 43 Ophion erato sp. n. (Figs 29, 52) Description. Mandibles stout, very weakly narrowed apically, with upper tooth of similar shape to but slightly longer than the lower tooth; outer mandibular surface polished, sparsely punctate, and with basal part of mandible slightly concave; distal segment of maxillary palp slender; malar space 0.1 times as long as basal mandibular width. Head in front view with greatest width across eyes 1.3 times length of head from vertex to clypeal margin; orbits ventrally subparallel; clypeus in profile moderately convex, with isolated punctures and with margin strongly impressed. Lower face centrally almost flat, polished and sparsely punctate. Head in dorsal view with genae rounded behind eyes; posterior ocellus separated from eye by about 0.1 times its own maximum diameter; occipital carina mediodorsally convex at centre, ventrally curved to join hypostomal carina well above base mandible. Antenna quite long and slender, with 48-55 flagellar segments; 20th segment 2.1—2.3 times as long as broad. Mesoscutum quite strongly polished and indistinctly microreticulate, in profile evenly rounded; notauli very weakly impressed near anterior end. Mesopleuron weakly polished, centrally finely microreticulate with minute sparse punctures; lower corner of epicnemial carina bluntly rounded after concavity, and with upper end abruptly curved towards anterior margin of pleuron which it reaches just above level of lower corner of pronotum. Scutellum in profile weakly convex, not laterally carinate. Metapleuron moderately convex, microreticulate with very fine punctures. Propodeum in profile abruptly declivous; anterior transverse carina more or less complete, posterior transverse carina present laterally; lateromedian longitudinal carinae weak, discernible posteriorly; lateral longitudinal carina from complete to present only in anterior half, joined to spiracular margin. Fore wing length 13-15 mm; CI = 0.40; ICI = 0.52-0.60; SDI = 1.29-1.31; cu-a proximal to the base Rs&M by about 0.3 times its own length; discosubmarginal cell sparsely hirsute, with a large glabrous area anteriorly, without a glabrous area along Rs&M; Rs+2r joining pterostigma close to base; 1st subdiscal cell anteriorly glabrous, posteriorly hirsute; 1m-cu centrally angled, ramellus unusually long, but shorter than abscissa of 1m-cu between its base and bulla. Hind wing with 5—6 hamuli on R1, the distal ones more tightly curved than the proximal ones; Ist abscissa of Rs proximally abruptly curved through about 60°; marginal cell proximally short, with junction between Rs and M far proximal to centre, and with distal abscissa of Rs long; proximal 0.3 of marginal cell very closely hirsute centrally, anteriorly narrowly glabrous, but otherwise without a distinct glabrous periphery; distal abscissa of Cu1 joining cu-a slightly closer to 1A than to M. Fore leg with tibia subcylindrical, without obvious spines on outer surface. Mid leg with longer tibial spur 1.4-1.6 times length of the shorter. Hind leg with coxa in profile 1.7—2.0 times as long as deep; trochantellus dorsally 0.5—0.7 times as long as broad; hind tarsus slender, the 4th segment 2.8—3.0 times as long as broad; claws of female very long and weakly curved, with long close pectinae, those of male similar but with pectinae much closer together and with apical tooth unusually short. Gaster quite slender; tergite 2 in profile about 4 times as long as posteriorly deep; thyridia elliptical and separated from anterior margin of tergite by about 0.5 times its own length. Female with subgenital plate long but unspecialized; ovipositor quite short and fairly stout; ovipositor sheath slender. Male with subgenital plate transverse, unspecialized; aedeagus apically simple, rounded. Colour generally bright yellowish with mesoscutal vittae, mesosternum and tergites 59 infuscate; legs and antennae golden;pterostigma centrally brown, peripherally pallid; wings almost hyaline. VARIATION. The 2nd discal cell of the male is unusually short and the bulla in 1m-cu is closer to 2m-cu than it is to the ramellus. These very unusual features are not shared by the female. RemaRKS. Ophion erato is most distinctive because of the slender flagellum, the very long ramellus and the posteriorly infuscate gaster. The marginal cell of the hind wing is more uniformly hirsute proximo- posteriorly than are other species. The claws of the male are longer than those of many other species and have unusually short apical teeth that are no longer than the pectinae. BIOLOGICAL INFORMATION. Only two specimens of Ophion erato are known. One, a male, was collected at a moderately high altitude site on the Cerro de la Muerte, Costa Rica, and the female was taken at a similar altitude in northern Panama. This female was the only specimen of O. erato collected in two years’ continuous sampling. Its host is not known. MATERIAL EXAMINED Holotype ©’, Costa Rica: San José Prov: Cerro de la Muerte, San Gerardo de Dota, 2430 m, xii.1981 (Janzen & Hallwachs) (BMNH). Paratype. Panama: 1 9, Chiriqui, Guadalupe Arriba, 2200 m, x.1985 (Wolda) (BMNH). 44 IAN D. GAULD Ophion terpsichore sp. n. (Fig. 53) Description. Mandibles stout, very weakly narrowed apically, with upper tooth slightly broader and slightly longer than the lower tooth; outer mandibular surface punctate, coriaceous between punctures, and with basal part of mandible slightly concave; distal segment of maxillary palp slender; malar space 0.1 times as long as basal mandibular width. Head in front view with greatest width across eyes 1.35 times length of head from vertex to clypeal margin; orbits ventrally slightly convergent; clypeus in profile moderately convex, coriaceous, with isolated punctures and with margin strongly impressed. Lower face centrally convex, polished and sparsely punctate. Head in dorsal view with genae rounded behind eyes; posterior ocellus contiguous with eye; occipital carina mediodorsally convex, ventrally curved to join hypostomal carina well above base mandible. Antenna quite long and stout, with 52 flagellar segments; 20th segment 1.5 times as long as broad. Mesoscutum weakly polished, granulate and indistinctly punctate, in profile abruptly rounded; notauli weakly impressed near anterior end. Mesopleuron weakly polished, granulate, with very obscure minute punctures that are separated by about twice their own diameters; lower corner of epicnemial carina bluntly rounded after concavity, and with upper end abruptly curved to join anterior margin of pleuron above level of lower corner of pronotum. Scutellum in profile quite strongly convex, not laterally carinate. Meta- pleuron weakly convex, granulo-punctate. Propodeum in profile abruptly rounded; anterior transverse carina more or less complete, posterior transverse carina only present laterally as crests; lateromedian longitudinal carinae weak, more or less absent; lateral longitudinal carina weak but complete, joined to spiracular margin. Fore wing length 14 mm; CI = 0.61; ICI = 0.50; SDI = 1.15; cu-a slightly proximal to the base Rs&M; discosubmarginal cell evenly hirsute, with a large glabrous area anteriorly, without a glabrous area along Rs&M; Rs+2r joining pterostigma near centre; 1st subdiscal cell very sparsely hirsute; 1m-cu centrally angled, ramellus very long. Hind wing with 8 hamuli on R1, the distal ones more tightly curved than the proximal ones; Ist abscissa of Rs proximally evenly curved through about 80°; marginal cell proximally short, with junction between Rs and M far proximal to centre, and with distal abscissa of Rs long; proximal 0.3 of marginal cell fairly evenly hirsute centrally, peripherally glabrous; distal abscissa of Cu1 joining cu-a almost equidistant between M and 1A. Fore leg with tibia quite strongly flattened, with isolated spines on outer surface. Mid leg with longer tibial spur 1.3 times length of the shorter. Hind leg with coxa in profile 1.7 times as long as deep; trochantellus dorsally 0.4 times as long as broad; hind tarsus slender, the 4th segment 2.6 times as long as broad; claws of male long and weakly curved, with long close pectinae. Gaster quite slender; tergite 2 in profile about 4 times as long as posteriorly deep; thyridia elliptical and separated from anterior margin of tergite by about 0.5.times its own length. Female unknown. Male with subgenital plate transverse, unspecialized; aedeagus apically simple, rounded. Head, lateral part of alitrunk and scutellum bright yellowish; mesoscutum, except for lateral mark, mesopleuron in a narrow band below subalar prominence, mesosternum and propodeum dark brown; gaster with segments 1-3 dark brown, remainder more orange-brown; interocellar area yellow; flagellum orange, distally slightly darker; pterostigma orange, distally very pale yellowish; wings more or less hyaline. REMARKS. Ophion terpsichore is most easily recognized by the combination of its coloration, venation and the sculpture of the mesoscutum. No other Mesoamerican species of Ophion exhibits this colour pattern. The very long ramellus and ventrally dark mesosternum are derived features which this species shares with O. erato and the two may be related. BIOLOGICAL INFORMATION. Ophion terpsichore is only known to occur at one moderately high altitude site in Costa Rica. Nothing is known of its biology. MATERIAL EXAMINED Holotype ©’, Costa Rica: San José Prov: Cerro de la Muerte, San Gerardo de Dota, 2430 m, xii.1981 (Janzen & Hallwachs) (BMNH). Ophion melpomene sp. n. (Figs 28, 31) DescripT1on. Mandibles stout, very weakly narrowed apically, teeth of equal length but with upper tooth very distinctly broader and less acute than the lower tooth, with its blade slightly indented apically; outer mandibular surface punctate, with granulation between punctures, and with basal part of mandible broadly OPHIONINAE OF TROPICAL MESOAMERICA 45 concave; distal segment of maxillary palp slender; malar space 0.1 times as long as basal mandibular width. Head in front view with greatest width across eyes 1.35—1.45 times length of head from vertex to clypeal margin; orbits ventrally slightly convergent; clypeus in profile moderately convex, weakly granulate with isolated punctures, apically slightly coriaceous and with margin strongly impressed. Lower face centrally convex, weakly granulate and sparsely punctate. Head in dorsal view with genae constricted behind eyes; posterior ocellus almost contiguous with eye; occipital carina mediodorsally convex, at centre slightly produced upwards, ventrally curved to join hypostomal carina well above base mandible. Antenna quite long and slender, with 51 flagellar segments; 20th segment 2.0-2.1 times as long as broad. Mesoscutum weakly polished microreticulate, in profile evenly rounded; notauli strongly impressed on anterior 0.3. Mesopleuron weakly polished, centrally finely and quite sparsely punctate, the intervening area microreticulate; lower corner of epicnemial carina bluntly rounded after concavity, and with upper end abruptly curved towards anterior margin of pleuron which it joins just above level of lower corner of pronotum. Scutellum in profile weakly convex, not laterally carinate. Metapleuron moderately convex, microreticulate, very finely punctate. Propodeum in profile evenly declivous; anterior transverse carina more or less complete, posterior transverse carina discernible only laterally as crests; lateromedian longitudinal carinae represented by a single median weak carina; lateral longitudinal carina strong anteriorly, joined to spiracular margin, posteriorly becoming a groove. Fore wing length 17-18 mm; CI = 0.76-0.82; ICI = 0.40-0.44; SDI = 1.23—1.26; cu-a more or less opposite to the base Rs&M; discosubmarginal cell sparsely hirsute, with moderately large glabrous area anteriorly, without a glabrous area along Rs&M; Rs+2r joining pterostigma near centre; lst subdiscal cell evenly hirsute; 1m-cu centrally angled, ramellus shorter than abscissa of 1m-cu between its base and bulla. Hind wing with 8 hamuli on R1, the distal ones more tightly curved than the proximal ones; Ist abscissa of Rs proximally abruptly curved through about 70°; marginal cell proximally short, with junction between Rs and M far proximal to centre, and with distal abscissa of Rs long; proximal 0.3 of marginal cell densely hirsute centrally, peripherally glabrous; distal abscissa of Cul joining cu-a equidistant between 1A and M. Fore leg with tibia slightly flattened, with isolated spines on outer surface. Mid leg with longer tibial spur 1.3—1.4 times length of the shorter. Hind leg with coxa in profile 1.9—2.0 times as long as deep; trochantellus dorsally 0.40.6 times as long as broad; hind tarsus slender, the 4th segment 2.8—3.0 times as long as broad; claws of male long and weakly curved, with pectinae moderately short and close together. Gaster quite slender; tergite 2 in profile about 4 times as long as posteriorly deep; thyridia elliptical and separated from anterior margin of tergite by about its own length. Female unknown. Male with subgenital plate transverse, unspecialized; aedeagus apically simple, rounded. Colour generally uniformly reddish brown with only head predominantly yellowish; interocellar area yellow; flagellum orange, distally infuscate; pterostigma orange; wings more or less hyaline. RemaRKS. This rather unremarkable species is most easily recognized by the finely microreticulate mesothorax and the venational features mentioned in the key. The strongly impressed notauli and the propodeal sculpture are features in which O. melpomene resembles O. polyhymniae and the two species may be related. BIOLOGICAL INFORMATION. In Costa Rica Ophion melpomene has only been collected in forest on the summit of Volcan Pods at an altitude of 2350 m. Nothing is known of the biology of this insect. MATERIAL EXAMINED Holotype ©, Costa Rica: Alajuela Prov: Volcan Pods National Park, 2350 m, xii.1981 (Janzen & Hallwachs) (BMNH) Paratype. 1 0’, same data as holotype (BMNH). Ophion arribai sp. n. (Figs 27, 32) DEscriPTIon. Mandibles stout, very weakly narrowed apically, with upper tooth slightly longer and broader than the lower tooth; outer mandibular surface punctate, weakly polished, with microreticulation between punctures only discernible basally, and with basal part of mandible broadly concave; distal segment of maxillary palp slender; malar space 0.1 times as long as basal mandibular width. Head in front view with greatest width across eyes 1.3-1.5 times length of head from vertex to clypeal margin; orbits ventrally slightly convergent; clypeus in profile moderately convex, with isolated punctures, microreticul- ate and with margin strongly impressed. Lower face centrally convex, weakly polished, granulate with very obscure punctures. Head in dorsal view with genae constricted behind eyes; posterior ocellus contiguous with eye; occipital carina mediodorsally convex, ventrally curved to join hypostomal carina well above 46 IAN D. GAULD base mandible. Antenna quite long and fairly slender, with 53-57 flagellar segments; 20th segment 1.8-1.9 times as long as broad. Mesoscutum weakly polished, granulate, with fine weak close punctures; mesoscutum in profile evenly rounded; notauli strongly impressed near anterior end. Mesopleuron polished, centrally finely punctate, with intervening area microreticulate; lower corner of epicnemial carina bluntly rounded after concavity, and with upper end inclined towards and joining anterior margin of pleuron well above level of lower corner of pronotum. Scutellum in profile quite strongly convex, not laterally carinate. Metapleuron moderately convex, finely punctate and with intervening areas microreticulate. Propodeum in profile rather abruptly declivous, weakly polished; anterior transverse carina strong, more or less complete, posterior transverse carina only represented by lateral vestiges; lateromedian longitudinal carinae ves- tigial, at most discernible as a weak median wrinkle; lateral longitudinal carina weak, usually more or less complete, joined to a rather remote spiracular margin by a raised ridge. Fore wing length 21-23 mm; CI = 0.50-0.55; ICI = 0.73-0.84; SDI = 1.14~1.17; cu-a slightly proximal to the base Rs&M; discosubmarginal cell closely hirsute, with a small glabrous area anteriorly, though the proximal part of the anterior corner is densely hirsute and without a glabrous area along Rs&M; Rs+2r basally incrassate, joining pterostigma close to its base; 1st subdiscal cell quite densely hirsute; 1m-cu centrally angled, ramellus very short, shorter than abscissa of 1m-cu between its base and bulla. Hind wing with 8-10 hamuli on R1, the distal ones moderately curved and unexceptional, the proximal ones almost straight and virtually parallel to R1, 1st abscissa of Rs proximally abruptly curved through about 80°; marginal cell proximally short, with junction between Rs and M far proximal to centre, and with distal abscissa of Rs long; proximal 0.3 of marginal cell centrally hirsute, peripherally glabrous; distal abscissa of Cu1 joining cu-a slightly closer to M than to 1A. Fore leg with tibia slightly flattened, with isolated spines on outer surface. Mid leg with longer tibial spur 1.3-1.4 times length of the shorter. Hind leg with coxa in profile 1.9-2.1 times as long as deep; trochantellus dorsally 0.40.5 times as long as broad; hind tarsus slender, the 4th segment 2.62.8 times as long as broad; claws of female exceptionally long and evenly curved, with long close pectinae; claws of male similar but slightly shorter with pectinae shorter and closer together. Gaster quite slender; tergite 2 in profile about 4 times as long as posteriorly deep; thyridia elliptical and separated from anterior margin of tergite by about 0.5 times its own length. Female with subgenital plate unspecialized; ovipositor moderately long and straight, its sheath slender. Male with subgenital plate transverse, unspecialized; aedeagus apically rounded. Colour generally orange-brown, with orbits, vertex, interocellar area and scutellum whitish; flagellum orange; pterostigma orange, posteriorly black margined; wings quite weakly infumate. Remarks. Ophion arribai may easily be recognized by the shape of the glabrous area in the fore wing, the basally slightly thickened Rs+2r, the characteristic form of the distal hamuli and by the unusually long claws. Superficially it resembles O. uraniae (with which it is sympatric) but O. arribai has the lower corner of the epicnemial carina rounded and lacks the extensive propodeal carination exhibited by O. uraniae. O. arribai resembles O. flavoorbitalis in the position and form of Rs+2r in the fore wing and the two species may be related. BIOLOGICAL INFORMATION. Ophion arribai is only known to occur at Guadalupe Arriba at an altitude of 2000 m in northern Panama. In two years’ sampling only four individual were taken, in April, June, July and September. Nothing is known of the biology of this species. MATERIAL EXAMINED Holotype 2, Panama: Chiriqui, Guadalupe Arriba, 2200 m, iv.1985 (Wolda) (BMNH) Paratypes. 1 0’, 2 9, same locality, vii.1984, ix.1984 & vi.1985 (Wolda) (BMNH, USNM). Ophion thaliae sp. n. (Figs 34, 46, 50) DEscriPTIoNn. Mandibles stout, very weakly narrowed apically, with upper tooth very slightly broader and longer than the lower tooth; outer mandibular surface punctate, coriaceous between punctures, and with basal part of mandible quite strongly concave; distal segment of maxillary palp slender; malar space 0.1 times as long as basal mandibular width. Head in front view with greatest width across eyes 1.3-1.4 times length of head from vertex to clypeal margin; orbits ventrally distinctly convergent; clypeus in profile moderately convex, polished, with isolated punctures and with margin strongly impressed. Lower face centrally convex, polished and sparsely punctate. Head in dorsal view with genae rounded behind eyes; OPHIONINAE OF TROPICAL MESOAMERICA 47 posterior ocellus almost contiguous with eye; occipital carina mediodorsally convex, ventrally curved to join hypostomal carina well above base mandible, though its lower end can be weak. Antenna quite long and stout, with 50-53 flagellar segments; 20th segment 2.0—2.1 times as long as broad. Mesoscutum polished, slightly granulate and indistinctly punctate, in profile abruptly rounded; notauli distinctly impressed near anterior end. Mesopleuron polished, with very obscure minute punctures that are separated by about twice their own diameters; lower corner of epicnemial carina bluntly rounded after concavity, and with upper end abruptly curved to join anterior margin of pleuron above level of lower corner of pronotum. Scutellum in profile evenly convex, not laterally carinate. Metapleuron moderately convex, punctate. Propodeum in profile abruptly rounded; anterior transverse carina more or less com- plete, posterior transverse carina usually only present laterally as crests; lateromedian longitudinal carinae weak but more or less complete; lateral longitudinal carina only present anteriorly, strong and curving back to join spiracular margin. Fore wing length 15-16 mm; CI = 0.51—70; ICI = 0.47-74; SDI = 1.27-1.32; cu-a slightly proximal to the base Rs&M; discosubmarginal cell evenly hirsute, with a large glabrous area anteriorly, without a glabrous area along Rs&M; Rs+2r joining pterostigma slightly proximal to centre; 1st subdiscal cell sparsely hirsute; 1m-cu centrally angled, ramellus from slightly to distinctly shorter than abscissa of 1m-cu between its base and bulla. Hind wing with 5—6 hamuli on R1, the distal ones more tightly curved than the proximal ones; Ist abscissa of Rs proximally evenly curved through about 70°; marginal cell proximally short, with junction between Rs and M far proximal to centre, and with distal abscissa of Rs long; proximal 0.3 of marginal cell broadly glabrous; distal abscissa of Cu1 joining cu-a slightly closer to 1A than to M. Fore leg with tibia weakly flattened, with isolated spines on outer surface. Mid leg with longer tibial spur 1.4-1.6 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally 0.40.5 times as long as broad; hind tarsus slender, the 4th segment 2.6—2.8 times as long as broad; claws of female long and weakly curved, with long moderately close pectinae, claws of male similar but with pectinae shorter and closer. Gaster quite slender; tergite 2 in profile about 4 times as long as posteriorly deep; thyridia oval, separated from anterior margin of tergite by 0.5—-1.0 times its own length. Female with subgenital plate long, unspecialized; ovipositor straight, its sheath narrow. Male with subgenital plate transverse, unspec- ialized; aedeagus apically simple, rounded. Colour generally orange-yellow, with mesoscutal vittae more orange; interocellar area yellow; flagellum orange, distally slightly darker; pterostigma orange; wings more or less hyaline. VARIATION. The posterior transverse carina of the propodeum is usually only discernible laterally, but some individuals have it complete except centrally between the lateromedian longitudinal carinae. RemarkKS. Ophion thaliae is most easily recognized by the combination of the glabrous distal part of the marginal cell of the hind wing, the form of the lateral propodeal carina, its size and coloration, and by the venation of the fore wing. Structurally it is rather unexceptional, and perhaps it most closely resembles O. flavoorbitalis from which it can be distinguished by the characters given in the key. BIOLOGICAL INFORMATION. Ophion thaliae is only known to occur at one moderately high altitude site in Costa Rica where it has been taken in moderately large numbers with O. uraniae. Nothing is known of the biology of this insect. MATERIAL EXAMINED Holotype 9, Costa Rica: San José Prov: Cerro de la Muerte, San Gerardo de Dota, 2430 m, xii.1981 (Janzen & Hallwachs) (BMNH) Paratypes. Costa Rica: San José Prov: 5 0’, 7 9, Cerro de la Muerte, San Gerardo de Dota, 2430 m, xii. 1981 (Janzen & Hallwachs) (BMNH, CNC, MNCR, TC). Ophion flavoorbitalis Cameron (Figs 33, 48, 51) Ophion flavo-orbitalis Cameron, 1886: 294. Lectotype 9, PANAMA (BMNH) designated by Morley (1912: 55) [examined]. Ophion flavoorbitalis Cameron; Hooker, 1912: 49. [Ophion intricatus Brullé; Morley, 1912: 55. Misidentification. ] Description. Mandibles stout, very weakly narrowed apically, with upper tooth of similar length to, but distinctly broader and blunter than the lower tooth; outer mandibular surface punctate, with microreticula- tion between punctures, and with basal part of mandible broadly concave; distal segment of maxillary palp 48 IAN D. GAULD slender; malar space 0.1 times as long as basal mandibular width. Head in front view with greatest width across eyes 1.4 times length of head from vertex to clypeal margin; orbits ventrally slightly convergent; clypeus in profile moderately convex, with isolated punctures, apically slightly coriaceous and with margin strongly impressed. Lower face centrally convex, polished and punctate. Head in dorsal view with genae constricted behind eyes; posterior ocellus contiguous with eye; occipital-carina mediodorsally convex, at centre slightly produced upwards, ventrally curved to join hypostomal carina well above base mandible. Antenna quite long and stout, with 52-53 flagellar segments; 20th segment 1.41.5 times as long as broad. Mesoscutum polished and distinctly punctate with traces of microreticulation between punctures, in profile weakly and evenly rounded, with notauli moderately weakly impressed near anterior end. Meso- pleuron polished, centrally finely and quite sparsely punctate; lower corner of epicnemial carina bluntly rounded after concavity, and with upper end inclined towards anterior margin of pleuron, which it reaches above level of lower corner of pronotum. Scutellum in profile weakly convex, not laterally carinate. Metapleuron moderately convex, obsoletely punctate with distinct microreticulation. Propodeum in profile abruptly rounded; anterior transverse carina strong, more or less complete, posterior transverse carina strong, complete, laterally strongly raised into crests; lateromedian longitudinal carinae strong and more or less complete to the anterior transverse carina, enclosing an elongate area superomedia; lateral longitudinal carina complete, joined to spiracular margin. Fore wing length 18-19 mm; CI = 0.43-0.61; ICI = 0.71-0.73; SDI = 1.16—1.23; cu-a proximal to the base Rs&M by about 0.2 times its own length; discosubmarginal cell evenly hirsute, with a rather small glabrous area anteriorly, and without a glabrous area along Rs&M; Rs+2r strongly thickened basally, joining pterostigma close to base; lst subdiscal cell unusual in being broad proximally and strongly tapered distally, sparsely hirsute; 1/m-cu centrally angled, the ramellus about as long as abscissa of 1m-cu between its base and bulla. Hind wing with 6 hamuli on R1, the distal ones more tightly curved than the proximal ones; Ist abscissa of Rs slightly incrassate, rather irregularly curved so it is slightly wavy, in total turned through about 60°; marginal cell proximally short, with junction between Rs and M far proximal to centre, and with distal abscissa of Rs long; proximal 0.3 of marginal cell glabrous; distal abscissa of Cu1 joining cu-a distinctly closer to M than to 1A. Fore leg with tibia slightly flattened, without obvious spines on outer surface. Mid leg with longer tibial spur 1.4 times length of the shorter. Hind leg with coxa in profile 1.7 times as long as deep; trochantellus dorsally 0.2—0.3 times as long as broad; hind tarsus slender, the 4th segment 2.6-2.7 times as long as broad; claws of female long and weakly curved, with long close pectinae; claws of male similar but with pectinae shorter and closer together. Gaster quite slender; tergite 2 in profile about 4 times as long as posteriorly deep; thyridia large, obovate and separated from anterior margin of tergite by about its own length. Female with subgenital plate unspecialized; ovipositor moderately long and straight, its sheath slender. Male with subgenital plate transverse, unspecialized; aedeagus apically simple, rounded. Colour generally orange-brown with head more yellowish; interocellar area yellow; flagellum brownish orange; pterostigma orange; wings very slightly yellowish. Remarks. Ophion flavoorbitalis is most easily recognized by the combination of distally strongly tapered 1st subdiscal cell in the fore wing, and the slightly wavy Rs and proximally glabrous marginal cell in the hind wing. The base of Rs+2r in the fore wing is slightly stouter than that of most other species though it is similar to that found in O. arribai (see that species). BIOLOGICAL INFORMATION. Ophion flavoorbitalis is only known to occur in Costa Rica and Panama. The paralectotype from Cordoba, Mexico is a different species, as are Morley’s specimens from Brazil (Morley, 1912). O. flavoorbitalis seems to occur at a rather lower altitude than many other species of Ophion as it has only been collected below 1400 m. It is a very uncommon species in collections and nothing is known of its biology. “MATERIAL EXAMINED Lectotype 9, Panama: Volcan de Chiriqui, 850-1300 m (BMNH). Costa Rica: Cartago Prov: 1 ©’, Tapanti, Rio Grande de Orosi, 1300-1400 m, 1.1985 (Janzen & Hallwachs) (BMNH). AGATHOPHIONA Westwood Agathophiona Westwood, 1882: 19. Type-species: Agathophiona fulvicornis Westwood, by monotypy. Diacnosis. Shining black to blue-black species with infumate wings; fore wing length 10-14 mm. Very similar to Ophion except that it has the glossae greatly elongated (Fig. 18), the ocelli are small and far ee OPHIONINAE OF TROPICAL MESOAMERICA 49 removed from the eyes, tergite 3 of the gaster is longer than tergite 2 and the female has a very large subgenital plate REMARKS. This genus contains a single species, Agathophiona fulvicornis, which is endemic to Mexico. It flies in bright sunlight and can often be seen feeding from the flowers of Compositae. No host records are known. The SICOPHION genus-group JANZOPHION Gauld Janzophion Gauld, 1985: 128. Type-species: Janzophion nebosus Gauld, by original designation. DiaGnosis. Predominantly yellowish brown insects with some black mottling; wings hyaline with infumate patch near pterostigma; fore wing length 14-16 mm. Mandibles not or barely twisted; maxillae and labium unspecialized; clypeus apically truncate or slightly convex; occipital carina absent. Notauli-absent; meso- pleural furrow vestigial; posterior transverse carina of mesosternum complete. Fore wing (Fig. 7) with Rs+2r centrally angled, basally broadened; discosubmarginal cell with a glabrous area anteriorly; pterostigma quite stout; lm-cu sinuous or bowed; hind wing with Rs bowed; R1 unusual in having penultimate hamulus greatly lengthened; distal abscissa of Cul basally much closer to 1A than it is to M. Fore tibial spur with membrane behind comb. Gaster slender; second segment with laterotergite folded under; tergite 2 longer than tergite 3. REMARKS. Janzophion is an enigmatic, small Central American genus whose component species, in appearance, resemble members of the Indo-Australian genus Leptophion. However, the two genera are probably not closely related, and the similarity between their species may be the result of evolutionary convergence (Gauld, 1985). Both occupy similar habitats. Janzophion comprises two species which occur in lower montane environments in Central America. The type-species inhabits cloud-forests in Costa Rica and Panama, and a second species, described below, occurs above 2000 m in northern central Mexico. Nothing is known of the host preferences of these species. Key to species of Janzophion 1 Head quite elongate, with lower face 0.65—0.70 times as broad as long; malar space greater than 0.5 times basal mandibular width (Fig. 55); hind wing with Rs weakly bowed (Fig. 57). nebosus Gauld (p. 49) —Head not unusually elongate, with lower face 0.90—0.95 times as broad as long; malar space about 0.3 times basal mandibular width (Fig. 54); hind wing with Rs very strongly bowed (PI O10) oh Bi ee eee ae Si ec cre ert a er a saxis sp. n. (p. 50) Janzophion nebosus Gauld (Figs 7, 55, 57) Janzophion nebosus Gauld, 1985: 129. Holotype co’, COSTA RICA (BMNH) [examined]. Description. Head unusually elongate, with lower face polished, 0.65—0.70 times as broad as long; malar space 0.50—0.55 times basal mandibular width; clypeus weakly convex, its margin truncate; head strongly narrowed behind eyes. Antenna slender, flagellum with 66-68 segments, the 20th segment 2.1—2.6 times as long as broad. Mesoscutum with margin slightly out-turned; scutellum finely shagreened; mesopleuron with upper part highly polished, finely and sparsely punctate, ventrally slightly more coriaceous; epicnemial carina strong; metapleuron finely punctate, weakly coriaceous. Propodeum in profile evenly declivous. Fore wing length 14-18 mm; AI = 0.81-1.25; CI = 0.43-0.47; ICI = 0.61-0.72; SDI = 1.14-1.19; marginal cell uniformly hirsute; 1m-cu sinuous; cu-a proximal to base of Rs&M by 0.3-0.4 times its own length. Hind wing with Rs weakly bowed. Gaster slender; male subgenital plate bearing long fine pubescence; gonosquama quite long, dorsally somewhat membranous. Pale yellowish species, with interocellar area, posterior part of mesoscutum, much of mesopleuron, metapleuron and propodeum blackish; gaster with posterior part of tergite 5 and tergites 6+ infuscate. Wing hyaline, pterostigma blackish and membrane adjacent to it infumate. RemaRKS. Janzophion nebosus is easily recognized by its rather long head, its long malar space and by the weakly curved vein Rs in the hind wing. Structurally it is otherwise very similar to J. saxis. Both species superficially resemble some species of Enicospilus, but they may easily be distinguished by their complete lack of an occipital carina, and by their non-twisted mandibles. 50 IAN D. GAULD Figs 54-57 Janzophion species. 54, 55, head, front view; (54) J. saxis; (55) J. nebosus. 56, 57, distal part of hind wing; (56) J. saxis; (57) J. nebosus. BIOLOGICAL INFORMATION. Janzophion nebosus is only known to occur in Costa Rica and Panama where it is restricted to humid montane cloud forests between 1500 and 2350 m. In Costa Rica only isolated specimens have been collected (December and February), but at Guadalupe Arriba, in northern Panama, two years light-trapping (1984-5) by H. Wolda yielded the following distributional data: Vie BoD Ao Mh ite Jag A a SO" see g ID Pel aS Se to US, ote, ed a The host of this species is not known. MATERIAL EXAMINED Holotype ©’, Costa Rica: Alajuela Prov., Volcan Pods National Park, 2350 m, xii.1982 (Janzen & Hallwachs) (BMNH) Paratypes. Costa Rica: 1 0’, same data as holotype (BMNH); 1 CO’, same locality as holotype, xii.1981 (Janzen & Hallwachs) (BMNH); 1 9, Puntarenas Prov., Monteverde Reserve, 1500 m, ii.1980 (Mason) (TC) Non-type material. Panama: 19 O’, 11 9, Chiriqui Prov., Guadalupe Arriba, 2200 m, 1984-5 (Wolda) (BMNH, CNC, MNCR, USNM). Janzophion saxis sp. n. (Figs 54, 56) DEscriPTION. Head not unusually elongate, with lower face polished, 0.90—0.95 times as broad as long; malar space 0.30-0.33 times basal mandibular width; clypeus strongly convex, apically impressed and with margin slightly concave; head moderately strongly narrowed behind eyes. Antenna slender, flagellum with 57-66 segments, the 20th segment about 2.3—2.8 times as long as broad. Mesoscutum with margin distinctly out-turned; scutellum finely shagreened; mesopleuron with upper part polished, finely coriaceous and sparsely punctate, ventrally slightly more coriaceous; epicnemial carina weak laterally; metapleuron finely punctate, weakly coriaceous. Propodeum in profile rather abruptly declivous. Fore wing length 14-16 mm; AI = 0.75-0.79; CI = 0.48-0.66; ICI = 0.72-0.78; SDI = 1.05—1.11; marginal cell uniformly hirsute; 1-cu bowed; cu-a from subopposite to proximal to the base Rs&M by 0.3-0.4 times its own length. Hind wing with Rs strongly bowed. Gaster slender; male subgenital plate with fine decumbent pubescence. Pale yellowish species, with interocellar area, posterior part of mesoscutum, much of mesopleuron, metapleuron and propodeum blackish; gaster with posterior part of tergite 5 and tergites 6+ slightly infuscate. Wing hyaline, pterostigma blackish and membrane adjacent to it infumate. OPHIONINAE OF TROPICAL MESOAMERICA Si REMARKS. Janzophion saxis is structurally similar to J. nebosus except for the differences outlined in the remarks section above. BIOLOGICAL INFORMATION. This species is only known to occur in Durango State, in northern Central Mexico where it has been collected at altitudes between 2300 and 3000 m. Its host is unknown. MATERIAL EXAMINED Holotype 9, Mexico: Durango, 16 km W. of El Salto, 3000 m, vii.1964 (Mason) (CNC) Paratypes. Mexico: 1 CO’ ,3 9, same locality as holotype, vi-vii.1964 (Mason) (BMNH, CNC); 1 9, Durango, 38 km W. La Cuidad, 2300 m, vi.1964 (Mason) (CNC). SICOPHION Gauld Sicophion Gauld, 1979: 71. Type-species: Sicophion pleuralis Gauld, by original designation. Diacnosis. Black insects with gaster brownish and wings more or less hyaline; fore wing length 16-19 mm. Mandibles twisted about 30° with lower tooth the longer; maxillae slightly lengthened; clypeus subtruncate apically; occipital carina only represented laterally, dorsally and ventrally incomplete. Notauli vestigial; mesopleural furrow transverse, extending from episternal scrobe to near upper end of epicnemial carina; posterior transverse carina of mesosternum present only laterally as vestiges. Fore wing (Fig. 9) with Rs+2r curved; Rs unusual in that it is dipped centrally; discosubmarginal cell anteriorly with an extensive glabrous area that bears traces of sclerites; pterostigma stout; lm-cu curved; hind wing with Rs slightly convex, distal abscissa of Cu1 basally much closer to M than to 1A. Gaster very slender; second segment with laterotergite narrow, membranous and pendant; tergite 2 slightly longer than tergite 3; female unusual in having the ovipositor proximally angled, apically slender and without a subapical notch. REMARKS. The type-species, S. pleuralis, was collected at high altitude in Bolivia. A second, very similar undescribed species occurs at altitudes above 2000 m in Costa Rica and Panama. It is described below. Sicophion fenestralis sp. n. (Figs 9, 13) Description. Mandibles with lower tooth about 1.5 times as long as the upper, apically elongately pointed. Clypeus apically truncate or even slightly concave; head elongate; lower face polished and finely punctate; ocelli very large, the posterior ones contiguous with the eyes. Antenna very long and slender; flagellum with 54-56 segments, the basal segment about 10 times as long as broad. Mesothorax weakly polished, finely and sparsely punctate; scutellum with lateral carina present on anterior 0.3. Propodeum in profile evenly declivous; anterior transverse carina centrally broadly complete, though laterally not reaching the lateral longitudinal carina; posterior transverse carina from absent to present laterally as weak crests and centrally as small tubercles; lateral longitudinal carina complete, not joined to margin of propodeal spiracle. Fore wing length 17-19 mm; AI = 1.04-1.25; CI = 0.57-0.92; ICI = 0.83-1.09; SDI = 1.36-1.47; fenestra very large, extending about 0.30.4 of way along Rs+2r; discosubmarginal cell glabrous in posteroventral corner. Hind wing with 8-9 slender hamuli on R1. Legs slender, otherwise unspecialized. Gaster very long and slender, with tergites 1-6 longer than posteriorly deep. Male with gonosquama apically elongate, produced into a tapered lobe; aedeagus slightly curved, apically slender and unspecialized. A predominantly blackish species with flagellum, legs distal to the trochanters, the posterior half of tergite 2 and tergites 3+ brownish, but with apex of gaster slightly infuscate; wings more or less hyaline, pterostigma yellowish brown. VARIATION. One male from Guadalupe Arriba has the alitrunk more or less entirely brownish. Both Costa Rica specimens lack vestiges of the posterior transverse carina of the propodeum. REMARKS. Sicophion fenestralis differs from the type-species of the genus in having a larger fenestra and a more strongly angulate Rs+2r, slightly more extensive scutellar carinae, an almost complete anterior transverse carina of the propodeum, a slightly longer lower mandibular tooth, and most usually in having the first segment of the gaster entirely black and the femora brownish. S. pleuralis has a smaller fenestra that only extends about 0.1 of the way along Rs+2r, and has this vein less conspicuous bent, and then so nearer its base. S. pleuralis also has very short scutellar carinae, only has a vestige of the anterior transverse carina of the propodeum discernible centrally, has the lower tooth of the mandible only slightly the longer, has the anterior part of tergite 1 yellow, and has blackish femora. 52 IAN D. GAULD BIOLOGICAL INFORMATION. Sicophion fenestralis has been collected in Panama and Costa Rica, where it is only known to occur at sites between 2200 and 2500 m. It occurs sympatrically and synchronously with Janzophion nebosus and some species of Ophion. At Guadalupe Arriba, Panama, S. fenestralis has the following seasonal distribution (based on light-trap catches for 1984-5): y fF MM tA Med i AS © On Ne VD LV {BPEL ie Ge 120 Sied2e S VB AO Deel Nothing is known of the hosts of S. fenestralis. The ovipositor of this species is quite unlike that of any other genus of ophionine, and obviously highly modified for a certain function. What this function is will not be understood until the host of this insect is discovered. MATERIAL EXAMINED Holotype 9, Panama: Chiriqui, Guadalupe Arriba, 2200 m, xii.1984-i.1985 (Wolda) (BMNH). Paratypes. Costa Rica: Alajuela Prov.: 1 o’, Volcan Pods National Park, 2350 m, xii.1981 (Janzen & Hallwachs) (BMNH): San José Prov.: 1 9, San Gerardo de Dota, Cerro de la Muerte, 2430 m, xii.1981 (Janzen & Hallwachs) (BMNH). Panama: 35 ©’, 17 9, Chiriqui, Guadalupe Arriba, 2200 m, 1984-85 (Wolda) (BMNH, CAS, CNC, MCZ, MNCR, PANS, TC, USNM, ZIL). The EREMOTYLUS genus-group EREMOTYLUS Foerster Eremotylus Foerster, 1869: 150. Type-species: Ophion marginatus Gravenhorst (= Anomalon marginatum Jurine), by subsequent monotypy, Thomson, 1888: 1193. Camptoneura Kriechbaumer, 1901a: 23. Type-species: Ophion marginatus Gravenhorst (= Anomalon marginatum Jurine), by subsequent designation, Viereck, 1914: 27. [Junior homonym of Camptoneura Agassiz, 1846.] Genophion Felt, 1904: 123. Type-species: Genophion gilletti Felt (= Ophion costale Cresson), by original designation. Camptoneuroides Strand, 1928: 52. [Replacement name for Camptoneura Kriechbaumer. ] Clistorapha Cushman, 1947: 450. Type-species: Ophion subfuliginosus Ashmead, by original designation. Boethoneura Cushman, 1947: 451. Type-species: Boethoneura arida Cushman, by original designation. Chilophion Cushman, 1947: 454. Type-species: Ophion abnormum Felt, by original designation. Chlorophion Townes, 1971: 55. Type-species: Chlorophion vitripennis Townes, by original designation. Diacnosis. Predominantly orange-brown insects with hyaline wings; fore wing length 9-17 mm. Mandibles not twisted but usually apically narrow; maxillae and labium unspecialized; clypeus rather flat, apically blunt, often slightly concave; occipital carina more or less complete. Notauli anteriorly weakly impressed; mesopleural furrow vestigial; posterior transverse carina of mesosternum present or absent. Fore wing (Fig. 22) with Rs+2r basally abruptly curved and thickened; discosubmarginal cell with a glabrous area anteriorly; pterostigma quite broad; 1m-cu curved to sinuous; hind wing with Rs usually bowed; distal abscissa of Cul more or less equidistant between M and 1A. Fore tibial spur with membranous flange behind comb. Gaster moderately slender to slender; second segment with laterotergite folded under; tergite 2 subequal to or slightly longer than tergite 3. Remarks. Eremotylus is a moderately large genus, which is most species-rich in semi-arid habitats, though a few species are widely distributed throughout the Holarctic realm. The centre of diversity of the genus appears to be the Mediterranean Basin in the Old World, and the south-western United States/northern Mexico in the New World. Eremotylus is represented in Central America by at least four species, E. arida (Cushman), E. subfuliginosus (Ashmead) and two undescribed species. Three of these are only known to occur in the drier parts of north and northern central Mexico, but one distinctive species, which is described below, occurs in Chiapas State in tropical southern Mexico. No hosts are recorded for Central American species, but one European species is known to parasitize a noctuid larva (Seyrig, 1926). Eremotylus tropicus sp. n. (Fig. 22) Description. Mandibles stout, long, weakly tapered and with the upper tooth very slightly the longer; outer mandibular surface with a distinct proximal concavity; malar space 0.2—0.3 times basal mandibular width. Clypeus in front view 2.1—2.3 times as broad as long, apically slightly protuberant and with margin OPHIONINAE OF TROPICAL MESOAMERICA 53 thin; face subquadrate, coarsely and closely punctate centrally but more sparsely and coarsely punctate on clypeus. Head in dorsal view with genae long, weakly rounded behind eyes in females, in male slightly inflated; ocelli large, the posterior ones almost contiguous with the margin of the eye. Occipital carina complete, ventrally joining the hypostomal carina some distance from the base of the mandible. Antenna very long and slender; flagellum with 63-65 segments; central segments 1.8—2.0 times as long as broad. Mesoscutum anteriorly rather irregularly rounded; notauli distinct near anterior scutal margin; scutellum with more or less complete lateral longitudinal carinae which are strongly convergent posteri- orly. Mesopleuron coarsely and rather sparsely punctate; epicnemial carina weak, but present laterally; metapleuron very weakly convex, coarsely and sparsely punctate; posterior transverse carina of mesoster- num only represented by lateral vestiges. Propodeum long, in profile evenly declivous; anterior transverse carina discernible laterally as vestiges, the posterior transverse carina more or less absent; anterior are unusually long (for a species of Eremotylus); posterior part of propodeum rather weakly and irregularly rugose; lateral longitudinal carina complete. Fore wing length 16-17 mm; alar pubescence fine and dense (normal for an ophionine); lm-cu rather unevenly curved, without a trace of a vein stub centrally; cu-a subopposite the base of Rs&M. Hind wing with 7-8 hamuli on R1; first abscissa of Rs strongly curved; marginal cell sparsely hirsute proximally. Legs slender and unspecialized; mid tibia with spurs slender, the longer 1.2—1.3 times the length of the shorter; tarsal claws long and weakly curved, those of the female coarsely and closely pectinate, those of the male more finely and closely pectinae. Gaster long and very slender; tergite 2 in profile more than 5 times as long as posteriorly deep; thyridia elliptical, separated from anterior margin of tergite by about own length; tergite 3 distinctly longer than posteriorly deep. Female with ovipositor unspecialized. Male with terminal sternites with fine unspec- ialized pubescence; apex of gonosquama produced slightly and obliquely truncate; aedeagus apically rounded. An orange-brown species with extreme posterior apex of gaster blackish. Wings very weakly and evenly infumate. REMARKS. This is the only tropical American species of Eremotylus and structurally it is one of the most distinctive species in the genus. The other Mexican species of Eremotylus are smaller and stouter with slender mandibles, and very short anterior propodeal areas. Usually they have short, sparse alar pubes- cence and are more uniformly reddish brown in colour. BIOLOGICAL INFORMATION. Eremotylus tropicus is only known to occur in Chiapas, Mexico where indivi- duals have been collected at an altitude of about 1000 m during June. MATERIAL EXAMINED Holotype 2, Mexico: Chiapas, 32-40 km N. Huixtla, 1000 m, vi.1969 (Peterson) (CNC) Paratypes. Mexico: 1 CO’, same data as holotype (CNC); 2 9, Chiapas, 32 km N. Huixtla, 1000 m, vi.1969 (Mason) (BMNH, CNC). The THYREODON genus-group RHYNCHOPHION Enderlein Rhynchophion Enderlein, 1912: 630. Type-species: Rhynchophion odontandroplax Enderlein, by original designation. Diacnosis. Large, stout, dark-coloured insects with violet or orange wings; fore wing length 18-29 mm. Mandible stout, not twisted; maxillae and labium lengthened, projecting by a distance equal to about half length of head (Fig. 17); clypeus apically slightly pointed; occipital carina more or less complete. Notauli vestigial; mesopleural furrow horizontal; unusual in that epicnemial carina does not extend laterally up mesopleuron; posterior transverse carina of mesosternum present only laterally as vestiges. Fore wing with Rs+2r almost straight; discosubmarginal cell evenly hirsute; pterostigma slender; 1m-cu curved; hind wing with Rs almost straight; distal abscissa of Cul basally closer to M than to 1A. Fore tibial spur without a membranous flange behind the comb. Gaster stout (Fig. 20); second segment with laterotergite folded under; tergite 2 shorter and in profile much smaller than tergite 3. Remarks. Rhynchophion is a small genus that comprises only three nominal species (Townes & Townes, 1966). Their cumulative range extends from the southern United States to southern Brazil. Only one, R. flammipennis (Ashmead), occurs in Mesoamerica. 54 IAN D. GAULD Rhynchophion flammipennis (Ashmead) (Figs 17, 20) Thyreodon flammipennis Ashmead, 1894: 125. Holotype 2, MEXICO: Baja California (CAS) [examined]. Rhynchophion flammipennis (Ashmead) Townes, 1945: 746. Description. Lower tooth of mandible stouter and slightly longer than the upper tooth; lower face about 0.8 times as broad as long, closely and coarsely punctate; ocelli small, the posterior ones separated from eye margins by more than their maximum diameter. Antenna stout, with 55—60 flagellar segments, the central segments transverse. Pronotum mediodorsally flat, unspecialized; mesoscutum, mesopleuron and metapleuron coarsely and closely punctate; metanotum laterally inflated. Propodeum without distinct transverse carinae, rather coarsely coriaceous. Fore wing covered with dense fine pubescence; marginal cell narrow, short; cu-a subopposite to the base of Rs&M; hind wing with 12-14 closely spaced hamuli on R1. Legs stout, fore tibia flattened; fore and mid tarsal claws long, evenly curved and with short close pectinae; hind tarsal claws more abruptly curved and with a longer apical point. Gaster exceptionally stout; hind margin of strongly sclerotized part of sternite 1 (the part fused with the tergite) anterior to the level of the spiracles; tergites 3+, in profile , posteriorly deeper than dorsally long. Female subgenital plate strongly sclerotized, longer than preceding sternite; male subgenital plate very short, with hind margin concave. A black species with antennae from black to bright yellowish; wings (of Costa Rica specimens) yellowish, basally and apically blackish. Remarks. Rhynchophion flammipennis is easily distinguished from all other ophionines in Mesoamerica by its very stout appearance. However, it may not be recognized as an ophionine unless its venation is closely examined BIOLOGICAL INFORMATION. This is a widespread insect whose range extends from southern Arizona to Ecuador. I have only seen it from two localities in Costa Rica, Santa Rosa National Park, and Casa Maritza at an altitude of 700 m on the lower western slopes of Volcan Orosi. In Santa Rosa National Park, R. flammipennis has occasionally been collected in Malaise traps during earlier months of the wet season (June/August). In flight and colour pattern R. flammipennis mimics species of the pompilid genus Pepsis. Rhynchophion species are diurnally active and may be observed feeding on flowers. They are not known to come to light. The hosts are not known, but species of the closely related Old World genus Dictyonotus parasitize sphingid larvae (Gauld, 1985). MATERIAL EXAMINED Holotype 9, Mexico: Baja California, El Taste, 1100 m (CAS). 16 &’, 13 Q, from the following localities: Costa Rica (Guanacaste); Ecuador; Mexico (Baja California, Durango, Guerrero, Oaxaca); Nicaragua; U.S.A. (Arizona, New Mexico) (BMNH, CNC, TC, USNM). THYREODON Brullé Thyreodon Brullé, 1846: 150. Type-species: Thyreodon cyaneus Brulle, by subsequent designation, Hooker, 1912: 107. Athyreodon Ashmead, 1900a: 87. Type-species: Athyreodon thoracicus Ashmead (= Ophion atriventris Cresson), by original designation. Tipulophion Kriechbaumer, 1901b: 75. Type-species: Tipulophion gigas Kriechbaumer (= Ophion atri- ventris Cresson), by monotypy. Macrophion Szépligeti, 1905: 32. Type-species: Macrophion ornatus Szépligeti (= Ophion atriventris Cresson), by subsequent designation, Wiereck, 1912: 640. Oleter Shestakov, 1926: 259. Type-species: Oleter selenaction Shestakov (= Thyreodon laticinctus Cres- son), by original designation. Diacnosis. Generally large black, orange and black or yellow and black species with infumate or mottled wings; fore wing length 16-28 mm. Mandible stout, not twisted; maxillae and labium unspecialized; clypeus apically slightly pointed; occipital carina mediodorsally complete. Pronotum unusual in having the anterior margin mediodorsally curved up, closely co-adapted with occiput, and often with a transverse keel behind this flange; notauli weak to very strong and long; mesopleural furrow weak, but if discernible then extending from episternal scrobe to subalar prominence; posterior transverse carina represented by lateral OPHIONINAE OF TROPICAL MESOAMERICA 55 and central vestiges; propodeum unusual in being strongly inflated and in side view dwarfing the meta- pleuron. Fore wing with Rs+2r curved basally, otherwise straight; discosubmarginal cell with or without a small glabrous area anteriorly; pterostigma very slender; 1m-cu bowed; hind wing with Rs straight or weakly bowed; distal abscissa of Cu1 basally either very much closer to M than to 1A or actually arising from M. Fore tibial spur without a membranous flange behind the comb. Gaster from slender to moderately stout; second segment with laterotergite pendant; tergite 2 longer than tergite 3. Remarks. Thyreodon is an American genus that contains about 40 large and conspicuous species. The majority are restricted to tropical America, but approximately five species occur in the southern part of the United States and one species, T. atricolor (Olivier), occurs as far north as southern Canada (Carlson, 1979). Thyreodon species are generally believed to be parasitoids of sphingids (Townes, 1971), but very few species have actually ever been reared, and one that has attacks saturniids Unlike most other ophionines many Thyreodon species are, at least to some extent, diurnally active and can be encountered during the day flying in forest clearings or feeding from flowers. Like other diurnal ichneumonids (Gauld, 1987), Thyreodon species are probably exposed to a variety of predators, par- ticularly asilids which may be very common in their forest habitats, and therefore it is not surprising to find that they have a defence against these predacious dipterans. Their pronotum is modified dorsally, furnished with keels and curved forward to reach the concave occiput, thus providing the cervical region (which is the part asilids try to penetrate) with a substantial and effective shield Although many Thyreodon species are diurnally active insects, and like other diurnal ichneumonids have small ocelli and eyes, some apparently are predominantly nocturnal and have enlarged ocelli and eyes. Some authors (Cushman, 1947; Porter, 1984) have placed the nocturnal species in a separate genus, Athyreodon. However, the diurnal and nocturnal species share a set of autapomorphic features that characterize them as a monophyletic lineage of Ophioninae (see Gauld, 1985) and I concur with Townes (1971) and suggest that the differences between the two groups are best expressed by according them the status of species-groups within a single genus, Thyreodon. This is concomitant with the general classifica- tion of the subfamily as a whole, as several other ophionine genera (e.g. Ophion, Enicospilus) include morphologically rather aberrant species which are diurnally active. Both species-groups are represented in Central America. The Thyreodon atriventris species-group (= Athyreodon) is characterized by having very large ocelli which touch or almost touch the margin of the eye, so that at most the orbital ocellar distance is less than 0.5 times as long as the maximum ocellar diameter. This species-group includes five described Mesoamerican taxa, T. atriventris (Cresson), which is widespread from tropical Mexico south to Panama and thence throughout much of tropical South America, T. flammiger Morley from Jamaica, T. fulvescens Cresson, which is endemic to Cuba, T. maculipennis Cresson, whose range extends from Mexico south to Panama, and T. rivinae Porter which is known from Costa Rica and the southern U.S.A. I have seen one additional and apparently undescribed species that occurs in Costa Rican forests. The Thyreodon laticinctus species-group (= Thyreodon s.s.) is characterized by having small ocelli, with the orbital ocellar distance equal to or greater than the maximum ocellar diameter. This species-group has been subdivided by Cushman (1947) and Porter (1984) into two lineages, the laticinctus subgroup and the atricolor subgroup. The Mesoamerican species of the former subgroup have recently been revised by Porter (1984) who recognized 10 species as occurring in the area: T. apricus Porter, whose range extends from Texas south to Costa Rica; T. elegans Cresson [Cuba]; T. erythrocera Cameron [Arizona, Mexico, Costa Rica]; T. ferrugineus Hooker [Mexico]; ; T. grandis Cresson [Cuba, Jamaica]; T. laticinctus Cresson [Belize, Bolivia, Colombia, Costa Rica, Guatemala, Honduras, Mexico, Nicaragua, Panama, Peru, Trinidad]; T. morosus Smith [Brazil, Costa Rica, Mexico, Panama, Peru]; 7. niger Cresson [Guatemala, Mexico]; T. robur Porter [Mexico]; T. ultor Porter [Bahamas, Cuba]. The atricolor subgroup is repre- sented in Mesoamerica by five described species, Thyreodon affinis Cresson [Cuba]; T. bicolor Morley [Central America?]; T. fernaldi Hooker [Mexico, southern U.S.A.]; T. ornatipennis Cresson [Mexico, southern U.S.A.]; T. santarosae Porter [Costa Rica] (Townes & Townes, 1966; Porter, 1986). Only the species of Thyreodon that occur in Costa Rica have been studied by me, and a synopsis of them is presented below. The Costa Rican species of Thyreodon Fairly extensive collecting has revealed the existence of nine species of Thyreodon in Costa Rica, though others may yet be discovered, especially in the wet forests on the eastern side of the country. The key given below should therefore be used with caution and, as far as possible, identifications should be verified by comparison with authenticated specimens. The keys and descriptions presented by Porter (1980; 1984; 56 IAN D. GAULD 1986) are also very informative. However, the simplified key given here will permit the most frequently encountered Costa Rica species to be recognized and some further critical features that will enable recognition are referred to under the species headings below. 1 Ocellivery large, touching or almost touching the margin of the eye, at most with orbital-ocellar distance 0.5 times as long as maximum ocellar diameter (Figs 63-64) ................... 2 — Ocelli small, separated from the margin of the eye by a distance that is at least equal to Maximunlocellarmdiameten (Bic t62)/he2 eee) oe ey eek ele city tel okie a2 eee ee 6 2 Alitrunk orange-brown; fore wings hyaline, with large blackish areas in basal and marginal cells. Notauli broad and weakly impressed (Fig. 65). ........... atriventris (Cresson) (most) (p. 56) — Alitrunk black or very dark brown; fore wings entirely black, or black with a central hyaline Circular ane a) os... ma erie e Sete Ms ee oA a ee ceteTn & cae ea, Se eto, Sys 3 3. Notauli broad and weakly impressed on anterior part of mesopleuron (Fig. 65); propodeum with a very strong tubercle just behind upper end of spiracle. atriventris (Cresson) (dark morph) (p. 56) — Notauli rather narrow but strongly impressed (Fig. 66); propodeum with a weak or indistinct crestor tuberclejustibehindyupper end of spiracles een eee ee i) ci ee 4 4 Posterior ocelli separated from eye margin, the orbital- ocellar distance about 0.3 times maxi- mum ocellar diameter (Fig. 64); antenna blackish and fore wing with a central hyaline area maculipennis Cresson (p. 59) — Posterior ocelli contiguous or almost contiguous with the eye margin (Fig. 63); either with antenna bright yellow, or with fore wing entirely black’... 0.2.0.0... .2.05. es ee semee 5 5 Antenna bright yellow; fore wing with a central hyaline area; hind femur with distal 0.5 stout and slightlyicompnesse ci (i100) lene tiie etree rivinae Porter (p. 60) — Antenna black; fore wing uniformly blackish; hind femur very slender, cylindrical throughout (Creo) een eer eT ee actrees aay c neem Toe Boge eee mene species 1 (p. 60) 6 Propodeum posterodorsally smooth and highly polished; wings generally centrally somewhat hyaline, basally and apically strongly infumate .................... santarosae Porter (p. 61) — Propodeum posterodorsally coarsely reticulate, rugose or coriaceous, variously polished; wings uniformly black :..2 0 F253 Ai atok Pelee mae ote e's ee beets a slats on chee ete one nee 7 7 Propodeum posterodorsally with a very strongly impressed median longitudinal furrow that is laterally bordered by broad shallow furrows posteriorly (Fig. 67); anterior end of notaulus withkaltransversemalsedicnesti(s10).109) er eisai emia eee ae apricus Porter (p. 61) — Propodeum posterodorsally with a broad, shallow median longitudinal furrow, laterally without secondary furrows (Fig. 68); anterior end of notaulus either without a crest, or if a crest is discernible then it is weak and parallel to notaulus (Fig. 70) ........................-4. 8 8 Occipital carina ventrally joining the hypostomal carina (Fig. 58); posterior part of propodeum COFLACCOUSHIN Es es SE Se ee ee ee erythrocera Cameron (p. 62) — Occipital carina ventrally incomplete, not reaching the hypostomal carina (Fig. 59); posterior part of propodeum generally with strong oblique wrinkles..........................0-. 9 9 Gaster black with segments 3—4 predominantly bright yellowish or orange; flagellum entirely bl aCe i..8 ccs. r nce Se acct aeico acaa: PaP eis ace” ie Seer laticinctus Cresson (p. 62) — Gaster entirely black; flagellum with proximal 0.6 white or whitish yellow, only distal apex BLACK ES Dissident te fe sou en demn SUR owe aero aaa at RS ICR y eae ice eras ko eT oF morosus Smith (p. 63) Thyreodon atriventris (Cresson) (Figs 16, 65) Ophion atriventris Cresson, 1874: 374. Holotype 2, MEXICO (PANS) [examined]. Thyreodon rufothorax Cameron, 1886: pl. 12. Holotype 9, PANAMA (BMNH) [examined]. [Syn- onymized by Hooker, 1912: 102.] Thyreodon rufithorax Cameron; Cameron, 1886: 290. [Mis-spelling. | Athyreodon thoracicus» Ashmead, 1900a: 87. Holotype 2, ECUADOR (USNM) [examined]. [Syn- onymized by Hooker, 1912: 102.] Thyreodon grenadensis Ashmead, 1900b: 270. Holotype 2, GRENADA (BMNH) [examined]. [Syn- onymized by Townes & Townes, 1966: 186.] OPHIONINAE OF TROPICAL MESOAMERICA 57 Figs 58-64 Thyreodon species. 58, 59, head, posteroventral view; (58) T. erythrocera; (59) T. morosus. 60, 61, hind femur; (60) Thyreodon sp. 1; (61) T. rivinae. 62-64, head, dorsal view; (62) T. laticinctus; (63) T. rivinae; (64) T. maculipennis. Tipulophion rufithorax (Cameron) Schulz, 1903: 249. [Mis-spelling. ] Macrophion ornatus Szépligeti, 1905: 33. Holotype &’, BELIZE (TM). [Synonymized by Morley, 1912: 15.] Athyreodon atriventris (Cresson) Hooker, 1912: 102. -[Macrophion fulvescens (Cresson) Morley, 1912: 14. Misidentification. ] Macrophion grenadensis (Ashmead) Morley, 1912: 15. Thyreodon atriventris (Cresson) Townes & Townes, 1966: 186. Diacnosis. Head, alitrunk and most of anterior two pairs of legs usually orange-brown; interocellar area, flagellum, gaster and most of hind legs black; wings hyaline, fore wing with basal and marginal cells partly infumate. Ocelli very large, the hind ones contiguous with the eyes; genae strongly constricted behind eyes; occipital carina ventrally incomplete, not turned towards hypostomal carina. Mesoscutum with notauli broad but weakly impressed, without an anterior crest (Fig. 65); mesopleuron finely and usually closely punctate; propodeum usually extensively coarsely reticulate, with prominent median and lateral tubercles, and posterodorsally with an indistinct median longitudinal furrow. Fore wing length 21-29 mm. Hind femur moderately slender, cylindrical. Gaster stout, tergite 2 less than 2 times as long as posteriorly deep; male with gonosquama apically subtruncate, with an elongate spine-like projection dorsally. VARIATION. Large numbers of individuals agree closely with the above description, but I have seen a number of aberrant individuals in Costa Rica which may represent distinct species. A female from Braulio Carrillo National Park is entirely black and has the wings strongly infumate except for a central hyaline band. A male from Corcovado National Park resembles typical specimens except that the gonosquamae are evenly but elongately tapered distally, and lack the spine-like process. A few individuals from Santa Rosa National Park are exceptionally small (fore wing length 17-18 mm), have the propodeum centrally striate rather than reticulate, lack a median propodeal tubercle and have a short, upcurved process on the distal apex of the gonosquama. The status of these morphs requires further investigation. REMARKS. Thyreodon atriventris is most easily recognized by its colour pattern, large size and the large ocelli. The dark morph may be separated from other species in the atriventris species-group by its weakly impressed notauli, robust gaster and large size. BIOLOGICAL INFORMATION. Thyreodon atriventris is one of the most common and conspicuous species of the genus in lowland and lower montane forest sites in Costa Rica and Panama. Occasional individuals may be encountered during the day, and specimens have been observed on the flowers of Forsteronia spicata (Apocynaceae) in Santa Rosa National Park, Costa Rica. However, T. atriventris appears principally to be 58 IAN D. GAULD —, 3 E overs~e : SE8N723 Figs 65-70 Thyreodon species. 65,66, mesoscutum, dorsal view; (65) T. atriventris; (66) T. maculipennis. 67-68, propodeum, dorsal view; (67) T. apricus; (68) T. laticinctus. 69, 70, mesoscutum, dorsal view; (69) T. apricus; (70) T. laticinctus. a nocturnally active species and specimens are frequently attracted to light. In Santa Rosa National Park it is common throughout most of the wet season (late May-December) and pooled seasonal distributional data from 1981-6 are: JimtFet Me Avs Mindinel eck iSaiOe Neb 104 15): 421) Se tases OPHIONINAE OF TROPICAL MESOAMERICA 59 On Barro Colorado Island, Panama, it seems to be less seasonal as individuals have been taken in light- traps by Dr H. Wolda most months of the year. The cumulative seasonal distributional data for 1983-S are: ar ww A MJ, J AS OF N B ee ee ee het T. atriventris has been reared by Dr D. H. Janzen from several species of sphingid caterpillars in Santa Rosa National Park, but most commonly from Pachylia ficus (Linnaeus) whose larvae feed on a variety of arborescent Moraceae (Janzen, 1984a). MaTERIAL EXAMINED Holotype 2 (Ophion atriventris Cresson), Mexico: Orizaba (PANS). Holotype 2 (Thyreodon rufothorax Cameron), Panama: Bugaba, 300-500 m (BMNH). Holotype 2 (Athyreodon thoracicus Ashmead), Ecuador (USNM). Holotype 9 (Thyreodon grenadensis Ashmead), Grenada: Baltasar, on windward side (BMNH). Costa Rica: Guanacaste Prov.: 1 9, 5 km NE. of Quebrada Grande, 600 m, vi.1986 (Gauld) (BMNH); 1 CO, Rincon de la Vieja National Park, Mirador, 900 m, iii.1984 (Gauld) (BMNH); 1 0’, 4 km W. Santa Cecilia, 300 m, iv.1983 (Janzen & Hallwachs) (BMNH); 17 0’, 25 9, Santa Rosa National Park, 300 m, _ dates as above (Janzen, Hallwachs & Gauld) (BMNH; MNCR); 3 Go’, 3 2, Volcan Orosi, Casa Mariksa _ [=Maritza], 800 m, vi-vii.1986 (Gauld) (BMNH): Heredia Prov.: 1 2, Finca La Selva, 3 km S. Puerto Veijo, vii. 1982 (Hespenheide) (BMNH): Lim6n Prov.: 1 0’, Tortuguero National Park, Cerro Tortuguero, 0-100 m, v.1984 (Janzen & Hallwachs) (BMNH): Puntarenas Prov.: 4 0’, 5 2, Corcovado National Park, Osa Peninsula, i-v. 1981-7 (Janzen & Hallwachs) (BMNH); 1 0’, 2 2, same locality, 20 m, v.1984 (Gauld) (BMNH); 1c’, 4 @, Fila Esquinas, 35 km S. Palmar Norte, 150 m, i.1983 (Janzen & Hallwachs) (BMNH): San José Prov.: 1 o’, 1 9, Braulio Carrillo National Park, Estacion Carrillo, 700 m, vii.1984, v.1985 (Chacon) (BMNH). Panama: 1 ©’, Barro Colorado Island, ix.1967 (Sexton) (UM); 25 OC’, 28 9, Barro Colorado Island, 120 m, dates as above (Wolda) (BMNH). Other specimens. Costa Rica: Guanacaste Prov.: 1 0’, 1 9, [small morph], Santa Rosa National Park, 300 m, v.1983 (Janzen & Hallwachs) (BMNH): Puntarenas Prov.: 1 0’, [pointed gonosquama morph], Corcovado National Park, iii.1978 (Janzen) (BMNH); San José Prov.: 1 9, [dark morph], La Montura, Braulio Carrillo National Park, 1100 m, xii.1981 (Janzen & Hallwachs) (BMNH). Thyreodon maculipennis Cresson (Figs 64, 66) Thyreodon maculipennis Cresson, 1874: 375. Lectotype 2, MEXICO, designated by Townes & Townes (1966: 189) (PANS) [examined]. Diacnosis. A uniformly black or blackish species; wings blackish, fore wing with a central hyaline circular area. Ocelli moderately large, the hind ones separated from the eyes by about 0.3 times their maximum diameter; genae rounded behind eyes; occipital carina ventrally incomplete, not turned towards hyposto- mal carina. Mesoscutum with notauli narrow but strongly impressed (Fig. 66), almost meeting each other before the scuto-scutellar groove, without an anterior crest; mesopleuron smooth, with isolated fine punctures; propodeum extensively irregularly reticulately wrinkled, with indistinct median and lateral tubercles, and posterodorsally with a broad shallow median longitudinal furrow. Fore wing length 18-20 mm. Hind femur moderately slender, distally weakly compressed. Gaster moderately stout, tergite 2 about twice as long as posteriorly deep; male with gonosquama apically elongately tapered, slightly upcurved. REMARKS. Thyreodon maculipennis can most easily be recognized by its colour pattern and its moderately large ocelli. It is most likely to be confused with T. rivinae and species 1, but can readily be distinguished by the characteristics given in the key. BIOLOGICAL INFORMATION. Thyreodon maculipennis seems to be associated with seasonally dry forests. In Costa Rica it has only been collected in or near to Santa Rosa National Park, frequently at light, although individuals may sometimes be found feeding on flowers during the day. T. maculipennis has been collected throughout the wet season (late May to December) though it seems to be most common in June and November, suggesting it may have two generations a year. The cumulative totals for its seasonal distribu- tion in 1983-6 are: Jala bAcoMeals eimvAcvS -O.°N .D Sea eS 20! Fern Dew aye ero 60 IAN D. GAULD Specimens have been reared by Dr D. H. Janzen from the larvae of the medium small sphingids Perigonia lusca (Fabricius) and Xylophanes turbata (Edwards) which feed on some arborescent Rubiaceae (Janzen, 1984a). MATERIAL EXAMINED Lectotype 9, Mexico: Orizaba (PANS); paralectotype 0’, Mexico: Cordoba (PANS). Costa Rica: Guanacaste Prov.: 1 9,2km E. Cuajiniquil, 300 m, vi.1986 (Gauld) (BMNH); 20 co’, 21 9, Santa Rosa National Park, 300 m, vi-vili, x-x1i. 1983-6 (Janzen, Hallwachs & Gauld) (BMNH, MNCR). Thyreodon rivinae Porter (Figs 19, 61, 63) Thyreodon rivinae Porter, 1980: 243. Holotype 9, U.S.A.: Texas (FSCA). Diacnosis. An entirely black or blackish species except for the antenna which is bright yellow; wings blackish, fore wing with a central hyaline circular area. Ocelli very large, the hind ones contiguous with the eyes; genae rounded behind eyes; occipital carina ventrally incomplete, not turned towards hypostomal carina. Mesoscutum with notauli strongly impressed, almost meeting before the scuto-scutellar groove, without a crest anteriorly; mesopleuron smooth with fine, very sparse punctures; propodeum extensively rather finely reticulate, with indistinct median and lateral tubercles, posterodorsally with a broad shallow median longitudinal furrow. Fore wing length 17-19 mm. Hind femur moderately stout, the distal 0.5 laterally compressed. Gaster moderately stout, tergite 2 about twice as long as posteriorly deep. REMARKS. Thyreodon rivinae is, to a large extent, sympatric with and resembles 7. maculipennis, but may be distinguished by its bright yellow antennae. Furthermore, the posterior ocelli of T. rivinae are contiguous with the eye margin whereas those of T. maculipennis are separated from the eye margin by about 0.3 times the ocellar diameter. BIOLOGICAL INFORMATION. Thyreodon rivinae is known to occur in Hidalgo county in the extreme south of Texas, and in north-western Costa Rica, so it is probably widely distributed throughout Central America. The single specimen from Texas was collected in July (a hot and dry period) in shady Celtis woodland flying over a patch of the herb Rivina humilis (Phytolaccaceae) (Porter, 1980). In Costa Rica T. rivinae appears to be associated with seasonally dry forests and it has only been collected in Santa Rosa National Park. [It is perhaps noteworthy that Rivina humilis also occurs in Santa Rosa (Janzen & Liesner, 1980).] Individuals are attracted to light, but it is rather less commonly encountered than 7. maculipennis. All the individuals of T. rivinae that I have seen were collected in June or early July. MATERIAL EXAMINED Costa Rica: Guanacaste Prov.: 1 2, Santa Rosa National Park, 300 m, vi.1980 (Janzen & Hallwachs) (BMNH); same locality, 1 2, vi.1984 (Janzen & Hallwachs) (BMNH); same locality, 2 2, vi.1985, 1 9, vii. 1986 (Gauld) (BMNH, MNCR). Thyreodon species 1 (Fig. 60) Diacnosis. A highly polished jet-black species; wings uniformly blackish. Ocelli quite large, the hind ones of the female more or less contiguous with the eyes, those of the male separated from the eye margin by about 0.1 of. the maximum ocellar diameter; genae constricted behind eyes; occipital carina ventrally incomplete, not turned towards hypostomal carina. Mesoscutum with notauli very strongly impressed, virtually meeting before scuto-scutellar groove, without an anterior crest; mesopleuron smooth with very sparse, fine punctures; propodeum extensively reticulate, centrally tending to irregularly longitudinally striate, deplanate, with a vestigial median tubercle, but with lateral tubercles well developed, and posterodorsally with a very weak median longitudinal furrow. Fore wing length 19-23 mm. Hind femur slender, cylindrical. Gaster quite slender, tergite 2 about 2.3 times as long as posteriorly deep; male with gonosquama apically elongately tapered, slightly upcurved. RemaRKS. I have compared this species with all available types of Mesoamerican species and I believe it is undescribed. However, species in this group exhibit a wide range of variation, and I believe it is best to refrain from describing this species until such time as a revision of all the Mesoamerican species of the atriventris species-group is undertaken. Species 1 may be separated from other taxa in this species-group by OPHIONINAE OF TROPICAL MESOAMERICA 61 the combination of the entirely black wings, the strongly impressed notauli, the well-developed lateral propodeal crests and its slender hind femora. It has a more slender gaster and is rather more polished than either T. rivinae or T. maculipennis. BIOLOGICAL INFORMATION. Thyreodon species 1 is only known to occur in Costa Rica in moist forests at altitudes between 600 and 800 m. It is thus allopatric with the rather similar species, T. rivinae and T. maculipennis. All examined specimens of species 1 were collected at light. Its host is unknown. MATERIAL EXAMINED Costa Rica: Alajuela Prov.: 1 &’, 2 2, San Ram6n Forest Reserve, Rio San Lorencito, 800 m, xi-xii.1986 (Chacon) (BMNH); 1 CO’, Finca Campana, 5 km NW. Dos Rios, 750 m, iii.1985 (Janzen & Hallwachs) (BMNH); 1 9, Finca San Gabriel, 16 km ENE. Quebrada Grande, 680 m, iii.1983 (Janzen & Hallwachs) (BMNH). Thyreodon santarosae Porter Thyreodon santarosae Porter, 1986: 133. Holotype o&', COSTA RICA (USNM). Diacnosis. A generally black or brownish black species, with antenna from blackish to yellowish; gastral tergites 2 to 3 and legs usually paler brownish-marked; wings from hyaline with bases and apices infumate, to extensively infumate basally and apically with only a central hyaline area. Ocelli small, the hind ones separated from the eyes by more than their maximum diameter; genae rounded or slightly expanded behind eyes; occipital carina ventrally incomplete, turned towards hypostomal carina. Mesoscutum with notauli broad and shallow, barely convergent posteriorly, impressed about 0.8 of mesoscutum and without an anterior crest; mesopleuron centrally quite closely punctate; propodeum extensively smooth and polished, with indistinct median and lateral tubercles. Fore wing length 15-19 mm. Hind femur moderately stout. Gaster stout, tergite 2 less than twice as long as posteriorly deep; male with gonosquama apically acute, slightly up-curved. REMARKS. Thyreodon santarosae is most easily distinguished from other species by the smooth and polished posterior face of the propodeum. Porter (1986) gives a detailed description of this species and describes its variation. BIOLOGICAL INFORMATION. Thyreodon santarosae is the only species of the atricolor subgroup that is known to occur in Costa Rica, and has only been collected in Santa Rosa National Park, in the seasonally dry north-eastern part of Guanacaste Province. This species has never been collected in Malaise traps or at light. All known specimens have been reared from hosts collected 3-20 m above the ground, so it is likely that this species only flies high in the canopy. Specimens have been reared by D. H. Janzen from the larvae of the ceratocampine saturniids Syssphinx molina (Cramer), Ptiloscola dargei Lemaire and Othorene purpurascens (Schaus) which feed in the crowns of trees of the families Leguminosae and Sapotaceae (Janzen, 1982; 1984a). The parasitoid adults emerged between late April and December, so the species is probably on the wing throughout the wet season. MATERIAL EXAMINED Costa Rica: Guanacaste Prov.: 1 0’, 1 9, Santa Rosa National Park, 300 m, 1984 (Janzen) (BMNH). Thyreodon apricus Porter (Figs 67, 69) Thyreodon apricus Porter, 1984: 57. Holotype &’, MEXICO (FSCA). Diacnosis. A jet-black species with distal 0.8 of flagellum blackish brown; wings entirely blackish. Ocelli small, the hind ones separated from the eyes by more than the maximum ocellar diameter; genae weakly narrowed behind eyes, those of male slightly inflated; occipital carina ventrally incomplete, its end slightly turned towards hypostomal carina. Mesoscutum with notauli very strongly impressed, rugose, anteriorly with a distinct transverse crest (Fig. 69); mesopleuron centrally smooth, with isolated fine punctures; propodeum finely rugose reticulate posteriorly, without distinct median and lateral tubercles, posterodor- sally with a deep median longitudinal furrow that is bordered posteriorly by broad shallow secondary furrows (Fig. 67). Fore wing length 16-19 mm. Hind femur moderately stout, distally slightly compressed. Male with segments 1—4 of hind tarsus ventrally bearing a row of long stout erect hairs. Gaster quite slender, tergite 2 2.2-2.7 times as long as posteriorly deep; male with gonosquama apico-dorsal angle produced into a moderately long acute process. 62 IAN D. GAULD Remarks. Thyreodon apricus is most easily recognized by the presence of the secondary dorsolateral furrows on the posterior face of the propodeum, and by the possession of strong, transverse crests anteriorly across the notauli. Porter (1984) remarked that this species appears to be closely related to T. ferrugineus Hooker, a Mexican species. BIOLOGICAL INFORMATION. Thyreodon apricus is a fairly widely distributed species whose range extends from Hidalgo County in the extreme south of Texas, U.S.A., south through Mexico and Guatemala to northern Costa Rica. Porter (1984) remarked that this species occurs in hot, low-altitude, semiarid habitats in the United States and Mexico. In Costa Rica this species is only known to occur in Santa Rosa National Park, where it has only been collected in very open regenerating woodland in early June. It has never been collected at light and its hosts in Santa Rosa are unknown. MATERIAL EXAMINED Costa Rica: 1 &’, 1 9, Guanacaste Prov., Santa Rosa National Park, 300 m, in Malaise trap in open woodland west of administration area, vi.1985 (Janzen & Gauld) (BMNH). Thyreodon erythrocera Cameron (Fig. 58) Thyreodon erythrocera Cameron, 1886: 288. Holotype 2, MEXICO (BMNH) [examined]. Thyreodon erythrocerus Cameron; Dalla Torre, 1901: 185. [Unjustified emendation. ] Diacnosis. A shining black species with legs brownish black; antenna of Costa Rican specimen dark brown, of other material orange-yellow; wings black with metallic purplish reflections. Ocelli small, the hind ones separated from the eyes by more than their own diameter; genae evenly constricted behind eyes; occipital carina ventrally complete, evenly convergent with and joining hypostomal carina far above base of mandible. Mesoscutum with notauli weakly impressed, broad (especially towards the anterior end) and reticulate with a low rugose crest anteriorly; mesopleuron smooth, with very sparse, minute punctures; propodeum extensively coriaceous posteriorly, without median and lateral tubercles, posterodorsally with a shallow narrow median longitudinal furrow. Fore wing length 18-23 mm. Hind femur stout, distally strongly laterally compressed. Gaster quite stout, tergite 2 less than twice as long as posteriorly deep; male with gonosquama apically obliquely truncate, upper angle acute. VARIATION. Porter (1984) characterizes this species by the ventrally complete occipital carina, a feature also possessed by the Costa Rican specimen, but the occipital carina of the holotype does not reach the hypostomal carina. The possibility thus exists that erythrocera (as represented by the holotype) is a different species, but more extensive material will need to be collected and examined before this matter can be resolved. The Costa Rican specimen differs from other individuals in having dark brown rather than orange-yellow antennae, and in having rather finer propodeal sculpture. RemaRkKs. Although I am not absolutely certain that the Costa Rican specimens are conspecific with either the holotype or the material determined by Porter as T. erythrocera, they are very similar and best treated under this species name until such time as more material is collected for study. The ventrally complete occipital carina distinguishes this species from all other Thyreodon species in Costa Rica. BIOLOGICAL INFORMATION. Two individuals of this species have been collected in Santa Rosa National Park during the first half of the wet season. It is otherwise only known from southern Arizona, in the south- western United States, and Mexico, where it apparently flies in deserts and thorn scrubs after late season rains (Porter, 1984). MATERIAL EXAMINED Holotype 2, Mexico: Yucatan, Valladolid (BMNH). Costa Rica: Guanacaste Prov.: 2 2, Santa Rosa National Park, 300 m, vi-vii.1984 (Janzen & Hallwachs) (BMNH). Mexico: Nuevo Leon: 1 9, Pedro Iturbide, 32 km W. Linares, x.1962 (Townes) (TC): Sonora: 1 oO, 1 9, Cananca, viii.1974 (Erickson) (TC); 1 9, ‘N. Sonora’ (Morrison) (BMNH). Thyreodon laticinctus Cresson (Figs 62, 68, 70) Thyreodon laticinctus Cresson, 1874: 376. Holotype 9, MEXICO (PANS) [examined]. Thyreodon principalis Smith, 1879: 230. Holotype 2, COSTA RICA (BMNH) [examined]. [Synonymized by Morley, 1912: 9.] OPHIONINAE OF TROPICAL MESOAMERICA 63 Thyreodon zonatus Szépligeti, 1906: 134. Lectotype o’, BOLIVIA (TM), designated by Townes & Townes (1966: 189). [Synonymized by Townes & Townes, 1966: 189.] Oleter selenaction Shestakov, 1926: 259. Lectotype 2 ,‘NOVA GRENADA’ (ZIL), designated by Townes & Townes (1966: 189). [Synonymized by Cushman, 1947: 427.] Diacnosis. A jet-black species with segments 3 and 4 of gaster predominantly bright yellow; wings uniformly blackish. Ocelli very small, the hind ones separated from the eyes by far more than the maximum ocellar diameter; genae evenly constricted behind eyes; occipital carina ventrally incomplete, its end turned towards hypostomal carina. Mesoscutum with notauli very broad and weakly impressed, rugose, with a weakly developed crest margining the mesal side anteriorly (Fig. 70); mesopleuron centrally with close, shallow punctures; propodeum extensively diagonally striate posteriorly, without a median tubercle, but with weak elongate lateral tubercles, posterodorsally with broad, quite deep median longitudinal furrrow (Fig. 68). Fore wing length 18-22 mm. Hind femur slender, cylindrical. Gaster quite slender, tergite 2 more than twice as long as posteriorly deep; male with gonosquama with upper corner apically produced into an elongate spine-like process. RemarKS. Thyreodon laticinctus is instantly recognizable on account of its colour pattern. It is the only Mesoamerican species that is black with a centrally bright yellow gaster. BIOLOGICAL INFORMATION. Thyreodon laticinctus is a widespread species in tropical Central America, and its range extends southwards into South America. Porter (1984: 63) described T. laticinctus as ‘the most hygrophilous and precinctively sylvan Middle American Thyreodon’. In Costa Rica T. laticinctus occurs in wet forests from sea level up to an altitude of about 2200 m and, at Monteverde (1300 m) W. Haber reared one specimen from the larva of Xylophanes anubus (Cramer). T. /aticinctus has never been collected in seasonally dry Pacific forests, despite the fact that its known host occurs commonly in this locality, feeding as larvae on rubiaceous shrubs (Janzen, 1984a) MATERIAL EXAMINED Holotype 2 (Thyreodon laticinctus Cresson), Mexico: Orizaba (PANS). Holotype 2 (Thyreodon principalis Smith), Costa Rica: Cachi (BMNH). Belize: 1 &’, Punta Gorda (BMNH). Costa Rica: Cartago Prov.: 1 0’, Turrialba (USNM); 1 2, Volcan Irazi, 2-2200 m (Rodgers) (BMNH): Guanacaste Prov.: 1 2, Volcan Orosi, Casa Maritza [= Mariksa], 700 m, vi.1986 (Gauld) (BMNH): Limon Prov.: 1 2, Guapiles (Schaus) (USNM): San José Prov.: 1 2, Braulio Carrillo National Park, 500 m, iv.1985 (Goulet & Masner) (CNC). Guatemala: 1 0’, Senahu, Vera Paz (Champion) (BMNH); 1 9, Zapote (Champion) (BMNH). Mexico: Orizaba: 1 CO’, Omealca (Trujillo) (BMNH). Nicaragua: 1 2, Chontales (BMNH). Panama: | CO’, Cerro Campana, vii.1970 (Howden) (TC). I have also examined specimens from Colombia, Ecuador and Peru. Thyreodon morosus Smith (Fig. 59) Thyreodon morosus Smith, 1879: 230. Holotype 2, COSTA RICA (BMNH) [examined]. Thyreodon pulchricornis Szépligeti, 1906: 133. Lectotype 2, PERU (TM), designated by Townes & Townes (1966: 190). [Synonymized by Hooker, 1912: 112.] Diacnosis. A jet-black species with proximal 0.6 or more of flagellum whitish or whitish yellow; wings entirely blackish. Ocelli small, the hind ones separated from the eyes by more than the maximum ocellar diameter; genae weakly constricted behind eyes; occipital carina ventrally incomplete, its end slightly turned towards hypostomal carina. Mesoscutum with notauli weakly impressed, quite broad and rugose, anteriorly without a distinct crest; mesopleuron centrally with close fine punctures; propodeum extensively diagonally striate posteriorly, without distinct median and lateral tubercles, posterodorsally with a shallow median longitudinal furrow. Fore wing length 18-23 mm. Hind femur moderately slender, distally cylindrical. Gaster quite slender, tergite 2 more than 2.5 times as long as posteriorly deep; male with gonosquama apically indented, with dorsal and ventral short acute processes. Remarks. Thyreodon morosus is immediately recognizable on account of the colour of its antennae. In freshly collected specimens the basal two-thirds of the flagellum is white, though this seems to become yellowish white with age after death. BIOLOGICAL INFORMATION. A widely distributed tropical species whose range extends from southern Mexico to Peru. In Costa Rica T. morosus has only been collected in or adjacent to humid forests. Its hosts are unknown. 64 IAN D. GAULD MATERIAL EXAMINED Holotype 2, Costa Rica: Cachi (BMNH). Costa Rica: Alajuela Prov.: 1 O’, 1.6 km S. La Marina, 600 m, x.1966 (Paulson) (FSCA): Guanacaste Prov.: 1 2, Estacion Mengo, SW. side Volcan Cacao, 1000 m, vi.1987 (Janzen) (BMNH): San José Prov.: 1 9, Braulio Carrillo National Park, 500 m, iv.1985 (Goulet & Masner) (CNC). Mexico: Yucatan: 1 C’, Valladolid (BMNH). Panama: 1 9, Anajuelo, iv.1911 (Buschk) (USNM). The ENICOSPILUS genus-group PRETHOPHION Townes Prethophion Townes, 1971: 74. Type-species: Prethophion latus Townes, by original designation. Diacnosis. Stout, orange-brown insects with weakly yellowed wings; fore wing length 17-20 mm. Mandi- bles weakly twisted but strongly narrowed; maxillae and labium unspecialized; clypeus apically convex; occipital carina absent. Notauli vestigial; mesopleural furrow extending from episternal scrobe to subalar prominence; unusual in that epicnemial carina absent laterally from the mesopleuron; posterior transverse carina of the mesosternum represented by vestiges laterally. Fore wing (Fig. 8) with Rs+2r almost straight; discosubmarginal cell without a hairless area in the anterior corner; pterostigma moderately broad; 1m-cu sinuous; hind wing with Rs almost straight; distal abscissa of Cu1 basally almost equidistant between M and 1A. Fore tibial spur without a membrane behind comb. Gaster stout; second segment with epipleuron usually folded under (rarely becoming pendant in some dried individuals); tergite 2 of similar length to tergite 3; ovipositor proximally extremely stout. RemarkS. Prethophion contains a single described species, P. latus, which is widely distributed throughout tropical South America from Bolivia north to central Costa Rica Prethophion latus Townes (Fig. 8) Prethophion latus Townes, 1971: 75. Holotype 2, BOLIVIA (TC) [examined]. Description. Mandibles small, malar space less than 0.2 times basal mandibular width; ocelli large, the posterior ones contiguous with the eye margin; occiput abruptly declivous behind posterior ocelli. Flagellum with 48-51 segments, the 20th segment 1.8—2.0 times as long as broad. Mesoscutum weakly polished, finely punctate; mesopleuron finely punctate, the area between the punctures microreticulate; scutellum without lateral carinae; lateral part of metanotum strongly inflated; metapleuron convex, sculptured like mesopleuron. Propodeum in profile short, abruptly declivous; anterior transverse carina complete centrally, lateral extremities absent; posterior transverse carina represented laterally by strong raised crests; lateral longitudinal carina complete, joined to spiracular margin by a raised ridge. Gaster stout; male with sternites unspecialized; female with ovipositor strongly laterally compressed. Orange-brown species with head paler yellowish; median mesoscutal vitta infuscate; antenna black; interocellar area slightly infuscate between posterior ocelli; pterostigma brownish, wings slightly yellowish. VARIATION. The Central and South American specimens are extremely similar in structure, sculpture and coloration. REMARKS. This large, robust species is superficially similar to some of the stouter species in the Enicospilus americanus species-complex. It differs most obviously from these taxa in lacking any trace of the occipital carina. BIOLOGICAL INFORMATION. In Central America Prethophion latus has occasionally been collected at light in lower montane wet forests at altitudes between 700 and 1100 m. Nothing is known of the host preferences of this insect. MATERIAL EXAMINED. Holotype 9, Bolivia: Cochabamba, Alto Palmar, 1100 m, ix.1960 (Walz) (TC). Paratype. Peru: 1 9, Cuzco, Paucartambo, 400 m, 11.1952 (Woytkowski) (TC). (Chacon) (BMNH). Panama: Chiriqui Prov.: 2 2, Fortuna, 1050 m, ii, v.1978 (Wolda) (RNH). OPHIONINAE OF TROPICAL MESOAMERICA 65 SIMOPHION Cushman _ Simophion Cushman, 1947: 446. Type-species: Simophion excarinatus Cushman, by original designation. Diacnosis. Slender, orange-brown insects with hyaline wings; fore wing length 13-16 mm. Mandibles not distinctly twisted, weakly narrowed; maxillae and labium unspecialized; clypeus apically slightly concave; occipital carina complete. Notauli vestigial; mesopleural furrow extending from episternal scrobe to subalar prominence; unusual in that epicnemial carina absent laterally from the mesopleuron; posterior transverse carina of the mesosternum represented by vestiges laterally. Fore wing with Rs+2r slightly bowed; discosubmarginal cell with a hairless area in the anterior corner; pterostigma moderately broad; 1m-cu strongly bowed; hind wing with Rs bowed; distal abscissa of Cu1 basally closer to 1A than to M. Fore tibial spur without a membrane behind comb. Gaster slender; second segment with epipleuron pendant (in Mesoamerican species); tergite 2 of similar length to tergite 3. REMARKS. Simophion is a small genus with a few species in arid areas of the south-western United States, the Middle East and Central Asia (Townes, 1971; Gauld, 1985). The damaged holotype and only known specimen of Ophion melanostigma Cameron seems to belong to Simophion as it possesses the apomorphic features that characterize the genus; it has no flange on the fore tibial spur, has no propodeal carinae, lacks the posterior transverse carina of the mesosternum and has no epicnemial carina. However, it also possesses a number of apomorphic features that have not been found to occur in other Simophion species, namely a blunt trochantellar tooth, a subcentral petiolar spiracle, a pendant laterotergite 2 and a strongly bowed 1Im-cu. As O. melanostigma and Simophion are only united by ‘loss’ apomorphies, its placement in this genus must be considered tentative. The holotype of S. melanostigma is damaged and lacks the posterior part of the gaster so I cannot be certain which sex it is. It is reputed to be a male (Townes & Townes, 1966), but the claws are more like those of a female. If it isa male then the form of the claw clearly separates it from Ophiogastrella, but if it is a female there is a possibility that it could be an extremely aberrant species of Ophiogastrella. Its systematic position will be more clearly resolvable if a male were known. Simophion melanostigma (Cameron) comb. n. Ophion melanostigma Cameron, 1886: 295. Holotype ? 9, PANAMA (BMNH) [examined]. Description. Mandibles moderately long, the upper tooth slightly longer and distinctly broader than the lower tooth; outer surface of mandible coarsely and shallowly punctate, with microreticulation between punctures, flat with a strong proximal concavity; malar space about 0.1 times as long as the basal mandibular width. Head in front view with width across the eyes 1.5 times the maximum height from the vertex to the clypeal apex; clypeus weakly convex, polished punctate; lower face unusually flat, rather closely punctate centrally; eyes ventrally slightly divergent. Posterior ocelli very close to eye margins; head posteriorly slightly buccate; occipital carina mediodorsally complete, slightly pointed, ventrally very strong, joining hypostomal carina. Antenna long and slender with 61 flagellar segments; 20th segment 2.5 times as long as broad. Mesoscutum highly polished, in profile evenly rounded; notauli very weak. Mesopleuron polished, finely and sparsely punctate. Scutellum flat, long, quite narrow but not strongly tapered posteriorly, carinate only at extreme anterior end. Metapleuron weakly convex, polished, obsoletely punctate. Pro- podeum smooth and polished, evenly declivous, dorsally rather flattened; propodeal carinae absent. Fore wing length about 16 mm; CI = 0.53; ICI = 0.67; SDI = 1.10; Rs+2r barely thickened or turned basally, centrally rather evenly bowed; 1m-cu evenly bowed, without a ramellus; discosubmarginal cell with a large glabrous area anteriorly; Ist subdiscal cell evenly sparsely hirsute; cu-a opposite base of Rs&M. Hind wing with Rs curved through about 45°; marginal cell proximally with a glabrous area. Legs slender, unspecialized except that the posterior two pairs have the distal margin of the trochantelli ventrally produced into a blunt tooth; mid tibial spurs unusual in being almost equal in length, the longer 1.05 times the length of the shorter; hind coxa slender, in profile about 2.4 times as long as deep; hind trochantellus mediodorsally about 0.8 times as long as broad. Gaster quite slender; tergite 1 unusual in that the spiracle is positioned quite far forwards, about 0.6 of the way along the segment; segment 2 with laterotergite pendant; umbo indistinct. Golden yellowish species with antennae reddish brown, interocellar area slightly infuscate; wings hyaline, pterostigma dark brown. REMARKS. Simophion melanostigma differs from all other species in the genus in having a pendant laterotergite, a very slender Rs+2r in the fore wing, more or less equal mid tibial spurs and no distinct umbo. Furthermore it is the only species of Simophion that occurs outside of arid habitats, and I have some doubt that it is correctly placed here. 66 IAN D. GAULD BIOLOGICAL INFORMATION. This species has only been taken at an altitude of between 650-1000 m on Volcan de Chiriqui in Panama. I have not seen this habitat, but presumably it would be an area of very wet forest. Although I have seen no other examples from Central America, there is no reason to doubt the locality data given on this specimen. MATERIAL EXAMINED Holotype ? 2, Panama: Volcan de Chiriqui, 650-1000 m (BMNH). OPHIOGASTRELLA Brues Ophiogastrella Brues, 1912: 201. Type-species: Ophiogastrella maculithorax Brues, by original designation. Brachyscenia Enderlein, 1921: 36. Type-species: Brachyscenia nigriventris Enderlein, by original designation. Diacnosis. Slender orange or yellowish, sometimes black-marked species; fore wing length 6-16 mm. Mandibles not twisted; maxillae and labium unspecialized; clypeus truncate or slightly concave; occipital carina complete. Notauli vestigial or absent; mesopleural furrow vestigial; posterior transverse carina of mesosternum represented by lateral vestiges. Fore wing generally with Rs+2r slender and not strongly curved, rarely basally incrassate and angulate; discosubmarginal cell with a hairless area in anterior corner; pterostigma moderately stout; 1mcu generally evenly curved; hind wing with Rs straight to bowed; distal abscissa of Cu1 basally from almost equidistant between M and 1A, to closer to 1A than to M. Fore tibial spur without a membrane behind comb. Gaster slender; second segment with laterotergite folded under, or narrow, membranous and pendant; tergite 2 longer than tergite 3; male unusual in having the pectinal teeth extending around apex of flattened claws (Fig. 15). REMARKS. Ophiogastrella is a small genus that is endemic to tropical America and whose cumulative range extends from tropical Central America to southern Brazil. Three species have been described from South America, and probably about eight more are undescribed in the whole Neotropical realm. In Central America several species of Ophiogastrella can be quite common in seasonally dry areas. In Pacific dry forest in Santa Rosa National Park, Costa Rica, four synchronous, sympatric species occur in fairly large numbers at the start of the wet season. All have a very short flight period of only a few weeks. Nothing is known of the host preferences of any of them. These species are keyed and described below. Key to Mesoamerican species of Ophiogastrella 1 Marginal cell of hind wing uniformly (but sometimes sparsely) hirsute proximally (Fig. 72); fore wing with Rs+2r basally quite strongly thickened and somewhat angulate (Fig. 75); occipital carina arched upwards into a point mediodorsally, male with distal margin of subgenital plate bearing a distinct median acute tooth (Fig. 80)...................... . gonzalezisp.n. (p. 68) — Marginal cell of hind wing with a large glabrous area proximally (Figs 71,73); fore wing with Rs+2r basally only slightly thickened, almost straight to weakly bowed (Fig. 74); occipital carina mediodorsally evenly convex or slightly flattened; male without a median acute tooth on distal margin of subgenital’ plate (Fig.79) .c .t-astir « 2yse/s/acdiaisierwhols Tawi. ew dda 6 sh a qe Rew aa ao nes — Hind tarsal claws moderately long, apically quite abruptly curved through 85—90+° (Fig. 98); face from elongate to subquadrate, that of male less than 0.90 times, and that of female Tenclngess than O.e5 times das broad! as Nighy sso cc. 2s» MWR Sone note ee ieee vine 22 Larger species, fore wing length 15+ mm, wings weakly to strongly infumate (Fig. 236); mandibles long, distally parallel sided; lateral longitudinal carina of propodeum not present Hence anelnOLD transverse CARMA... 0.:c.ccecccscvacne sc + csp texanus (Ashmead) (p. — Smaller species, fore wing length 10-15 mm, wings hyaline (Fig. 237); mandibles rather short, evenly tapered; lateral longitudinal carina of propodeum usually complete. cushmani Gauld (p. 25euoOre wine with Rs-+-2r very weakly sinuate (Figs 238, 239) ......... 00d sescaeeeeceen cess — Fore wing with Rs+2r moderately to strongly sinuate (Figs 241-248) ..................00. 24 Scutellum with lateral longitudinal carinae present only at extreme anterior end; genae dis- tinctly buccate (Fig. 97); fore wing with Im-cu distal to bulla 0.56 times as long as 3s-m (Fig. 238). Male with posterior sternites bearing fine decumbent pubescence.... halffteri sp. n. (p. — Scutellum with lateral longitudinal carinae extending at least 0.5 of its length; genae evenly rounded behind eye (Fig. 96); fore wing with Im-cu distal to bulla more than 0.75 times as long as 3rs-m (Fig. 239). Male, where known, with posterior sternites bearing long stout erect MEE tice: SOci 0 = 3h Rca, Set ea vias a> Oma echo HAG > Vial awh Ss G's» waves Om ere 25 Propodeum with posterior area immediately behind the anterior transverse carina smooth and polished, and with anterior and spiracular areas forming a broad shallow groove; postscutellum with a median longitudinal keel. .................... sarukhani sp. n. (p. — Propodeum with posterior area coarsely sculptured right up to anterior transverse carina, and with anterior and spiracular areas forming a short, deeply U-shaped groove (Fig. 161); postscutellum without a distinct median longitudinal keel. ...................0.2.0000. 26 Fore wing with fenestra bearing a weakly sclerotized, oval or elliptical mark centrally; 2nd discal cell not exceptionally elongate, with abscissa of Cula between Culb and 2m-cu less than 1.35 times as long as first abscissa of Cul (Fig. 240); a pallid yellow species. aktites Gauld (p. — Fore wing with fenestra more or less hyaline, without a sclerotized elliptical or oval mark; 2nd discal cell very elongate, with abscissa of Cula between Culb and 2m-cu more than 1.35 times as long as first abscissa of Cul (Fig. 233); a darker yellowish brown species. hallwachsae sp. n. (in part) (p. 27 Fenestra very large, reaching close to junction of Rs and 3rs-m, margined peripherally by a weak and broadly U-shaped sclerite (Fig. 241); abscissa of Im-cu distal to bulla short, 0.50—-0.75 times as long as 3rs-m; 1m-cu quite evenly bowed. Propodeum with anterior area long, almost smooth; posterior area with weak irregular REMMI LEME (ate OZ) en gains artis ee one Steel oxera cers feces sri gomezpompai sp. n. (in part) (p. — Fenestra usually smaller, its margin not close to junction of Rs and 3rs-m, or if close, then without a peripheral U- shaped sclerite (Figs 242-248); sclerite otherwise present or absent, usually the latter; abscissa of 1m-cu distal to bulla of various lengths, usually more than 0.80 times as long as 3rs-m; 1m-cu from fairly evenly bowed to sinuous ...................4. 28 Fore wing with fenestra bearing two distinct, similar-sized weakly sclerotized quadrae (Fig. 242). Male with sternites 7-9 bearing long stout erect pubescence (Fig. 163); clypeus in profile TLE ome aM mer RN oe cate trae Pete eee sc cose, Susie is Gi. ict ks wien tie 95 visleiaee er _ lebophagus Gauld (p. 81 147) 150) 22 23 154) 155) 26 156) 146) 157) 28 160) 82 IAN D. GAULD ee ee 141 Figs 132-146 Enicospilus species. 132, 133, posterior part of mesothorax, ventral view; (132) E. guin- dont; (133) E. leoni. 134, 135, metapleuron and propodeum, lateral view; (134) E. erasi; (135) E. brenesiae. 136, 137, lower part of head and mandibles, lateral view; (136) E. erasi; (137) E. brenesiae. 138-140, hind trochanteral segments; (138) E. burgosi; (139) E. gallegosi; (140) E. dispilus. 141-144, hind wing; (141) E. masoni; (142) E. howdenorum; (143) E. dispilus; (144) E. georginae. 145, 146, metapleuron and propodeum, lateral view; (145) E. howdenorum; (146) E. sondrae. OPHIONINAE OF TROPICAL MESOAMERICA — Fore wing with fenestra without quadrae, or with one quadra discernible (Figs 243-248) .... 29 Fenestra quite long, its distal side separated from base of Rs by about length of 3rs-m (Fig. 243); genal carina more or less reaching hypostomal carina; apex of clypeus truncate to slightly concave (Fig. 164). Male with sternites 7-9 bearing long stout erect hairs; female with ovipositor sheath paler mowimisn. only shehtly infuseate apically... 2... 6 ee eee eee ee eee ugaldei sp. n. ( — Fenestra short, its distal side separated from base of Rs by more than length of 3rs-m (Figs 244— 248); genal carina ventrally evanescent, not reaching hypostomal carina; apex of clypeus from cet PERU OXEC AROSE COMICAL crys telat «++. = Seeieemios aiNare etre aya wicistcle le, «- «puss cicielel Ss 30 Metapleuron characteristically convex posterodorsally, flattened anteroventrally (Fig. 165); propodeum with posterior area dorsally irregularly rugose to reticulate, never concentrically rugose/striate. Male with sternites 7—9 bearing fine scattered erect hairs ............ umanai sp. n. (p. — Metapleuron evenly convex, not anteroventrally flattened (Fig. 165); propodeum with pos- terior area concentrically rugose/striate, at least near posterior end..................... 31 Fore wing with 3rs-m conspicuously longer than abscissa of M between 3rs-m and 2m-cu (Fig. 245). Male with sternites 7-9 bearing stout erect hairs.................. quintanai sp. n. (p. — Fore wing with 3rs-m shorter than abscissa of M between 3s-m and 2m-cu (Figs 247-248). ... 32 Antenna with central segments 1.7 or less times as long as broad; clypeus in profile weakly to strongly out-flared and with genae somewhat swollen. Mainly present in extra- tropical areas. americanus (Christ) (p. — Antenna with central segments 1.8 or more times as long as broad; clypeus in profile generally flat to weakly convex, if slightly out- flared then genae not swollen; genae otherwise from 33 Male with subgenital plate conspicuously Rvs than the preceding sternite (Fig. 167); meta- pleuron finely and closely striate (Fig. 166); mandibles of moderate length, fairly evenly TAOVACIN LOT LOG st. tea a's Saher. tae Veda eles whleenees tenuigena (Kriechbaumer) (p. — Male with subgenital plate shorter than the preceding sternite; metapleuron punctate to punctostriate mandibles long, stout, distally parallel sided..................0..2.00005. 34 Genal carina ventrally strong, joining hypostomal carina; male with hind tarsal claws very abruptly curved with long apical point; sternites 7-9 of male bearing close, long stout erect Se ES cr 5. 4, 2 SRR Sino tecles + oi « «1 Malte cate venezuelanus (Szépligeti) (p. — Genal carina ventrally evanescent, not joining hypostomal carina; male with hind tarsal claws unusually long, quite abruptly curved, but with short apical point; sternites 7-9 of male bearing fine scattered semierect pubescence. Fore wing with Rs+2r bearing a sharp angulation ventrally near proximal 0.2 (Fig. 248). peigleri Gauld (p. 35 Outer surface of mandible with an impressed diagonal furrow extending from near upper proximal corner to between bases of apical teeth, this groove bearing close pubescence (Fig. Tee, PR. i. Foe a ach. . +s». saengiinciy tenon Und eles 2b bb wade as — Outer surface of mandible flat, weakly convex or slightly concave, never with a distinct hirsute diagonal furrow (Figs 172, 176), but occasionally with a shallow longitudinal furrow (Fig. 217) which does not bear hairs 36 Fore wing with central sclerite entirely absent; vein Rs+2r centrally evenly bowed forwards emerremes tna. Jere Mee itil oto: M, Unetcier, Casas useing ta ti eo kiehe a neotropicus Hooker (p. — Fore wing with a well-defined central sclerite; vein Rs+2r more or less straight or weakly sinuous (Figs 250, 251) 37 Metapleuron anteroventrally flattened (Fig. 169); fore wing with central sclerite more or less kite-shaped, with proximal side pointed and weakly sclerotized (Fig. 250). doylei sp. n. (p. — Metapleuron rather evenly convex (Fig. 170); fore wing with central sclerite subcircular to D-shaped, the inner side convex or straight, strongly sclerotized (Fig. 251). purgatus (Say) (p. > 7S) 38 Central sclerite present but confluent with the distal sclerite (Fig. 252).... luquillo sp. n. (p 83 29 p. 161) 164) 170) 175) 84 IAN D. GAULD Figs 147-152 Enicospilus species. 147, 148, subgenital plate of male; (147) E. bozai; (148) E. enigmus. 149, 150, metapleuron and propodeum, lateral view; (149) E. bozai; (150) E. enigmus. 151, 152, hind tarsal claws of female; (151) E. major; (152) E. clarkorum. OPHIONINAE OF TROPICAL MESOAMERICA — Central sclerite absent or, if present, separated from the distal sclerite.................... 39 Mandible with upper tooth about 0.7 or less times as long as the lower tooth (Fig. 172), and with EI TRPIRAIIT MUR LY COMVON foo 2 oo. 0)s.555 nisin Gis = wots vie atalefeie sain 4ixis me ei alb aE aielsls hae cle ies — Mandible with upper tooth subequal to or longer than the lower tooth or, if slightly shorter, then Semerpeus i profile exceptionally convex (Fig. 116)... 0.6... c cece cette eee eee 40 Fore wing with central sclerite vestigial, indistinctly differentiated (Fig. 253); 3rs-m less than 0.45 times as long as abscissa of M between 2m-cu and 3rs-m,; flagellum with 60 or fewer ME MEN foo ea iar e oe al ela ie (Si oher a's + «inte diutsinies ieee sab arerece dajaboni sp. n. (p 45 Interocellar area yellowish; fore wing usually with proximal sclerite subcircular, and generally with distal sclerite elliptical, with its longest axis parallel to Rs+2r trilineatus (Brullé) (in part) (p — Interocellar area black; fore wing with proximal sclerite more or less triangular, distal sclerite absent, or if present, then usually not elliptical and with longest axis parallel to Rs+2r.... 46 Central sclerite oval, weak, positioned close to distal margin of the fenestra (Fig. 258); male with sternites 7-9 bearing dense long pubescence. Metapleuron dorsally coarsely rugose (Fig. 103); central flagellar segments more than MPMRIE NN OTD AG ULOAG Vs cee eco. eae cles ccs Serceccccernes porteri sp. n. (p — Central sclerite absent, or if present then positioned at or proximal to centre of fenestra (Figs 259-261); male with sternites 7-9 bearing fine semidecumbent pubescence (Fig. 101) ..... 47 Fore wing with proximal sclerite narrow, very strongly acute anteriorly (Fig. 259); alitrunk extensively black-marked especially on mesoscutum and mesosternum. lupemejia sp. n. (p. — Fore wing with proximal sclerite rounded anteriorly or, if acute, then entire sclerite is almost equilaterally triangular (Figs 260, 261); alitrunk generally not black- marked, rarely with an Mlimesu the Tdatk Ol MICSOSCUCUM CENUIANY 2.5... ce ence ete an Sek sees sintigmen nae anaes 48 Male with aedeagus bearing a distinct collar (Fig. 175); either with hind trochantellus less than 0.2 times as long as broad mediodorsally, and/or with metapleuron coriaceous to striate. trilineatus (Brullé) (in part) (p 40 4] . 183) . 185) . 186) _ 189) 46 . 187) 47 188) 48 . 189) 86 IAN D. GAULD Zz 5 4 Figs 153-158 Enicospilus species. 153, 154, propodeum, dorsal view; (153) E. alvaroi; (154) E. chiri- quensis; 155, 156, subgenital plate of male; (155) E. mayi; (156) E. chiriquensis. 157, 158, mandibles; (157) E. robertoi; (158) E. brevis. OPHIONINAE OF TROPICAL MESOAMERICA — Male with aedeagus without a distinct collar, at most only with lateral flanges (Fig. 101); hind trochantellus 0.4 times as long as broad mediodorsally and metapleuron polished, punctate 87 dirzoi sp. n. (p. 194) 49 Central sclerite absent, or with only an indistinct, unpigmented thickening present in fenestra ase NI ey te UT ERTS. on 5.0 « eee MAM tea nc yis eed. alata gr ora v's Medias, bea ees =wieemiral sclerite present, distinctly pigmented (Figs 278-352) ............cccccecccccceeee Slmiumenecelian area uniformly black or dark brown ........2. 00. .00020 eee cee saee cc cceecaaee — Interocellar area yellowish, at most with a black mark between the posterior ocelli......... [Intermediate specimens will key either way] 51 Cells in distal part of both fore and hind wings with broad glabrous areas adjacent to veins (Figs 106, 262); fore wing with proximal corner of marginal cell infumate; outer hind corner of 2nd discal cell very acute (Fig. 262); hind wing with abscissa of Rs between R and rs-m more than 50 67 51 54 2.2 times as long as rs-m, bowed; distal abscissa of Cul strongly sinuate masneri sp. n. (p. 195) — Cells in distal part of fore and hind wings more or less uniformly hirsute (Figs 263, 264); fore wing with proximal corner of marginal cell not infumate; outer hind corner of 2nd discal cell obtuse or about a right-angle (Figs 263, 264); hind wing with abscissa of Rs between R and rs-m less than 2.2 times as long as rs-m, more or less straight (Fig. 107); distal abscissa of Cul Se LEMON E COE NGA SIU AGE i ciaien « « .. «+ ae ee inteiee M Aileeartimle Git wale bie Seca cerns ote 52 Fore wing with vein 3rs-m equal to or longer than abscissa of M between 2m-cu and 3rs-m (Fig. 221); gaster stout, tergite 2 in profile about 2.5 times as long as posteriorly deep (cf. Fig. 112); 52 female with ovipositor sheath very stout. ......... exoticus (Morley) (few specimens) (p. 216) — Fore wing with vein 3rs-m less than 0.6 times the length of M between 2m-cu and 3rs-m (Figs 263, 264); gaster slender, tergite 2 in profile more than 4.5 times as long as deep posteriorly; MREMMOSUOL SHOAL SICNGET ..........cscdactesceucsiectccicasvevecdeucucucns 53 Fore wing with Rs+2r virtually straight; fenestra short; distal sclerite absent; first abscissa of Cula more than 1.10 times as long as first abscissa of Cul (Fig. 263). 53 randalli sp. n. (most specimens) (p. 196) — Fore wing with Rs+2r slightly sinuate; fenestra moderately long; distal sclerite weak but discernible; first abscissa of Cula less than 1.05 times as long as first abscissa of Cul (Fig. CSG. os etic elec es Reels 2 aneplepne bad ele anee ane cressoni Hooker (in part) (p. 198) 54 Lower tooth of mandible broader than the upper tooth and ventrally swollen, with ventral margin sharp (Fig. 176); mandible twisted 20-30°. ieceta row Species With termites 3+- black (022 ....2-- conc pees se sveeccevceuseus — Lower tooth of mandible of similar breadth to, or more slender than the upper, or if broader then the ventral margin lacks a sharp ventral margin and the mandibular apex is twisted more Pa MAncdiole Otmerwise VarlOusly twisted: . iu... c.dc0 ees cc ee ere cesses cccescess 55 Metapleuron regularly punctate (Fig. 177); fore wing with 3rs-m more than 1.00 times as long as abscissa of M between 2m-cu and 3rs-m (Fig. 265); distal sclerite obsolescent, if discernible confluent with proximal sclerite; posterior transverse carina of mesosternum weak or 56 eB OE tev COMITE AUN sc uc Pee Gs... .. wes biedaulatediaevects laurenae sp. n. (p. 199) — Metapleuron posterodorsally striate (Fig. 178); fore wing with 3rs-m less than 1.00 times as long as abscissa of M between 2m-cu and 3rs-m (Fig. 266); distal sclerite discernible distally, not confluent with proximal sclerite; posterior transverse carina of mesosternum complete. vegai sp. n. (p. 200) 56 Fore wing with 3rs-m more than 0.75 times as long as abscissa of M between 3rs-m and 2m-cu OPCS ZS SEG Se al at OS Oar ee, ee ee — Fore wing with 3rs-m less than 0.70 times as long as abscissa of M between 3rs-m and 2m-cu (Figs ee eet aera Re ere es ee cre ees EO TA we oer eis oe dee 57 Proximal sclerite barely broader than and confluent with a similarly sclerotized distal sclerite (Fig. 267); mandible twisted less than 45°; lateral crest of propodeum absent. (Fig. 104). a7 Sf baltodanorum sp. n. (p. 201) — Proximal sclerite very much broader than and far more strongly sclerotized than distal sclerite (Figs 268, 269); mandible twisted 45° or more; propodeum with a weak to strong lateral crest Wee LO Erne nee SP Set body cs ORS oa Ba iad eves oes ca SOR. be 88 IAN D. GAULD Figs 159-164 Enicospilus species. 159-162, propodeum, dorsal view; (159) E. cameronii; (160) E. gamezi; (161) E. aktites; (162) E. gomezpompai. 163, subgenital plate of male, E. lebophagus. 164, face, E. ugaldei. OPHIONINAE OF TROPICAL MESOAMERICA 58 Fore wing with 3rs-m less than 1.20 times as long as abscissa of M between 3rs-m and 2m-cu (Fig. 268); occiptal carina mediodorsally complete, strong; ventral corner of epicnemial carina rounded (Fig. 108); hind tarsal claws long ................22.0005 carri sp. 0. (p. — Fore wing with 3rs-m more than 1.30 times as long as abscissa of M between 3rs-m and 2m-cu (Fig. 269); occipital carina mediodorsally incomplete or weak; ventral corner of epicnemial carina usually extended backwards slightly and acutely angled (Fig. 109); hind tarsal claws eat ee fae on ey Noe ols Sv oo eR aie ear rete le apes a eae eves colini sp. n. (p. 59 Fore wing with marginal cell proximally strongly infumate; abscissa of Cul between Im-cu and Cula more than 0.40 times as long as Cu1b (Fig. 270); pterostigma centrally black, proximally RC iene maiam os ne + ee se ey een ess maculipennis (Cameron) (p. — Fore wing with marginal cell hyaline or uniformly weakly infumate; abscissa of Cul between 1m-cu and Cula less than 0.35 times as long as Culb (Figs 271-274); pterostigma orange or IEICE REE Se TTT Coss «no ARMA RA ww ah alow halyard ae wapelan sce a se 60 Fore wing with Rs+2r bowed for most of its length (Fig. 271); fenestra long, reaching close to TPRIPINE IOUS ACS +22 fh; ATIGUIIGS) «. Sraye ae: cre oe 2) oer eieneieidisieieraiviesn’ @ guatemalensis (Cameron) (p. — Fore wing with Rs+2r straight or sinuous (Figs 272-274); fenestra shorter, its distal margin Revataredinom base of Rs by length’of 3rs-mm or MOTe ..... 22.02 e cece eee ee cee eet ness 61 Fore wing with first abscissa of Cula equal in length or shorter than abscissa of Cul between Rs&M and 1m-cu (Fig. 264); hind wing with first abscissa of Rs weakly but distinctly bowed (cf. Fig. 127), and with distal sclerite separated from proximal sclerite. cressoni Hooker (in part) (p. — Fore wing with first abscissa of Cula 1.05 or more times as long as abscissa of Cul between Rs&M and 1m-cu (Figs 272- 274); hind wing with first abscissa of Rs straight or, if weakly bowed, then distal sclerite confluent with proximal sclerite; distal sclerite otherwise various 62 Posterior transverse carina of mesosternum broadly incomplete centrally; margin of clypeus truncate (Fig. 179) or even slightly concave; antennae golden; small species, fore wing length MOM ten Nee Tee occa e ree e tees Corer eres karrensis sp. n. (p. — Posterior transverse carina of mesosternum complete or rarely weak centrally; margin of clypeus slightly convex (Fig. 180); antennae basally blackish or dark brown; larger species, Pe CMe OTM OTe Aten CHAM LS TIT jy... «<< + ausi'vtnlaby ayaa ore (ehauapwyeuelansiraye lah n/enaior sielaraievers:« » 63 Mandibles twisted 60° or more (Fig. 180); metapleuron strongly convex; propodeum postero- dorsally very strongly flattened, laterally with a rather sharp crest formed from the vestige of REN RERE MGRAMISUGTSC CAMA Tinea pip gis «+ «orc neh dean vine d ety sedeeen ci daeaenen ees — Mandibles twisted 40° or less (Fig. 173); metapleuron weakly to moderately convex; pro- podeum generally weakly flattened dorsally, if deplanate then laterally without a sharp crest mame trom vestige of posterior transverse Carina .........4..202- cece cece cecenecnes 64 Large species, fore wing length 19+ mm; female with ovipositor sheath exceptionally stout (Fig. meer easier pale with tergites 5+ infuscate. ..........cc eee aeces tees stevensi sp. n. (p. — Smaller species, fore wing length 14-16 mm; female with ovipositor sheath relatively slender tiie. 110); gaster orange with tergites 3+ blackish................... pamelae sp. n. (p. 65 Fore wing with Rs+2r very straight; fenestra small; distal sclerite absent or extremely weak, represented only by slight infumation (Fig. 275)....... randalli sp. n. (few specimens) (p. — Fore wing with Rs+2r slightly sinuous; fenestra of moderate size; distal sclerite present but often weak (Figs 276, 277) 66 Metapleuron striate (cf. Fig. 178); fore wing with abscissa of 1m-cu distal to bulla more than 1.10 times as long as 3rs-m; proximal sclerite gradually tapered distally so that the junction between it and the distal sclerite is not clear (Fig. 276); distal sclerite evanescent distally, not temeIe INVENT MONIES. Uo: Poe rcs cease lester iv csscarecensas vilmari sp. n. (p — Metapleuron more or less punctate (cf. Fig. 177); fore wing with abscissa of 1m-cu distal to bulla less than 1.00 times as long as 3rs-m; proximal sclerite abruptly tapered distally, confluent with the slender distal sclerite, but with junction between then clearly recognizable (Fig. 277); distal sclerite more or less complete, extending around most of periphery of fenestra. 1 o:tne: 48 610 G GIB al Oe) OO "OG ONE CHO ONC LO A Ce NC ee ee estradarum sp. n. (p. 89 202) 203) 205) 60 206) 61 198) 62 209) 63 64 219) 213) 90 IAN D. GAULD Figs 165-170 Enicospilus species. 165, 166, metapleuron and propodeum, lateral view; (165) E. umanai; (166) E. tenuigena. 167, 168, E. tenuigena; (167) subgenital plate of male; (168) mandible. 169, 170, metapleuron and propodeum, lateral; (169) E. doylei; (170) E. purgatus. OPHIONINAE OF TROPICAL MESOAMERICA 91 67 Central sclerite crescentic, positioned very close to and of similar surface area to the distal gE) le ely Ari cette ag aR PS 2 cubensis (Norton) (in part) (p.183) — Central sclerite of various shapes, if crescentic never both close to and of similar surface area MURS RO SIU TAMSCIEMMCC Utter ric TEE: = «os RET eyterc Siale Walle wien es sia wma emai oai leds we 68 rma aed UIMIGLMY(DIACK 044. as!» - 0. cWebae Woe esa v sieeedee ud detGeceve senate 69 — Interocellar area yellowish to brownish, at most infuscate between hind ocelli ............. 84 69 Clypeus in profile nasute, strongly produced and with a flattened ventral region (Fig. 116); mandibles weakly tapered, with lower tooth slightly the longer. ....... teodorae sp. n. (p. 214) — Clypeus in profile flat to moderately convex, never nasute, and never with a flattened ventral region; mandibles usually moderately to strongly tapered, generally with upper tooth the Bn aa See Prey eo wc un ENS 5 ene a Paton ere Saha wa wad Fawe et 70 70 Fore wing with 3rs-m at least 0.75 times as long as abscissa of M between 2m-cu and 3rs-m (Figs 280-282); stout species, gaster with tergite 2 in profile generally less than 4 times as long as Kis icSiehe GlEG yo) (QRS 7) I Ee tet Rar ener ane Rear en Soe eae 7) — Fore wing with 3rs-m less than 0.70 times as long as abscissa of M between 2m-cu and 3rs-m (Figs 283-290); slender species, gaster with tergite 2 in profile more than 4 times as long as a TAC LU NeCee rH (HN Oerm IED) aca Hah ar ic cs... 6 ais s © « cmmetsievdlnssjslslo Sieteve celtic me ere bie eee tie la's ere ¢ 73 71 Flagellum stout, 20th segment 1.4-1.6 times as long as broad; tarsal claws of female with pectinae centrally widely interspaced, proximally and distally close together (Fig. 216); very large species, fore wing length 20-23 mm; mesoscutum uniformly reddish brown, or rarely Me EAM LET OULCOILALIV a tomvaitawiiviog « «+s Lieibeget vaddreleeciens dss maritzai sp. n. (p. 215) — Flagellum less stout, 20th segment 1.9-2.2 times as long as broad; tarsal claws of female quite uniformly pectinate (Fig. 218); moderately large species, fore wing length 13-15 mm; meso- scutum usually with black vittae or extensively dark-marked.................02.-00000- 72 72 Fore wing with alar sclerites very weak; 3rs-m equal to or longer than abscissa of M between 3rs-m and 2m-cu (Fig. 281) tergite 4 of gaster anterolaterally pale-marked exoticus (Morley) (in part) (p. 216) — Fore wing with alar sclerites strongly pigmented; 3rs-m equal to or shorter than abscissa of M between 3rs-m and 2m-cu (Fig. 282); tergite 4 of gaster usually entirely black, rarely pale- aI eee Tal 20) oP tte ake bones baa tae urn’ = unle cater wienierareneah ns pacha Jesicae sp. n. (in part) (p. 218) 73 Fore wing with distal sclerite strongly pigmented, complete, extending from proximal sclerite around margin of fenestra to nearly reach Rs+2r (Figs 283-285) ...........0.0.cc ee eeee 74 — Fore wing with distal sclerite absent, or weakly pigmented, or if strongly pigmented then not bordering virtually the entire margin of the fenestra (Figs 286-290)..................... 76 74 Mandibles twisted 30-S0°, with lower tooth broad, with a pronounced convex ventral cutting surface (Fig. 120); head, in dorsal view moderately narrow (Fig. 130). coercovadoi sp. n. (p. 220) — Mandibles twisted 70° or more, with lower tooth a little compressed but without a ventral cutting edge (Fig. 119); head, in dorsal view strongly transverse (Fig. 131)..................... 75 75 Fore wing with central sclerite oval/elliptical (Fig. 284); upper hind part of mesopleuron with a blackish mark; metapleuron finely punctate (Fig. 181)................ lovejoyi sp. n. (p. 221) — Fore wing with central sclerite somewhat stoutly S-shaped (Fig. 285); upper hind part of mesopleuron not black-marked; metapleuron striate or rugose-vermiculate (Fig. 182). hacha sp. n. (p. 223) 76 Meso- and metapleuron highly polished, smooth, at most with fine, sparse punctures (Fig. 183); proximal sclerite abruptly tapered distally, with a very slender confluent distal sclerite (Figs Ze lOhy ZA) sesceeprabelitec cache ato ac ah Sete MER ARC RS ERE ee a SPAN Paar ee Pn Uh — Meso and/or metapleuron striate or variously coarsely sculptured (Fig. 184), sometimes closely punctate, but then the punctures separated by about their own diameters or less; if distal sclerite is confluent with the proximal sclerite then the latter is evenly tapered to join the RONG Tn (STS SO) Rae eee eee cee eso le hey cat ahc aloaa vega aes o cyaicem. Viscum ancjetsts ine ete devacn 78 77 Mandibles twisted 70° or more (Fig. 118); occipital carina joining hypostomal carina (Fig. 114); hind trochantellus, in dorsal view, projecting beyond apex of trochanter by about 0.2 of its breadth (Fig. 122); posterior transverse carina of mesosternum complete. xanthostigma (Szépligeti) (p. 225) 92 IAN D. GAULD Figs 171-176 Enicospilus species. 171, 172, mandibles; (171) E. purgatus; (172) E. columbianus. 173, 174, face; (173) E. randalli; (174) E. trilineatus. 175, aedeagus, E. trilineatus. 176, mandibles, E. laurenae. OPHIONINAE OF TROPICAL MESOAMERICA — Mandibles twisted 30° or less (Fig. 117); occipital carina ventrally evanescent, not joining hypostomal carina (Fig. 115); hind trochantellus, in dorsal view, projecting beyond apex of trochanter by about 0.5 of its breadth (Fig. 121); posterior transverse carina of mesosternum CT en ot Saisie ic og 05s «Tapers (oie Scnceielne's «vw teas wee’ cepillo sp. n. (p. 78 Fore wing with central sclerite very slender and characteristically shaped like an inverted letter J LS sien lo RES 6 epee oe i ae persimilis (Szépligeti) (p. — Fore wing with central sclerite various, usually elliptical, subcircular or linear, never J-shaped ete Se ee LR... ate nn hme aud suiahe LRG dja ele bv eae ws 79 Antenna with at least the five basal flagellar segments yellowish brown or orange ........... — Antenna with five basal flagellar segments blackish or dark brown ..................-.4.. 80 Central sclerite subcircular to oval, its longest axis at 90° to Rs+2r (Fig. 289); marginal cell proximally less densely hirsute than centrally; outer surface of mandible without a distinct proximal concavity; gaster usually with tergites 5+ black, the preceding ones yellowish BT OLN MNES ot OPN S Ste SILL t a ASS valx abdicate Si wlals b ave eiahe Slallaheie oe wine wea Me fernaldi Hooker (p. — Central sclerite oval-elliptical, its longest axis almost parallel to fore margin of wing (Fig. 290); marginal cell uniformly hirsute; outer surface of mandible with a well- developed proximal concavity; gaster usually uniformly yellowish brown, rarely with tergites 4+ infuscate. flavus (Fabricius) (in part) (p. 81 Pterostigma in dorsal view centrally black; distal sclerite absent (Figs 291-293) and lower corner of epicnemial carina abruptly angled, slightly raised (Fig. 186). A reddish species with gaster with at least tergites 3+ black; central sclerite varying in form from circular to I-shaped, see Figs 291-293....... monticola (Cameron) (in part) (p. — Pterostigma in dorsal view brownish or orange; either with distal sclerite present, or if absent then with lower corner of epicnemial carina evenly rounded and not prominent (Fig. 185). 82 Mandibles twisted 70° or more (Fig. 187); fore wing with Rs+2r almost straight, incrassate near proximal 0.3 (Fig. 294); marginal cell broadly glabrous proximally. madrigalae sp. n. (p. — Mandibles twisted 45° or less (Fig. 188); fore wing with Rs+2r slightly sinuous, with proximal 0.3—0.5 slightly broadened (Figs 295, 296); marginal cell proximally usually evenly hirsute, at Meio AICO UV SONAL LIS real su ofere cabs) (6s ceeauSli (ere i6in o, si uo,iviel Gt v wlehe e/a eineallg dies sje ae o'¥in oe Seeing os 83 Fore wing with central sclerite elongately oval, characteristically with longest axis subtending an angle of 45—70° to Rs+2r; cu-a proximal to base of Rs&M by 0.2 or more times its own length (Fig. 295); mesopleuron almost always with a band of quite coarse rugose-striate sculpture extending from lower proximal corner to near centre (Fig. 189) xanthocarpus (Szépligeti) (p. — Fore wing with central sclerite strongly pigmented, broadly oval to subcircular; fore wing with cu-a more or less opposite base of Rs&M (Fig. 296); mesopleuron ventrally punctate, at most with fine, indistinct wrinkling (Fig. 190) ..................-....-- duckworthi sp. n. (p. 84 Fore wing with fenestra bearing 2 separate central sclerites (Fig. 297)..... devriesi sp. n. (p. SECeERUIneawatntenestra bearing 1 centrallsclerite 6... 2.6566. ccc ene cee eee ce we bene 85 Fore wing with fenestra partially obliterated by an area of fine hairs which extends beyond the distal sclerite, and almost reaches the central sclerite (Fig. 298)....... donahuei sp. n. (p. — Fore wing with fenestra glabrous, without an area of pubescence between the distal and central RERIVESY (lal OSTA 99__S02) Memernene aetrtert seit -isaeee mot lels wre Csi ceeds cde vue sea ho nie esis 86 Mandible barely narrowed, but twisted about 80°, subequally bidentate, and with teeth strongly compressed (Fig. 192). Large species, fore wing length 19-26 mm; 3rs-m 0.80 or more times as long as abscissa of IM! OWCHESN ZA CUTPENITO STO (Ore ae Oe OO SUD BRNe FON 6 ROONe co eMror simoni sp. n. (p. — Mandible weakly to very strongly narrowed, variously twisted, usually with upper tooth dis- tinctly the longer, and generally without both teeth compressed, never exactly as above... 87 Mandibles quite short, very stout, strongly narrowed and twisted outwards so that upper tooth is forwards; upper mandibular tooth more than 2 times length of the lower tooth (Fig. 191). Central sclerite large, obovate, weakly sclerotized (Fig. 300).......... orosii sp. n. (p. — Mandibles moderately to very long, usually evenly narrowed, twisted so that lower tooth is forwards; upper mandibular tooth less than 2 times length of the lower tooth............ 93 226) 227) 79 80 81 228) 230) 243) 87 246) 94 IAN D. GAULD Figs 177-182 Enicospilus species. 177, 178, metapleuron and propodeum, lateral view; (177) E. lau- renae; (178) E. vegai. 179, 180, face; (179) E. karrensis; (180) E. stevensi. 181, 182, metapleuron and propodeum, lateral view; (181) E. lovejoyi; (182) E. hacha. OPHIONINAE OF TROPICAL MESOAMERICA 88 Clypeus medioapically produced and slightly out-curved so its apical margin is more or less U-shaped in front view (Fig. 193); epicnemial carina laterally not reaching onto mesopleuron I fader rt Tt ciisreintals «os. + + « bine Pie OGaiew dyaie ad ves ome bima sp. n. (p. — Clypeus not medioapically produced, in front view weakly arcuate (Fig. 215); epicnemial carina SEMEN HE UMGPIEOT TALCEAIN bares) ice oo = = fas win ike Wiese s onion ews ies nlev eee aueeee we Teil part Of antenna (at least dorsally) black ......0.......0. cece ccc eee ESELoOximal part of antenna dark brown, orange or yellow............2.0c ccm ee cece seewece 90 Fore wing with Rs+2r strongly sinuous, so that in part its anterior margin is almost parallel to fore margin of wing; central sclerite positioned in anterodistal corner of fenestra (Fig. 302). barbarae sp. n. (p. — Fore wing with Rs+2r weakly sinuous, its anterior margin evenly convergent to fore margin of wing; central sclerite subcentral or positioned mediodistally (Figs 303-306).............. 91 Fore wing with a band of hair along anterior margin of the fenestra, below Rs+2r (Fig. 303). Proximal sclerite elongately oval, often weakly sclerotized; 1st subdiscal cell very narrow. fosteri sp. n. (p. — Fore wing without a band of hair along anterior margin of fenestra, below Rs+2r (Figs 304-306) 92 Propodeum long, with posterior area almost smooth, at most with only vestigial rugae (Fig. 195). Scutellum with lateral carinae present only anteriorly; central sclerite obovate; fore wing with first abscissa of Cula less than 1.05 times as long as first abscissa of Cul NE errs ge aT a... Uaia diad daed na a cornifuscus nom. n. (p. — Propodeum shorter, with posterior area strongly sculptured, with distinct rugae............ 93 Posterior transverse carina of mesosternum broadly incomplete centrally, the gap between the two ends at least as wide as thickness of hind trochantellus...................-00eeeeee — Posterior transverse carina of mesosternum more or less complete, at most only narrowly Perera OSERCANSCH TALIM Cleans MOTE Y SL Aner ere « « « « +» aBiiles eis a eicivirel aisiaphe slelejeiiw se iaie saad ae'ers « 94 Scutellum rounded, with lateral longitudinal carinae only discernible as an anterior vestige (Fig. 198); mandible twisted 40° or more, with lower tooth swollen and with a sharp, convex ventral Tle (Tee, JMOL adhe hie alone ira Iai ad Aa ae ieee i aie a a kelloggae sp. n. (in part) (p. g g p — Scutellum slightly flattened centrally, with distinct lateral longitudinal carinae extending more than 0.6 of its length (as in Fig. 215); mandible twisted 20°-or less, with lower tooth neither moticeably swollen nor with a sharp, convex ventral edge .................0.eeeeeeaees 95 Fore wing with 3rs-m short, about 0.35 times as long as the abscissa of M between 3rs-m and 2m-cu (Fig. 305); lower face broad, 0.83 times as broad as long; anterior transverse carina of NN REE TMUCIO CEO cogs nie. MRA EM 4 6 6 +h cedglee ea whose aa catemacoi sp. n. (p. — Fore wing with 3rs-m quite long, 0.75 times the length of abscissa of M between 3rs-m and 2m-cu (Fig. 306); lower face quite narrow, 0.74 times as long as broad; anterior transverse carina of PESMOCEUMMOLOAGIVMMCOMIPIElG. Can ccdwce-. +--+ senccreteetehecneeeee gabrieli sp. n. (p. 96 Upper surface of fore wing with distal abscissa of 1A bearing a row of long, and very fine hairs 2 LSE eo CE all eee, ON nn 2 on cre luisi sp. n. (p. — Upper surface of fore wing with distal abscissa of 1A glabrous, without long fine hairs (Figs 309- 314) 97 Fore wing with anterior corner of discosubmarginal cell with a cluster of hairs on membrane near base of Rs+2r, sometimes with this area of hairs separated by a glabrous band from rest of cell (cf. Fig. 307). Central sclerite small and slender, similar to that of preceding species (Fig. 307) opleri sp. n. (p. — Fore wing with anterior corner of discosubmarginal cell glabrous, this glabrous area confluent with fenestra (Figs 310-314) or, if hairs are apparently present, then they actually arise from vein, not membrane (Fig. 309) 98 Scutellum with lateral carinae present only on anterior half and posterior transverse carina of mesosternum centrally produced forward so entire carina has the form of a chevron (Fig. 132): Fore wing with central sclerite very small and indistinctly delineated (Fig. 309). guindoni sp. n. (p. 95 247) 89 90 114 248) 9] 249) 02 259) 254) 98 256) 96 IAN D. GAULD Figs 183-188 Enicospilus species. 183, 184, metapleuron and propodeum, lateral view; (183) E. xanthostigma; (184) E. fernaldi. 185, mesopleuron, E. duckworthi. 186, alitrunk, lateral view, E. monticola. 187, 188, mandibles; (187) E. madrigalae; (188) E. xanthocarpus. OPHIONINAE OF TROPICAL MESOAMERICA — Scutellum usually with lateral carinae extending 0.7 or more of its length, or if with these carinae very short, then posterior transverse carina of mesosternum straight (or rarely incomplete centrally); posterior transverse carina of mesosternum otherwise various, usually not pro- eM UALCCENITAlly (tO SS)iis. ccna... o).c +.» MAE a os Wiad ams ced eunte sis os ace cae 99 Fore wing with central sclerite very slender, more or less I- shaped, with longest axis perpen- MME MME CUE ISIOL VEINS 1-27 (OS SOG 1)... . . .p.cetnetete viele mee clan se sae 06 Sinise die tue ow see — Fore wing with central sclerite from narrowly comma-shaped, to subcircular or irregularly oval, if slender then with longest axis subparallel to Rs+2r (Figs 312-322).................... 100 Mandibles strongly narrowed, twisted about 70°; fore wing with distal sclerite absent; marginal cell proximally uniformly hirsute (Fig. 310); mesoscutum unicolorous reddish brown; pterostigma black. Mietapleuromistronely striate. «2... 2.0.56. e cee we monticola (Cameron) (in part) (p. — Mandibles evenly tapered, twisted about 45° or less; fore wing with distal sclerite discernible; marginal cell proximally glabrous or notably more sparsely hirsute than it is centrally (Fig. 311); mesoscutum with three longitudinal black marks; pterostigma brownish yellow. leoni sp. n. (p. 101 Fore wing with abscissa of Cul between Im-cu and Cula more than 0.60 times as long as Culb (Fig. 339). Central sclerite large, distinctly delineated; distal sclerite complete, confluent with prox- PLAN SILEIMIG.S 205 cece Ate rR coc er masoni sp. n. (in part) (p. — Fore wing with abscissa of Cul between Im-cu and Cula less than 0.50 times as long as Culb SE) i RRR oS SSR SP egg 102 Lower tooth of mandible swollen, very much stouter than the upper and ventrally with a convex margin (Fig. 199). Metapleuron usually smooth and punctate; distal sclerite discernible (Fig. 312). kelloggae sp. n. (in part) (p. — Lower tooth of mandible not swollen, of similar stoutness to the upper one, or if slightly eee Hien mever With VENiral MlATBiN.CONVES ofa ses c ssn ssc rent tcc er eee sete seus 103 Propodeum with posterior area weakly sculptured, laterally almost coriaceous, and witha single ‘median longitudinal ridge; frons with a black mark extending from median ocellus to between OSSES GIP EITETIE SG pytrc 0 OO ne ERE Oe CRE RRR 7c COME Ch aCe mengoi sp. n. (p. — Propodeum coarsely rugose, often with rugae posteriorly concentrically striate, or if weaker then always with at least 3 longitudinal ridges centrally; frons generally unicolorous yellowish 104 Distal sclerite strong, separated from proximal sclerite; proximal sclerite characteristically eee aa Ome I ER PI eR crdcest ... s . « Rerad nent ad bwlesmcaiiee he & haberi sp. n. (p. — Distal sclerite absent or weak, or if quite strongly sclerotized then it is confluent with the proximal sclerite; proximal sclerite variously triangular, not hastate (Figs 315-325)....... 105 Flagellum with central segments exceptionally elongate, 2.6- 2.8 times as long as broad. Meso- and metapleurae regularly punctate, though rarely the latter may have isolated rugae dorsally; lateral longitudinal carina of propodeum complete; fore wing with central sclerite variously comma-shaped (Figs 316-318); male with lateromedian rows of close erect hairs on sternite 7 and anterior margin of sternite 8, rest of sternite 8 glabrous (Fig. 200). lacsa sp. n. (in part) (p — Flagellum with central segments normally long, 2.4 or less times as long as broad .......... 106 Central sclerite subtriangular, with distal side acutely angled and distal sclerite absent (Fig. 319); male with lateromedian rows of close erect hairs on sternite 7 and anterior margin of sternite 8, rest of sternite 8 glabrous, sternite 9 with fine decumbent pubescence (cf. Fig. 200). echeverri sp. n. (in part) (p. — Central sclerite generally oval to D-shaped, rarely crescentic, or if very rarely subtriangular then with distal sclerite distinct; distal sclerite otherwise present or absent (Figs 320-325); Male nwith lone stout erect hairsion stemmites J—9) «100. 52. s nce oc eee sess eee sss ems t 107 Central sclerite narrowly crescentic (Fig. 315). Metaplenromucpularly punctate (Fig. 135). ... 20... cece ee ce eee brenesiae sp. n. (p. — Central sclerite usually either oval or D-shaped, or very rarely subtriangular, never crescent- Sede GEL DE Gal <2 27 est ot ht eis bate ting anit lg a a ie ee 97 233) 257) 292) 102 259) 103 260) 104 261) 105 . 262) 106 300) 98 IAN D. GAULD Figs 189-194 Enicospilus species. 189, 190, mesopleuron; (189) E. xanthocarpus; (190) E. duckworthi. 191, 192, mandibles; (191) E. orosii; (192) E. simoni. 193, 194, E. bima; (193) face; (194) mesopleuron. . OPHIONINAE OF TROPICAL MESOAMERICA 108 Mesoscutum more or less uniformly reddish brown; alitrunk and tergites 1-2 of gaster reddish RE AS AGE er, Ge...» » » er ete saaiaiane sin 9 siglaiy Dawaieg ad oie Dame ine ss — Mesoscutum almost always with dark brown or black longitudinal vittae which are separated by pale yellowish brown lines; alitrunk and much of gaster brownish yellow to dirty yellowish brown, at most with posterior tergites weakly infuscate, sometimes with distal most blackish 109 Dorsal surface of posterior part of propodeum concentrically striate (Fig. 201); metapleuron weakly rugulose-punctate posterodorsally (Fig. 202); distal sclerite more or less absent NT ee ini, Sn aia nak + i+ a dgeanne med acneedeic forsythei sp. n. (p. — Dorsal surface of posterior part of propodeum rugose (Fig. 203); metapleuron posterodorsally coarsely rugose (Fig. 204); distal sclerite complete (Fig. 321) .... fogdenorum sp. n. (p. 110 Fore wing with 3rs-m less than 0.50 times as long as abscissa of M between 2m-cu and 3rs-m (Fig. 322); smaller species, fore wing length less than 16 mm........ burgosi sp. n. (in part) (p. — Fore wing with 3rs-m more than 0.52 times as long as abscissa of M between 2m-cu and 3rs-m aiicais2s—3525)2 large species, fore wing length 17+ mm... .. cc. eee ce sees cee cet eecus 111 Mandible apically strongly twisted through 50° or more; propodeum with posterior transverse carina represented laterally by a distinct raised crest (Fig. 134) ...................0000- — Mandible apically weakly twisted, turned through 45° or less (Fig. 208); propodeum without any trace of a crest formed from a vestige of the posterior transverse carina ................. 112 Mandible with a distinct ridge extending from the upper proximal corner to base of upper tooth (Figs 136, 217); gaster slender, tergite 2 in profile more than 5 times as long as posteriorly ey eR RG 8 S20 RM erasi sp. n. (p. — Mandible without a ridge extending from the upper proximal corner to the base of the upper tooth; gaster moderately stout, tergite 2 in profile 4 or less times as long as posteriorly deep. Jjesicae sp. n. (in part) (p. 113 Fore wing with Rs+2r virtually straight (Fig. 324); marginal cell proximally rather sparsely hirsute; gaster slender, tergite 2 more than 6.0 times as long as posteriorly deep; propodeum Pee ROCOUSAIV ITRESMATIV TUSOSCLc .cte. eens ee ce bree se ecne ee eae galilea sp. n. (p. — Fore wing with Rs+2r distinctly bowed centrally (Fig. 325); marginal cell proximally almost evenly hirsute; gaster quite stout, tergite 2 in profile less than 5.0 times as long as posteriorly deep; propodeum dorsally with striae subconcentric posterolaterally, usually extended Rr AMELIE INT CHUTE (EEO AL) casts ca ain aie a «02h ichaie, ¥ esapnisimardin 800) m8» 0905 hubbelli sp. n. (p. 114 Fore wing with central sclerite elongately linear, positioned in centre of fenestra, and with longest axis parallel to Rs+2r (Fig. 326). Alitrunk laterally polished and rather weakly sculptured; an extremely common and DES) SPECS 4 ielenead Cn Oat atne nen RR oo cn flavoscutellatus (Brullé) (p. — Fore wing with central sclerite subcircular, oval or crescent-shaped, or if linear, then with Mmseamei si perpendicular to usar (PIGS 327-330) ca... ce pce sree ese en eect seew ans REM SHOT, TWIStEO 70° OF MONG. ... 2... .. «cece cece ence ba ntcccuenvevvsccuces Eeeancibles moderately long to long, twisted 10-60°.............. cece cee cece ee eect eenes 116 Fore wing with proximal corner of marginal cell uniformly hirsute; anterior part of discosub- marginal cell hirsute between proximal sclerite and anterior corner; fenestra very long, its distal end reaching nearly to base of Rs (Fig. 327); a more or less uniformly pale yellow TN MMO TR EES reo RRC UCU u ns babe aay woldai sp. n. (p. — Fore wing with proximal corner of marginal cell glabrous or rather sparsely hirsute; anterior part of discosubmarginal cell with few or no hairs between proximal sclerite and anterior corner; distal margin of fenestra separated from base of Rs by more than length of M between Rs and 3rs-m (Figs 328-330); pale yellow, but often with mesoscutum and/or terminal segments of gaster marked with black or dark brown............... 0... c eee eee cence 117 Posterior transverse carina of mesosternum broadly incomplete centrally or with central portion emVMIOSCIN Cn aaa ate UMA e tins thant cee ce ae oa cet ee sree ees fees — Posterior transverse carina of mesosternum complete, with central part strongly raised ..... 118 Proximal sclerite more or less triangular, with proximal side forming an angle (Fig. 328); fore wing with 3rs-m more than 0.42 times as long as abscissa of M between 2m-cu and 3rs-m; head posteriorly very short, so that the hind ocelli are separated from occipital carina by less than 99 109 110 265) 266) 267) 111 112 113 269) 218) 270) 271) 272) 1S 116 120 275) 117 118 119 100 IAN D. GAULD Figs 195-200 Enicospilus species. 195, propodeum, dorsal view, E. cornifuscus. 196, 197, E. leoni; (196) mandibles; (197) metapleuron and propodeum, lateral view. 198, 199, E. kelloggae; (198) scutellum; (199) mandibles. 200, subgenital plate of male, E. lacsa. OPHIONINAE OF TROPICAL MESOAMERICA their smallest diameter (Fig. 128); propodeum with posterior area finely and weakly SEES) CNSR Ms OCP A A ae en eee liesneri sp. n. (p. — Proximal sclerite more or less quadrate, with proximal side flat (Fig. 329): fore wing with 3rs-m less than 0.42 times as long as abscissa of M between 2m-cu and 3rs-m; head posteriorly only moderately short, so that hind ocelli are separated from occipital carina by more than their own smallest diameter (cf. Fig. 129); propodeum with posterior area rugose, reticulate, or TEM od SNe hee ola ans - 5 « dora Vie weiss 6% 40:2 2.0,0 x marini sp. n. (p 119 Dark mark on mesopleuron extending ventrally along pleural suture below level of episternal scrobe, then usually broadly across pleuron (Fig. 125); second segment of gaster usually with laterotergite pendant; hind wing with first abscissa of Rs weakly bowed (Fig. 127); fore wing generally with distal sclerite either narrowly separated from proximal sclerite or with them joined by a weak bridge (Fig. 330); mesoscutum usually posteriorly Te aS oye. i. f ce c's in sho asin «> « ye eae anh ete haere es sanchezi sp. n. (p. — Dark mark on mesopleuron extending from upper corner to episternal scrobe and not extending ventrally along pleural suture (Fig. 124); second segment of gaster with the laterotergite folded under margin of tergite; hind wing with first abscissa of Rs more or less straight (Fig. 126); fore wing with distal sclerite confluent with proximal sclerite (Fig. 331); mesoscutum maunentee dark vittae present anteriorly...............ccereseacens pescadori sp. n. (p 120 Hind wing with cu-a greater than 0.4 times the length of the first abscissa of Cul which is bowed (Fig. 142); fore wing with cu-a subopposite the base of Rs&M or proximal to it by its own thickness, and with distal sclerite entirely absent (Figs 332, 333). SATAN MCMRACI AN AIC CATIDUGAM SIIECICS . iia cis canis cecr ten waeces suc svees wees — Hind wing with cu-a less than 0.4 times the length of the first abscissa of Cul which is generally more or less straight (Figs 141, 143, 144); fore wing either with cu-a proximal to base of Rs&M by 0.1 or more of its length, or with distal sclerite present, or both 121 Fore wing with marginal cell proximally uniformly hirsute; Rs+2r not incrassate centrally (Fig. 332); abscissa of 1m-cu distal to bulla less than 0.90 times as long as 3rs-m; metapleuron very strongly convex, polished and finely punctate (Figs 145, 211); male with sternites 7—9 bearing Senttered, long, fine semidecumbent hairs. ................e00- howdenorum sp. 0. (p. — Fore wing with marginal cell proximally very sparsely hirsute; Rs+2r strongly incrassate centrally (Fig. 333); abscissa of 1m-cu distal to bulla more than 0.95 times as long as 3rs-m; metapleuron moderately convex, weakly polished coarsely punctate and generally with diagonal rugae discernible (Fig. 146); male with sternite 7 with lateral longitudinal rows of long close erect hairs, the sternite centrally glabrous; sternite 8 more or less glabrous; sternite iveanine short sine decumbent pubescence. «.....0.-2 0000+ sess cnve- sondrae sp. n. (p 122 Marginal cell of fore wing proximally broadly glabrous, at most with isolated scattered hairs Pmenion periphery (Pips 334-336, 352)... ck ccc ccc seer caer eccesenesecenseens — Marginal cell of fore wing not broadly glabrous proximally, either uniformly hirsute, or more sparsely hirsute (Figs 337-350), or occasionally with a glabrous area adjacent to Rs+2r, but if such an area is present then with a distinct area of hairs extending to corner centrally [Intermediate specimens will key either way] 123 Hind wing with only 4 distal hamuli on vein R; fore wing with cu-a very slightly distal to base of Rs&M (Fig. 334); mesopleuron with a large dark mark posteriorly that extends down the pleural suture to the level of the episternal scrobe (Fig. 123)........... oduberi sp. n. (p — Hind wing with 5 or more distal hamuli on vein R; fore wing with cu-a proximal to base of Rs&M (Figs 335,336); mesopleuron with, at most a small dark mark in upper posterior corner ... 124 Fore wing with distal sclerite strong, broadly separated from the proximal sclerite (Fig. 335); central sclerite oval; abscissa of Cul between Im-cu and Cula 0.20-0.37 times as long as Cul1b; hind trochantellus unusually long, dorsally projecting well beyond apex of trochanter (Fig. 139); small species, fore wing length 9-12 mm.................. gallegosi sp. n. (p — Fore wing with distal sclerite weak or indistinct, when visible confluent with the proximal sclerite (Fig. 336); central sclerite more or less crescentic; abscissa of Cul between 1m-cu and Cula more than 0.40 times as long as Cu1b; hind trochantellus short, not or barely projecting beyond the apex of the trochanter (Fig. 140); larger species, fore wing length 17+ mm.... 125 Propodeum with anterior transverse carina usually centrally incomplete, sometimes absent; metapleuron punctate, more or less evenly convex; gaster never narrowly black dorsally. dispilus (Szépligeti) (few specimens) (p. 101 276) . 278) 279) . 281) 121 122 . 284) 123 126 a5) 124 . 286) 125 296) 102 IAN D. GAULD — Propodeum with anterior transverse carina centrally complete, at most only obsolescent at lateral extremities; metapleuron posterodorsally rugose-striate, usually flattened ventrally (Fig. 210); gaster usually narrowly black-marked dorsally ............. parkeri sp. n. (p 126 Central sclerite small and more or less circular, its maximum diameter subequal to maximum thickness of Rs+2r (Fig. 337) and with distal sclerite present......... ulfstrandi sp. n. (p — Central sclerite moderately large to large, often oval or crescent-shaped, if subcircular then its diameter much greater than thickness of Rs+2r (Figs 338-343) or, if subcircular and relatively small’, then’ distal sclerite absent-(Fig, 344)... 50 5 seers = teeta tate ae 127 Fore wing with abscissa of Cul between 1m-cu and Cula more than 0.55 times as long as Culb (Pigs 3385-339) eee setae wre pore e Sle au «0m 5» sills Cae oie «eee — Fore wing with abscissa of Cul between 1m-cu and Cula less than 0.50 times as long as Culb (Figs: 340-352). oo cia. eca aris ew meye, 6 blbselvistee « » « « «0 » ShGlmlinn eectnitvinie ete Sheela ea 128 Central sclerite crescent-shaped (Fig. 338); hind wing with first abscissa of Rs distinctly bowed (Fig. 144); wings strongly infumate; gaster with tergites 3-7 posteriorly and ventrally dark, centrally pallid so the gaster has a mottled appearance.............. georginae sp. n. (p — Central sclerite elongately D-shaped (Fig. 339); hind wing with first abscissa of Rs more or less straight (Fig. 141); wings hyaline; gaster with posterior tergites uniformly infuscate so that it does notappear to be mottled. ic -...9s.25,.... «cane ae masoni sp. n. (in part) (p 129 Fore wing with central sclerite rather narrow and comma- shaped (Fig. 340). Vein Rs+2r distinctly sinuous; metapleuron diagonally striate (Fig. 212); upper mandibular tooth slender; distal sclerite usually absent. . 75: .2-. «29s eee kleini sp. n. (p — Fore wing with central sclerite oval, hastate, circular, D- shaped or sometimes crescentic or very broad and commia- shaped (Figs'341-352):. 2... 00.06 occ nee ease n ce ieee 130 Interocellar area distinctly and rather uniformly brownish, darker brown than vertex; central sclerite elliptical; distal sclerite quite long (Fig. 341)....... flavus (Fabricius) (in part) (p — Interocellar area yellowish orange, concolorous with vertex, at most slightly infuscate only between hind ocelli; central sclerite oval, subcircular, subtriangular hastate or crescentic; distal sclerite usually regularly triangular (Figs 342-352) .............. ee eeeeeee icsecteatete 131 Distal sclerite distinct, strongly sclerotized and narrowly separated from proximal sclerite Cte 2 i ne ae ey ee ee ee! ee hemicrescellae sp. n. (p — Distal sclerite absent, or if discernible (and it may be quite strongly sclerotized) then it is broadly confluent with the distal sclerite (Figs 343-352) 132 Hind trochantellus dorsally 0.2-0.4 times as long as broad (Fig. 138); metapleuron coriaceous- striate (Fig. 205). Distal sclerite absent (Fig. 343); terminal sternites of male gaster with scattered long erect TYAS o's etree eters teate vo cae ere hc MRE cscs Lae VET ue eee burgosi sp. n. (in part) (p — Hind trochantellus dorsally 0.1 or less times as long as broad (Fig. 140); metapleuron punctate, punctostriate or partially mugose (Fic. 213)... ec ae eee ene nite eee 133 Fore wing with central sclerite subcircular, D-shaped, comma- shaped, or less commonly crescentic (Figs 344, 347-352); male with sternites 7-9 with quite dense long erect pubescence (Pig: 214)... 0:5 6. cies ies wns srordse Bu pale a Rie by « Bly rake eres ele Seo el Seeger — Fore wing with central sclerite more or less hastate, with inner side concave, and with outer side strongly curved or distally pointed (Figs 345, 346); male with sternite 7 and anterior margin of sternite 8 bearing lateromedian rows of close short hairs, remainder of sternite 8 glabrous; stemnite)9 with fine decumbent pubescence: (Gis Eig, 200) he ray eee 134 Distal sclerite absent; proximal sclerite hastate; central sclerite subcircular, indistinctly defined 287) _ 288) 127 128 129 _ 291) _ 292) _ 293) 130 230) 131 _ 294) 132 267) 133 134 135 (Fig.(344):... sss1idenanstdbve nh ee, eae iol tiveness aie ceciliae sp. n. (p. 295) — Distal sclerite present; proximal sclerite simply triangular; central sclerite various, if subcircular then distinctly delineated (Figs 347-352). ...... dispilus (Szépligeti) (most specimens) (p. 135 Central flagellar segments very slender, 2.5—3.0 times as long as broad; fore wing with central sclerite distally rounded, fenestra of moderate length (Fig. 345); lower face generally less than!0}7O0timestas broadtasilongas a aaeeee ore ae eee eee lacsa sp. n. (in part) (p. — Central flagellar segments not exceptionally slender, 2.0—2.2 times as long as broad; fore wing with central sclerite distally acutely angled, fenestra unusually long (Fig. 346); lower face generally greater than 0.70 times as broad as long........... echeverri sp. n. (in part) (p 296) 262) _ 300) OPHIONINAE OF TROPICAL MESOAMERICA 103 | _ Figs 201-206 Enicospilus species. 201, 202, E. forsythei; (201) propodeum dorsal; (202) metapleuron _ and propodeum, lateral view. 203, 204, E. fogdenorum; (203) propodeum dorsal; (204) metapleuron and propodeum, lateral view. 205, metapleuron and propodeum, lateral view, E. burgosi. 206, alitrunk, lateral view, E. flavoscutellatus. 104 IAN D. GAULD Figs 207-212 Enicospilus species. 207, 208, E. hubbelli; (207) propodeum dorsal; (208) mandibles. 209, propodeum dorsal, E. liesneri. 210-212, metapleuron and propodeum, lateral, view; (210) E. parkeri; (211) E. howdenorum; (212) E. kleini. OPHIONINAE OF TROPICAL MESOAMERICA 105 & Se Be 213 fee 214. Figs 213-218 Enicospilus species. 213, 214, E. dispilus; (213) metapleuron and propodeum, lateral view; (214) subgenital plate and aedeagus of male. 215, scutellum, EF. flavoscutellatus. 216, hind tarsal claw of female, E. maritzai. 217, mandibles, E. erasi. 218, hind tarsal claw of female, E. exoticus. 106 IAN D. GAULD Figs 219-226 Enicospilus species, central part of right fore wing in dorsal view; (219) E. bozai; (220) E. enigmus ; (221) E. exoticus; (222) major; (223) E. clarkorum; (224) E. glabratus; (225) E. alvaroi; (226) E. chiriquensis. OPHIONINAE OF TROPICAL MESOAMERICA 107 228 Figs 227-234 Enicospilus species, central part of right fore wing in dorsal view; (227) E. mayi; (228) E. robertoi; (229) E. brevis; (230) E. scuintlei; (231) E. mexicanus; (232) E. abelardoi; (233) E. hall- wachsae; (234) E. cameronii. 108 IAN D. GAULD Figs 235-242 Enicospilus species, central part of right fore wing in dorsal view; (235) E. gamezi; (236) E. texanus ; (237) E. cushmani, (238) E. halffteri; (239) E. sarukhani; (240) E. aktites; (241) E. gomezpom- pai; (242) E. lebophagus. OPHIONINAE OF TROPICAL MESOAMERICA 109 Figs 243-250 Enicospilus species, central part of right fore wing in dorsal view; (243) E. ugaldei; (244) E. umanai; (245) E. quintanai,; (246) E. americanus; (247) E. tenuigena; (248) E. peigleri; (249) E. neotropicus; (250) E. doylei. 110 IAN D. GAULD Figs 251-258 Enicospilus species, central part of right fore wing in dorsal view; (251) E. purgatus; (252) E. luquillo; (253) E. martae; (254) E. columbianus; (255) E. cubensis; (256) E. carlota; (257) E. dajaboni; (258) E. porteri. OPHIONINAE OF TROPICAL MESOAMERICA 111 260 Figs 259-266 Enicospilus species, central part of right fore wing in dorsal view; (259) E. lupemejia; (260) E. trilineatus; (261) E. dirzoi; (262) E. masneri; (263) E. randalli; (264) E. cressoni; (265) E. laurenae; (266) E. vegai. by IAN D..GAULD 271 | 272 Figs 267-274 Enicospilus species, central part of right fore wing in dorsal view; (267) E. baltodanorum, | (268) E..carri; (269) E. colini; (270) E. maculipennis; (271) E. guatemalensis; (272) E. karrensis; (273) E. stevensi; (274) E. pamelae. OPHIONINAE OF TROPICAL MESOAMERICA 113 Figs 275-282 Enicospilus species, central part of right fore wing in dorsal view; (275) E. randalli; (276) = vilmari; (277) E. estradarum; (278) E. cubensis; (279) E. teodorae; (280) E. maritzai; (281) E. exoticus; (282) E. jesicae. 114 IAN D. GAULD 289 Figs 283-290 Enicospilus species, central part of right fore wing in dorsal view; (283) E. corcovadoi; (284) E. lovejoyi; (285) E. hacha; (286) E. xanthostigma; (287) E. cepillo; (288) E. persimilis; (289) E. fernaldi; (290) E. flavus. OPHIONINAE OF TROPICAL MESOAMERICA DS 292 Figs 291-298 Enicospilus species, central part of right fore wing in dorsal view; (291-293) E. monticola; (294) E. madrigalae, (295) E. xanthocarpus; (296) E. duckworthi; (297) E. devriesi; (298) E. donahuei. 116 IAN D. GAULD 300° Figs 299-306 Enicospilus species, central part of right fore wing in dorsal view; (299) E. simoni; (300) E. orosit; (301) E. bima; (302) E. barbarae; (303) E. forsteri ; (304) E. cornifuscus; (305) E. catemacot; (306) E. gabrieli. OPHIONINAE OF TROPICAL MESOAMERICA 117 Figs 307-314 Enicospilus species, central part of right fore wing in dorsal view; (307, 308) E. luisi; (309) E. guindoni; (310) E. monticola; (311) E. leoni; (312) E. kelloggae; (313) E. haberi; (314) E. mengoi. 118 IAN D. GAULD Sa Bigs as sake a Figs 315-322 Enicospilus species, central part of right fore wing in dorsal view; (315) E. brenesiae; (316— 318) E. lacsa; (319) E. echeverri; (320) E. forsythei; (321) E. fogdenorum; (322) E. burgosi. OPHIONINAE OF TROPICAL MESOAMERICA 119 . SA SL Figs 323-330 Enicospilus species, central part of right fore wing in dorsal view; (323) E. erasi; (324) E. galilea; (325) E. hubbelli; (326) E. flavoscutellatus; (327) E. woldai; (328) E. liesneri; (329) E. marini; (330) E. sanchezi. 120 IAN D. GAULD Figs 331-338 — Enicospilus species, central part of right fore wing in dorsal view; (331) E. pescadori; (332) E. howdenorum; (333) E. sondrae; (334) E. oduberi; (335) E. gallegosi; (336) E. parkert; (337) E. ulfstrandi; (338) E. georginae. OPHIONINAE OF TROPICAL MESOAMERICA (eal | : ae aes A specimen in the USMN is purported to have been reared from Hylesia sp. (Lepidoptera: Saturniidae). MATERIAL EXAMINED Holotype & (Eurycamptus calcator Morley), Brazil: Ega (= Tefé) on the Amazon (Bates) (BMNH). Holotype 2 (Ophion chiriquensis Cameron), Panama: Volcan de Chiriqui, 1700-2000 m (BMNH). Belize: 1 9, Toledo, Columbia Forest Station, vii.1968 (Hasse) (BMNH). Brazil: 1 2, Nova Teutonia, Santa Catarina, xii.1952 (Plaumann) (TC); Mato Grosso; 1 CO’, Sinop, 12°31’S 55°37’°W, x.1975 (Alvarenga) (TC); 1 @, dry forest, 12°50’S 51°47’W, ix.1938 (Richards) (BMNH). Costa Rica: Alajuela Prov.: 3 O', 4 Q, Finca Campana, 5 km NW. Dos Rios, 750 m, iii.1985 (Janzen & Hallwachs) (BMNH); 1 Q, Finca San Gabriel, 3 km W. Dos Rios, 850 m, vi.1986 (Gauld) (BMNH): Guanacaste Prov.: 1 9, Rincon de la Vieja, 900 m, iii.1984 (Gauld) (BMNH); 2 C’, 8 9, Santa Rosa National Park, 300 m, xi.1981, i-ii. 1982, ii,iv-v.1983, iii & vi.1984 (Janzen & Hallwachs) (BMNH); 1 2, Volcan Cacao, Estacion Mengo on SW. side, 1100 m, vi.1987 (Janzen) (BMNH); 1 2, Volcdn Orosi, Casa Mariksa [Maritza], 700 m, vii. 1986 (Gauld) (BMNH): Heredia Prov.: 2 9 , Puerto Viejo de Sarapiqui, Finca La Selva, xi.1986 (Janzen & Hallwachs) (BMNH): Limon Prov.: 2 0’, 1 2, Cerro Tortuguero, N. edge Tortuguero National Park, 0-100 m, v.1984 (Janzen & Hallwachs) (BMNH). Guatemala: 1 9, Cayuga, iv.1915 (Cush) (USNM). Honduras: 1 @, Trujillo, vii.1968 (Dozier) (FSCA). Mexico: Chiapas: 1 9, 32 km N. Huixtla, 1000 m, vi.1969 (CNC); 1 9, Palenque, viii.1969, (Kelton) (CNC). Panama: 2 9, Barro Colorado Island, vii.1963 (Cavagnaro & Irwin) (CAS); 1 9, same locality, iv.1965 (Duckworth & Duckworth) (USNM); 1 2, same locality, x-xi.1941 (Zetek) (USNM); 4 2, same locality, iv, vi, vii, xi.1978 (Wolda) (RNH); 10,2 2, Barro Colorado Island, Gatun Lake, iv.1979 & i-iv.1983 (Wolda) (TC); 1 0’, 15 9, Barro Colorado {sland, i-ix, xii. 1984-5 (Wolda) (BMNH); 1 9, Canal Zone, ex Hylesia sp. (Dunn) (USNM); 1 9, Trinidad River, vi.1912 (Busck) (USNM). Surinam: 1 9, Paramaribo, v.1944 (Geijskes) (RNH). Venezuela: 1 2, San Esteban, nr Puerto Cabello, i.1940 (Anduze) (TC). Enicospilus brevis (Morley) (Figs 158, 229) Allocamptus brevis Morley, 1912: 24. Holotype 2, BRAZIL (BMNH) [examined]. Enicospilus brevis (Morley) Townes & Townes, 1966: 174. DEscRIPTION. Mandibles moderately short, proximally strongly narrowed, distally weakly narrowed, apically twisted 45—60° (Fig. 158); upper mandibular tooth compressed, quite stout, 1.2-1.4 times as long as the lower tooth; outer mandibular surface flat, shortly and sparsely hirsute, with a weak proximal concavity. Labrum 0.3—0.4 times as long as broad; malar space 0.3—0.4 times as long as basal mandibular width. Clypeus in profile flat, margin sharp; clypeus in front view 1.5—1.7 times as broad as long, margin truncate apically. Lower face 0.76—-0.91 times as broad as long, polished, finely punctate. Head in dorsal view with genae evenly rounded; posterior ocellus more or less contiguous with eye; FI = 70-75%; occipital carina mediodorsally generally centrally interrupted, rarely complete, ventrally abruptly curved to join hypostomal carina but with lower end evanescent. Antenna moderately long, with (44) 51-56 flagellar segments; 20th segment 1.7—2.0 times as long as broad. Mesoscutum weakly polished, finely coriaceous, in profile abruptly rounded; notauli vestigial. Meso- pleuron polished, the upper part punctate, the lower part punctostriate; epicnemial carina inclined towards anterior margin of pleuron, its upper end evanescent. Scutellum in profile weakly convex, laterally carinate for all of its length; scutellum in dorsal view 1.3—1.4 times as long as anteriorly broad, smooth, with isolated punctures and posteriorly with a few irregular wrinkles. Metapleuron quite strongly convex, weakly to quite strongly polished, finely punctate or punctogranulate, grading to weakly striate posterodorsally; submetapleural carina weakly anteriorly broadened; posterior transverse carina of mesosternum usually centrally interrupted, if complete then centrally rather weak. Propodeum in profile abruptly declivous; anterior transverse carina from complete to present only as acentral vestige, or in the holotype, completely absent; posterior transverse carina absent; anterior area moderately long, at most weakly wrinkled; spiracular area short, smooth; posterior area generally rather weakly wrinkled, often almost smooth, if more coarsely sculptured then wrinkles tend to be subconcentric posteriorly; lateral longitudinal carina usually entirely absent, not joined to spiracular margin by a short carina. OPHIONINAE OF TROPICAL MESOAMERICA 137 Fore wing length usually 17-20 mm, though sometimes as little as 11 mm; discosubmarginal cell as in Fig. 229; AI = 0.79-1.07; CI = (0.39) 0.73-0.95; ICI = 0.35—0.78; SDI = 1.23-1.62; cu-a from subopposite to Rs&M to proximal to it by 0.3 times its own length; marginal cell proximally more or less evenly hirsute; 1st subdiscal cell with anterior 0.2—0.4 and posterodistal corner hirsute. Hind wing with 5—9 hamuli on R1; 1st abscissa of Rs more or less straight, 2nd abscissa weakly arcuate. Fore leg with tibia slightly flattened, with scattered fine spines on outer surface. Mid leg with longer tibial spur 1.3—1.4 times length of the shorter. Hind leg with coxa in profile 1.6-1.7 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.3-2.7 times as long as broad; claws of female with moderately long, close pectinae, those of male similar. Gaster slender; tergite 2 in profile 4.0-4.5 times as long as posteriorly deep, laterotergite generally folded under though it may be partially pendant in some individuals, thyridia elliptical and separated from anterior margin of tergite by about 2.5—3.5 times its own length. Ovipositor slender, its sheath moderately stout. Male with sternites 7-9 bearing scattered fine erect hairs amongst short decumbent pubescence; gonosquama strongly distally tapered, but terminally rounded. Colour generally rather pale brownish yellow with mesoscutal vittae darker brown, the central one often blackish; interocellar area yellowish brown; antenna orange-brown, often infuscate distally; pterostigma orange-brown; wings weakly infumate. VARIATION. Some specimens have tergites 3+ weakly infuscate to blackish and in one of these the anterior gastral segments have a distinct greenish hue. The holotype and a male from Corcovado National Park are unusually small (fore wing lengths 11 and 13 mm respectively) and have only 44 flagellar segments. The antennae of the Corcovado specimen and several others are black. One individual from Braulio Carrillo National Park, Costa Rica is unusual in having CI very small, but in the form of the mandibles and the sculpture of the propodeum it is a typical example of this species. Remarks. Enicospilus brevis is most easily recognized by the possession of two very unusual features — having the mandibles strongly twisted and tapered right to the end, and (usually) having the cubital index (CI) exceptionally large. The majority of specimens are also distinctive in having the propodeal sculpture very weak, and in frequently having the anterior transverse carina discontinuous. The characteristic pubescence of the terminal male sternites and the convex metapleuron are features that E. brevis shares with E. robertoi and E. scuintlei and the three species are probably closely related. BIOLOGICAL INFORMATION. Enicospilus brevis is a widely distributed species whose range extends from Costa Rica south to southern Brazil (Map 6). It has been collected in sites from near sea-level up to 2400 m in Bolivia. In Central America this species is associated with wet forests from sea-level up to an altitude of about 1400 m. It has never been found in dry forests of north-western Guanacaste. MATERIAL EXAMINED Holotype 9, Brazil: Villa Nova on the Amazon (Bates) (BMNH). Non-type material. Bolivia: 1 CO, Yungas, 2400 m, xii.1984 (Pena) (TC). Brazil: 1 0’, 1 9, Santa Catarina, Nova Teutonia, 27°11’S, 52°23’W, x.1935 & ii.1939 (Plaumann) (BMNH). Costa Rica: Alajuela Prov.: 1 0’, 1 , Finca San Gabriel, 16 km ENE. of Quebrada Grande, 650 m, iii. 1983 & v.1984 (Janzen & 10,1 9, Virgen de Socorro, 800 m, ii.1987 (Janzen & Hallwachs) (BMNH): Heredia Prov.: 1 2, Puerto Viejo de Sarapiqui, Finca La Selva, iv.1986 (BMNH): Puntarenas Prov.: 1 9, 1 o’, Corcovado National Park, 20 m, v.1984 (Gauld) (BMNH); 1 0’, Monteverde, 1300-1400 m, vii.1982 (Janzen & Hallwachs) (BMNH); 1 @, same locality, iv-v.1984 (Gauld) (BMNH); 2 ’, same locality, xi.1985 & i.1986 (Haber) (BMNH): San José Prov.: 2 9, Estacion Carrillo, Braulio Carrillo National Park, 700 m, iii & v.1985 (Chacon & Chacon) (BMNH). Panama: 1 o’, 1 , Barro Colorado Island, iv.1965 (Duckworth & Duckworth) (USNM); 2 9, same locality, x.1977, iv.1978 (Wolda) (RNH); 2 0’, 5 Q, Chiriqui, Fortuna, 1050 m, i-v, x.1978 (Wolda) (RNH); 1 9, Las Cumbres, vi.1984 (Wolda) (BMNH). Peru: 1 9, Divisoria, Huanuco, 1700 m, ix.1946 (Woytkowski) (TC); 1 &’, Machu Picchu, xi.1965, (Townes & Townes) (TC). Enicospilus robertoi sp. n. (Figs 157, 228) Enicospilus species; Janzen, 1984b: 509. Description. Mandibles evenly tapered, apically twisted 10—-15° (Fig. 157); upper mandibular tooth slightly compressed, 1.2-1.3 times as long as the lower tooth; outer mandibular surface centrally flat, proximally with a shallow concavity. Labrum 0.3 times as long as broad; malar space 0.3-0.4 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt to subacute; clypeus in front 138 IAN D. GAULD Nenen, eueee” 22ees ha Pend eee v= Map 6 Localities at which Enicospilus brevis has been collected. view 1.3-1.4 times as broad as long, margin truncate. Lower face 0.68—0.74 times as broad as long, polished, punctate, sometimes tending to punctostriate centrally. Head in dorsal view with genae evenly rounded posteriorly; posterior ocellus virtually touching eye; FI = 75-80% ; occipital carina mediodorsally slightly flattened, often weak, sometimes interrupted, ventrally curved to join hypostomal carina but with lower end evanescent. Antenna long and slender, with 54-57 flagellar segments; 20th segment 1.9-2.0 times as long as broad. Mesoscutum polished, puncto-granulate, in profile quite steeply rounded; notauli weak, shallow. Mesopleuron polished, the upper part superficially punctostriate, the lower part coarsely and closely punctostriate; epicnemial carina inclined towards anterior margin of pleuron, its upper end curved, rather evanescent. Scutellum in profile moderately convex, laterally carinate for 0.9+ of its length; scutellum in | OPHIONINAE OF TROPICAL MESOAMERICA 139 dorsal view 1.5—1.6 times as long as anteriorly broad, finely granulate, posteriorly rugose to wrinkled. Metapleuron strongly convex, diagonally striate, polished; submetapleural carina evenly anteriorly broad- ened; posterior transverse carina of mesosternum complete, strong, often slightly raised centrally. Pro- podeum in profile evenly declivous; anterior transverse carina complete, weak, posterior transverse carina indistinct, usually absent; anterior area moderately long, steeply declivous, sparsely striate; spiracular area short, granulate; posterior area strongly coarsely concentrically striate; lateral longitudinal carina present only anteriorly, not joined to spiracular margin by a short carina. Fore wing length 14-17 mm; discosubmarginal cell as in Fig. 228; AI= 0.75—1.20; CI = 0.37-0.55; ICI = 0.42-0.56; SDI = 1.16-1.27; cu-a subopposite or slightly proximal to base of Rs&M; marginal cell proximally evenly hirsute; 1st subdiscal cell with anterior 0.6 sparsely hirsute. Hind wing with 7-8 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa weakly arcuate. Fore leg with tibia subcylindrical to weakly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.3—1.5 times length of the shorter. Hind leg with coxa in profile 1.6—-1.7 times as long as deep; trochantellus dorsally not exposed centrally; 4th segment of tarsus 2.5—2.7 times as long as broad; claws of female large, with long stout pectinae, those of male similar but with pectinae much shorter. Gaster long and slender; tergite 2 in profile 4.0-5.0 times as long as posteriorly deep, laterotergite upturned, thyridia elliptical and separated from anterior margin of tergite by about 3-5 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing very sparse, long fine erect hairs amid fine decumbent pubescence; gonosquama evenly rounded. Colour generally pale yellowish, tergites 3+ infuscate, mesoscutum slightly infuscate; interocellar area yellow; antenna yellowish; pterostigma yellowish; wings hyaline. VARIATION. Smaller specimens tend to be very pale yellow in general colour. REMARKS. This species is named in honour of Roberto Espinoza of Cuajiniquil, who has faithfully tended the Malaise traps, sorted Hymenoptera and reared caterpillars for years in Santa Rosa National Park. Enicospilus robertoi closely resembles E. scuintlei in structure and colour, but differs in having Rs+2r more evenly bowed and possessing a larger oval-triangular sclerite in the fenestra. The similarity between E. robertoi and E. scuintlei in the form of the mandibles and metapleurae, the male subgenital plate and colour suggest they are sister-species. BIOLOGICAL INFORMATION. Enicospilus robertoi is a Mesoamerican species that occurs from Belize south to Costa Rica. It has been collected in lowland and lower montane forest up to an altitude of about 1400 m. In Santa Rosa National Park, Costa Rica, it appears to fly at the beginning and end of the wet season (pooled data for 1979-84): Pee Ne ee Pe AS AN BPI ES! PS 2 Ape Se * ts At Monteverde, Puntarenas Province, Costa Rica, it has only been encountered between January and May (all data pooled): Diskehinwke) Abby ¢diagk iSoO~ Ni. BD My ee a Ee ae eee ee E. robertoi is a larval endoparasitoid of Hylesia lineata Druce (Lepidoptera: Saturniidae). This small, polyphagous hemileucine saturniid has urticating larvae which live gregariously up to the third instar. It is not known what instar larvae are parasitized, but E. robertoi has emerged from mature larvae that had been collected during their 4th instar. In Santa Rosa National Park H. lineata usually has two generations a year (Janzen, 1984b) Rearing data: 79-SRNP-55C; 79-SRNP-315; 79-SRNP-345; 79-SRNP-55H. MATERIAL EXAMINED Holotype ©’, Costa Rica: Guanacaste Prov., Santa Rosa National Park, ex Hylesia lineata, 1979 (Janzen & Hallwachs) (BMNH). Paratypes. Belize: 1 2, Middlesex, 125 m, v.1963 (Welling) (CNC). Costa Rica: Alajuela Prov.: 2 9, Finca Campana, 5 km NW. Dos Rios, 750 m, iii. 1985 (Janzen & Hallwachs) (BMNH); Guanacaste Prov.: 4 2, Volcan Cacao, Finca La Luz [Estacion Mengo], 1100 m, viii.1986, i,iii.1987 (Janzen & Hallwachs) (BMNH); 1 9, Volcan Orosi, Casa Mariksa [Maritza], 700 m, i.1986 (Janzen & Hallwachs) (BMNH);3 @, same locality, 700 m, vii.1986 (Gauld) (BMNH); 5 0’, 6 9, Santa Rosa National Park, 300 m, i, vi, vii & xii. 1979 (Janzen) (TC); 7 2, same locality, v & vii.1980, ii.1984, iii.1985 & i.1986 (Janzen & Hallwachs) (BMNH); 1 9, same locality, SE. of Entrada, vi.1986 (Godfrey) (BMNH): Puntarenas Prov.: 1 9, 140 IAN D. GAULD Monteverde, 1300 m, ii.1967 (Palmer) (TC); 1 9, same locality, ii.1980 (Mason) (TC); 1 2, same locality, iv.1984 (Gauld) (BMNH); 1 9, same locality, v.1980 (Janzen & Hallwachs) (BMNH); 2 co’, 14 9, same locality, i-iii, v.1986 (Haber) (BMNH); 14 Q, same locality, i-iii.1986 (Forsyth) (CNC). Enicospilus scuintlei sp. n. (Fig. 230) Description. Mandibles moderately long, relatively slender, apically twisted 10-30°; upper mandibular tooth slightly compressed, 1.7—2.0 times as long as the lower tooth; outer mandibular surface flat, proximal concavity moderately deep. Labrum 0.2-0.3 times as long as broad; malar space 0.20.3 times as long as basal mandibular width. Clypeus in profile almost flat, margin quite sharp; clypeus in front 1.3—1.5 times as broad as long, margin subtruncate. Lower face 0.72-0.78 times as broad as long, centrally obsoletely punctate, or punctostriate. Head in dorsal view with genae evenly constricted; posterior ocellus very close to eye; FI = 70-75%; occipital carina mediodorsally complete or obsolescent, ventrally incomplete, not curved towards hypostomal carina. Antenna slender, with 56-61 flagellar segments; 20th segment 1.9-2.1 times as long as broad. Mesoscutum polished, with fine shallow punctures, in profile evenly rounded, with anterior margin slightly turned forwards; notauli vestigial. Mesopleuron polished, the upper and lower part punctostriate; epicnemial carina abruptly curved towards anterior margin of pleuron, but with upper end evanescent. Scutellum in profile weakly convex, laterally carinate for all of its length; scutellum in dorsal view 1.4-1.5 times as long as anteriorly broad, from finely shagreened to virtually smooth. Metapleuron quite strongly convex, generally finely diagonally striate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile fairly abruptly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area moderately long, almost smooth, coriaceous or striate; spiracular area short, almost smooth; posterior area strongly rugose, the rugae tending to be concentric dorsally; lateral longitudinal carina present at least anteriorly, not distinctly joined to spiracular margin by a short carina. Fore wing length 14-16 mm; discosubmarginal cell as in Fig. 230; AI = 0.84-1.00; CI = 0.54-0.67; ICI = 0.52-0.61; SDI = 1.14-1.28; cu-a from subopposite to base of Rs& M to proximal to it by about 0.2 ofits own length; marginal cell proximally evenly hirsute; 1st subdiscal cell with anterior 0.4 sparsely hirsute. Hind wing with 7-8 hamuli on R1; Ist abscissa of Rs straight, 2nd abscissa straight. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.5-1.6 times length of the shorter. Hind leg with coxa in profile 1.7-1.9 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.4—2.6 times as long as broad; claws of female with long stout pectinae, those of male with pectinae slightly shorter. Gaster slender; tergite 2 in profile 4.5—5.5 times as long as posteriorly deep, laterotergite upturned, thyridia oval to elliptical and separated from anterior margin of tergite by about 3-4 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing scattered erect hairs; gonosquama distally truncate to rounded, rarely pointed. Colour generally pale yellowish brown, mesoscutum generally dark-marked and posterior segments of gaster infuscate; interocellar area yellowish; antenna golden; pterostigma golden; wings hyaline. VARIATION. Most specimens have three dark brownish longitudinal marks on the mesoscutum, though in a few individuals these are absent, and occasional other specimens have the entire mesoscutum dark brown. Most specimens have tergites 3+ of the gaster infuscate, but a few individuals also have tergite 2 slightly infuscate also. Almost every individual has a small oval/triangular ill-defined sclerotized patch in the fenestra, but in a few specimens this area is very weakly represented. ReEmaRKS. This species is named after one of the great characters of Santa Rosa (Agouti paca III) who will be remembered by toothmarks on the boots of a generation of field biologists Enicospilus scuintlei is structurally very similar to E. robertoi. Both species have similarly convex, striate metapleurae, similar mandibles, and have the 2nd subdiscal cell of the fore wing slightly broader than normal. The males of both species have a similar arrangement of rather sparse, long erect hairs on the posterior sternites. The colour pattern of the two species is also very similar. They differ principally in the form of the fenestra and the degree of sinuation of Rs+2r. This vein is less strongly sinuate in E. robertoi, but the sinuation extends virtually to the base of Rs; in E. scuintlei this vein is very strongly sinuate, but the most distal part is straight. The fenestra of E. scuintlei is broadly separated from the base of Rs, whilst in E. robertoi the distal fenestral margin reaches almost to Rs. E. robertoi has one and sometimes two distinct pigmented areas in the centre of the fenestra; the larger of these is oval/triangular and is quite large; the homologous sclerite in E. scuintlei is smaller and narrower. OPHIONINAE OF TROPICAL MESOAMERICA 141 BIoLoGIcAL INFORMATION. Enicospilus scuintlei is a common and widely distributed species whose range extends from the southern states of Mexico southwards to northern Argentina (Map 7). In Central America it is most commonly encountered in relatively forested sites, below 1000 m. Despite relatively intensive collecting it has never been collected in lower montane forest at Monteverde, Costa Rica (1300 m). In Santa Rosa National Park, Costa Rica, it has been collected during the driest months of the year, and at the start of the wet season. Individuals have been collected later in the year. The pooled distribu- tional data for 1981—4 at Santa Rosa are: TONEY WM “Ar iM 1 Dee AstS: Oy SN oD ras DIE 2 1 * 3 " + ‘ 2 y t B) € 2 » Map 7 Localities at which Enicospilus scuintlei has been collected. 142 IAN D. GAULD On Barro Colorado Island, Panama, E. scuintlei has been collected in most months, but is absent at the start of the year. Pooled distributional data are: jrowE MA Meds « Ju) (= SchOm i mB => tab Sresi BStenly siete dal ol eoik aro) Ide? A single specimen of this species has been reared by D. H. Janzen in Santa Rosa National Park from an unidentified species of Lasiocampidae (87-SRNP-1197) found feeding on Ocotea. This is an unusual host for species in this species-group as most others are only attack larvae of Saturniidae. MATERIAL EXAMINED Holotype 9, Costa Rica: Alajuela Prov., Finca San Gabriel, 16 km E. of Quebrada Grande, vi.1986 (Gauld) (BMNH). Paratypes. Argentina: 1 9, La Plata, xii.1965 (Townes & Townes) (TC); 1 CO’, Punta Lara, i.1966 (Townes & Townes) (TC). Belize: 1 2 , Augustine, vii. 1968 (Hasse) (BMNH); 1 0’, 3 9, Middlesex, 125 m, v.1963 (Welling) (CNC); 1 9, Rio Temas, vii.1937 (White) (BMNH). Bolivia: 1 2 , Rio San Pedro, Yungas de La Paz, 850 m, i.1976 (Pena) (TC). Brazil: 4 0’, 43 9, Bahia, Encruzilhada, 960-980 m, xi.1972 & xi.1974 (Alvarenga) (TC); 1 9, Ceara, Barbalha, 400 m, v.1969 (Alvarenga) (TC); 1 9, Caruaru, 900 m, vi.1972 (Lima) (TC); 1 9, Goias, Jatai, xi.1972 (Oliveira) (TC); 1 2, Mato Grosso, 12°49’§ 51°45’W, gallery forest, xi.1968 (Knight) (BMNH); 1 0’, 1 9, Mato Grosso, Sinop, 12°31’S 55°37W, x.1974 & x.1975 (Alvarenga) (TC); 1 9, Minas Gerais, Pedra Azul, 800 m, xi.1972 (Alvarenga & Seabra) (TC); 10, Paranda, Quatro Barros, nr Curitiba, ii.1966 (Townes & Townes) (TC); 1 9, Rio de Janeiro, Itatiaia National Park, x.1969 (Otero) (TC); 7 O, 6 2, Santa Catarina, Nova Teutonia, 27°11’S 52°23’W, x.1935, i & xi.1938, 1.1939, iii.1948, x.1951, xii.1953, iii.1956, i & x.1968 (Plaumann) (BMNH). Colombia: 1 9, Magdalena, 800 m, 1°10’N 74°8’W, iv.1973 (Helava) (BMNH). Costa Rica: Alajuela Prov.: 1 ©’, Finca Campana, 5 km NW. Dos Rios, 750 m, iii.1985 (Janzen & Hallwachs) (BMNH); 1 GC’, Finca San Gabriel, 16 km E. Quebrada Grande, 630 m, iii.1983 (Janzen & Hallwachs) (BMNH); 1 9, same locality, vi.1986 (Gauld) (BMNH): Cartago Prov.: 1 2, Turrialba, iii.1965 (Duckworth & Duckworth) (USNM); 1 9, Tapanti, Rio Grande de Orosi, 9°46’N 83°50’W, 1300-1400 m, 1.1985 (Janzen & Hallwachs) (BMNH): Guanacaste Prov.: 5 0’, 5 9, Santa Rosa National Park, 300 m, iii. 1980, viii. 1981, viii. 1982, iii & x. 1983, ii, iv-vi.1984 (Janzen & Hallwachs) (BMNH, TC); 4 9, Volcan Cacao, Estacion Mengo on SW. side, 1100 m, viii. 1986, v-vi.1987 (Janzen & Hallwachs) (BMNH): Heredia Prov.: 1 2, Puerto Viejo de Sarapiqui, Finca La Selva, iii.1986 (BMNH): Limon Prov.: 1 0’, Cerro Tortuguero, N. edge Tortuguero National Park, 0-100 m, v.1984 (Janzen & Hallwachs) (BMNH): Puntarenas Prov.: 1 2 , Corcovado National Park, 20 m, v.1984 (Gauld) (BMNH); 1 0’, Corcovado National Park, Sirena, i.1981, (Janzen & Hallwachs) (TC); 1 O',1 9, San Vito de Las Cruces, xi.1987 (Chacon) (BMNH): San José Prov.: 2 0’, 2 2, Estacion Carrillo, Braulio: Carrillo National Park, 700 m, vii & x.1984, iv-v.1985 (Chacon) (BMNH). Ecuador: 1 o’, 1 9, Cerro Tinajillas, Cumbaratza, 3100 m, iv.1965 (Peria) (BMNH); 1 9, Pichilingue, i.1974 (BMNH); 10’, Pichincha, nr Nanegal, 1100 m, ix.1977 (Pefia) (BMNH); 1 9, Rio Palenque, Los Rios, vi.1974 (Longino) (FSCA). Mexico: Chiapas: 1 9 , Chiapas de Corzo, viii. 1969 (Kritsch) (CNC): Quintana Roo: 1 9, X-can, viii.1963 (Welling) (CNC). Panama: 1 9, Barro Colorado Island, iv.1941 (Zetek) (USNM); 1 9, same locality, iv.1963 (Rettenmeyer) (TC); 1 2, same locality, vii.1963 (Cavagnaro & Irwin) (CAS); 12 9, same locality, iv-vii.1978 (Wolda) (RNH); 2 2, same locality, ix & xi.1982 (Wolda) (TC); 1 0’, 6 9, same locality, viii, xii.1983, v, vii. 1984, vi.1985 (Wolda) (BMNH); 2 0’, 6 2, Cerro Campana, nr Chica, iv.1965 (Duckworth & Duckworth) (USNM); 1 2, Cerro Campana, xi.1959 (Hanson) (TC); 1 CO’, 1 9, same locality, viii.1970 (Howden) (TC); 1 9, Las Cumbres, i.1982 (Wolda) (TC). Peru: 1 0’, Loreto, Pucallpa, 1.1962 (Schunke) (BMNH). Surinam: 2 9, Krakka, xii.1962 (Vreden) (RNH); 1 9, Zanderij, ix.1964 (Geijskes) (TC). Venezuela: 1 9, El Junquito, 1040 m, iv.1938 (Berthier) (TC). Enicospilus mexicanus (Cresson) stat. n. (Figs 93, 95, 231) Ophion mexicanus Cresson, 1874: 374. Holotype 9, MEXICO (PANS) [examined]. Ophion (Enicospilus) mexicanus Cresson; Cameron, 1886: 290. Enicospilus mexicanus (Cresson) Ashmead, 1895: 547. Henicospilus mexicanus (Cresson) Morley, 1912: 36. Description. Mandibles moderately long, basally strongly tapered, distally more evenly tapered, apically twisted 25—S0°; upper mandibular tooth slightly compressed, 1.4-1.6 times as long as the lower tooth; outer mandibular surface centrally bearing fine, short, scattered hairs, with a weak proximal concavity. Labrum OPHIONINAE OF TROPICAL MESOAMERICA 143 0.2-0.3 times as long as broad; malar space 0.20.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.3-1.5 times as broad as long, with margin truncate. Lower face 0.68—0.74 times as broad as long, centrally obsoletely punctate, the areas between the punctures slightly roughened. Head in dorsal view with genae rounded behind the eyes; posterior ocellus contiguous with the eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally quite strongly raised, curved towards and almost reaching hypostomal carina about 1.0—1.2 times the basal mandibular width away from mandible. Antenna slender, with 57-60 flagellar segments; 20th segment 1.7—1.8 times as long as broad. Mesoscutum matt, microreticulate, in profile evenly rounded, margin slightly out-turned; notauli weak but discernible. Mesopleuron polished, the upper part with specular area obsoletely punctostriate, posteri- orly grading to striate, the lower part more uniformly punctostriate/striate; epicnemial carina inclined towards anterior margin of pleuron, its upper end absent. Scutellum in profile weakly convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.4~—1.5 times as long as anteriorly broad, virtually smooth. Metapleuron moderately strongly convex, finely striate; submetapleural carina ante- riorly quite broad, but almost parallel-sided; posterior transverse carina of mesosternum complete, or slightly weaker medially. Propodeum in profile abruptly declivous; anterior transverse carina complete, posterior transverse carina present laterally; anterior area short, steeply declivous, with scattered striae; spiracular area short, almost smooth; posterior area coarsely rugose, the rugae tending to be concentric mediodorsally; lateral longitudinal carina present anteriorly, not joined to spiracular margin by a distinct short carina. Fore wing length 18-26 mm; discosubmarginal cell as in Fig. 231; AI = 0.87—1.11; CI = 0.53-0.62; ICI = 0.88-1.30; SDI = 1.19-1.54; cu-a proximal to base of Rs&M by 0.10.3 times its own length; marginal cell proximally evenly hirsute; 1st subdiscal cell with anterior 0.5 sparsely hirsute. Hind wing with 6-10 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa weakly bowed. Fore leg with tibia quite strongly flattened, with strong scattered spines on outer surface. Mid leg with longer tibial spur 1.4~-1.5 times length of the shorter. Hind leg with coxa in profile 1.5—1.7 times as long as deep; trochantellus dorsally <0.1 times as long as borad; 4th segment of tarsus 2.3-2.4 times as long as broad; claws of female large with close stout pectinae, those of male similar, but with pectinae a little shorter. Gaster moderately slender; tergite 2 in profile 3.1-3.5 times as long as posteriorly deep, laterotergite pendant, thyridia oval and separated from anterior margin of tergite by about 3-4 times its own length. Female unusual in having subgenital plate large, in profile longer than sternite 5 (Fig. 93); ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing long stout erect pubescence; gonosquama distally acute. Colour generally yellowish brown, generally with head, mesoscutal stripes and scutellum paler yellow, gaster only very slightly infuscate; interocellar area yellow; antenna generally brownish, though in some specimens they are blackish; pterostigma golden; wings almost hyaline. VARIATION. The most striking variation is in the torsion of the mandibles. Although the range given above extends from 25—S0° only a very few specimens have the mandibles weakly twisted; most have them turned 40-50°. A few specimens have the mesoscutal vittae dark brown. The range of variation in the coloration of the antenna is quite unusual, but seems to occur within a population. Individuals with both black and brown antennae have been collected at the same site on Barro Colorado Island in May/June. REMARKS. Enicospilus mexicanus is one of the most distinctive large species of the E. americanus complex. It can easily be recognized by the pendant laterotergite 2. Females are unusual in having a very large subgenital plate. Unlike many other species in this complex the genal carina (lower end of the occipital carina) often almost reaches the hypostomal carina. In this feature it resembles E. cameronii (see p. 147). Hooker (1912) erroneously treated E. mexicanus as a junior synonym of E. americanus and ever since this synonymy has been accepted. However, the two species are quite clearly distinct, especially in the features mentioned in the preceding paragraph. BIOLOGICAL INFORMATION. Enicospilus mexicanus is a widely distributed Neotropical species whose range extends from Mexico south of the tropic of Cancer to northern Argentina (Map 8). In Central America it is quite commonly collected in lowland rain forest, both on the eastern and western sides of the continent, and in humid lower montane forests up to an altitude of about 1400 m. In Braulio Carrillo National Park, Costa Rica, E. mexicanus has been collected in February, April to June, August and November. Although it is not common at Monteverde, a small number of specimens have been collected in various months, mostly in the first half of the year. The pooled seasonal data for Monteverde are: lieben tiw ApeSi.vOe NSD eR i: las ones) te Ai" dhe sik Giew se ood 144 IAN D. GAULD ” Map 8 Localities at which Enicospilus mexicanus has been collected. On Barro Colorado Island it is most common between May and September and only isolated individuals have been collected at other times of the year. The pooled data for Barro Colorado Island for 1978 and 1981-85 are: ee NMEA eM eA SO ND | gp cs ta I amps ie! PR a? “sits ‘al A tat E. mexicanus is uncommon in drier areas and only four individuals have been collected in Santa Rosa National Park. These were taken in mid June, early in the rainy season. OPHIONINAE OF TROPICAL MESOAMERICA 145 MaTERIAL EXAMINED Holotype 2, Mexico: Veracruz: Cordoba (PANS). Argentina: 1 9, Misiones, Panambi, xii.1957 (Walz) (TC). Belize: 1 9, Punta Gorda, x.1933 (White) (BMNH). Brazil: 2 2 , Bahia, Encruzilhada, 960-980 m, xi.1972 & xi.1974 (Alvarenga) (TC); 1 oO’, Parana, Quatro Barros, nr Curitiba, ii.1966 (Townes & Townes) (TC); 1 92, ROndania, Vilhena, xi.1973 x.1954 (Plaumann) (TC). Costa Rica: Alajuela Prov: 2 0’, 1 9, San Ram6n Reserve, Rio San Lorencito, xi. 1986, ii.1987 (Chacon) (BMNH): Cartago Prov.: 1 9, Tapanti, iv.1984 (Gauld) (BMNH): Guanacaste Prov.: 1 2, Santa Rosa National Park, 300 m, vi.1979 (Janzen) (BMNH); 2 9, same locality, vi.1984 (Janzen & Hallwachs) (BMNH); 1 2, same locality, vi. 1986 (Gauld) (BMNH); 1 2, Volcan Cacao, Finca La Luz [Estacion Mengo], 1100 m, i.1987 (Janzen & Hallwachs) (BMNH); 1 Q, same locality, vi.1987 (Janzen) (BMNH): Puntarenas Prov.: 1 9, Finca Las Cruces, 6 km S. San Vito de Java, 1400 m, x.1986 (Eger) (BMNH); 1 2, Monteverde, xii.1961 (Palmer) (TC); 2 2, same locality, 1300-1400 m, v.1980 & vii.1982 (Janzen & Hallwachs) (BMNH); 1 9, same locality, v.1984 (Fogden) (BMNH); 1 co’, 2 2, same locality, i, ili, v.1986 (Haber) (BMNH); 2 o’, 4 2, same locality, i-iii. 1986 (Forsyth) (CNC); 1 9, Sirena, Corcovado National Park, Osa Peninsula, i.1981 (Janzen & Hallwachs) (BMNH): San José Prov.: 1 6’, 7 Q, Estacion Carrillo, Braulio Carrillo National Park, 700 m, viii & xi.1984, ii, iv, v & vi.1985 (Chacon) (BMNH). Ecuador: 1 9, Pichincha, Tinlandia, 12 km E. Santo Domingo de los Colorados, 800 m, v.1986 (Eger) (BMNH). Guatemala: 1 2, Zacapa, San Lorenzo, 1700 m, xi.1986 (Sharkey) (CNC). Mexico: Chiapas: 1 9, Palenque, viii.1969 (Kelton) (CNC); 2 9, 35 km N. Huixtla, 1000 m, vi.1969 (Peterson) (CNC): Quintana Roo: 1 ©’, X-can, vi.1962 (Welling) (CNC). Panama: 1 9, Barro Colorado Island, Gatun Lake, iv.1941 (Zetek) (USNM); 4 0’, 14 9, same locality, i, iii, v-vii, ix, x, xi.1978 (Wolda) (RNH); 10°, 4 Q, same locality, ix, xi-xii.1982 & v.1983 (Wolda) (TC); 2 0’, 13 2, same locality, v, vi.1983, v, vi, vii, Vili, ix, x.1984, iv, vi, vii, viii.1985 (Wolda) (BMNH); 1 2, Boquete, Alto Lino, 1300 m, viii.1977 (Wolda) (RNH); 1 0’, Cerro Campana, nr Chica, iv.1965 (Duckworth & Duckworth) (USNM);10,1 9, Chiriqui, Fortuna, 1050 m, iii.1978 (Wolda) (RNH). Surinam: 1 2, Brownsweg, km 116, x.1961 (Pypers) (RNH). Enicospilus abelardoi sp. n. (Figs 94, 232) Description. Mandibles long, proximally very strongly narrowed, especially ventrally where margin curves out slightly distally so narrowed point of mandible is near the centre, the extreme distal end parallel- sided; mandibles apically twisted 30-40°; upper mandibular tooth compressed, 1.4~1.6 times as long as the lower; outer mandibular surface flat or weakly concave, with hairs scattered along central part, with a moderately strongly developed proximal concavity. Labrum 0.3-0.4 times as long as broad; malar space 0.2-0.4 times as long as basal mandibular width. Clypeus in profile flat, margin acute; clypeus in front view 1.2-1.4 times as broad as long, with margin truncate. Lower face 0.67—0.74 times as broad as long, weakly polished, punctate shallowly, centrally grading to punctostriate. Head in dorsal view with genae rounded; posterior ocellus very close to eye; FI = 70-75%; occipital carina mediodorsally narrowly interrupted, ventrally curved towards hypostomal carina but lower end evanescent. Antenna long and slender, with 69-71 flagellar segments; 20th segment 1.5—1.7 times as long as broad. Mesoscutum granulo-coriaceous, matt, in profile abruptly rounded; notauli weak. Mesopleuron matt, the upper part punctate with microreticulation on area between punctures, the lower part similar but coarser, tending to granulate; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile pyramidal, its anterior part strongly raised, almost conical, and posteriorly abruptly declivous (Fig. 94); scutellum laterally carinate for 0.7+ of its length; scutellum in dorsal view 1.4-1.6 times as long as anteriorly broad, puncto-granulate, posteriorly wrinkled. Metapleuron moderately convex, upper part rugose, the lower part granulo-coriaceous; submetapleural carina strongly anteriorly broadened, usually with anterior corner quite sharp; posterior transverse carina of mesosternum complete. Propodeum in profile abruptly declivous, posterodorsally deplanate; anterior transverse carina more or less complete, strongly sinuous, posterior transverse carina discernible only as lateral crests; anterior area abruptly declivous, quite short, strongly striate; spiracular area short and granulate; posterior area very coarsely rugose-reticulate; lateral longitudinal carina present only anteriorly, not joined to spiracular margin by a short carina. Fore wing length 23-28 mm; discosubmarginal cell as in Fig. 232; AI = 0.75-0.98; CI = 0.39-0.56; ICI = 0.45-0.87; SDI = 1.39-1.51; cu-a proximal to base of Rs&M by 0.1-0.2 times its own length; marginal cell proximally evenly hirsute; 1st subdiscal cell with anterior 0.6 and posterodistal corner hirsute. Hind wing with 9-13 hamuli on R1; Ist abscissa of Rs straight, 2nd abscissa weakly arcuate. 146 IAN D. GAULD Fore leg with tibia quite strongly flattened, with long scattered spines on outer surface. Mid leg with longer tibial spur 1.3—1.4 times length of the shorter. Hind leg with coxa in profile 1.6—-1.7 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.2-2.4 times as long as broad; claws of female and male similar, strongly curved, with quite short, close pectinae. Gaster long and slender; tergite 2 in profile more than 5.0 times as long as posteriorly deep, laterotergite normally folded under, thyridia small, obovate to cordate and separated from anterior margin of tergite by about 5-6 times its own length. Female with subgenital plate of moderate size, in profile only slightly longer than sternite 5 and posteromedially membranous; ovipositor exceptionally slender, its sheath narrow. Male with sternites 7-9 bearing scattered fine semidecumbent pubescence; gonosquama distally evenly rounded. Colour generally quite uniformly brownish yellow with indistinct pale mesoscutal stripes; interocellar area yellowish; antenna dark brown, paler distally; pterostigma yellowish brown; wings weakly infumate. VARIATION. None remarkable. REMARKS. This species is named in honour of Abelardo Chacon, who has collected many of the ophionines examined in this work, and who has worked so hard on the moth fauna of Costa Rica. Enicospilus abelardoi is a distinctive species on account of its very large size and its weakly sinuous Rs+2r; all the other very large species in the E. americanus complex have Rs+2r more strongly sinuous. E. abelardoi is particularly distinctive in the form of its scutellum which, in profile appears pyramidal, and is quite unlike that of any other Mesoamerican species. The form of the mandibles is also quite distinctive but is rather difficult to appreciate unless comparative material is at hand. Although the form of Rs+2r suggests E. abelardoi may be related to the complex of species that includes E. aktites (all of which have Rs+2r very weakly sinuate), the form of the propodeal sculpture and the shape of the claws suggest it may be more closely related to E. cameronii. The males of E. cameronii, however, have erect hairs on the posterior sternites. BIOLOGICAL INFORMATION. Enicospilus abelardoi is only known from wet forests at altitudes between 700 and 800 m in Costa Rica. Its host is unknown, but its phylogenetic affinities and very large size suggest it parasitizes one of the largest saturniines; the exceptionally slender ovipositor suggests it may be ovipositing in relatively early instar larvae. MATERIAL EXAMINED Holotype 9, Costa Rica: Alajuela Prov., San Ramon Reserve, Rio San Lorencito, 800 m, v.1987 (Chacon & Chacon) (BMNH). Paratypes. Costa Rica: 3 O’, San José Prov., Estacion Carrillo, Braulio Carrillo National Park, 700 m, iii & v.1985 (Chacon & Chacon) (BMNH). Enicospilus hallwachsae sp. n. (Fig. 233) Description. Mandibles moderately long, strongly narrowed so apex is slender, apically twisted 15—20°; upper mandibular tooth subcylindrical, 1.2-1.4 times as long as the lower tooth; outer mandibular surface with a broad shallow proximo-ventral concavity. Labrum 0.2-0.3 times as long as broad; malar space 0.2- 0.3 times as long as basal mandibular width. Clypeus in profile very weakly convex, margin blunt; clypeus in front view 1.2—1.4 times as broad as long, margin truncate. Lower face 0.67-0.73 times as broad as long, centrally with very fine, sparse punctures. Head in dorsal view with genae rounded behind eyes; posterior ocellus contiguous with eye; FI = 65-70%; occipital carina mediodorsally complete, ventrally not reaching hypostomal carina. Antenna slender, with 60-62 flagellar segments; 20th segment 1.6-1.7 times as long as broad. Mesoscutum weakly polished, sparsely punctate, in profile abruptly rounded; notauli vestigial. Meso- pleuron polished, the upper part punctostriate, the lower part striate; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for all of its length; scutellum in dorsal view 1.3—1.5 times as long as anteriorly broad, anteriorly smooth, posteriorly rugulose. Metapleuron convex, granulate or finely diagonally striate, or exceptionally, rugose; submetapleural carina generally evenly anteriorly broadened; posterior transverse carina of mesosternum usually com- plete, or in Panamanian individuals weak to incomplete. Propodeum in profile abruptly declivous; anterior transverse carina complete, usually centrally raised, posterior transverse carina from absent to present laterally as a vestige; anterior area moderately long, striate; spiracular area short, abruptly declivous, punctate finely; posterior area rugose, the rugae tending to be concentric posterodorsally; lateral long- itudinal carina present only anteriorly, not joined to spiracular margin by a short carina. | | | | | | | | | | OPHIONINAE OF TROPICAL MESOAMERICA 147 Fore wing length 15-18 mm; discosubmarginal cell as in Fig. 233; AI = 0.79-1.20; CI = 0.51-0.59; ICI = 0.76—-1.06; SDI = 1.38-1.61; cu-a proximal to base of Rs&M by 0. 1—0.4 times its own length; marginal cell proximally more or less evenly hirsute; 1st subdiscal cell with anterior 0.3 hirsute. Hind wing with 5-8 hamuli on R1; 1st abscissa of Rs almost straight, 2nd abscissa straight. Fore leg with tibia slightly flattened, with a few strong spines on outer surface. Mid leg with longer tibial spur 1.3-1.4 times length of the shorter. Hind leg with coxa in profile 1.6-1.7 times as long as deep; | trochantellus dorsally <0.1 times as long as broad; claws of female large, with short close pectinae, those of _ male similar. Gaster slender; tergite 2 in profile 3.74.3 times as long as posteriorly deep, laterotergite turned under, | thyridia small, obovate and separated from anterior margin of tergite by about 4—5 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing long stout erect hairs; gonosquama | long, apically somewhat acute, usually with upper distal corner slightly produced. Colour generally yellowish brown, head brighter yellow, mesoscutum with weak to quite strong darker | brownish vittae, gaster slightly infuscate; interocellar area yellowish; antenna brownish orange, generally | infuscate or even blackish proximally; pterostigma golden; wings hyaline. | VarIaTION. Occasional specimens have the antennae almost entirely black. The female from Belize has the metapleuron punctate and so have the five specimens from Chiriqui, Panama. These five specimens may | possibly represent a separate species for, in addition to having the metapleuron punctate, they have the | posterior transverse carina of the mesosternum centrally weak or incomplete, their general colour is darker brown, and the males have slightly coarser erect hairs on the posterior sternites. I have seen a single damaged male labelled ‘Trinidad’ (in RNH), which runs to this species in the key. It differs in having a more finely sculptured alitrunk, with the metapleuron striate, and an exceptionally large male genital capsule. It appears to be a distinct species but I have refrained from formally describing it until more material is available (and which ‘Trinidad’ it was collected in becomes apparent). REMARKS. This species is named in honour of Winifred Hallwachs who has baked many butterscotch brownies, collected ichneumonids at hours when most sane people are asleep, and inspired the moth study that led to this ichneumonid study. Enicospilus hallwachsae belongs to a species-complex that is characterized by having an almost straight Rs+2r. Within this group it appears to be very closely related to E. aktites which it resembles in having slender mandibles and similar venation. E. hallwachsae differs in having Rs+2r very slightly more slender, in lacking the distinct small central sclerite and in being rather darker in colour. BIOLOGICAL INFORMATION. Enicospilus hallwachsae is a rarely collected species that is widely distributed from Belize south to Brazil. It occurs in rather wetter habitats than does E. aktites and in Costa Rica it is associated with rainforests between 600 and 1100 m, mostly on the Atlantic side of the continent. In northern Costa Rica it has been collected near the continental divide at 630 m. Despite extensive collecting it has not been collected in lowland forest on Barro Colorado Island, or in lower montane forest at Monteverde (1300 m). MATERIAL EXAMINED Holotype 0’, Costa Rica: Alajuela Prov., Finca San Gabriel, 16 km E. Quebrada Grande, 630m, iii.1983 (Janzen & Hallwachs) (BMNH). Paratypes. Belize: 1 0’, Camp Sibim, Coyo Dist., v.1963 (CNC). Brazil: 1 O’, Santa Catarina, Nova Teutonia, v.1953 (Plaumann) (TC). Costa Rica: Alajuela Prov.: 1 0, Finca San Gabriel, 16 km E. Quebrada Grande, 630 m, iti.1983 (Janzen & Hallwachs) (BMNH); | 0’, same locality, vi.1986 (Gauld) (BMNH); 1 9, Caché, 1914 (Rogers) (BMNH): San José Prov.: 1 co’, 3 2, Estacion Carrillo, Braulio Carrillo National Park, 700 m, vii-viii. 1984, iii.1985 (Chacon) (BMNH). Panama: 3 ©’, 2 9, Chiriqui, | Fortuna, 1050 m, i-iv.1978 (Wolda) (RNH). Non-paratypic material. Trinidad: 1 O'(de Voogd) (RNH). Enicospilus cameronii (Dalla Torre) (Figs 159, 234) | Ophion curvinervis Cameron, 1886: 293. Lectotype 9, GUATEMALA, (BNMH) designated by Townes & Townes (1966: 175) [examined]. Homonym of Ophion curvinervis Kriechbaumer, 1878. Ophion cameronii Dalla Torre, 1901: 188. Replacement name for curvinervis Cameron. Ophiomorpha curvinervis (Cameron) Szépligeti, 1905: 35. Ophion latilineatus Cameron, 1911: 179. Holotype 2, GUYANA (BMNH) [examined]. Synonymized by Morley, 1914: 409. 148 IAN D. GAULD Allocamptus renovatus Morley, 1912: 23. Replacement name for curvinervis Cameron. Cymatoneura renovata (Morley) Bréthes, 1927: 323. Enicospilus cameronii (Dalla Torre) Cushman, 1947: 466. Description. Mandibles moderately long, proximally strongly narrowed, distally almost parallel-sided, apically twisted 20-30°; upper mandibular tooth slightly compressed, 1.2-1.4 times as long as the lower tooth, and slightly divergent from it; outer mandibular surface centrally finely pubescent, proximally almost flat. Labrum 0.2-0.3 times as long as broad; malar space 0.30.4 times as long as basal mandibular width. Clypeus in profile almost flat, margin blunt to subacute; clypeus in front view 1.4-1.6 times as broad as long, with margin truncate. Lower face 0.70—0.83 times as broad as long, centrally obsoletely punctate. Head in dorsal view with genae rounded behind eye; posterior ocellus contiguous with eye; FI = 70-75%; occipital carina mediodorsally generally slightly depressed, sometimes narrowly interrupted, ventrally curved towards the hypostomal carina but usually not joining it, though in a few individuals these carinae do join. Antenna moderately slender, with 61-70 flagellar segments; 20th segment 1.7—1.9 times as long as broad. Mesoscutum weakly polished, finely punctate, in profile abruptly rounded; notauli vestigial. Meso- pleuron weakly polished, the upper part punctostriate, grading in the lower part to striate; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile moderately convex, laterally usually carinate for 0.9 or more of its length, rarely with carinae only extending 0.5 of its length; scutellum in dorsal view 1.4-1.5 times as long as anteriorly broad, anteriorly smooth, posteriorly often striate. Metapleuron posterodorsally convex, ventrally flattened, more or less matt, finely rugulose, or granulate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete, centrally produced into a squared-off lobe. Propodeum in profile abruptly declivous; anterior transverse carina sinuous, usually complete, rarely with lateromedian discontinuities, posterior transverse carina represented laterally by crests; anterior area deeply impressed, striate; spiracular area very short and almost smooth; posterior area coarsely rugose (Fig. 159), the rugae posteriorly tending to be concentric; lateral longitudinal carina present anteriorly, not joined to spiracular margin by a short carina. Fore wing length 17-27 mm; discosubmarginal cell as in Fig. 234; AI = 0.78-1.40; CI = 0.59-0.61; ICI = 0.51-0.80; SDI = 1.25-1.46; cu-a proximal to Rs&M by 0.2-0.4 times its own length; marginal cell proximally usually evenly hirsute, sometimes with a small glabrous area; 1st subdiscal cell with anterior and distal margins hirsute. Hind wing with 9-14 hamuli on R1; Ist abscissa of Rs straight, 2nd abscissa weakly bowed. Fore leg with tibia strongly flattened, with scattered stout spines on outer surface. Mid leg with longer tibial spur 1.3—-1.5 times length of the shorter. Hind leg with coxa in profile 1.6-1.7 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.5—2.7 times as long as broad; claws of female large, with strong, close pectinae, those of male similar but with pectinae slightly closer and shorter. Gaster quite slender; tergite 2 in profile 4.4-5.6 times as long as posteriorly deep, laterotergite folded under, or in a few individuals with extreme posterior part slightly pendant; thyridia obovate and separated from anterior margin of tergite by about 3.5-4.5 times its own length. Female with subgenital plate large, conspicuously longer than sternite 5; ovipositor slender, its sheath a little stouter than is normal for species in this complex. Male with sternites 7-9 bearing dense long erect pubescence; gonosquama distally elongately rounded or acute. Colour generally yellowish brown with head paler yellowish; upper part of mesopleuron, mesoscutal stripes and scutellum often yellow; interocellar area yellow; antenna blackish; pterostigma golden; wings very weakly infumate. VARIATION. Enicospilus cameronii is a morphologically rather uniform species that shows most variation in the sculpture of the metapleuron. Generally the metapleuron is more or less granulate to rugulose but occasional individuals have it somewhat striate, or even rugose. Specimens from Fortuna, Chiriqui, Panama have the alitrunks slightly darker than other material. REMARKS. Enicospilus cameronii appears to be closely related to E. mexicanus. Both species have a comparatively large female subgenital plate and have the lower part of the occipital carina reaching or almost reaching to the hypostomal carina. E. cameronii differs from E. mexicanus in having the epipleuron 2 more or less completely turned under, and in having a smaller ICI (<0.80). E. cameronii is most easily distinguished from other species in the E. americanus complex by the possession of a black flagellum and in having the propodeum very coarsely sculptured. BIOLOGICAL INFORMATION. The range of Enicospilus cameronii extends from Oaxaca, Mexico south to equatorial South America (Map 9). It seems to be associated with wet forests from sea-level up to altitudes of about 1400 m. Most specimens have been collected in the first six months of the year. 49 1 OPHIONINAE OF TROPICAL MESOAMERICA ‘pajoa][oo udaq sey Muosauvs snpidsorUy YOY ye sayyyeooT 6 dey 150 IAN D. GAULD MATERIAL EXAMINED Lectotype 9 (Ophion curvinervis Cameron), Guatemala: Las Mercedes, 1000 m (Champion) (BMNH); paralectotype 1 CG’, Guatemala: Senahu, Vera Paz (Champion) (BMNH). Holotype 9 (Ophion latilineatus Cameron), Guyana (BMNH). Belize: 1 &', 1 2, Columbia Forest, vii.1968 (Hasse) (BMNH); 1 2, Rio Temas, iv.1937, (White) (BMNH). Costa Rica: Alajuela Prov.: 1 0’, 1 2, Finca Campana, 5 km NW. Dos Rios, 750 m, iii.1985 & i.1986 (Janzen & Hallwachs) (BMNH); 1c’, 3 km W. Dos Rios, Finca San Gabriel, 800 m, vi.1986 (Gauld) (BMNH): Guanacaste Prov.: 1 &’, 4 km E. Casetilla, Rincon National Park, 750 m, 1.1982, (Janzen & Hallwachs) (BMNH); 1 2, Volcan Orosi, Casa Mariksa [Maritza], v.1986 (Gauld) (BMNH): Puntarenas Prov.: 1 &', 2 9, Monteverde, 1300 m, ii, vii.1961, ii.1962 (Palmer) (TC); 1 9, same locality, iv-v.1984 (Gauld) (BMNH); 3 9, 3 ,, same locality, i-ii, v.1986 (Haber) (BMNH); 1 9, Sirena, Corcovado National Park, Osa Peninsula, iii.1981 (Janzen & Hallwachs) (BMNH): San José Prov.: 1 0’, 2 2, Estacion Carrillo, Braulio Carrillo National Park, 700 m, ix.1984, iii & v.1985 (Chacon) (BMNH); 2 9, La Montura, Braulio Carrillo National Park, 1100 m, xii.1981 (Janzen & Hallwachs) (BMNH); 2 2, San Antonio de Escazi, 1300 m, v-vi.1981 (Eberhard) (UM). Ecuador: 1 ©’, Otavalo to Apuela, 2200 m, ix.1977 (Pena) (TC). Guatemala: 1 9, Tabril, 1931 (MCZ). Mexico: Chiapas: 1 CO’, 32 km N. Huixtla, 1000 m, vi.1969 (Mason) (CNC); 10',1 9, 32kmN. Huixtla, 1000 m, vi.1969 (Peterson) (CNC); 2 2, 32 km N. Huixtla, 1000 m, vi.1969 (CNC): Oaxaca: 1 9, 140 km on Highway 175, 1200 m, v.1969 (Howden) (CNC); 1 2, Temascal, viii. 1966 (Lau) (USNM); 1 9, 20 km S. Valle Nacional, 1000 m, v.1971 (Howden) (TC). Panama: 1 O’, Barro Colorado Island, vii.1961 (Campbell) (CNC); 5 oO’, 17 9, Chiriqui, Fortuna, 1050 m, i, iv-vi, x-xi.1978 (Wolda) (RNH); 1 2, Las Cumbres, 150 m, vi.1984 (Wolda) (BMNH). Venezuela: 1 9, El Blanquito, Lara, x.1978 (Osorio) (FSCA). Enicospilus gamezi sp. n. (Figs 160, 235) Description. Mandibles quite stout, long, distally almost parallel-sided, apically twisted 10—20°; upper mandibular tooth subcylindrical, 1.8—2.0 times as long as the lower tooth; outer mandibular surface almost flat centrally, with a weak proximal concavity. Labrum 0.2-0.3 times as long as broad; malar space 0.2-0.3 times as long as basal mandibular width. Clypeus in profile flat to slightly out-flared, margin slightly blunt; clypeus in front view 1.4—1.5 times as broad as long, its margin truncate to weakly convex. Lower face 0.77— 0.85 times as broad as long, centrally regularly but finely punctate. Head in dorsal view with genae evenly rounded behind eye; posterior ocellus close to eye; FI = 60-65%; occipital carina mediodorsally narrowly incomplete, ventrally evanescent, remote from the hypostomal carina. Antenna slender, with 57-69 flagellar segments; 20th segment 2.2—2.3 times as long as broad. Mesoscutum polished, finely granulate with obsolescent punctures, in profile steeply but evenly rounded; notauli vestigial. Mesopleuron polished, the upper part punctate with a few striae posteriorly, the lower part punctostriate; epicnemial carina curved towards anterior margin of pleuron, but with upper end evanescent. Scutellum in profile moderately convex, laterally carinate for 0.9 of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, rather smooth, with striae posteriorly. Metapleuron weakly convex, diagonally striate; submetapleural carina narrow, weakly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly rounded; anterior transverse carina present but rather weak, posterior transverse carina absent; anterior area moderately long, striate or rugulose; spiracular area short, almost smooth; posterior area transversely rugulose (Fig. 160); lateral — longitudinal carina present only anteriorly, not joined to spiracular margin by a short carina. ( Fore wing length 13-19 mm; discosubmarginal cell as in Fig. 235; AI = 0.80-1.23; CI = 0.37-0.52; ICI = 0.54—0.84; SDI = 1.21-1.27; cu-a from slightly distal to the base of Rs&M to proximal to Rs&M by 0.10.2 times its own length; marginal cell proximally very slightly more sparsely hirsute; 1st subdiscal cell with anterior 0.5—0.7 hirsute. Hind wing with 5-8 hamuli on R1; Ist abscissa of Rs straight, 2nd abscissa — weakly bowed. Fore leg with tibia weakly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.3-1.4 times length of the shorter. Hind leg with coxa in profile 1.6-1.8 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.6-3.1 times as long as broad; claws of female long, distally abruptly curved, with fine close pectinae, those of male similar. Gaster slender; tergite 2 in profile 4.14.3 times as long as posteriorly deep, laterotergite folded under, thyridia oval and separated from anterior margin of tergite by about 3-4 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing fine decumbent pubescence; gonosquama distally slightly acute. OPHIONINAE OF TROPICAL MESOAMERICA 151 Colour generally yellowish brown with head paler yellowish and gaster very weakly infuscate posteri- orly; interocellar area yellowish; antenna blackish; pterostigma golden brown; wings hyaline. VARIATION. None remarkable. REMARKS. This species is named in honour of Professor Rodrigo Gamez who has done so much in welding the political and academic community to the conservation cause in Costa Rica. Enicospilus gamezi is rather similar to, though generally smaller than, E. cameronii. E. gamezi differs in that the female does not have quite such an elongate subgenital plate, the male has the terminal sternites finely pubescent, both sexes have more slender central flagellar segments and stouter, longer mandibles. The propodeal sculpture of E. gamezi is far less strong than that of E. cameronii, and it lacks the lateral vestiges of the posterior transverse carina. BIOLOGICAL INFORMATION. Enicospilus gamezi is only known from north-western Costa Rica. It is appar- ently restricted to rather open deciduous forest. The adults are active throughout the wet season from the end of May until November. Pooled data for 1977-85 are: [ca eo ae emi gag as gr ia a Sere ee tae ne About one-quarter of the specimens examined have very worn mandibles, suggesting that this species may emerge from a hard pupal cell in the ground. MATERIAL EXAMINED Holotype ©’, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, vili.1983 (Janzen & Hallwachs) (BMNH). Paratypes. Costa Rica: Guanacaste Prov.: 7 0’, 6 2, Santa Rosa National Park, 300 m, viii, x, xi.1977 vi, Vii, viii. 1978, vi, ix.1983, vi.1984, vi.1985 (Janzen & Hallwachs) (BMNH, MNCR). Enicospilus texanus (Ashmead) (Figs 99, 236) Thyreodon texanus Ashmead, 1890: 422. Holotype 0’, U.S.A.: Texas (USNM) [examined]. Eremotylus texanus (Ashmead) Ashmead, 1896: 23. Macrophion texanus (Ashmead) Morley, 1912: 14. Enicospilus texanus (Ashmead) Townes, 1945: 745. Description. Mandibles long, weakly tapered proximally, distally almost parallel-sided, apically twisted 10-15°; upper mandibular tooth subcylindrical, 1.2—1.7 times as long as the lower tooth; outer mandibular surface centrally with fine sparse pubescence proximally, generally weakly concave, with a weak proximal concavity. Labrum 0.2-0.3 times as long as broad; malar space sexually dimorphic, of 2 0.3-0.5, and of C 0.5—0.8 times as long as basal mandibular width. Clypeus in profile flat, sometimes slightly out-flared, margin rather blunt; clypeus in front view 1.5—1.8 times as broad as long, its margin truncate. Lower face of 2 0.86-1.05, of O' 0.94-1.25 times as broad as long; face centrally smooth and polished, with scattered coarse punctures. Head in dorsal view with genae slightly inflated in 2, buccate in ©’; posterior ocellus from close to eye to separated from it by about 0.2 its own maximum diameter in ?, in CO’ separated from eye by 0.2-0.3 times the maximum ocellar diameter; FI of O’ 45-50%, of 2 50-55%; occipital carina mediodorsally narrowly interrupted, ventrally curved to nearly join hypostomal carina about 0.8-0.9 times the basal mandibular width away from mandible, but with lower end evanescent so the two carinae do not actually meet. Antenna moderately slender, with 53-61 flagellar segments; 20th segment 1.5—1.8 times as long as broad. Mesoscutum polished, with close punctures, in profile steeply rounded; notauli vestigial. Mesopleuron polished, rather coarsely punctate, though posterodorsally and medioventrally tending to punctostriate; epicnemial carina curved towards anterior margin of pleuron but with upper end evanescent. Scutellum in profile weakly to moderately convex, laterally carinate for most of its length though often with carina degenerating into a series of wrinkles posteriorly, sometimes with the carinae weak; scutellum in dorsal view 1.2-1.3 times as long as anteriorly broad, with deep punctures, posteriorly somewhat striate. Metapleuron convex, punctate, rarely punctostriate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum more or less complete, sometimes a little protuberant cen- trally. Propodeum in profile abruptly declivous, posterodorsally deplanate; anterior transverse carina present, often weak and discontinuous, posterior transverse carina represented by a lateral vestige; IS, IAN D. GAULD anterior area short, deeply impressed, rugose/striate; spiracular area short and punctate, posterior area rugose/reticulate, the rugae tending to concentric posterodorsally; lateral longitudinal carina only present anteriorly, not joined to spiracular margin by a short carina. Fore wing length 15-18 mm; discosubmarginal cell as in Fig. 236; AI = 0.64-1.46; CI = 0.46-0.73; ICI = 0.61-0.86; SDI = 1.35—1.50; cu-a proximal to base of Rs&M by 0. 1-0.3 times its own length; marginal cell proximally generally slightly more sparsely hirsute distally; 1st subdiscal cell generally extensively hirsute, often only with centre or posterior margin glabrous. Hind wing with 8-12 hamuli on R1; 1st abscissa of Rs almost straight, 2nd abscissa weakly curved. Fore leg with tibia weakly flattened, with numerous slender spines on outer surface. Mid leg with longer tibial spur 1.2-1.4 times length of the shorter. Hind leg with coxa in profile 1.5—1.7 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.1—2.6 times as long as broad; claws of female long and weakly curved, with stout pectinae (Fig. 99), those of male similar, with pectinae slightly closer. Gaster moderately stout; tergite 2 in profile 2.5-3.8 times as long as posteriorly deep, laterotergite turned under, thyridia elongately obovate and separated from anterior margin of tergite by about 1.5—2.0 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing rather fine, long,, erect hairs amid decumbent pubescence; gonosquama distally rounded. Colour generally dark reddish or orange-brown, usually with head and scutellum slightly paler; inter- ocellar area orange; antenna orange-brown slightly infuscate apically; pterostigma orange; wings quite strongly infumate. VARIATION. I have seen three specimens (in BMNH and USNM) from Arizona, U.S.A., all reared, which differ from typical examples of the species in being slightly smaller (fore wing length about 14 mm) and having almost hyaline wings. REMARKS. Enicospilus texanus is easily recognized by the dark wings and the sexually dimorphic head. The broad face, long malar space and buccate genae of the male are quite distinctive. Structurally this species is similar to E. halffteriin sculpture, venation and form of the mandibles. However, E. texanus has a broader face, wider malar space, more spinose hind tarsi and, usually, darker wings than does E. halffteri. BIOLOGICAL INFORMATION. This species is widespread throughout the southern states of the U.S.A. from Virginia westwards to the Pacific coast, and it is reputed to extend as far north as Washington (Carlson, 1979). In the east there are occasional records of it from as far north as Ohio. In the south its range extends into northern Mexico, though it does not seem to occur in tropical Central America. Enicospilus texanus is a larval endoparasitoid of (mostly) hemileucine saturniids. In the United States it commonly has been reared as a larval parasitoid of Hemileuca maia (Drury), a saturniid that is common in drier scrubby areas of the eastern United States (Ferguson, 1971). E. texanus has also been reared from H. peigleri Lemaire (Lemaire, 1981). R. Peigler informs me that this species also parasitizes H. magnifica (Rotger). In the USNM are two specimens that are similar to texanus, but may not be conspecific with it (see variation above), which have been reared from Hemileuca tricolor (Packard), and in the BMNH is a very similar specimen that has been reared from Hemileuca juno Packard. In northern Mexico, E. texanus has been reared from the larva of Hemileuca oliviae Cockerell, an occasional pest of range lands (Fritz et al., 1986). R. Peigler has also reared E. texanus from the saturniine, Agapema galbina (Clemens). In the USNM is a single specimen of texanus that is labelled as having been reared from the ennomine geometrid Eucaterva variaria (Grote). This record is erroneous as the purported host is too small to support such a large parasitoid. Fritz et al. (1986) observed that the larva of E. texanus killed its larval host after the saturniid had spun a cocoon. The ichneumonid constructed a tough ovoid fibrous cocoon in the host cocoon, underneath the remains of the saturniid larva. This larval skin, which bears urticating spines, may serve to protect the ichneumonid cocoon. The single specimen I have seen from a saturniine (Agapema galbina) also con- structed its cocoon within the rather loose host cocoon. The cocoon of E. texanus is particularly thick and, like other ophionines, internally impregnated with a cellophane-like substance that presumably serves to reduce water-loss and protects the larva from micro-organisms. The cocoon of texanus is remarkable in having a well-developed, free, inner cellophane-like envelope that completely ensheaths the prepupa and pupa. This has its own cap, separate from the usual external cap. Possibly this additional layer, another barrier to water-loss, allows texanus to diapause as larvae for long periods in relatively dry localities. In some cases individuals of E. texanus enter prolonged diapause. For example, the individual reared from Agapema spun a cocoon in May 1981, but did not emerge until January 1983. MATERIAL EXAMINED Holotype 0’, U.S.A.: Texas (USNM). OPHIONINAE OF TROPICAL MESOAMERICA 153 Mexico: Chihuahua: 1 0’, 1 9, 3 km W. Anehuac, ex H. oliviae, x.1983 (Fritz) (BMNH). U.S.A.: Arizona: | Q, Pima County, E. of Tucson, iv.1987 (Hyatt) (BMNH); 1 o&’, 1 2, Tombstone, x.1934 (Wehrle) (USNM); 2 9, Tucson, iii.1936, iv.1937 (Bryant) (CAS): California: 1 Go’, Lake Tahoe, ix.1915 (Dyar) (USNM); 5 9, Sequioa Natl. Park (CAS): Florida: 4 2, Highlands Co., Archbold Biological Station, iii.1962 (Ferguson) (USNM); 1 Q, Lake Placid, v.1968 (Heinrich) (CNC); 3 2, Lake Placid iv.1979 (CNC): Georgia: 1 CO’, Waycross, iii. 1952 (Gillis) (CNC): New Mexico: 1 2, Taos, vii. 1934 (Craig) (CAS): North Carolina: 1 9, Highlands, iv.1957 (Mason) (CNC): Texas: 1 0’, Brewster Co., i.1969 (Kendall) (USNM); 1 9, Reeves Co., US Highway 285, 3 km SE. of Orla, ex A. galbina larva feeding on Condalia ericoides (Kendall & Kendall) (BMNH); 1 2, San Antonio, iii. 1930 (Seaton) (USNM); 1 2, San Antonio, iii.1957 (Kendall) (USNM): Virginia; 1 2, Mountain L. vii.]940 (Milne) (USNM). Enicospilus cushmani Gauld (Fig. 237) Enicospilus cushmani Gauld, 1988: 41. Holotype, @ U.S.A.: Florida (USNM) [examined]. DEscriPTION. Mandibles moderately short, fairly evenly tapered from base to apex, apically twisted 10—20°; upper mandibular tooth slightly compressed, 1.8-2.1 times as long as the lower tooth; outer mandibular surface sparsely pubescent, centrally flat, and with a weak proximal concavity. Labrum 0.2-0.3 times as long as broad; malar space 0.20.5 times as long as basal mandibular width (males tend to have a longer malar space than females). Clypeus in profile flat or slightly out-flared, margin blunt to subacute; clypeus in front view 1.8—1.9 times as broad as long, its margin truncate. Lower face of 2 0.80—1.10, of co’ 0.90-1.20 times as broad as long, centrally smooth and polished, with fine scattered punctures. Head in dorsal view with genae slightly inflated behind eyes; posterior ocellus close to or contiguous with eye; FI = 55-65%; occipital carina mediodorsally absent or weak, ventrally evanescent before joining the hypostomal carina. Antenna moderately long and slender, with 51—S9 flagellar segments; 20th segment 1.5-1.8 times as long as broad, that of male tending to be more slender than that of the female Mesoscutum polished, punctate, in profile steeply rounded; notauli vestigial. Mesopleuron polished, punctate but sometimes with some punctostriation centrally; epicnemial carina weak, curved towards anterior margin of pleuron, but with upper end evanescent. Scutellum in profile moderately convex, laterally carinate for most of its length, but often with carinae weak; scutellum in dorsal view 1.4~1.5 times as long as anteriorly broad, generally rather smooth. Metapleuron moderately convex, weakly punctostriate; submetapleural carina slightly broadened anteriorly; posterior transverse carina of mesosternum from complete to broadly indistinct centrally. Propodeum in profile abruptly declivous, dorsally deplanate; anterior transverse carina from complete to completely absent, posterior transverse carina present laterally as short crests; anterior area deeply impressed, striate; spiracular area short, almost smooth; posterior area weakly sculptured, with only a few weak concentric rugae dorsally; lateral longitudinal carina generally complete though occasionally weak posteriorly, not joined to spiracular margin by a short carina. Fore wing length 10-15 mm; fore wing rather more sparsely hirsute than is normal for species in this genus; discosubmarginal cell as in Fig. 237; AI = 0.45—1.17; CI = 0.48-0.57; ICI = 0.39-0.70; SDI = 1.27- 1.46; cu-a somewhat oblique, proximal to the base of Rs&M by about 0.10.2 times its own length; marginal cell proximally slightly more sparsely hirsute; 1st subdiscal cell sparsely hirsute. Hind wing with 7-10 hamuli on R1; 1st abscissa of Rs almost straight, 2nd abscissa weakly bowed. Fore leg with tibia barely flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.3—-1.5 times length of the shorter. Hind leg with coxa in profile 1.5—1.7 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 1.92.0 times as long as broad; claws of female long, weakly curved with close fine pectinae, those of male very similar. Gaster moderately slender; tergite 2 in profile 2.8-3.5 times as long as posteriorly deep, laterotergite folded under, thyridia large, obovate, separated from anterior margin of tergite by about 1.5—2.0 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing fine semierect pubes- cence; gonosquama quite long, acutely rounded. Colour generally yellowish brown, often with orbits and scutellum paler yellowish; interocellar area yellowish; antenna orange, distally slightly infuscate; pterostigma yellowish orange; wings hyaline. VaRIATION. Enicospilus cushmani is a morphologically rather uniform species. The most striking variation is in the degree of pubescence of the wings. Typically this pubescence is sparser than in other species, but in some specimens the central part of the fore wing bears very isolated, short hairs. Such specimens often tend to have slightly thickened veins. 154 IAN D. GAULD RemarkKS. Enicospilus cushmani has not been collected in any country south of the United States, and therefore does not strictly fall within the area of this study. However, it does occur in southern Florida in company with many Neotropical species, and it may occur further south. Furthermore, it is structurally quite similar to E. texanus and some other species that do occur in the region being studied. Therefore I have included it so that it may be differentiated from its relatives. Enicospilus cushmani is probably very closely related to E. texanus. Both species have long, weakly curved tarsal claws (almost certainly a specialized characteristic), sexually dimorphic heads and subquadr- ate to transverse faces (two further probable synapomorphies). It is very easy to separate the typical specimens of E. texanus from E. cushmani since the former species is larger and has infumate wings. However, there are a few specimens of E. texanus (or possibly a very closely related, undescribed species that also parasitizes hemileucine saturniids) that have almost hyaline wings. The following features will enable them to be separated. E. cushmani Hairs in centre of fore wing sparse, separated by more than their own lengths Lateral longitudinal carina of propodeum present behind anterior transverse carina Rs+2r tending to be evenly bowed between pterostigma and 3rs-m Mandibles short and evenly tapered with upper tooth at least 1.8 times length of the lower Parasitoids of Lymantriidae E. texanus Hairs in centre of fore wingclose together, sep- arated by less than their own lengths Lateral longitudinal carina of propodeum absent behind anterior transverse carina Rs+2r more sinuous, the part near to 3rs-m more or less straight Mandibles long, distally parallel-sided, the upper less than 1.7 times length of lower Parasitoids of Saturniidae E. cushmani also superficially resembles a Mexican species, E. halffteri, but the latter species has a narrower face and the claws more abruptly rounded. BIOLOGICAL INFORMATION. Enicospilus cushmani is widely distributed throughout eastern North America, from Ontario south to Highlands county, Florida. It is a common parasitoid of the larvae of Malacosoma americana (Fabricius) and M. disstria (Hiibner). E. cushmani appears to have a single generation per year. It flies as early as January in Florida, though it is most common in March/April and individuals have been collected as late as mid May. Further north it is most common in June. The parasitoid larva apparently kills its host before the caterpillar spins a cocoon. The ichneumonid emerges from its host larva and spins a thick fibrous ovoid cocoon 12-15 mm in length in which it pupates. This cocoon is more whitish and woolly externally than are the cocoons of many species of Enicospilus. MATERIAL EXAMINED Holotype 9, U.S.A.: Florida, Alachua Co., Gainesville ii.1955 (Patton) (USNM). Paratypes. 34 0, 61 9, Canada (Ontario); U.S.A. (Florida, Massachusetts, New Jersey) (BMNH, CNC, FSCA, TC, USNM) as detailed by Gauld (1988). Enicospilus halffteri sp. n. (Figs 97, 238) Description. Mandibles moderately long, evenly tapered, apically twisted 10°; upper mandibular tooth slightly compressed, 1.8 times as long as the lower tooth; outer mandibular surface with a broad shallow proximal concavity, distally centrally rugulose. Labrum 0.3 times as long as broad; malar space 0.5 times as long as basal mandibular width. Clypeus in profile flat, margin quite sharp; clypeus in front view 1.5 times as broad as long, with its margin truncate. Lower face 0.86 times as broad as long, centrally obsoletely punctostriate. Head in dorsal view with genae slightly swollen (Fig. 97); posterior ocellus very close to eye; FI = 63%; occipital carina mediodorsally obsolescent, ventrally inclined towards hypostomal carina but with lower end absent. Antenna moderately slender, with 61 flagellar segments; 20th segment 2.1 times as long as broad. Mesoscutum weakly polished with obsolescent punctures, in profile rather steeply rounded; notauli vestigial. Mesopleuron weakly polished, the upper part punctate, grading ventrally to punctostriate; epicnemial carina inclined towards anterior margin of pleuron, its upper end obsolescent. Scutellum in profile moderately convex, laterally carinate for 0.3 of its length; scutellum in dorsal view 1.4 times as long as anteriorly broad, anteriorly punctate, posteriorly longitudinally striate. Metapleuron strongly convex, coarsely and irregularly striate; submetapleural carina evenly anteriorly broadened; posterior transverse OPHIONINAE OF TROPICAL MESOAMERICA 155 carina of mesosternum strong, with a broad V-shaped cleft centrally. Propodeum in profile fairly abruptly declivous; anterior transverse carina complete except at extreme lateral ends, posterior transverse carina absent; anterior area striate, spiracular area short, finely punctate; posterior area reticulate; lateral longitudinal carina present anteriorly, not joined to spiracular margin by a short carina. Fore wing length 21 mm; discosubmarginal cell as in Fig. 238; AI = 0.56; CI = 0.46; ICI = 0.67; SDI = 1.34; cu-a proximal to base of Rs&M by about its own thickness; marginal cell proximally evenly hirsute; 1st subdiscal cell with anterior 0.5 hirsute. Hind wing with 11 hamuli on R1; 1st abscissa of Rs almost straight, 2nd abscissa straight. Fore leg with tibia weakly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.4 times length of the shorter. Hind leg with coxa in profile 1.8 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 1.9 times as long as broad; claws of male closely pectinate. Gaster moderately slender; tergite 2 in profile 4.6 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 2.5 times its own length. Female unknown. Male with sternites 7—9 bearing fine decumbent pubescence; gonosquama distally acute. Colour generally pale yellowish, mesoscutum with obscure light brownish vittae, gaster posteriorly weakly infuscate; interocellar area yellow; antenna yellowish brown, infuscate at extreme apices; pterostigma golden; wings hyaline. VaRIATION. A second specimen from Chiapas differs from the holotype in being larger (fore wing length 24 mm), having the scutellum more convex, the metapleuron ventrally flattened, the gaster infuscate and the wings infumate. It is only tentatively associated with this species. Remarks. This species is named in honour of Dr Gonzalo Halffter, who has done so much to further conservation biology in Mexico E. halffteri belongs to a complex of species that have Rs+2r weakly sinuous. However, the absence of well-developed scutellar carinae enable E. halffteri to be distinguished easily from the other species. In venation, coloration and general appearance E. halffteri closely resembles E. aktites. However, the genae of E. aktites are narrower and the propodeum is posterodorsally concentrically striate; in E. halffteri the propodeum is posterodorsally reticulate. Furthermore, the pubescence on the male postericr sternites is finer and more decumbent in E. halffteri. BIOLOGICAL INFORMATION. Enicospilus halffteri is only known to occur in the southern Mexican state of Chiapas. A tentatively associated specimen was taken in cloud forest. MATERIAL EXAMINED Holotype ©’, Mexico: Chiapas, junction of highways 190-195, 6.vi.1969 (Howden) (CNC). Non-paratypic material. Mexico: Chiapas: 1 0’, Santa Rosa, cloud forest, 1260 m, v.1967 (Halffter & Reyes) (CNC). Enicospilus sarukhani sp. n. (Figs 96, 239) Description. Mandibles moderately long, rather evenly tapered, apically twisted 20-25°; upper mandibu- lar tooth subcylindrical to slightly compressed, 1.3—1.5 times as long as the lower tooth; outer mandibular surface with fine close pubescence, centrally flat, and with a weak proximal concavity. Labrum 0.3 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile flat, margin subacute; clypeus in anterior aspect 1.41.5 times as broad as long, the margin weakly convex apically. Lower face 0.72-0.80 times as broad as long, centrally obsoletely punctate. Head in dorsal view with genae rounded behind eyes (Fig. 96); posterior ocellus contiguous with eye; FI = 65-70%; occipital carina mediodorsally weak or obsolescent, ventrally evanescent, not curved to join hypostomal carina. Antenna rather stout, with 65—66 flagellar segments; 20th segment 1.5 times as long as broad. Mesoscutum polished, finely punctate, in profile abruptly rounded; notauli obsolescent. Mesopleuron polished, the upper part punctate, the lower part punctate or punctostriate; epicnemial carina with upper end curved towards anterior margin of pleuron. Scutellum in profile quite strongly convex, laterally carinate for most of its length; scutellum in dorsal view 1.4 times as long as anteriorly broad, anteriorly matt, with fine punctures amongst weak rugae, posteriorly becoming strongly longitudinally rugose, with central part slightly raised into a weak crest; postscutellum exceptional in being centrally raised and bearing a weak to strong longitudinal crest. Metapleuron convex, rather smooth and polished, with fine punctures; submetapleural carina quite strongly anteriorly broadened; posterior transverse carina of 156 IAN D. GAULD mesosternum complete. Propodeum in profile fairly abruptly declivous; anterior transverse carina com- plete, sinuous, posterior transverse carina present as strong lateral vestiges; anterior area long, smooth with isolated striae and together with the smooth spiracular area forming a broad shallow depression; posterior area deplanate, with anterior 0.3 behind anterior transverse carina smooth, the part behind this smooth except for a few scattered rugae; lateral longitudinal carina present anteriorly, not joined to spiracular margin by a short carina. Fore wing length 23-24 mm; discosubmarginal cell as in Fig. 239; AI = 0.89-0.94; CI = 0.43-0.51; ICI = 0.66—0.80; SDI = 1.44-1.61; cu-a proximal to base of Rs&M by 0.1—0.2 times its own length; marginal cell proximally more or less evenly hirsute; 1st subdiscal cell with anterior and distal sides hirsute. Hind wing with 9-11 hamuli on R1; Ist abscissa of Rs almost straight, 2nd abscissa almost straight. Fore leg with tibia slightly flattened, with few scattered spines on outer surface. Mid leg with longer tibial spur 1.4 times length of the shorter. Hind leg with coxa in profile 1.6-1.7 times as long as deep; trochantellus dorsally <0.1 times as broad; 4th segment of tarsus 2.2—2.3 times as long as broad; claws of female large, strongly curved with close pectinae. Gaster slender; tergite 2 in profile 6.0—6.3 times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by about 3 times its own length. Ovipositor slender, its sheath narrow. Male unknown. Colour generally yellowish brown, with head paler yellowish, gaster with tergites 3+ infuscate; inter- ocellar area yellowish or whitish; antenna and pterostigma golden; wings hyaline. VARIATION. The specimen from California has Rs+2r basally slightly more incrassate than occurs in the Mexican material. REmMaRKS. This species is named in honour of Dr José Sarukhan, who has done so much for the develop- ment of field biology in Mexico. Enicospilus sarukhani belongs to the complex of species that are characterized by the possession of an almost straight Rs+2r. Within this complex it may be recognized by the combination of the specialized postscutellum and the smooth propodeum. The metapleuron is much smoother and polished than that of other species with a fairly straight Rs+2r. BIOLOGICAL INFORMATION. Enicospilus sarukhani has been taken at high altitude sites between 2300 and 3000 m in Mexico, and a single female has been collected near San Diego, California in the United States. MATERIAL EXAMINED Holotype 9, Mexico: Durango, 16 km W. El Salto, 3000 m, vii.1964 (Martin) (CNC). Paratypes. Mexico: Hidalgo: 1 9, San Vicente, 2300 m, vi.1963 (Woodruff) (FSCA). U.S.A.: California: 1 2, San Diego (USNM). Enicospilus aktites Gauld (Figs 161, 240) Enicospilus aktites Gauld, 1988: 36. Holotype 9, COSTA RICA (BMNH) [examined. ] Description. Mandibles moderately long, strongly narrowed so apex is slender, apically twisted 15-20°; upper mandibular tooth subcylindrical, 1.2-1.4 times as long as the lower tooth; outer mandibular surface with a broad shallow proximo-ventral concavity. Labrum 0.2-0.3 times as long as broad; malar space 0.2— 0.3 times as long as basal mandibular width. Clypeus in profile flat or very weakly convex, margin blunt; clypeus in front view 1.2—1.4 times as broad as long, its margin truncate. Lower face 0.65—0.72 times as broad as long, centrally with distinct fine punctures. Head in dorsal view with genae rounded behind eyes; posterior ocellus contiguous with eye; FI = 65-70% ; occipital carina mediodorsally complete, ventrally not reaching hypostomal carina. Antenna moderately slender, with 55—62 flagellar segments; 20th segment 1.3-1.7 times as long as broad. Mesoscutum weakly polished, closely but shallowly punctate, in profile abruptly rounded; notauli vestigial. Mesopleuron polished, punctostriate, grading more towards striate ventrally; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile weakly to moderately convex, laterally carinate for all of its length; scutellum in dorsal view 1.3—1.5 times as long as anteriorly broad, anteriorly smooth, posteriorly rugulose. Metapleuron convex, generally punctate or coarsely punctostriate, excep- tionally granulate; submetapleural carina generally evenly anteriorly broadened; posterior transverse carina of mesosternum complete or centrally obsolescent. Propodeum in profile abruptly declivous; anterior transverse carina complete, usually centrally raised, occasionally laterally absent; posterior transverse carina from absent to present laterally as a vestige; anterior area short, striate or rarely smooth; OPHIONINAE OF TROPICAL MESOAMERICA SW) spiracular area short, punctate finely; posterior area rugose, the rugae tending to be concentric posterodor- sally (Fig. 161); lateral longitudinal carina present at least anteriorly, not joined to spiracular margin by a short carina. Fore wing length 13-17 mm; discosubmarginal cell as in Fig. 240; AI = 0.69-1.21; CI = 0.36-0.53; ICI = 0.70-1.02; SDI = 1.15-1.32; cu-a from subopposite base of Rs&M to proximal toit by 0.3 times its own length; marginal cell proximally more or less evenly hirsute except for narrow glabrous band adjacent to Rs+2r; 1st subdiscal cell with anterior 0.3—0.5, and posterodistal corner hirsute. Hind wing with S—9 hamuli on R1; 1st abscissa of Rs almost straight, 2nd abscissa straight. Fore leg with tibia slightly flattened, with a few strong spines on outer surface. Mid leg with longer tibial spur 1.3—1.5 times length of the shorter. Hind leg with coxa in profile 1.6-1.7 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.0—2.4 times as long as broad; claws of female large, with short close pectinae, those of male similar. Gaster slender; tergite 2 in profile 3.45.1 times as long as posteriorly deep, laterotergite turned under, thyridia small, obovate and separated from anterior margin of tergite by about 4—5 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing long stout erect hairs; gonosquama apically acute. Colour generally pale yellowish, mesoscutum with weak brownish vittae, gaster slightly infuscate; interocellar area yellowish; antenna golden; pterostigma golden; wings hyaline. VARIATION. Enicospilus aktites is a morphologically rather uniform species except that the larger specimens tend to have the metapleurae more strongly punctate than in smaller individuals. All specimens have a quite distinct and elongate quadra present in the fenestra, but a number of individuals have a second weak quadra paralleling this. Remarks. Enicospilus aktites belongs to a complex of species that are characterized by the possession of an _almost straight Rs+2r. This complex includes E. aktites, E. abelardoi, E. hallwachsae, E. halffteri and E. sarukhani. Of these species E. abelardoi can easily be distinguished by its pyramidal scutellum and large size, and E. halffteri is recognizable because it lacks well-developed scutellar carinae. E. sarukhani is distinctive in having an anteriorly smooth posterior propodeal area. E. aktites most closely resembles the Mesoamerican species E. hallwachsae. Both have similarly modified, rather slender mandibles and large values for AI and ICI, suggesting they may be closely related. E. hallwachsae does not have a distinctly thickened quadra, is darker than E. aktites, and has a more elongate 2nd discal cell BIOLOGICAL INFORMATION. Enicospilus aktites is a very common species in Santa Rosa National Park where individuals have been collected during all months of the year. Pooled data for 1980-S are: ey ieee A IM oD 6 Jo Bet,» Or uN. oD Ae etl OAD, Gi Ore ie Ops oT, Elsewhere this species is apparently very uncommon. I have seen two individuals from Cerro el Hacha, in Guanacaste National Park. Apart from these, and a single specimen collected at Monteverde, Costa Rica in 1962 (and intensive subsequent collecting has failed to find any more examples of this species at this locality) all other (3) specimens have been collected at coastal sites (Map 10). The Guanacaste National Park/Santa Rosa site is within 8 km of the coast, so possibly this species is normally restricted to drier coastal areas and dry forests adjacent to such sites. MATERIAL EXAMINED Holotype 9, Costa Rica: Guanacaste Prov.: Santa Rosa National Park, 300 m, light-trap overlooking deciduous forest, vi.1985 (Gauld) (BMNH) Paratypes. Belize: 1 o’, Punta Gorda, x.1913 (Norton) (BMNH). Costa Rica: Guanacaste Prov.: 27 C’, 48 2, Santa Rosa National Park, 300 m, months as enumerated above, 1980-85 (Janzen & Hallwachs) (BMNH, CNC, TC, USNM): Puntarenas Prov.: 1 9, Monteverde, ii.1962 (Palmer) (TC). Mexico: Tamaulipas: 1 9, Municip. de Aldama, Barra Coma, v.1979 (Gicca) (FSCA). U.S.A.: Florida: 1 CO’, Monroe Co., No Name Key, vi.1974 (Heppner) (FSCA). Non-paratypic material. Costa Rica: Guanacaste Prov.: 1 o’, 1 2, Guanacaste National Park, Cerro el Hacha, 400 m, x-xi.1987 (Chacon) (BMNH). Enicospilus gomezpompai sp. n. (Figs 162, 241) DescriPTion. Mandibles moderately long, proximally abruptly narrowed, distally rather slender and parallel-sided, apically twisted 10—20°; upper mandibular tooth very slightly compressed, 1.4—1.6 times as long as the lower tooth; outer mandibular surface with fine, scattered pubescence, almost flat centrally, IAN D. GAULD 158 ‘payoa]joo usaq sey sayy snjidsooiuq yoy ye sanyesoT oT dep OPHIONINAE OF TROPICAL MESOAMERICA 159 and with a shallow proximal concavity. Labrum 0.2 times as long as broad; malar space 0.2-0.3 times as long as basal mandibular width. Clypeus in profile from almost flat to weakly convex, margin blunt to subacute; clypeus in front view 1.4-1.6 times as broad as long, its margin weakly convex. Lower face 0.69- 0.74 times as broad as long, centrally with obsolescent punctures. Head in dorsal view with genae rounded behind eyes; posterior ocellus contiguous with eye; FI = 70-75% ; occipital carina mediodorsally complete, usually weak, sometimes very faint and obsolescent, ventrally evanescent, not joining hypostomal carina. Antenna long and slender, with 64-66 flagellar segments; 20th segment 1.8—2.0 times as long as broad. Mesoscutum weakly polished with fine punctures, in profile steeply rounded; notauli vestigial. Meso- pleuron polished, the upper part punctate ventrally grading to punctostriate; epicnemial carina with upper end weak but curved towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.7 or more of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, anteriorly punctate with microreticulations, posteriorly slightly rugose. Metapleuron weakly convex with fine punctures on a generally weakly shagreened surface; submetapleural carina usually evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile abruptly declivous; anterior transverse carina weak but usually complete, posterior transverse carina vestigial; anterior area quite long, with a few wrinkles but not striate; spiracular area short, smooth; posterior area dorsally deplanate, with weak irregular sculpture (Fig. 162); lateral longitudinal carina present only as a vestige anteriorly, not joined to spiracular margin by a short carina. Fore wing length 17-22 mm; discosubmarginal cell as in Fig. 241; AI = 0.42-0.75; CI = 0.55-0.66; ICI = 0.49-0.55; SDI = 1.28-1.35; cu-a proximal to the base of Rs&M by 0.1-0.3 times its own length; marginal cell proximally very narrowly glabrous; 1st subdiscal cell anteriorly and distally sparsely hirsute. Hind wing with 8-11 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa weakly bowed. Fore leg with tibia weakly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.3—-1.4 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.4~2.6 times as long as broad; claws of female strongly curved, with long stout pectinae, those of male similar but with pectinae slightly finer. Gaster slender; tergite 2 in profile 5.5—6.0 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 2.5-3.5 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing fine, erect hairs; gonosquama long, distally evenly rounded. Colour generally pale yellowish brown, head yellow, mesoscutum centrally weakly infuscate; interocel- lar area yellow; antenna golden; pterostigma golden; wings hyaline. VARIATION. The specimens from Cordoba, Veracruz and Lagos des Calores, Chiapas differ from the series collected in Guerrero in that tergites 3+ of the gaster are somewhat infuscate. A second specimen from Chiapas differs from the other material in being very much darker, with blackish antennae and gaster and dark brownish alitrunk. This specimen has the metapleuron more punctate than normal and the mandibles more twisted. The male from Veracruz has the propodeum more coarsely sculptured than normal and the U-shaped alar sclerite is very weak. REMARKS. This species is named in honour of Dr Arturo Gomez-Pompa, who has aggressively developed Mexican biology for more than two decades. Enicospilus gomezpompai may most easily be recognized by the possession of a very large fenestra bounded by a weak, almost U-shaped vestigial sclerite. The subgenital plate of the female is larger than in many allied species. I have excluded the dark specimen from the paratype series as I am not convinced it is conspecific. However, with only a single specimen to hand any other placement would also have been uncertain and in this complex of closely related species conservatism at the species-level is essential BIOLOGICAL INFORMATION. Enicospilus gomezpompai has only been collected at moderately high altitude in southern Mexico. Nothing is known about its biology. MATERIAL EXAMINED Holotype ?, Mexico: Guerrero, Amula, 2000 m, viii.1904 (Smith) (BMNH). Paratypes. Mexico: Chiapas: 1 9, Rt 17, Lagos des Calores, v.1969 (Howden) (CNC): Guerrero: 1 0’, 3 2, Amula, 2000 m, viii, ix.1904 (Smith) (BMNH); 1 9, Xucumanatlan, 2300 m, vii.1904 (Smith) (BMNH): Veracruz: 1 0’, Cordoba, x.1963 (Lau) (RNH). Non-paratypic material. Mexico: Chiapas: 1 9, San Cristobal de las Casas, v.1964 (Martin) (CNC). 160 IAN D. GAULD Enicospilus lebophagus Gauld (Figs 163, 242) Enicospilus lebophagus Gauld, 1988: 45. Holotype 2, COSTA RICA (BMNH) [examined]. DEscrIPTION. Mandibles of moderate length, proximally strongly narrowed, distally more weakly so, apically twisted 15—25°; upper mandibular tooth subcylindrical, 1.2-1.5 times as long as the lower tooth; outer mandibular surface finely and sparsely hirsute, though more densely hirsute on proximoventral lobe; outer surface flat centrally, with a broad, shallow proximal concavity. Labrum 0.2-0.3 times as long as broad; malar space 0.3-0.4 times as long as basal mandibular width. Clypeus in profile flat, at very most only slightly out-flared, margin blunt; clypeus in front view truncate or weakly convex apically, 1.4-1.6 times as broad as long. Lower face 0.66—0.88 times as broad as long, centrally polished, virtually smooth. Head in dorsal view with genae short, rounded behind the eyes; posterior ocellus contiguous with eye; FI = 70-80%; occipital carina mediodorsally from complete to narrowly interrupted, ventrally curved to approach, but not actually join hypostomal carina about 1.0 times the basal mandibular width away from mandible. Antenna long and quite slender, with 60-67 flagellar segments; 20th segment 1.6—2.0 times as long as broad. Mesoscutum polished, finely punctate, in profile evenly rounded; notauli very shallow, but extending back to level of hind corner of pronotum. Mesopleuron polished, the upper part punctate to punctostriate, the lower part punctostriate to striate; epicnemial carina with upper end weak, curved towards anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.3—1.5 times as long as anteriorly broad, anteriorly generally relatively smooth with weak to very strong longitudinal striae developed posteriorly. Metapleuron moderately convex, varying considerably in sculpture from punctate or punctostriate to rugulose; submetapleural carina quite narrow, not or only slightly broadened anteriorly; posterior transverse carina of mesosternum from complete to with central part effaced. Propodeum in profile rather abruptly declivous, posterodorsally deplanate; anterior transverse carina usually complete, posterior transverse carina present only as a vestigial keel; anterior area deeply impressed, striate, spiracular area short and smooth or punctate, posterior area coarsely rugose to rugose-reticulate, often with the rugae concentric posteriorly; lateral longitudinal carina present anteriorly, and often extending back for half the length of the propodeum, not joined to spiracular margin by a short carina. Fore wing length 20-25 mm; discosubmarginal cell as in Fig. 242; AI = 0.77-1.61; CI = 0.55-0.74; ICI = 0.60-0.90; SDI = 1.35-1.61; cu-a proximal to the base of Rs&M by 0.1-0.3 times its own length; marginal cell proximally fairly evenly hirsute, at most with a narrow glabrous band adjacent to Rs+2r; 1st subdiscal cell with at least the anterior 0.3 hirsute, often almost entirely hirsute except for the posteroproxi- mal part. Hind wing with 9-13 hamuli on R1; Ist abscissa of Rs straight to weakly bowed, 2nd abscissa almost straight. Fore leg with tibia weakly flattened, with numerous scattered spines on outer surface. Mid leg with longer tibial spur 1.1—1.2 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.8—3.0 times as long as broad; claws of female long, abruptly curved with long, strong close pectinae, those of male similar but with pectinae slightly shorter and closer together. Gaster moderately slender; tergite 2 in profile 3.54.0 times as long as posteriorly deep, laterotergite folded under, thyridia oval to obovate and separated from anterior margin of tergite by about 2-4 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing moderately long, very stout erect hairs (Fig. 163); gonosquama apically subacute. Colour generally orange-brown, sometimes with the orbits yellow; interocellar area orange; antenna brownish orange; pterostigma orange; wings hyaline or weakly infumate. VARIATION. Morphologically this is a very uniform species. Occasional individuals may have the gaster slightly infuscate, but I have not seen a single specimen with the gaster black posteriorly. All specimens have two thickened patches (quadrae) in the alar fenestra, but the degree to which these are pigmented is somewhat variable. A single, possibly conspecific female from Barro Colorado, has rather short mandibles. Remarks. Enicospilus lebophagus is a large species that has frequently been confused with E. americanus, which it closely resembles in size and general appearance. However, the two differ consistently in a number of morphological features, are apparently allopatric and have different hosts. The following table contrasts the critical characters of the two taxa. OPHIONINAE OF TROPICAL MESOAMERICA 161 E. lebophagus Genae evenly rounded behind eyes Sternites 6-9 of male bearing numerous stout truncated erect hairs Fenestra with two distinct quadrae Clypeus weakly out-flared or flat, even in large specimens E. americanus Genae barely narrowed behind eyes Sternites 6-9 of male bearing scattered fine erect hairs, or hairs with close stout hairs (var. 1) Fenestra without quadrae, or with an indistinct one Clypeus usually strongly out-flared, in small speci- mens almost flat Flagellum stout, central segments 1.41.7 times as long centrally broad Flagellum slender, central segments 1.6—2.0 times as long as centrally broad BIOLOGICAL INFORMATION. Enicospilus lebophagus occurs in the extreme south of Texas in Cameron and Hidalgo counties, and thence southwards throughout Central America to northern Costa Rica (Map 11). I have examined extensive collections from Panama and southern Costa Rica, but I have only found one questionable specimen of this species to occur south of about 11°N. In Santa Rosa National Park, Costa Rica, this species has been taken at light in the following months (cumulative total 1979-1985) Pir MA Mot InyA& § 0 »-D Se ee tc ar 28 a2 Oe Me ed It is acommon solitary endoparasitoid of the larvae of Rothschildia lebeau lebeau (Guérin-Ménéville) and R. lebeau forbesi (Benjamin). Oviposition is apparently into a relatively mature larva and the ophionine destroys its host caterpillar after the host has constructed a cocoon. The Enicospilus cocoon is a typical ovoid ophionine cocoon, but it is spun within the cocoon of the saturniid. The adult parasitoid emerges by pushing out through the cocoon entrance, just as does the adult moth. MATERIAL EXAMINED Holotype 9, Costa Rica: Guanacaste Prov.: Santa Rosa National Park, 300 m, ex Rothschildia lebeau, 83.SRNP.123 (Janzen) (BMNH) Paratypes. Costa Rica: Guanacaste Prov: 1 0’, W. of Carmona Nicoya, 6-700 m, viii.1982 (Janzen & Hallwachs) (BMNH); 30 0’, 37 2, Santa Rosa National Park, 300 m, ex Rothschildia lebeau 1979-1985 (Janzen) (BMNH, CAS, CNC, MCZ, PANS, TC, USNM); 10", 1 2, same locality, v.1979 (Janzen) (TC); 1 Q, vi.1979 (Janzen) (BMNH); 1 0’, 1 Q, vi.80; 2 Q, viii.1980; 1 9, vii.1981; 1 9, vi.1982; 1 CO’, vii.1982; 10, x.1982; 1 2, xii.1982; 1 2, vi.1983; 1 0, ix.1983; 3 9, vi.1984; 107, 1 9, vi.1985 (Janzen & Hallwachs) (BMNH); 1’, vi.1985 (Gauld) (BMNH). Guatemala: 1 9, Solola, 5 km NE. Panajachel, 1740 m, viii.1975 (Fisher) (CAS). Honduras: 1 ©’, Trujillo, vii. 1968 (Dozier) (FSCA). Mexico: Baja California: 1 0’, 10 km W. Santiago, viii.1959 (Radford & Werner) (CAS): Chiapas: 1 0’, 2 2, 30km N. Huixtla, 1000 m, vi.1969 (Mason & Peterson) (CNC); 1 2, 20 km SW. El Salto, 2300 m, vii.1964 (Madson) (CNC): Guerrero: 1 0’, 1 9, Xucumanatlan, 2300 m, vii.1904 (Smith) (BMNH): San Luis Potosi: 1 0’, 1 2, El Salto Falls, vi.1963 (Woodruff) (FSCA): Tamaulipas: 1 0’, Hacienda Sta Eugracra, vii.1939 (Haag) (MCZ): 2 2, ‘Mexico’, no further data (MCZ). U.S.A.: Texas: 1 2, Brownsville, x.1916 (Vickery) (PANS); 3 9, Harlingen, ex Rothschildia lebeau forbesi, em. 111.1981, v.1981, x.1981 (Peigler) (BMNH); 1 Oo’, Hidalgo Co., Bentsen- Rio Grande State Park, ex R. /. forbesiem. x.1981 (Peigler) (BMNH); 1 0’, 2 2, Hidalgo Co. south, ex R. 1. forbesi, em. ii.1980 (indoors) (Agnew & Eger) (BMNH); 1 9, same data, em. ix.1980 (Eger) (BMNH). Non-paratypic material. Panama: 1 9, Barro Colorado Island, vi.1978 (Wolda) (RNH). Enicospilus ugaldci sp. n. (Figs 164, 243) | Description. Mandibles moderately long, proximally strongly narrowed, distally weakly tapered, apically __ twisted 15—25°; upper mandibular tooth slightly compressed, 1.3-1.5 times as long as the lower tooth; outer | mandibular surface- with fine sparse hair, more or less flat centrally, with a weak proximal concavity. _ Labrum 0.2-0.3 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile flat to somewhat out-flared, margin flat, subacute; clypeus in front view 1.3-1.5 times as broad as long, the margin truncate to slightly concave (Fig. 164). Lower face 0.71-0.78 times as broad as long, centrally weakly punctate, rarely punctostriate. Head in dorsal view with gena rounded behind the eyes; posterior ocellus contiguous with eye; FI = 60-65%; occipital carina mediodorsally from complete to IAN D. GAULD 162 "poyda][09 usaq sey snsvydogay snjidsoaiuq yoy ye saytesoT =, dew OPHIONINAE OF TROPICAL MESOAMERICA 163 narrowly interrupted, ventrally not curved to, but very nearly reaching or actually joining the hypostomal carina about 0.9 times the basal mandibular width away from mandible. Antenna long and slender, with 61-71 flagellar segments; 20th segment 2.1—2.3 times as long as broad. Mesoscutum polished, with close shallow punctures, in profile evenly rounded, notauli vestigial. Mesopleuron polished, the upper part weakly punctostriate, the lower part more strongly punctostriate, sometimes almost punctoreticulate; epicnemial carina with upper end curved towards but not reaching anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for all of its length; scutellum in dorsal view 1.4-1.6 times as long as anteriorly broad, anteriorly smooth but strongly longitudinally wrinkled posteriorly. Metapleuron moderately convex, matt, punctogranulate with traces of striae; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum slightly sinuous, complete. Propodeum in profile abruptly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area short, deep, striate; spiracular area short and more or less smooth; posterior area, deplanate, coarsely rugose/reticulate, posteriorly with rugae tending to be con- centric; lateral longitudinal carina present anteriorly, not joined to spiracular margin by a short carina. Fore wing length 17-19 mm; discosubmarginal cell as in Fig. 243; AI = 0.74-1.05; CI = 0.49-0.55; ICI = 0.58-0.74; SDI = 1.39-1.45; cu-a proximal to the base of Rs&M by 0.1 or more times its own length; marginal cell proximally virtually glabrous adjacent to Rs+2r; 1st subdiscal cell anteriorly and posterodistally hirsute. Hind wing with 8-10 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa weakly bowed. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.2-1.4 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.4-2.8 times as long as broad; claws of female long, strongly curved with long stout pectinae, those of male similar. Gaster slender; tergite 2 in profile 5.2-6.2 times as long as posteriorly deep, laterotergite folded under, thyridia obovate to rather broadly elliptical, separated from anterior margin of tergite by about 34 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing long stout erect hairs; gonosquama distally evenly rounded. Colour generally brownish orange with head, scutellum and sometimes also with mesoscutal stripes yellowish, gaster irregularly weakly infuscate, unusual in having the ovipositor sheaths brownish yellow, at most slightly infuscate apically; interocellar area yellow; antenna orange; pterostigma golden; wings hyaline. VARIATION. One specimen from Chiapas is paler than the others. It has pronounced dark vittae on the mesoscutum. A male from Volcan Orosi, Costa Rica resembles other specimens of this species in the structure of the mandible and clypeus, having a broad face, and in general sculpture, but differs in having the fenestra somewhat shorter. It is tentatively included here, but excluded from the paratype series. REMARKS. This species is named in honour of Senor Ugalde, the first director of Santa Rosa National Park, a person who has devoted his life to the growth of Costa Rica’s national park system. Enicospilus ugaldei belongs to the E. americanus species-complex. It may be distinguished from other taxa by the long fenestra which extends nearly to the base of Rs, and by the long sinuation on Rs+2r. The clypeus of this species is very subtly (but invariably) different from other species in this complex in being either truncate, or more usually slightly convex. The clypea of most species are very slightly convex in anterior aspect. Another unusual feature of this species is that the lower end of the genal (occipital) carina virtually meets or actually joins the hypostomal carina. In most species of the E. americanus complex the lower end of the genal carina is evanescent and it does not reach the hypostomal carina. BioLoGicaL INFORMATION. Enicospilus ugaldei is an uncommon Mesoamerican species whose range extends from central Mexico south to north-western Costa Rica. It has been reared once by D. H. Janzenin Santa Rosa National Park from the hemileucine saturniid Automeris tridens (Herrich-Schaeffer) (81.SRNP.892) (known in older literature as A. rubrescens (Walker)). This moth, whose larvae feed on a variety of conspicuous trees (see Janzen, 1982), is one the most common species of Automeris in Santa and though it has often been reared only a single instance of parasitism by E. ugaldei has been observed. MATERIAL EXAMINED Holotype ©’, Costa Rica: Guanacaste Prov., Santa Rosa National Park, ex Automeris tridens, 1981 (Janzen & Hallwachs) (BMNH). Paratypes. Costa Rica: Guanacaste Prov.: 1 9, Cerro el Hacha, 3-400 m, xi.1986-i.1987 (Janzen & Hallwachs) (BMNH); 1 o’, Cerro el Hacha, Casa Oeste, 400 m, x.1987 (Chacon) (BMNH); 1 oc’, Santa Rosa National Park, 300 m, xii.1980 (Janzen & Hallwachs) (BMNH). Mexico: Chiapas: 1 9, 40 km N. of Huixtla, 1000 m, vi.1969 (Peterson) (CNC); 1 2, Huixtla, Muste, 440 m, ix.1970 (Welling) (CNC): 164 IAN D. GAULD Quintana Roo: 1 9, X-can, viii.1963 (Welling) (CNC): San Luis Potosi: 2 0’, El Salto Falls, vi.1963 (Woodruff) (FSCA). Non-paratypic material. Costa Rica: 1 &’, Guanacaste Prov.: Volcan Orosi, Casa Mariksa [Maritza], 700 m, v.1986 (Gauld) (BMNH). Enicospilus umanai sp. n. (Figs 165, 244) Description. Mandibles moderately long, proximally abruptly narrowed, distally more or less parallel- sided, apically twisted 20—-25°; upper mandibular tooth subcylindrical to slightly compressed, 1.3—1.5 times as long as the lower tooth; outer mandibular surface bearing fine sparse hairs, almost flat. Labrum 0.2-0.3 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile very weakly convex, margin flat, subacute; clypeus in front view 1.5—1.7 times as broad as long, with margin weakly convex. Lower face 0.78—0.82 times as broad as long, centrally sparsely punctate. Head in dorsal view with genae rounded behind eyes; posterior ocellus contiguous with eye; FI = 70-75%; occipital carina mediodorsally weak or interrupted, ventrally abruptly curved to approach but not join the hypostomal carina about 1.0 times the basal mandibular width away from mandible. Antenna long and quite stout, with 66-68 flagellar segments; 20th segment 1.7—1.9 times as long as broad. Mesoscutum polished, with sparse shallow punctures, in profile evenly but steeply rounded; notauli vestigial. Mesopleuron polished, the upper part punctate, the lower part closely punctate, rarely punc- tostriate; epicnemial carina curved towards but not reaching anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for most of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, obsoletely punctate, sometimes a little wrinkled posteriorly. Metapleuron characteristically convex posterodorsally and flattened anteroventrally, closely punctate (Fig. 165); sub- metapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum strong laterally, medially produced slightly into a rounded lobe, often weak or obsolescent either side of this lobe. Propodeum in profile abruptly declivous; anterior transverse carina complete or interrupted later- omedially, posterior transverse carina absent; anterior area deeply impressed, striate; spiracular area short, almost smooth; posterior area irregularly rugose to reticulate; lateral longitudinal carina present anteriorly, not joined to spiracular margin by a short carina. Fore wing length 21-23 mm; discosubmarginal cell as in Fig. 244; AI = 0.54-0.98; CI = 0.44-0.52; ICI = 0.64—0.83; SDI = 1.23—1.40; cu-a proximal to base of Rs&M by 0.1-0.3 times its own length; marginal cell proximally with a narrow glabrous band; 1st subdiscal cell anteriorly and distally broadly hirsute. Hind wing with 9-11 hamuli on R1; 1st abscissa of Rs almost straight, 2nd abscissa weakly bowed. Fore leg with tibia barely flattened, with scattered stout spines on outer surface. Mid leg with longer tibial spur 1.2—1.5 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.2—2.3 times as long as broad; 4th segment of tarsus 2.2—2.3 times as long as broad; claws of female stout, abruptly curved, with long close pectinae, those of male similar but with pectinae shorter. Gaster slender; tergite 2 in profile 5.0-5.7 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 34 times its own length. Ovipositor apically very slender, its sheath narrow. Male with sternites 7-9 bearing fine scattered erect hairs; gonosquama long, distally subtruncate. Colour generally yellowish brown, generally with mesoscutum bearing dark brownish vittae and with gastral tergites and sternites irregularly infuscate; interocellar area yellow; antenna golden; pterostigma yellowish brown; wings hyaline. VARIATION. In most individuals the mesopleuron is regularly punctate, with the punctures closer ventrally. The female from Mexico has the lower part of the mesopleuron distinctly punctostriate. REMARKS. This species is named in honour of Dr Alvaro Umana, who has done so much to organize the governmental care of the conservation cause in Costa Rica. Enicospilus umanai closely resembles E. americanus and several other large species. However, unlike many others in the E. americanus complex none of the specimens of this species has any distinct trace of concentric striae or rugae present on the propodeum. Concentric striae or rugae are almost always present in E. americanus. A further characteristic feature of E. umanai is the distribution of hairs on the fore wing. In E. umanai the extreme anterior corner of the discosubmarginal cell is uniformly hirsute, whilst in E. americanus a small glabrous area is usually present. The convexity of the metapleuron differs in the two species; that of E. americanus is far more uniform, whilst in E. umanai the greatest convexity is along the posterodorsal margin, and anteroventrally the metapleuron is flattened. E. umanai is unusual in often | ea ee ee, re = eineTasaeetsee seo OPHIONINAE OF TROPICAL MESOAMERICA 165 having the distal corner of the discosubmarginal cell very acute (50-55°); in most species this corner is greater than 60°. | BIOLOGICAL INFORMATION. Enicospilus umanai is a Mesoamerican species whose range extends from Durango, Mexico, south to Costa Rica. In Costa Rica it is only known to occur in lower montane wet forest at altitudes between 1000 and 1400 m. At Monteverde it has only been collected between November and February even though most collecting at this locality has been undertaken between April and June. The pooled monthly catch is: Tete hhee Alle tele ALS CN OD 5 I i Br mabe 0 Nothing is known of its biology. MATERIAL EXAMINED Holotype 9, Costa Rica: Puntarenas Prov., Monteverde, 1300 m, xi.1985 (Haber) (BMNH). Paratypes. Costa Rica: Puntarenas Prov.: 20", 5 9, Monteverde, 1350 m, xi, xii.1961, i, ii. 1962 (Palmer) (TC); 1 2, same locality, xii.1979 (Janzen) (BMNH); 1 Q, same locality, xii.1985 (Haber) (BMNH). Mexico: Chiapas: 1 ©’, San Cristobal, viii.1969 (Kritsch) (CNC): Durango: 1 9, Tepalcates, 50 km W. Durango, vii.1964 (Howden) (CNC). Enicospilus quintanai sp. n. (Fig. 245) Description. Mandibles moderately long, quite stout being proximally weakly tapered and distally more or less parallel-sided, apically twisted about 30°; upper mandibular tooth slightly compressed, 1.3-1.5 times as long as the lower tooth; outer mandibular surface more or less flat with scattered pubescence. Labrum 0.3 times as long as broad; malar space 0.30.4 times as long as basal mandibular width. Clypeus in profile flat, margin acute; clypeus in front view 1.4—1.5 times as broad as long, the margin truncate to weakly convex. Lower face 0.64-0.71 times as broad as long, finely granulate. Head in dorsal view with genae rounded behind eyes; posterior ocellus contiguous with eye; FI = 68-73%; occipital carina medi- odorsally weak or narrowly interrupted, ventrally curved to almost join hypostomal carina about 0.8 times the basal mandibular width away from mandible. Antenna long and slender, with 66—68 flagellar segments; 20th segment 1.9-2.0 times as long as broad. Mesoscutum weakly polished, finely granulate, in profile abruptly rounded; notauli vestigial. Meso- pleuron weakly polished, the upper part obsoletely puncto-granulate, the lower part punctostriate; epicnemial carina inclined towards, but not reaching the anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.9 ofits length; scutellum in dorsal view 1.4 times as long as anteriorly broad, granulate. Metapleuron weakly convex, granulate; submetapleural carina evenly anteriorly broad- ened; posterior transverse carina of mesosternum obsolescent on midline. Propodeum in profile abruptly declivous; anterior transverse carina sinuous, complete, posterior one vestigial; anterior area short and deeply impressed, with isolated striae; spiracular area short, smooth; posterior area coarsely rugose, with the rugae tending to be concentric posteriorly; lateral longitudinal carina from complete to present only anteriorly, not joined to spiracular margin by a distinct short carina. Fore wing length 16-18 mm; discosubmarginal cell as in Fig. 245; AI = 0.68-0.95; CI = 0.52-1.10; ICI = 1.09-1.75; SDI = 1.47-1.59; cu-a proximal to the base of Rs&M by about 0.1-0.2 of its own length; marginal cell proximally slightly more sparsely pubescent adjacent to Rs+2r; 1st subdiscal cell with anterior 0.3—-0.4 sparsely hirsute. Hind wing with 9-10 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa straight. Fore leg with tibia weakly flattened, with scattered stout spines on outer surface. Mid leg with longer tibial spur 1.3-1.4 times length of the shorter. Hind leg with coxa in profile 1.6-1.7 times as long as deep; trochantellus dorsally obscured; 4th segment of tarsus 2.5-2.6 times as long as broad; claws of male abruptly rounded, with short, close pectination. Gaster moderately slender, though a little stouter than many related species; tergite 2 in profile 3.3-3.7 times as long as posteriorly deep, laterotergite folded under, thyridia oval and separated from anterior margin of tergite by about 1.5 times its own length. Female unknown. Male with sternites 7-9 bearing close long stout erect hairs; gonosquama apically quite acutely pointed. Colour generally yellowish orange; interocellar area yellowish; antenna slightly infuscate apically; pterostigma orange; wings more or less hyaline. VARIATION. None remarkable. 166 IAN D. GAULD RemaRKS. Enicospilus quintanai can most easily be distinguished from others in the E. americanus complex by the large ICI, its rather matt appearance, the densely hirsute male sternites and its slightly stouter gaster. Structurally it is quite similar to E. ugaldei from which it differs in being less polished and not punctate, generally having a narrower face and shorter fenestra, and in having the larger ICI. The metapleuron of E. ugaldei is more evenly convex than that of E. quintanai. BIOLOGICAL INFORMATION. Only three isolated specimens of Enicospilus quintanai are known. All were collected in lowland wet forest. Nothing is known about its biology. MATERIAL EXAMINED Holotype 0’, Mexico: Quintana Roo, X-can, vi.1962 (Welling) (CNC). Paratypes. Panama: 1 ©’, Barro Colorado Island, xi.1978 (Wolda) (RNH); 1 0’, same locality, iv.1979 (Wolda) (TC). Enicospilus americanus (Christ) (Figs 98, 246) [Ichneumon macrurus Linnaeus; Drury, 1773: 1. Misidentification. ] Ichneumon luteus americanus Christ, 1791: 358. Holotype 9, U.S.A.: New York (lost). [Ophion macrurum (L.) Westwood in Drury, 1837: 92. Misidentification. | Ophion rugosus Brullé, 1846: 138. Holotype 2, ‘North America’ (MNHN). [Synonymized by Hooker, 1912: 148.] [Ophion cecropiae Scudder, 1863: 188. Nomen nudum. | Ophion cecropiae Sanborn, 1863: 169. Holotype ? sex, U.S.A.: Mass. (lost). [Ophion undulatus Gravenhorst; Taschenberg, 1875: 430. Misidentification. ] Eremotylus Druryi Kriechbaumer, 1901c: 152. Syntypes 3 2, 1 &', U.S.A.: New York (ZSBS). [Syn- onymized by Hooker, 1912: 149.] [Eremotylus macrurus (L.) Felt, 1904: 101. Misidentification. | [Allocamptus macrurus (L.) Morley, 1912: 24. Misidentification. ] [Enicospilus macrurus (L.) Essig, 1926: 792. Misidentification. ] Enicospilus americanus (Christ) Townes, 1945: 737. Description. Mandibles of moderate length, proximally strongly narrowed, distally almost parallel-sided, apically twisted 15—30°; upper mandibular tooth subcylindrical, 1.2—1.5 times as long as the lower tooth; outer mandibular surface finely and sparsely hirsute centrally, more densely hirsute on proximoventral lobe; outer surface flat centrally, with a broad shallow proximal concavity. Labrum 0.2-0.4 times as long as broad; malar space 0.2-0.3 times as long as basal mandibular width. Clypeus in profile usually distinctly out-flared, margin relatively blunt; clypeus in front view 1.4-1.6 times as broad as long, the margin truncate. Lower face 0.75—0.90 times as broad as long, centrally with scattered fine punctures, the area between the punctures often finely microreticulate. Head in dorsal view with genae barely narrowed behind eyes, in larger individuals distinctly buccate; posterior ocellus contiguous with eye; FI = 70-80%; occipital carina mediodorsally from complete and slightly dipped, to narrowly incomplete, ventrally curved to approach but not actually join hypostomal carina about 1.0 times the basal mandibular width away from mandible. Antenna long but relatively stout, with 63-77 flagellar segments; 20th segment 1.4— 1.7 times as long as broad. Mesoscutum polished, with close shallow punctures, in profile evenly rounded; notauli very shallow. Mesopleuron weakly polished, the upper part punctate to punctostriate, the lower part punctostriate; epicnemial carina curved towards anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.3—1.5 times as long as anteriorly broad, anteriorly generally punctate grading to striate posteriorly. Metapleuron moderately convex, rather variable, from regularly punctate to puncto-reticulate, irregularly rugose or even somewhat striate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile rather abruptly declivous, posterodorsally deplanate; anterior transverse carina complete or effaced lateromedially, posterior transverse carina generally discernible as a lateral crest; anterior area deeply impressed, striate, spiracular area very short, smooth or punctate, posterior area coarsely rugose, often with rugae concentric posterodorsally; lateral longitudinal carina present only anteriorly, not joined to spiracular margin by a short carina. Fore wing length 18-26 mm; discosubmarginal cell as in Fig. 246; AI = 0.83-1.55; CI = 0.41-0.69; ICI = 0.50-0.80; SDI = 1.20-1.35; cu-a from subopposite the base of Rs&M, to proximal to it by about 0.3 times its own length; marginal cell proximally more or less uniformly hirsute; 1st subdiscal cell broadly hirsute. Hind wing with 10-12 hamuli on R1; 1st abscissa of Rs almost straight, 2nd abscissa weakly bowed. OPHIONINAE OF TROPICAL MESOAMERICA 167 Fore leg with tibia very weakly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.2—1.4 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.42.9 times as long as broad; claws of female strongly curved with long strong, close pectinae (Fig. 98), those of male similar but with pectinae shorter. Gaster moderately slender; tergite 2 in profile 3.7—5.8 times as long as posteriorly deep, laterotergite turned under, thyridia obovate and separated from anterior margin of tergite by about 2-3 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing fine erect pubescence; gonosquama long, distally rounded. Colour generally brownish orange with the head paler yellowish; interocellar area yellow; antenna orange; pterostigma brownish or yellowish orange; wings from very weakly to quite strongly yellowish. VARIATION. There is a considerable range of morphological variation within this species, and it is possible that more than one sibling species may be confused here. Typical examples of E. americanus are large insects (fore wing length 23+ mm) with slightly yellow wings, the clypeus out-flared and the male only has scattered fine erect hairs on the posterior gastral sternites. In the southern part of the U.S.A. and northern Mexico, the specimens have almost hyaline wings. A number of smaller individuals have the clypeus almost flat, but these are otherwise morphologically more or less indistinguishable from typical E. americanus and have the same host range. I conclude that these are simply small individuals of the species. The sculpture of the metapleuron is so variable as to be of no value in characterizing the species The major taxonomic problem concerns a group of male specimens (fore wing length 19-23 mm) that have long stout dense pubescence on the posterior sternites. In all other species I have seen, I have never encountered variation of this extent in this character, and this suggests that this group of specimens represents a different species. I have seen a series of it from Aurora, Colorado, U.S.A. (Peigler, 1985) reared from Automeris io, a hemileucine saturniid (all host records for typical E. americanus are from saturniine saturniids) further suggesting that it may be a separate species. However, I have not been able to find characters which would reliably allow me to separate the females (reared from Automeris) from females of typical E. americanus. I (Gauld, 1988) opted to treat the ‘hairy males’ as an unnamed variety of E. americanus, pending further study. Remarks. Until the publication of Townes’ (1945) catalogue, this species was generally known as Eremotylus or Ophion macrurus. The true macrurus of Linnaeus is a species of Megarhyssa (Townes & Townes, 1960; Fitton, 1978). E. mexicanus (Cresson) has been treated as a synonym of E. americanus since 1912, but examination of the holotype has shown that E. mexicanus is a distinct species (see p. 142) Enicospilus americanus has been confused with other species of the genus both in the United States and in Latin America. In the north it has not clearly been separated from either E. texanus or E. glabratus. It differs from the former species in being larger, having more strongly curved claws, a generally narrower face and narrower malar space, and longer antennae. The curvature of Rs+2r in the fore wing differs between the two species (see Figs 236, 237, 246). E. americanus differs from E. glabratus in the form of the alar fenestra and in not having a distinctive cluster of hairs proximal to this fenestra (see Figs 224, 246). The clypeus of E. glabratus is never out-flared and E. americanus has the lower end of the genal carina obsolescent, not joining the hypostomal carina as does that of E. glabratus. In tropical America E. americanus has been confused with a group of other species that include E. mexicanus, E. tenuigena, E. lebophagus, E. brevis and other taxa without alar sclerites. The present key should allow these taxa to be differentiated. BIOLOGICAL INFORMATION. Enicospilus americanus is a common species throughout much of eastern North America. It range extends into southern Canada and west to California. It occurs as far south as Argentina (Townes & Townes, 1966), and I have seen specimens from Tucuman and southern Brazil. Almost certainly the references to Enicospilus undulatus (a European species) as Neotropical (Taschenberg, 1875; Bréthes, 1909; Schrottky, 1913) are based on misidentifications of E. americanus. E. americanus is, however, extremely rare in tropical America and is generally replaced by a complex of very similar species that include E. lebophagus, E. mexicanus, E. cameronii and E. tenuigena. There are numerous early records of E. americanus attacking many hosts, but as it was commonly confused, not only with other species of Ophioninae, but also with Netelia species, all the early records are best disregarded unless authenticated material can be examined. The rearings I have seen suggest that E. americanus is a polyphagous parasitoid of large saturniids that construct thick-walled cocoons. This includes most species of Saturniinae in North America. I have seen authenticated specimens reared in the United States from Antheraea polyphemus (Cramer), Callosamia promethea (Drury), Callosamia securifera (Maassen), Hyalophora cecropia (L.), Hyalophora euryalis (Boisduval), Rothschildia orizaba (Westwood) and Samia cynthia (Drury). It is recorded additionally from Actias luna (L.) (Carlson, 1971), 168 IAN D. GAULD and in South America in is purported to parasitize Rothschildia maurus (Burmeister) (Blanchard, 1940), R. arethusa (Walker) (Costa Lima, 1962). Bourquin (1947) recorded it from the lasiocampid Tolype pauperata (Burmeister), but this host record is highly suspect. I have seen a series of the ‘hairy male morph’ of E. americanus reared from Automeris io (F.). Although this moth is a hemileucine, not a saturniine, it spins a more sturdy cocoon than do most other hemileucines (Ferguson, 1971). Peigler (1977) observed that the adults of E. americanus appear about the same time as the adults of their saturniid host. Oviposition is into a very young larva (Price, 1975) and the ophionine kills its host after the latter has formed a cocoon. Some individuals enter diapause as a prepupa and do not emerge for two years. For example, in CAS is a female that emerged in September 1925 from a cocoon of Antheraea polyphemus collected in late 1923. Peigler (1985) suggested that populations of E. americanus may move, following fluctuations in populations of Callosamia promethea and C. securifera. MATERIAL EXAMINED 76 &, 89 9, from the following: Argentina (Tucum4n); Bolivia (Cochabamba); Brazil (Nova Teutonia); Canada (Ontario, Quebec); Mexico (Chiapas, Chihuahua, Durango, Mexico State, Nuevo Leén, Tlax- cala, Veracruz); U.S.A. (California, Colorado, Delaware, Florida, Georgia, Maryland, Massachusetts, Michigan, Minnesota, New Hampshire, New Jersey, New York, North Carolina, Ohio, South Carolina, Texas, Virginia) (BMNH, CAS, CNC, FSCA, PANS, TC, USNM). Enicospilus tenuigena (Kriechbaumer) (Figs 166, 167, 168, 247) Eremotylus tenuigena Kriechbaumer, 1901c: 153. Holotype 2, BRAZIL (TMP) [examined]. Enicospilus tenuigena (Kriechbaumer) Townes & Townes, 1966: 183. DescripTIon. Mandibles moderately stout, fairly evenly narrowed, apically twisted 10—20° (Fig. 168); upper mandibular tooth compressed, sometimes a little upcurved, 1.5—1.7 times as long as the lower tooth; outer mandibular surface centrally flattish, proximally with a broad shallow concavity which bears close fine pubescence. Labrum 0.1-0.2 times as long as broad; malar space 0.2-0.3 times as long as basal mandibular width. Clypeus in profile flat, margin blunt; clypeus in front view 1.3—1.4 times as broad as long, margin truncate4 apically. Lower face 0.66—0.74 times as broad as long, centrally punctate to punctostriate. Head in dorsal view with genae rounded behind eye; posterior ocellus very close to eye; FI = 75-80% ; occipital carina mediodorsally complete, ventrally curved to approach but not join hypostomal carina about 0.9 times the basal mandibular width away from mandible. Antenna long and slender, with 58-66 flagellar segments; 20th segment 1.82.0 times as long as broad. Mesoscutum finely punctate, in profile evenly rounded; notauli distinct but shallow and short. Meso- pleuron polished, the upper part with obsolescent sculpture, the lower part striate; epicnemial carina with upper end curved towards anterior margin of pleuron, but then effaced. Scutellum in profile moderately to weakly convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.3—1.5 times as long as anteriorly broad, smooth with some wrinkling posteriorly. Metapleuron moderately convex, striate (Fig. 166); submetapleural carina weakly broadened anteriorly; posterior transverse carina of mesoster- num strong, complete, with a central projecting V-shaped part. Propodeum in profile evenly but steeply rounded; anterior transverse carina complete, posterior transverse carina indistinct; anterior area short, deeply impressed, striate; spiracular area short, more or less smooth, posterior area dorsally deplanate, concentrically striate; lateral longitudinal carina present anteriorly, not joined to spiracular margin by a short carina. Fore wing length 16-25 mm; discosubmarginal cell as in Fig. 247; AI = 0.95—1.06; CI = 0.45-0.67; ICI = 0.77-0.93; SDI = 1.33-1.69; cu-a proximal to the base of Rs&M by 0.1-0.3 times its own length; marginal cell proximally more or less evenly hirsute except for extreme proximal corner which is virtually glabrous; 1st subdiscal cell anteriorly sparsely hirsute. Hind wing with S—10 hamuli on R1; 1st abscissa of Rs more or less straight, 2nd abscissa weakly bowed. Fore leg with tibia slightly flattened, with scattered stout spines on outer surface. Mid leg with longer tibial spur 1.4~1.6 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.3-3.6 times as long as broad; claws of female long, strongly curved, with close, moderately long pectinae, those of male similar but slightly shorter and more strongly curved. Gaster slender; tergite 2 in profile 4.6—5.4 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical to obovate and separated from anterior margin of tergite by about 34 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing fine decumbent pubescence, a a ce ee OPHIONINAE OF TROPICAL MESOAMERICA 169 the last sternite very unusual in being far longer than the penultimate one (Fig. 167); gonosquama _ proximally narrowed, distally parallel-sided, apically truncate. Colour generally pale brownish yellow, with a median or a median and pair of lateral vittae on mesoscutum dark brown; gaster with tergites 3+ infuscate; interocellar area yellow; antenna yellowish brown, distally infuscate, centrally golden; pterostigma golden; wings hyaline. Variation. Enicospilus tenuigena is morphologically a rather uniform species that shows some variation in the colour of the mesoscutum as outlined above. Most specimens have the submetapleural carina only weakly broadened anteriorly, but in isolated larger specimens it can be more abruptly widened. REMARKS. The male of this species is very easily recognizable on account of the long subgenital plate and the characteristic gonosquamae. The female resembles the male in colour pattern, and the combination of this, the striate metapleuron and the rather marked ventral protuberance near the base of Rs+2r in the fore wing should enable the female to be recognized. However, it is sometimes difficult to separate from females of some other species in the E. americanus complex. BIOLOGICAL INFORMATION. Enicospilus tenuigena is a widely distributed species whose range extends from central Costa Rica southwards to Ecuador and Brazil. In Central America it is only known from wet forest areas from sea-level up to about 700 m. The scattered collecting dates suggest it may occur as an adult throughout the year. MATERIAL EXAMINED Holotype 9, Brazil: Santos (TMP). Colombia: 1 2 , Valle, Lago Calima, in tropical wet forest (FSCA). Costa Rica: Alajuela Prov.: 1 2, Finca San Gabriel, 3 km W. Dos Rios, 850 m, vi.1986 (Gauld) (BMNH): Cartago Prov.: 1 2, Turrialba, 700 m, vii. 1965 (Real) (CAS): Lim6n Prov.: 1 oO’, Cerro Tortuguero, N. edge of Tortuguero National Park, 0-100 m, v.1984 (Janzen & Hallwachs) (BMNH): San José Prov.: 3 0’, 1 9, Estacion Carrillo, Braulio Carrillo National Park, 700 m, i, ii, v.1985 (Chacon) (BMNH). Ecuador: 1 Q, Pichincha, nr Nanegal, 1200 m, ix.1977 (Peria) (BMNH). Panama: 1 9, Barro Colorado Island, iv.1978 (Wolda) (RNH); 1 ©’, same locality, ii.1983 (Wolda) (TC); 10,2 2, same locality, 120m, xii.1983, vii. 1984, ix.1985 (Wolda) (BMNH) Enicospilus venezuelanus (Szépligeti) Allocamptus venezuelanus Szépligeti, 1906: 149. Lectotype &', VENEZUELA (TM), designated by Townes & Townes (1966: 184) [examined]. | Enicospilus venezuelanus (Szépligeti) Townes & Townes, 1966: 184. Description. Mandibles of moderate length, proximally evenly narrowed, distally almost parallel-sided, apically twisted 10—20°; upper mandibular tooth slightly compressed, 1.4 times as long as the lower tooth; outer mandibular surface rather flat, with fine inconspicuous pubescence, and with a shallow and indistinct proximal concavity. Labrum 0.2 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile slightly out-flared, margin centrally blunt; clypeus in front view 1.4 times as broad as long, with margin apically truncate. Lower face 0.76 times as broad as long, centrally finely and sparsely punctate. Head in dorsal view with genae constricted behind the eyes; posterior ocellus contiguous with the eye; Fl = 75%; occipital carina mediodorsally narrowly interrupted, ventrally strong, curved to join hypostomal carina about 0.7 times the basal mandibular width away from mandible. Antenna long and slender, with 68 flagellar segments; 20th segment 2.0 times as long as broad. Mesoscutum weakly polished and finely puncto-granulate, in profile abruptly rounded; notauli vestigial. Mesopleuron weakly polished, the upper part punctate, the lower part more closely punctate with the punctures tending to coalesce, becoming punctostriate; epicnemial carina abruptly curved towards ante- rior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for 0.9 of its length; scutellum in dorsal view 1.6 times as long as anteriorly broad, punctate. Metapleuron evenly convex, coarsely punctate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina quite weak with lateromedian and lateral parts hardly represented, posterior one absent; anterior area short, striate; spiracular area granulate; posterior area coarsely transversely concentrically rugose-striate; lateral long- itudinal carina present anteriorly, not joined to spiracular margin by a short carina. Fore wing length 24 mm; discosubmarginal cell similar to that shown in Fig. 247; AI = 0.96; CI = 0.46; ICI = 0.62; SDI = 1.31; cu-a proximal to the base of Rs&M by about 0.3 times its own length; marginal cell proximally narrowly glabrous adjacent to Rs+2r; 1st subdiscal cell with anterior and distal parts broadly hirsute. Hind wing with 10 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa straight. 170 IAN D. GAULD Fore leg with tibia slightly flattened, with isolated fine spines on outer surface. Mid leg with longer tibial spur 1.4 times length of the shorter. Hind leg with coxa in profile 1.9 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.5 times as long as broad; claws of male of . moderate length, abruptly rounded with a long stout apical point. Gaster long and slender; tergite 2 in profile 5.9 times as long as posteriorly deep, laterotergite folded Ft under, thyridia elliptical and separated from anterior margin of tergite by about 3 times its own length. 1 Female unknown. Male with sternites 7-9 bearing long stout close erect pubescence; gonosquama long, distally rounded. Colour generally pale yellowish brown with segments 3-5 of gaster darker (or possibly discoloured due {ii to age); interocellar area yellowish; antenna orange; pterostigma yellowish; wings more or less hyaline. RemaRKS. Enicospilus venezuelanus is not known to occur in the study region delimited, but as it occursin Venezuela and is structurally very similar to many Central American species, it has been included here for the sake of comparison. This species belongs to the E. americanus species-complex, and most closely | resembles E. peigleri. Both have similar shaped mandibles, similarly sculptured alitrunks and rather _ similar venation. However, EF. venezuelanus is much larger, has the genal and hypostomal carinae meeting, _ and quite a different arrangement of pubescence on the sternites. | BIOLOGICAL INFORMATION. Despite the fact that I have examined a considerable number of South American | specimens of the E. americanus species-complex, I have seen only a single example of E. venezuelanus, the — lectotype from Venezuela. Nothing is known about its biology. MATERIAL EXAMINED Lectotype 0’, Venezuela: Merida (TM). Enicospilus peigleri Gauld (Fig. 248) Enicospilus peigleri Gauld, 1988: 46. Holotype 2 U.S.A.: Florida (FSCA) [examined]. Description. Mandibles long and stout, proximally abruptly narrowed, distally parallel-sided, apically twisted 10—20°; upper mandibular tooth subcylindrical or slightly compressed, 1.4~1.6 times as long as the lower tooth; outer mandibular surface finely hirsute, centrally rather flat and with a weak proximal concavity. Labrum 0.3 times as long as broad; malar space 0.2-0.3 times as long as basal mandibular width. Clypeus in profile almost flat, margin blunt; clypeus in front view 1.4—1.5 times as broad as long, with margin apically weakly convex. Lower face 0.71—-0.80 times as broad as long, centrally finely punctate. Head in dorsal view with genae very slightly inflated; posterior ocellus very close to eye; FI = 60-65%; | occipital carina mediodorsally complete, or weak or interrupted, ventrally evanescent, not curved to join hypostomal carina. Antenna very long and slender, with 62-65 flagellar segments; 20th segment 1.9-2.2 times as long as broad. | Mesoscutum polished, obsoletely punctate, in profile evenly rounded; notauli vestigial. Mesopleuron polished, the upper part punctate, ventrally grading to punctostriate; epicnemial carina with upper end curved to approach anterior margin of pleuron, its lower corner produced backwards, acute. Scutellum in profile weakly convex, laterally strongly carinate for all of its length; scutellum in dorsal view 1.4—1.5 times . as long as anteriorly broad, rather smooth. Metapleuron polished, convex, punctate to punctostriate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile abruptly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area short, with scattered striae; spiracular area very short, more or less smooth; posterior area dorsally deplanate, concentrically rugose-striate; lateral longitudinal carina present at least ante- riorly, not joined to spiracular margin by a short carina. Fore wing length 15-18 mm; discosubmarginal cell as in Fig. 248; AI = 0.76-1.28; CI = 0.34-0.62; ICI = 0.59-0.72; SDI = 1.20-1.28; cu-a subopposite to the base of Rs&M; marginal cell proximally evenly hirsute except for a small glabrous area in extreme proximal corner; 1st subdiscal cell hirsute anteriorly and posterodistally. Hind wing with 6-8 hamuli on R1; 1st abscissa of Rs almost straight, 2nd abscissa bowed. Fore leg with tibia slightly flattened, with numerous slender spines on outer surface. Mid leg with longer tibial spur 1.2-1.3 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.42.6 times as long as broad; claws of female very long and slender, apically abruptly turned, but apical tooth short and the pectinae short and scattered; claws of male similar but with pectinae finer and closer. Gaster slender; tergite 2 in profile 4.2—-5.3 times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by about 2-3 times its own length. 1 17 OPHIONINAE OF TROPICAL MESOAMERICA ‘pajoayjoo usaq sey iapsiad snjidsooiuq yoy ye sayyesoT «Zp dey We IAN D. GAULD Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing fine scattered semierect hairs; gonosquama rounded apically. Colour generally yellowish brown, head and scutellum more yellow, extreme posterior end of gaster slightly infuscate; interocellar area yellow; antenna orange; pterostigma brownish yellow; wings hyaline. VARIATION. Some of the Mexican specimens have the propodeal sculpture weaker and finer than the majority of individuals. REMARKS. Enicospilus peigleri is likely to have been confused with E. americanus which it resembles in general appearance. It differs in having more slender antennae, unusually robust mandibles and, in most cases, in having a small protuberance basally on the posterior margin of Rs+2r. The long slender claws of this species are quite unlike those of any other sympatric Enicospilus. BIOLOGICAL INFORMATION. Enicospilus peigleri is a fairly widespread species whose range extends from about 35°N in eastern North America, south to about 17°N in southern Mexico (Map 12). It has not been reared, but several of the specimens have the mandibular teeth almost completely worn away, suggesting this insect emerges from a cocoon in hard earth. Many of the specimens examined were collected in the hottest, driest part of the summer. MATERIAL EXAMINED Holotype 9, U.S.A.: Florida: Alachua Co., Gainesville, viii.1970 (Mead) (FSCA) Paratypes. Mexico: Chiapas: 1 ©’, San Cristobal de las Casas, vii.1969 (Kritsch) (CNC): Durango: 1 9, 11 km W. Durango, 2200 m, viii.1964 (Mason) (CNC): Mexico Ste: 1 0’, Ixtapan la Sal, 2200 m, viii.1962 (Evans) (MCZ): San Luis Potosi: 1 2, El Salto Falls, vi.1963 (Woodruff) (FSCA). U.S.A.: Florida: 2 9, Gainesville, xi.1971, x.1972 (Mead) (FSCA): South Carolina: 1 9, Greenville, viii.1982 (Peigler) (BMNH). Enicospilus neotropicus Hooker (Fig. 249) Enicospilus neotropicus Hooker, 1912: 69. Lectotype 2, DOMINICAN REPUBLIC (USNM), desig- nated by Townes & Townes (1966: 182) [examined]. Description. Mandibles long, proximally narrowed, distally parallel-sided, apically twisted 10—20°; upper mandibular tooth cylindrical, about 2.0 times as long as the lower tooth; outer mandibular surface with an impressed hirsute groove running from near upper proximal corner to base of teeth; proximal concavity strong. Labrum 0.4—0.5 times as long as broad; malar space 0.4-0.5 times as long as basal mandibular width. Clypeus in profile convex, margin impressed, acute; clypeus in front view 1.5—1.7 times as broad as long, margin weakly convex apically. Lower face 0.69-0.82 times as broad as long, centrally coarsely punctate. Head in dorsal view with genae rounded; posterior ocellus close to eye; FI = 57-65%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 1.0 times the basal mandibular width away from mandible. Antenna long and slender, with 63-66 flagellar segments; 20th segment 1.9—2.0 times as long as broad. Mesoscutum polished, closely punctate, in profile evenly rounded; notauli short, but quite deeply impressed. Mesopleuron polished, the upper part punctate to punctostriate, the lower part more coarsely sculptured, punctogranulate to punctostriate; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for all of its length; scutellum in dorsal view 1.8-2.0 times as long as anteriorly broad, shallowly, closely punctate. Metapleuron weakly convex, coarsely and closely punctate to rugose; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly rounded; anterior transverse carina complete, posterior transverse carina absent; anterior area quite long, irregularly rugose/striate; spiracular area quite long, punctate; posterior area reticulate/wrinkled; lateral longitudinal carina from present anteriorly to complete, usually not joined to spiracular margin by a short carina. Fore wing length 13-14 mm; discosubmarginal cell as in Fig. 249; AI = 0.42-0.76; CI = 0.27-0.54; ICI =0.36-0.45; SDI = 1.18-1.23; cu-a from more or less opposite to, to proximal to base of Rs&M; marginal cell sparsely, but extensively hirsute; 1st subdiscal cell with anterior and distal parts hirsute. Hind wing with 7-8 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa almost straight. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.5-1.6 times length of the shorter. Hind leg with coxa in profile 1.7-1.9 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.9-3.1 times as long as broad; claws of female long, weakly curved and with quite close pectinae, those of male similar but with pectinae finer and very much more closer together. | | OPHIONINAE OF TROPICAL MESOAMERICA 173 Gaster long and slender; tergite 2 in profile 5 or more times as long as posteriorly deep, laterotergite upturned, thyridia elliptical and separated from anterior margin of tergite by about 3 times its own length. Ovipositor slender, its sheath narrow. Males with sternites 7-9 bearing fine semidecumbent pubescence; gonosquama obliquely truncate. Colour generally orange-brown with most of head paler yellowish; interocellar area yellowish; antenna _ orange; pterostigma orange-brown; wings hyaline. VARIATION. A male from the Dominican Republic has the head, scutellum and sides of the alitrunk whitish. | Remarks. Hooker (1912) claimed to have specimens of this species from Chile, but I have been unable to locate these and I have seen no specimens from outside the general area of the Caribbean and Florida. However, I have seen several specimens of a similar, but undescribed species from Chile, which has much broader genae, and another, similar undescribed species occurs in Bolivia. The missing Hooker paralec- totype(s) may be one of these species. The possession of the mandibular groove and closely pectinate male claws indicate that Enicospilus neotropicus belongs to the E. ramidulus species-group. It differs from any other described New World species in this group in lacking a central sclerite, in having a larger fenestra and in having a more strongly sinuous Rs+2r. BIOLOGICAL INFORMATION. Enicospilus neotropicus is only known to occur in the Caribbean and in southern Florida (Map 13). Its host is unknown. MATERIAL EXAMINED Lectotype 2 , Dominican Republic: San Francisco, ix.1905 (Busck) (USNM). Paralectotypes, 2 0’, same data as lectotype (USNM). Dominican Republic: 1 o', La Cumbre, Santiago, 1000 m, iv.1978 (Woodruff) (FSCA); 1 2, Los Hidalgros, vi.1969 (Flint & Gémez) (USNM). Jamaica: 1 ©’, Kingston, iii.1900 (Taylor) (USNM); 2 oc, Liguanen Plain, xi-xii.1911 (Brues) (MCZ); 2 2, Mandeville, xi.1970 (Frank) (BMNH). U.S.A.: Florida: 10, S. Miami, ix (Graenicher) (MCZ). Enicospilus doylei sp. n. (Figs 169, 250) Description. Mandibles very long, proximally strongly narrowed, distally parallel-sided, apically twisted 10—20°; upper mandibular tooth compressed, more than 3.0 times as long as the rather small lower tooth; outer mandibular surface with a deep hirsute diagonal groove extending from upper proximal corner to near base of teeth. Labrum 0.40.5 times as long as broad; malar space 0.3-0.4 times as long as basal mandibular width. Clypeus in profile weakly convex, margin impressed, acute; clypeus in front view 1.9- 2.0 times as broad as long, the margin apically truncate. Lower face 0.81-0.88 times as broad as long, polished, centrally punctate to punctostriate. Head in dorsal view with genae rounded; posterior ocellus very close to eye; FI = 58-68%; occipital carina mediodorsally complete, ventrally curved to join hyposto- mal carina about 0.8 times the basal mandibular width away from mandible. Antenna moderately long, with 51-53 flagellar segments; 20th segment 1.7—2.1 times as long as broad. Mesoscutum polished, very sparsely and shallowly punctate, in profile evenly rounded; notauli weak but discernible. Mesopleuron polished, the upper part from punctate to striate, the lower part coarsely punctostriate; epicnemial carina inclined towards anterior margin of pleuron, but with upper end evanes- cent. Scutellum in profile weakly convex, laterally carinate for 0.9 or more of its length; scutellum in dorsal view 1.7—1.9 times as long as anteriorly broad, finely punctate. Metapleuron characteristically flattened, coarsely irregularly punctostriate (Fig. 169); submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina medially strong, arched slightly forwards, laterally weak or evanescent; posterior transverse carina absent; anterior area moderately long, striate; spiracular area quite short, especially centrally, punctate; posterior area coarsely irregularly wrinkled; lateral longitudinal carina complete, not joined to spiracular margin by a short carina. Fore wing length 12-14 mm; discosubmarginal cell as in Fig. 250; AI = 0.58-0.91; CI = 0.40-0.51; ICI = 0.50—0.58; SDI = 1.10-1.15; cu-a opposite to the base of Rs&M or proximal to it by about its own thickness; marginal cell proximally slightly more sparsely hirsute than centrally; 1st subdiscal cell with anterior and distal parts hirsute. Hind wing with 5-6 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa straight. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.6-1.7 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; ‘payoaj[oo useq sey snoidowoau snjidsoougq YoY ye soiyyeooT ET dey 34 IAN D. GAULD 174 OPHIONINAE OF TROPICAL MESOAMERICA 7S) trochantellus dorsally 0.1—0.2 times as long as broad; 4th segment of tarsus 2.2—2.4 times as long as broad; claws of female large, distally quite strongly curved, with close short pectinae; claws of male similar, but with pectinae finer and closer together. Gaster long and slender; tergite 2 in profile more than 4 times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by 2 or more times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7—9 bearing fine decumbent pubescence; gonosquama obliquely truncate. Colour generally yellowish brown; interocellar area yellow; antenna orange-brown; pterostigma yellow- ish; wings hyaline. VARIATION. A few specimens have pale longitudinal stripes on the mesoscutum. REMARKS. This species is named in honour of Doyle McKey for his enthusiasm for unravelling tropical animal-plant interactions. The form of the mandibles and male claws indicate that Enicospilus doylei belongs to the E. ramidulus species-group. It differs most obviously from all other New World members of this group in the form of the central sclerite, which is almost kite-shaped rather than oval or D-shaped, and in having the metapleuron flattened. These are not the only autapomorphic features of this species; it has longer and more bowed mandibles than other taxa, the head is subtly shorter, and the face slightly constricted ventrally. BIOLOGICAL INFORMATION. Enicospilus doylei is widely distributed throughout Florida where it occurs from the Keys to almost as far north as the Georgia State line. I have also seen a single specimen from Baton Rouge, Louisiana, so it may occur all along the northern periphery of the Gulf of Mexico. Outside of the United States E. doylei has also been collected in the Bahamas, and on Bermuda. On the mainland of Central America this species is only known to occur in agricultural areas in Nicaragua (Map 14). This peculiar distribution pattern suggests E. doylei may be associated with a synanthropic lepidopteran host, and it may have been spread by the agency of man. However, it has never been reared. MATERIAL EXAMINED Holotype ?, U.S.A.: Florida, Gadsden Co., Quincy, viii.1970 (Hasse) (FSCA). Paratypes. Bahamas: 1 ?, Grand Bahama Is., 8 Mile Rock, iv (French) (MCZ). Bermuda: 1 9, Bermuda Is., vi.1962 (Maynard) (TC). Nicaragua: 2 0’, 3 9, Leon, vii.1985 (Mays) (BMNH; ULN). U.S.A.: Florida: 1 9, Alachua Co., Gainesville, iv.1952 (Walley) (CNC); 1 9, same locality, vii.1953 (Patton) (FSCA); 1 O’, same locality, x.1963 (Platt) (FSCA); 1 0’, same locality, vii. 1969 (Mead) (FSCA); 29, same locality, v.1970 (Mead) (FSCA); 1 9, same locality, x.1971 (Mead) (FSCA); 1 0’, same locality, i.1972 (Mead) (FSCA); 1 O’, same locality, vi.1972 (MacGowan) (FSCA); 1 9, same locality, viii.1972 (Mead) (FSCA); 3 2, Dade Co., Miami, vi.1960 (Briggs) (FSCA); 1 0’, 4 9, Gadsden Co., Quincy, vii- viii.1970 (Hasse) (BMNH, FSCA); 3 co’, 2 9, same locality, viii.1971 (Reid) (FSCA); 9 0’, 9 2, Highlands Co., Archbold Biological Station, i-iv.1962 (Frost) (TC); 3 2, same locality, iv.1967 (Peterson) (CNC); 30°, 4 9, Highlands Co., Lake Placid, v-vi.1967 (Heinrich) (CNC); 2 ©’, Hillsborough Co., St Petersburg, vi.1962 (Forsyth) (FSCA); 1 9, Lake Co., Leesburg, x.1961 (Felshaw) (FSCA); 1 2, Manatee Co., Bradenton, xii.1963 (Frederic) (FSCA); 1 0’, Monroe Co., Key Largo, v.1973 (Wyles) (FSCA); 1 9, Monroe Co., Key West, vi.1960 (Warner) (FSCA); 1 oO’, same locality, iii.1962 (Buchanan) (FSCA); 1 °, same locality and collector v.1963 (FSCA); 1 2, Polk Co., Winter Haven, viii.1960 (Hayward) (FSCA): Louisiana: 1 9, Baton Rouge, xi.1961 (Arnold) (TC). Enicospilus purgatus (Say) (Figs 170, 171, 251) Ophion purgatus Say, 1836: 239. Syntypes, U.S.A. (destroyed). Ophion lateralis Brullé, 1846: 141. Holotype 2, U.S.A. (MNHN) [examined]. Synonymized by Hooker, 1912: 79. Ophion flaviceps Brullé, 1846: 142. Holotype 2, BRAZIL (MNHN) [examined]. Synonymized by Townes & Townes, 1966: 181. Ophion volubilis Holmgren, 1868: 410. Lectotype co’, ARGENTINA (NR) designated by Townes & Townes (1966: 181). Synonymized by Townes & Townes, 1966: 181. [Ophion merdarius Gravenhorst; Taschenberg, 1875: 435. Misidentification. | Henicospilus flaviceps (Brullé) Szépligeti, 1905: 27. Henicospilus merdarius var. volubilis (Holmgren) Roman, 1910: 165. Enicospilus purgatus (Say) Hooker, 1912: 79. Enicospilus flaviceps (Brullé) Hooker, 1912: 85. IAN D. GAULD 176 "pajoa][oo useq sey 1ajAop snjidsooiugq yoiym ye saytyeooy «py dey OPHIONINAE OF TROPICAL MESOAMERICA 177 Henicospilus purgatus (Say) Morley, 1912: 33. Henicospilus volubilis (Holmgren) Morley, 1912: 34. Enicospilus purgatus (Say); Townes, 1945: 473. [Henicospilus merdarius (Gravenhorst); Schrottky, 1913: 130. Misidentification. ] [Enicospilus merdarius (Gravenhorst); Townes & Townes, 1966: 181. Misidentification.] [Enicospilus merdarius (Gravenhorst); Carlson, 1979: 703. Misidentification. | Description. Mandibles with a weakly developed proximoventral lobe, proximally abruptly narrowed, distally long, parallel-sided, apically twisted 10—20°; upper mandibular tooth cylindrical, 2.0-3.0 times as long as the lower tooth; outer mandibular surface with a diagonal furrow extending from upper proximal corner to between bases of teeth, this furrow bearing close pubescence. (Fig. 171). Labrum 0.3-0.4 times as long as broad; malar space 0.4—0.5 times as long as basal mandibular width. Clypeus in profile moderately strongly convex, margin impressed, acute; clypeus in front view 1.3—1.7 times as broad as long, the apical margin truncate. Lower face 0.75—0.85 times as broad as long, polished, finely punctate. Head in dorsal view with genae fairly short, slightly swollen; posterior ocellus close to but generally not touching eye; FI = 60-70% ; occipital carina mediodorsally complete, ventrally usually complete, joining hypostomal carina about 0.8 times the basal mandibular width away from mandible. Antenna moderately long and slender, with 53-57 flagellar segments; 20th segment 2.1-2.5 times as long as broad. Mesoscutum weakly to moderately strongly polished, with fine punctures, in profile evenly rounded; notauli absent. Mesopleuron polished, the upper part finely punctate to punctostriate, the lower part more coarsely sculptured, either punctate or punctostriate; epicnemial carina curved towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.5—1.7 times as long as anteriorly broad, punctate, often posteriorly wrinkled. Metapleuron weakly convex, punctate (Fig. 170); submetapleural carina evenly anteriorly broadened; posterior trans- verse carina of mesosternum complete. Propodeum in profile fairly evenly rounded; anterior transverse carina generally complete but with lateral extremities evanescent, posterior transverse carina absent; anterior area moderately long, strongly impressed, striate to rugose/coriaceous; spiracular area of moder- ate length, generally fairly smooth; posterior area finely reticulately wrinkled; lateral longitudinal carina present anteriorly, posteriorly either complete or effaced, usually joined to spiracular margin by a short carina, but this carina may be weak or absent. Fore wing length 12-14 mm; discosubmarginal cell as in Fig. 251; AI = 0.33-0.99; CI = 0.32-0.48; ICI = 0.36-0.51; SDI = 1.19-1.31; cu-a proximal to the base of Rs&M by 0. 1-0.4 times its own length, rarely Virtually opposite; marginal cell proximally fairly evenly hirsute; lst subdiscal cell with anterior 0.3 or more hirsute. Hind wing with 6-7 hamuli on R1; Ist abscissa of Rs straight, 2nd abscissa straight. Fore leg with tibia subcylindrical, with scattered spines on outer surface. Mid leg with longer tibial spur 1.4-1.6times length of the shorter. Hind leg with coxa in profile 1.8—2.0 times as long as deep; trochantellus dorsally 0.1—0.2 times as long as broad; 4th segment of tarsus 2.7—2.9 times as long as broad; claws of female long, distally evenly curved, fairly closely pectinate; claws of male similar, but with pectinae extremely close together. Gaster long and slender; tergite 2 in profile more than 6 times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by about 2.5—3.5 times its own length. Ovipositor apically slender, its sheath slender. Male with sternites 7-9 bearing fine semierect pubescence; gonosquama quite long and strongly tapered, distally obliquely truncate and dorsally with margin slightly excavate. Colour generally yellowish brown with terminal segments of gaster weakly infuscate; interocellar area, vertex, genal orbit and most of face paler yellowish; pterostigma orange; wings hyaline. VARIATION. Despite the fact that Enicospilus purgatus has an extensive geographical range it is a remarka- bly morphological uniform species. The most striking variation is in density of the punctures on the metapleuron. In most specimens these punctures are very close, but isolated individuals from many areas may have them small and scattered. There is comparatively little variation in the form of the alar sclerites though a few individuals may have the distal sclerite either more or less heavily sclerotized than the figure. The central sclerite varies in shape from circular to D-shaped though occasional specimens have the distal side angularly rounded. There are in the BMNH a few females from various localities (including Costa Rica) which seem to belong to this species except that they are rather larger with a fore wing length of up to 18 mm and possess up to 69 flagellar segments. In other features these individuals are typical of the species. Amongst material in North American institutions I have seen a number of specimens which are Structurally unusual for this species. I have not investigated these further, and it is possible that several Nearctic species may be confused under the name purgatus. These do not occur in the Neotropics. 178 IAN D. GAULD REMARKS. Enicospilus purgatus has been considered to be synonymous with the Palaearctic species E. merdarius (Gravenhorst) and treated under that name in several catalogues (e.g. Townes & Townes, 1966; Carlson, 1979). However, it is noteworthy that the name merdarius is actually a junior synonym of E. ramidulus (L.) (see Townes et al., 1965), and thus, if one were to treat the New World species as being conspecific with the Old World one, the name ramidulus should be used However, I am not certain that the New and Old World species are indeed conspecific, as they differ subtly, but consistently, in the shape of the alar sclerites and in the density of the thoracic punctures. Structurally they are otherwise very similar and both seem to parasitize similar hosts, but then many other members of the ramidulus species-group (including the Old World tropical species E. capensis) are morphologically similar and parasitize noctuids in agroecosystems. It is possible to reliably separate ‘typical’ members of each of the nominal species E. purgatus, E. ramidulus and E. capensis, but the ranges of variation do somewhat overlap, making definition of the species difficult. At present I believe that the best course of action is to treat the New World species (E. purgatus) as distinct from the temperate (E. ramidulus) and tropical (E. capensis) Old World ones. E. purgatus is the commonest New World species of the E. ramidulus (= capensis) species-group, a complex of species characterized by possessing a long, diagonally grooved mandible and strongly sexually dimorphic claws (those of the male have extremely fine, close pectination). E. purgatus may easily be separated from other Neotropical species by the characters given in the key E. purgatus is obviously very closely related to several Old World species (including E. ramidulus and E. capensis), and it seems reasonable to regard the position and number of alar sclerites, and the general sculpture of this complex of species as representing the plesiomorphic condition for the species-group, because they share these features with their sister-group, the E. antefurcalis species-group (Gauld, 1982). Assuming that E. purgatus is the most plesiomorphic of the New World species of the ramidulus species- group, all the others can be regarded as having arisen from it. E. neotropicus is distinguished by having lost the central sclerite, whilst E. doylei has acquired a specialized flattened metapleuron and a modified central sclerite. The endemic Galapagos E. vidor species-complex is characterized by the possession of fine thoracic sculpture and by the reduction of the alar sclerites (Gauld & Carter, 1983). Various other, undescribed, South American species can similarly be assumed to have been derived from E. purgatus, leaving E. purgatus, as it is here recognized, as a ‘paraphyletic species’. BIOLOGICAL INFORMATION. Enicospilus purgatus is a very widely distributed species whose range extends from northern Canada throughout virtually the entire United States, the Caribbean, and Central America, south to Argentina. However, it seems to be uncommon in many tropical areas of South America, and it is apparently absent from Chile, where it seems to be replaced by a related but undescribed species which has a stouter head. Throughout most of its range E. purgatus seems to be most common in disturbed habitats and agricultural areas. E. purgatus is a relatively infrequently collected species in much of Costa Rica. I have seen isolated specimens from Tortuguero National Park in Limon Province and Monteverde Reserve in Puntarenas Province, but most specimens have been collected in Santa Rosa National Park in Guanacaste Province, or on the periphery of the park in disturbed areas, notably on the Cerro el Hacha, and at Estacion Mengo. At these sites most individuals have been collected between May and July during the earlier part of the wet season. Further north, in the agricultural lowlands of Nicaragua, E. purgatus is one of the most commonly collected species in the genus. It might be widespread and common in lowland agricultural areas throughout Central America; it is just that few people have ever light-trapped in such bleak areas! No specimens of E. purgatus have been collected in lowland forest sites such as Barro Colorado Island. E. purgatus is known to parasitize a variety of noctuid larvae that feed on rather low-growing vegetation. In the Neotropical region it is recorded as a parasitoid of the following species of Noctuidae: Alabama argillacea (Hubner), Faronta albilinea (Hubner), Helicoverpa zea (Boddie), Mythimna unipuncta (Haworth), Peridromia saucia (Hubner), Spodoptera frugiperda (Smith & Abbott) and Spodoptera ornithogalli (Guenée) (Gowdey, 1926; Blanchard, 1940; Bruner et al., 1945; Costa Lima, 1962). Townes (1945) and Carlson (1979) list North American host records, but several of these must be regarded with suspicion as this species has previouly been confused with other Enicospilus. I have seen no authenicated examples of E. purgatus reared from the larvae of Notodontidae, Sphingidae or Saturniidae. Oviposition seems to be into half-grown larvae, and the parasitoid larva emerges after the host larva has ceased to feed. The parasitoid cocoon is usually found in the soil or in leaf-litter. MATERIAL EXAMINED Holotype 9 (Ophion flaviceps Brullé), Brazil (MNHN). 286 O', 339 9, from the following localities- Anguilla; Antigua; Argentina (Buenos Aires, Chaco, Tucumén); Barbados; Bermuda; Bolivia (Santa Cruz); Brazil (Parana, Santa Catarina); Canada (most OPHIONINAE OF TROPICAL MESOAMERICA 179 provinces); Cayman Islands (Grand Cayman); Colombia (Valle); Costa Rica (Guanacaste, Limon and Puntarenas Provinces); Cuba; Dominica; Dominican Republic (Altagracia); Ecuador (Esmeraldas, Pichincha); Guyana; Jamaica; Mexico (Baja California, Chiapas, Chihuahua, Durango, Sinaloa, Sonora, Veracruz); Nicaragua (Le6n); Panama (Canal Zone); Puerto Rico; Peru (La Libertad, Lima, Loreto); Saint Vincent; Surinam; Trinidad; U.S.A. (almost all states); Venezuela (Miranda, Sucre); Virgin Islands. Enicospilus luquillo sp. n. (Fig. 252) Description. Mandibles moderately long, proximally moderately narrowed, distally almost parallel-sided, apically twisted 10—20°; upper mandibular tooth slender, subcylindrical, 1.3—1.6 times as long as the lower tooth; outer mandibular surface centrally very slightly concave, finely pubescent, with a very weak proximal concavity. Labrum 0.3 times as long as broad; malar space 0.4 times as long as basal mandibular width. Clypeus in profile weakly convex, margin inturned, blunt; clypeus in front view 1.3-1.6 times as broad as long, with margin weakly convex. Lower face 0.75—0.78 times as broad as long, centrally finely punctate. Head in dorsal view with genae evenly constricted behind eyes; posterior ocellus close to eye; FI = 65-70%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 1.0 times the basal mandibular width away from mandible. Antenna long and slender, with 65—68 flagellar _ segments; 20th segment 2.1—2.3 times as long as broad. Mesoscutum polished, sparsely punctate, in profile evenly rounded; notauli vestigial. Mesopleuron polished, the upper and lower parts alutaceous; epicnemial carina evenly curved towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.6—1.7 times as long as anteriorly broad, sparsely and shallowly punctate. Metapleuron weakly convex, virtually smooth; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete or rarely with a very narrow median discontinuity. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area striate; spiracular area moderately long, finely punctate; posterior area irregularly wrinkled, centrally with the wrinkles tending to be longitudinal; lateral longitudinal carina usually complete, joined to spiracular margin by a short carina. Fore wing length 17-18 mm; discosubmarginal cell as in Fig. 252; AI = 0.47-0.61; CI = 0.34-0.36; ICI = 0.52-0.58; SDI = 0.98-1.02; cu-a proximal to base of Rs&M by about 0.2 times its own length; | marginal cell proximally evenly hirsute; 1st subdiscal cell anteriorly rather sparsely but distally broadly hirsute. Hind wing with 7-8 hamuli on R1; 1st and 2nd abscissae of Rs straight. _ Fore leg with tibia slightly flattened, with scattered fine spines on outer surface. Mid leg with longer tibial | spur 1.2-1.5 times as long as the shorter. Hind leg with coxa in profile 1.8-1.9 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of hind tarsus 2.6—2.7 times as long as broad; claws of female with short, stout pectinae, those of male similar. Gaster slender; tergite 2 in profile more than 6 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical, separated from anterior margin of tergite by 34 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing dense long fine semierect pubescence; gonos- quama distally acutely rounded. ' Colour generally yellowish, sometimes with mesoscutum darker, rarely with mesoscutum with three dark longitudinal vittae; interocellar area yellow; antenna yellowish; pterostigma golden; wings hyaline. VaRIATION. The only unusual variation exhibited by the specimens at hand is in the colour of the mesoscutum as detailed above. Remarks. This species is named after the Luquillo National Forest, a unique and endangered habitat. Enicospilus luquillo is immediately recognizable by the highly modified, apomorphic alar sclerites (Fig. 252). It otherwise closely resembles E. flavoscutellatus and may have arisen from this species by becoming geographically isolated on Puerto Rico, the only place it is known to occur. BioLocicaL INFoRMATION. Enicospilus luquillo is only known to occur in forests on the island of Puerto Rico. No details of its biology are known. | MateriaAL EXAMINED Holotype 9, Puerto Rico: Luquillo Forest, El Yunque Biology Station, 700 m, i.1963 (Spangler & | Spangler) (USNM). Paratypes. Puerto Rico: 1 9 , Barranquitas, vi.1969 (Flint) (USNM); 1 2, Catano, x.1943 (USNM); 1 9, Gurabo, xi.1914 (USNM); 2 9, Luquillo Forest, El Yunque Biology Station, 700 m, i.1963 (Spangler & Spangler) (BMNH, USNM); 1 G’, same locality, vii.1969 (Howden) (CNC); 1 9, Santa Rita, xii.1913 (Merrill) (FSCA). 180 IAN D. GAULD Enicospilus martae sp. n. (Fig. 253) Description. Mandibles moderately long, fairly evenly narrowed, apically twisted 30—-40°; upper mandibu- lar tooth slightly compressed, 0.7—0.8 times as long as the lower tooth; outer mandibular surface centrally flat with isolated fine hairs, and with a weak proximal concavity. Labrum 0.2 times as long as broad; malar space 0.4 times as long as basal mandibular width. Clypeus in profile weakly convex, margin rather sharp; clypeus in anterior aspect 1.5—1.6 times as broad as long, with margin weakly convex. Lower face 0.70-0.75 times as broad as long, centrally closely punctate, fairly weakly polished. Head in dorsal view with genae rounded; posterior ocellus contiguous with eye; FI = 60-65%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.8-0.9 times the basal mandibular width away from mandible. Antenna slender with 64-65 flagellar segments; 20th segment 2.3—2.4 times as long as broad. Mesoscutum polished, finely punctate, in profile steeply rounded; notauli weak but discernible. Meso- pleuron polished, the upper part with obsolescent punctures, the lower part similar but with area between punctures finely alutaceous; epicnemial carina inclined towards but not reaching anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for most of its length, but posteriorly the carinae are rather weak; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, polished with fine scattered punctures. Metapleuron alutaceous, with close weak punctures; submetapleural carina weakly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area quite deeply impressed, irregularly wrinkled; spiracular area of moderate length, smooth; posterior area coriaceous; lateral longitudinal carina complete, joined to spiracular margin by a short carina. Fore wing length 15-17 mm; discosubmarginal cell as in Fig. 253; AI = 0.60-0.97; CI = 0.33-0.39; ICI = 0.61-0.66; SDI = 1.21—1.25; cu-a slightly proximal to base of Rs&M; marginal cell sparsely hirsute proximally; 1st subdiscal cell hirsute anteriorly and distally. Hind wing with 6-7 hamuli on R1; 1st abscissa of Rs weakly curved, 2nd abscissa almost straight. Fore leg with tibia weakly flattened, with scattered fine spines on outer surface. Mid leg with longer tibial spur 1.5—1.6 times the length of the shorter. Hind leg with coxa in profile 1.8—-1.9 times as long as deep; hind trochantellus in dorsal view about 0.1 times as long as broad; 4th segment of hind tarsus 2.3—2.4 times as long as wide; hind tarsal claws of female with short stout pectinae, those of male similar but with pectinae closer and finer. Gaster long and slender; tergite 2 in profile about 6 times as long as posteriorly deep, its laterotergite folded under and with elliptical thyridium separated from anterior margin of tergite by about 3 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing dense long erect pubescence; gonosquama truncated. Colour generally yellowish, mesoscutum with three dark longitudinal vittae; interocellar area pale yellow; antenna proximally black, distally paler; pterostigma brownish yellow; wings hyaline. VARIATION. None noteworthy. RemaRKS. This species is named in honour of Marta Boza in recognition of her contributions to Costa Rican conservation biology. Enicospilus martae is obviously very closely related to E. columbianus, which it resembles in having the mandibles unusually modified, with the lower tooth distinctly the longer. The two are probably sister- species. E. martae may be recognized on account of its larger size, longer antennae and larger ICI as well as by differences in the alar sclerites (compare Figs 253 and 254). BIOLOGICAL INFORMATION. Enicospilus martae is a Central American species whose range is only known to extend from Costa Rica south to central Panama. It has only been collected in wet forests from near sea- level up to about 750 m. Its host is unknown. MATERIAL EXAMINED Holotype ©’, Panama: Barro Colorado Island, xii.1984 (Wolda) (BMNH). Paratypes. Costa Rica: Guanacaste Prov.: 1 0’, Rincon de la Vieja National Park, 4 km E. of Casetilla, 750 m, xii.1981 (Janzen & Hallwachs) (BMNH). Panama: 1 9, Barro Colorado Island, vi.1985 (Wolda) (BMNH). Enicospilus columbianus (Enderlein) (Figs 172, 254) Henicospilus columbianus Enderlein, 1921: 32. Holotype 9, COLOMBIA (IZPAN) [examined]. Enicospilus columbianus (Enderlein) Townes & Townes, 1966: 176. OPHIONINAE OF TROPICAL MESOAMERICA 181 DEscriPTION. Mandibles moderately short, fairly evenly narrowed, apically twisted 10—20°; upper man- dibular tooth subcylindrical, 0.7—0.8 times as long as the lower tooth (Fig. 172); outer mandibular surface centrally flat, with a weak proximal concavity. Labrum 0.2 times as long as broad; malar space 0.4 times as long as basal mandibular width. Clypeus in profile weakly convex, margin rather sharp; clypeus in front view 1.5—1.6 times as broad as long, with margin weakly convex. Lower face 0.70—0.75 times as broad as long, centrally closely punctate, fairly weakly polished. Head in dorsal view with genae rounded; posterior ocellus contiguous with eye; FI = 60-65%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.8—0.9 times the basal mandibular width away from mandible. Antenna slender with 55-60 flagellar segments; 20th segment 2.3—2.4 times as long as broad. Mesoscutum polished, finely punctate, in profile steeply rounded; notauli vestigial. Mesopleuron polished, the upper part with obsolescent punctures, the lower part similar but with area between punctures finely alutaceous; epicnemial carina inclined towards but not reaching anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for most of its length, but posteriorly the carinae are rather weak; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, polished with fine scattered punctures. Metapleuron with close weak punctures; submetapleural carina weakly ante- riorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area irregularly wrinkled; spiracular area quite long, smooth; posterior area coriaceous, somewhat more weakly sculp- tured than is usual for species of this genus; lateral longitudinal carina complete, sometimes joined to spiracular margin by a short carina. Fore wing length 11-13 mm; discosubmarginal cell as in Fig. 254; AI = 0.52-1.25; CI = 0.15-0.39; ICI = 0.28-0.44; SDI = 1.10-1.16; cu-a from opposite to slightly proximal to Rs&M; marginal cell proximally very slightly more sparsely hirsute than it is centrally, sometimes narrowly glabrous; 1st subdiscal cell with sparse, scattered hairs. Hind wing with 6-7 hamuli on R1; 1st and 2nd abscissae of Rs straight. Fore leg with tibia subcylindrical, with numerous spines on outer surface. Mid leg with longer tibial spur 1.6—1.8 times length of the shorter. Hind leg with coxa in profile 1.8—1.9 times as long as deep; trochantellus dorsally 0.2 times as long as broad; 4th segment of tarsus 2.1—2.3 times as long as broad; claws of female quite long with close long pectinae, those of male with pectinae short and stout. Gaster slender; tergite 2 in profile at least 5 times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by 24 times its own length. Ovipositor slender, its sheath moderately slender. Male with sternites 7—9 bearing scattered long erect hairs; gonos- quama distally from quite evenly rounded to subtruncate. Colour generally yellowish brown, mesoscutal vittae blackish; interocellar area yellow, usually with a small blackish mark between the posterior ocelli; antenna blackish; pterostigma yellowish brown; wings hyaline. VARIATION. The female from Tabasco, Mexico is rather distinctive and may represent a separate species. It has a slightly stouter mandible, and the central sclerite is of similar shape but rotated so its longest axis is at 45° to Rs+2r, rather than at 90° to this vein. Remarks. Enicospilus columbianus is one of only two species in Central America (the other being E. martae) that have the lower tooth of the mandible substantially longer than the upper. E. columbianus also resembles E. martae in having a rather quadrate gonosquama and in having similarly pubescent posterior sternites in the male. E. columbianus differs from E. martae in being smaller, having shorter antennae and a smaller ICI as well as by differences in the alar pubescence and sclerites (compare Figs 253 and 254). BIoLoGicaL INFoRMATION. Enicospilus columbianus is widely distributed from about 18°N in Tabasco, southern Mexico south to about 23°S in Brazil (Map 15). Most specimens have been collected in lowland sites, and despite extensive collecting it has never been taken at Monteverde, Costa Rica (1350 m). Generally only isolated specimens are ever to be collected in one locality. Between 1980 and 1985 three were collected in Santa Rosa National Park, Costa Rica, and three years light-trap samples (1983-5) from Barro Colorado Island, Panama, yielded two further specimens. In Santa Rosa the specimens were collected between June and July (early wet season), whilst in lowland rainforest sites, such as Corcovado National Park, Costa Rica, and on Barro Colorado Island, isolated individuals have been collected between May and August. No details of the biology of this species are known. MATERIAL EXAMINED Holotype 2, Colombia: Rio Magdalena (Pehlke) (IZPAN). Brazil: 1 9 , Rio de Janeiro, iii. 1966 (Townes) (TC); 1 @, Vila Vera, x.1973 (Alvarenga) (TC). Colombia: 1, Valle, Anchicaya, vii.1970 (Howden) (TC). Costa Rica: Cartago Prov.: 1 9, Turrialba, viii.1983 182 IAN D. GAULD Map 15 Localities at which Enicospilus columbianus has been collected. (Porter) (MCZ): Guanacaste Prov.: 2 9, Santa Rosa National Park, 300 m, vi-vii.1984 (Janzen & Hallwachs) (BMNH); 1 ©’, same locality, vi.1985 (Gauld) (BMNH): Puntarenas Prov.: 1 ©’, Sirena, Corcovado National Park, viti.1980 (Janzen & Hallwachs) (BMNH). Mexico: Tabasco: 1 9, Teapa, iii. 1904 (Godman-Salvin) (BMNH). Panama: 1 ©’, Barro Colorado Island, vi.1978 (Wolda) (RNH); 2 ?, same locality, v.1983, vii.1984 (Wolda) (BMNH). Peru: 1 9, Satipo, vii.1943 (Paprzyck) (TC). Venezuela: 10,1 Q, nr Puerto Cabello, xi-xii.1939 (Anduze) (TC). OPHIONINAE OF TROPICAL MESOAMERICA 183 Enicospilus cubensis (Norton) (Figs 255, 278) Ophion cubensis Norton, 1863: 358. LECTOTYPE ©, CUBA (PANS), here designated [examined]. Enicospilus cubensis (Norton) Ashmead, 1900b: 270. Henicospilus cubensis (Norton) Morley, 1912: 33. Enicospilus cubensis (Norton); Townes, 1946: 47. DEscriPTION. Mandibles quite long, weakly and evenly narrowed, apically twisted 10—20°; upper mandibu- lar tooth slightly compressed, about 2.0 times as long as the lower tooth; outer mandibular surface centrally more or less flat, with scattered pubescence, and with a strong proximal concavity. Labrum 0.3-0.4 times as long as broad; malar space 0.3—0.4 times as long as basal mandibular width. Clypeus in profile moderately convex, margin apparently blunt, but extreme margin narrowly impressed; clypeus in front view 1.3—1.4 times as broad as long, with margin very weakly convex. Lower face 0.65—0.70 times as broad as long, with obsolescent fine punctures. Head in dorsal view with genae constricted behind eyes; posterior ocellus very close to eye; FI = 65-70%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 1.0 times the basal mandibular width away from mandible. Antenna long and slender, with 60-66 flagellar segments; 20th segment 2.42.5 times as long as broad. Mesoscutum weakly to strongly polished, finely punctate, in profile evenly rounded; notauli virtually indistinguishable. Mesopleuron polished, the upper part rather sparsely punctate, the lower part, at least in part striately wrinkled; epicnemial carina curved or inclined towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for all of its length; scutellum in dorsal view 1.6-1.9 times as long as anteriorly broad, relatively smooth. Metapleuron closely and shallowly punctate to punctostriate to rather irregularly wrinkled; submetapleural carina evenly but very weakly broadened anteriorly; posterior transverse carina of mesosternum strong, complete. Propodeum in profile evenly rounded; anterior transverse carina more or less complete, posterior transverse carina absent; anterior area long, rather shallowly impressed, striate; spiracular area smooth; posterior area coriaceous to finely reticulate; lateral longitudinal carina present anteriorly, often complete, sometimes joined to spiracular margin by a weak short carina, sometimes without a carina. Fore wing length 12-14 mm; discosubmarginal cell as in Figs 255, 278; AI = 0.92-1.38; CI = 0.30-0.40; ICI = 0.32—0.44; SDI = 1.15—1.25; cu-a more or less opposite the base of Rs&M; marginal cell proximally slightly more sparsely hirsute, often narrowly glabrous adjacent to Rs+2r; 1st subdiscal cell with anterior 0.7 and distal end hirsute. Hind wing with 5—7 hamuli on R1; 1st abscissa of Rs more or less straight, 2nd abscissa straight. Fore leg with tibia slightly flattened, with scattered, fine spines on outer surface. Mid leg with longer tibial spur 1.3—1.4 times length of the shorter. Hind leg with coxa in profile 1.7—1.9 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.4—2.7 times as long as broad; claws of female with long stout pectinae, of male similar with pectinae slightly shorter. Gaster slender; tergite 2 in profile more than 6 times as long as posteriorly deep, laterotergite narrow, membranous, generally pendant, thyridia elliptical, separated from anterior margin of tergite by about 4-6 times its own length. Ovipositor slender, its sheath narrow. Males with sternites 7—9 bearing fine semierect pubescence; aedeagus apically simply rounded; gonosquama rather elongately rounded. Colour generally yellowish, usually with the posterior segments of the gaster black; interocellar area usually black; antenna yellowish or orange, sometimes distally infuscate; pterostigma brownish; wings more or less hyaline. VaRIATION. Enicospilus cubensis shows a considerable range of variation in the colour of the interocellar area. Specimens from Florida in the U.S.A. and Cuba often have this almost entirely yellow though in many examples it is slightly infuscate. In Costa Rican specimens this area is black. Some Floridan and Cuban specimens have the gaster entirely yellowish. There is also a considerable variation in the sculpture of the alitrunk laterally. All specimens seen have some longitudinal wrinkling on the mesopleuron, but the extent of this area is variable, and at its greatest the entire ventral half of the pleuron is so sculptured. Most specimens have a second very faint curved central sclerite in the fenestra. This sclerite together with the more clearly defined central sclerite form a medioventrally interrupted U-shape. In some specimens the second sclerite is quite strongly pigmented. Remarks. Cresson (1928) published a collective designation of lectotypes for many ichneumonid species in the Philadelphia Academy of Natural Sciences. Such collective designations are not valid under the Code (Art. 74c) and, as no subsequent author has validly designated a lectotype for this species, I designate one here. I have selected the specimen labelled as type by Cresson (type number 79 in PANS). This is a male from ‘Cuba O.S.’ There are no other definite syntypes in the Norton collection, but three unlabelled IAN D. GAULD 184 ‘pajoa]]Oo Udaq sey sisuaqgno snjidsonUT Yi ye sattyeso7T oT dep OPHIONINAE OF TROPICAL MESOAMERICA 185 specimens may be syntypic. However, these cannot positively be identified as authentic Norton material. Cresson (1928: 15) stated that the specimen he labelled as type was female, but this is incorrect. Enicospilus cubensis is an extremely distinctive species on account of the unique form of its alar sclerites (see Fig. 255). It is probably closely related to E. trilineatus which it resembles in possessing the following specialized features: a pendant membranous laterotergite 2; and a rather long clypeus. The aedeagus is not flanged, unlike those of some taxa in the E. trilineatus species-group. BIOLOGICAL INFORMATION. Enicospilus cubensis is a very widely distributed species whose cumulative range extends from Florida throughout the Caribbean and Central America, south to Venezuela and Ecuador (Map 16). The most northern record I have seen is of a single female collected in Opelika, Alabama (32.5°N); all other U.S. records to hand are from Florida south of 30°N. This species is most commonly encountered in disturbed habitats such as on the edges of agricultural areas and in suburban gardens. It is very uncommon in pristine forest habitats, and, despite intensive collecting, it has not been collected on Barro Colorado Island, Panama. In Costa Rica E. cubensis is widely distributed, but it is most common in disturbed mid altitude sites between 600 and 1400 m (e.g. at Monteverde or San Antonio de Escazu). In the seasonally dry forests of Santa Rosa National Park, lowland Guanacaste, this species is seldom collected and despite intensive collecting only isolated specimens have been taken in January, May and June. Although this species is common around agricultural areas it has never been reared and no hosts are known. MATERIAL EXAMINED Lectotype 0’, Cuba: ‘O.S.’ (PANS). Bahamas: | 9, Eleuthera, Rainbow Bay, v.1984 (Wiley) (FSCA); 1 co’, Grand Bahama, Pine Ridge, v.1953 (Hayden & Giovannoli) (USNM); 1 2, Grand Bahama, West End, v.1953 (Hayden & Giovannoli) (USNM). Belize: 1 9, W. Highway, mile 15, vi.1968 (Hasse) (FSCA). Costa Rica: Alajuela Prov.: 1 0’, 2 @, Finca Campana, 5 km NW. Dos Rios, 750 m, iii.1985 (Janzen & Hallwachs) (BMNH); 1 9, Finca San Gabriel, 16 km ENE. Quebrada Grande, 650 m, v.1984 (Janzen & Hallwachs); 1 2, same locality, vi.1986 2, 8.2 km N. of Vara Blanca, iv.1984 (Janzen & Hallwachs) (BMNH): Guanacaste Prov.: 1 2, Cerro el Hacha, 3-400 m, xi.1986-i.1987 (Janzen & Hallwachs) (BMNH); 1 9, Rincon de la Vieja National Park, 4km E of Casetilla, 750 m, ii.1982 (Janzen & Hallwachs) (BMNH); 1 CO’, Santa Rosa National Park, 300 m, v. 1980 (Janzen & Hallwachs) (TC); 1 9, same locality and collectors, i.1982 (BMNH); 1 2, same locality and collectors, v.1985 (BMNH); 2 9, same locality, vi.1985 (Gauld) (BMNH): Puntarenas Prov.: 3 Q, Monteverde 1350 m, ii.1962 (Palmer) (TC); 1 2, same locality and collector, ix.1962 (TC); 1 9, same locality, ii. 1963 (Rettenmeyer) (TC); 2 2, same locality, ii.1980 (Mason) (TC); 3 2, same locality, ii.1984 (Cameron) (WC); 1 O’, 2 9, same locality, i-iii.1986 (Haber) (BMNH); 2 9, same locality, i-ii.1986 (Forsyth) (CNC): San José Prov.: 4 9, San Antonio de Escazt, 1300 m, v-vi.1981 (Eberhard) (UMC);1 1.00; having the lower tooth of mandible swollen ventrally; having the posterior transverse carina of mesoster- num centrally weak or interrupted; having the metapleuron punctate; lacking a central sclerite. Struc- turally it resembles E. vegai (see that species). BIOLOGICAL INFORMATION. Enicospilus laurenae is widely distributed from Costa Rica south to Ecuador. It is an uncommon insect in collections so little is known of its habitat preferences, but the Costa Rican and Panamanian specimens were collected in wet forest. It is perhaps noteworthy that E. /aureni has never been collected in seasonally dry forests, despite the fact that this was the most intensively sampled habitat in this study. This species has never been reared. MATERIAL EXAMINED Holotype CO’, Costa Rica: Limon Prov., Cerro Tortuguero, N. edge Tortuguero National Park, 0-100 m, v.1984 (Janzen & Hallwachs) (BMNH). Paratypes. Costa Rica: Cartago Prov.: 1 0’, IICA, Turrialba, viii. 1963 (Porter) (USNM). Ecuador: 1’, Balzapomba, vi.1938 (MacIntyre) (TC). Panama: 1 9, Barro Colorado Island, xi.1941 (Zetek) (USNM). Enicospilus vegai sp. n. (Figs 178, 266) Description. Mandibles quite long, proximally evenly tapered, distally more or less parallel-sided, its apex twisted 20-30°; upper mandibular tooth slightly depressed, 1.3-1.5 times as long as the lower which is stouter, slightly compressed, and swollen ventrally; outer mandibular surface sparsely pubescent, more or less flat with a weak proximal concavity. Labrum 0.2 times as long as broad; malar space 0.2 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.31.4 times as broad as long, its margin weakly convex. Lower face 0.65—0.68 times as broad as long, polished, punctate. Head in dorsal view with genae rounded; posterior ocellus contiguous eye; FI = 75-80; occipital OPHIONINAE OF TROPICAL MESOAMERICA 201 carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.9 times the basal mandibular width away from mandible. Antenna long and slender, with 66-69 flagellar segments; 20th segment 1.9-2.1 times as long as broad. Mesoscutum polished, finely punctate, in profile evenly rounded; notauli vestigial. Mesopleuron pol- ished, the upper part punctate, the lower part punctostriate grading centrally to striate; epicnemial carina inclined towards anterior margin of pleuron, its upper end weak and curved. Scutellum in profile weakly convex, laterally carinate for all of its length; scutellum in dorsal view 1.6-1.7 times as long as anteriorly broad, punctate, posteriorly wrinkled. Metapleuron weakly convex, striate grading anteroventrally to punctate (Fig. 178); submetapleural carina not distinctly broadened anteriorly; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina complete except lateral extremities, posterior transverse carina absent; anterior area deeply excavate, striate; spiracular area moderately long, smooth, posterior area reticulate laterally, centrally with rugae tending to be concentric; lateral longitudinal carina present only as an anterior vestige, sometimes joined to spiracular margin by a short weak carina. Fore wing length 17-18 mm; discosubmarginal cell as in Fig. 266; AI = 0.61-1.00; CI = 0.28-0.39; ICI = 0.69-0.83; SDI = 1.20-1.25; cu-a proximal to the base of Rs&M by about 0.2-0.3 times its own length; marginal cell proximally more or less uniformly hirsute; 1st subdiscal cell hirsute except for posterior and proximal margins. Hind wing with 7 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa almost straight. Fore leg with tibia slightly flattened, with numerous, slender spines on outer surface. Mid leg with longer tibial spur 1.4—1.5 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally. VARIATION. Structurally this is a remarkably uniform species and the Brazilian specimens are virtually identical to the Mesoamerican individuals. REMARKS. This species is named in honour of Gerardo Vega for his tireless efforts as a research assistant for Daniel Janzen. Enicospilus vegai can be recognized by the combination of the stout, ventrally swollen lower tooth of the mandible, striate metapleuron and absence of central sclerite. A number of other Mesoamerican and Neotropical species have very similar mandibles to vegai, and these species also resemble each other in coloration and, usually, in the sculpture of the propodeum. Possibly they comprise a holophyletic group. In Central America this species-complex includes E. laurenae and E. kelloggae. Both may be distinguished from E. vegai because they have punctate metapleurae; E. kelloggae has a distinct central sclerite. E. laurenae may easily be separated from E. vegai by the features given in couplet 55. BIOLOGICAL INFORMATION. Enicospilus vegai is a widely distributed species whose range extends from southern Costa Rica (9°N) south to Rio de Janeiro Province, Brazil (23°S). Nothing is known of the biology of this species. MATERIAL EXAMINED Holotype ©’, Costa Rica: Puntarenas Prov., Esquinas, nr Golfito (Allen) (MCZ). Paratypes. Brazil: 2 2, Rio de Janeiro, Silva Jardin, viii.1974 (Oliveira) (TC). Panama: 1 ©’, Las Cumbres, v.1982 (Wolda) (TC). Enicospilus baltodanorum sp. n. (Figs 102, 104, 267) DescriPTIon. Mandibles moderately long, evenly narrowed, apically twisted 30-40°; upper mandibular tooth slender, depressed, 1.4 times as long as the lower tooth; outer mandibular surface finely pubescent, flat, but with a broad shallow proximal concavity. Labrum 0.3 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin subacute; clypeus in front view 1.5 times as broad as long, the margin almost truncate (Fig. 102). Lower face 0.76 times as broad as long, centrally slightly granulate, closely punctate. Head in dorsal view with genae constricted behind eyes; posterior ocellus contiguous with eye; FI = 70%; occipital carina mediodorsally complete but weak, ventrally curved to join hypostomal carina about 0.5 times the basal mandibular width away from mandible. Antenna slender, with 61 flagellar segments; 20th segment 1.8 times as long as broad. Mesoscutum weakly polished, shallowly punctate, in profile evenly rounded; notauli shallow, but extending back to level of tegulae. Mesopleuron polished, the upper part finely and closely punctate, the lower part with punctures more superficial and larger; epicnemial carina curved towards anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for 0.6 of its length; scutellum in dorsal view 1.2 times as long as anteriorly broad, regularly punctate. Metapleuron rather weakly convex, 202 IAN D. GAULD punctate, with fine granulation between punctures (Fig. 104); submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum more or less complete. Propodeum in profile abruptly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area deep and long, finely striate; spiracular area short, punctate; posterior area finely irregularly wrinkled; lateral longitudinal carina present only anteriorly, but joined to spiracular margin by a short carina. Fore wing length 18 mm; discosubmarginal cell as in Fig. 267; AI = 0.95; CI = 0.25; ICI = 0.89; SDI = 1.21; cu-a proximal to the base of Rs&M by 0.3 times its own length; marginal cell proximally evenly hirsute; 1st subdiscal cell with anterior and distal margins broadly hirsute. Hind wing with 8 hamuli on R1; 1st abscissa of Rs almost straight, 2nd abscissa straight. Fore leg with tibia quite strongly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.5 times length of the shorter. Hind leg with coxa in profile 1.7 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus missing; claws of female with close even pectinae. Gaster moderately stout; tergite 2 in profile 3.7 times as long as posteriorly deep, laterotergite folded under, thyridia oval and separated from anterior margin of tergite by about 3.5 times its own length. Ovipositor slender, its sheath moderately stout. Male unknown. Colour generally pale yellowish, legs darker brownish yellow and with gaster weakly infuscate posteri- orly; mesoscutum with three darker longitudinal stripes; interocellar area slightly infuscate centrally; antenna brownish, paler distally; pterostigma dark brown; wings weakly infumate. RemakkS. This species is named after Jorge, Jorge Enrique and Aristides Baltodano who have been very supportive of the Guanacaste National Park project. Enicospilus baltodanorum is a rather stout species that is most easily recognized by its almost quadrate scutellum and very characteristic alar sclerite. It is apparently closely related to E. estradarum, which it resembles in the form of the mandibles and possession of a complete distal sclerite. It also resembles E. colini in sculpture, but differs in having less strongly twisted mandibles, no lateral crest on the propodeum and a smaller ICI. BIOLOGICAL INFORMATION. Enicospilus baltodanorum is only known to occur in Costa Rica. Nothing is known about its biology. MATERIAL EXAMINED Holotype 2, Costa Rica: Puntarenas Prov., Esquinas, nr Golfito (Allen) (MCZ). Enicospilus carri sp. n. (Figs 105, 108, 268) Description. Mandibles moderately long, proximally abruptly narrowed, distally weakly narrowed, apically twisted 60—70°; upper mandibular tooth rather slender, slightly depressed, 1.3-1.5 times as long as the slightly compressed lower tooth; outer mandibular surface with long fine hair centrally, rather flat with a weak proximal concavity. Labrum 0.2-0.3 times as long as broad; malar space 0.20.3 times as long as basal mandibular width. Clypeus in profile very weakly convex, margin subacute; clypeus in front view 1.4— 1.5 times as broad as long, apically truncate. Lower face 0.63—-0.66 times as broad as long, centrally shallowly punctate. Head in dorsal view with genae constricted behind eyes; posterior ocellus contiguous with eye; FI = 80-85% ; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.9 times the basal mandibular width away from mandible. Antenna long and slender, with apical flagellar segments missing; 20th segment 1.9-2.0 times as long as broad. Mesoscutum polished with shallow fine scattered punctures, in profile evenly rounded with anterior margin strongly out-curved; notauli shallow but conspicuous. Mesopleuron polished, the upper part sparsely punctate, the lower part punctostriate to striate; epicnemial carina inclined towards anterior margin of pleuron, but with upper end evanescent (Fig. 108). Scutellum in profile weakly convex, laterally carinate for 0.9 or more of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, punctate, with isolated rugae posteriorly. Metapleuron moderately convex, diagonally striate (Fig. 105); submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile abruptly rounded; anterior transverse carina complete except at lateral extremities, posterior transverse carina represented by a weak lateral crest; anterior area deeply excavate, with isolated rugae; spiracular area short, finely and closely punctate; posterior area rugose; lateral longitudinal carina present anteriorly, joined to spiracular margin by a short weak carina. Fore wing length 18-19 mm; discosubmarginal cell as in Fig. 268; AI = 0.76-0.80; CI = 0.21-0.35; ICI = 0.81-1.00; SDI = 1.22-1.27; cu-a proximal to the base of Rs&M by 0.2-0.3 times its own length; OPHIONINAE OF TROPICAL MESOAMERICA 203 marginal cell proximally slightly more sparsely hirsute than it is centrally; 1st subdiscal cell with anterior and distal periphery hirsute. Hind wing with 7-8 hamuli on R1; 1st abscissa of Rs almost straight, 2nd abscissa straight. Fore leg with tibia slightly flattened, with scattered conspicuous spines on outer surface. Mid leg with longer tibial spur 1.3—1.4 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.3—2.4 times as long as broad; claws of female unusually long, bearing long fine pectinae and distally abruptly rounded. Gaster long and slender; tergite 2 in profile 5.5—6.0 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 4.0-4.5 times its own length. Ovipositor slender, its sheath moderately narrow. Male unknown. Colour generally brownish yellow, with three dark brown longitudinal vittae on the mesoscutum; gaster reddish brown; interocellar area yellowish, infuscate between posterior ocelli; antenna proximally black- ish, distally brown; pterostigma brown; wings hyaline. VARIATION. None remarkable. REMARKS. This species is named in honour of Archie Carr for his intense conservation activities. Enicospilus carri is quite similar to E. stevensi in general appearance but differs most conspicuously in colour pattern. E. carri has dark mesoscutal vittae and a more or less uniformly reddish brown gaster; E. stevensi has a more or less uniformly brownish mesoscutum and a yellowish gaster with the posterior tergites black. E. stevensi has a far more convex metapleuron than E. carri and the females have much broader ovipositor sheaths. BIOLOGICAL INFORMATION. Enicospilus carri has a range which extends from Belize (17°N) south to about 3°S in Amazonas Province in Brazil. The species has only been collected in lowland rain forest. Its hosts are not known. MATERIAL EXAMINED Holotype , Belize: Toledo, Columbia Forest Station, vii.1968 (Hasse) (BMNH). Paratype. Brazil: 1 2, Amazonas Prov., Reserva Ducke, km 26 Manaus-Itacoatiara highway, v.1972 (Munroe family) (CNC). Enicospilus colini sp. n. (Figs 109, 269) Description. Mandibles of moderate length, proximally strongly narrowed, apically twisted 50-60°; upper mandibular tooth subcylindrical, 1.5—1.9 times as long as the lower tooth; outer mandibular surface sparsely pubescent, relatively flat, with a small proximal concavity. Labrum 0.3 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile convex, margin blunt; clypeus in front view 1.5—1.6 times as broad as long, with margin very weakly convex. Lower face 0.74-0.78 times as broad as long, closely, finely punctate. Head in dorsal view with genae rounded behind the eyes; posterior ocellus very close to eye; FI = 60-70%; occipital carina mediodorsally weak or, more usually, narrowly interrupted, ventrally curved to join hypostomal carina about 0.6—0.8 times the basal mandibular width away from mandible. Antenna long but stout, with 62-64 flagellar segments; 20th segment 1.6-1.9 times as long as broad. Mesoscutum polished, finely punctate, in profile evenly rounded; notauli weakly impressed. Meso- pleuron polished, the upper part closely punctate ventrally grading to punctostriate or even striate; epicnemial carina inclined towards anterior margin of pleuron, its lower corner acute (Fig. 109). Scutellum in profile moderately convex, laterally carinate to centre, behind this the carinae evanescent; scutellum in dorsal view 1.5-1.6 times as long as anteriorly broad, punctate. Metapleuron convex, anteroventrally punctate becoming coarsely striate or even rugose posterodorsally; submetapleural carina quite strongly broadened anteriorly; posterior transverse carina of mesosternum well developed, complete. Propodeum in profile abruptly declivous; anterior transverse carina complete, posterior transverse carina represented by lateral ridges; anterior area deeply impressed, striate; spiracular area short, smooth; posterior area reticulately wrinkled; lateral longitudinal carina present only anteriorly only as a vestige, not joined to spiracular margin by a short carina. Fore wing length 21-25 mm; discosubmarginal cell as in Fig. 269; AI = 0.80-1.00; CI = 0.28-0.32; ICI = 1.35-1.45; SDI = 1.25-1.35; cu-a proximal to base of Rs&M by 0. 1-0.3 times its own length; marginal cell proximally evenly hirsute; 1st subdiscal cell with anterior and distal margins hirsute. Hind wing with 6-8 hamuli on R1; 1st abscissa of Rs very weakly bowed, 2nd abscissa from straight to slightly sinuous. Fore leg with tibia slightly flattened, with numerous long, slender spines on outer surface. Mid leg with 204 IAN D. GAULD longer tibial spur 1.3—1.5 times length of the shorter. Hind leg with coxa in profile 1.6—-1.7 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.2—2.4 times as long as broad; claws of female short, abruptly rounded apically, with close pectinae; claws of male similar but with pectinae slightly shorter. Gaster quite slender; tergite 2 in profile 4—S times as long as posteriorly deep, laterotergite folded under, thyridia oval and separated from anterior margin of tergite by about 5—6 times its own length. Ovipositor moderately slender, its sheath broad. Male with sternites 7-9 bearing long, stout, erect hairs; gonosquama evenly rounded. Colour generally yellowish brown with head, scutellum and mesoscutal stripes paler yellowish; inter- ocellar area yellow; antennae blackish at least on basal 0.3; pterostigma orange; wings weakly infumate. VARIATION. There is considerable variation in the colour of the mesoscutum. In most specimens it is yellowish brown, with lateral and lateromedian longitudinal pale stripes. The majority of the specimens from Mexico and occasional individuals from Santa Rosa have the mesoscutum blackish, although the pale stripes are still present. The Panamanian specimen is generally darker in colour than any others. The male from Virgen de Socorro has a rather atrophied alar sclerite, a longer fenestra, an anteriorly more hirsute discosubmarginal cell and the occipital carina complete. Possibly it represents a separate species, so I have excluded it from the paratype series. REMARKS. This species is named after the anthropologist Dr Colin Chapman in gratitude for his help and companionship in the field. Enicospilus colini may easily be recognized by the combination of the following features: the large ICI (>1.30); no central sclerite; epicnemial carina sharply angled ventrally; its large size (fore wing length >20 mm). E. colini is superficially similar to E. stevensi, but differs not only in the form of the venation, but usually in having an incomplete or weak occipital carina and no carina between the propodeal spiracle and the lateral carina. BIOLOGICAL INFORMATION. Enicospilus colini is a widespread Neotropical species whose range extends from about 15°N in southern Mexico, south to about 26°S in north-eastern Argentina. It occurs from near sea- level up to an altitude of about 1400 m. In Santa Rosa National Park relatively few individuals have ever been collected at once, but isolated individuals turn up at light regularly. Their seasonal distribution may be summarized thus: I F M A Melegra Ss oO NM BD Se a ee ee ee | The June/July specimens were all females whilst those collected in December were all males. In the higher, wetter parts of Costa Rica I have seen females collected between October and March. Despite intensive collecting this species has never been taken on Barro Colorado Island, Panama. The hosts of this insect are unknown. MATERIAL EXAMINED Holotype 2, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, vi.1985 (Gauld) (BMNH). Paratypes. Argentina: 1 9, Misiones, x.1960 (Foerster) (CNC). Colombia: 1 ©’, Valle, Anchicaya, nr Buenaventura, 300 m, iv.1972 (Cooper) (BMNH). Costa Rica: Cartago Prov.: 1 2, 2 0’, Turrialba, iii.1965 (Duckworth) (USNM): Guanacaste Prov.: 1 2, Rincén de la Vieja, 900 m, iii.1984 (Janzen, Hallwachs & Gauld) (BMNH); 2 ©’, Santa Rosa National Park, 300 m, xii.1979 (Janzen) (BMNH); 2 o’, 4 9, same locality, vii.1982, v,vi,xii.1983 & vi.1984 (Janzen & Hallwachs) (BMNH); 1 Q, same locality, vi.1986 (Gauld) (BMNH);2 9, same locality, SE. of Entrada, vi.1986 (Godfrey) (BMNH): Heredia Prov.: 1 9, La Selva Biological Station, Puerto Viejo de Sarapiqui, 40 m, ii.1986@ (Chavarris & Chacon) (BMNH): Puntarenas Prov.: 1 9, Monteverde, xii.1962 (Palmer) (TC); 1 9, Monteverde, 1350 m, i.1986 (Forsyth) (CNC): San José Prov.: 2 9, Estacion Carrillo, Braulio Carrillo National Park, 700 m, x.1984 (Chacon) (BMNH); 1 co’, San José, i.1961 (Palmer) (TC). Mexico: Chiapas: 2 0’, 32 km N. Huixtla, 1000 m, vi. 1969 (Masan) (CNC); 10’, 30-40 km N. Huixtla, 1000 m, vi.1969 (Peterson) (CNC); 2 0, 30-40 km N. Huixtla, 1000 m, vi.1969 (CNC); 1 2, Muste, nr Huixtla, 440 m, x.1970 (Welling) (CNC); 1 Oo’, Municipio de Tzimol, 30 km SSE. Pugiltic, 762m, x.1981 (Breedlove) (CAS): Quintana Roo: 1 CO’, X-can, viii.1963 (Welling) (CNC). Panama: 1 9, Chiriqui, Fortuna, 1050 m, ii.1978 (Wolda) (RNH). Peru: 1 9, Quincemil, Cuzco, xi.1962 (Pena) (CNC). Non-paratypic material. Costa Rica: Alajuela Prov.: 1 0’, Virgen de Socorro, 800 m, ii.1987 (Janzen & Hallwachs) (BMNH). OPHIONINAE OF TROPICAL MESOAMERICA 205 Enicospilus maculipennis (Cameron) (Fig. 270) Ophion (Enicospilus) maculipennis Cameron, 1886: 292. Holotype 9, PANAMA (BMNH) [examined]. Enicospilus maculipennis (Cameron) Ashmead, 1895: 547. Henicospilus maculipennis (Cameron) Dalla Torre, 1901: 182. Enicospilus parvifasciatus Cameron, 1911: 180. Holotype 9, GUYANA (BMNH) [examined]. Syn. n. Enicospilus parvifasciatus Cameron; Hooker, 1912: 87. Stauropodoctonus maculipennis (Cameron) Morley, 1912: 18 Description. Mandibles quite long, proximally narrowed, distally almost parallel-sided, apically twisted 15—25°; upper mandibular tooth slightly compressed, 1.1—1.3 times as long as the lower; outer mandibular surface slightly concave, with scattered fine hairs. Labrum 0.3—0.4 times as long as broad; malar space 0.3— 0.4 times as long as basal mandibular width. Clypeus in profile weakly convex, margin impressed, acute; clypeus in front view 1.25—1.45 times as broad as long, with margin apically weakly convex. Lower face 0.70-0.75 times as broad as long, polished with very fine punctures. Head in dorsal view with genae rounded behind eyes; posterior ocellus very close to eye; FI = 60-65%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.8—0.9 times the basal mandibular width away from mandible. Antenna long and slender, with 60-65 flagellar segments; 20th segment 2.8-3.3 times as long as broad. Mesoscutum weakly polished, finely and inconspicuously punctate, in profile evenly rounded; notauli vestigial. Mesopleuron polished, the upper part finely punctate, the lower part similar but with a shallowly rugose band of sculpture extending from epicnemial carina towards base of mid coxa; epicnemial carina curved towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.7 or more of its length; scutellum in dorsal view 1.6—1.8 times as long as anteriorly broad, smooth or with striae posteriorly. Metapleuron moderately convex, punctate; submetapleural carina barely broadened ante- riorly; posterior transverse carina of mesosternum complete. Propodeum in profile evenly rounded; anterior transverse carina complete, posterior transverse carina absent; anterior area short and quite shallowly impressed, with a few coarse striae; spiracular area long, with obsolescent punctures; posterior area coarsely and irregularly wrinkled; lateral longitudinal carina complete, at least anterior to transverse carina, joined to spiracular margin by a short and often weak carina. Fore wing length 13-15 mm; discosubmarginal cell as in Fig. 270; AI = 1.05-1.67; CI = 0.44-0.64; ICI = 0.26-0.34; SDI = 1.19-1.39; cu-a opposite or slightly proximal to base of Rs&M; marginal cell proximally uniformly hirsute; 1st subdiscal cell with anterior and distal parts sparsely hirsute. Hind wing with 5—6 hamuli on R1; 1st abscissa of Rs weakly bowed, 2nd abscissa almost straight. Fore leg with tibia not distinctly flattened, with slender scattered spines on outer surface. Mid leg with longer tibial spur 1.6—1.8 times length of the shorter. Hind leg with coxa in profile 1.7—1.9 times as long as deep; trochantellus dorsally 0.1—-0.2 times as long as broad; 4th segment of tarsus 2.6-2.7 times as long as broad; claws of female short, strongly but evenly curved apically, with close regular pectinae; claws of male similar. Gaster slender; tergite 2 in profile at least 6 times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by about 3 or more times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing long fine erect pubescence; gonosquama quite long, distally rounded. Colour generally pale yellow; antennae and interocellar area yellow; frons between antennae, occiput, propleuron, mesoscutal vittae, mesothorax ventrally and laterally on epicnemium, lower part of meta- pleuron, bases of mid and ventral parts of hind coxae black; pterostigma black, proximally and distally yellow; wings hyaline, proximal corner of marginal cell conspicuously infumate. VARIATION. There is a considerable range of variation in the extent of the areas of black pigmentation. Some Costa Rican specimens have virtually all the mesopleuron black except for the upper 0.3 and a small area in the lower hind corner. The propodeum, pronotum, posterior 0.3 of tergite 1 and much of tergite 3 may be blackish also. Specimens from further south (Venezuela) are much less extensively dark and generally lack all gastral and coxal markings and frequently also those on the metathorax, propodeum and prothorax. The alar sclerite and pterostigma of Venezuelan specimens is also paler, and the punctures on the metathorax are finer and sparser. These specimens also lack the characteristic band of rugose sculpture on the mesopleuron. RemarKS. Enicospilus maculipennis is most easily recognized by the characteristic coloration of the alitrunk, gaster and pterostigma, and by the infumation present at the base of the marginal cell. It is the only Neotropical species without a central sclerite that has the marginal cell infumate. The shape of the alar 206 IAN D. GAULD sclerite and the venation are also quite characteristic. In sculpture E. maculipennis resembles E. flavo- scutellatus but it is not clear whether this similarity is due to evolutionary convergence or results from a close phylogenetic relationship. Morley (1912) placed maculipennis in Stauropodoctonus (Morley’s name for the genus now called Stauropoctonus) on account of its colour pattern; he subsequently (Morley, 1914) examined the holotype of E. parvifasciatus which he correctly recognized as being synonymous with maculipennis. However, the holotype of E. parvifasciatus was inadvertently labelled by Cameron as ‘Enicospilus parvimaculatus’ and Morley (1914) used this name when suggesting the synonymy. Consequently the synonymy which Morley correctly recognized has not been published, so I have formally established it above. BIOLOGICAL INFORMATION. Enicospilus maculipennis is a Mesoamerican and northern South American species whose range extends from about 15°N in southern Mexico southwards to Ecuador, Colombia and Venezuela. It has been collected from near sea-level up to an altitude of 2300 m. It is known to occur ina variety of habitats, but despite quite intensive collecting it has not been collected in lowland rainforest on Barro Colorado Island, though it does occur in a more disturbed habitat at Las Cumbres. In Santa Rosa National Park E. maculipennis is regularly collected, but never in large numbers. The specimens from this site have the following aggregated seasonal distribution: TEs NiGAY (REST eI APS MOT aire 1D were ee I I Sr 2 a No hosts are known for this species. MATERIAL EXAMINED Holotype 2 (Ophion (Enicospilus) maculipennis Cameron), Panama: Bugaba (BMNH). Holotype 2 (Enicospilus parvifasciatus Cameron), Guyana (BMNH). Colombia: 3 2, Monterredondo, Cundinamarca, ii.1957, i-ii.1959 (Foerster) (CNC); 19,41 kmS. Santa Marta Magdalena, 2300 m, v.1973 (Howden & Campbell) (TC). Costa Rica: Guanacaste Prov.: Santa Rosa National Park, 300 m, 1 9, xii.1976, 3 Q, viii-ix.1977, 1 2, v.1978, 1 Q, vii.1978, 1 ©’, xii.1979 (Janzen) (TC); 3 0’, 5 Q, same locality, iii.1981, viii, xi-xii.1982, iv.1983 (Janzen & Hallwachs) (BMNH); 2 9, Volcan Cacao, Finca La Luz [Estacion Mengo], 1100 m, i, iii.1987 (Janzen & Hallwachs) (BMNH); 2 °, Volcan Orosi, Casa Mariksa [= Maritza], 800 m, v-vi. 1986 (Gauld) (BMNH): Heredia Prov.: 1 0’, Finca La Selva, 3 km S. Puerto Viejo, vi.1986 (Hespenheide) (BMNH): Puntarenas Prov.: 1 9, Monteverde, 1350 m, viii.1985 (Haber) (BMNH). Ecuador: 1 9, San Mateo, Esmeraldas, vi.1956 (Foerster) (CNC). El Salvador: 1 2, 10 km W. Quezaltepeque, viii. 1963 (Cavagnaro & Irwin) (CAS). Mexico: Chiapas: 1 2, 3 km N. Huixtla, 750 m, vi.1969 (Mason) (CNC); 1 9, Muste, nr. Huixtla, 440 m, viii.1970 (CNC). Panama: 10,2 9, Las Cumbres, v-vii.1982 (Wolda) (TC). Venezuela: 1 0’, 1 2, Antimano, 950 m, vi.1938, vi.1939 (Berthier) (TC); 3 9, nr Caracas, x.1938, 1942 (TC); 1 OC’, Caripito, vii.1947 (Anduze) (TC); 1 oO, El Encanto, Mir., v.1939 (Lopez) (TC); 1 2, Tucuco, Zulia, iv.1981 (Townes) (TC). Enicospilus guatemalensis (Cameron) (Fig. 271) Ophion (Enicospilus) guatemalensis Cameron, 1886: 293. Holotype C’ [not 9 as stated in description], GUATEMALA (BMNH) [examined]. Henicospilus guatemalensis (Cameron) Dalla Torre, 1901: 182. Enicospilus guatemalensis (Cameron) Hooker, 1912: 68. Description. Mandibles quite long, proximally quite strongly narrowed, distally more or less parallel- sided, apically twisted 30-45°; upper mandibular tooth slightly compressed, 1.6-1.9 times as long as the lower tooth; outer mandibular surface sparsely pubescent, centrally slightly convex, with a very weak proximal concavity. Labrum 0.2-0.4 times as long as broad; malar space 0.30.4 times as long as basal mandibular width. Clypeus in profile almost flat, margin rather acute, not impressed; clypeus in front view 1,2-1.4 times as broad as long, with margin weakly convex apically. Lower face 0.65—0.75 times as broad as long, polished, with fine punctures. Head in dorsal view with genae strongly narrowed behind eyes; posterior ocellus very close to eye; FI = 60-70% ; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6—-0.8 times the basal mandibular width away from mandible. Antenna long but quite stout, with 55-60 flagellar segments; 20th segment 1.6-2.0 times as long as broad. Mesoscutum polished with shallow close punctures, in profile evenly rounded; notauli vestigial. Meso- pleuron weakly to strongly polished, the upper part sparsely and finely punctate, the lower part with OPHIONINAE OF TROPICAL MESOAMERICA 207 transverse wrinkles; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for all of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, smooth or irregularly wrinkled. Metapleuron moderately convex, punctostriate to striate or slightly rugose posterodorsally; submetapleural carina weakly and evenly anteriorly broadened; pos- terior transverse carina of mesosternum usually weak or indistinct centrally or absent either side of the mid line. Propodeum in profile abruptly declivous; anterior transverse carina more or less complete, posterior transverse carina absent or represented as lateral vestiges; anterior area quite short and deeply impressed, coriaceous-striate; spiracular area short, smooth; posterior area reticulately wrinkled; lateral longitudinal carina more or less complete, joined to spiracular margin by a short carina. Fore wing length 12-16 mm; discosubmarginal cell as in Fig. 271; AI = 0.73-1.15; CI = 0.29-0.37; ICI = 0.38-0.53; SDI = 1.00-1.16; cu-a from subopposite to base of Rs&M, to proximal to it by up to 0.3 times its own length; marginal cell proximally slightly more sparsely hirsute; 1st subdiscal cell with distal end and anterior 0.6 hirsute. Hind wing with 6-8 hamuli on R1; Ist abscissa of Rs straight, 2nd abscissa weakly arcuate. Fore leg with tibia subcylindrical, with fine scattered spines on outer surface. Mid leg with longer tibial spur 1.3-1.5 times length of the shorter. Hind leg with coxa in profile 1.7-1.8 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 1.9-2.4 times as long as broad; claws of female with long close pectinae, those of male similar but with shorter pectinae. Gaster long and slender; tergite 2 in profile at least 5 times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by about 3.5 or more times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing scattered erect hairs; gonosquama evenly rounded. Colour generally yellowish orange with or without darker marks as discussed below; interocellar area and antennae yellow; pterostigma orange; wings hyaline. VARIATION. This species exists in two distinctive colour morphs. The widespread form (which includes the holotype) is uniformly yellowish orange, and slightly more robust, whilst there is an apparently conspecific form in Ecuador which is black-marked on the mesoscutum, mesopleuron and mesosternum, propodeum, mid and hind coxae, hind femur, posterior parts of tergites 1-2 and most of 3+. The dark form is slightly more slender, and tends to be more striately sculptured laterally than does the widespread form. Speci- mens from the Dominican Republic have dark mesoscutal vittae. A relatively large (fore wing length 17 mm) female from the Dominican Republic is tentatively referred to this species. It differs from other specimens in that the alitrunk is dark reddish brown with large irregular pale yellowish markings. The alar pubescence is sparser and the distal sclerite is slightly broadened distally. RemaRrkS. The long, large fenestra distinguishes Enicospilus guatemalensis from all other species that have a yellow interocellar area and lack a central sclerite. Its phylogenetic position is uncertain. BIOLOGICAL INFORMATION. Enicospilus guatemalensis is an extremely widespread species whose range extends throughout Mexico, from about 26°N, south into Guatemala and from about 30°N in Florida southwards to the Bahamas and onto the older, larger Caribbean islands (Map 19). In South America it is known from Venezuela westwards into Ecuador and south to Argentina. Despite intensive collecting it has not been taken in either Costa Rica or Panama. Whether this indicates a real break in the distribution of this species, or whether it occurs in habitats so far not sampled only further collecting will reveal. E. guatemalensis has been collected from almost sea level up to 1700 m. It has often been collected in rather open forests, but in Florida it has been taken in suburban situations. No hosts are known for this species. MATERIAL EXAMINED Holotype oO’, Guatemala: San Geronimo (BMNH). Argentina: 1 0, Yuto, xi.1966 (Townes & Townes) (TC). Bahamas: | ©’, Great Abaco, Simons Pt, i.1984 (Teale) (USNM); 2 &’, 1 2, Man-o-War Cay, viii.1971 (Howden) (TC). Brazil: 1 2, Minas Gerais, Pedra Azul, 800 m, xi.1972 (Alvarenga & Seabra) (TC). Cuba: 1 9, Guantanamo Bay, ii.1965 (Porter) (FSCA). Dominican Republic: 1 0’, Monte Cristi Prov., 10 km S. Monte Cristi, 5 m, v.1973 (Davis & Davis) (USNM); 2 @,, San Lorenzo, v.1915 (USNM); 1c’, 1 Q, Santiago, iii.1936 (Rosario) (MCZ); 2 9, Santiago, La Cumbre, 1000 m, vi.1978 (Woodruff) (FSCA). Ecuador: 4 2, Rio Leon, 1700 m, iii.1965 (Peria) (TC). Guatemala: 1 2, San Geronimo (BMNH). Jamaica: 1 C’, Trelawny Parish, Trelawny Beach Hotel, viii.1985 (Eger) (BMNH). Mexico: Baja California: 1 CO’, El Sargento, vi.1983 (Stange & Miller) (FSCA); 10°, 8kmN. La Paz on highway 1, xii.1978 (Weissman et al.) (CAS); 19, 10kmN. Todos Santos, road to La Paz, xii.1958 (Leech) (CAS); 1 CO’, St Nicolas Bay, Gulf of California, v.1921 (Chamberlin) (CAS): Chiapas: 1 2, San Cristobal, 2000 m, v.1969 (Teskey) (CNC): Mexico State: 1 0’, Cuernavaca, viii.1951 (Hull) (CNC): Morelos: 1 ©’, nr Tijalpa, vi.1963 (Woodruff) (FSCA): Nayarit: 1 2, Vic. 208 IAN D. GAULD Map 19 Localities at which Enicospilus guatemalensis has been collected. Compostela, vi.1934 (TC): Nuevo Leén: 1 9, 8 km S. Monterrey, vii.1963 (Howden) (BMNH): Oaxaca: 1 GO, Ixtpec, viii.1939 (Townes) (TC): Quintana Roo: 1 O’, X-can, viii.1963 (Welling) (CNC): San Luis Potosi: 1 9,2 km W. Tamazanchale, viii.1972 (Hevel & Hevel) (USNM): Sinaloa: 2 9, Mazatlan, viii.1971 (Davis) (TC); 1 Oo’, Mazatlan, vii.1939 (Townes) (TC); 1 9, 43 km E. Villa Union, vii.1964 (Howden & Howden) (CNC); 10,45 km E. Villa Union, viii.1964 (Martin) (CNC): Tamulipas: 2 9 , Ciudad Victoria, vili.1972 (Hevel & Hevel) (USNM). U.S.A.: Florida: 5 9, Alachua Co., Austin Cary Memorial Forest, iv-vii.1975 (Fairchild) (FSCA); 1 9, same locality and collector, iv.1982 (FSCA); 1 9, Alachua Co., Gainesville, iv.1971 (Mead) (FSCA); 2 9, same locality, iv.1975 (Mead) (FSCA); 1 9, same locality, xi.1975 (Weems) (FSCA); 1 OO’, Highlands Co., Archbold Biological Station, i.1962 (Frost) (TC); 1 9, OPHIONINAE OF TROPICAL MESOAMERICA 209 Jackson Co., Florida Caverns State Park, xii.1951 (Denmark) (FSCA); 2 9, same locality, xii.1957 (Weems) (TC); 1 9, Marion Co., 14km SSW. Ocala, Kingslane Country Est. , x.1975 (Wiley) (FSCA);1 9, Monroe Co., Bahia Honda State Park, vii.1973 (Woodruff) (FSCA); 1 9, Seminole Co., Longwood, ii.1975 (Mason) (CNC). Venezuela: 1 9, Aragua, Rancho Grande, iii-iv.1960 (Test) (UM). Enicospilus karrensis sp. n. (Figs 179, 272) Description. Mandibles rather short, proximally strongly narrowed, apically twisted 10—20°; upper man- dibular tooth subtriangular, 1.4~1.7 times as long as the lower tooth; outer mandibular surface sparsely punctate, concave, this concave area often slightly coriaceous. Labrum 0.3 times as long as broad; malar space 0.2-0.3 times as long as basal mandibular width. Clypeus in profile slightly out-flared, margin blunt; clypeus in front view 1.5—1.9 times as broad as long with apical margin truncate to slightly concave (Fig. 179). Lower face 0.70-0.75 times as broad as long, polished, with sparse fine punctures. Head in dorsal view with genae constricted behind the eyes; posterior ocellus contiguous with eye; FI = 55-60%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6-0.7 times the basal mandibular width away from mandible. Antenna moderately long and slender, with 49-53 flagellar segments; 20th segment 1.8-2.0 times as long as broad. Mesoscutum in profile abruptly rounded; notauli absent. Mesopleuron polished, the upper part with rather coarse but shallow punctures, the lower part punctate or punctostriate; epicnemial carina curved towards but generally not reaching anterior margin of pleuron. Scutellum in profile weakly convex, strongly laterally carinate for all of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, polished. Metapleuron centrally punctate tending to punctostriate peripherally; submetapleural carina evenly broadened anteriorly; posterior transverse carina of mesosternum centrally broadly incomplete, at most represented by a weak wrinkle. Propodeum in profile abruptly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area steep, weakly coriaceous; spiracular area moderately long, smooth; posterior area finely coriaceous; lateral longitudinal carina complete, not joined to spiracular margin by a short carina. Fore wing length 10-12 mm; discosubmarginal cell as in Fig. 272; AI = 1.35-2.07; CI = 0.20-0.25; ICI = 0.25-0.35; SDI = 1.10—1.15; cu-a slightly proximal to Rs&M; marginal cell proximally very sparsely pubescent; Ist subdiscal cell with scattered hairs distally. Hind wing with 5-6 hamuli on R1; Ist abscissa of Rs straight, 2nd abscissa weakly arcuate. Fore leg with tibia subcylindrical, with scattered spines on outer surface. Mid leg with longer tibial spur 1.7 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 1.9-2.1 times as long as broad; claws of female quite long, apically evenly curved, with short close pectinae; claws of male similar, but with pectinae finer. Gaster slender; tergite 2 in profile 4-5 times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by about 4 times its own length. Ovipositor moderately slender, its sheath moderately slender. Male with sternites 7-9 bearing long, sparse, erect pubescence amid short, decumbent pubescence; gonosquama obliquely truncate. Colour generally pale yellow, mesoscutum with three dark longitudinal vittae and mesoscutum with a weak to strongly developed brownish mark along upper hind margin; tergites 5+ of gaster infuscate; interocellar area usually yellow, rarely with infuscation between hind ocelli; antennae golden; pterostigma orange; wings hyaline. VaRIATION. Morphologically this is a very uniform species. REMARKS. This species is named in honour of James Karr in recognition of his contribution to the understanding of rain forest bird biology. Enicospilus karrensis is easily distinguishable by the incomplete posterior transverse mesosternal carina, by the form of the clypeus and mandibles, and the pattern of the alar sclerite. The form of the mouthparts of this species bears some superficial similarity to the Old World species E. sesamiae Delobel and E. sakaguchii (Matsumura & Uchida). However, the Old World species have the clypeal margin thin and the mandibles diagonally grooved suggesting there is no real phylogenetic affinity between these three species. The concave clypeus and short mandibles of E. karrensis are probably a parallel adaptation for dealing with a similar type of host. The Old World species parasitize grass-feeding noctuids (Moutia, 1934; Nagatomi, 1972), but the host of E. karrensis is not known. 210 IAN D. GAULD BIOLOGICAL INFORMATION. Enicospilus karrensis has only been collected in lowland forest in Central Panama, but at this locality is is one of the commoner species of Enicospilus in light-trap samples. Its pooled seasonal distributional data (1983-5) on Barro Colorado Island are: Joo Ee vie Awe Mee eAS Se OO: INGeD = =| Oe Ge Sao alee | SA eS Its host is unknown. MATERIAL EXAMINED Holotype CO’, Panama: Barro Colorado Island, 120 m, vi.1984 (Wolda) (BMNH). Panama: 2’, 4 9, Barro Colorado Island, Gatun Lake, iv.1979, xi. 1982, iv-v.1983 (Wolda) (TC); 100’, 14 @, Barro Colorado Island, 120 m, v-ix, xi. 1983-85 (Wolda) (BMNH; CNC). Enicospilus stevensi sp. n. (Figs 111, 180, 273) DescriPTIon. Mandibles moderately long, fairly evenly narrowed, apically twisted about 80°; upper mandibular tooth subcylindrical, 1.4-1.6 times as long as the slightly compressed and slightly broader lower tooth (Fig. 180); outer mandibular surface flat, with a weak proximal concavity. Labrum 0.2 times as long as broad; malar space 0.2 times as long as basal mandibular width. Clypeus in profile flat, margin rather blunt; clypeus in front view 1.4—1.5 times as broad as long, with its margin apically truncate. Lower face 0.70—0.75 times as broad as long, finely punctate. Head in dorsal view with genae rounded behind the eyes; posterior ocellus contiguous with eye; FI = 70%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.5—0.8 times the basal mandibular width away from mandible. Antenna long, relatively slender, with 61-63 flagellar segments; 20th segment 2.1—2.2 times as long as broad. Mesoscutum polished, without obvious punctures, in profile evenly rounded; notauli weak. Meso- pleuron polished, the upper part weakly and sparsely punctate, ventrally becoming more closely punctate or punctostriate, and with the lower part punctostriate to striate; epicnemial carina curved towards anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for most of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, punctate, posteriorly wrinkled. Metapleuron very convex, rugose-striate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete or narrowly interrupted centrally. Propodeum in profile abruptly declivous, dorsally very strongly flattened; anterior transverse carina complete, posterior trans- verse carina represented laterally by a pair of anteriorly directed ridges; anterior area steep, quite long, striate; spiracular area moderately long, finely punctate; posterior area coarsely irregularly reticulate; lateral longitudinal carina present on anterior 0.6, joined to spiracular margin by a short carina. Fore wing length 19-22 mm; discosubmarginal cell as in Fig. 273; AI = 1.00-1.05; CI = 0.19-0.34; ICI = 0.56-0.58; SDI = 1.05—1.09; cu-a proximal to base of Rs&M by 0.40.7 times its own length; marginal cell proximally with a small glabrous area adjacent to Rs+2r opposite proximal sclerite; 1st subdiscal cell with anterior and distal margins hirsute. Hind wing with 8-10 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa very weakly arcuate. Fore leg with tibia subcylindrical, with scattered spines on outer surface. Mid leg with longer tibial spur 1.4-1.5 times length of the shorter. Hind leg with coxa in profile 1.7 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.2—2.3 times as long as broad; claws of female stout, abruptly curved, bearing long, close pectinae; those of male similar but with pectinae finer. Gaster long and slender; tergite 2 in profile 6 times as long as posteriorly deep, laterotergite turned under, thyridia obovate and separated from anterior margin of tergite by about 4 times its own length. Ovipositor slender, its sheath exceptionally broad (Fig. 111). Male with sternites 7-9 bearing long erect hairs; gonosquama distally bluntly rounded. Colour generally brownish orange; interocellar area yellow; antennae blackish; mesoscutal vittae blackish. Gaster paler yellowish brown, with tergites 5+ blackish and often with posterior margin of tergite 2 slightly infuscate; pterostigma orange; wings almost hyaline. VARIATION. The specimens from Bolivia are very similar to the Costa Rican specimens but have the meso- and metapleurae more regularly punctate. REMARKS. This species is named in honour of George Stevens for his tireless activity on behalf of dry forest research in Santa Rosa National Park. OPHIONINAE OF TROPICAL MESOAMERICA 211 Enicospilus stevensi can be recognized by the following combination of features: absence of any central sclerite; having a strongly twisted mandible; possessing an exceptionally convex metapleuron; having the very broad ovipositor sheath. The relatively small value of ICI is unusual in a species of this size and the colour pattern of the gaster is also quite unusual and characteristic. In the venation, form of the mandible, sculpture and shape of the metapleuron E. stevensiis very similar to E. pamelae. Females of the two species can easily be distinguished by the shape of the ovipositor sheath (that of E. pamelae is slender). The male of E. pamelae is unknown so I cannot yet give features for distinguishing it from males of E. stevensi. BIOLOGICAL INFORMATION. Enicospilus stevensi is a very widely distributed Neotropical species whose range extends from Chiapas in southern Mexico, south to Bolivia. However, it is relatively rarely captured, and despite extensive collecting in Costa Rica only three individuals have been caught. Two of these were taken in Santa Rosa National Park during the wet season. Nothing is known of the hosts of this species. MATERIAL EXAMINED Holotype ©’, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, viii.1984 (Janzen & Hallwachs) (BMNH). Paratypes. Bolivia: 1 2, Chapare, El Limbo, 2200 m, i.1962 (BMNH); 1 9, Santa Cruz, Ingenio la Belgica, 38 km N. Santa Cruz, 1.1980, (Stange) (FSCA). Costa Rica: Guanacaste Prov.: 1 o’, Santa Rosa National Park, 300 m, vi.1980 (Janzen & Hallwachs) (BMNH): San José Prov.: 1 9, Estacion Carrillo, Braulio Carrillo National Park, 700 m, iv.1985 (Chacon) (BMNH). Mexico: Chiapas: 1 0’, 30-40 km N. Huixtla, 1000 m, vi.1969 (Peterson) (CNC). Enicospilus pamelae sp. n. (Figs 110, 274) Description. Mandibles of moderate length, evenly narrowed, apically twisted 80°; upper mandibular tooth compressed, very acute, 1.4-1.7 times as long as the lower tooth; outer mandibular surface sparsely pubescent, flat with a broad, shallow proximal concavity. Labrum 0.1-—0.2 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile flat, margin subacute; clypeus in front view 1.2—1.4 times as broad as long, its apex virtually truncate. Lower face 0.63-0.70 times as broad as long, centrally with very sparse fine punctures. Head in dorsal view with genae evenly constricted behind eye; posterior ocellus contiguous with eye; FI = 65-70%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.8—0.9 times the basal mandibular width away from mandible. Antenna long and slender, with 59-61 flagellar segments; 20th segment 1.9-2.1 times as long as broad. Mesoscutum polished and sparsely punctate, in profile evenly rounded; notauli vestigial. Mesopleuron polished, the upper part finely punctate, the lower part punctate with weak longitudinal wrinkling; epicnemial carina curved towards anterior margin of pleuron, its upper end evanescent. Scutellum in profile moderately convex, laterally carinate for all of its length; scutellum in dorsal view 1.6 times as long as anteriorly broad, smooth with scattered punctures, and posteriorly with a few weak wrinkles. Meta- pleuron strongly convex, with diagonal wrinkling anterodorsally; submetapleural carina weakly broad- ened anteriorly; posterior transverse carina of mesosternum complete. Propodeum in profile quite abruptly declivous, flattened; anterior transverse carina complete, posterior one present as a lateral vestige that forms a sharp crest laterally on the posterior area; anterior area deeply excavate, sparsely striate; spiracular area short, smooth; posterior area coarsely irregularly reticulate; lateral longitudinal carina generally complete, joined to spiracular margin by a short carina. Fore wing length 14-16 mm; discosubmarginal cell as in Fig. 274; AI = 0.73-0.93; CI = 0.30-0.33; ICI = 0.41-0.68; SDI = 1.10—-1.19; cu-a proximal to the base of Rs&M by 0.1-0.3 times its own length; marginal cell proximally slightly more sparsely hirsute proximally; 1st subdiscal cell with distal part sparsely hirsute. Hind wing with 6-8 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa weakly bowed. Fore leg with tibia subcylindrical, with fine scattered spines on outer surface. Mid leg with longer tibial spur 1.4-1.5 times length of the shorter. Hind leg with coxa in profile 1.9-2.0 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.5 times as long as broad; claws of female large, strongly curved and with short close pectinae. Gaster slender; tergite 2 in profile more than 5 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 4 times its own length. Ovipositor slender, its sheath narrow (Fig. 110). Male unknown. Colour generally pale yellowish, with legs golden and anterior gastral segments orange; tergite 3+ infuscate; mesoscutum anteriorly slightly darker; interocellar area pale yellow; antenna proximally black, distally brown; pterostigma yellow brown; wings weakly infumate. 212 IAN D. GAULD VaRIATION. The smaller specimens have weaker striation on the thorax laterally and only have the scape blackish. In the Townes’ collection is a female from Bolivia which resembles this species except that it is altogether paler. The propodeal sculpture of this specimen is finer and it is not clear whether or not it is truly conspecific so I have excluded it from the paratype series. REMARKS. This species is named in honour of Pamela Mitchell, as a gesture of thanks for all her help in sorting 36 Malaise trap-years of samples. Enicospilus pamelae resembles E. stevensi in the form of the mandibles, alar sclerites, venation and form of the metapleuron, but differs in several features emphasized in the key. E. stevensi is altogether larger and more robust; it has the area of the fore wing anterior to the proximal sclerite fairly evenly hirsute so there is only a narrow anterior extension of the fenestra, whilst E. pamelae has a very broad anterior fenestral extension. BIOLOGICAL INFORMATION. In Mesoamerica Enicospilus pamelae is only known from lowland rainforest on Barro Colorado Island where occasional specimens have been collected between May and July. Nothing is known of its hosts. MATERIAL EXAMINED Holotype 2, Panama: Barro Colorado Island, 120 m, v.1985 (Wolda) (BMNH). Paratypes. Panama: 4 9, Barro Colorado Island, 120 m, v-vii.1984—5 (Wolda) (BMNH). Non-paratypic material. Bolivia: 1 9 , Santa Cruz, Buena Vista, iv.1950 (Peria) (TC). Enicospilus vilmari sp. n. (Fig. 276) Description. Mandibles quite short, strongly narrowed and with a ventrobasal lobe, apically twisted 30°; upper mandibular tooth depressed, 1.4 times as long as the lower; outer mandibular surface flat, sparsely pubescent. Labrum 0.2 times as long as broad; malar space 0.4 times as long as basal mandibular width. Clypeus in profile very weakly convex, margin blunt; clypeus in front view 1.3 times as broad as long, its margin weakly convex. Lower face 0.7 times as broad as long, polished, with shallow punctostriations centrally. Head in dorsal view with genae abruptly constricted; posterior ocellus close to eye; FI = 65%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 1.0 times the basal mandibular width away from mandible. Antenna long and slender, with 56 flagellar segments; 20th segment 2.5 times as long as broad. Mesoscutum polished, sparsely but distinctly punctate, in profile evenly rounded; notauli vestigial. Mesopleuron polished, the upper part punctate, the lower part punctate but centrally tending to punc- tostriate; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for all of its length; scutellum in dorsal view 1.5 times as long as anteriorly broad, anteriorly smooth, posteriorly with a few striae. Metapleuron moderately convex, diagonally striate; submetapleural carina strongly anteriorly broadened; posterior transverse carina of mesosternum com- plete. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area deeply excavate, striate; spiracular area short, smooth; posterior area reticul- ate laterally, centrally tending to concentrically rugose; lateral longitudinal carina present anteriorly, joined to spiracular margin by a short carina. Fore wing length 14mm; discosubmarginal cell as in Fig. 276; AI = 1.17; CI = 0.23; ICI = 0.44SDI = 1.09; cu-a proximal to the base of Rs&M by about 0.1 times its own length; marginal cell proximally sparsely pubescent; 1st subdiscal cell with anterior and distal parts sparsely hirsute. Hind wing with 7 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa weakly curved. Fore leg with tibia slightly flattened, with long slender spines on outer surface. Mid leg with longer tibial spur 1.7 times length of the shorter. Hind leg with coxa in profile 1.9 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.5 times as long as broad; claws of female long, apically evenly curved with long stout pectinae. Gaster long and slender; tergite 2 in profile 7 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 4 times its own length. Ovipositor laterally compressed, rather stout, its sheath moderately broad. Male unknown. Colour generally yellowish, mesoscutal vittae blackish and tergites 3+ of gaster infuscate; interocellar area yellow; antenna black; pterostigma brown; wings hyaline. REMARKS. This species is named in honour of Vilmar Rodriguez in recognition of his tireless efforts at protecting Cerro el Hacha, Guanacaste National Park, from fire. OPHIONINAE OF TROPICAL MESOAMERICA 213 Enicospilus vilmari may easily be distinguished from all other Mesoamerican species by the form of the proximal alar sclerite (Fig. 276) which is rather elongately tapered distally. BIOLOGICAL INFORMATION. Enicospilus vilmari is only known to occur in north-western Costa Rica where a single individual has been collected in dry forest during the dry season. It is not known what hosts this species parasitizes. MATERIAL EXAMINED Holotype 9, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, ii.1982 (Janzen & Hallwachs) (BMNH). Enicospilus estradarum sp. n. (Fig. 277) DEscriPTION. Mandibles moderately long, evenly narrowed, very weakly curved, apically twisted 20°; upper mandibular tooth slender, 1.5 times as long as the lower tooth; outer mandibular surface sparsely pubescent, distally flat, but with a weak proximal concavity. Labrum 0.3 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.3—1.4 times as broad as long, the margin weakly convex. Lower face 0.73-0.76 times as broad as long, polished, sparsely punctate. Head in dorsal view with genae evenly narrowed; posterior ocellus contiguous with eye; FI = 62%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6 times the basal mandibular width away from mandible. Antenna long and slender, with 56 flagellar segments; 20th segment 2.4 times as long as broad. Mesoscutum polished, finely punctate, in profile evenly rounded; notauli vestigial. Mesopleuron pol- ished, the upper and lower parts finely punctate; epicnemial carina curved towards anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for 0.6 of its length; scutellum in dorsal view 1.6 times as long as anteriorly broad, smooth, posteriorly wrinkled. Metapleuron convex punctate; submetapleural carina weakly broadened anteriorly; posterior transverse carina of mesosternum com- plete. Propodeum in profile evenly rounded; anterior transverse carina complete, posterior transverse carina represented laterally by a weak to vestigial crest; anterior area quite long, striate; spiracular area finely punctate; posterior area reticulate; lateral longitudinal carina present on anterior 0.4, joined to spiracular margin by a short, very weak carina. Fore wing length 16-17 mm; discosubmarginal cell as in Fig. 277; AI = 0.60-0.80; CI = 0.30-0.32; ICI = 0.48-0.55; SDI = 1.11-1.16; cu-a slightly proximal to the base of Rs&M; marginal cell proximally more or less evenly hirsute; 1st subdiscal cell with anterior and distal periphery hirsute. Hind wing with 7 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa weakly curved. Fore leg with tibia weakly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.2-1.3 times length of the shorter. Hind leg with coxa in profile 1.7 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus missing; claws of male missing. Gaster slender; tergite 2 in profile 6 times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by about 3.5 times its own length. Female unknown. Male with sternites 7-9 bearing long, dense, erect pubescence; gonosquama evenly rounded. Colour generally yellowish orange; interocellar area yellow; antennae blackish; terminal segments of gaster slightly infuscate; pterostigma orange; wings hyaline. VARIATION. None remarkable. REMARKS. This species is named in honour of Rosy and Alejandro Estrada for their efforts in biology at the Los Tuxtlas Biology Station, Mexico. The mandibles of Enicospilus estradarum are rather different from those of any other species that lacks a central sclerite in that they are slightly broader and angled downwards. This difference is rather difficult to appreciate unless reference can be made to several species. E. estradarum is most easily distinguished from other Mesoamerican species of Enicospilus by the presence of a band of hairs in the discosubmarginal cell close to the base of Rs+2r; these hairs are bounded proximally by an extension of the fenestra (Fig. 277). BIOLOGICAL INFORMATION. Enicospilus estradarum is only known to occur in Costa Rica where it has been collected in the seasonally dry forests in Santa Rosa National Park. Despite extensive collecting only two specimens have been taken, in June and July, at the start of the wet season. Nothing is known of the hosts of this species. 214 IAN D. GAULD MATERIAL EXAMINED Holotype ©’, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, vii.1980 (Janzen & Hallwachs) (TC). Paratype. Costa Rica: Guanacaste Prov.: 1 ©’, Santa Rosa National Park, 300 m, vi.1982 (Janzen & Hallwachs) (BMNH). Enicospilus teodorae sp. n. (Figs 116, 279) DescripeTion. Mandibles quite long, proximally weakly narrowed, distally almost parallel-sided and apically twisted 10°; upper mandibular tooth compressed, 0.8—0.9 times as long as the lower tooth; outer mandibular surface almost flat, bearing long scattered hairs. Labrum 0.3—0.4 times as long as broad; malar space 0.3-0.4 times as long as basal mandibular width. Clypeus in profile nasute, margin blunt (Fig. 116); clypeus in front view 1.1—1.2 times as broad as long, with margin slightly V-shaped. Lower face 0.6—0.7 times as broad as long, polished, obsoletely punctostriate. Head in dorsal view with genae strongly constricted behind eyes; posterior ocellus very close to eye; FI = 60-65%; occipital carina mediodorsally complete, ventrally joining hypostomal carina about 0.8 times the basal mandibular width away from mandible. Antenna exceedingly long and slender, with 60-65 flagellar segments; 20th segment 2.9-3.0 times as long as broad. Mesoscutum weakly polished, with scattered weak punctures, in profile evenly rounded; notauli absent. Mesopleuron polished, the upper part centrally smooth, the lower part punctostriate to striate; epicnemial carina evenly curved towards, but usually not quite reaching anterior margin of pleuron. Scutellum in profile evenly rounded, laterally carinate for all of its length; scutellum in dorsal view 1.2—1.3 times as long as anteriorly broad, anteriorly finely punctate, posteriorly striate. Metapleuron moderately convex, with obsolescent fine scattered punctures, generaliy rather smooth and shining; submetapleural carina narrow, barely broadened anteriorly; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area weakly impressed, centrally striate; spiracular area long, smooth; posterior area transversely striate to irregularly wrinkled; lateral longitudinal carina complete, joined to spiracular margin by a short carina, or sometimes with this carina discontinuous. Fore wing length 12-13 mm; discosubmarginal cell as in Fig. 279; AI = 1.43-2.05; CI = 0.17-0.29; ICI = 0.34-0.40; SDI = 1.12-1.19; cu-a opposite or very slightly distal to the base of Rs&M; marginal cell proximally broadly glabrous, or with isolated hairs; 1st subdiscal cell rather sparsely hirsute, except near proximal end which is glabrous. Hind wing with 5—6 hamuli on R1; Ist abscissa of Rs straight, 2nd abscissa very slightly arcuate. Fore leg with tibia weakly flattened, without spines on outer surface. Mid leg with longer tibial spur 1.6 times length of the shorter. Hind leg with coxa in profile 1.9-2.0 times as long as deep; trochantellus dorsally 0.4—0.5 times as long as broad; 4th segment of tarsus 2.3—2.4 times as long as broad; claws of female symmetrical, long, with short, fairly closely spaced pectinae; claws of male similar but wiih pectinae slightly shorter. Gaster slender; tergite 2 in profile more than 7 times as long as posteriorly deep, laterotergite up-turned, thyridia elliptical and separated from anterior margin of tergite by about 5.0 times its own length. Ovipositor apically quite slender, its sheath narrow. Male with sternites 7-9 bearing fine semierect pubescence that is longer, closer and more erect mediolongitudinally; gonosquama evenly rounded. Colour generally pale yellow; interocellar area and three mesoscutal vittae black; antennae brownish, distally infuscate, scape often blackish; mesosternum dark brown; legs golden; terminal segments of gaster slightly infuscate, tergite 3 very pale; pterostigma brownish yellow; wings hyaline. VARIATION. The holotype female from Panama has the alitrunk extensively black-marked and the gaster uniformly brown. Structurally it is otherwise very like the paratypes. REMARKS. This species is named in honour of Teodora Rodriguez for her courageous stand against threats to Cerro el Hacha, Guanacaste National Park. Enicospilus teodorae is easily recognized by the peculiar form of the clypeus. The axis of the mandible is twisted slightly forwards so the mandibles open outwards as well as downwards. The function of these modifications to the mouthparts is not known. The phylogenetic relationships of this species are not clear even though the clypeus of E. teodorae resembles that of E. bima (see that species). As these two taxa do not share any other morphological specializations I do not think they are closely related. BIOLOGICAL INFORMATION. Enicospilus teodorae is a widely distributed, but rarely collected species, whose range extends from Central Panama south throughout lowland South America to the tropic of Capricorn. OPHIONINAE OF TROPICAL MESOAMERICA 215 The extremely elongate antennae suggest that this species may fly in the canopy of forests. Its hosts are unknown. MATERIAL EXAMINED Holotype Y, Panama: Las Cumbres, iv-v.1982 (Wolda) (TC). Paratypes. Brazil: 1 ©’, Goias, Jatai, xi.1971 (Oliveira) (TC); 1 CO, Rio de Janeiro, Mangaratiba, iii.1972 (Alvarenga) (BMNH; TC). Surinam: 1 ©’, Paramaribo, Lelydorp, iii. 1964 (Geijskes) (TC). Enicospilus maritzai sp. n. (Figs 112, 216, 280) Description. Mandibles moderately long, evenly narrowed, apically twisted 70-80°; upper mandibular tooth slightly compressed, 1.4-1.8 times as long as the lower tooth which is rather more strongly com- pressed; outer mandibular surface sparsely pubescent, more or less flat. Labrum 0.3-0.4 times as long as broad; malar space 0.2-0.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.25—1.45 times as broad as long, its apical margin virtually truncate. Lower face 0.55—0.65 times as broad as long, submatt, finely but closely punctate, centrally tending to punctostriate. Head in dorsal view with genae constricted behind the eyes; posterior ocellus contiguous with eye; FI = 80-90% ; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.7 times the basal mandibular width away from mandible. Antenna long but relatively stout, with 63-65 flagellar segments; 20th segment 1.4~1.5 times as long as broad. Mesoscutum weakly polished, with fine punctures, in profile fairly abruptly rounded; notauli quite long but very weakly impressed. Mesopleuron polished, the upper part finely punctate, the lower part punc- tostriate; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for all of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, polished, smooth with wrinkles posteriorly. Metapleuron strongly convex, finely punctate, posterodorsally coarsely strigose to reticulate; submetapleural carina anteriorly strongly broadened; posterior transverse carina of mesosternum broadly incomplete centrally. Propodeum in profile abruptly declivous; anterior transverse carina complete, posterior transverse carina strong, stretching from near hind coxa forwards to near centre of anterior transverse carina; anterior area strongly impressed, striate; spiracular area quite short, finely punctogranulate; posterior area usually centrally irregularly rugose to reticulate, laterally strigose-reticulate; lateral longitudinal carina present only as a vestige anteriorly, joined to spiracular margin by a short carina. Fore wing length 20-23 mm; discosubmarginal cell as in Fig. 280; AI = 0.65-0.92; CI = 0.25-0.30; ICI = 0.78-0.88; SDI = 1.14-1.27; cu-a slightly proximal to the base of Rs&M; marginal cell proximally evenly hirsute; 1st subdiscal cell with anterior 0.5 hirsute. Hind wing with 9-10 hamuli on R1; 1st abscissa of Rs more or less straight, 2nd abscissa weakly arcuate. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.1—1.3 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.2—2.3 times as long as broad; claws of female large and apically abruptly curved, with long fine pectinae, the central ones more widely interspaced than the distal or proximal ones (Fig. 216); claws of male similar but with pectinae shorter. Gaster quite stout (Fig. 112); tergite 2 in profile 2.8-3.5 times as long as posteriorly deep, laterotergite folded under, thyridia subcircular to oval and separated from anterior margin of tergite by at least 3 times its own length. Ovipositor quite stout, its sheath slightly broadened. Males with sternites 7-9 bearing a dense covering of long stout erect hairs; gonosquama evenly rounded apically. Colour generally brownish orange with tergites 3+ blackish; mesoscutum usually uniformly reddish brown, sometimes with a medial anterior black mark; interocellar area and antenna blackish; pterostigma brown; wings weakly infumate. VARIATION. Most specimens have tergites 3+ of the gaster uniformly black, but some Brazilian specimens have a small pale mark laterally on tergite 3. There is considerable variation in the sculpture of the posterior area of the propodeum. The posterior transverse carina is always strong laterally, but the area between may be almost smooth in a few individuals, though it is most usually reticulate or irregularly rugose. The Peruvian specimen has a well-developed median longitudinal ridge. REMARKS. Enicospilus maritzai together with E. exoticus and E. jesicae form a closely knit species complex. The three are characterized by the several apomorphic features. They are the only species with a black interocellar area, central sclerite and a large ICI. All have a dorsally flattened propodeum with the 216 IAN D. GAULD posterior transverse carina represented laterally by at least two weak crests. Tergites 3+ of the gaster are generally black, though there is a tendency for pale markings to be present laterally on tergite 3 and sometimes 4. All species have the posterior transverse carina of the mesosternum broadly interrupted centrally, and lack most of the lateral carina of the propodeum. E. maritzai differs from E. exoticus in possessing distinct alar sclerites and having more twisted mandibles. It is also larger, has stouter antennae and generally has a more coarsely sculptured propodeum than either EF. exoticus or E. jesicae. The arrangement of pectinae on the tarsal claw of the female of E. maritzai (Fig. 216) is quite unlike that of either of the other two species (see Fig. 218). BIOLOGICAL INFORMATION. Enicospilus maritzai is a widely distributed species whose range extends from northern Costa Rica (ca 11°N) south throughout South America to about 18°S in Bolivia. It is rather uncommon in Costa Rica. Only five specimens have been collected in Santa Rosa National Park, a male and a female in May at the start of the wet season, and two females and a male in December at the end of the wet season. Elsewhere in the country two specimens have been collected at light in cloud forest in Monteverde Reserve during February and March and a single female during May in forest on Volcan Orosi. Nothing is known of the host range of this species. MATERIAL EXAMINED Holotype @, Costa Rica: Guanacaste Prov., Volcan Orosi, Casa Mariksa (= Maritza), 7-800 m, v.1986 (Gauld) (BMNH). Paratypes. Bolivia: 1 2, Cochabamba, Alto Palmar, 1100 m, ix.1960 (Walz) (TC). Brazil: 16 9, Bahia, Encruzilhada, 960 m, xi.1972 & xi.1974 (Alvarenga) (TC). Costa Rica: Guanacaste Prov.: 1 0’, 2 9, Santa Rosa National Park, 300 m, xii.1979 (Janzen) (TC); 10’, 1 9, same locality, v.1980 (Janzen & Hallwachs) (BMNH): Puntarenas Prov.: 1 &’, Monteverde, iii.1961 (Palmer) (TC); 1 9, Monteverde, ii.1986 (For- syth) (CNC). Peru: 1 2, Cusco, Santa Isabel, Rio Ceosnipata, xii.1951 (Woytkowski) (TC). Enicospilus exoticus (Morley) (Figs 92, 107, 218, 221, 281) Ophiomorpha bicolor Szépligeti, 1905: 35. Holotype 2, BRAZIL (TM) [examined]. [Junior secondary homonym of Enicospilus bicolor (Taschenberg, 1875).] Henicospilus exoticus Morley, 1912: 36. Holotype 2, BRAZIL (BMNH) [examined]. Syn. n. Enicospilus dichromus Townes & Townes, 1966: 176. [Replacement name for bicolor Szépligeti.] Enicospilus exoticus (Morley) Townes & Townes, 1966: 177. Description. Mandibles moderately long, quite strongly but evenly narrowed, apically twisted 50-60°; upper mandibular tooth subcylindrical, 1.2-1.4 times as long as the lower tooth which is slightly com- pressed; outer mandibular surface sparsely pubescent, distally almost flat, proximally with a weak concave area. Labrum 0.3 times as long as broad; malar space 0.2-0.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.5—1.6 times as broad as long, with apical margin weakly convex. Lower face 0.60.7 times as broad as long, weakly polished, finely punctate. Head in dorsal view with genae quite strongly constricted behind eyes; posterior ocellus contiguous with eye; FI = 75-80%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.7 times the basal mandibular width away from mandible. Antenna moderately long, stout, with 57-59 flagellar segments; 20th segment 1.9-2.0 times as long as broad. Mesoscutum weakly polished with very fine scattered punctures, in profile evenly rounded; notauli vestigial. Mesopleuron polished, the upper part finely punctate, the lower part similar but more closely punctate, sometimes tending to punctostriate anteriorly; lateral part of epicnemial carina straight, its upper end evanescent, abruptly curved towards anterior margin of pleuron. Scutellum in profile evenly rounded, laterally carinate for all of its length; scutellum in dorsal view 1.6—1.7 times as long as anteriorly broad, very sparsely punctate, posteriorly wrinkled. Metapleuron convex, with coarse striae radiating from propodeum, grading anteroventrally to punctate; submetapleural carina strongly broadened ante- riorly; posterior transverse carina of mesosternum usually broadly incomplete centrally, rarely narrowly incomplete but then with carina very weak. Propodeum in profile rather short, abruptly declivous; anterior transverse carina strong, most usually strongly raised centrally and less so laterally; posterior transverse carina represented by blunt lateral tubercles; anterior area short, strongly impressed, from almost smooth to coarsely striate; spiracular area short, smooth; posterior area rather flattened dorsally, centrally with one or sometimes more longitudinal ridges, laterally coarsely transversely striate, the striae often extend- ing on to the metapleuron; lateral longitudinal carina present only as a vestige anteriorly, joined to spiracular margin by a short weak carina. OPHIONINAE OF TROPICAL MESOAMERICA 217 Fore wing length 15—17 mm; discosubmarginal cell as in Figs 221, 281; AI = 0.69-0.83; CI = 0.29-0.40; ICI = 1.00-1.31; SDI = 1.08-1.23; cu-a proximal to base of Rs&M by about 0.2 times its own length; marginal cell proximally evenly hirsute; 1st subdiscal cell with anterior 0.6 and distal corner hirsute. Hind wing with 9-10 hamuli on R1, the distal three or four a little shorter than the proximal ones; Ist abscissa of Rs very weakly curved, 2nd abscissa arcuate. Fore leg with tibia barely flattened, with about 10 scattered spines on outer surface. Mid leg with longer tibial spur 1.5—1.7 times length of the shorter. Hind leg with coxa in profile 1.7 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.42.5 times as long as broad; claws of female long, distally evenly curved, rather evenly pectinate (Fig. 218); claws of male similar. Gaster quite stout; tergite 2 in profile about 2.5 times as long as posteriorly deep, laterotergite folded under, thyridia large, oval and separated from anterior margin of tergite by about 3 times its own length; tergite 7 unusual, in that in profile it is strongly concave posteriorly (Fig. 92). Ovipositor proximally stout, distally slender and slightly decurved, its sheath quite broad. Male with sternites 7-9 bearing dense thick erect pubescence; gonosquama posteriorly truncate. Colour generally yellowish orange; mesoscutum with three dark longitudinal vittae; interocellar area usually black; antennae, especially basally, infuscate; gaster orange-brown, usually with tergites 3+ blackish brown and unusual in having a pair of lateral spots on tergite 4 and sometimes 3 pale yellow; pterostigma dark brown; wings weakly infumate. VaRIATION. The colour of the gaster is quite variable in this species. Some specimens have only small pale spots on tergite 4, in others these areas are large and confluent with similar areas on tergite 3. They are not confluent dorsally as there is always a dark dorsal stripe. Many of the Panamanian males have the gaster more or less uniformly pale yellowish orange, but females are darker. Such sexual differences in coloration have not been observed in other populations. One of the Costa Rican specimens examined differs from the remainder in having a slightly granulate metapleuron with barely any striae present, and in having the submetapleural carina barely expanded anteriorly. There is some variation in the degree of sclerotization of the alar sclerites; they are always more weakly sclerotized in this species than they are in related taxa, but those of some individuals may be extremely weak and possibly ignored. For this reason E. exoticus has been taken out in the key in several places. This allows for the central sclerite or all sclerites being overlooked. REMARKS. Enicospilus exoticus shares a number of apomorphic features with E. maritzai and E. jesicae, suggesting the three species are closely inter-related (see discussion on E. maritzai). It can easily be distinguished from the other two species as it has very weak alar sclerites (Fig. 281). BIOLOGICAL INFORMATION. Enicospilus exoticus is a quite widely distributed species whose range extends from about 10°N in Costa Rica, south to about 15°S in Peru and Brazil (Map 20). Most specimens have been collected between sea-level and 1000 m. The Costa Rican specimens were collected at light in a forest in Braulio Carrillo National Park during February and April. No others are known from this far north. In Panama in forest on Barro Colorado island E. exoticus has been collected only rather rarely. The cumulative seasonal data for 1982-5 are: Je EM AS Me J a tALeS= (O. Ne iD Se eS ee a ee a ee ee Nothing is known of the hosts of this species. E. exoticus may be mimicing certain aculeates. When the gaster is viewed dorsally the pale patches on tergites 3 and 4 ‘disappear’ if the insect is on a pale background, and the only parts of the gaster that are visible are the darker anterior and posterior tergites together with the conspicuous dark dorsal stripe on tergites 3-4. Thus, in dorsal aspect the gaster appears to be elongately petiolate, and resembles the gasters of several common and ferocious vespids. MATERIAL EXAMINED Holotype 2 (Henicospilus exoticus Morley) Brazil: Para (Bates) (BMNH). Holotype 2 (Ophiomorpha bicolor Szépligeti) Brazil: Minas Gerais (TM). Brazil: 1 2, Aguas Vermethas, 800 m, xii.1983 (Alvarenga) (TC); 4.0’, 11 9, Bahia, Encruzilhada, 960- 980 m, xi.1972 & xi.1974 (Alvarenga) (TC); 1 ©’, Jacareacanga, vii.1970 (Barbosa) (TC); 1 2, Minas Gerais, Pedra Azul, 800 m, xi.1972 (Alvarenga & Seabra) (TC). Colombia: 1 9 , Magdalena, 800 m, iv.1973 (Helava) (BMNH). Costa Rica: San José Prov.: 1 &’, Estacion Carrillo, Braulio Carrillo National Park, 700 m, ii.1985 (Chacon & Chacon) (BMNH); 1 9, Braulio Carrillo National Park, 500 m, iv.1985 (Goulet & Masner) (CNC). Panama: 2 ©’, 1 2, Barro Colorado Island, iv, vi, xi.1978 (Wolda) (RNH); 1 9, same 218 IAN D. GAULD Map 20 Localities at which Enicospilus exoticus has been collected. locality, ix.1982 (Wolda) (TC); 1 ot, 4 9, same locality, vi.1983, ii, vi, vii & x.1985 (Wolda) (BMNH). Peru: 1 2, Quincemil, nr Macapata, 30 m, ix.1962 (Pera) (TC). Venezuela: 1 9 , Aragua, Rancho Grande, iv.1960 (Test) (UM); 1 9, Caracas, Colonia Tovar 16.5 km, 1700 m, x.1938 (TC). Enicospilus jesicae sp. n. (Fig. 282) DEscrIPTION. Mandibles of moderate length, strongly but evenly narrowed, apically twisted 70—85°; upper mandibular tooth slightly compressed, 1.41.6 times as long as the lower tooth; outer mandibular surface weakly concave. Labrum 0.3 times as long as broad; malar space 0.3 times as long as basal mandibular OPHIONINAE OF TROPICAL MESOAMERICA 219 width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.2—1.4 times as broad as long, its apical margin more or less truncate. Lower face 0.5—0.6 times as broad as long, polished, finely punctate. Head in dorsal view with genae evenly narrowed; posterior ocellus contiguous with eye; FI = 85-87% ; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.8 times the basal mandibular width away from mandible. Antenna quite long, moderately stout, with 57-60 flagellar segments; 20th segment 2.1—2.2 times as long as broad. Mesoscutum weakly polished, with fine scattered punctures, in profile evenly rounded; notauli vestigial. Mesopleuron polished, the upper part finely punctate, the lower part irregularly punctostriate; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for all of its length; scutellum in dorsal view 1.4-1.5 times as long as anteriorly broad, finely punctate, posteriorly wrinkled. Metapleuron moderately to strongly convex, punctostriate, sometimes posterodor- sally rugose; submetapleural carina quite strongly broadened anteriorly; posterior transverse carina of mesosternum from quite narrowly to broadly interrupted centrally. Propodeum in profile abruptly declivous; anterior transverse carina strong, often broadened slightly centrally; posterior transverse carina represented by weak to strong lateral tubercles; anterior area short, strongly impressed, striate; spiracular area short, smooth; posterior area generally with median longitudinal carina, laterally coarsely striate, the striae generally extending onto the metapleuron; lateral longitudinal carina present only as a vestige anteriorly, usually joined to spiracular margin by a short carina. Fore wing length 13-15 mm; discosubmarginal cell as in Fig. 282; AI = 0.63-0.91; CI = 0.10-0.20; ICI = 0.79-0.97; SDI = 1.16—1.19; cu-a proximal to base of Rs&M by 0.1-0.2 times its own length; marginal cell proximally slightly more sparsely hirsute than it is centrally; lst subdiscal cell with anterior and distal margins hirsute. Hind wing with 7 hamuli on R1, the distal 3 or 4 shorter than the others; 1st abscissa of Rs more or less straight, 2nd abscissa weakly arcuate. Fore leg with tibia barely flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.4-1.5 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.3—2.5 times as long as broad; claws of female quite long, evenly curved, with stout close pectinae; claws of male similar. Gaster moderately stout; tergite 2 in profile 34 times as long as posteriorly deep, laterotergite folded under, thyridia oval and separated from anterior margin of tergite by 2.5 or more times its own length. Ovipositor proximally stout, distally slender, with fairly conspicuous fine hairs, slightly decurved and with its sheath slightly broadened. Male with sternites 7-9 bearing dense long erect pubescence; gonosquama quite slender, elongately tapered. Colour generally yellowish brown, mesoscutum generally dark-marked; interocellar area usually black; antennae proximally black; gaster with tergites 1 and 2 reddish brown, 3+ infuscate; pterostigma brown; wings weakly infumate. VARIATION. There is considerable variation in the colour of the mesoscutum. Most specimens have three dark longitudinal stripes, but occasionally these appear to have coalesced so the mesoscutum is entirely black anteriorly. A few individuals have the mesoscutum brown. The specimens from Santa Rosa National Park include some of the darkest individuals to hand; frequently the black interocellar area extends down the frons onto the central part of the lower face. The two Panamanian specimens are rather pale and have well-developed pale areas on tergites 3 and 4. Some individuals have the interocellar area rather weakly infuscate and allowance has been made for them to run out in two places in the key. REMARKS. This species is named after Jesica Rodriguez, who will grow up to protect Cerro el Hacha, Guanacaste National Park. Enicospilus jesicae belongs to the E. maritzai species-complex (see discussion of that species). It strongly resembles E. exoticus in general structure, especially in the form of the propodeum, but differs in having much more strongly defined sclerites, more slender flagellar segments and (generally) lacking the pale mark on tergite 4. BIOLOGICAL INFORMATION. The range of Enicospilus jesicae extends further north than those of other species in the E. maritzai complex. E. jesicae is known to occur in southern Mexico, and from there south to Central Brazil and Ecuador. In Costa Rica isolated specimens have been collected in Santa Rosa National Park during the wet season between May and November. Isolated individuals have also been taken in cloud forest at Monteverde and on Volcan Cacao. Nothing is known about the biology of this species. MATERIAL EXAMINED Holotype 9, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, v.1986 (Gauld) (BMNH). 220 IAN D. GAULD Paratypes. Brazil: 1 9, Bahia, Encruzilhada, 980 m, xi.1974 (Alvarenga) (TC). Costa Rica: Guanacaste Prov.: 1 9, Cerro el Hacha, Casa Oeste, 400 m, x.1987 (Chacon) (BMNH); 1 o’, Estacion Mengo, SW. side of Volcan Cacao, 1100 m, vi.1987 (Janzen) (BMNH); 3 o’, 1 2, Santa Rosa National Park, 300 m, vii. 1980, xi.1982, x.1983 & v.1985 (Janzen & Hallwachs) (BMNH): Puntarenas Prov.: 1 2, Monteverde, 1350 m, xi.1985 (Haber) (BMNH); 1 9, same locality, ii.1986 (Forsyth) (CNC). Ecuador: 1 9, Playas de Montalvro, 15 m, iii.1938 (Clarke & McIntyre) (TC); 2 2, Quevedo, v.1976 (Fritz) (TC). Mexico: Chiapas: 10,32 kmN. Huixtla, 1000 m, vi.1969 (CNC). Panama: 2 9, Barro Colorado Island, iv.1941 (Zetek) (USNM); 2 9, same locality, 120 m, v.1984, vi.1985 (Wolda) (BMNH). Enicospilus corcovadoi sp. n. (Figs 120, 130, 283) Description. Mandibles rather short, strongly and evenly narrowed, apically twisted 30-5S0°; upper mandibular tooth slender, 1.3—1.4 times as long as the lower tooth which is very stout, swollen and has a sharp cutting edge ventrally (Fig. 120); outer mandibular surface more or less flat, with long fine scattered hairs. Labrum 0.1 times as long as broad; malar space 0.2—0.3 times as long as basal mandibular width. Clypeus in profile moderately convex, margin blunt; clypeus in front view 1.3—1.4 times as broad as long, the apical weakly convex. Lower face 0.6—0.7 times as broad as long, finely punctate. Head in dorsal view rather more quadrate than that of most species, with genae rather long, weakly constricted behind eyes (Fig. 130); posterior ocellus contiguous with eye; FI = 68-70%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6 times the basal mandibular width away from mandible. Antenna slender, with 61 flagellar segments; 20th segment 2.3—2.4 times as long as broad. Mesoscutum polished, finely punctate, in profile evenly rounded; notauli vestigial. Mesopleuron pol- ished, the upper part finely but quite closely punctate, the lower part similar but with indications of striae present anteriorly; epicnemial carina inclined towards but not reaching anterior margin of pleuron, its lower corner rather sharp and slightly raised. Scutellum in profile weakly convex, laterally carinate for 0.6 of its length; scutellum in dorsal view 1.6 times as long as anteriorly broad, finely alutaceous. Metapleuron rather weakly convex, posterodorsally diagonally striate, anteroventrally punctate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly rounded; anterior transverse carina complete or narrowly incomplete at lateral extremities, pos- terior transverse carina absent; anterior area moderately long, quite steep, striate; spiracular area moder- ately long, smooth; posterior area rugose, with the rugae tending to be concentric dorsally; lateral longitudinal carina complete at least on anterior 0.5, sometimes joined to spiracular margin by a short, very weak carina. Fore wing length 14mm; discosubmarginal cell as in Fig. 283; AI = 1.04-1.13; CI = 0.31-0.32; ICI = 0.49- 0.50; SDI = 1.10—1.13; cu-a proximal to the base of Rs&M by about 0.2 times its own length; marginal cell proximally slightly more sparsely pubescent; 1st subdiscal cell with anterodistal part sparsely hirsute. Hind wing with 5 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa almost straight. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.4-1.7 times length of the shorter. Hind leg with coxa in profile 1.7 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.42.6 times as long as broad; claws of female moderately long, evenly curved, with long close pectinae; claws of male similar, but with pectinae slightly shorter. Gaster slender; tergite 2 in profile 5—6 times as long as posteriorly deep, laterotergite folded under, thyridia oval and separated from anterior margin of tergite by 4-9 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing dense, long stout erect pubescence. Colour generally orange-yellow, with mesoscutum reddish brown; interocellar area blackish, often with a dark mark extending ventrally on frons between bases of antennae; antenna blackish, but with scape, pedicel and part of first flagellar segment brownish; terminal segments of gaster weakly to moderately infuscate; hind tibia and tarsus slightly infuscate. Pterostigma blackish; wings weakly infumate. VARIATION. The terminal segments of the gaster of the male are less infuscate than are those of the female. REMARKS. The rather small, somewhat quadrate head is particularly characteristic of this species, though E. fernaldi has a rather similar head form. The two species may be related, but E. corcovadoi can easily be distinguished by the form of the mandibles. Those of E. fernaldi are less strongly twisted and have a much more slender lower tooth than those of E. corcovadoi. BIOLOGICAL INFORMATION. Enicospilus corcovadoi is only known to occur in wet lowland forests in the southern part of Central America. It has only rarely been collected and nothing is known of its hosts. OPHIONINAE OF TROPICAL MESOAMERICA 221 MATERIAL EXAMINED Holotype ©’, Costa Rica: Osa Peninsula, Corcovado National Park, 20 m, v.1984 (Gauld) (BMNH). Paratype. Panama: 1 9, Darién, 1967 (Triplehorn) (TC). Enicospilus lovejoyi sp. n. (Figs 181, 284) Description. Mandibles short, quite strongly narrowed, apically twisted 75—80°; upper mandibular tooth slightly compressed, 1.2—1.4 times as long as the lower tooth; outer mandibular surface finely pubescent, with a weak basal concavity. Labrum 0.2 times as long as broad; malar space 0.2 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.2-1.3 times as broad as long, its apical margin more or less truncate. Lower face 0.6—0.7 times as broad as long, polished, virtually smooth. Head in dorsal view with genae strongly constricted; posterior ocellus very close to eye; FI = 60-65%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6 times the basal mandibular width away from mandible. Antenna long and slender, with 47-50 flagellar segments; 20th segment 1.9-2.1 times as long as broad. Mesoscutum polished, with indistinct punctures, in profile abruptly rounded; notauli very weak. Mesopleuron polished, the upper part very sparsely punctate, the lower part punctate to punctostriate; epicnemial carina slightly curved towards anterior margin of pleuron, but with upper end evanescent. Scutellum in profile evenly rounded, laterally carinate for all of its length; scutellum in dorsal view 1.3-1.4 times as long as anteriorly broad, smooth with isolated punctures, occasionally with a few striae posteri- orly. Metapleuron weakly to moderately convex, finely punctate (Fig. 181); submetapleural carina weakly broadened anteriorly; posterior transverse carina of mesosternum usually complete, sometimes narrowly interrupted centrally. Propodeum in profile evenly rounded; anterior transverse carina present, weak laterally, posterior transverse carina absent; anterior area moderately long, sparsely striate; spiracular area long, smooth; posterior area with weak irregular transverse striations; lateral longitudinal carina usually complete, sometimes joined to spiracular margin by a short weak carina, or sometimes absent. Fore wing length 9-12 mm; discosubmarginal cell as in Fig. 284 (but note many of the hairs have been rubbed off the central part of the discosubmarginal cell in this specimen); AI = 0.55-1.00; CI = 0.25-0.42; ICI = 0.29-0.41; SDI = 1.14-1.18; cu-a opposite or slightly proximal to the base of Rs&M; marginal cell proximally sparsely pubescent, generally with a distinct glabrous area adjacent to Rs+2r; 1st subdiscal cell with anterior and distal parts sparsely hirsute. Hind wing with 4-6 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa almost straight. Fore leg with tibia slightly flattened, with fine scattered spines on outer surface. Mid leg with longer tibial spur 1.4-1.5 times length of the shorter. Hind leg with coxa in profile 1.8-1.9 times as long as deep; trochantellus dorsally 0.1—0.3 times as long as broad; 4th segment of tarsus 3.0-3.1 times as long as broad; claws of female long, quite strongly curved apically, with short pectinae. Gaster slender; tergite 2 in profile 5 or more times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by about 3.54.0 times its own length. Ovipositor moderately slender, its sheath slightly broadened. Male unknown. Colour generally yellowish brown with profuse pale irregular markings on alitrunk; mesoscutal vittae blackish and upper posterior corner of mesopleuron bearing an irregular black mark; head yellow, interocellar area black; antenna brown, basally slightly infuscate; pterostigma brown; wings hyaline. VARIATION. Enicospilus lovejoyi is a morphologically rather uniform species, but southern Brazilian specimens have paler alitrunks than those from the northern end of its range. RemakkKsS. This species is named in honour of Thomas Lovejoy in recognition of his many contributions to Neotropical conservation biology. Enicospilus lovejoyi is a small, distinctive species which may easily be recognized by the combination of the form of the alar sclerites and the black interocellar area. The strong continuous distal sclerite, the mesopleural dark mark and the short, strongly twisted mandibles are apomorphic features that are also found in E. liesneri, E. marini and E. pescadori, and it is probable all are closely related. E. lovejoyi and E. liesneri have very similar modified heads (see remarks under E. tiesneri) suggesting they are sister species. BIOLOGICAL INFORMATION. Enicospilus lovejoyi is a widely distributed species whose range extends from Central Panama south to Peru and Central Brazil (Map 21). The Panamanian specimen was collected in lowland rainforest, but the general habitat preference and host range of this species are not known. 222 IAN D. GAULD Noone, Map 21 Localities at which Enicospilus lovejoyi has been collected. MATERIAL EXAMINED Holotype 9, Brazil: Para, Santarem, x-xi.1966 (Knowles) (BMNH). Paratypes. Brazil: 2 9, Aguas Vermelhas, 800 m, xii.1983 (Alvarenga) (TC); 1 9, Bahia, Encruzilhada, 980 m, xi.1974 (Alvarenga) (TC); 1 9, Guanabara, Represa Rio Grande, xii.1969 (Alvarenga) (TC). Colombia: 1 9, Camp Sautata, Rio Atrato, xi-xii.1967 (TC). Guyana: 2 2, R. Essequibo, Rockstone, iii.1913 (Bodkin) (BMNH). Panama: 1 @, Barro Colorado Island, Gatun Lake, iv.1979 (Wolda) (TC). Peru: 1 9, Loreto, Pucallpa, viii.1963 (Schunke) (BMNH). Surinam: 1 9 , Marowijne R., vii.1965 (Gale) (BMNH). OPHIONINAE OF TROPICAL MESOAMERICA 223 Enicospilus hacha sp. n. (Figs 113, 119, 131, 182, 285) DescriPTIon. Mandibles moderately long, with a pronounced proximoventral lobe, distally evenly nar- rowed and apically twisted 75—85° (Fig. 119); upper mandibular tooth slightly depressed, 1.4-1.6 times as long as the lower tooth; outer mandibular surface rather finely and sparsely pubescent, with a shallow proximal concavity. Labrum 0.2 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin slightly impressed, quite sharp; clypeus in front view 1.3— 1.4 times as broad as long, its apical margin more or less truncate. Lower face 0.6-0.7 times as broad as long, with fine sparse punctures. Head in dorsal view with genae strongly constricted behind eyes (Fig. 131); posterior ocellus more or less contiguous with eye; FI = 65-70%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.8—1.0 times the basal mandibular width away from mandible. Antenna long and slender, with 55-60 flagellar segments; 20th segment 2.3—2.5 times as long as broad. Mesoscutum weakly polished, without obvious punctures, in profile abruptly rounded; notauli absent. Mesopleuron polished, the upper part finely and sparsely punctate, the lower part usually more closely punctate, often with incipient traces of striations observable; epicnemial carina inclined towards anterior margin of pleuron, but its upper end not reaching this margin. Scutellum in profile weakly convex, laterally carinate for all of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, from smooth with isolated punctures to weakly coriaceous. Metapleuron moderately strongly convex, from transversely striate to irregularly vermiculate (Fig. 182); submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina absent or indicated by weak wrinkling; anterior area short and quite deeply impressed, striate; spiracular area moderately long, smooth; posterior area rather coarsely reticulate; lateral longitudinal carina with at least anterior 0.7 complete, joined to spiracular margin by a short carina. Fore wing length 12-14 mm; discosubmarginal cell as in Fig. 285; AI = 1.00-1.32; CI = 0.25-0.42; ICI = 0.44-0.54; SDI = 1.09-1.31; cu-a distal to the base of Rs&M by about 0.3 times its own length; marginal cell proximally rather sparsely hirsute; 1st subdiscal cell with anterior and distal parts sparsely hirsute. Hind wing with 6 hamuli on R1; 1st and 2nd abscissa of Rs more or less straight. Fore leg with tibia somewhat flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.3-1.5 times length of the shorter. Hind leg with coxa in profile 1.7-1.8 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.4—2.5 times as long as broad; claws of female of moderate length, abruptly curved apically, with close long pectinae; claws of male similar but with pectinae shorter. Gaster slender; tergite 2 in profile more than 6 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 3 or more times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing moderately dense long erect pubescence; gonosquama quite slender, dorsally more strongly tapered than it is ventrally. Colour generally yellowish orange; interocellar area, antenna and tergites 3+ of gaster black; meso- scutum infuscate. Pterostigma brownish; wings more or less hyaline. VaRIATION. Enicospilus hacha is a morphologically rather uniform species that shows a small amount of variation in the sculpture of the meso- and metapleurae, as detailed above. The Jamaican specimen is almost uniformly pale yellowish, and lacks the dark marks present on individuals from other localities. REMARKS. Enicospilus hacha may be recognized most easily by the characteristic form of the central sclerite (Fig. 285) and the possession of a strongly sclerotized complete, distal sclerite. In this latter feature, and in the form of the mandibles, E. hacha resembles E. lovejoyi and it is possible that the two species are related. BIoLocicaL INForMaTION. Enicospilus hacha is a Central American and tropical South American species whose range extends from central Mexico south to Colombia (Map 22). It is also known to occur on the island of Jamaica. In Costa Rica this species has only been collected in the north-western part of the country, in Guanacaste or just across the watershed in northern Alajuela. It is present at more humid sites in December, February and March, but it has only been collected in dry forest sites during the wetter months of year. For example, in Santa Rosa National Park it has been collected in May and August, and on Cerro el Hacha it was taken between August and January. Nothing is known of the host range of this species. MATERIAL EXAMINED Holotype 2, Costa Rica: Guanacaste Prov., Guanacaste National Park, Cerro el Hacha, 3-400 m, viii- xi.1986 (Janzen & Hallwachs) (BMNH). IAN D. GAULD 224 ‘pa}d9]]09 Ud0q sey vYIvY snpidsoolUT YoIyM ye sayesoT + 7 dey OPHIONINAE OF TROPICAL MESOAMERICA 225 Paratypes. Colombia: 1 2 , Magdalena, 800 m, iv.1973 (Heleva) (BMNH). Costa Rica: Alajuela Prov.: 1 Q , Finca San Gabriel, 750 m, iii. 1984 (Janzen) (BMNH): Guanacaste Prov.: 1 9, Cerro el Hacha, 3-400 m, i.1987 (Janzen & Hallwachs) (BMNH); 10’, 1 9, Rinc6n de la Vieja National Park, 4km E. Casetilla, 750 m, xii.1981, ii.1983 (Janzen & Hallwachs) (BMNH); 1 9, 5 km NE. of Quebrada Grande, 600 m, vi.1986 (Gauld) (BMNH); 1 o’, 1 9, Santa Rosa National Park, 300 m, v & viii.1983 (Janzen & Hallwachs) (BMNH). Jamaica: 1 9, Port Antonio, Bonnie View, vii.1952 (FSCA). Mexico: Veracruz: 1 CO’, L. Catemaco, 11 km N. Hotel Playa Azul, ca 400 m, viii.1963 (Weems) (FSCA). Panama: 1 9, Cerro Campana, vii.1970 (Howden & Howden) (TC) Enicospilus xanthostigma (Szépligeti) (Figs 114, 118, 122, 183, 286) Henicospilus xanthostigma Szépligeti, 1906: 147. Holotype 0’, BRAZIL (TM) [examined. ] Enicospilus xanthostigma (Szépligeti) Hooker, 1912: 91. Description. Mandibles moderately long, with a proximoventral lobe, distally evenly narrowed and apically twisted 70—75°; upper mandibular tooth slightly compressed, 1.2—1.4 times as long as the lower tooth (Fig. 118); outer mandibular surface sparsely pubescent, more or less flat. Labrum 0.2-0.3 times as long as broad; malar space 0.3—0.4 times as long as basal mandibular width. Clypeus in profile weakly convex, margin acute; clypeus in front view 1.3—1.5 times as broad as long, the margin apically weakly convex. Lower face 0.60.7 times as broad as long, polished, virtually impunctate. Head in dorsal view with genae rounded behind eye; posterior ocellus very close to but not touching eye; FI = 60-65%; occipital carina mediodorsally complete, ventrally joining hypostomal carina about 0.7 of basal width away from the mandible (Fig. 114). Antenna very long and slender, with 56-65 flagellar segments; 20th segment 3.0-3.2 times as long as broad. Mesoscutum polished, with minute inconspicuous punctures, in profile abruptly rounded anterior margin out-turned; notauli absent. Mesopleuron polished, the upper part smooth and shining, the lower part similar but with fine punctures; epicnemial carina evenly curved towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for all of its length; scutellum in dorsal view 1.5 times as long as anteriorly broad, finely and indistinctly punctate, striate posteriorly. Metapleuron very weakly convex, polished with obsolescent punctures (Fig. 183); submetapleural carina evenly anteriorly broad- ened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area moderately long, rather shallowly impressed, striate; spiracular area long, smooth; posterior area with very weak coriaceous sculpture; lateral longitudinal carina complete, usually joined to spiracular margin by a short carina. Fore wing length 10-11 mm; discosubmarginal cell as in Fig. 286; AI = 0.59-1.06; CI = 0.26-0.37; ICI = 0.23-0.40; SDI = 0.86-1.07; cu-a from distal to the base of Rs&M by about 0.2 times its own length to slightly proximal to this vein; marginal cell proximally glabrous; 1st subdiscal cell distally sparsely pubescent. Hind wing with 4-5 hamuli on R1; Ist abscissa of Rs straignt, 2nd abscissa weakly arcuate. Fore leg with tibia very weakly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.8—1.9 times length of the shorter. Hind leg with coxa in profile 1.8—1.9 times as long as deep; trochantellus dorsally 0.2 times as long as broad (Fig. 122); 4th segment of tarsus 2.6-2.7 times as long as broad; claws of female moderately long, fairly abruptly curved apically, with stout close pectinae. Gaster slender; tergite 2 in profile at least 6 times as long as posteriorly deep, laterotergite turned under, thyridia oval and separated from anterior margin of tergite by about 3.0-3.5 times its own length. Ovipositor apically slender, its sheath narrow. Male with sternites 6-9 bearing very long, erect pubescence; gonosquama apically acute. Colour generally golden yellowish with irregular pale markings on the head and on the alitrunk laterally; mesoscutum brown or infuscate; mesopleuron with a very weak to quite distinct dark mark in posterodor- sal corner; gaster with tergites 5+ infuscate; interocellar area and vertex extensively black; antenna infuscate; pterostigma brown; wings hyaline. VARIATION. The Mexican specimens have the entire anterior part of the mesoscutum blackish whereas in South American specimens this is brown. REMARKS. Enicospilus xanthostigma is a small species that is most easily recognized by the characteristic form of the alar sclerites. It is structurally very similar to Enicospilus cepillo, which it closely resembles in venation and form of the proximal sclerite. The two are most easily distinguished by the mandibles; those of E. xanthostigma are more strongly twisted. E. xanthostigma also has a shorter hind trochantellus than E. cepillo. It also resembles an undescribed Brazilian species (in TC), which differs most obviously in having infumate markings in the wing. 226 IAN D. GAULD BIOLOGICAL INFORMATION. Enicospilus xanthostigma is a widely distributed, but seldom collected species whose range extends from southern Mexico, south into Ecuador, Peru and Brazil. Although it has not been collected in Central America its known distribution suggests it may occur there. Most specimens have been collected at altitudes between 400 and 1000 m. MATERIAL EXAMINED Holotype OC’, Brazil: Blumenau (TM). Ecuador: 2 9, Napo & Coca Rivers, v.1965 (Pera) (TC); 1 9, Pichincha, Tinalandia, 16 km SE. Santo Domingo, vi.1976 (Peck & Peck) (TC). Mexico: Chiapas: 1 9, 32 km N. Huixtla, vi.1969 (CNC); 3 Q, Muste, nr Huixtla, 440 m, ix.1970 (Welling) (CNC). Peru: 2 2, Quincemil, nr Marcapata, 750 m, xi.1962 (Pena) (TC). Enicospilus cepillo sp. n. (Figs 115, 117, 121, 287) Description. Mandibles short, quite strongly and evenly tapered, apically twisted 10—20°; upper mandibu- lar tooth slightly depressed, 1.3—-1.5 times as long as the lower tooth (Fig. 117); outer mandibular surface with a weak longitudinal concavity. Labrum 0.2 times as long as broad; malar space 0. 1—0.2 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.5-1.6 times as broad as long, the margin apically truncate. Lower face 0.70-0.76 times as broad as long, smooth and polished, centrally punctate. Head in dorsal view with genae constricted behind eyes; posterior ocellus contiguous with eye; FI = 65-70%; occipital carina mediodorsally complete, ventrally evanescent, not joining hypostomal carina (Fig. 115). Antenna very long and slender, with 52-56 flagellar segments; 20th segment 2.5—2.8 times as long as broad. Mesoscutum polished, smooth, in profile steeply rounded; notauli absent. Mesopleuron highly polished, the upper and lower parts finely and sparsely punctate; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.8—-0.9 of its length; scutellum in dorsal view 1.3-1.5 times as long as anteriorly broad, smooth. Metapleuron very weakly convex, polished, finely and sparsely punctate; submetapleural carina evenly broadened anteriorly; posterior transverse carina of mesosternum broadly incomplete centrally. Propodeum in profile evenly rounded; anterior transverse carina complete, posterior transverse carina absent; anterior area quite short, weakly impressed, from smooth to slightly coriaceous; spiracular area exceptionally long, smooth; posterior area virtually smooth, at most with very weak rugosities; lateral longitudinal carina weak, but usually discerni- ble anteriorly, sometimes complete, not joined to spiracular margin by a short carina. Fore wing length 8-9 mm; discosubmarginal cell as in Fig. 287; AI = 0.98-1.29; CI = 0.17-0.24; ICI = 0.39-0.47; SDI = 1.05-1.15; cu-a virtually opposite to the base of Rs&M; marginal cell with proximal corner broadly glabrous ; 1st subdiscal cell with scattered hairs anteriorly and distally. Hind wing with 5 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa straight. Fore leg with tibia moderately flattened, with scattered weak spines on outer surface. Mid leg with longer tibial spur 1.4—1.6 times length of the shorter. Hind leg with coxa in profile 1.8-1.9 times as long as deep; trochantellus dorsally 0.5—0.6 times as long as broad (Fig. 121); 4th segment of tarsus 2.3-2.4 times as long as broad; claws of female short, abruptly curved with short close pectinae, those of male similar. Gaster slender; tergite 2 in profile more than 5 times as long as posteriorly deep, laterotergite generally folded under, thyridia oval to elliptical and separated from anterior margin of tergite by about 2—3 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing long fine sparse pubescence and with a few long stout erect bristles along posterior margin; gonosquama evenly rounded. Colour generally with head, scutellum and lateral parts of alitrunk yellow, gaster yellowish brown; maxillary and labial palps infuscate; mesoscutal vittae and upper posterior margin of mesopleuron dark brown; interocellar area black; antenna dark brown; pterostigma yellowish brown; wings hyaline. VARIATION. This is structurally a rather uniform species, but the flagellum of one of the Panamanian specimens is slightly paler than those of the other specimens. REMARKS. Enicospilus cepillo is a very distinctive species on account of the ventrally incomplete occipital carina, broadly incomplete posterior transverse carina of the mesosternum and long hind trochantellus. The male is unusual in having exceptionally stout bristles only along the hind margin of the sternites. Structurally it otherwise resembles E. xanthostigma, and the two species may be closely related. BIOLOGICAL INFORMATION. Enicospilus cepillo is a rarely collected species whose range extend from the Isthmus of Panama south to Peru and Central Brazil. In Panama it has been collected in lowland rainforest, but its hosts are not known. OPHIONINAE OF TROPICAL MESOAMERICA 2H MATERIAL EXAMINED Holotype 2, Panama: Barro Colorado Island, 120 m, vii.1985 (Wolda) (BMNH). Paratypes. Brazil: 1 2, Bahia, Encruzilhada, 980 m, xi.1974 (Alvarenga) (TC). Panama: 1 9, Barro Colorado Island, 120 m, v.1985 (Wolda) (BMNH); 1 9, Tucaman, i.1953 (USNM). Peru: 1 0’, Loreto, Pucallpa, 1.1952 (Schunke) (BMNH). Enicospilus persimilis (Szépligeti) (Fig. 288) Henicospilus persimilis Szépligeti, 1906: 147. Holotype co’, PERU (TM) [examined]. [Enicospilus fuscipennis (Szépligeti) Hooker, 1912: 89. In part, misidentification. ] Enicospilus persimilis (Szépligeti) Townes & Townes, 1966: 183. Description. Mandibles moderately long, quite strongly and evenly tapered, apically twisted 20—30°; upper mandibular tooth slightly depressed 1.2—1.4 times as long as the lower tooth which is distinctly the stouter of the two; outer mandibular surface with a few long scattered hairs proximally, distally flat but with a weak proximal concavity. Labrum 0.2 times as long as broad; malar space 0.2 times as long as basal mandibular width. Clypeus in profile almost flat, margin subacute; clypeus in front view 1.3—-1.4 times as broad as long, the margin weakly convex. Lower face 0.68—0.70 times as broad as long, polished, centrally striate. Head in dorsal view with genae constricted behind eyes; posterior ocellus contiguous with eye; FI = 65-70% ; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 1.0 times the basal mandibular width away from mandible. Antenna long and slender, with 58-61 flagellar segments; 20th segment 2.3—2.7 times as long as broad. Mesoscutum polished, finely punctate, in profile steeply rounded; notauli absent. Mesopleuron pol- ished, the upper part punctate, the lower part punctostriate to striate; epicnemial carina inclined towards anterior margin of pleuron, its upper end evanescent. Scutellum in profile weakly convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.6 times as long as anteriorly broad, anteriorly smooth, posteriorly striate. Metapleuron weakly to moderately convex, diagonally striate, weakly pol- ished; submetapleural carina quite strongly anteriorly broadened; posterior transverse carina of mesoster- num complete. Propodeum in profile evenly rounded; anterior transverse carina complete except at extreme lateral extremities, posterior transverse carina absent; anterior area quite steep, shagreened with a few irregular striae; spiracular area moderately long, smooth; posterior area from coarsely concentrically striate to irregularly transversely wrinkled; lateral longitudinal carina complete, sometimes joined to spiracular margin by a short weak carina. Fore wing length 14-15 mm; discosubmarginal cell as in Fig. 288; AI = 1.19-1.72; CI = 0.44-0.50; ICI = 0.42-0.45; SDI = 1.13-1.27; cu-a proximal to the base of Rs&M by 0.3-0.4 times its own length; marginal cell with proximal corner entirely glabrous; 1st subdiscal cell with anterior 0.4 hirsute, proximal end entirely glabrous. Hind wing with 5-6 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa almost straight. Fore leg with tibia very weakly flattened, with slender, inconspicuous spines on outer surface. Mid leg with longer tibial spur 1.7—1.8 times length of the shorter. Hind leg with coxa in profile 1.8-1.9 times as long as deep; trochantellus dorsally 0.1 or less times as long as broad; 4th segment of tarsus 2.5—2.6 times as long as broad; claws of female long, abruptly curved, with stout incurved pectinae; claws of male similar but with pectinae finer. Gaster long and slender; tergite 2 in profile 6 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 3 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing short fine decumbent pubescence; gonosquama apically evenly rounded. Colour generally orange brown with head paler yellowish and tergites 3+ of gaster infuscate; interocellar area black; antenna blackish; pterostigma blackish brown; wings weakly infumate. VARIATION. The Panamanian and Surinamese specimens are extremely similar, so similar in fact that they would pass as having come from the same locality. REMARKS. The characteristic inverted J-shaped central sclerite (Fig. 288) makes Enicospilus persimilis one of the most easily recognized of all Neotropical species. In general structure, venation and form of the proximal sclerite it resembles E. madrigalae and E. duckworthi, and these species may well be closely related. The central sclerite of E. duckworthi is oval and that of E. madrigalae is linear. BIOLOGICAL INFORMATION. Enicospilus persimilis is an uncommon species in collections and is only known from three specimens, the holotype male from Peru, and two females, one collected in Surinam, and a 228 IAN D. GAULD second collected in lowland rainforest on Barro Colorado Island, Panama. Nothing is known about the biology. MATERIAL EXAMINED Holotype CO’, Peru: Pachitea (TM). Panama: 1 9, Barro Colorado Island, v.1941 (Zetek) (USNM). Surinam: 1 9 , Lavva Anapaike, xi.1963 (Ligorie) (TC). Enicospilus fernaldi Hooker (Figs 184, 289) Enicospilus fernaldi Hooker, 1912: 63. Lectotype 2, DOMINICAN REPUBLIC (USNM), designated by Townes & Townes (1966: 177) [examined]. Enicospilus fernaldi Hooker; Brues & Richardson, 1913: 496. Description. Mandibles fairly short, evenly narrowed, apically twisted 15—20°; upper tooth strongly depressed, slender, 1.2—1.5 times as long as the lower tooth; outer mandibular surface finely pubescent, distally weakly concave, but without a distinct proximal concavity. Labrum 0.2 times as long as broad; malar space 0.3—0.4 times as long as basal mandibular width. Clypeus in profile weakly to moderately convex, margin blunt; clypeus in front view 1.2-1.4 times as broad as long, the apical margin weakly convex. Lower face 0.65—0.75 times as broad as long, finely punctate, often with punctures centrally grading to striae. Head in dorsal view with genae evenly constricted behind eyes; posterior ocellus very close to eye; FI = 60-65%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.9-1.0 times the basal mandibular width away from mandible. Antenna slender, with 55—57 flagellar segments; 20th segment 2.1—2.2 times as long as broad. Mesoscutum polished, finely and inconspicuously punctate, in profile evenly rounded; notauli vestigial. Mesopleuron weakly polished, the upper part centrally smooth, peripherally finely punctate to punctostri- ate, the lower part punctostriate to striate; epicnemial carina inclined towards but not reaching anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.6 or more of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, smooth or finely punctate, grading posteriorly into irregularly wrinkled. Metapleuron moderately convex, quite coarsely punctostriate to coriaceous (Fig. 184); submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile fairly abruptly declivous; anterior transverse carina com- plete except at lateral extremities, posterior transverse carina absent; anterior area moderately long and quite strongly impressed, striate; spiracular area fairly short, almost smooth; posterior area irregularly reticulately wrinkled; lateral longitudinal carina from complete to weak and discontinuous, usually not joined to spiracular margin by a strong, short carina though sometimes a weak one may be present and rarely it may be well developed. Fore wing length 11-13 mm; discosubmarginal cell as in Fig. 289; AI = 0.75-1.00; CI = 0.18-0.37; ICI = 0.38-0.51; SDI = 1.07-1.27; cu-a slightly proximal to the base of Rs&M; marginal cell proximally rather sparsely hirsute, sometimes with a glabrous area; 1st subdiscal cell with distal 0.6 hirsute, especially centrally. Hind wing with 5—7 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa more or less straight. Fore leg with tibia subcylindrical, without obvious spines on outer surface, or with very slender, scattered spines. Mid leg with longer tibial spur 1.4-1.5 times length of the shorter. Hind leg with coxa in profile 1.7—1.9 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.7—2.9 times as long as broad; claws of female moderately long, apically strongly curved, with rather stout long pectinae; claws of male similar but with short pectinae. Gaster slender; tergite 2 in profile 5 or more times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by about 3 or more times its own length. Ovipositor slender, its sheath fairly narrow. Male with sternites 7-9 bearing scattered erect pubescence; gonosquama distally rounded. Colour generally brownish orange with paler stripes on mesoscutum; interocellar area and tergites 5+ of gaster black; antenna proximally reddish brown, distally infuscate; pterostigma orange brown; wings hyaline. VARIATION. Some of the Caribbean specimens are paler yellowish in colour and have a bright yellow scutellum. Specimens from Belize all have the terminal segments of the gaster slightly infuscate, not black as do most individuals. The distal sclerite is usually confluent with the proximal sclerite but evanescent before reaching the level of the central sclerite. In a few individuals it is continued just distal to level of central sclerite. OPHIONINAE OF TROPICAL MESOAMERICA 229 Remarks. Enicospilus fernaldi is easily recognized by the rather characteristic colour pattern — black interocellar area and tip of the gaster. Its head, in dorsal view, is slightly more quadrate than most other species. Structurally E. fernaldi most closely resembles E. flavus and E. corcovadoi, but can be separated from either by the form of the alar sclerites (compare Figs 283, 289, 290). BIOLOGICAL INFORMATION. Enicospilus fernaldi is a widespread species whose range extends from the southern tip of Florida, south throughout the Caribbean to Trinidad, and from Mexico south to central Brazil (Map 23). It has been collected most frequently on some Caribbean islands, where it seems to be one of the commoner species of the genus. E. fernaldi is much less frequently collected on the Central American mainland. For example, in Costa Rica extensive collecting has yielded only two individuals in worse phe ad ee ira Map 23 Localities at which Enicospilus fernaldi has been collected. 230 IAN D. GAULD Santa Rosa National Park, a female in July 1982 and another female the following year in May, a male from Cerro el Hacha, and a single specimen from Turrialba. No specimens were collected on Barro Colorado Island, Panama, despite extensive sampling in lowland forest. The hosts of this species are unknown. MaTERIAL EXAMINED Lectotype 2, Dominican Republic: San Francisco (in mountains near San Cristobal) ix.1905 (Busck) (USNM),; paralectotypes: 1 0’, 1 2, same data as lectotype (USNM). Belize: 1 2 , Coyo Dist., Camp Sibim, v.1963 (CNC); 1 9, Punta Gordas, vii.1935 (White) (BMNH);1 9, Rio Grande, xi.1935 (White) (BMNH); 1 2, Rio Temas, 1935 (White) (BMNH). Brazil: 1 9, Amazonas, Ducke Res., 25 km E. of Manaus, iii.1973 (Tyson) (CNC); 1 9, Ceara, Barbalha, 400 m, v.1969 (Alvarenga) (TC); 4 2, Guanabara, Represa Rio Grande, v.1967 (Alvarenga) (TC). Colombia: 1 9 , Meta, Villavicencio, 450 m, 1936 (Bequaert) (MCZ). Costa Rica: Cartago Prov.: 1 O&, Turrialba, viii.1977 (Hansen) (CNC): Guanacaste Prov.: 1 0’, Cerro el Hacha, 3-400 m, i-iv.1987 (Janzen & Hallwachs) (BMNH); 2 9, Santa Rosa National Park, 300 m, vii.1982, v.1983 (Janzen & Hallwachs) (BMNH). Cuba: 1 2, Baragua, x.1928 (Rambousen) (MCZ); 1 2, Cienfuegos, Soledad, Limones Seboruco, ix.1930 (Dow) (MCZ); 3 2, Cienfuegos, Soledad, Vilches Potrero, viii-ix.1930 (Dow) (MCZ); 2 OC’, Pinard R. ix.1913 (USNM); 2 9, Santa Clara, San Blas, viii.1932 (Leavitt) (MCZ); 1 9, Santa Clara, San José, Trinidad Mts, viii.1930 (Dow) (MCZ);2 0,2 2, 7kmN. of Vinales, ix.1913 (USNM). Dominica: 1 0’, 1 2, Clarke Hall, xii. 1964-1.1965 (Spangler & Wirth) (USNM); 1 9, Roseau, vi.1911 (MCZ). Dominican Republic: 4 9, Altagracia, Nisibon, v.1978 (Fairchild) (FSCA); 1 0’, Greenhill Est., vii.1941 (Fennah) (USNM). Gre- nada: 2 Q, Saint Georges Parish, Grand Anse, ii.1977 (Tanaka) (CAS). Mexico: 2 9, Jesus Carranza, viii. 1939 (Townes) (TC). Nicaragua: 1 2, Zelaya, El Recreo, x.1964 (MCZ). Panama: 1 9, Darién, 1967 (Triplehorn) (TC). Trinidad: 2 2, Caroni R., xi.1952 (Simmonds) (CNC); 1 2, Curepe, x.1952 (Sim- monds) (CNC); 2 2, Orange Grove, x.1952 (Simmonds) (CNC); 1 9, San Juan, x.1952 (Simmonds) (CNC); 2 2, no further data (Voogd) (RNH). U.S.A.: Florida: 1 2, Monroe Co., No Name Key, xii.1972 (Dodge) (FSCA). Venezuela: 1 2, San Esteban, nr Puerto Cabello, xi.1939 (Anduze) (TC); 1 9, Zulia, Tucuco, iv.1981 (Townes) (TC). Enicospilus flavus (Fabricius) (Figs 290, 341) Ichneumon flavus Fabricius, 1775: 341. Lectotype 9, ‘AMERICA’, designated by Townes & Townes (1966: 178) (Kiel). Ophion flavus (Fabricius) Fabricius, 1798: 236. Ichneumon flavarius Thunberg, 1822: 262. [Unnecessary replacement name. ] Ophion concolor Cresson, 1865: 56. LECTOTYPE ©’, CUBA, here designated (PANS) [examined]. [Synonymized by Townes in Wolcott, 1948: 765.] Ophion concolor Cresson; Fox, 1891: 337. Enicospilus concolor (Cresson) Ashmead, 1900b: 271. Enicospilus guyanensis Cameron, 1911: 179. Holotype 2, GUYANA (BMNH) [examined]. [Syn- onymized by Morley, 1914: 409.] Enicospilus flavus (Fabricius); Hooker, 1912: 71. [In part.] Henicospilus concolor (Cresson) Morley, 1912: 33. Henicospilus flavoscutellatus var. concolor (Cresson); Enderlein, 1921: 82. Enicospilus concolor (Cresson); Walcott, 1923: 65. Henicospilus concolor (Cresson); Ogilvie, 1928: 48. Enicospilus flavus (Fabricius); Townes & Townes, 1966: 178. DescriPTION. Mandibles of moderate length, quite strongly narrowed, apically twisted 25—40°; upper mandibular tooth subcylindrical, 1.2—1.5 times as long as the lower tooth; outer mandibular surface weakly convex, but with a distinct proximal concavity. Labrum 0.2 times as long as broad; malar space 0.2-0.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt to moderately sharp, not impressed; clypeus in front view 1.2—1.4 times as broad as long, its margin truncate to weakly convex. Lower face 0.6-0.7 times as broad as long, finely punctate laterally, centrally with traces of punctostriation. Head in dorsal view with genae strongly constricted behind eye; posterior ocellus very close to eye; FI = 60-65%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.8—1.0 times the basal mandibular width away from mandible. Antenna moderately slender, with 55—59 flagellar segments; 20th segment 2.2-2.3 times as long as broad. Mesoscutum polished with inconspicuous punctures, in profile evenly rounded; notauli absent. Meso- pleuron weakly polished, the upper part quite closely punctate, grading ventrally to punctostriate, the OPHIONINAE OF TROPICAL MESOAMERICA 231 lower part punctostriate to striate; epicnemial carina curved towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for all of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, polished, usually rather smooth, sometimes with a few striae posteriorly. Metapleuron moderately convex, punctate, rarely in a few individuals grading to punctostriate; sub- metapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile fairly evenly declivous; anterior transverse carina usually complete, posterior transverse carina absent; anterior area strongly impressed, striate; spiracular area moderately long, smooth; posterior area reticulate; lateral longitudinal carina usually complete, usually joined to spiracular margin by a short carina. Fore wing length 11-14 mm; discosubmarginal cell as in Figs 290, 341; AI = 0.66-1.13; CI = 0.18-0.41; ICI = 0.55—0.68; SDI = 1.14-1.33; cu-a proximal to the base of Rs&M by 0.1-0.3 times its own length; marginal cell proximally uniformly hairy; 1st subdiscal cell with distal 0.8 hirsute. Hind wing with 5—7 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa weakly arcuate. Fore leg with tibia weakly flattened, with isolated spines on outer surface. Mid leg with longer tibial spur 1.3-1.5 times length of the shorter. Hind leg with coxa in profile 1.8—-1.9 times as long as deep; trochantellus dorsally 0.10.2 times as long as broad; 4th segment of tarsus 2.7—3.1 times as long as broad; claws of female moderately long, apically strongly curved, with long stout pectinae; claws of male similar but with pectinae slightly shorter. Gaster slender; tergite 2 in profile 5 or more times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by about 3 or more times its own length. Ovipositor slender, its sheath very narrow. Male with sternites 7-9 bearing scattered long erect hairs; gonosquama distally elongately rounded. Colour generally uniformly orange-brown; interocellar area from black to brownish, contrasting with remainder of vertex; antenna reddish orange; pterostigma yellowish orange; wings hyaline. VARIATION. This species shows considerable variation in the sculpture of the lateral parts of the alitrunk. The metapleuron may be regularly and rather sparsely punctate, closely punctate or even coarsely punctostriate, almost grading to striate. This range of variation can occur within apparently a single population (e.g. the Anguilla sample), but is most pronounced in the Caribbean examples. Most mainland South American species have the metapleuron punctate. One or two individuals, including the lectotype of E. concolor, have the interocellar area yellowish brown, and only infuscate adjacent to the posterior ocelli. Remarks. In PANS there are three syntypic specimens (76-1, 76-2 and 76-3) determined by Cresson as ‘Ophion concolor’. The first is a male, and this was labelled as ‘type’ by Cresson. The remaining two specimens (both females) were labelled as paratypes. In 1916 Cresson (invalidly) designated lectotypes for a large number of his species; he listed each species together with the sex of the lectotype. However, in the case of E. concolor alone the sex symbol was omitted; presumably this was a printer’s error and I believe Cresson intended to designate the male as lectotype. Townes & Townes (1966) state that the lectotype (which they acknowledge as being designated by Cresson) is female. Perhaps this mistake arose from reading the entry above concolor (Ophion), which is concolor (Mesoleptus) for which the lectotype sex is given as female. Whatever, Cresson’s blanket designation of lectotypes in invalid under Article 74(c) of the Code. I hereby select as lectotype the male labelled as ‘type’ by Cresson (76-1). In the literature published between 1837 and 1948 there are many references to E. flavus (see Townes & Townes, 1966), but a large proportion are almost certainly misidentifications of E. trilineatus. This latter species was figured in three publications (Guérin-Ménéville & Pecheron, 1835; Cameron, 1886; Cushman, 1947) under the name flavus. When I examined the collection in the United States National Museum I discovered that virtually all the specimens determined as E. flavus were in fact E. trilineatus. E. flavus is rather similar to and can be confused with E. fernaldi. The most obvious differences are in the form of the alar sclerites. The central sclerite of E. fernaldiis more nearly circular and larger with its longest axis at 90° to Rs+2r (Fig. 289); that of E. flavus is smaller, more elliptical with its longest axis subtending an angle of about 70° to Rs+2r (Fig. 290). The marginal cell of E. flavus is uniformly finely and closely hirsute whilst that of E. fernaldi is more coarsely hirsute and is proximally far less densely pubescent than it is centrally. The mandibles of the two species are quite different, though this difference is difficult to appreciate unless both species are at hand. The upper mandibular tooth of E. fernaldi is definitely depressed, whilst that of E. flavus is more cylindrical. E. fernaldi lacks the basal concavity possessed by the mandible of E. flavus, but has the distal part of the outer face concave, not slightly convex as in E. flavus. BIOLOGICAL INFORMATION. Enicospilus flavus is widely distributed from Florida south throughout the Caribbean and thence southwards into South America as far south as northern Argentina (Map 24). In Central America it apparently only extends northwards to the Isthmus of Panama. Despite extensive 232 IAN D. GAULD Map 24 Localities at which Enicospilus flavus has been collected. collecting I have not found it to be present in Costa Rica. The records of E. flavus from Mexico, Guatemala and Nicaragua seem to be based on specimens of E. trilineatus and I have seen no authentic specimens of E. flavus from these countries. Although it is extremely widely distributed throughout much of the Neotropi- cal region, E. flavus appears to be most common on the islands of the Caribbean. On many islands it is the most commonly collected species of the genus. E. flavus appears to be most common in open habitats, and it has been described as ‘abundant in grass’ in Puerto Rico (Walcott, 1923). MATERIAL EXAMINED Lectotype OC’ (Ophion concolor Cresson), Cuba (PANS); paralectotypes 2 9, same locality (PANS). OPHIONINAE OF TROPICAL MESOAMERICA 233 Holotype 2 (Enicospilus guyanensis Cameron), Guyana (BMNH). 38 0’, 57 Q, from the following localities - Anguilla; Argentina (Tucuman); Bahamas (Long Island); Bolivia (Chapare, Santa Cruz); Brazil (Bahia, Santa Catarina); Cayman Islands (Grand Cayman); Colom- bia (Atlantico, Valle); Cuba; Dominica; Dominican Republic; Ecuador (Napo, Pichincha); Grenada; Guyana; Haiti; Jamaica; Monserrat; Panama (Canal Zone); Peru (Loreto, Quiroz); Puerto Rico; Saint Kitts & Nevis (Nevis); Saint Lucia; Saint Vincent; Surinam; U.S.A. (Florida — Alachua, Broward, Dade, Highlands, Lee, Leon, Manatee, Marion, Monroe, Palm Beach, Putnam and Suwannee counties); Venezuela; Virgin Islands (BMNH, CAS, CNC, FSCA, MCZ, TC, USNM). This species has also been recorded from Bermuda (Ogilvie, 1928) and Paraguay (Schrottky, 1913), but in view of its widespread misidentification these records require confirmation. Enicospilus monticola (Cameron) (Figs 186, 291, 292, 293, 310) Ophion (Enicospilus) monticola Cameron, 1886: 292. Lectotype 9, GUATEMALA (BMNH), desig- nated by Townes & Townes (1966: 182) [examined]. Henicospilus monticola (Cameron) Dalla Torre, 1901: 182. Henicospilus elegans Szépligeti, 1906: 146. Lectotype 2, BRAZIL (TM), designated by Townes & Townes (1966: 177) [examined]. Syn. n. Henicospilus fuscipennis Szépligeti, 1906: 147. Lectotype 9, BOLIVIA (TM), designated by Townes & Townes (1966: 180) [examined]. Syn. n. Enicospilus fuscipennis (Szépligeti) Hooker, 1912: 58, 89. Enicospilus monticola (Cameron) Hooker, 1912: 64. Enicospilus elegans (Szépligeti) Hooker, 1912: 88. [Henicospilus nigricornis (Brullé) Morley, 1912: 34. Misidentification. | Henicospilus antomelas Enderlein, 1921: 35. Holotype 2, COLOMBIA (IZPAN) [examined]. Syn. n. Enicospilus antomelas (Enderlein) Townes & Townes, 1966: 174. Description. Mandibles moderately long, proximally strongly constricted and with a well-developed proximoventral lobe, distally weakly narrowed, apically twisted 45—65°; upper mandibular tooth cylindri- cal, 1.3-1.7 times as long as the lower tooth; outer mandibular surface finely pubescent, more or less flat. Labrum 0.2-0.3 times as long as broad; malar space 0.3—0.4 times as long as basal mandibular width. Clypeus in profile weakly convex, margin usually blunt; clypeus in front view 1.2-1.5 times as broad as long, with margin apically weakly convex. Lower face 0.55—0.65 times as broad as long, finely punctate. Head in dorsal view with genae evenly constricted behind eyes; posterior ocellus contiguous with eye; FI = 75-80% ; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6— 0.8 times the basal mandibular width away from mandible. Antenna slender, with 59-62 flagellar seg- ments; 20th segment 1.7—2.4 times as long as broad. Mesoscutum rather weakly polished, with fine inconspicuous punctures, in profile evenly rounded; notauli vestigial. Mesopleuron polished, the upper part finely punctate or rarely punctostriate, the lower part from finely punctate to punctostriate; epicnemial carina usually slightly inclined towards anterior margin of pleuron, often with upper end evanescent, and most frequently with lower corner somewhat produced as a weak tubercle (Fig. 189). Scutellum in profile weakly convex, laterally carinate for 0.7 or more of its length; scutellum in dorsal view 1.5—1.7 times as long as anteriorly broad, from smooth with fine punctures to wrinkled. Metapleuron moderately strongly convex, generally diagonally striate, sometimes with strong diagonal rugae present posterodorsally, sometimes with anteroventral region punctate, at the most extreme almost entirely punctate, with traces of striae or rugae anterodorsally; submetapleural carina quite broad anteriorly; posterior transverse carina of mesosternum from complete to centrally weak and even sometimes broadly discontinuous medially. Propodeum in profile fairly abruptly declivous; anterior transverse carina complete or with lateral extremities evanescent, posterior transverse carina usually absent, in largest specimens sometimes represented by weak lateral crests; anterior area moderately long, strongly impressed, striate or coriaceous; spiracular area fairly short, generally smooth or sometimes finely punctate; posterior area reticulate to somewhat concentrically rugose, sometimes with traces of lateral crests or a median longitudinal wrinkle; lateral longitudinal carina usually complete, sometimes indistinct posteriorly, joined to spiracular margin by a short carina. Fore wing length 15-19 mm; discosubmarginal cell as in Figs 291-293, 310; AI = 0.86-1.61; CI = 0.21- 0.46; ICI = 0.35-0.55; SDI = 1.07—-1.44; cu-a proximal to the base of Rs&M by 0. 1—0.3 times its own length; marginal cell usually proximally evenly hirsute; 1st subdiscal cell with anterior and distal margins broadly hirsute. Hind wing with 6-7 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa weakly curved. 234 IAN D. GAULD Fore leg with tibia slightly flattened, with numerous slender spines on outer surface. Mid leg with longer tibial spur 1.4-1.5 times length of the shorter. Hind leg with coxa in profile 1.7—1.9 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.4—2.7 times as long as broad; claws of femalé moderately long, distally abruptly curved, with close long pectination; claws of male similar, but with slightly shorter closer pectination. Gaster slender; tergite 2 in profile at least 5 times as long as posteriorly deep, laterotergite folded under, thyridia oval/elliptical and separated from anterior margin of tergite by at least 4 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing long dense erect hairs; gonosquama quite elongate, distally rounded. Colour generally reddish brown, with gaster (usually) from tergite 3 (rarely from tergites 1 or 2) black; interocellar area black with the black area generally extending between bases of antennae and often onto lower face; antenna black, distally slightly paler, often with scapes brownish; orbits sometimes paler yellowish, and sometimes with face pallid; pterostigma black centrally; wings weakly but evenly infumate. VaRIATION. Although this species is rather uniform in general structure, it is exceptional in having three ‘forms’ of central sclerite. The I morph (Figs 291, 310), the most common form in Costa Rica, has the central sclerite elongate, almost I-shaped, with its longest axis positioned at 90° to Rs+2r. The holotype of Enicospilus antomelas has the sclerite of this pattern. The O morph (Fig. 293) (of which the lectotype of E. monticola is an example) has the central sclerite circular; generally it is very slightly smaller than the I morph. The O morph is also common in Costa Rica. The H morph (of which the lectotypes of E. fuscipennis and E. elegans are examples) has the central sclerite hastate, with an angular side (Fig. 292). This morph is relatively uncommon in Costa Rica, but it becomes more common in South America. For example, in Brazil it constitutes a majority of the material to hand. Some individuals of the H-morph (including the lectotype of E. elegans) have the gaster and the hind legs distal to the trochantellus entirely black. It is questionable whether or not these morphs represent distinct species, but at present I favour the idea that they are conspecific because, despite the fact that they do not seem to intergrade, I can find no other consistent morphological difference between them. Furthermore, all seem to show a rather similar range of variation and all occur sympatrically and synchronously in some sites. Both the I and O morphs have been reared from Gonodonta sp. in Costa Rica The true status of the three morphs is only likely to be resolved by further study of their biology, and it is to facilitate this (and draw attention to the problem) that I have accorded them different epithets. There is some variation in the extent of the black marking on the gaster. Virtually all specimens have tergites 3+ black, but some have tergite 2 and even the posterior part of tergite 1 black. Ihave seen isolated specimens from Venezuela and Colombia with the gaster entirely black and with the hind leg distal to the trochantellus black also. A few individuals may have the interocellar area weakly infuscate and allowances for this have been made in the key. REMARKS. Enicospilus monticola is a rather distinctive species that, with practice, may immediately be recognized by colour alone. It is a richer reddish brown than most other species and has a glossy black gaster (at least tergites 3+), black pterostigma, interocellar area and flagellum. BIOLOGICAL INFORMATION. Enicospilus monticola is a widespread species whose range extends from about 22°N in Mexico to 25°S in Brazil and northern Argentina (Map 25). As yet it has not been collected on any Caribbean island. In Mesoamerica it is one of the more commonly collected species with a black interocel- lar area, and it has been collected in many forested sites from sea-level up to an altitude of about 1800 m. It is frequently taken in Costa Rica, and I have seen specimens from Finca Campana, Finca San Gabriel and San Ramon Forestry Reserve in Alajuela Province; 3 km S. of Casa Mata and Turrialba in Cartago Province; Cerro el Hacha, Estacion Mengo, Rincén de la Vieja and Santa Rosa National Parks in Guanacaste Province; Tortuguero National Park in Lim6én Province; Corcovado National Park, Finca Las Cruces (6 km S. San Vito de Java) and Monteverde Reserve in Puntarenas Province; Braulio Carrillo National Park and San Antonio de Escazt in San José Province. In the best-collected study site, Santa Rosa National Park, Costa Rica, E. monticola has frequently been collected at a light overlooking dry forest. The majority of specimens seem to fly following the start of the rainy season in May. Pooled seasonal distributional data from this site for 1980-87 are: J FM AM Ie eA) Se 4Om Nie D QS Sik 28) 248 220) ele alee 02 In Santa Rosa this species probably has several generations a year as a reared specimen [85.SRNP.396] spun a cocoon on 15 June and emerged 7 July of the same year OPHIONINAE OF TROPICAL MESOAMERICA 235 Map 25 Localities at which Enicospilus monticola has been collected. On the Cerro el Hacha, Costa Rica, in mature dry forest, E. monticola is quite commonly collected between August and November. Although it has never been seen in dry forests from February through April (the very driest months of the year), it has been collected at neighbouring sites (Finca Campana and San Gabriel) and at Monteverde in February and March. Despite intensive collecting on Barro Colorado Island, few specimens of E. monticola have been taken. Two were collected in May 1984 and two more in July 1985. Two specimens have been reared in Santa Rosa National Park. Both parasitized Gonodonta species (Noctuidae) [81.SRNP.712; 85.SRNP.396]. 236 IAN D. GAULD MATERIAL EXAMINED Lectotype 9 (Ophion (Enicospilus) monticola Cameron), Guatemala: Las Mercedes, 1000 m (Cham- pion) (BMNH). Holotype 2 (Henicospilus antomelas Enderlein), Colombia: Rio Magdalena (IZPAN). Lectotype Q (Henicospilus elegans Szépligeti), Brazil: Blumenau (TM); paralectotype, 1 9, same data as lectotype (TM). Lectotype 9 (Henicospilus fuscipennis Szépligeti), Bolivia: Mapiri (TM); paralectotype, 1 9, Brazil: Minas Gerais, 1897 (Fruhstofter) (TM). 82 0,179 9, from the following localities- Argentina (Tucuman); Belize; Bolivia (Cochabamba, Santa Cruz); Brazil (Amazonas, Bahia, Espiritu Santo, Goias, Guanabara, Mato Grosso, Rio de Janeiro, Santa Catarina, Sao Paulo); Colombia (Amazonas, Choco, Meta, Putumayo, Valle); Costa Rica (Alajuela, Cartago, Guanacaste, Limon, Puntarenas and San José Provinces); Ecuador (Esmeraldas, Pichincha); Guatemala; Mexico (Chiapas, Oaxaca, Queretaro, Quintana Roo, Tabasco, Veracruz); Panama (Canal Zone, Chiriqui); Paraguay; Peru (Loreto, Madre de Dios); Surinam; Venezuela (Aragua, Yaracuy, Zulia). (BMNH, CAS, CNC, FSCA, TC, UM, USNM.) Enicospilus madrigalae sp. n. (Figs 187, 294) Description. Mandibles moderately long, evenly narrowed, apically twisted 75—80°; upper mandibular tooth slender, slightly depressed, 1.41.7 times as long as the slightly stouter lower tooth (Fig. 187); outer mandibular surface sparsely pubescent, distally flat and with a weak proximal concavity. Labrum 0.2 times as long as broad; malar space 0.2—0.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin fairly blunt; clypeus in front view 1.3—1.4 times as broad as long, the margin weakly convex apically. Lower face 0.6-0.7 times as broad as long, smooth with very superficial, small punctures. Head in dorsal view with genae strongly constricted behind eye; posterior ocellus contiguous with eye; FI = 60-65% ; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6— 0.7 times the basal mandibular width away from mandible. Antenna slender, with 49-52 flagellar seg- ments; 20th segment 1.9-2.1 times as long as broad. Mesoscutum polished, with extremely fine close punctures, in profile strongly rounded; notauli absent. Mesopleuron polished, the upper part shallowly punctate, the lower part punctostriate to striate; epic- nemial carina curved towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.9 or more of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, anteriorly smooth, posterior striate or wrinkled. Metapleuron weakly convex, rather weakly diagonally striate or punctostriate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile abruptly declivous; anterior transverse carina more or less complete, posterior transverse carina absent; anterior area of moderate length, strongly impressed, striate; spiracular area quite long, virtually smooth; posterior area more or less transversely concentrically striate; lateral longitudinal carina usually present only anteriorly, less frequently represented by discontinuities posteriorly, not joined to spiracular margin by a short carina, or with this carina weak. Fore wing length 12-13 mm; discosubmarginal cell as in Fig. 294; AI = 1.17-1.40; CI = 0.09-0.21; ICI = 0.45-0.61; SDI = 1.14-1.22; cu-a from subopposite to slightly proximal to the base of Rs&M; marginal cell proximally glabrous; 1st subdiscal cell with scattered isolated hairs distally. Hind wing with 5-6 hamuli on R1; 1st and 2nd abscissae of Rs straight. Fore leg with tibia weakly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.7—1.8 times length of the shorter. Hind leg with coxa in profile 1.6-1.7 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.2-2.6 times as long as broad; claws of female moderately long, abruptly but shortly rounded apically, with rather short, close pectinae; claws of male similar but with slightly shorter pectinae. Gaster slender; tergite 2 in profile more than 4.5 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by at least 3 times its own length. Ovipositor slender, its sheath moderately narrow. Male with sternites 7-9 bearing long stout erect hairs; gonosquama distally evenly rounded. Colour generally orange, but with most of head, prothorax and mesothorax laterally paler yellow, mesoscutal vittae brownish; interocellar area black; antenna blackish, distally brown; pterostigma brownish; wings very weakly infumate. VaRIATION. Many specimens have the scutellum and mesoscutal stripes pale yellow. Remarks. This species is named in honour of Liliana Madrigal in recognition of her numerous contribu- tions to Costa Rican conservation biology. OPHIONINAE OF TROPICAL MESOAMERICA 237 Enicospilus madrigalae is most easily recognized by the straight vein Rs+2r in the fore wing, the proximally glabrous marginal cell and the characteristic central sclerite (Fig. 294). It is quite similar to E. xanthocarpus from which it may be separated by the characters given in the key. BIOLOGICAL INFORMATION. Enicospilus madrigalae is a widely distributed species whose range extends from Guatemala south to central Brazil. It is relatively rare in collections, but this may be because the adults seem to have both a short flight season and may only be common in certain years. For example, in Santa Rosa National Park, Costa Rica, only isolated specimens were collected between 1978 and 1983, but in 1984 the species was very common. Only a few individuals were collected in 1985 and 1987, despite the fact that an intensive collecting effort was made at what should have been the height of their season. Seasonal data for 1984 are: A A a OLIN oD - - - - 1 18 1 -'’- - = = The only other Santa Rosa specimens collected are — vi.78 (1), vi.80(1), vii.82(1), vi.85(3) and vi.87(1). Nothing is known of the hosts of E. madrigalae. MATERIAL EXAMINED Holotype 9, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, vi.1984 (Janzen & Hallwachs) (BMNH). Paratypes. Brazil: 1 9, Aguas Vermelhas, 800 m, xii.1983 (Alvarenga) (TC); 5 CO’, 21 9, Bahia, Encruzilhada, 960-980 m, xi. 1972, xi.1973 & xi.1974 (Alvarenga) (TC); 1 &’, Mato Grosso, Sinop, 12°31’°S 55°37'W, x.1974 (Alvarenga) (TC); 3 2, Minas Gerais, Pedra Azul, 800 m, xi.1972 (Alvarenga & Seabra) (TC). Costa Rica: Guanacaste Prov.: 15 0’, 19 2, Santa Rosa National Park, 300 m, vi.1978, vi.1980, vii. 1982, v-vii.1984 & vi.1987 (Janzen & Hallwachs) (BMNH; CNC; MNCR; PANS); 3 Q, same locality, vi.1985 (Gauld) (BMNH). Guatemala: 1 2, Moga Guatalon, 1000 m, iii-iv.1931 (Bequaert) (MCZ). Enicospilus xanthocarpus (Szépligeti) (Figs 188, 189, 295) Henicospilus xanthocarpus Szépligeti, 1906: 146. Holotype 2, BOLIVIA (TM) [examined]. Enicospilus xanthocarpus (Szépligeti) Hooker, 1912: 91. Henicospilus flavosignatus Enderlein, 1921: 34. Holotype 9, ECUADOR (IZPAN) [examined]. Syn. n. Enicospilus flavosignatus (Enderlein) Townes & Townes, 1966: 178. Description. Mandibles moderately long, proximally fairly strongly tapered, distally more parallel-sided, apically twisted 10—15°; upper mandibular tooth slightly depressed, 1.2—1.3 times as long as the lower tooth (Fig. 188); outer mandibular surface slightly concave, with scattered fine pubescence. Labrum 0.2 times as long as broad; malar space 0.3—0.4 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.25—1.35 times as broad as long, with the apical margin weakly convex. Lower face 0.60—0.65 times as broad as long, finely but quite closely punctate, the punctures fusing centrally to give striations. Head in dorsal view with genae strongly constricted posteriorly; posterior ocellus contiguous with eye; FI = 65—75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.7—0.8 times the basal mandibular width away from mandible. Antenna slender, with 57-61 flagellar segments; 20th segment 2.7—2.8 times as long as broad. Mesoscutum polished, closely and finely punctate, in profile strongly rounded; notauli vestigial. Meso- pleuron polished, the upper part finely but closely punctate, the lower part rugose-striate, especially in a band from lower corner forwards to the epicnemial carina (Fig. 189); epicnemial carina curved or inclined towards but not reaching anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for more or less all of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, often irregularly wrinkled, sometimes smooth. Metapleuron weakly convex, punctate, punctostriate or striate; submetapleural carina quite strongly but evenly broadened anteriorly; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area short and strongly impressed, striate; spiracular area quite long, smooth; posterior area centrally more or less concentrically striate, usually also with a median longitudinal ridge, rarely more evenly reticulate; lateral longitudinal carina from complete to present only anteriorly, joined to spiracular margin by a short carina. Fore wing length 13-14 mm; discosubmarginal cell as in Fig. 295; AI = 0.85-1.36; CI = 0.22-0.34; ICI = 0.31-0.48; SDI = 1.17—1.29; cu-a proximal to the base of Rs&M by about 0.2 of its length; marginal 238 IAN D. GAULD cell proximally fairly uniformly hirsute; 1st subdiscal cell with scattered hairs distally. Hind wing with 6 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa almost straight. Fore leg with tibia weakly flattened, with fine scattered spines on outer surface. Mid leg with longer tibial spur 1.6 times length of the shorter. Hind leg with coxa in profile 1.8-1.9 times as long as deep; trochantellus dorsally 0.1—-0.4 times as long as broad; 4th segment of tarsus 2.6—2.7 times as long as broad; claws of female moderately long, evenly rounded apically, with long close pectinae; claws of male similar, but with shorter slightly closer pectination. Gaster slender; tergite 2 in profile more than 6 times as long as posteriorly deep, laterotergite folded under, thyridia oval/elliptical and separated from anterior margin of tergite by about 4-6 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing long stout erect hairs; gonosquama long, distally rounded. Colour generally orange-brown, mesoscutal vittae darker and tergites 4+ of gaster infuscate; interocel- lar area black, and with black markings extending down between antennal bases; antenna black, distally brownish; pterostigma brown; wings hyaline to weakly infumate. VARIATION. This species exhibits a considerable range of morphological variation, and I am not completely certain that the Mesoamerican specimens (which are very similar to the holotype of E. xanthocarpus) are conspecific with the majority of the lowland South American examples (which are structurally like E. flavosignatus). Firstly, the Mesoamerican and a few Brazilian specimens have a fairly long hind trochan- tellus (0.3—0.4 times as long dorsally as broad) but many from further south have a shorter one (0.1) and also a stouter coxa. These specimens often also have a stouter lower tooth of the mandible than do Costa Rican specimens, and some have the mandible more apically twisted (up to about 30°). There is some variation in the size and degree of sclerotization of the central sclerite. In most Costa Rican specimens it is moderately sclerotized and medium-sized, but specimens from further south tend to have this sclerite slightly smaller and more strongly sclerotized. Typical examples of E. xanthocarpus have the metapleuron punctostriate to striate, but some Brazilian material and the holotype of EF. flavosignatus have the matapleuron regularly punctate. However, except for these differences all the specimens are very similar to each other, and as the variation seems to be more or less continuous, I suggest that for the present flavosignatus is treated as a synonym of xanthocarpus. Remarks. Enicospilus xanthocarpus is rather similar and probably closely related to E. kleini. The two have similar wings and alar sclerites, though the central sclerite of E. kleini is usually somewhat curved. The most obvious differences between these two species is in coloration: the antennae, interocellar area and terminal gastral segments of E. xanthocarpus are blackish whereas in E. kleini they are yellowish brown. The slightly impressed, rugose/striate area present on the mesopleuron of EF. xanthocarpus is absent in E. kleini, and the Costa Rican specimens of E. kleini have a short hind trochantellus, whereas that of Costa Rican examples of E. xanthocarpus is long. BIOLOGICAL INFORMATION. Enicospilus xanthocarpus is a widely distributed species whose range extends from northern Costa Rica south into Paraguay and southern Brazil. Although it is a relatively commonly collected species in South America, it is much more rarely taken in Costa Rica where it has only been collected in Santa Rosa National Park. Isolated specimens have been collected at this site from the middle of the wet season to early dry season, between July and January. Nothing is known of the host range of this species. MATERIAL EXAMINED Holotype @ (Henicospilus xanthocarpus Szépligeti), Bolivia: Mapiri (TM). Holotype 9 (Henicospilus flavosignatus Enderlein), Ecuador: Bucay (IZPAN). Long trochantellar morph material. Brazil: 1 0’, 3 9, Aguas Vermelhas, 800 m, xii.1983 (Alvarenga) (BMNH). Costa Rica: Guanacaste Prov.: 6 9, Santa Rosa National Park, 300 m, vii, viii, x & xi.1982, i.1984 (Janzen & Hallwachs) (BMNH). Mexico: Chiapas: 1 2, Muste, near Huixtla, 440 m, ix.1970 (Welling) (CNC). Short trochantellar morph material. 29 0’, 56 9, from — Brazil (Amazonas, Bahia, Goias, Guanabara, Mato Grosso, Minas Gerais, Para, Rio de Janeiro, Santa Catarina, SAo Paulo); Colombia (Amazonas, Magdalena, Valle); Panama; Paraguay; Peru (Loreto); Surinam. Enicospilus duckworthi sp. n. (Figs 185, 190, 296) Description. Mandibles moderately long, proximally strongly tapered, distally more weakly narrowed, apically twisted 30—-40°; upper mandibular tooth cylindrical, 1.3—1.5 times as long as the lower tooth; outer OPHIONINAE OF TROPICAL MESOAMERICA 239 mandibular surface flat with scattered fine hairs basally. Labrum 0.3 times as long as broad; malar space 0.3—0.4 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.5—1.6 times as broad as long, with apical margin weakly convex. Lower face 0.74—0.76 times as broad as long, punctate, centrally tending to striate. Head in dorsal view with genae rounded behind eyes; posterior ocellus close to but not touching eye; FI = 55-60%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.9 times the basal mandibular width away from mandible. Antenna long and slender, with 50-52 flagellar segments; 20th segment 2.2 times as long as broad. Mesoscutum polished, finely punctate, in profile evenly rounded; notauli vestigial. Mesopleuron pol- ished, the upper and lower parts finely punctate, ventrally sometimes punctostriate; epicnemial carina inclined towards anterior margin of pleuron (Fig. 185), its upper end usually evanescent. Scutellum in profile weakly convex, laterally carinate for all of its length; scutellum in dorsal view 1.8 times as long as anteriorly broad, anteriorly punctate, posteriorly slightly wrinkled. Metapleuron rather weakly convex, with coarse shallow punctures that tend to coalesce irregularly so in some individuals the metapleuron is irregularly wrinkled (Fig. 190); submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area short, strongly impressed, slightly rugose; spiracular area moderately long, smooth; posterior area finely reticulate; lateral longitudinal carina present anteriorly but weak, not joined to spiracular margin by a short carina. Fore wing length 14-16 mm; discosubmarginal cell as in Fig. 296; AI = 0.87-1.00; CI = 0.32-0.40; ICI = 0.53-0.57; SDI = 1.09-1.15; cu-a opposite or proximal to the base of Rs&M by less than its own thickness; marginal cell proximally narrowly glabrous to evenly hirsute; 1st subdiscal cell centrally and distally hirsute. Hind wing with 6-7 hamuli on R1; 1st abscissa of Rs weakly curved, 2nd abscissa virtually straight. Fore leg with tibia slightly flattened, with scattered fine spines on outer surface. Mid leg with longer tibial spur 1.5—1.7 times length of the shorter. Hind leg with coxa in profile 1.7—-1.8 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2. 1—2.2 times as long as broad; claws of female quite long and evenly curved, with close stout pectinae. Gaster long and slender; tergite 2 in profile more than 6 times as long as posteriorly deep, laterotergite up-turned, thyridia elliptical and separated from anterior margin of tergite by about 3 or more times its own length. Ovipositor moderately stout, its sheath slender. Male unknown. Colour generally orange-brown with tergites 3+ of gaster black, gastral segments 3 and 4 ventrally pallid; interocellar area black; antenna blackish; pterostigma dark brown; wings weakly infumate. VARIATION. The holotype is slightly darker in colour than any of the paratypes, but structurally all are very alike. REMARKS. This species is named in honour of Donald Duckworth in recognition of his many contributions to tropical entomology. Enicospilus duckworthiis a very distinctive species on account of its characteristic alar sclerites (see Fig. 296). The second discal cell in the fore wing is slightly deeper and 1m-cu seems more sinuate than other species which key out near this one. The phylogenetic relationships of this species are unclear. BIOLOGICAL INFORMATION. Enicospilus duckworthi is a widely distributed species whose range extends from Panama south to Bolivia. It has not been collected in the lowlands of eastern South America, and from the very limited data to hand, it seems to occur in mid altitude forests between 1100 and 2200 m. The hosts of this species are unknown. MATERIAL EXAMINED Holotype 9, Panama: Cerro Campana, nr Chica, iv.1965 (Duckworth & Duckworth) (USNM). Paratypes. Bolivia: 1 9, Chapare, Alto Palmar, 1100 m, ii.1961 (CNC); 1 2, Chapare, El Limbo, 2200 m, i.1962 (CNC). Colombia: 1 9, Cundinamarca, Monteredondo, iv.1961 (Foerster) (TC). Enicospilus devriesi sp. n. (Fig. 297) [Henicospilus trimaculatus (Taschenberg); Morley, 1912: 35, in part. Misidentification. | Description. Mandibles quite long, rather evenly narrowed proximally and somewhat parallel-sided distally, apically twisted 10—20°; upper mandibular tooth subcylindrical, 1.3—1.4 times as long as the lower tooth; outer mandibular surface centrally rather flat, without a discernible proximal concavity. Labrum 0.2-0.3 times as long as broad; malar space 0.30.4 times as long as basal mandibular width. Clypeus in 240 IAN D. GAULD profile moderately convex, margin inturned, usually very narrowly impressed; clypeus in front view 1.4— 1.6 times as broad as long, margin weakly convex. Lower face 0.65—0.70 times as broad as long, usually polished, with scattered weak punctures. Head in dorsal view with genae constricted behind the eyes; posterior ocellus very close to the eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.9 times the basal mandibular width away from mandible. Antenna long and slender with 59-62 flagellar segments; 20th segment 2.0—2.3 times as long as broad. Mesoscutum polished, with shallow sparse punctures, in profile steeply rounded anteriorly; notauli vestigial. Mesopleuron highly polished, the upper part with fine inconspicuous scattered punctures, the lower part similar though sometimes smoother; epicnemial carina very strong and often crenulate behind, curved towards anterior margin of pleuron, usually with upper end reaching a small scabrous patch. Scutellum in profile weakly convex, laterally carinate for anterior 0.2—0.4 of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, relatively smooth. Metapleuron polished, smooth except for minute scattered punctures; submetapleural carina barely broadened anteriorly; posterior transverse carina of mesosternum complete. Propodeum in profile quite long, evenly declivous; anterior transverse carina complete, posterior transverse carina sometimes represented by lateral vestiges; anterior area moderately long, shallow, striate; spiracular area long, smooth; posterior area from irregularly reticulate to relatively smooth, somewhat alutaceous, often with a weak median longitudinal ridge; lateral longitudi- nal carina generally complete, joined to spiracular margin by a short carina. Fore wing length 13-17 mm; discosubmarginal cell as in Fig. 297; AI = 0.42-0.68; CI = 0.29-0.39; ICI = 0.33-0.45; SDI = 1.20-1.33; cu-a proximal to the base of Rs&M; marginal cell proximally uniformly hirsute; 1st subdiscal cell with anterior and distal parts extensively hirsute. Hind wing with 6-7 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa from almost straight to slightly sinuous. Fore leg with tibia slightly flattened, with scattered fine spines on outer surface. Mid leg with longer tibial spur 1.2-1.4 times length of the shorter. Hind leg with coxa in profile 1.9-2.1 times as long as deep; trochantellus dorsally 0.3—0.6 times as long as broad; 4th segment of tarsus 2.2—2.4 times as long as broad; claws of female quite large, with moderately short, fairly closely spaced pectinae, those of male similar. Gaster slender; tergite 2 in profile at least 5 times as long as posteriorly deep, laterotergite upturned, thyridia elliptical and separated from anterior margin of tergite by about 4 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing long, relatively fine, semierect pubescence; gonosquama evenly rounded. Colour generally pale yellowish, with mesoscutal vittae, mesosternum, anterior part of propodeum infuscate; gaster pale yellow with hind part of tergites 1-3 and most of tergites 4+ infuscate so that anterior part of gaster appears to be banded; interocellar area yellowish; antenna golden; pterostigma yellow; wings hyaline. VARIATION. There is some variation in the degree of integumental infuscation. Many specimens have the lower part of the mesopleuron extensively infuscate; some also have the hind coxa blackish brown. REMARKS. This species is named in honour of Phil DeVries in recognition for his work on the Costa Rican butterfly fauna. Enicospilus devriesi is one of the most easily recognized species of Enicospilus as it is the only one in ‘Central America with two central sclerites (Fig. 297). It is also unusual in having very reduced lateral longitudinal scutellar carinae, and its colour pattern is quite distinctive. E. devriesi is probably closely related to E. flavoscutellatus as both species have similar sculpture, venation and mandibles BIOLOGICAL INFORMATION. Enicospilus devriesi is a widely distributed species whose range extends from about 24°N in Mexico south through Central America and northern South America to Ecuador (3°S) (Map 26). Itis relatively frequently collected in cloud forests at altitudes between 1200 and 2500 m, and has never been collected below 800 m. E. devriesi seems to gradually replace E. flavoscutellatus at altitudes above 2000 m, whilst at intermediate elevations E. flavoscutellatus predominates (e.g. at Monteverde, 1350 m). In Panama, at Guadalupe Arriba (2300 m), E. devriesi is the most commonly collected species of Enicospilus and accounts for about 85% of all specimens of the genus collected at light. In such habitats it occurs with several species of Ophion, Sicophion fenestralis and Janzophion nebosus. Its monthly occur- rence at Guadalupe Arriba for 1984-5 is: Jon FeiecM. @Ay (Mito: sy wAveSam, Oe NiegeD) =e ott ah Grea OS ell © F2dcee Lge cael 241 OPHIONINAE OF TROPICAL MESOAMERICA ‘pajsa]]oo usaq sey isazaap snjidsoouy YoY ye sanesoT 97 dey = ~~~ ee eee 242 IAN D. GAULD E. devriesi is comparatively less common at Monteverde, Costa Rica (1350 m). The cumulative seasonal distribution for 1962-86 is: J CB a Ba, Nd ee Se OL ING, Sp LL de wile Ai i Lire etn Geen! ieee It is not known which insects serve as hosts for this species. MATERIAL EXAMINED Holotype 9, Costa Rica: Alajuela Prov., San Ramon Reserve, Rio San Lorencito, 800 m, ii.1987 (Chacon & Chacon) (BMNH). Paratypes. Colombia: 1 9, Cundinamarca, Monterredondo, iv.1961 (Foerster) (CNC). Costa Rica: Alajuela Prov.: 1 0’, San Ramon Forestry Reserve, 5 km N. of Col. Palmarena, 900 m, v.1985 (Chacon) (BMNH); 2 o’, 3 9, Volcan Pods National Park, 2350 m, xii.1982 (Janzen & Hallwachs) (BMNH): Puntarenas Prov.: 1 0’, 13 2, Monteverde, 1350 m, i-iii.1962 (Palmer) (TC); 1 9, same locality, ii.1984 (Cameron) (WC); 2 9, same locality, iv-v.1984 (Gauld) (BMNH); 1 9, same locality, vi.1984 (Fogden) (BMNH); 2 0’, 2 2, same locality, xi.1985, i, v, viii.1986 (Haber) (BMNH; MNCR); 3 9, same locality, i.1986 (Forsyth) (CNC): San José Prov.: 1 0’, 4 9, San Gerardo de Dota, Cerro de la Muerte, 2430 m, xii.1981 (Janzen & Hallwachs) (BMNH); 1 @, San Antonio de Escazt, 1300 m, v-vi.1981 (Eberhard) (UMC); 1 9, San José, i.1961 (Palmer) (TC). Ecuador: 1 2, N. Perucho (Ota), 2000 m, i.1971 (Pena) (TC). El Salvador: 1 9, Cerro Verde, 2000 m, v.1971 (Howden) (TC). Guatemala: 1 ?, Solola, Pana- jachel, 1550 m, iii.1955 (Stuart) (UMC). Mexico: Chiapas: 2 2, 35 km NE. Huixtla, 1000 m vi.1969 (Teskey & Mason) (CNC); 10’, 4 9, San Cristobal de las Casas, 2400 m, vi-vii.1969 (Bright & Campbell) (CNC); 1 ©, Yerba Buena, 32 km N. Bochil, 1900 m, vi.1969 (CNC): Durango: 3 9, 48 km W. La Cuidad, 2200 m, vii.1964 (Mason) (CNC): Guerrero: 1 2, Amula, 2000 m, viii.1904 (Smith) (BMNH); 2 2, Chilpancingo, 1500 m, vi.1904 (Smith) (BMNH): Hidalgo: 1 2, Jacala, 1500 m, vii.1939 (Haag) (MCZ): Veracruz: 10’, Atoyac, v.1904 (Smith) (BMNH). Panama: 1 9, Chiriqui, Fortuna, 1050 m, iv.1978 (Wolda) (RNH); 12 Oo’, 16 Q, Chiriqui, Guadalupe Arriba, 2300 m, dates as above (Wolda) (BMNH). Venezuela: 1 9, Aragua, Rancho Grande, iv.1960 (Test) (UMC). Enicospilus donahuei sp. n. (Fig. 298) [Ophion (Enicospilus) flavo-scutellatus Brullé; Cameron, 1886. Misidentification. | DEscrIPTION. Mandibles moderately long, weakly narrowed, apically twisted 10—20°; upper mandibular tooth subcylindrical, 1.2—1.4 times as long as the lower tooth; outer mandibular surface flat, with scattered long hairs. Labrum 0.3 times as long as broad; malar space 0.3—0.4 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.45—1.55 times as broad as long, with margin weakly convex. Lower face 0.70—0.75 times as broad as long, weakly polished, centrally finely and quite closely punctate. Head in dorsal view with genae constricted behind eyes; posterior ocellus con- tiguous with eye; FI = 68-73%; occipital carina mediodorsally complete, ventrally curved to join hyposto- mal carina about 0.8—0.9 times the basal mandibular width away from mandible. Antenna long but rather stout, with 64-70 flagellar segments; 20th segment 1.82.1 times as long as broad. Mesoscutum weakly polished, with very fine punctation, in profile steeply rounded; notauli vestigial. Mesopleuron moderately polished, the upper part finely and closely punctate, the lower part similar but with punctures generally weaker, and usually with weak coriaceous sculpture centrally; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.6 or more of its length; scutellum in dorsal view 1.4-1.5 times as long as anteriorly broad, finely punctate. Metapleuron finely but rather weakly punctate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina represented laterally by vestiges; anterior area long, shallow, with scattered striae; spiracular area smooth with some fine punctures; posterior area weakly irregularly wrinkled, with a median longitudinal wrinkle; lateral longitudinal carina complete, joined to spiracular margin by a weak short carina. Fore wing length 19-20 mm; discosubmarginal cell as in Fig. 298; AI = 0.36-0.65; CI = 0.30-0.40; ICI = 0.39-0.51; SDI = 1.01—-1.11; cu-a proximal to the base of Rs&M; marginal cell proximally uniformly hirsute; 1st subdiscal cell with anterior and distal parts densely hirsute. Hind wing with 7-11 hamuli on R1; 1st abscissa of Rs almost straight, 2nd abscissa weakly arcuate. OPHIONINAE OF TROPICAL MESOAMERICA 243 Fore leg with tibia noticeably flattened, with numerous spines on outer surface. Mid leg with longer tibial spur 1.3-1.4 times length of the shorter. Hind leg with coxa in profile 1.8-1.9 times as long as deep; trochantellus dorsally 0.2 times as long as broad; 4th segment of tarsus 1.7—2.0 times as long as broad; claws of female large, with close, short, slightly inwardly inclined pectinae, those of male similar. Gaster quite slender; tergite 2 in profile at least 6 times as long as posteriorly deep, laterotergite upturned, thyridia large, oval/elliptical and separated from anterior margin of tergite by about 3 or more times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-8 bearing long stout erect hairs, which appear to be arranged in two subterminal clusters; gonosquama evenly rounded. Colour generally yellowish, with mesoscutal vittae and posterior and dorsal parts of gastral tergites infuscate; interocellar area yellowish; antenna golden; pterostigma yellowish brown; wings very weakly infumate. VaRIATION. The specimens from Mexico are paler yellow with darker markings on the mesopleuron, mesosternum and propodeum. One individual also has a much larger alar sclerite than the other speci- mens, but otherwise it appears to be conspecific. REMARKS. This species is named in honour of Julian Donahue for his enthusiastic support of Costa Rican conservation plans. Enicospilus donahuei is easily recognized because it has part of the fenestra, inside the distal sclerite, bearing a wide band of fine pubescence (Fig. 298). Only one other species, E. fosteri, has part of the fenestra hirsute, and in that case it is the anterior margin of the fenestra, adjacent to Rs+2r that is hairy (Fig. 303). The phylogenetic affinities of this species are not known, but in colour and sculpture of the propodeum it is similar to E. devriesi. Possibly this is convergence because both species inhabit similar areas, and several cloud-forest species in a variety of localities have banded gasters and relatively smooth alitrunks. BIOLOGICAL INFORMATION. Enicospilus donahuei is only known to occur in Central America, from about 18°N in southern Mexico south to Costa Rica. It is a relatively rarely collected species, and most specimens have been taken in cloud forests at moderately high altitudes (1200-2400 m). Despite intensive collecting it has never been taken in lowland forests. MATERIAL EXAMINED Holotype 9, Costa Rica: Alajuela Prov., 8.2 km N. Vara Blanca, iv.1984 (Janzen & Hallwachs) (BMNH). Paratypes. Costa Rica: Puntarenas Prov.: 1 0’, Monteverde, 1350 m, vii.1982 (Janzen & Hallwachs) (BMNH); 1 9, same locality and collectors, 1500 m, i.1984 (MNCR); 1 0’, same locality, vi.1985 (Gauld) (BMNH). Unknown Province: 1 9, 14kmN. Urena, vi.1974 (Donahue) (TC). Guatemala: 1 0’, Cerro Zunil, 1700 m (Champion) (BMNH). Mexico: Chiapas: 1 2, San Cristobal de las Casas, 2400 m, vi.1969 (Peterson) (CNC); 1 &’, Yerba Buena, 32 km N. Bochil, v.1969 (Mason) (CNC): Oaxaca: 1 2, Mpio Comaltepec, Vista Hermosa, 1450 m, ix.1962 (CNC); 1 9, 51 km S. Valle Nacional, 2300 m, v.1971 (Howden & Howden) (TC). Enicospilus simoni sp. n. (Figs 192, 299) Description. Mandibles moderately long, very weakly narrowed, apically twisted 80—90°; upper mandibu- lar tooth strongly compressed, 0.9-1.1 times as long as the lower tooth (Fig. 192); outer mandibular surface flat, finely punctate. Labrum 0.2-0.3 times as long as broad; malar space 0.3—0.4 times as long as basal mandibular width. Clypeus in profile weakly convex, margin fairly sharp; clypeus in front view 1.5—1.6 times as broad as long, with margin weakly convex. Lower face 0.70—0.75 times as broad as long, polished, centrally punctate. Head in dorsal view with genae rounded, sometimes slightly inflated; posterior ocellus very close to eye; FI = 65-70%; occipital carina mediodorsally weak, obsolescent or incomplete, if complete then usually dipped, ventrally curved to join hypostomal carina about 0.7—0.8 times the basal mandibular width away from mandible. Antenna long and relatively slender, with 66-70 flagellar seg- ments; 20th segment 1.9-2.0 times as long as broad. Mesoscutum in profile evenly rounded; notauli shallow but discernible. Mesopleuron polished, the upper part closely and finely punctate, the lower part similar but also wrinkled; epicnemial carina with upper end curved towards but not reaching anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.7 or more of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, sparsely punctate. Metapleuron convex, coarsely wrinkled-rugose; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile 244 IAN D. GAULD abruptly declivous; anterior transverse carina complete, posterior transverse carina vestigial or absent; anterior area short, deeply impressed, striate; spiracular area short, relatively smooth; posterior area reticulate to coarsely rugose; lateral longitudinal carina sometimes complete, usually always present anteriorly, joined to spiracular margin by a very short carina. Fore wing length 19-26 mm; discosubmarginal cell as in Fig. 299; AI = 0.70-0.75; CI = 0.19-0.35; ICI = 0.84-1.05; SDI = 1.05—1.25; cu-a proximal to the base of Rs&M; marginal cell proximally fairly evenly hirsute, sometimes slightly more sparsely so adjacent to Rs+2r; 1st subdiscal cell with anterior 0.4 and distal corner hirsute. Hind wing with 7-11 hamuli on R1; 1st and 2nd abscissae of Rs more or less straight. Fore leg with tibia quite strongly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.3—1.6 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally 0.1—-0.2 times as long as broad; 4th segment of tarsus 2.2—2.4 times as long as broad; claws of female with close, rather short pectinae, those of male similar but with pectinae closer and slightly shorter. Gaster slender; tergite 2 in profile at least 5 times as long as posteriorly deep, laterotergite upturned, thyridia oval to obovate and separated from anterior margin of tergite by about 4-5 times its own length. Ovipositor slender, its sheath moderately narrow. Male with sternites 7-9 bearing numerous long stout erect hairs; gonosquama moderately long, usually with a weak dorsal subapical notch. Colour generally yellowish brown, usually with dark brown mesoscutal vittae; interocellar area yellow- ish; antenna black, distally dark brown; pterostigma orange-brown; wings hyaline. VARIATION. This is a morphologically rather uniform species over its extensive geographical range. There is some variation in the degree of pigmentation of the distal sclerite. It varies from complete, confluent with the proximal sclerite, to being virtually undiscernible. There is a little variation in the size of the central sclerite, but it is always quite small. As is usual for large species there is considerable variation in the sculpture of the posterior part of the propodeum. Specimens with coarsely rugose propodea generally have the posterior transverse carina present as a pair of lateral crests. REMARKS. This species is named after Simon Harrison, in recognition of his enthusiastic volunteer efforts in Guanacaste National Park, Costa Rica. Enicospilus simoni is a robust, large species that can most easily be recognized by the characteristic form of the mandibles (Fig. 192); they are much stouter than the similarly strongly twisted mandibles that some other Central American species possess. E. simoni also has an unusually large value for ICI in the fore wing. The phylogenetic affinities of this species are not known. BIOLOGICAL INFORMATION. Enicospilus simoni is an extremely widespread species whose range extends from Sinaloa and San Luis Potosi in Central Mexico (23—24°N) south to Paraguay (25°S). It is present on Trinidad, but does not seem to occur on the more isolated Caribbean islands (Map 27). E. simoni has been collected at a variety of altitudes from almost sea-level (on Barro Colorado Island) up to 2300 m in Guerrero, Mexico. In Santa Rosa National Park it has two peaks of abundance, at the begining and end of the wet season. The pooled data for 1977-1986 are: i MICAS Ne ee. SO Nay 1 .= = = ple — Mee 2 3) oS It is not known what species serve as hosts for E. simoni. MATERIAL EXAMINED Holotype ©’, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, xi.1982 (Janzen & Hallwachs) (BMNH). Paratypes. Brazil: 2 O’, Santa Catarina, Nova Teutonia, ii.1937, ii.1939 (Plaumann) (BMNH); 1 9, same locality and collector, v.1943 (TC); 1 0’, 1 2, same locality and collector, viii & xii.1952 (TC); 1 , same locality and collector, ix.1970 (TC); 1 2, Sao Paulo State, Casa Grande, Boraceia Field Stn, i.1975 (Rodgers) (TC). Costa Rica: Alajuela Prov.: 1 0’, 1 9, San Ramon Reserve, Rio San Lorencito, 800 m, v.1987 (Chacon) (BMNH): Cartago Prov.: 1 Oo’, Turrialba, 1000 m, vii.1965 (Real) (CAS): Guanacaste Prov.: 1 &', 1 9, Santa Rosa National Park, 300 m, xi-xii.1977 (Janzen)(TC); 1 CO’, same locality and collector, xii.1979 (TC); 1 2, same locality xii.1980 (Janzen & Hallwachs) (TC); same locality and collectors, 1 9, vi.1981; 1 9, x.1982; 2 0’, xi.1982; 107, 1 9, ix.1983; 1 0’, 1.1984; 1 9, v.1984; 1 9, vi.1984 (BMNH; MNCR); 2 9, same locality, vi.1985 (Gauld) (BMNH); 2 9, same locality and collector, v-vi.1986 (BMNH); 2 2, Volcan Cacao, Estacion Mengo, 1100 m, vii-ix.1987 (Janzen) (BMNH); 1 9, Volcan Orosi, Hda Mariksa [= Maritza], 550 m, i.1986 (Hallwachs & Janzen) (BMNH): Puntarenas Prov.: 4 2, Monteverde, 1350 m, xii.1961, i.1962 (Palmer) (TC); 1 9, same locality, v.1980 (Janzen & Hallwachs) (BMNH); 3 o’, 7 @, same locality, i-iti.1986 (Forsyth) (CNC): San José Prov.: 2 9, Estacion Carrillo, OPHIONINAE OF TROPICAL MESOAMERICA 245 Map 27 Localities at which Enicospilus simoni has been collected. Braulio Carrillo National Park, 700 m, iii & v.1985 (Chacon) (BMNH). Ecuador: 1 2, Abitagua, 1000 m, x.1939 (MacIntyre) (TC). El Salvador: 1 ©’, Quezaltepeque, 500 m, viii.1963 (Cavagnaro & Irwin) (CAS). Mexico: Chiapas: 7 2, 32 km N. Huixtla, 1000 m, vi.1969 (Peterson) (CNC): Guerrero: 3 GC’, 2 9, Xucumanatlan, 2300 m, vii.1904 (Smith) (BMNH): San Luis Potosi: 1 co’, El Naranjo, 24 km W. Nuevo Morelos, x.1962 (Townes) (TC): Sinaloa: 1 9, Porterillos, 24 km W. El Palmito, 1700 m, vii.1964 (McAlpine) (CNC). Panama: 1 ©’, Barro Colorado Island, vii.1963 (Cavagnaro & Irwin) (CAS); same locality, 2 Q, viii.1983; 1 Q, xii.1983; 1 2, ix.1984; 1 2, vi.1985 (Wolda) (BMNH); 1 9, Chiriqui, Fortuna, 1050 m, iv.1978 (Wolda) (RNH); 3 2, Las Cumbres, 150 m, iv-v, 1983-85 (Wolda) (BMNH). Paraguay: 1, Sapucay, 1904 (Foster) (BMNH). Trinidad: 1 9 , Blanchisseuse Ward, Morne Bleu, 18 km N. Simla, vi.1977 (Grissell) (USNM). 246 IAN D. GAULD Enicospilus orosii sp. n. (Figs 191, 300) Description. Mandibles unusually short and stout, very strongly tapered, apically twisted outwards 10—-15° so that upper tooth is forwards (Fig. 191); upper mandibular tooth cylindrical, 2-3 times as long as the lower tooth; outer mandibular surface rather flat, with short pubescence. Labrum 0.2-0.3 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile quite strongly convex, margin impressed, acute; clypeus in front view 1.3-1.6 times as broad as long, margin apically truncate. Lower face 0.70—0.75 times as broad as long, polished, finely punctate. Head in dorsal view with genae slightly inflated, rounded behind eyes; posterior ocellus contiguous with eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.8 times the basal mandibular width away from mandible. Antenna long and slender, with 56-60 flagellar segments; 20th segment 1.7-2.3 times as long as broad. Mesoscutum weakly polished, with obsolescent punctation, in profile evenly rounded; notauli weak but discernible. Mesopleuron polished, the upper part finely punctate, the lower part striate, wrinkled; epicnemial carina inclined towards anterior margin of pleuron, its posterior edge often crenulate. Scutellum in profile weakly convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.4-1.6 times as long as anteriorly broad, punctate, posteriorly with a few longitudinal wrinkles. Meta- pleuron strongly convex, anteroventrally smooth, posterodorsally rugose, very strongly so in large speci- mens; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile abruptly declivous; anterior transverse carina complete, posterior trans- verse carina absent, or in larger specimens represented by lateral crests; anterior area rather short, deeply impressed, striate; spiracular area short, smooth; posterior area from reticulate to coarsely irregularly rugose, often with a median longitudinal ridge; lateral longitudinal carina usually complete, not joined to spiracular margin by a short carina. Fore wing length (16) 18-24 mm; discosubmarginal cell as in Fig. 300; AI = 0.51-0.79; CI = 0.41-0.54; ICI = 0.51-0.61; SDI = 1.20-1.35; cu-a proximal to the base of Rs&M; marginal cell proximally more sparsely hirsute than it is centrally; 1st subdiscal cell with distal 0.8 almost entirely hirsute. Hind wing with 7-9 hamuli on R1; 1st abscissa of Rs weakly bowed, 2nd abscissa slightly sinuous. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.4-1.5 times length of the shorter. Hind leg with coxa in profile 1.7—-1.8 times as long as deep; trochantellus dorsally 0.2 times as long as broad; 4th segment of tarsus 2. 1—2.4 times as long as broad; claws of female with long close pectinae, those of male similar but with pectinae shorter. Gaster moderately slender; tergite 2 in profile at least 4 times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by about 3 or more times its own length. Ovipositor slender, its sheath moderately narrow. Male with sternites 7-9 bearing long stout erect hairs; gonosquama evenly rounded. Colour generally yellowish brown, with tergites 3+ of gaster infuscate; mesoscutal vittae dark brown; interocellar area yellow; antenna proximally blackish, distally brownish; pterostigma brownish; wings almost hyaline. VaRIATION. Enicospilus orosii is morphologically a rather uniform species except for variation in the sculpture of the lateral part of the alitrunk and the dorsal region of the propodeum, as outlined above. Specimens from Mexico are more yellowish than yellowish brown, but they have dark markings like other specimens. The single male from Florida is virtually entirely pale yellow, with only the antennae infuscate. This specimen is also unusually small, with a fore wing length of only 16 mm. The specimen from Monteverde, which is also the largest individual, has the central sclerite about proportionately half as large as other individuals. RemMaRKS. Enicospilus orosii may easily be recognized by the form of the mandibles (Fig. 191). No other Central American species has them twisted so the upper tooth is outermost, or has them so stout and strongly tapered. The phylogenetic relationships of this species are unclear. BIOLOGICAL INFORMATION. Enicospilus orosii is a widely distributed species whose range extends from tropical Mexico south to Guyana. I have seen a single male that was collected in Fort Myers, Florida, but apart from this there are no other records from Florida or from the Caribbean. In Costa Rica and Panama E. orosii is associated with wet forests from sea level up to about 1400 m. In Guanacaste Province, Costa Rica, it has been collected in the moister forest on the lower slopes of Volcan Orosi, but despite intensive collecting, it has never been taken in the nearby seasonally dry forests of Santa Rosa National Park. E. orosiiis generally very seldom collected even in sites where it does occur, and there has been relatively OPHIONINAE OF TROPICAL MESOAMERICA 247 intensive collecting. For example, only a single specimen has been taken at Monteverde, and several years’ light-trapping on Barro Colorado Island have yielded seven individuals. The majority of the Barro Colorado specimens were collected either in May or in October. Nothing is known about the biology. MATERIAL EXAMINED Holotype ©’, Costa Rica: Guanacaste Prov., Volcan Orosi, Casa Mariksa [= Maritza], 650 m, vii. 1986 (Gauld) (BMNH). Paratypes. Costa Rica: Puntarenas Prov.: 1 2, Monteverde, 1350 m, xi.1985 (Haber) (BMNH). Guyana: 1 O’, Essequibo River, Morabali Ck, viii-x.1929 (BMNH). Mexico: Guerrero: 2 0’, Amula, 2000 m, ix.1904 (Smith) (BMNH): Morelos: 1 2, Cuernavaca, vi.1904 (Smith) (BMNH). Panama: 1 9, Barro Colorado Island, x.1982 (Wolda) (TC); 1 CO’, same locality and collector, v.1983 (TC); same locality, 1 9, v.1983; 2 2, x.1983; 1 9, v.1984; vii.1985 (Wolda) (BMNH). U.S.A.: Florida: 1 0’, Fort Myers, vi.1968 (Heinrich) (CNC). Venezuela: 1 9, San Esteban, nr Puerto Cabello, xii.1939 (Anduze) (TC). Enicospilus bima sp. n. (Figs 193, 194, 301) Description. Mandibles moderately long, proximally rather strongly narrowed, distally almost parallel- sided, apically twisted 45—60°; upper mandibular tooth slender, subcylindrical to slightly depressed, 1.6— 2.0 times as long as the lower tooth; outer mandibular surface with a weak median longitudinal concavity that bears scattered hairs, and proximally is confluent with a weak basal depression. Labrum rather inflated, 0.3 times as long as broad; malar space 0.3—-0.4 times as long as basal mandibular width. Clypeus in profile out-flared, with a sharp protuberant subapical margin, true margin more or less concealed; clypeus in front view 1.1—1.2 times as broad as long, with margin deeply U-shaped apically (Fig. 193). Lower face 0.70-0.76 times as broad as long, centrally punctate, often with fine striae. Head in dorsal view with genae rounded; posterior ocellus close to eye; FI = 67-73%; occipital carina mediodorsally complete, ventrally curved to approach but not actually join the hypostomal carina about 1.0 times the basal mandibular width away from mandible. Antenna slender, with 55—57 flagellar segments; 20th segment 1.8—2.0 times as long as broad. Mesoscutum polished, with obsolescent punctures, in profile evenly rounded and with margin out- turned; notauli vestigial. Mesopleuron weakly polished, the upper part finely punctate with weak striae, the lower part punctostriate to finely striate; epicnemial carina absent above lower corner of mesopleuron (Fig. 194). Scutellum in profile moderately convex, laterally carinate for virtually all of its length; scutellum in dorsal view 1.4—1.5 times as long as anteriorly broad, anteriorly punctate, posteriorly with longitudinal wrinkles. Metapleuron moderately convex, finely and rather irregularly striate to coriaceous; sub- metapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum broadly incomplete centrally. Propodeum in profile abruptly declivous; anterior transverse carina complete, strong, posterior transverse carina represented laterally by strong blunt prominences; anterior area short, strongly impressed, often almost smooth; spiracular area short, more or less smooth; posterior area generally weakly coriaceous or with fine irregular sculpture, and with a weak median longitudinal ridge; lateral longitudinal carina only represented by an anterior vestige, which is usually joined to spiracular margin by a short carina. Fore wing length 14-18 mm; discosubmarginal cell as in Fig. 301; AI = 0.64-0.73; CI = 0.21-0.27; ICI = 0.88-1.17; SDI = 1.07—-1.20; cu-a from opposite to the base of Rs&M to distal to it by about 0.1 times its own length; marginal cell proximally uniformly hirsute; 1st subdiscal cell with anterior and distal margins broadly hirsute. Hind wing with 6-9 hamuli on R1; 1st abscissa of Rs more or less straight, 2nd abscissa very weakly bowed. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.6-1.7 times length of the shorter. Hind leg with coxa in profile 1.8-1.9 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.0—2.2 times as long as broad; claws of female abruptly curved, with stout regularly spaced pectinae; those of male similar but with pectinae shorter. Gaster moderately slender; tergite 2 in profile 2.6-3.0 times as long as posteriorly deep, laterotergite folded under, thyridia oval and separated from anterior margin of tergite by about 3 times its own length. Ovipositor rather stout, slightly decurved, its sheath moderately broad. Male with sternites 7-9 bearing numerous long, erect, slightly curved hairs; gonosquama stout, distally rounded. Colour generally brownish yellow, with three dark longitudinal vittae on mesoscutum, and with tergites 3+ somewhat infuscate ; interocellar area yellow; antenna brownish, basally slightly darker; legs golden; pterostigma brownish; wings very slightly yellow, especially near hind margin of fore wing and along fore margin of hind wing. 248 IAN D. GAULD VARIATION. The specimens from Barro Colorado Island are more generally yellowish than other indivi- duals; one also has the mesoscutal vittae paler brownish. REMARKS. This species is named in honour of Bill Eberhard and Mary-Jane West Eberhard in recognition of their great contribution to tropical biology, and as a gesture of thanks for their hospitality and help. Enicospilus bima is immediately recognizable by the unusually protuberant clypeus and by the lack of an epicnemial carina on the lateral region of the alitrunk. The phylogenetic relationships of E. bima are not clear. The only other Central American species with a rather similarly modified clypeus is E. teodorae, but otherwise these two species have little in common, so possibly the similarly modified clypeus is an evolutionary parallelism, and not the result of common ancestry. BIOLOGICAL INFORMATION. Enicospilus bima is a widespread species whose range extends from Costa Rica (11°N) south to about 22°S in Brazil. Very few specimens have been collected in South America, but it is more common in collections from Central America. In Costa Rica E. bima is a lowland species and has never been collected at altitudes above 700 m. Specimens have been collected in Santa Rosa National Park in some of the driest months of the year (January-February), when no hosts are available. MATERIAL EXAMINED Holotype 9, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, i.1983 (Janzen & Hallwachs) (BMNH). Paratypes. Brazil: 1 2, Rio de Janeiro, Conceicaode, Macabu, i.1978 (Alvarenga) (TC). Costa Rica: Guanacaste Prov.: 1 0’, 1 9, W. of Carmona, Nicoya, 6-700 m, viii. 1982 (Janzen & Hallwachs) (BMNH), 1 9, Santa Rosa National Park, 300 m, xi.1982 (Janzen & Hallwachs) (BMNH); same locality and collectors, 1 Q, ii.1983; 1 CO’, vii.1984 (BMNH; MNCR): Heredia Prov.: 1 2, Finca La Selva, Puerto Viejo de Sarapiqui, 40 m, ii.1986 (Chavarria & Chacon) (BMNH); 1 9, same locality, xi.1986 (BMNH): Limon Prov.: 5 9, Cerro Tortuguero, N. edge of Tortuguero National Park, 0-100 m, v.1984 (Janzen & Hallwachs) (BMNH): San José Prov.: 1 0’, 1 9, Estacion Carrillo, Braulio Carrillo National Park, 700 m, xi.1984, v.1985 (Chacon) (BMNH). Panama: 2 ©’, Barro Colorado Island, vi.1983, v.1984 (Wolda) (BMNH) Enicospilus barbarae sp. n. (Fig. 302) DEscripTION. Mandibles moderately long, proximally strongly narrowed, with a basal lobe, distally more weakly tapered, apically twisted 40—50°; upper mandibular tooth depressed, 1.3—1.6 times as long as the lower tooth; outer mandibular surface centrally almost flat, sparsely pubescent, and with a weak proximal concavity. Labrum 0.2-0.3 times as long as broad; malar space 0.30.4 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.3—1.4 times as broad as long, the margin weakly convex. Lower face 0.66—-0.70 times as broad as long, finely punctate. Head in dorsal view with genae evenly narrowed behind eyes; posterior ocellus very close to eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.7—0.8 times the basal mandibular width away from mandible. Antenna slender, with 52-54 flagellar segments; 20th segment 1.9- 2.1 times as long as broad. Mesoscutum polished, with vestigial punctures, in profile abruptly rounded and with anterior margin slightly out-turned; notauli weak. Mesopleuron polished, the upper and lower parts uniformly punctate; epicnemial carina strong, curved towards anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.4-1.5 times as long as anteriorly broad, smooth, but posteriorly with some longitudinal wrinkles. Metapleuron weakly convex, punctate, but with the punctures rather coarse and very close posterodorsally; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile quite long and evenly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area moderately long, with scattered striae; spiracular area quite short, smooth; posterior area with irregular wrinkling; lateral longitudinal carina more or less complete, usually joined to spiracular margin by a short carina. Fore wing length 13-14 mm; discosubmarginal cell as in Fig. 302; AI = 1.31-1.50; CI = 0.24-0.38; ICI = 0.41-0.50; SDI = 1.29-1.52; cu-a proximal to the base of Rs&M by 0.2-0.3 times its own length; marginal cell proximally from evenly hirsute to narrowly glabrous close to Rs+2r; 1st subdiscal cell with anterior and distal margins hirsute. Hind wing with 6-7 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa almost straight. Fore leg with tibia subcylindrical, with scattered slender spines on outer surface. Mid leg with longer tibial spur 1.4~1.6 times length of the shorter. Hind leg with coxa in profile 1.8—-1.9 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.2—2.5 times as long as broad; OPHIONINAE OF TROPICAL MESOAMERICA 249 claws of female rather long, distally abruptly but shortly curved, with close, short pectinae; those of male similar but with pectinae closer together. Gaster slender; tergite 2 in profile 6-7 times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by about 2-3 times its own length. Ovipositor slender, more, or less straight, its sheath slender. Male with sternites 7-9 bearing scattered, long, erect slightly curved hairs; gonosquama distally evenly rounded. Colour generally yellowish, with three dark brown mesoscutal vittae, and with posterior segments of gaster slightly infuscate; interocellar area yellowish, sometimes slightly infuscate close to posterior ocellus; antenna blackish, with distal apices paler; pterostigma yellowish brown; wings slightly yellowish. VARIATION. Enicospilus barbarae is a morphologically and chromatically uniform species. REMARKS. This species is named in honour of Barbara Haber for her tireless efforts on behalf of education in Monteverde. Enicospilus barbarae is most easily recognized by the rather strongly sinuous Rs+2r in the fore wing (Fig. 302). No other species with black antennae and a central sclerite has this vein quite so sinuous. The form of the central sclerite — being obovate and positioned in the anterodistal margin of the fenestra — is also quite distinctive. The phylogenetic relationships of this species are not known. BIOLOGICAL INFORMATION. Enicospilus barbarae has only been collected at medium elevation sites at altitudes between 1100 and 1300 m in Costa Rica and Bolivia. Nothing is known about its natural history. MATERIAL EXAMINED Holotype 9, Costa Rica: Puntarenas Prov., Monteverde, 1350 m, no date or collector (TC). Paratypes. Bolivia: 5 9, Chapare, Alto Palmar, 1100 m, ii.1961 (BMNH; CNC). Costa Rica: Puntarenas Prov.: 1 ©’, Monteverde, 1350 m, 1962 (Palmer) (TC). Enicospilus fosteri sp. n. (Fig. 303) DEscriPTion. Mandibles moderately long, distally evenly tapered, apically twisted 60—70°; upper mandibu- lar tooth depressed, 1.3—1.5 times as long as the lower tooth which is slightly compressed; outer mandibular surface more or less flat, finely hirsute and with a weak proximal concavity. Labrum 0.2-0.3 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.3—1.4 times as broad as long, the margin weakly convex. Lower face 0.61—0.69 times as broad as long, with fine scattered punctures. Head in dorsal view with genae evenly constricted; posterior ocellus very close to eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.7-0.8 times the basal mandibular width away from mandible. Antenna slender, with 57-58 flagellar segments; 20th segment 1.9 times as long as broad. Mesoscutum polished, finely punctate, in profile steeply rounded; notauli vestigial. Mesopleuron polished, the upper and lower parts from uniformly finely and sparsely punctate, to increasingly punctostri- ate ventrally; epicnemial carina strong, curved towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.8 of its length; scutellum in dorsal view 1.5 times as long as anteriorly broad, smooth. Metapleuron moderately convex, from finely punctate with a few rugae posteriorly to diagonally striate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina represented by lateral vestiges; anterior area short, deeply impressed, striate; spiracular area short, smooth; posterior area quite coarsely and irregularly wrinkled, with a tendency to have longitudinal wrinkles centrally; lateral longitudinal carina complete, joined to spiracular margin by a short carina. Fore wing length 15-16 mm; discosubmarginal cell as in Fig. 303, very unusual in that the fenestra is margined anteriorly by a band of hair; AI = 0.67-0.81; CI = 0.24-0.28; ICI = 0.68-0.77; SDI = 1.21-1.30; cu-a proximal to base of Rs&M by about 0.2 times its own length; marginal cell proximally evenly hirsute or slightly more sparsely hirsute adjacent to Rs+2r; 1st subdiscal cell with antero-distal half hirsute. Hind wing with 5—6 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa noticeably bowed near distal end. Fore leg with tibia subcylindrical, with scattered, slender spines on outer surface. Mid leg with longer tibial spur 1.5—1.6 times length of the shorter. Hind leg with coxa in profile 1.9 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 1.6-1.7 times as long as broad; claws of female moderately long, abruptly curved, with short regularly spaced pectinae; claws of male similar. Gaster slender; tergite 2 in profile about 6 times as long as posteriorly deep, laterotergite folded under, thyridia oval and separated from anterior margin of tergite by about 6 times its own length. Ovipositor 250 IAN D. GAULD straight, apically very slender, its sheath narrow. Male with sternites 7-9 bearing dense moderately stout, semierect pubescence. Colour generally golden yellowish brown, with three dark brown vittae on the mesoscutum, and with the terminal segments of the gaster slightly infuscate; interocellar area yellow; antenna black; pterostigma brown; wings faintly yellowish. VARIATION. The male has the lateral region of the alitrunk more striate than the two females. REMARKS. This species is named in honour of Robin Foster in recognition of his devotion to helping all of us understand tropical tree biology. Enicospilus fosteri is immediately recognizable because of the presence of a band of hair bordering the fenestra, immediately adjacent to Rs+2r (Fig. 303). No other Central American species has such a hirsute area. Structurally, and in the form of the alar sclerites, E. fosteri resembles E. Luisi. BIOLOGICAL INFORMATION. Enicospilus fosteri is only known to occur in Panama where it has occasionally been collected in lowland rainforest on Barro Colorado Island. No details are known of its natural history. MATERIAL EXAMINED Holotype 9, Panama: Barro Colorado Island, 120 m, v.1985 (Wolda) (BMNH). Paratypes. Panama: 1 9, Barro Colorado Island, xi.1977 (Wolda) (RNH); 1 oO’, same locality, vi.1978 (Wolda) (RNH). Enicospilus cornifuscus nom. n. (Figs 195, 304) Ophion (Enicospilus) fuscicornis Cameron, 1886: 291. LECTOTYPE 2, GUATEMALA (BMNH), here designated [examined]. [Primary homonym of Ophion fuscicornis Erichson, 1842. ] Henicospilus fuscicornis (Cameron) Dalla Torre, 1901: 182. Enicospilus fuscicornis (Cameron) Hooker, 1912: 60. Description. Mandibles moderately long, proximally strongly tapered, distally weakly tapered, apically twisted 15—25°; upper mandibular tooth subcylindrical, 1.6-1.9 times as long as the lower tooth; outer mandibular surface centrally weakly concave, sparsely hirsute, and proximally with a weak concavity. Labrum 0.3 times as long as broad; malar space 0.30.4 times as long as basal mandibular width. Clypeus in profile weakly convex, margin subacute; clypeus in front view 1.3—1.5 times as broad as long, the margin almost truncate. Lower face 0.68—0.72 times as broad as long, centrally with scattered fine punctures. Head in dorsal view with genae evenly constricted behind the eyes; posterior ocellus contiguous with the eye; FI = 65-70%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.8 times the basal mandibular width away from mandible. Antenna slender, with 53-54 flagellar segments; 20th segment 1.9-2.4 times as long as broad. Mesoscutum polished, finely punctate, in profile abruptly rounded; notauli vestigial. Mesopleuron highly polished, the upper part very finely punctate, the lower part punctate to punctostriate; epicnemial carina strong, curved towards anterior margin of pleuron, but often with upper end evanescent. Scutellum in profile weakly convex, laterally carinate for less than 0.5 of its length; scutellum in dorsal view 1.4-1.5 times as long as anteriorly broad, smooth. Metapleuron weakly convex, highly polished, finely punctate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete, or in some specimens weak centrally. Propodeum in profile long, weakly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area moderately short, coriaceous or with scattered striae; spiracular area long, smooth; posterior area virtually smooth and highly polished, at most with weak indistinct sculpture centrally (Fig. 195); lateral longitudinal carina from complete to completely absent, not joined to spiracular margin by a short carina. Fore wing length 14-16 mm; discosubmarginal cell as in Fig. 304; AI = 0.78-1.24; CI = 0.17-0.30; ICI = 0.33-0.37; SDI = 0.91-1.03; cu-a proximal to the base of Rs&M by 0.1—0.2 times its own length; marginal cell proximally from slightly less densely hirsute than centrally to with a small glabrous area adjacent to Rs+2r; 1st subdiscal cell hirsute except at proximal end. Hind wing with 6-7 hamuli on R1; Ist abscissa of Rs slightly curved, 2nd abscissa weakly bowed. Fore leg with tibia subcylindrical, with numerous fine spines on outer surface. Mid leg with longer tibial spur 1.5—1.7 times length of the shorter. Hind leg with coxa in profile 1.9-2.0 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.3—2.5 times as long as broad; claws of female long, weakly curved, with short stout pectinae, those of male similar but with pectinae finer and closer together. OPHIONINAE OF TROPICAL MESOAMERICA 251 Gaster slender; tergite 2 in profile 6.0—7.0 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 3-4 times its own length. Ovipositor slender, straight, its sheath narrow. Male with sternites 7-9 bearing scattered stout erect hairs; gonosquama distally subacute. Colour generally brownish yellow, with most of head, irregular mesoscutal markings, periphery of mesoscutum, mesoscutal stripes, and scutellum brighter yellow, and with terminal segments of gaster weakly infuscate; interocellar area yellow; antenna blackish, distally slightly paler; pterostigma yellowish brown; wings hyaline. VARIATION. The specimens from Guatemala City are slightly darker than the others. The specimen from San Lorenzo lacks longitudinal scutellar carinae, but has in their place longitudinal wrinkles. REMARKS. Townes & Townes (1966) state that the type of Ophion (Enicospilus) fuscicornis Cameron is lost, but I located a syntype in the BMNH which is labelled as ‘ex Cameron collection’. This I have designated as lectotype. Unfortunately it is asomewhat damaged specimen, lacking a fore and a hind wing, most of the legs and the posterior tergites of the gaster. The head has been badly glued onto the body. My supposition that this specimen is female is based on the examination of the fore tarsal claws; those of the single male I have seen have closer pectination. Enicospilus cornifuscus is a distinctive species on account of the long and very weakly sculptured propodeum (Fig. 195). The usually small value of the SDI, the virtual absence of lateral longitudinal carinae on the scutellum and the characteristic form of the central sclerite (Fig. 304) are also important diagnostic features. E. cornifuscus is one of the very few Central American species with dark antennae and without black stripes on the mesoscutum. I do not know which species E. cornifuscus is most closely related to. The most striking specialized features it exhibits are characters found in many tropical montane species from various regions, including New Guinea and Central Africa (Gauld & Mitchell, 1978; 1981), so I surmise these are adaptive parallelisms. BIOLOGICAL INFORMATION. The majority of specimens I have seen were collected in southern Mexico and Guatemala, but a single female in the BMNH was collected in southern Brazil, indicating that Enicospilus cornifuscus extends over a considerable geographical range. The Central American specimens were all collected in sites above 1000 m. Nothing is known of the biology of this species. MATERIAL EXAMINED Lectotype 2, Guatemala: ‘S. Geronimo’ (= San Jeronimo) (Champion) (BMNH). Brazil: 1 9, Santa Catarina, Nova Teutonia, 27°11’S, 52°23’W, i.1939 (Plaumann) (BMNH). Guatemala: 4 2, Guatemala City (Rodriguez) (BMNH); 1 9, Zacapa, San Lorenzo, 1700 m, xi.1986 (Sharkey) (CNC). Mexico: Chiapas: 3 2, San Cristobal de las Casas, 2400 m, v.1969 (CNC); 4 9, same locality, v.1969 (Martin) (CNC); 1 o’, same locality, vii.1969 (Kritsch) (CNC); 1 9, Yerba Buena, 32 km N. Bochil, v.1969 (Mason) (CNC). Enicospilus catemacoi sp. n. (Fig. 305) DescriPTION. Mandibles moderately long, rather evenly narrowed, apically twisted 15°; upper mandibular tooth slender, 1.5 times as long as the lower tooth; outer mandibular surface almost flat, with scattered pubescence. Labrum 0.4 times as long as broad; malar space 0.2 times as long as basal mandibular width. Clypeus in profile almost flat, margin subacute; clypeus in front view 1.7 times as broad as long, margin truncate apically. Lower face 0.83 times as broad as long, centrally punctate. Head in dorsal view with genae slightly inflated behind eyes; posterior ocellus contiguous with eye; FI = 60%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.9 times the basal mandibular width away from mandible. Antenna slender, with 62 flagellar segments; 20th segment 1.6 times as long as broad. Mesoscutum polished, sparsely punctate, in profile abruptly rounded, with anterior margin out-turned; notauli distinctly impressed near anterior margin. Mesopleuron polished, the upper part punctate, the lower part punctostriate; epicnemial carina curved towards anterior margin of pleuron, but with upper end evanescent. Scutellum in profile almost flat, laterally carinate for 0.8 of its length; scutellum in dorsal view 1.5 times as long as anteriorly broad, punctate, with a few longitudinal wrinkles posteriorly. Metapleuron weakly convex, anteriorly striate, posteriorly rugose-striate; submetapleural carina weakly anteriorly broadened; posterior transverse carina of mesosternum broadly incomplete centrally. Propodeum in profile abruptly declivous; anterior transverse carina complete but weak, posterior transverse carina 252 IAN D. GAULD absent; anterior area short, deeply impressed, coriaceous; spiracular area moderately long, finely punc- tate; posterior area coarsely irregularly wrinkled; lateral longitudinal carina absent, and therefore not joined to spiracular margin by a short carina. Fore wing length 15 mm; discosubmarginal cell asin Fig. 305; AI = 1.00; CI = 0.39; ICI = 0.35; SDI = 1.10; cu-a subopposite to base of Rs&M; marginal cell proximally more or less evenly hirsute; 1st subdiscal cell with anterior periphery and posterodistal corner hirsute. Hind wing with 8 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa straight. Fore leg with tibia weakly flattened, with slender scattered spines on outer surface. Mid leg with longer tibial spur 1.3 times length of the shorter. Hind leg with coxa in profile 1.9 times as long as deep; trochantellus dorsally 0.2 times as long as broad; 4th segment of tarsus 2.1 times as long as broad; claws of female abruptly curved, with close long stout pectinae. Gaster slender; tergite 2 in profile 4.3 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 4 times its own length. Ovipositor moderately slender, its sheath narrow. Male unknown. Colour generally mottled yellowish and reddish brown, without dark mesoscutal stripes; gaster brownish, with tergites 3+ darker brown; interocellar area yellow; antenna blackish; pterostigma dark reddish brown; wings hyaline. REMARKS. Enicospilus catemacoi may be recognized by the combination of black antennae, more or less completely carinate scutellum, incomplete posterior mesosternal transverse carina, small value of ICI and broad face. Structurally, especially in the form of the mandibles, it is quite similar to E. gabrieli, but the two may be separated, not only on the characters given in the key, but also by the form of the alar sclerites and the shape of the 2nd discal cells (compare Figs 305 and 306). The central sclerite of E. catemacoi is subtriangular and positioned antero-distally, and the 2nd discal cell is quite short and broad; the central sclerite of E. gabrieli is more ovate and positioned mediodistally, and the 2nd discal cell in long and rather narrow. These two species also differ in the sculpture of the posterior area of the propodeum; that of E. catemacoi is irregularly wrinkled whereas that of E. gabrieli is more or less concentrically rugose. BIOLOGICAL INFORMATION. The single specimen was collected in July at a black-light run near Lake Catemaco, Veracruz, Mexico. MATERIAL EXAMINED Holotype 2, Mexico: Veracruz, ‘Coyame’, Lake Catemaco, vii.1965 (Woodruff) (FSCA). Enicospilus gabrieli sp. n. (Fig. 306) Description. Mandibles moderately long, rather evenly narrowed, apically twisted 20°; upper mandibular tooth subcylindrical, 1.6 times as long as the lower tooth; outer mandibular surface more or less flat, but with a weak dorsal ridge, centrally sparsely hirsute and without a distinct proximal concavity. Labrum 0.3 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile almost flat, margin blunt; clypeus in front view 1.7 times as broad as long, the margin subtruncate apically. Lower face 0.74 times as broad as long, centrally punctate. Head in dorsal view with genae evenly narrowed behind the eyes; posterior ocellus contiguous with the eye; FI = 65%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.9 times the basal mandibular width away from mandible. Antenna slender, with apical flagellar segments missing; 20th segment 1.8 times as long as broad. Mesoscutum polished, punctate, in profile weakly rounded; notauli distinctly impressed anteriorly. Mesopleuron polished, the upper part punctate, the lower part punctostriate; epicnemial carina curved towards anterior margin of pleuron, but with its upper end complete. Scutellum in profile almost flat, laterally carinate for 0.8 of its length; scutellum in dorsal view 1.5 times as long as anteriorly broad, punctate anteriorly, posteriorly with longitudinal wrinkles. Metapleuron moderately convex, rugose- striate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum broadly incomplete centrally. Propodeum in profile abruptly declivous; anterior transverse carina sinuous, lateromedially discontinuous, posterior transverse carina vestigial; anterior area long, shallow and striate; spiracular area short, punctate; posterior area strongly deplanate and abruptly declivous laterally, cen- trally rugose with the rugae tending to be concentric; lateral longitudinal carina present only as a vestige anteriorly, indistinctly joined to spiracular margin by a short carina. Fore wing length 18 mm; discosubmarginal cell as in Fig. 306; AI = 0.79; CI = 0.44; ICI = 0.75; SDI = 1.31; cu-a proximal to the base of Rs&M by 0.1 times its own length; marginal cell proximally very slightly more OPHIONINAE OF TROPICAL MESOAMERICA 253 sparsely hirsute than it is centrally; 1st subdiscal cell with anterior 0.5 and distal periphery hirsute. Hind wing with 8 hamuli on R1; Ist abscissa of Rs more or less straight, 2nd abscissa weakly arcuate. Fore leg with tibia slightly flattened, with scattered, dark spines on outer surface. Mid leg with longer tibial spur 1.6 times length of the shorter. Hind leg with coxa in profile 1.7 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 1.9 times as long as broad; claws of female abruptly curved, with close pectinae. Gaster slender; tergite 2 in profile 4.6 times as long as posteriorly deep, laterotergite turned under, thyridia oval and separated from anterior margin of tergite by about 5 times its own length. Ovipositor slender, its sheath narrow. Male unknown. Colour generally with head and alitrunk bright yellowish, except for mesoscutum which bears three black longitudinal stripes; legs and gaster golden; interocellar area yellow with slight infuscation between posterior ocelli; antenna black; pterostigma brown; wings virtually hyaline. Remarks. Enicospilus gabrieli may be distinguished from the other species with black antennae and an incomplete posterior transverse carina of the mesosternum (E. catemacoi) by the incomplete anterior transverse carina of the propcdeum, the propodeal sculpture, the laterally carinate scutellum and the coloration (see also remarks under E. catemacoi p. 252). BIOLOGICAL INFORMATION. The single female was collected in June at 850 m at a black-light overlooking a small area of wet forest, just on the Atlantic side of the continental divide in northern Costa Rica. It is perhaps noteworthy that only about ten nights have been spent light-trapping in this locality. E. gabrieli has not been collected in other Costa Rican wet forest sites (such as Monteverde and Braulio Carrillo) which have been more intensively sampled. MATERIAL EXAMINED Holotype 9 ,Costa Rica: Aiajuela Prov.: Finca San Gabriel, 3 km W. of Dos Rios, 850 m, vi.1986 (Gauld) (BMNH) Enicospilus luisi sp. n. (Figs 307, 308) Description. Mandibles moderately long, proximally strongly narrowed and with a proximoventral lobe; mandibles distally weakly narrowed, apically twisted 45—5S0°; upper tooth depressed, 1.5—1.7 times as long as the lower tooth which is ventrally slightly swollen; outer mandibular surface slightly concave, with an angular ridge extending from upper corner to between teeth. Labrum 0.2-0.3 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin inturned, slightly emarginate; clypeus in front view 1.3—1.5 times as broad as long, with margin almost truncate apically. Lower face 0.68-0.71 times as broad as long, polished, punctate, centrally sometimes slightly punctostriate. Head in dorsal view with genae evenly rounded behind eyes; posterior ocellus contiguous with eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.8 times the basal mandibular width away from mandible. Antenna slender, with 64-68 flagellar segments; 20th segment 2.0—2.1 times as long as broad. Mesoscutum polished, quite closely punctate, in profile abruptly rounded; notauli vestigial. Meso- pleuron polished, the upper part finely and closely punctate, the lower part with irregular transverse wrinkles; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.4—1.5 times as long as anteriorly broad, anteriorly smooth, posteriorly wrinkled. Metapleuron moderately convex, striate or with irregular rugae that extend dorsally onto propodeum; submetapleural carina quite strongly anteriorly broadened; posterior transverse carina of mesosternum complete, generally slightly produced centrally so entire carina is somewhat chevronate. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina vestigial, at most represented by weak lateral keels; anterior area quite long, sparsely striate; spiracular area short, smooth; posterior area rather flat, with quite coarse irregular rugae, tending towards concentrically striate laterally, centrally more reticulate; lateral longitudi- nal carina present only anteriorly, joined to spiracular margin by ashort, sometimes discontinuous carina. Fore wing length 18-20 mm; discosubmarginal cell as in Figs 307, 308; AI = 0.91-1.06; CI = 0.20-0.31; ICI = 0.55—0.69; SDI = 1.25—1.51; distal abscissa of 1A unusual in that it bears a dorsal row of long fine hairs; cu-a proximal to the base of Rs&M by 0.2-0.4 times its own length; marginal cell proximally uniformly hirsute; 1st subdiscal cell with anterior and distal parts hirsute. Hind wing with 7-8 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa weakly arcuate. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.4~1.5 times length of the shorter. Hind leg with coxa in profile 1.8-1.9 times as long as deep; 254 IAN D. GAULD trochantellus dorsally 0.1—0.2 times as long as broad; 4th segment of tarsus 2.3-2.6 times as long as broad; claws of female abruptly curved, with short stout pectinae, those of male similar but with pectinae slightly finer. Gaster slender; tergite 2 in profile from 5.5—6.8 times as long as posteriorly deep, laterotergite turned under, thyridia obovate to elliptical and separated from anterior margin of tergite by about 4—5 times its own length. Ovipositor slender, slightly decurved, its sheath quite broad. Male with sternites 7-9 bearing long stout, erect pubescence; gonosquama distally rounded. Colour generally pale yellowish brown, mesoscutal vittae dark brown, the gaster reddish brown to dark brownish; interocellar area yellowish, sometimes with slight infuscation between posterior ocelli; antenna proximally black, distally paler; pterostigma brown; wings weakly infumate. VaRIATION. There is slight variation in the size and shape of the alar sclerites (Figs 307, 308). The proximal one may be truncated anteriorly whilst the central one is always small, but may be circular or oval. This is otherwise a morphologically very uniform species. REMARKS. This species is named in honour of Luis Diego Gomez, in recognition of his diverse contributions to Costa Rican biology and conservation. Enicospilus luisi can be distinguished from all other Neotropical species by the presence of a row of long fine hairs on the dorsal surface of 1A in the fore wing. Structurally, particularly in the form of the mandible and in sculpture, FE. Juisi resembles E. fosteri and E. opleri. BIOLOGICAL INFORMATION. Enicospilus luisi is a widely distributed species whose range extends from Chiapas in southern Mexico south to Ecuador. It has been collected in forests at altitudes from 300 to 1350 m,but is most commonly encountered at the lower elevation sites. In Costa Rica, in Santa Rosa National Park, E. /uisiis the most frequently collected Enicospilus species in Malaise traps from August until December (the latter part of the wet season). Dr D. H. Janzen has reared a single female in Santa Rosa from an unidentified species of Notodontidae found feeding on Malpighia glabra L. (Malpighiaceae) (rearing ref. number 84.S5RNP.1456). The caterpillar was collected on 14th July and the ichneumonid emerged on the Sth August. MATERIAL EXAMINED Holotype 9, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, viii.1984 (Janzen & Hallwachs) (BMNH). Paratypes. Costa Rica: Guanacaste Prov.: 1 9,4 kmE. Casetilla, Rincén de la Vieja National Park, 750 m, xii.1981 (Janzen & Hallwachs) (BMNH); 1 o’, Santa Rosa National Park, 300 m, vii.1982 (Janzen & Hallwachs) (BMNH); same locality and collectors, 1 2, vi.1984; 6 Q, viii.1984; 8 Q, viii-xii.1984 (BMNH): Puntarenas Prov.: 1 0’, Monteverde, 1350 m, ii.1986 (Haber) (BMNH); 1 9, same locality, i.1986 (Forsyth) (CNC). Ecuador: 1 2, Zamora, iv.1965 (Peria) (TC). Mexico: Chiapas: 2 2, 32 km N. Huixtla, 1000 m, vi.1969 (CNC). Panama: 1 ©’, Chiriqui, Fortuna, 1050 m, i.1978 (Wolda) (RNH). Venezuela: 2 9, Tucuco, Zulia, iv.1981 (Townes) (TC). Enicospilus opleri sp. n. Description. Mandibles moderately long, distally fairly evenly narrowed, apically twisted 40—50°; upper mandibular tooth depressed, slender, 1.3—1.5 times as long as the lower tooth which is swollen and rather convex ventrally; outer mandibular surface with a weak longitudinal concavity that is margined dorsally by a small ridge that extends from base of upper tooth to upper corner of mandibular base, this depression sparsely pubescent; proximal concavity weak. Labrum 0.2-0.3 times as long as broad; malar space 0.2-0.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt to subacute; clypeus in front view 1.3-1.5 times as broad as long, with margin apically truncate. Lower face 0.65—0.69 times as broad as long, centrally finely punctate, sometimes virtually smooth. Head in dorsal view with genae evenly rounded behind eyes; posterior ocellus contiguous with eye; FI = 75-80%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.7—0.9 times the basal man- dibular width away from mandible. Antenna slender, with 62-65 flagellar segments; 20th segment 1.7—1.8 times as long as broad. Mesoscutum polished, finely punctate, in profile rather abruptly rounded, with anterior margin out- turned; notauli weak but discernible anteriorly. Mesopleuron highly polished, the upper part regularly punctate, the lower part often similar to the upper but generally with some tendency to punctostriation; epicnemial carina curved towards but not joining anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for 0.7 or more of its length; scutellum in dorsal view 1.4—1.5 times as long as anteriorly broad, anteriorly smooth or punctate, posteriorly generally with longitudinal wrinkling. OPHIONINAE OF TROPICAL MESOAMERICA 255 Metapleuron weakly convex, with strong rugae posterodorsally; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile abruptly declivous; anterior transverse carina complete, posterior transverse carina vestigial, at most represented by lateral crests; anterior area quite long, deep, striate; spiracular area rather short, smooth; posterior area coarsely reticulate-rugose, often with rugae tending to be longitudinal anterocentrally, and sometimes even subconcentric posteriorly; lateral longitudinal carina present anterior to anterior transverse carina, to which it is often joined, and joined to spiracular margin by a short carina. Fore wing length 16-20 mm; discosubmarginal cell very like those shown in Figs 307, 308, except that vein 1A does not bear long hairs; AI = 0.70—0.81; CI = 0.25—0.30; ICI = 0.59-0.88; SDI = 1.31-1.49; cu-a proximal to the base of Rs&M by 0.1-0.3 times its own length; marginal cell proximally fairly uniformly hirsute; 1st subdiscal cell extensively hirsute except for posteroproximal corner. Hind wing with 7-8 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa weakly bowed. Fore leg with tibia distinctly flattened, with numerous scattered spines on outer surface. Mid leg with longer tibial spur 1.3—1.6 times length of the shorter. Hind leg with coxa in profile 1.8—1.9 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.0—2.2 times as long as broad; claws of female abruptly curved, with long stout pectinae; those of male similar but with pectinae closer and shorter. Gaster slender; tergite 2 in profile 5.2—-5.8 times as long as posteriorly deep, laterotergite turned under, thyridia oval and separated from anterior margin of tergite by about 4—5 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing numerous long stout erect, slightly curved hairs; gonosquama distally evenly rounded. Colour generally yellowish to reddish brown, usually with tergites 3+ of gaster somewhat infuscate, always with three black longitudinal vittae on the mesoscutum; interocellar area yellowish, sometimes indistinctly blackish posteriorly, adjacent to ocelli; antenna black; pterostigma brown; wings weakly infumate. VARIATION. The specimens from Florida are generally more orange-yellow than others, but structurally they are otherwise typical examples of the species. REMARKS. This species is named in honour of Paul Opler in recognition of his many efforts on behalf of insect conservation. The most distinctive feature of Enicospilus opleri is the presence of a group of hairs in the extreme anterior corner of the discosubmarginal cell, adjacent to the base of Rs+2r (cf. Fig. 307). This feature in combination with the characteristic alar sclerites, slightly ridged mandibles and black antenna serve to distinguish E. opleri from all other species except E. /uisi. In many other features, such as the form of the mandibles, general sculpture, alar indices, and position of the alar sclerites, E. opleri and E. luisi are very similar, but E. opleri does not have slender hairs on the upper surface of the distal abscissa of 1A. BIOLOGICAL INFORMATION. Enicospilus opleri is a widespread species (Map 28) whose range extends from northern Costa Rica (11°N) south to Bahia, Central Brazil (ca 10°S). Several individuals have also been collected in the extreme southern tip of Florida and on the Florida Keys, though I have seen no other material from the Caribbean. In Central America E. opleri has been collected from almost sea level up to about 1800 m. In Santa Rosa National Park, Costa Rica, despite intensive sampling, only a single specimen has been collected, in June, at the start of the wet season. Nothing is known of the natural history of this species. MATERIAL EXAMINED Holotype 9, Costa Rica: Cartago Prov., 3 km S. Casa Mata, 21.6 km S. San Isidro de Tejar, 1800 m, xii.1983 (Janzen & Hallwachs) (BMNH). Paratypes. Brazil: 1 9, Bahia, Encruzilhada, 900 m, xi.1972 (Alvarenga) (TC). Colombia: 1 C, Monterredondo, Cundinamarca, xii.1956 (Foerster) (TC). Costa Rica: Cartago Prov.: 1 2, 3 km S. Casa Mata, 21.6 kmS. San Isidro de Tejar, 1800 m, xii.1983 (Janzen & Hallwachs) (BMNH): Guanacaste Prov.: 1 9, Santa Rosa National Park, 300 m, vi.1984 (Janzen & Hallwachs) (BMNH): Heredia Prov.: 1 CO’, Puerto Viejo de Sarapiqui, Finca la Selva, iv.1986 (BMNH): Puntarenas Prov.: 1 oO’, 1 9, Finca Las Cruces, 6 km S. San Vito de Java, 1400 m, ix.1986 (Eger) (BMNH). Guyana: 1 2, Kuraubam Ck, Cattle trail survey, ix.1919 (Abraham) (BMNH). Panama: 3 Q, Barro Colorado Island, xi.1983 & vi.1985 (Wolda) (BMNH). U.S.A.: Florida: 1 9, Dade Co., Fuch’s Hammock, nr Homestead, v.1979 (Dickel & Weems) (FSCA); 1 2, Monroe Co., Big Pine Key, Alligator Ford, vii. 1978 (Stange) (FSCA); 2 2, Monroe Co., Key Largo Key, 16 km N. Key Largo City, i.1974 (Heppner) (FSCA); 1 9, Monroe Co., No Name Key, xii.1972 (Dodge) (FSCA). Venezuela: 1 2, Tucuco, Zulia, iv.1981 (Townes) (TC). 256 IAN D. GAULD Map 28 Localities at which Enicospilus opleri has been collected. Enicospilus guindoni sp. n. (Figs 132, 309) DescriPTION. Mandibles moderately long, proximally abruptly narrowed, so a distinct ventral lobe is discernible, distally slender and weakly narrowed, apically twisted about 40°; upper mandibular tooth slender, depressed, 1.4-1.6 times as long as the lower tooth; outer mandibular surface almost flat, with scattered pubescence, proximally with a weak concavity. Labrum 0.2 times as long as broad; malar space 0.2-0.3 times as long as basal mandibular width. Clypeus in profile very weakly convex, margin blunt; clypeus in front view 1.3—1.5 times as broad as long, the margin truncate apically. Lower face 0.70-0.73 OPHIONINAE OF TROPICAL MESOAMERICA Di. times as broad as long, centrally finely, sparsely punctate. Head in dorsal view with genae evenly rounded behind eyes; posterior ocellus very close to eye; FI = 70-75; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.7—0.8 times the basal mandibular width away from mandible. Antenna slender, with 56—S8 flagellar segments; 20th segment 2.0—2.2 times as long as broad. Mesoscutum polished, very finely punctate, in profile evenly rounded; notauli vestigial. Mesopleuron polished, the upper part finely punctate, the lower part similar but with some striation present; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile almost flat, laterally carinate for 0.3 or less of its length; scutellum in dorsal view 1.5 times as long as anteriorly broad, rather smooth with scattered punctures, without wrinkling posteriorly. Metapleuron moderately convex, generally punctate, sometimes with some rugae posterodorsally; submetapleural carina weakly broadened anteriorly; pos- terior transverse carina of mesosternum complete, projecting forward on midline so that in ventral view the carina appears to form a chevron (Fig. 132). Propodeum in profile fairly eveniy declivous; anterior transverse carina strong and complete, posterior transverse carina represented laterally by crests; anterior area long and shallow, striate; spiracular area rather short, smooth; posterior area irregularly wrinkled- rugose, with a median longitudinal wrinkle present anteriorly, the other rugae often tending to be subconcentric, but generally with these usually weakly developed; lateral longitudinal carina from com- plete to interrupted centrally, joined to spiracular margin by a short carina. Fore wing length 18-19 mm; discosubmarginal cell as in Fig. 309; AI = 1.03—1.32; CI = 0.21-0.36; ICI = 0.51-0.60; SDI = 1.27-1.39; cu-a proximal to the base of Rs&M by about 0.2 times its own length; marginal cell proximally more sparsely hirsute than it is centrally; 1st subdiscal cell with anterodistal part hirsute. Hind wing with 7-8 hamuli on R1; Ist abscissa of Rs almost straight, 2nd abscissa straight. Fore leg with tibia weakly flattened, with scattered slender spines on outer surface. Mid leg with longer tibial spur 1.41.5 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 1.6—2.0 times as long as broad; claws of female quite long, distally abruptly rounded, with rather short, close pectinae. Gaster very slender; tergite 2 in profile 7-8 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 4 times its own length. Ovipositor quite slender, slightly decurved, its sheath narrow. Male unknown. Colour generally yellowish, with three longitudinal dark brown stripes on mesoscutum; gaster with tergites 3+ laterally rather pale, dorsally and ventrally very slightly infuscate; interocellar area yellow, or slightly infuscate close to posterior ocelli; antenna black; pterostigma yellowish brown; wings weakly infumate. VaRIATION. The Costa Rican and Bolivian specimens are remarkably alike, except that the mandibles of the former are very slightly more slender than are those of Bolivian examples. REMARKS. This species is named in honour of Wolf Guindon, in recognition for his devotion to con- servation in the Monteverde area, and for his help in the field. Enicospilus guindoni may be recognized by the following combination of features: the centrally pro- duced, chevronate posterior transverse carina of the mesosternum; the black antennae; the short lateral carinae of the scutellum; the weak rugae on the propodeal posterior area; and the small, perpendicular central sclerite. No other Central American species shares all these features. The form of the alar sclerites is similar to that found in E. leoni (cf. Figs 309 and 311) and possibly the two species are closely related. BIOLOGICAL INFORMATION. Enicospilus guindoni is only known from two sites at mid altitude (1100-1300 m) in Costa Rica and Bolivia, but presumably it occurs at similar elevations along the Andes in intervening countries. At Monteverde the two females were collected at a black light in cloud forest. Although other workers have collected many specimens in Monteverde adjacent to the forest (between 1984 and 1986), they have not taken any further examples of this species. Possibly EF. guindoni is restricted just to the forest. MATERIAL EXAMINED Holotype 2, Costa Rica: Puntarenas Prov., Monteverde, 1300 m, iv-v.1984 (Gauld) (BMNH). Paratypes. Bolivia: 2 9, Chapare, Alto Palmar, 1100 m, ii.1961 (CNC). Costa Rica: Puntarenas Prov.: 1 2, Monteverde, 1300 m, iv-v.1984 (Gauld) (BMNH). Enicospilus leoni sp. n. (Figs 133, 196, 197, 311) DescriPTion. Mandibles moderately long, evenly tapered, but with a very small basal lobe, apically twisted 20°; upper mandibular tooth slender, depressed, 1.3—1.5 times as long as the lower tooth (Fig. 196); outer 258 IAN D. GAULD mandibular surface centrally weakly concave, with a weak ridge extending from base of upper tooth to mandibular base, the concave area bearing close pubescence. Labrum 0.2-0.3 times as long as broad; malar space 0.2 times as long as basal mandibular width. Clypeus in profile very weakly convex, margin blunt; clypeus in front view 1.3-1.4 times as broad as long, with margin weakly convex apically. Lower face 0.70-0.74 times as broad as long, polished, centrally sparsely punctate. Head in dorsal view with genae strongly constricted behind eyes; posterior ocellus contiguous with eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.8 times the basal mandibular width away from mandible. Antenna long and slender, with 53-56 flagellar segments; 20th segment 2.2-2.4 times as long as broad. Mesoscutum polished, with fine punctures, in profile abruptly rounded; notauli absent. Mesopleuron polished, the upper part finely punctate, the lower part punctate to punctostriate; epicnemial carina inclined towards anterior, but not reaching margin of pleuron. Scutellum in profile weakly convex, laterally carinate for most of its length; scutellum in dorsal view 1.6—1.7 times as long as anteriorly broad, anteriorly smooth, posterior weakly wrinkled. Metapleuron weakly convex, more or less striate (Fig. 197); submetapleural carina weakly anteriorly broadened; posterior transverse carina of mesosternum com- plete. Propodeum in profile evenly rounded; anterior transverse carina complete, posterior transverse carina absent; anterior area short, and deep, striate to coriaceous; spiracular area quite long, smooth; posterior area irregularly reticulate; lateral longitudinal carina from present only as a small vestige anteriorly, to incomplete posteriorly anteriorly joined to spiracular margin by a short carina. Fore wing length 12-14 mm; discosubmarginal cell as in Fig. 311; AI = 1.00-1.50; CI = 0.23-0.30; ICI = 0.40—0.45; SDI = 1.00—-1.17; cu-a from subopposite to proximal to the base of Rs&M by 0.1 times its own length; marginal cell proximally narrowly glabrous; 1st subdiscal cell with anterior 0.3 sparsely hirsute. Hind wing with 5—7 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa almost straight or weakly bowed. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.5—1.7 times length of the shorter. Hind leg with coxa in profile 1.8-1.9 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.0—2.4 times as long as broad; claws of female with close, long pectinae, those of male with pectinae very short and closely interspaced. Gaster slender; tergite 2 in profile 5 or more times as long as posteriorly deep, laterotergite folded under, thyridia obovate and separated from anterior margin of tergite by about 5—6 times its own length. Ovipositor slender, very slightly decurved, its sheath moderately narrow. Males with sternites 7-9 bearing numerous rather fine, erect hairs; gonosquama with a weak dorsal notch, distally rounded. Colour generally brownish yellow with dark brown mesoscutal vittae and with tergites 3+ of gaster infuscate; interocellar area yellow; antenna proximally black, distally yellowish brown; pterostigma orange; wings hyaline. VARIATION. The majority of specimens have the distal sclerite very weakly sclerotized and confluent with the proximal sclerite. Several individuals have the medial part of the distal sclerite thickened. REMARKS. This species is named in honour of Pedro Leon in recognition of his efforts to improve the quality of Costa Rican high school biology texts. Enicospilus leoni can most easily be recognized by the I-shaped central sclerite, a feature it shares with E. guindoni and E. monticola. These three species can easily be separated by the characters given in the key. BIOLOGICAL INFORMATION. Enicospilus leoni is a widespread species whose range extends from northern Costa Rica south to Bolivia and central Brazil. The majority of specimens have been collected below 1000 m, although one Bolivian individual is purported to have been collected at 2200 m. In Central America E. leoni has only been collected infrequently. Despite a considerable collecting effort in Costa Rica only three specimens have been taken, and all these are from Santa Rosa National Park, possibly the northern extremity of the species’ range. Nothing is known about the biology of this insect. MATERIAL EXAMINED Holotype 9, Panama: Barro Colorado Island, 120 m, vi.1984 (Wolda) (BMNH). Paratypes. Bolivia: 1 9 , Chapare Prov., El Limbo, 2200 m, i.1962 (CNC); 1 9, La Paz, Yungas, 1600 m, xli.1984 (Pera) (TC). Brazil: 9 2, Bahia, Encruzilhada, 960 m, xi.1972 (Alvarenga) (TC); 1 2, Minas Gerais, Aguas Vermelhas, 800 m, xii.1983 (Alvarenga) (TC); 3 2, Minas Gerais, Pedras Azul, 800 m, xi.1972 (Alvarenga & Seabra) (TC). Colombia: 1 2 , Magdalena, 11°10’N 74°8’W, 800 m, iv.1973 (Helava) (BMNH). Costa Rica: Guanacaste Prov.: 1 9, Santa Rosa National Park, 300 m, viii.1982 (Janzen & Hallwachs) (BMNH); 1 Oo’, 1 , same locality and collectors, v-vi.1983 (BMNH). Panama: 1 ©’, Barro Colorado Island, vii.1983 (Wolda) (BMNH); same locality, 1 ©’, ix.1983, 1 Oo’, v.1984, 1 CO’, vi.1985 OPHIONINAE OF TROPICAL MESOAMERICA 259 (Wolda) (BMNH). Venezuela: 1 9, Aragua, Rancho Grande, v.1960 (Test) (UMC); 1 9, San Esteban, nr Puerto Cabello, xii.1939 (Anduze) (TC). Enicospilus kelloggae sp. n. (Figs 198, 199, 312) Description. Mandibles of moderate length, proximally abruptly narrowed, distally weakly narrowed, apically twisted 40—60°; upper mandibular tooth rather slender, 1.3—1.5 times as long as the lower tooth which is swollen and has a sharp, convex ventral margin (Fig. 199); outer mandibular surface almost flat, sparsely hirsute. Labrum 0.2-0.3 times as long as broad; malar space 0.30.4 times as long as basal mandibular width. Clypeus in profile weakly convex, almost flat, margin blunt; clypeus in front view 1.3— 1.5 times as broad as long, the margin apically subtruncate. Lower face 0.65—0.73 times as broad as long, centrally punctate. Head in dorsal view with genae narrowed behind eyes; posterior ocellus contiguous with eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.70.8 times the basal mandibular width away from mandible. Antenna slender, with 54-60 flagellar segments; 20th segment 2.0—2.4 times as long as broad. Mesoscutum polished, finely punctate, in profile abruptly rounded with anterior margin out-turned; notauli weak but distinctly impressed anteriorly. Mesopleuron polished, the upper part finely punctate, the lower part from punctate to punctostriate; epicnemial carina curved towards anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for less than 0.6 of its length (Fig. 198); scutellum in dorsal view 1.4—1.5 times as long as anteriorly broad, generally uniformly smooth. Metapleuron weakly to moderately convex, polished, rather uniformly finely and sparsely punctate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum usually complete. Propodeum in profile evenly declivous; anterior transverse carina from complete to interrupted lateromedially, posterior transverse carina absent, or more usually vestigial and discernible as an oblique ridge laterally; anterior area quite long, striate; spiracular area short to moderately long, smooth or finely punctate; posterior area with weak rugosities, anterolaterally almost coriaceous, centrally with one or more weak to distinct longitudinal wrinkles; lateral longitudinal carina from complete to present only anteriorly, joined to spiracular margin by a short carina. Fore wing length 15—18 mm; discosubmarginal cell as in Fig. 312; AI = 0.85-1.00; CI = 0.29-0.44; ICI = 0.50-0.79; SDI = 1.11—1.29; cu-a proximal to the base of Rs&M by 0.2-0.4 times its own length; marginal cell proximally from uniformly hirsute to with a narrow glabrous area adjacent to Rs+2r; 1st subdiscal cell with anterior and distal parts broadly hirsute, often with only a narrow glabrous area posteriorly. Hind wing with 6-7 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa weakly bowed. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.3—-1.6 times length of the shorter. Hind leg with coxa in profile 1.7—1.9 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 1.9-2.5 times as long as broad; claws of female moderately long, abruptly curved, with close pectinae; claws of male similar but with pectinae slightly shorter and closer together. Gaster slender; tergite 2 in profile 4.6-6.0 times as long as posteriorly deep, laterotergite folded under, thyridia oval to oval/elliptical and separated from anterior margin of tergite by about 4-5 times its own length. Ovipositor quite stout, strongly tapered, its sheath slender. Male with sternites 7-9 bearing long stout erect pubescence; gonosquama distally rounded. Colour generally orange-brown, with most of head and scutellum paler yellowish and with mesoscutal vittae blackish; interocellar area yellowish, generally blackish between posterior ocelli, often with blackish marks on the frons between the median ocellus and the antennal socket; antenna black; pterostigma brownish; wings weakly infumate. VARIATION. The specimens from San Ramon, Monteverde and Fortuna have tergites 3+ of the gaster distinctly infuscate. The specimen from Venezuela is generally paler yellowish brown, and does not have the interocellar area dark. Occasional individuals have the sculpture on the posterior area of the pro- podeum very weak. One specimen from Braulio Carrillo only has anterior vestiges of the scutellar carina discernible, and has the posterior transverse carina of the mesosternum incomplete centrally. There seem to be two forms of central sclerite. Most individuals have a subcircular, quite large and weakly sclerotized central sclerite whereas a number of other individuals have this sclerite smaller, more strongly sclerotized and somewhat comma-shaped. REmakkKS. This species is named in honour of Liz Kellogg for her untiring efforts on behalf of Guanacaste National Park, Costa Rica, and in gratitude for her assistance in staying mobile. 260 IAN D. GAULD Enicospilus kelloggae is not a particularly easy species to recognize. Its most distinctive feature is the swollen lower tooth of the mandible, but this requires practice to appreciate. It may be separated from other species by the combination of short scutellar carinae and by the more or less complete distal sclerite. Structurally it is quite similar to E. lacsa; both species are similarly sculptured, particularly on the metapleuron, but E. /acsa has more slender antennae and lacks a distinct distal sclerite. E. kelloggae has a rather similar shaped mandible to E. /uisi. E. luisi may most easily be distinguished from E. kelloggae by the long hairs present on vein 1A in the fore wing; such hairs are not present on the wings of E. kelloggae. BIOLOGICAL INFoRMATION. Enicospilus kelloggae is a fairly widely distributed species whose range extends from near Huixtla, about 15°N in southern Mexico, south to Venezuela. It has most frequently been collected in humid forests at altitudes between 700 and 1350 m. Nothing is known of the biology of this species. MATERIAL EXAMINED Holotype 9, Costa Rica: Alajuela Prov., Virgen de Socorro, 800 m, ii.1987 (Janzen & Hallwachs) (BMNH). Paratypes. Costa Rica: Alajuela Prov.: 1 O&’, Finca Campana, 5 km NW. Dos Rios, 750 m, iii.1985 (Janzen & Hallwachs) (BMNH); 1 9, Finca San Gabriel, 3 km W. Dos Rios, 850 m, vi.1986 (Gauld) (BMNH); 1 co’, 1 2, San Ramon Reserve, Rio San Lorencito, 800 m, ii.1987 (Chacon) (BMNH); 1 Cc’, same locality, v.1987 (Chacon) (BMNH); 2 o’, San Vito, Las Cruces, xi.1987 (Chacon) (BMNH); 1 9, Virgen de Socorro, 800 m, ii.1987 (Janzen & Hallwachs) (BMNH): Cartago Prov.: 1 9, Turrialba, 700 m, vii.1965 (Real) (CAS): Guanacaste Prov.: 1 Oo’, Estacion Mengo on SW. side of Volcan Cacao, 1100 m, v.1987 (Janzen) (BMNH): Puntarenas Prov.: 1 2, Monteverde, 1350 m, xii.1985 (Haber) (BMNH): San José Prov.: 2 2, Estacion Carrillo, Braulio Carrillo National Park, 700 m, x. 1984 & v.1985 (Chacon) (BMNH). Mexico: Chiapas: 1 2, 32 km N. Huixtla, 1000 m, vi.1969 (Mason) (CNC). Panama: 1 2, Barro Colorado Island, 120 m, x.1984 (Wolda) (BMNH); 2 9, Chiriqui, Fortuna, 1050 m, iv & x.1978 (Wolda) (RNH). Venezuela: 1 9, El Junquito, D.F., i.1939 (Berthier) (TC). Enicospilus mengoi sp. n. (Fig. 314) Description. Mandibles rather short, distally fairly evenly tapered, apically twisted 20—-30°; upper man- dibular tooth slender, 1.3 times as long as the lower tooth; outer mandibular surface centrally almost flat, proximally with a weak concavity, and sparsely pubescent. Labrum 0.2 times as long as broad; malar space 0.2 times as long as basal mandibular width. Clypeus in profile moderately convex, margin blunt; clypeus in front view 1.3—1.4 times as broad as long, the margin weakly convex apically. Lower face 0.61—0.70 times as broad as long, rather flat, centrally sparsely punctate. Head in dorsal view with genae constricted behind eyes; posterior ocellus contiguous with eye; FI = 80%; occipital carina mediodorsally complete, flattened slightly, ventrally curved to join hypostomal carina about 0.8 times the basal mandibular width away from mandible. Antenna slender, with 52-54 flagellar segments; 20th segment 2.3—2.4 times as long as broad; distal flagellar segments apparently with shorter sensilla than is usually encountered in species of this genus. Mesoscutum polished, with vestigial punctures, in profile evenly rounded but with anterior margin out- turned; notauli weak. Mesopleuron polished, the upper part finely punctate, with a few irregular striae, the lower part with numerous wrinkle-like striae present; epicnemial carina strong, curved towards anterior margin of pleuron, but with upper end evanescent. Scutellum in profile weakly rounded, laterally carinate for most of its length; scutellum in dorsal view 1.6 times as long as anteriorly broad, rather smooth. Metapleuron weakly convex, smooth with very fine punctures; submetapleural carina fairly evenly ante- riorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area quite long, striate; spiracular area long, smooth; posterior area with weak irregular wrinkling, with a moderate to strong median longitudinal ridge; lateral longitudinal carina complete, joined to spiracular margin by a short weak carina. Fore wing length 15-17 mm; discosubmarginal cell as in Fig. 314; AI = 0.64-0.85; CI = 0.28-0.33; ICI = 0.34-0.41; SDI = 1.05; cu-a proximal to the base of Rs&M by 0.2-0.3 times its own length; marginal cell proximally fairly uniformly hirsute; 1st subdiscal cell anterodistally broadly hirsute. Hind wing with 6—7 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa almost straight. Fore leg with tibia slightly flattened, with fine scattered spines on outer surface. Mid leg with longer tibial spur 1.5—-1.6 times length of the shorter. Hind leg with coxa in profile 1.9 times as long as deep; OPHIONINAE OF TROPICAL MESOAMERICA 261 trochantellus dorsally <0.1 times as long as broad; 4th segment of hind tarsus 2.2 times as long as broad; claws of female quite long with rather close pectinae. Gaster slender; tergite 2 in profile 5.1—5.6 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 4 times its own length. Ovipositor concealed, its sheath narrow. Male unknown. Colour generally orange-brown, with head, pronotum, mesoscutal stripes, upper part of mesopleuron and scutellum yellowish; gaster with tergites 3+ blackish; interocellar area yellowish, with part slightly infuscate, and with an infuscate band extending from median ocellus to between antennal sockets; antenna black; pterostigma yellowish brown; wings slightly yellowish. VARIATION. None remarkable. REMARKS. This species is named after Pedro ‘Mengo’ Mejilla in recognition of his pioneer spirit in establishing the Mengo Biological Station on Volcan Cacao, Costa Rica. Enicospilus mengoi can be recognized by the form of the alar sclerites — the central sclerite is rather small and circular (see Fig. 314) and by the sculpture on the propodeum. Its relationship with other species is not clearly understood. BIOLOGICAL INFORMATION. Enicospilus mengoi is only known to occur in Costa Rica where two females have been collected at light in forests at altitudes between 700 and 1100 m. Nothing is known about the biology of this insect. MATERIAL EXAMINED Holotype 2, Costa Rica: Guanacaste Prov.: 1 2, Casa [= Estacion] Mengo, SW. side of Volcan Cacao, 1100 m, vii.1987 (Janzen) (BMNH). Paratype. Costa Rica: San José Prov.: 1 9, Estacion Carrillo, Braulio Carrillo National Park, 700 m, iv.1985 (Chacon & Chacon) (BMNH). Enicospilus haberi sp. n. (Fig. 313) Description. Mandibles moderately long, distally evenly narrowed, apically twisted 40-50°; upper man- dibular tooth slightly depressed, 1.5 times as long as the lower tooth; outer mandibular surface flat, with scattered hairs; proximal concavity weak. Labrum 0.3 times as long as broad; malar space 0.2 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.41.5 times as broad as long, the margin apically weakly convex. Lower face 0.68—0.74 times as broad as long, punctate. Head in dorsal view with genae constricted behind eyes; posterior ocellus very close to eye; FI = 75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.8-0.9 times the basal mandibular width away from mandible. Antenna slender, with 58 flagellar segments; 20th segment 2.1—2.2 times as long as broad. Mesoscutum polished, finely punctate, in profile quite abruptly rounded; notauli weakly impressed near anterior margin. Mesopleuron polished, the upper part punctate, the lower part punctostriate; epicnemial carina inclined towards anterior margin of pleuron, its upper end evanescent. Scutellum in profile weakly convex, laterally carinate for 0.8 of its length; scutellum in dorsal view 1.4 times as long as anteriorly broad, anteriorly smooth, posteriorly with isolated wrinkles. Metapleuron moderately convex, anteroventrally punctate, posterodorsally striate with a few rugae present also; submetapleural carina barely broadened anteriorly; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area quite long, striate; spiracular area moderately long, smooth; posterior area irregularly rugose, with the central rugae tending to be longitudinally aligned; lateral longitudinal carina present only anteriorly, joined to spiracular margin by a short carina. Fore wing length 14-15 mm; discosubmarginal cell as in Fig. 313; AI = 1.00-1.05; CI = 0.26-0.28; ICI = 0.46-0.47; SDI = 1.10—1.23; cu-a proximal to the base of Rs&M by 0.1-0.2 times its own length; marginal cell proximally slightly more sparsely hirsute than it is centrally; 1st subdiscal cell with scattered hairs centrally and distally. Hind wing with 8 hamuli on R1; 1st abscissa of Rs almost straight, 2nd abscissa slightly bowed. Fore leg with tibia slightly flattened, with isolated spines on outer surface. Mid leg with longer tibial spur 1.4-1.5 times length of the shorter. Hind leg with coxa in profile 1.7 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of hind tarsus 2.2—2.3 times as long as broad; claws of female abruptly curved, with long close pectinae. 262 IAN D. GAULD Gaster slender; tergite 2 in profile 6.7—7.2 times as long as posteriorly deep, laterotergite folded under, thyridia shortly elliptical and separated from anterior margin of tergite by about 4-5 times its own length. Ovipositor quite stout, slightly decurved, its sheath slender. Male unknown. Colour generally orange-brown, with head paler yellow and with mesoscutum bearing three longitudinal dark brown vittae; gaster posteriorly slightly infuscate; interocellar area yellowish; antenna black, with distal apices brownish; pterostigma brown; wings more or less hyaline. VARIATION. None remarkable. REMARKS. This species is named in honour of Bill Haber in recognition of his work on Costa Rican insects, and as a gesture of thanks for the numerous ophionines he has collected for this study. Enicospilus haberi is a rather delicate, small species that can be distinguished from other taxa that have black antennae, a yellow interocellar area and a central sclerite by the fact that the distal sclerite is strongly sclerotized and separated from the proximal sclerite (Fig. 313). Furthermore the proximal sclerite is rather more elongately triangular than that of other species. The entire complex of alar sclerites and fenestra is longer than that of many other species with dark antennae. In this respect E. haberi resembles E. forsythei, but E. forsythei does not have a discrete distal sclerite. BIOLOGICAL INFORMATION. Enicospilus haberi is only known from Monteverde, Costa Rica. The two females were collected at light near the edge of an area of cloud forest at an altitude of 1350 m. Nothing is known of their biology. MATERIAL EXAMINED Holotype 9, Costa Rica: Puntarenas Prov., Monteverde, 1350 m, v.1986 (Haber) (BMNH). Paratype. 1 2, same locality and collector as holotype, ii.1986 (BMNH). Enicospilus lacsa sp. n. (Figs 200, 316, 317, 318, 345) Description. Mandibles moderately long, quite strongly narrowed, with a weak proximoventral lobe, apically twisted 20-45°; upper mandibular tooth slender, depressed, 1.3—1.6 times as long as the lower tooth; outer mandibular surface bearing isolated pubescence, almost flat. Labrum 0.2-0.3 times as long as broad; malar space 0.2-0.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin subacute; clypeus in front view quite long, 1.1—-1.3 times as broad as long, its margin convex apically. Lower face narrow, 0.61—0.68 times as broad as long, centrally with isolated punctures. Head in dorsal view with genae strongly constricted behind eye; posterior ocellus close to eye; FI = 60-70%; occipital carina mediodorsally complete, ventrally sinuously curved to join hypostomal carina about 0.5— 0.7 times the basal mandibular width away from mandible. Antenna very long and slender, with 56-61 flagellar segments; 20th segment 2.5—3.0 times as long as broad. Mesoscutum polished, finely and sparsely punctate, in profile abruptly rounded and with anterior margin often slightly out-turned; notauli vestigial. Mesopleuron polished, from uniformly punctate to ventrally with traces of punctostriation; epicnemial carina very strong, curved towards anterior margin of pleuron which it almost reaches. Scutellum in profile weakly convex, laterally carinate for 0.5 or more of its length; scutellum in dorsal view 1.4-1.6 times as long as anteriorly broad, punctate with posterior end bearing some longitudinal wrinkles. Metapleuron weakly to moderately convex, generally regularly punctate, occasionally punctostriate; submetapleural carina evenly anteriorly broadened; posterior trans- verse carina of mesosternum complete, distinctly produced anteriorly centrally so it has a V-shaped notch on midline. Propodeum in profile rather long and weakly declivous posteriorly; anterior transverse carina complete, posterior transverse carina vestigial or absent; anterior area moderately long, striate; spiracular area long, more or less smooth; posterior area rather finely irregularly wrinkled-rugose, with rugae centrally subparallel, longitudinally arranged; lateral longitudinal carina usually more or less complete, rarely interrupted posteriorly, joined to spiracular margin by a short carina. Fore wing length 12-16 mm; discosubmarginal cell as in Figs 316-318, 345; AI = 0.90-1.68; CI = 0.16— 0.46; ICI = 0.35—0.47; SDI = 1.13-1.49; cu-a proximal to the base of Rs&M by 0. 1-0.3 times its own length; marginal cell proximally more or less evenly hirsute; 1st subdiscal cell with distal 0.6 sparsely hirsute. Hind wing with 5-8 hamuli on R1; 1st abscissa of Rs almost straight, 2nd abscissa more or less straight. Fore leg with tibia slightly flattened, with scattered fine spines on outer surface. Mid leg with longer tibial spur 1.5-1.7 times length of the shorter. Hind leg with coxa in profile 1.7-1.9 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.0—2.2 times as long as broad; claws of female evenly curved with quite long, moderately close pectinae, those of male similar but with pectinae shorter. OPHIONINAE OF TROPICAL MESOAMERICA 263 Gaster slender; tergite 2 in profile 5.1—6.9 times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by about 34 times its own length. Ovipositor slender, its sheath narrow. Male with sternite 7 and extreme anterior part of sternite 8 bearing lateral longitudinal rows of close stout erect hairs; remainder of sternite 8 glabrous, with apical margin strongly concave; sternite 9 bearing fine decumbent pubescence (Fig. 200); gonosquama distally rounded. Colour particularly variable, generally reddish brown, with head paler yellow and gastral tergites 3+ variously infuscate; interocellar area yellowish; antenna from brownish yellow to black; pterostigma brown; wings more or less hyaline. VARIATION. The most striking variation observable in this species is in colour. The colour pattern described above (reddish brown without mesoscutal vittae and with posterior segments of the gaster blackish) applies to about half the individuals studied. Several of the others are generally paler orange yellow with darker brown mesoscutal vittae, and with the terminal segments of the gaster only indistinctly infuscate. Others are darker, but have discernible mesoscutal vittae. A few individuals have the antennae brownish yellow rather than black. There is also considerable variation in the size of the alar sclerites. The central sclerite of the specimen from Finca La Selva, Costa Rica, is small and quite slender, and the distal sclerite is absent (Fig. 318). Most other specimens have the central sclerite stouter, but several lack the distal sclerite (Fig. 317), whilst others have it clearly discernible as a small fleck distally. The specimens from Santa Rosa National Park are somewhat atypical in that they have a slightly longer fenestra, a more heart-shaped central sclerite (Fig. 316) and brown antennae. Remarks. This species is named after LACSA, the national airline of Costa Rica in recognition of the quality service rendered to biologists for many years. Enicospilus lacsa can be distinguished from all other species with black antennae and a yellowish interocellar area by the combination of a comma-shaped central sclerite and the long slender antennae. The only other Central American species that has a similar arrangement of hair on sternites 7-8 is E. echeverri and these two species are probably closely related (see that species). Structurally E. lacsa also resembles E. kelloggae (see that species). BIOLOGICAL INFORMATION. Enicospilus lacsa is a widespread species (Map 29) whose range extends from about 19° in southern Mexico south to Santa Catarina Province in Brazil (27°S). It has been collected at low and medium elevation sites between 100 and 1400 m. In Central America E. lacsa is comparatively rarely collected, and in Santa Rosa National Park only isolated individuals have been taken at the end of the wet season (December-January), later than its relative E. echeverri. Nothing is known about the biology of this species. MATERIAL EXAMINED Holotype ©’, Costa Rica: Heredia Prov., El Angel waterfall, 8.2 km downhill Vara Blanca, 1350 m, i.1981 (Janzen & Hallwachs) (BMNH). Paratypes. Bolivia: 1 9 , Santa Cruz, General Saavedra, xi.1974 (Porter & Stange) (FSCA). Brazil: 1 2, Bahia, Encruzilhada, 980 m, xi.1973 (Alvarenga) (TC); 1 9, Minas Gerais, Pedra Azul, xi.1970 (Oliveira) (TC); Santa Catarina, Nova Teutonia, 27°11’S, 52°23’W, 300-500 m, 1 CO’, ii.1936, 1 Q, ii.1938, 107,19, xi.1938, 1 CO’, ii. 1939, 1 O’, xii.1968, 1 9, i.1966 (Plaumann) (BMNH & CNC); same locality, 1 2, xi.1952, 2 Q, 1.1953, 1 O, 3 Q, xi-xii.1953, 1 9, xi.1970, 1 9, iv.1971 (Plaumann) (TC). Colombia: 1 C’, Buenaventura, Zabaleta — Rio Dagua forest road, 100 m, ix.1971 (Cooper) (BMNH); 1 0’, Caqueta, Florencia, i.1979 (Cooper) (BMNH); 1 9, Valle, Anchicaya, xii.1977 (Tidwell) (FSCA); 1 9, Valle, Anchicaya, 30 km E. Buenaventura, tropical wet forest, 560 m, vii.1975 (Wilkerson) (FSCA); 10,1 9, Valle, Rio Zabletas, Las Piedras, viii.1975 (Wilkerson) (FSCA). Costa Rica: Alajuela Prov.: 1 0’, San Ramon Forestry Reserve, 5 km N. Col. Palmarena, 900 m, v.1985 (Chacon) (BMNH); 1 2, San Ramon Forestry Reserve, Rio San Lorencito, 800 m, iii.1987 (Chacon) (BMNH); 1 @, Virgen de Socorro, 800 m, 1.1987 (Janzen & Hallwachs) (BMNH): Cartago Prov.: 1 0’, Tapanti, Rio Grande de Orosi, 13-1400 m, xi. 1982 (Janzen & Hallwachs) (BMNH); Cartago Prov.: 1 9, Turrialba, iii.1965 (Duckworth) (USNM): Guanacaste Prov.: 2 2, Santa Rosa National Park, 300 m, xii.1979 (Janzen) (TC); same locality, 1 0,19, 1.1983, 1 CO, 1.1984 (Janzen & Hallwachs) (BMNH): Heredia Prov.: 1 9, Finca La Selva, 3 km S. Puerto Viejo, vii. 1986 (Hespenheide) (BMNH): Puntarenas Prov.: 3 9, Monteverde, 1350 m, vii.1982 (Janzen & Hallwachs) (BMNH); 1 9, same locality, iii.1986 (Haber) (BMNH); 1 2, San Vito de C., S. of Las Cruces, 1200 m, vii.1983 (Gill) (TC): San José Prov.: 1 O’, Estacion Carrillo, Braulio Carrillo National Park, 700 m, ix.1984 (Chacon) (BMNH); 1 ©’, same locality and collector, i.1985 (BMNH); 1 o’, 2 9, Braulio Carrillo National Park, 500 m, iv.1985 (Goulet & Masner) (CNC). Ecuador: 1 2, Los Rios, Rio Palenque, vi.1974 (Longino) (FSCA); 1 O’, 1 9, Pichincha, Santo Domingo, 680 m, v.1975 (Peck) (TC); 1 9, 264 IAN D. GAULD Map 29 Localities at which Enicospilus lacsa has been collected. Pichincha, 47 km S. Santo Domingo, Rio Palenque Sta., vii.1975 (Forsyth) (CNC). Guyana: 2 9, Essequibo River, Moraballi Creek, viii-x.1929 (BMNH). Mexico: Chiapas: 1 0’, 32 km N. Huixtla, 1000 m, vi. 1969 (Mason) (CNC); 1 9, Muste, nr Huixtla, 440 m, (Welling) (CNC). Panama: Barro Colorado Island, 1 Q, iv.1978, 1 CO’, ii.1984, 1 9, v.1985 (Wolda) (BMNH & RNH); Chiriqui, Fortuna, 1050 m, 2 C’, i-ii.1978, 1 9, iv.1978, 1 Q, iv.1979 (Wolda) (RNH); 1 2, Las Cumbres, xii.1981 (Wolda) (TC). Peru: 2 9, Madre de Dios, Avispas, 400 m, ix.1962 (Peria) (CNC); 10,2 2, Quincemil, nr Marcapata, 750 m, xi. 1962 (Pena) (TC). Surinam: 1 ©’, Maratakka River, upper course, iii.1971 (Geijskes) (RNH). OPHIONINAE OF TROPICAL MESOAMERICA 265 Enicospilus brenesiae sp. n. (Figs 135, 137, 315) Description. Mandibles long, proximally abruptly narrowed, and with a pronounced proximoventral lobe; mandibles distally parallel-sided, apically twisted 20° (Fig. 137); upper mandibular tooth subcylindrical, 1.2 times as long as the lower tooth; outer mandibular surface with a weak median longitudinal concavity that bears scattered pubescence. Labrum 0.3 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile almost flat, margin blunt; clypeus in front view 1.4 times as broad as long, with margin apically truncate. Lower face 0.83—0.85 times as broad as long, closely and rather coarsely punctate, centrally tending to punctostriate. Head in dorsal view with genae rounded behind eyes; posterior ocellus contiguous with eye; FI = 67%; occipital carina mediodorsally complete, but weak, ventrally curved to join hypostomal carina about 0.6 times the basal mandibular width away from mandible. Antenna slender, with 64—65 flagellar segments; 20th segment 1.5 times as long as broad. Mesoscutum polished, closely and coarsely punctate, in profile abruptly rounded; notauli vestigial. Mesopleuron polished, the upper part punctate, ventrally grading to punctostriate; epicnemial carina curved towards anterior margin of pleuron, but with its upper end evanescent. Scutellum in profile weakly convex, laterally carinate for 0.8 of its length; scutellum in dorsal view 1.5 times as long as anteriorly broad, punctate. Metapleuron moderately convex, very closely and coarsely punctate (Fig. 135); submetapleural carina narrow, barely broadened anteriorly; posterior transverse carina of mesosternum complete. Pro- podeum in profile evenly declivous; anterior transverse carina weak, lateromedially discontinuous, pos- terior transverse carina vestigial; anterior area moderately long, shallow, striate; spiracular area short, punctate; posterior area weakly rugose, the rugae tending to be longitudinal centrally; lateral longitudinal carina absent except for a short anterior vestige, not joined to spiracular margin by a short carina. Fore wing length 18 mm; discosubmarginal cell as in Fig. 315; AI = 1.00-1.25; CI = 0.37-0.44; ICI = 0.42- 0.56; SDI = 1.23-1.29; cu-a proximal to the base of Rs&M by about 0.1 times its own length; marginal cell proximally broadly glabrous; 1st subdiscal cell with anterior and distal margins sparsely hirsute. Hind wing with 7 hamuli on R1; 1st abscissa of Rs almost straight, 2nd abscissa weakly arcuate. Fore leg with tibia slightly flattened, with isolated slender spines on outer surface. Mid leg with longer tibial spur 1.4 times length of the shorter. Hind leg with coxa in profile 1.8 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.3 times as long as broad; claws of female moderately long, with close, stout pectinae. Gaster quite slender; tergite 2 in profile 4.6 times as long as posteriorly deep, laterotergite folded under, thyridia oval and separated from anterior margin of tergite by about 3.5 times its own length. Ovipositor moderately stout, apically slightly decurved, its sheath narrow. Male unknown. Colour generally reddish brown, mesoscutum with three blackish vittae; gaster with tergites 3+ dark brown; interocellar area slightly infuscate close to ocelli; antenna basally black, but with flagellum distal to segment 3 brown; pterostigma reddish brown; wings weakly infumate. REMARKS. This species is named in honour of Liz Brenes for her efforts in the establishment of the education programme at Santa Rosa National Park, Costa Rica. Enicospilus brenesiae can be recognized most easily by the form of the alar sclerites — the narrowly crescentic central sclerite is very characteristic (Fig. 315), and by the stout mandibles (Fig. 137). It is generally a stouter and more coarsely sculptured species than many others with black antennae. BIOLOGICAL INFORMATION. Enicospilus brenesiae has only been collected during May in cloud forest at 1350 m in Monteverde Reserve, Costa Rica. Nothing is known about its biology. MATERIAL EXAMINED Holotype 9, Costa Rica: Puntarenas Prov., Monteverde, 1350 m, v.1980 (Janzen & Hallwachs) (BMNH). Paratype. Costa Rica: 1 2, same locality as holotype, v.1983 (Haber) (BMNH). Enicospilus forsythei sp. n. (Figs 201, 202, 320) Description. Mandibles moderately long, proximally strongly narrowed, distally weakly tapered, apically twisted 20—-30°; upper mandibular tooth quite slender, 1.3—1.5 times as long as the lower tooth; outer mandibular surface centrally slightly concave, with a broad shallow proximal concavity, the whole bearing fine sparse pubescence. Labrum 0.2-0.3 times as long as broad; malar space 0.2 times as long as basal mandibular width. Clypeus in profile moderately convex, margin blunt; clypeus in front view 1.3-1.4 times 266 IAN D. GAULD as broad as long, with margin very weakly convex apically. Lower face 0.70—-0.75 times as broad as long, laterally punctate, centrally punctostriate. Head in dorsal view with genae constricted behind eyes; posterior ocellus very close to eye; FI = 65-70%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6-0.7 times the basal mandibular width away from mandible. Antenna long and slender, with 53-55 flagellar segments; 20th segment 2.1—2.4 times as long as broad. Mesoscutum weakly polished, with close shallow punctures, in profile fairly evenly rounded but with anterior margin slightly out-turned; notauli weak. Mesopleuron polished, the upper part finely punctate, the lower part punctostriate; epicnemial carina strong, curved towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.5 times as long as anteriorly broad, generally smooth, with a few striae posteriorly. Metapleuron weakly convex, anteroventrally puncto-coriaceous becoming weakly rugulose-punctate posterodorsally (Fig. 202); submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina sinuous, generally complete, rarely interrupted lateromedially, posterior transverse carina absent; anterior area moderately long, striate; spiracular area short, finely punctate; posterior area coarsely rugose-striate, generally with the rugae more or less concentric (Fig. 201); lateral longitudinal carina present anteriorly, rarely almost complete, not joined to spiracular margin by a short carina. Fore wing length 15-16 mm; discosubmarginal cell as in Fig. 320; AI = 0.76-1.04; CI = 0.17-0.22; ICI = 0.48-0.53; SDI = 1.00-1.07; cu-a usually opposite to the base of Rs&M but occasionally proximal to it by 0.1 times its own length; marginal cell proximally uniformly hirsute; 1st subdiscal cell extensively but sparsely hirsute anterodistally. Hind wing with 6-8 hamuli on R1; 1st abscissa of Rs weakly curved, 2nd abscissa weakly bowed. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.4~-1.5 times length of the shorter. Hind leg with coxa in profile 1.6-1.7 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.5—2.6 times as long as broad; claws of female quite long, apically evenly rounded and bearing moderately long, regularly spaced pectinae; claws of male similar, but with pectinae shorter. Gaster very slender; tergite 2 in profile 6.1—6.9 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 4 times its own length. Ovipositor quite stout though with a slender, slightly decurved apex, its sheath narrow. Male with sternites 7-9 bearing very long scattered erect hairs amid shorter, semidecumbent pubescence. Colour generally reddish brown, with head slightly more yellowish, tergites 3+ of gaster black, often with ventral margin of tergite 3 and sometimes also of 4 yellowish; interocellar area yellowish, generally slightly infuscate between posterior ocelli; antenna black; pterostigma brownish; wings weakly infumate. VARIATION. A morphologically rather uniform species except for the variation outlined above. ReMaRKS. This species is named in honour of Adrian Forsythe for his efforts to raise conservation conciousness in the visitor to Costa Rica. Enicospilus forsythei may be recognized by the combination of concentrically striate propodeum, weakly twisted mandibles, characteristic alar sclerites (see Fig. 320), alar indices and colour pattern. Structurally it resembles E. burgosi and the two species may be closely related. BIOLOGICAL INFORMATION. Enicospilus forsythei is a Central American species whose range extends from southern Mexico to Costa Rica. The Costa Rican specimens have all been collected at a moderate elevation site adjacent to cloud forest. MATERIAL EXAMINED Holotype 9, Costa Rica: Puntarenas Prov., Monteverde, ii.1986 (Haber) (BMNH). Paratypes. Costa Rica: Puntarenas Prov.: 3 2, Monteverde, 1350 m, ix.1985-ii.1986 (Haber) (BMNH); 10, 1 Q, same locality, i-ii.1986 (Forsyth) (CNC). Mexico: Chiapas: 1 9, 3.2 km NE. Bochil, vi.1969 (Campbell) (CNC). Enicospilus fogdenorum sp. n. (Figs 203, 204, 321) DescriPTIoNn. Mandibles moderately long, strongly narrowed proximally, with a distinct basal lobe, distally weakly narrowed, apically twisted 35—45°; upper mandibular tooth slender, 1.2-1.5 times as long as and slightly divergent from the lower tooth; outer mandibular surface flat, punctate, bearing fine hairs. Labrum 0.2-0.3 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile almost flat, margin blunt; clypeus in front view 1.4-1.6 times as broad as long, the margin almost OPHIONINAE OF TROPICAL MESOAMERICA 267 truncate apically. Lower face 0.67—-0.70 times as broad as long, polished, centrally punctostriate. Head in dorsal view with genae evenly constricted behind eye; posterior ocellus contiguous with eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.5 times the basal mandibular width away from mandible. Antenna long and slender, with 60—65 flagellar segments; 20th segment 1.9-2.0 times as long as broad. Mesoscutum polished, finely punctate, in profile abruptly rounded; notauli vestigial. Mesopleuron polished, the upper part finely punctate, the lower part similar but with a few wrinkles; epicnemial carina with upper end curved towards anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for 0.7 or more of its length; scutellum in dorsal view 1.6—1.7 times as long as anteriorly broad, punctate. Metapleuron posterodorsally with coarse rugae that are confluent with rugae on pro- podeum, anteroventrally punctate (Fig. 204); submetapleural carina evenly anteriorly broadened; pos- terior transverse carina of mesosternum complete. Propodeum in profile abruptly declivous; anterior transverse carina complete, posterior transverse carina represented by oblique lateral crests; anterior area quite long, with isolated rugae; spiracular area short, generally rather smooth; posterior area coarsely rugose wrinkled, with three longitudinal rugae centrally (Fig. 203); lateral longitudinal carina only present from anterior margin to just behind anterior transverse carina, behind this absent, anteriorly not joined to spiracular margin by a short carina. Fore wing length 18-21 mm; discosubmarginal cell as in Fig. 321; AI = 0.70—0.77; CI = 0.25-0.26; ICI = 0.55-0.57; SDI = 1.15—1.17; cu-a proximal to the base of Rs&M by about 0.2 times its own length; marginal cell proximally fairly uniformly hirsute though with a narrow glabrous area adjacent to Rs+2r; 1st subdiscal cell with anterior 0.5 and distal corners hirsute. Hind wing with 8 hamuli on R1; 1st and 2nd abscissae of Rs straight. Fore leg with tibia weakly flattened, with fine, scattered spines on outer surface. Mid leg with longer tibial spur 1.5 times length of the shorter. Hind leg with coxa in profile 1.9 times as long as deep; trochantellus dorsally occluded; 4th segment of tarsus 2.3—2.4 times as long as broad; claws of female large, strongly curved with close short pectinae, those of male very similar. Gaster slender; tergite 2 in profile about 6 times as long as posteriorly deep, laterotergite turned under, thyridia oval and separated from anterior margin of tergite by about 5 times its own length. Female with ovipositor rather stout, apically slender and slightly decurved; ovipositor sheath narrow. Male with sternites 7-9 bearing dense long erect pubescence; gonosquama long, but evenly rounded distally. Colour generally reddish brown with tergites 3+ blackish; interocellar area yellowish; antenna black, distally brownish; pterostigma brown; wings weakly infumate. VARIATION. None remarkable. REMARKS. This species is dedicated to Mike and Tricia Fogden as a gesture of thanks for their hospitality and help collecting specimens at light. Enicospilus fogdenorum can most easily be recognized by its large size, distinctive colour pattern, propodeal crests and by the presence of a more or less D-shaped central sclerite in the fenestra of the fore wing. E. fogdenorum is similar in many features to several other species with black antennae, including E. erasi, E. galilea and E. hubbelli. All have large value of ICI, somewhat similar mesopleural sculpture, claws and male terminal sternites. Of these species, E. fogdenorum most closely resembles E. erasi which also possesses propodeal crests. It differs from E. erasi in colour pattern and in having less twisted mandibles. BIOLOGICAL INFORMATION. Enicospilus fogdenorum is only known to occur in Costa Rica where it has been collected in lower montane cloud forest at Monteverde. Nothing is known of the host range of this species. MATERIAL EXAMINED Holotype ©’, Costa Rica: Puntarenas Prov., Monteverde, 1300-1400 m, vii.1982 (Janzen & Hallwachs) (BMNH). Paratype. 1 2, same locality and collectors, 1500 m, i.1984 (BMNH). Enicospilus burgosi sp. n. (Figs 138, 205, 322, 343) Description. Mandibles moderately long, proximally strongly constricted, distally weakly narrowed, apically twisted 25—35°; upper mandibular tooth slightly depressed, quite slender, 1.3—1.5 times as long as the lower tooth; outer mandibular surface with a broad shallow proximal concavity that continues distally into a weakly concave area extending about half way along the mandible, and with this area sparsely pubescent; upper edge of mandible often margined by a very weak ridge. Labrum 0.2-0.3 times as long as broad; malar space 0.3—-0.4 times as long as basal mandibular width. Clypeus in profile weakly convex, 268 IAN D. GAULD margin blunt; clypeus in front view 1.2—1.4 times as broad as long, with margin subtruncate apically. Lower face 0.66-0.77 times as broad as long, centrally sparsely punctate. Head in dorsal view with genae rounded behind eyes; posterior ocellus close to eye; FI = 60-70%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6—0.7 times the basal mandibular width away from mandible. Antenna long and slender, with 52-57 flagellar segments; 20th segment 1.8-2.3 times as long as broad. Mesoscutum polished, finely punctate, in profile abruptly rounded with margin out-turned; notauli weakly impressed anteriorly. Mesopleuron polished, the upper part punctate, the lower part grading from punctate to punctostriate to striate; epicnemial carina inclined towards anterior margin of pleuron, with its upper end evanescent. Scutellum in profile weakly convex, usually laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.4—1.5 times as long as anteriorly broad, smooth, with isolated punctures, but with a few wrinkles posteriorly. Metapleuron weakly convex, diagonally striate or coriaceous to coriaceous-striate (Fig. 205); submetapleural carina fairly evenly broadened anteriorly; posterior trans- verse carina of mesosternum complete. Propodeum in profile evenly rounded; anterior transverse carina complete, posterior transverse carina absent; anterior area quite long, striate; spiracular area fairly short, smooth with fine punctures; posterior area rugose-reticulate to rugose-striate, with the rugae tending to be concentric posteriorly; lateral longitudinal carina present before anterior transverse carina, incomplete posteriorly, joined to spiracular margin by a short carina. Fore wing length 11-15 mm; discosubmarginal cell as in Figs 322, 343; AI = 0.90-1.05; CI = 0.18-0.26; ICI = 0.38-0.49; SDI = 1.08-1.26; cu-a proximal to the base of Rs&M by 0.1-0.2 times its own length; marginal cell proximally sparsely to very sparsely pubescent; 1st subdiscal cell with anterodistal half hirsute. Hind wing with 4~7 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa weakly curved. Fore leg with tibia weakly flattened, with fine, scattered spines on outer surface. Mid leg with longer tibial spur 1.4-1.7 times length of the shorter. Hind leg with coxa in profile 1.6-1.8 times as long as deep; trochantellus dorsally 0.1-0.4 times as long as broad (Fig. 138); 4th segment of tarsus 1.8-2.4 times as long as broad; claws of female moderately long, abruptly curved, with regular, moderately long pectinae, those of male similar but with pectinae shorter. Gaster slender; tergite 2 in profile 6.0—7.0 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 4-5 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 usually bearing scattered long, slightly curved, erect hairs; gonosquama slender, distally almost pointed. Colour generally brownish yellow, with mesoscutal vittae dark brown, and with posterior tergites of gaster from very weakly to strongly infuscate; interocellar area yellowish; antenna brown to black; pterostigma brownish; wings hyaline. VARIATION. This is a morphologically rather variable species and possibly several taxa may be confused under a single name here. The typical form is exhibited by the Costa Rica specimens from Santa Rosa National Park. These have the hind trochantellus longer than other individuals, have a ventrally striate mesopleuron, have the fenestra quite long and with a virtually indistinguishable distal sclerite, have elongate pubescence on the male terminal sternites and have the metapleuron coriaceous-striate. Several have the antenna basally brownish, though most usually it is black. The Brazilian specimens have the hind trochantellus dorsally shorter (0.1—0.2 times as long as broad), the metapleuron more finely striate and the distal sclerite discernible. A pair from Belize and a male from Santa Rosa differ in having the mesopleuron almost entirely punctate, the fenestra margined by a strong distal sclerite; the males have rather fine decumbent hair on the posterior sternites. A male from Braulio Carrillo National Park, Costa Rica, has short lateral carinae on the propodeum, and the posterior sternites bear dense long erect pubescence. I am not certain that all these individuals are conspecific, but until more material is available it seems reason- able, at present, to include all together, though I have excluded the more atypical specimens from the paratype series. REMARKS. This species is named in honour of Senor Mario Burgos for his patience in the development of Guanacaste National Park. Enicospilus burgosi can be recognized by the venation and by the mandibles which have a weak but characteristic ridge along their upper margin. BIOLOGICAL INFORMATION. Enicospilus burgosi is a widely distributed but rarely collected species whose range extends from Belize to southern Brazil. It has been collected from sea level up to about 1000 m. In Costa Rica most specimens have been collected in seasonally dry forest in Santa Rosa National Park, the majority, between April (the end of the dry season) and July (well into the beginning of the wet season), though an aberrant male was taken in January. The cumulative seasonal data for this site are: OPHIONINAE OF TROPICAL MESOAMERICA 269 Tift ada Mite site Ty ye Titty Au S24 OtgeN sD Memes mead Ser linet Sp aes ods) dene = os Nothing is known of the biology of this species. MATERIAL EXAMINED Holotype ©’, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, vii.1984 (Janzen & Hallwachs) (BMNH). Paratypes. Belize: 1 9, Augustine, vii.1968 (Hasse) (BMNH). Costa Rica: Guanacaste Prov.: 3 2, Santa Rosa National Park, 300 m, vii & x.1977 (Janzen) (TC); 1 0’, same locality, x.1982 (Janzen & Hallwachs) (BMNH); 1 9, same locality, iv.1983 (Janzen & Hallwachs) (BMNH); 2 9, same locality, iv.1984 (Gauld) (BMNH); 1 9, same locality, v.1984 (Janzen & Hallwachs) (BMNH); 3 9, same locality vi.1985 (Gauld) (BMNH). Non-paratypic material. Belize: 1 0’, 1 9, Middlesex, 125 m, v.1963 (Welling) (CNC). Brazil: 2 9, Bahia, Encruzilhada, 980 m, xi.74 (Alvarenga) (TC); 1 9, Santa Catarina, Nova Teutonia, xi.1970 (Plaumann) (TC). Costa Rica: Guanacaste Prov.: 1 O’, Santa Rosa National Park, 300 m, i.1984 (Janzen & Hallwachs) (BMNH): Puntarenas Prov.: 1 oO’, Finca Las Cruces, 6 km S. San Vito de Java, 1400 m, ix.1986 (Eger) (BMNH): San José Prov.: 1 0’, 1 9, Estacion Carrillo, Braulio Carrillo National Park, 700 m, Enicospilus erasi sp. n. (Figs 134, 136, 217, 323) Description. Mandibles moderately long, distally fairly strongly but evenly narrowed, apically twisted 55-75° (Fig. 136); upper mandibular tooth slightly depressed, 1.4~1.5 times as long as the lower tooth; outer mandibular surface with a weak ridge running from upper proximal corner to near base of upper tooth, the part below this ridge slightly concave and bearing scattered hairs (Fig. 217). Labrum 0.2 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile weakly convex to almost flat, margin subacute; clypeus in front view 1.3—1.4 times as broad as long, the margin apically truncate. Lower face 0.63—-0.66 times as broad as long, centrally sparsely punctate. Head in dorsal view with genae constricted behind eyes; posterior ocellus contiguous with eye; FI = 75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.7 times the basal mandibular width away from mandible. Antenna slender, with 63—64 flagellar segments; 20th segment 1.7—2.0 times as long as broad. Mesoscutum polished, finely punctate, in profile abruptly rounded; notauli weakly impressed ante- riorly. Mesopleuron polished, the upper part punctate, the lower part striate; epicnemial carina curved towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.5—0.7 of its length; scutellum in dorsal view 1.5 times as long as anteriorly broad, anteriorly sparsely punctate, posteriorly with weak longitudinal wrinkles. Metapleuron moderately convex, diagonally striate with coarse rugae present posterodorsally (Fig. 134); submetapleural carina narrow, rather abruptly broadened anteriorly; posterior transverse carina of mesosternum complete, but weak centrally. Propodeum in profile abruptly declivous; anterior transverse carina complete, posterior transverse carina present as vestigial keels laterally; anterior area quite long, striate; spiracular area short, almost smooth; posterior area coarsely irregularly rugose-reticulate, with rugae tending to be longitudinal centrally; lateral long- itudinal carina posteriorly incomplete, joined or not joined to spiracular margin by a short carina. Fore wing length 18-20 mm; discosubmarginal cell as in Fig. 323; AI = 0.76-0.84; CI = 0.25-0.28; ICI = 0.87-0.97; SDI = 1.21-1.36; cu-a proximal to the base of Rs&M by 0.2-0.3 times its own length; marginal cell proximally rather sparsely hirsute adjacent to Rs+2r; 1st subdiscal cell distally hirsute. Hind wing with 8 hamuli on R1; 1st abscissa of Rs almost straight, 2nd abscissa weakly bowed. Fore leg with tibia weakly flattened, with isolated spines on outer surface. Mid leg with longer tibial spur 1.5—1.6 times length of the shorter. Hind leg with coxa in profile 1.8 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.3—2.4 times as long as broad; claws of female abruptly curved, stout, with many, close pectinae; claws of male similar but with pectinae shorter. Gaster slender; tergite 2 in profile 5.4~5.6 times as long as posteriorly deep, laterotergite folded under, thyridia more or less circular and separated from anterior margin of tergite by about 7 times its own length. Ovipositor apically slender, its sheath narrow. Male with sternites 7—9 bearing long close erect pubescence; gonosquamae long, distally tapered, apically rounded. Colour generally golden yellowish brown, with mesoscutal vittae blackish and distal segments of gaster slightly infuscate; interocellar area brownish yellow; antenna black dorsally near bases, distally paler brownish; pterostigma brown; wings weakly infumate. 270 IAN D. GAULD VARIATION. One specimen from Panama has the gaster slightly darker than the other, but this individual is extremely dirty, and its true colour may not be apparent. The Costa Rican specimen has the head and much of the alitrunk quite bright yellowish, weak scutellar carinae and the lateral carina of the propodeum is more or less complete. REMARKS. This species is named in honour of Alejandro Eras for his tireless efforts to protect Guanacaste National Park, Costa Rica. Enicospilus erasi can most easily be recognized by the twisted, ridged mandible (Fig. 217), the large value of ICI and the characteristic alar sclerites (Fig. 323). E. erasi may be confused with E. galilea and E. hubbelli which it superficially resembles. Both E. galilea and E. erasi have the anterior margin of vein Rs+2r almost straight (Figs 323, 324), not weakly centrally bowed like that of E. hubbelli (Fig. 325), and E. hubbelli has the posterior area of the propodeum more regularly concentrically striate. The differences between E. galilea and E. erasi are, however, more subtle. The former has weakly twisted, stout mandibles and lacks propodeal crests, whilst E. erasi has quite strongly tapered mandibles twisted 55° or more, and distinct lateral crests present on the propodeum. BIOLOGICAL INFORMATION. Enicospilus erasi is only known to occur in Costa Rica and Panama where two specimens have been collected at black light, at 1050 m in Fortuna (8°12’N, 82°13’W), northern Panama during February and April, and a third taken in rainforest at 800 m in Costa Rica. Nothing is known about the natural history of this species. MATERIAL EXAMINED Holotype 0’, Costa Rica: Alajuela Prov., Virgen de Socorro, 800 m, ii.1987 (Janzen & Hallwachs) (BMNH). Paratypes. Panama: 1 9, Chiriqui, Fortuna, 1050 m, ii.1978 (Wolda) (RNH); 1 2, same locality, iv.1978 (Wolda) (RNH). Enicospilus galilea sp. n. (Fig. 324) DEscrIPTION. Mandibles stout, moderately long, weakly and evenly narrowed, apically twisted 25—35°; upper mandibular tooth quite slender, subcylindrical, 1.2—1.4 times as long as the compressed and slightly broader lower tooth; outer mandibular surface bearing fine scattered hairs, more or less flat centrally, but with a distinct shallow proximoventral concavity. Labrum 0.40.5 times as long as broad; malar space 0.2— 0.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.4-1.6 times as broad as long, with margin subtruncate apically. Lower face 0.69-0.74 times as broad as long, centrally finely and closely punctate. Head in dorsal view with genae rounded behind eyes; posterior ocellus contiguous with eye; FI = 67-73%; occipital carina mediodorsally complete or narrowly interrupted, ventrally curved to join hypostomal carina about 0.7—0.8 times the basal mandibular width away from mandible. Antenna long and slender, with 64-66 flagellar segments; 20th segment 2.0—2.1 times as long as broad. Mesoscutum polished, virtually impunctate, in profile evenly rounded; notauli weak but discernible anteriorly. Mesopleuron polished, the upper part punctate, the lower part punctate with transverse rugae or striae present; epicnemial carina strongly curved towards, but generally not reaching, anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.3—-1.4 times as long as anteriorly broad, anteriorly punctate, somewhat rugose posteriorly. Metapleuron weakly convex, but posterodorsally raised, this raised part being transversely rugose to striate, the flatter anteroventral part being punctate; submetapleural carina weakly broadened anteriorly; posterior transverse carina of mesosternum complete. Propodeum in profile abruptly declivous; anterior transverse carina centrally complete, its lateral extremities obsolescent, posterior transverse carina absent; anterior area short, steep, striate; spiracular area quite long, more or less smooth; posterior area coarsely irregularly rugose-reticulate; lateral longitudinal carina usually present anteriorly as a vestige, its extreme anterior end usually joined to spiracular margin by a weak carina. Fore wing length 19-21 mm; discosubmarginal cell as in Fig. 324); AI = 0.77-1.32; CI = 0.35-0.48; ICI = 0.54-0.68; SDI = 1.31-1.42; cu-a proximal to the base of Rs&M by 0.1—0.3 times its own length; marginal cell proximally distinctly more sparsely hirsute than it is centrally; 1st subdiscal cell anteriorly and distally broadly hirsute. Hind wing with 7-10 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa weakly bowed. Fore leg with tibia slightly flattened, with scattered inconspicuous spines on outer surface. Mid leg with longer tibial spur 1.3—1.6 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of hind tarsus 2.4—2.5 times as long as OPHIONINAE OF TROPICAL MESOAMERICA 27 broad; claws of female moderately long, apically abruptly rounded, with fine close pectinae; those of male similar. Gaster long and slender; tergite 2 in profile 6.0 or more times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 3.54.0 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing very dense long stout pubescence; gonosquama apically rounded. Colour generally with head and alitrunk yellowish, the mesoscutum bearing three longitudinal black vittae; legs, propodeum dorsally and gaster darker brownish yellow; interocellar area yellow or slightly infuscate; antenna proximally black, distally becoming blackish brown or even brown; pterostigma golden; wings more or less hyaline. VARIATION. Most specimens have the alar sclerites similar to those illustrated, but in some individuals the distal sclerite is evanescent distally, or even occasionally totally indistinct. REMARKS. This species is named after the Hotel Galilea, San José, Costa Rica, provider of first-rate hospitality for tropical biologists over the years. Enicospilus galilea is similar in general appearance to and likely to be confused with E. erasi (p. 269). BIOLOGICAL INFORMATION. Enicospilus galilea is a Mesoamerican species that is known to occur from Belize south to Central Panama. It has been collected from sea-level up to about 1300 m. Nothing is known about the natural history of this species. MATERIAL EXAMINED Holotype 9, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, 1.1984 (Janzen & Hallwachs) (BMNH). Paratypes. Belize: 1 2, Middlesex, 125 m, iii.1965 (Welling) (CNC). Costa Rica: Cartago Prov.: 1 9, Turrialba, 700 m, vii.1965 (Real) (CAS): Guanacaste Prov.: 1 2, Estacion Mengo on SW. side of Volcan Cacao, 1100 m, vii.1987 (Janzen & Hallwachs) (BMNH): Puntarenas Prov.: 2 2, Monteverde, 1300 m, ix.1985, vi.1986 (Haber) (BMNH). Panama: 1 ©’, Barro Colorado Island, 120 m, viii.1983 (Wolda) (BMNH). Enicospilus hubbelli sp. n. (Figs 207, 208, 325) Description. Mandibles moderately stout, quite long, distally moderately strongly and evenly narrowed, apically twisted 35—45° (Fig. 208); upper mandibular tooth quite slender, subcylindrical, 1.2—1.4 times as long as the compressed and very slightly broader lower tooth; outer mandibular surface bearing long fine scattered hairs, more or less flat centrally, but with a distinct shallow and rather finely pubescent proximoventral concavity. Labrum 0.3—0.4 times as long as broad; malar space 0.20.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.4-1.6 times as broad as long, with margin subtruncate apically. Lower face 0.70—-0.76 times as broad as long, centrally finely and closely punctate. Head in dorsal view with genae rounded behind eyes; posterior ocellus contiguous with eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.7—0.8 times the basal mandibular width away from mandible. Antenna long and slender, with 59-64 flagellar segments; 20th segment 1.8—2.0 times as long as broad. Mesoscutum polished, virtually impunctate, in profile evenly rounded; notauli weak but discernible anteriorly. Mesopleuron polished, the upper part punctate to punctostriate, the lower part punctate with transverse rugae or striae present; epicnemial carina strongly curved towards, but generally not reaching, anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.3—1.4 times as long as anteriorly broad, anteriorly punctate, somewhat rugose posteriorly. Metapleuron quite strongly convex, punctate anteroventrally, posterodorsally with isolated irregular rugae; submetapleural carina weakly broadened anteriorly; posterior transverse carina of mesosternum complete. Propodeum in profile abruptly declivous; anterior transverse carina sinuous, centrally complete, though often weak, its lateral extremities always obsolescent, posterior transverse carina absent; anterior area moderately long, steep, striate; spiracular area short, finely and closely punctate; posterior area more or less concentrically rugose-striate, the striae tending to become longitudi- nal on midline (Fig. 207); lateral longitudinal carina fine but present, usually joined to spiracular margin by a short carina. Fore wing length 19-21 mm; discosubmarginal cell as in Fig. 325; AI = 0.62-0.78; CI = 0.36-0.41; ICI = 0.78-0.83; SDI = 1.25-1.46; cu-a proximal to the base of Rs&M by 0.10.2 times its own length; marginal cell proximally from evenly hirsute to narrowly glabrous; 1st subdiscal cell anteriorly and distally broadly hirsute. Hind wing with 9-10 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa weakly bowed. D2, IAN D. GAULD Fore leg with tibia slightly flattened, with scattered slender spines on outer surface. Mid leg with longer tibial spur 1.4-1.6 times length of the shorter. Hind leg with coxa in profile 1.8-1.9 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.2—2.4 times as long as broad; claws of female moderately long, apically abruptly rounded, with fine close pectinae. Gaster from rather stout to long and slender; tergite 2 in profile 4. 1—6.8 times as long as posteriorly deep, laterotergite folded under, thyridia oval and separated from anterior margin of tergite by about 4.5-4.9 times its own length. Ovipositor slender, its sheath narrow. Male unknown. Colour generally with head and alitrunk yellowish, the mesoscutum bearing three longitudinal black vittae; legs, propodeum dorsally and gaster darker brownish yellow; interocellar area yellow or slightly infuscate; antenna almost entirely black, extreme distal apex pale brown; pterostigma golden; wings more or less hyaline. VARIATION. None remarkable. REMARKS. This species is named in honour of Steve Hubbell in recognition of his unflagging interest in tropical tree biology. Enicospilus hubbelli is similar to E. galilea and E. erasi (see the remarks under the last species). BIOLOGICAL INFORMATION. Enicospilus hubbelli is only known to occur in Panama where it has been collected in lowland rainforest on Barro Colorado Island. Nothing is known about its biology. MATERIAL EXAMINED Holotype 9, Panama: Barro Colorado Island, 120 m, vii.1985 (Wolda) (BMNH). Paratypes. Panama: 3 9, Barro Colorado Island, 120 m, vi & x.1984, vii.1985 (Wolda) (BMNH). Enicospilus flavoscutellatus (Brullé) (Figs 206, 215, 326) Ophion flavo-scutellatus Brullé, 1846: 140. Holotype 0’, BRAZIL (MNHN) [examined]. Ophion thoracicus Cresson, 1865: 55. LECTOTYPE 9, CUBA, (PANS) here designated [examined]. [Synonymized by Townes & Townes, 1966: 177.] Ophion trimaculatus Taschenberg, 1875: 433. Holotype 2, BRAZIL (Halle). [Synonymized by Townes & Townes, 1966: 177.] Ophion (Enicospilus) flavo-scutellatus Brullé; Cameron, 1886: 291. [Ophion (Enicospilus) concolor Cresson; Cameron, 1886: 291. Misidentification. ] Enicospilus thoracicus (Cresson) Ashmead, 1900b: 271. Henicospilus flavoscutellatus (Brullé) Dalla Torre, 1901: 181. Henicospilus thoracicus (Cresson) Dalla Torre, 1901: 184. Henicospilus trispilus Szépligeti, 1906: 145. LECTOTYPE Oo’, VENEZUELA, (TM) here designated [examined]. Syn. n. Enicospilus flavo-scutellatus (Brullé) Hooker, 1912: 68. Enicospilus trispilus (Szépligeti) Hooker, 1912: 90. Henicospilus trimaculatus (Taschenberg) Morley, 1912: 35. Henicospilus attritus Enderlein, 1921: 33. Holotype 0’, PERU (IZPAN) [examined]. Syn. n. Enicospilus attritus (Enderlein) Townes & Townes, 1966: 175. Enicospilus flavoscutellatus (Brullé); Townes & Townes, 1966: 177. DescriPTion. Mandibles long, proximally moderately narrowed, distally broad and more or less parallel- sided, apically twisted 15—25°; upper mandibular tooth compressed, 1.2—1.6 times as long as the lower tooth; outer mandibular surface bearing long fine sparse hairs, distally flat to slightly concave and proximally with a weak ventrobasal concavity. Labrum 0.3—0.4 times as long as broad; malar space 0.3—0.4 times as long as basal mandibular width. Clypeus in profile moderately to quite strongly convex, margin generally subacute; clypeus in front view 1.3—1.5 times as broad as long, with margin subtruncate or weakly convex apically. Lower face 0.73—0.78 times as broad as long, polished, punctate. Head in dorsal view with genae constricted behind the eyes; posterior ocellus close to eye; FI = 65-70%; occipital carina mediodor- sally complete, ventrally curved to join hypostomal carina about 0.60.8 times the basal mandibular width away from mandible. Antenna long and slender, with 60-65 flagellar segments; 20th segment 2. 1—2.4 times as long as broad. Mesoscutum polished, almost impunctate, in profile evenly rounded (Fig. 206); notauli vestigial. Mesopleuron highly polished, the upper part very finely punctate, the lower part similar but with occasional wrinkles ventrally; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.7 or more of its length (Fig. 215); scutellum in dorsal view OPHIONINAE OF TROPICAL MESOAMERICA 273 1.4-1.6 times as long as anteriorly broad, polished and punctate, occasionally wrinkled posteriorly. Metapleuron moderately convex, highly polished and punctate finely; submetapleural carina evenly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina usually absent, rarely represented later- ally; anterior area long, striate; spiracular area quite short, smooth; posterior area generally rather weakly and irregularly rugose, often with the rugosities very weak so the surface is nearly smooth, very rarely almost reticulate, but in each case a more or less distinct median longitudinal wrinkle is usually discernible, and occasionally this may be particularly strong, resembling a carina; lateral longitudinal carina complete, joined to spiracular margin by a short carina. Fore wing length 15-20 mm; discosubmarginal cell as in Fig. 326; AI = 0.71-0.77; CI = 0.27-0.39; ICI = 0.44-0.54; SDI = 1.06-1.18; cu-a proximal to the base of Rs&M by 0.1—0.3 times its own length; marginal cell proximally more or less evenly hirsute; 1st subdiscal cell with distal margin broadly hirsute. Hind wing with 5—7 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa more or less straight. Fore leg with tibia slightly flattened, with slender, scattered spines on outer surface. Mid leg with longer tibial spur 1.4~1.5 times length of the shorter. Hind leg with coxa in profile 1.8-1.9 times as long as deep; trochantellus dorsally 0.1—0.2 times as long as broad; 4th segment of tarsus 2.2—2.5 times as long as broad; claws of female moderately long, distally abruptly curved, bearing fine, close pectinae; claws of male similar but with pectinae slightly finer and shorter. Gaster long and slender; tergite 2 in profile more than 6.5 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 2-4 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing fine decumbent pubescence, but with pubescence denser and more erect on posteromedian margin of sternites 7-8; gonosquama apically rounded. Colour rather variable but most generally quite brightish yellow with mesoscutum bearing three dark longitudinal vittae; gaster usually with upper and lower margins of segments 3-5 slightly infuscate and tergite 6+ more uniformly weakly infuscate; interocellar area yellow; antenna golden; pterostigma brownish yellow; wings hyaline. VARIATION. Enicospilus flavoscutellatus exhibits a considerable range of colour variation. Many individuals from lowland ‘disturbed’ sites are more or less entirely yellowish with the mesoscutal vittae slightly darker brown. Individuals from higher altitude sites (1200+ m) tend to be more extensively dark-marked than the norm, having additionally dark areas on the posterodorsal corner of the mesopleuron, mesosternum, metanotum and posterior area of the propodeum. Quite often specimens have the central part of tergites 3-5 pale and the periphery dark so the gaster appears banded or mottled in side view. Occasional specimens are generally darker brownish. One series of individuals collected at 2200 m at Guadelupe Arriba, Panama, are very distinctive in being smooth and extremely highly polished, and in having distinct lateral parts of the posterior transverse carina of the propodeum, the lower corner of the epicnemial carina acute, the petiole black and whitish marks on the scutellum, mesopleuron, face and gena. These individuals also have the wings weakly infumate and the flagellum distally infuscate, but with the extreme apex pale. These individuals closely resemble the holotype of E. attritus, and I consider them all to be conspecific with typical examples of E. flavoscutellatus. REMARKS. Townes & Townes (1966: 178) state that in 1916 Cresson designated a lectotype for Ophion thoracicus. However, Cresson’s collective designation of numerous lectotypes in this paper is invalid under the Code (Article 74c). Cresson stated that the [lecto]type was specimen ‘77 Cuba’. In the collection of the Philadelphia Academy I could find only a single specimen (number 77.1) labelled as ‘type’ and this is the specimen I here designate as lectotype. The form of the alar sclerites, in particular the elongate central sclerite, make E. flavoscutellatus one of the most distinctive and easily recognizable Neotropical species (Fig. 326). The structure of the mandibles and the rather polished appearance are also distinctive but require experience to appreciate. Otherwise, in structure, this is a rather unexceptional species. Morphologically, it most closely resembles E. devriesi (see that species) and an undescribed Argentinian/Southern Brazilian species which differs only in having a small circular central sclerite. BIOLOGICAL INFORMATION. Enicospilus flavoscutellatus is a widespread Neotropical species (Map 30) whose range extends from about 18°N in southern Mexico south throughout Central and South America to southern Brazil and northern Argentina (ca 28°S) where it is apparently gradually replaced by a closely related species. E. flavoscutellatus has been taken in a wide variety of habitat types from about 300 m up to about 2300 m in Central America, and up to 3000 m in the Andes. At such high altitudes it is rather scarce, and it comprises a much smaller proportion of the total light-trap catch than does its close relative E. 274 IAN D. GAULD Map 30 Localities at which Enicospilus flavoscutellatus has been collected. devriesi. These two species are sympatric between 1200 and 2300 m, but at these lower elevation sites E. flavoscutellatus is much more frequently collected than E. devriesi. E. flavoscutellatus is one of the commonest nocturnal ichneumonids in lower montane forests in Central America. At many sites individuals are extremely abundant, often outnumbering all other species com- bined and literally hundreds of specimens have been collected at Monteverde, between January and April. It has been collected at almost every mid-altitude site that has been sampled, though it is absent from high altitude sites (2500+ m) in Costa Rica. In Costa Rica this species has been collected at the following sites: Finca Campana at 5 km NW. Dos Rios, Finca San Gabriel at 3 km W. of Dos Rios and Virgen de Socorro in Alajuela Province; 16 km S. San Isidro de Tejar, Tapanti, Turrialba and Volcan Irazi in Cartago Province; Cerro el Hacha, Rincon de la Vieja National Park, 4km W. Santa Cecilia, Santa Rosa National Park, 5 km OPHIONINAE OF TROPICAL MESOAMERICA 275 NE. Quebrada Grande, Estacion Mengo [= Finca La Luz] on the upper slopes of Volcan Cacao and Casa Maritza on the lower slopes of Volcan Orosi in Guanacaste Province; Monteverde area in Puntarenas Province; Braulio Carrillo National Park, San Antonio de Escazt and San Isidro in San José Province. Despite intensive collecting E. flavoscutellatus has never been collected on Barro Colorado Island in lowland wet forest. Neither has it been found to be present in similar habitats in Costa Rica (such as Tortuguero or Corcovado National Parks) though these last two sites are only poorly collected. In lower montane humid forest sites such as Monteverde (1350 m) and Braulio Carrillo National Park (800 m) E. flavoscutellatus is present throughout the year, though in both sites it seems to be most abundant between January and April. It appears to be rather less common in Santa Rosa National Park where it has a different phenology. At this site virtually all specimens were collected from late May to July or from November to early January. These distributional peaks, which correspond to the beginning and end of the wet season, suggest this species may have two generations in deciduous forest. I have seen no specimens from Santa Rosa collected in February-April and only one in August to October. I have dissected 10 females from Santa Rosa National Park and Monteverde. One had no eggs at all in either lateral oviduct; the remainder had between 9 and 17 eggs present (mean = 12.75) in each of the two lateral oviducts and the largest number of eggs found altogether in any one individual was 32. All eggs seemed to be mature and no small developing eggs were observed. Despite the fact that this is an exceptionally common species at many sites, there is not a single host record available. MATERIAL EXAMINED Lectotype oO (Enicospilus trispilus Szépligeti), Venezuela: Merida (TM); paralectotypes: Bolivia: 1 9, Mapiri (TM); Mexico: 1 9, ‘Verebelyi’ (TM). Holotype 2 (Ophion flavo-scutellatus Brullé), Brazil: Rio Grande (MNHN). Lectotype 2 (Ophion thoracicus Cresson), Cuba ‘77.1’ (PANS). Holotype (Henicospilus attritus Enderlein), Peru: Chanchamayo (IZPAN). 583 o’, 649 9, from the following localities: Argentina (Buenos Aires, Misiones, Tucuman); Barbados; Bolivia (Chapare, Cochabamba, Ichilo, La Paz); Brazil (Bahia, Guanabara, Santa Catarina); Colombia (Antioquia, Cundinamarca, Huila, Magdalena, Valle); Costa Rica (Alajuela, Cartago, Guanacaste, Puntarenas, San José); Cuba; Dominica; Dominican Republic; Ecuador (Cotopaxi, Esmeraldas, Imbabura, Loja, Napo, Pichincha, Tungurahua, Zamora); Grenada; Guadeloupe; Guatemala; Guyana; Jamaica; Martinique; Mexico (Chiapas, Guerrero, Oaxaca, Tabasco); Panama; Peru (Cuzco, Huanuco, Lima, Loreto); St Kitts; St Lucia; Surinam; Venezuela (Caracas, Lara, Merida, Yaracuy). (BMNH, CNC, FSCA, PANS, TC, UMC, USNM.) Enicospilus woldai sp. n. (Fig. 327) Description. Mandibles short, proximally strongly narrowed, with a small basal lobe, distally more weakly narrowed, apically twisted 80°; upper mandibular tooth slightly depressed, 1.4-1.5 times as long as the lower tooth; outer mandibular surface centrally convex, very sparsely pubescent, proximally with a narrow crescentic concavity. Labrum 0.3 times as long as broad; malar space 0.3—-0.4 times as long as basal mandibular width. Clypeus in profile flat, margin sharp; clypeus in front view 1.3—1.4 times as broad as long, with margin truncate. Lower face 0.72-0.77 times as broad as long, centrally sparsely punctate. Head in dorsal view with genae rounded behind eyes; posterior ocellus close to eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6-0.8 times the basal man- dibular width away from mandible. Antenna slender, with 55—57 flagellar segments; 20th segment 2.0-2.2 times as long as broad. Mesoscutum polished, finely and closely punctate, in profile abruptly rounded; notauli vestigial. Mesopleuron highly polished, the upper and lower parts punctate; epicnemial carina inclined towards anterior margin of pleuron, its upper end abruptly turned forward and evanescent. Scutellum in profile weakly convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, anteriorly smooth and punctate, posteriorly longitudinally wrinkled. Metapleuron weakly convex, very finely coriaceous with weak punctures; submetapleural carina weakly broadened anteriorly; posterior transverse carina of mesosternum complete. Propodeum in profile evenly rounded; anterior transverse carina complete, posterior transverse carina absent; anterior area long, shallow, with irregular rugae; spiracular area quite long, smooth; posterior area finely wrinkled/rugulose, usually with a strong median longitudinal wrinkle; lateral longitudinal carina complete anteriorly, sometimes entire, not joined to spiracular margin by a short carina. Fore wing length 15-16 mm; discosubmarginal cell as in Fig. 327; AI = 0.57-0.80; CI = 0.34-0.38; ICI = 0.42-0.47; SDI = 1.10—1.19; cu-a from subopposite to the base of Rs&M to proximal to it by 0.2 times 276 IAN D. GAULD its own length; marginal cell proximally uniformly hirsute; 1st subdiscal cell with anterior 0.4 very sparsely hirsute. Hind wing with 6-7 hamuli on R1; 1st abscissa of Rs almost straight, 2nd abscissa straight. Fore leg with tibia slightly flattened, with fine, scattered spines on outer surface. Mid leg with longer tibial spur 1.3—1.5 times length of the shorter. Hind leg with coxa in profile 1.9—2.0 times as long as deep; trochantellus dorsally 0.5 times as long as broad; 4th segment of tarsus 2.3—2.4 times as long as broad; claws of female long, abruptly curved with short stout pectinae, those of male similar. Gaster slender; tergite 2 in profile 4.9-5.5 times as long as posteriorly deep, laterotergite folded under, thyridia elongately oval and separated from anterior margin of tergite by about 4—S times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 densely pubescent, bearing many long, erect hairs; subgenital plate unusual in having a small angular projection in the middle of the posterior margin; gonosquama large, distally rounded; aedeagus unusual in having a broad membranous collar proximal to apex. * Colon generally pale yellow; interocellar area yellow; antenna yellowish; pterostigma golden yellow; wings hyaline. VARIATION. This is a morphologically very uniform species. REMARKS. This species is named in honour of Dr Henk Wolda as a gesture of thanks for collecting virtually all the the ophionines known from Panama. Enicospilus woldai is one of the most distinctive Central American species of Enicospilus. The large fenestra and trapezoidal proximal sclerite (Fig. 327) are quite unlike those of any other species. The highly modified male genitalia are similar to those of some species in the trilineatus species-group, but E. woldai does not have a pendant second epipleuron or the stout mandibles characteristic of taxa in this species- group. BIOLOGICAL INFORMATION. Enicospilus woldai is only known to occur in lowland rainforest on Barro Colorado Island, Panama and in Ecuador. In Panama isolated individuals have been collected at light between the end of March and early August. The hosts of this species are unknown. MATERIAL EXAMINED Holotype 9, Panama: Barro Colorado Island, 120 m, iv.1983 (Wolda) (BMNH) Paratypes. Ecuador: 1 9, Pichincha Prov., Tinlandia, 12 km E. Santo Domingo de Los Colorados, 800 m, v.1986 (Eger) (BMNH). Panama: 1 9, Barro Colorado Island, Gatun Lake, iii.1979 (Wolda) (TC);1 9, same locality, v.1982 (Wolda) (TC); Barro Colorado Island, 120 m, 1 9, v.1985; 1 CO’, vii.1984; 1 Co’, viii. 1984; 2 9, v.1985 (Wolda) (BMNH). Enicospilus liesneri sp. n. (Figs 128, 209, 328) Description. Mandibles short, proximally strongly narrowed with a small proximoventral lobe, distally more evenly tapered, apically twisted 75—80°; upper mandibular tooth slightly depressed, 1.4-1.7 times as long as the lower tooth which is slightly compressed and unevenly tapered, with distal apex unusually slender; outer mandibular surface centrally flat, with scattered pubescence, proximally with a strong narrow concavity. Labrum 0.3—0.4 times as long as broad; malar space 0.10.2 times as long as basal mandibular width. Clypeus in profile flat, margin blunt to subacute; clypeus in front view 1.4-1.5 times as broad as long, with margin truncate or evenly slightly concave. Lower face 0.65—0.70 times as broad as long, convex, centrally punctate. Head in dorsal view with genae very short and strongly constricted (Fig. 128); posterior ocellus close to eye; FI = 65-70%; occiput unusually strongly impressed so that head is centrally very short, and abruptly declivous behind ocelli; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6—0.8 times the basal mandibular width away from mandible. Antenna long, but rather stout, with 47-50 flagellar segments; 20th segment 1.6-1.9 times as long as broad. Mesoscutum polished, finely punctate, in profile abruptly rounded; notauli vestigial. Mesopleuron polished, the upper part sparsely finely punctate, the lower part slightly more closely punctate; epicnemial carina inclined towards anterior margin of pleuron, its upper end evanescent. Scutellum in profile weakly convex, laterally carinate for almost all of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, anteriorly smooth, posteriorly with isolated striae. Metapleuron flat except for a convex ridge dorsally, punctate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum broadly incomplete centrally. Propodeum in profile evenly declivous; anterior transverse carina generally complete, though often weak or evanescent laterally, posterior transverse carina absent; anterior area quite long, coriaceous with scattered longitudinal striae; spiracular area long, smooth; OPHIONINAE OF TROPICAL MESOAMERICA 277 posterior area weakly sculptured (Fig. 209), mainly coriaceous, but sometimes with a weak median longitudinal ridge; lateral longitudinal carina often complete, though sometimes weak posteriorly, indis- tinctly or not joined to spiracular margin by a short carina. Fore wing length 11-14 mm; discosubmarginal cell as in Fig. 328; AI = 1.00-1.14; CI = 0.18-0.24; ICI = 0.45-0.54; SDI = 1.16—1.27; cu-a from subopposite to the base of Rs&M to proximal to it by 0.1 times its own length; marginal cell proximally broadly glabrous; 1st subdiscal cell with scattered pubescence over distal surface. Hind wing with 6-7 hamuli on R1; Ist abscissa of Rs straight, 2nd abscissa straight. Fore leg with tibia weakly flattened, with isolated weak spines on outer surface. Mid leg with longer tibial spur 1.5—1.6 times length of the shorter. Hind leg with coxa in profile 1.7—-1.8 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.0—2.3 times as long as broad; claws of female rather small with fine close pectinae, those of male similar but with pectinae slightly shorter. Gaster moderately slender; tergite 2 in profile 4.5—5.0 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 2.5—3.0 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-8 bearing lateromedian rows of long hairs which are sparse anteriorly and terminate in a large tuft posteriorly, tergite 9 with fine decumbent pubescence; gonosquama distally subacute. Colour generally pale yellow, with three mesoscutal vittae and 5+ blackish; interocellar area yellow; antenna golden; pterostigma yellow; wings hyaline. VARIATION. Enicospilus liesneri is a morphologically very uniform species. Some of the smaller individuals may have the mesoscutal vittae medium brown, or even paler, and a few have the posterior tergites of the gaster only very weakly infuscate. REMARKS. This species is named in honour of Ron Liesner for his helpfulness in identifying Neotropical plants for biologists. Enicospilus liesneri may easily be recognized by the shape of the head, which in dorsal view is extremely short centrally, and abruptly declivous behind the ocelli (Fig. 209). The antennae are much stouter than other small species with strongly twisted mandibles, and the arrangement of the pubescence on the male terminal sternites is quite characteristic. The characteristically modified form of the head of this species is also shared by E. lovejoyi which differs from E. liesneri most obviously in having a black interocellar area. The two are almost certainly sister-species, but their relationship to other taxa remains unclear. Possibly they are related to other species with a complete distal sclerite. BIOLOGICAL INFORMATION. Enicospilus liesneri is a widely distributed species whose range extends from Veracruz in southern Mexico (ca 20°N) south to about 28°S in southern Brazil. It also has been recorded in Cuba, but it is not known to occur on any other Caribbean island. E. liesneri has been collected most frequently in the seasonally dry lowland forests of Guanacaste, Costa Rica, but it has been taken at altitudes up to 1350 m (e.g. at Monteverde), where it seems to be relatively rare. In the seasonally dry forest in Santa Rosa National Park E. liesneri is one of the most frequently collected Enicospilus species at the beginning of the wet season. The cumulative light-trap data for the years 1980-87 are: Vive ere Mie AM Jad Ass ON, iD Bed Stel Die ones oIOB, Filey a= 1 - 4 It has also beeen collected on San José Island, one of the small Murciélagos Islands off the coast of the Santa Elena Peninsula, in north-western Costa Rica. E. liesneri is rather uncommon at light in wetter lowland forests such as on Barro Colorado Island, but even at this site it still seems to have a similar seasonal distribution. The cumulative data for Barro Colorado Island are: Seite AGM Tee AS. ©. N. 2D 2 ON a Dr D. H. Janzen has reared E. liesneri on two occasions in Santa Rosa National Park from the larvae of a species of Eulepidotis (Noctuidae) found feeding on Luehea sp. (Tiliaceae) (rearing reference numbers 84-SRNP-473; 84-SRNP-547). The two parasitized caterpillars were collected on the 12th and 14th of June; they spun cocoons about four days later and the adult ichneumonids eclosed on the 1st and 3rd of July, suggesting this species may have more than one generation per year. 278 IAN D. GAULD MATERIAL EXAMINED Holotype 2, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, vi.1985 (Gauld) (BMNH),. Paratypes. Brazil: 2 2, Minas Gerais, Pedra Azul, 800 m, xi.1972 (Alvarenga & Seabra) (TC); 1 9, Minas Gerais, Aguas Vermelhas, 800 m, xii. 1983 (Alvarenga) (TC); 1 0’, Nova Teutonia, Santa Catarina, xi.1936 (Plaumann) (BMNH); 3 @, Para, Tucuruf, i.1979 (Alvarenga) (TC). Costa Rica: Guanacaste Prov.: 1 2, Highway, 2 km E. Cuajiniquil, vi.1986 (Gauld) (BMNH); 6 oO’, 4 9, Islas Murciélagos, San José Is, viii.1987 (Janzen & Hallwachs) (BMNH); 97 Oo’, 120 9, Santa Rosa National Park, 300 m, months as enumerated above, 1980-1986 (Janzen & Hallwachs/Gauld) (BMNH, CNC, MNCR, TC, USNM); 3 9, Volcan Cacao, Finca La Luz [= Estacion Mengo], 1100 m, viii. 1986 (Janzen & Hallwachs) (BMNH);8 @, same locality and collectors, vi-vii.1987 (BMNH); 3 2, Volcan Orosi, Casa Mariksa [= Maritza], 800 m, vii. 1986 (Gauld) (BMNH): Puntarenas Prov.: 1 0’, Monteverde, 1350 m, viii.1985 (Haber) (BMNH);1<0, (FSCA); 1 2, Cayuga, v.1915 (Schaus) (USNM). Honduras: 1 9, Lancetilla (Bates) (MCZ). Mexico: Chiapas: 1 CO’, 32 km N. Huixtla, 1000 m, vi.1969 (Mason) (CNC): Veracruz: 2 9, Lake Catemaco, Coyame, viii.1963 (Woodruff) (FSCA). Panama: 1 0’, 1 9, Barro Colorado Island, vi.1978 (Wolda) (RNH); Barro Colorado Island, Gatun Lake, 1 9, 11.1979; 5 Q, iv.1979; 1 CO’, i.1983; 1 CO’, iti.1983 (Wolda) (TC); 2 0’, 9 9, same locality, 120 m, i-vii.1983—4 (Wolda) (BMNH). Venezuela: 1 Q , Junquito, viii.1940 (TC). Enicospilus marini sp. n. (Fig. 329) DescriPtion. Mandibles short, proximally strongly narrowed, with a small proximoventral lobe, distally more evenly narrowed, apically twisted 75—80°; upper mandibular tooth slightly depressed, 1.41.7 times as long as the lower tooth which is compressed and apically very acute; outer mandibular surface with a strong proximal concavity, distally convex with scattered pubescence. Labrum 0.2-0.3 times as long as broad; malar space 0.1—0.2 times as long as basal mandibular width. Clypeus in profile from almost flat to very weakly convex, margin blunt; clypeus in front view 1.2—1.4 times as broad as long, the margin truncate apically. Lower face 0.66—0.70 times as broad as long, convex, centrally finely punctate. Head in dorsal view with genae constricted behind eyes; posterior ocellus close to eye; FI = 60-65%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6—0.8 times the basal man- dibular width away from mandible, but with lower end very weak. Antenna long and slender, with 50—52 flagellar segments; 20th segment 2.2—2.3 times as long as broad. Mesoscutum polished, sparsely punctate, in profile abruptly rounded with anterior margin out-turned; notauli vestigial. Mesopleuron polished, the upper part punctate to punctostriate, the lower part punc- tostriate to striate; epicnemial carina curved towards anterior margin of pleuron, with upper end evanes- cent. Scutellum in profile weakly convex, laterally carinate for almost all of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, smooth, with isolated rugae posteriorly. Metapleuron weakly convex, from punctate with scattered weak striae to diagonally striate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum centrally interrupted. Pro- podeum in profile abruptly declivous; anterior transverse carina complete, laterally sometimes weak, posterior transverse carina absent; anterior area short, with isolated rugae; spiracular area long, smooth; posterior area rugose, often with a median longitudinal ridge and with almost concentric transverse rugae, but in some specimens with this sculpture weakly developed; lateral longitudinal carina generally com- plete, not joined to spiracular margin by a distinct short carina. Fore wing length 10-11 mm; discosubmarginal cell as in Fig. 329; AI = 1.00—-1.68; CI = 0.19-0.27; ICI = 0.36-0.41; SDI = 1.14-1.51; cu-a opposite to the base of Rs&M; marginal cell proximally broadly glabrous; 1st subdiscal cell with scattered hairs. Hind wing with 5-6 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa almost straight. Fore leg with tibia weakly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.5—-1.7 times length of the shorter. Hind leg with coxa in profile 1.7—-1.9 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 3.0—3.1 times as long as broad; claws of female quite long, with short pectinae, those of male similar, but with pectinae finer. Gaster slender; tergite 2 in profile 5.5—6.1 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 3.54.0 times its own length. Ovipositor apically quite slender, its sheath narrow. Male with sternites 7-9 bearing numerous long, slightly curved, erect hairs; gonosquama apically rounded. OPHIONINAE OF TROPICAL MESOAMERICA 279 Colour generally pale yellowish brown, with head, pro- and mesothorax paler yellow; mesoscutum with three blackish longitudinal stripes; alitrunk laterally with a crescentic dark mark near the upper hind corner of the mesopleuron, and with an indistinct dark patch between the propodeum and the metapleuron; gaster with tergites 5+ infuscate; interocellar area yellow, blackish between the posterior ocelli; antenna yellowish brown; pterostigma yellowish brown; wings more or less hyaline. VARIATION. Some of the Brazilian specimens examined are paler yellowish with tergites 2+ of the gaster infuscate. This specimens often, but not always, have the meso- and metapleurae almost smooth, and highly polished. REMARKS. This species is named in honour of Sigifredo Marin for his great efforts to promote positive interactions between Costa Rican National Parks and their neighbours. Enicospilus marini is a small species that is most easily recognized by the form of the proximal sclerite (Fig. 329); a similar one is not found in any other Central American species. E. marini is one of a group of about six rather similar small species (herein called the marini species-complex) that are widely distributed throughout tropical America. Some have only been collected in Brazil, but three others E. oduberi, E. pescadori and E. sanchezi, occur in Central America. All these species are characterized by being small, having similar pleural and propodeal scuplture, and by having similar venation. All usually also have a crescent-shaped dark mark on the mesopleuron. BIOLOGICAL INFORMATION. Enicospilus marini is a widely distributed species whose range extends from El Salvador (ca 16°N), south to central Brazil (14°S). The majority of specimens are labelled as having been collected below 1000 m and none has been labelled as being from above this altitude (some have no altitude appended). In the seasonally dry forests of Santa Rosa National Park EF. marini has been collected intermittently between April and December with most at the end of the dry season (April/May). The pooled data for all captures at this site are: ie tne neuen aU AN ig SRD Pe re eT One A OR ee Fil eek a7 Despite intensive light-trapping in lowland rainforest on Barro Colorado Island E. marini has only been collected on two occasions, in January and June. The hosts of this species are unknown. MATERIAL EXAMINED Holotype ¢, Costa Rica: Guanacaste Prov., 300 m, iv.1983 (Janzen & Hallwachs) (BMNH). Paratypes. Brazil: 6 2, Bahia, Encruzilhada, 960 m, xi.1972 (Alvarenga) (TC); 4 9, same locality and collector, 980 m, xi.1973, xi.1974 (TC); 1 2, Minas Gerais, Aguas Vermelhas, 800 m, xii. 1983 (Alvarenga) (TC). Costa Rica: Guanacaste Prov.: 1 2, Santa Rosa National Park, 300 m, xii.1976 (Janzen) (TC); same locality and collector, 1 9, x.1977 (TC); 1 @, vii.1978 (BMNH); 1 9, same locality, v.1980 (Janzen & Hallwachs) (TC); same locality and collectors, 3 9, iv.1983 (BMNH); 1 9, iv.1984 (BMNH). El Salvador: 10’, Lake Coatapeque, viii.1972 (Hevel) (USNM). Panama: 2 9, Barro Colorado Island, 120 m, vi.1983, i.1984 (Wolda) (BMNH). Venezuela: 1 9, Aragua, Portachuelo Pass, nr Rancho Grande, iv.1960 (Test) (UMC). Enicospilus sanchezi sp. n. (Figs 125, 127, 330) Description. Mandibles quite short, distally evenly narrowed, apically twisted about 70°; upper mandibu- lar tooth strongly depressed, 1.2—1.3 times as long as the laterally compressed lower tooth; outer mandibu- lar surface sparsely pubescent, flat with a broad shallow proximal concavity. Labrum 0.2-0.3 times as long as broad; malar space 0.4 times as long as basal mandibular width. Clypeus in profile very weakly convex, margin flat; clypeus in front view 1.4~1.5 times as broad as long, the apical margin truncate. Lower face 0.70-0.75 times as broad as long, centrally polished, finely punctate. Head in dorsal view with genae constricted behind the eyes; posterior ocellus contiguous with the eye; FI = 60-65%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.7—0.8 times the basal man- dibular width away from mandible. Antenna long and slender, with 47—S2 flagellar segments; 20th segment 2.42.7 times as long as broad. Mesoscutum polished, sparsely punctate, in profile abruptly rounded; notauli absent. Mesopleuron polished, the upper part punctate, the lower part punctostriate; epicnemial carina strong, inclined towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.9 or more of its 280 IAN D. GAULD length; scutellum in dorsal view 1.5—-1.6 times as long as anteriorly broad, polished with scattered punctures, with a few longitudinal striae posteriorly. Metapleuron weakly convex, punctate to punctostri- ate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete, strong centrally. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area short, steep, bearing isolated striae; spiracular area long, smooth; posterior area with distinct irregular wrinkles; lateral longitudinal carina complete or evanescent posteriorly, joined to spiracular margin by a short carina. Fore wing length 7-11 mm; discosubmarginal cell as in Fig. 330; AI = 1.00—-1.50; CI = 0.25-0.43; ICI = 0.25—-0.37; SDI = 1.07-1.24; cu-a more or less opposite to the base of Rs&M; marginal cell proximally narrowly glabrous close to Rs+2r; 1st subdiscal cell with centre hirsute, this hirsute area reaching to distal end, the anterior and posterior margins very sparsely hirsute; unusual in having 2nd discal cell posteriorly glabrous, and with a glabrous arc in 2nd subdiscal cell. Hind wing with 5-6 hamuli on R1; 1st abscissa of Rs weakly but distinctly bowed centrally (Fig. 127), 2nd abscissa almost straight. Fore leg with tibia barely flattened, with isolate fine spines on outer surface. Mid leg with longer tibial spur 1.5-1.6 times length of the shorter. Hind leg with coxa in profile 1.6-1.7 times as long as deep; trochantellus dorsally about 0.3 times as long as broad; 4th segment of tarsus 2.6-2.8 times as long as broad; claws of female evenly curved, with close long stout pectinae, those of male similar. Gaster long and slender; tergite 2 in profile more than 7 times as long as posteriorly deep, laterotergite very narrow, membranous, pendant, thyridia elliptical and separated from anterior margin of tergite by about 3 times its own length. Ovipositor moderately stout, straight, its sheath narrow. Male with sternites 7-9 bearing long, rather sparse, slightly curved, erect hairs; gonosquama apically rounded. Colour generally pale yellowish with slightly darker mesoscutal vittae and with the posterior part of the mesoscutum infuscate; upper part of mesopleuron with a large dark mark posteriorly, that extends down pleural suture beyond level of episternal scrobe, and with its lower anterior end extending forward to or almost to epicnemial carina (Fig. 125); anterior concavity of metapleuron frequently with an infuscate mark; gaster yellowish, at the most with a general slight darkening posteriorly; interocellar area yellowish, or slightly infuscate between posterior ocelli, and often with frons below median ocellus infuscate; antenna yellowish, slightly infuscate distally; pterostigma yellow; wings hyaline. VARIATION. One female from Santa Rosa is slightly ‘gummy’ but appears to have the laterotergites turned under. In other characters it is a typical member of this species, but I have excluded it from the paratype series. The Venezuelan specimen has dark mesoscutal vittae. REmakkKS. This species is named in honour of Pablo Sanchez in recognition of his administrative efforts at the Museo Nacional de Costa Rica. Enicospilus sanchezi is very similar to E. pescadori but, with the exception of the single specimen referred to above, the characters given in the key work well for the material at hand. The pubescence of the 2nd discal and 2nd subdiscal cells of the fore wing is rather more uniform in E. pescadori than it is in E. sanchezi (cf. Figs 330 and 331). E. sanchezi belongs to the E. marini species-complex. BIOLOGICAL INFORMATION. Enicospilus sanchezi is a widely distributed tropical American species whose range extends from the extreme south of Mexico (about 15°N) south to Venezuela. Isolated specimens have been collected in Santa Rosa National Park between April and November. The seasonal distribution for cumulative catch in Santa Rosa (1977-86) is: J oko. MigeAs Melia iterAaSieO.0oNa oD ee i eee Nothing is known about the biology of this species. MATERIAL EXAMINED Holotype 2, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, v.1986 (Gauld) (BMNH). Paratypes. Costa Rica: Guanacaste Prov.: 4 9, Santa Rosa National Park, 300 m, vii-viii, xi.1977 (Janzen) (TC); 1 9, same locality, iv.1983 (Janzen & Hallwachs) (BMNH); 3 @, same locality, vi.1985, v & vii.1986 (Gauld) (BMNH); 1 ©’, Volcan Orosi, Casa Mariksa [= Maritza], 550 m, i.1986 (Janzen & Hallwachs) (BMNH). Mexico: Chiapas: 1 0’, 32 kmN. Huixtla, 1000 m, vi.1969 (Mason) (CNC);10',3 9, OPHIONINAE OF TROPICAL MESOAMERICA 281 Muste, nr Huixtla, 440 m, ix.1970 (Welling) (CNC). Panama: 1 9, Canal Zone, Margarita, v.1960 (Breeland) (TC). Venezuela: 1 2, Rancho Grande, ii.1970 (Howden) (TC). Non-paratypic material. Costa Rica: 1 9 , Guanacaste Prov., Santa Rosa National Park, vi.1980 (Janzen & Hallwachs) (BMNH). Enicospilus pescadori sp. n. (Figs 124, 126, 331) DescriPTion. Mandibles quite short, distally evenly narrowed, apically twisted about 70°; upper mandibu- lar tooth strongly depressed, 1.3—1.5 times as long as the laterally compressed lower tooth; outer mandibu- lar surface sparsely pubescent, flat with a broad shallow proximal concavity. Labrum 0.2-0.3 times as long as broad; malar space 0.4 times as long as basal mandibular width. Clypeus in profile very weakly convex, margin flat; clypeus in front view 1.4—1.5 times as broad as long, the apical margin truncate. Lower face 0.65—0.70 times as broad as long, centrally polished, finely punctate. Head in dorsal view with genae constricted behind the eyes; posterior ocellus contiguous with the eye; FI = 60-65%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6-0.8 times the basal man- dibular width away from mandible. Antenna long and slender, with 48-52 flagellar segments; 20th segment 2.0-2.4 times as long as broad. Mesoscutum polished, sparsely punctate, in profile abruptly rounded; notauli absent. Mesopleuron polished, the upper part punctate, the lower part punctostriate; epicnemial carina strong, inclined towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.9 or more of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, polished with scattered punctures, with a few longitudinal striae posteriorly. Metapleuron weakly convex, punctate to punctostri- ate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete, strong centrally. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area short, steep, bearing isolated striae; spiracular area long, smooth; posterior area with distinct irregular wrinkles; lateral longitudinal carina complete or evanescent posteriorly, joined to spiracular margin by a short carina. Fore wing length 10-12 mm; discosubmarginal cell as in Fig. 331; AI = 0.95-1.25; CI = 0.16-0.21; ICI = 0.30-0.35; SDI = 1.05—1.27; cu-a more or less opposite to the base of Rs& M; marginal cell proximally narrowly glabrous close to Rs+2r; 1st subdiscal cell with distal part sparsely hirsute. Hind wing with 5-6 hamuli on R1; 1st abscissa of Rs straight (Fig. 126), 2nd abscissa more or less straight. Fore leg with tibia subcylindrical; with isolated spines on outer surface. Mid leg with longer tibial spur 1.5—1.6 times length of the shorter. Hind leg with coxa in profile 1.6—1.7 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.2-2.4 times as long as broad; claws of female moderately long, evenly curved, with close fine pectinae; claws of male shorter and more abruptly rounded, with slender pectinae. Gaster long and slender; tergite 2 in profile more than 6 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 3 times its own length. Ovipositor slender, straight, its sheath narrow. Male with sternites 7-9 bearing close erect pubescence; gonosquama apically rounded. Colour generally bright yellowish, with three longitudinal dark vittae on the mesoscutum; mesopleuron bearing a small infuscate mark that extends from upper hind corner to near episternal scrobe, but does not extend down margin of pleural suture (Fig. 124); gaster yellow with posterior part of tergite 5 and the following tergites infuscate; interocellar area yellow; antenna golden; pterostigma yellowish; wings hyaline. VARIATION. The specimen from Brazil has the meso- and metapleurae smoother than those of other specimens. REMARKS. This species is named in honour of Alfonso Pescador for his studies of the moth fauna of the Chamela Biological Station, Mexico. Enicospilus pescadori is morphologically very similar to E. sanchezi, but the two species differ consist- ently in a number of quite subtle features. Most obviously they differ in colour: E. pescadori has a much less extensively dark-marked mesopleuron than E. sanchezi, never has dark metapleural marks and has the posterior half of tergite 5 and tergites 6+ infuscate, contrasting with the generally pale yellow gaster. The gaster of E. sanchezi is uniformly yellowish with, at most, an indistinct infuscation on tergites 3+. Most specimens of FE. sanchezi have only the posterior half of the mesoscutum dark-marked, whilst the 282 IAN D. GAULD mesoscutal vittae of E. pescadori are blackish. Correlating with these colour differences are some structural differences. These are tabulated below. pescadori sanchezi value of CI 0.16-0.21 0.25-0.43 length of 20th flagellar segment to its width 2.0-2.4 2.42.7 laterotergite 2 upturned pendant 1st abscissa of Rs in hind wing straight bowed distal sclerite near proximal sclerite strong weak Enicospilus pescadori belongs to the E. marini species-complex (see that species). BIOLOGICAL INFORMATION. The range of this species extends from northern Costa Rica (11°N) south to Bahia, Brazil (13°S). In Costa Rica it has only been collected in the seasonally dry forest in Santa Rosa National Park where isolated individuals have occasionally been taken between April and December. A single female has been reared from a larva which might be a species of Eulepidotis (Noctuidae) (rearing reference number: 82-SRNP-344A). MATERIAL EXAMINED Holotype 2, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, vii.1982 (Janzen & Hallwachs) (BMNH). Paratypes. Brazil: 1 9, Bahia, Encruzilhada, 960 m, xi.1972 (Alvarenga) (TC). Costa Rica: Guanacaste Prov.: 4 2, Santa Rosa National Park, dry hill, vi, viii, x, xi.1977 (Janzen) (TC); 1 9, Santa Rosa National Park, 300 m, xii.1982 (Janzen & Hallwachs) (BMNH); 1 CO’, 1 9, same locality and collectors, iv, v.1983 (BMNH). Enicospilus howdenorum sp. n. (Figs 142, 145, 211, 332) Description. Mandibles long, proximally strongly narrowed, distally parallel-sided, apically twisted 15—25°; upper mandibular tooth cylindrical, 1.6-1.8 times as long as the lower tooth; outer mandibular surface very slightly concave, the concave area weakly coriaceous and bearing long fine pubescence; proximal concavity weak. Labrum 0.2-0.3 times as long as broad; malar space 0.3-0.4 times as long as basal mandibular width. Clypeus in profile moderately convex, margin impressed, acute; clypeus in front view 1.4-1.5 times as broad as long, with margin weakly convex apically. Lower face 0.68-0.73 times as broad as long, centrally polished, finely punctate. Head in dorsal view with genae rounded behind eye; posterior ocellus close to eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.8 times the basal mandibular width away from mandible. Antenna long and slender, with 64-66 flagellar segments; 20th segment 2.1—2.4 times as long as broad. Mesoscutum polished, finely and inconspicuously punctate, in profile evenly rounded; notauli vestigial. Mesopleuron polished, more or less evenly finely punctate; epicnemial carina inclined towards anterior margin of pleuron, its upper end often weak or obsolescent. Scutellum in profile moderately convex, laterally carinate for 0.9 or more of its length; scutellum in dorsal view 1.6—-1.7 times as long as anteriorly broad, finely punctate, with isolated striae posteriorly. Metapleuron strongly convex, polished and finely punctate (Fig. 211); submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly rounded (Fig. 145); anterior transverse carina complete or evanescent at lateral extremities, posterior transverse carina absent; anterior area steep, striate; spiracular area moderately long, smooth; posterior area closely irregularly reticulate, becoming almost longitudinally striate close to anterior transverse carina; lateral longitudinal carina usually com- plete, joined to spiracular margin by a short carina. Fore wing length 12-14 mm; discosubmarginal cell as in Fig. 332; AI = 0.65-0.75; CI = 0.28-0.45; ICI = 0.37—-0.45; SDI = 1.10—1.19; cu-asubopposite to the base of Rs&M, at the most proximal to it by about its own thickness; marginal cell proximally uniformly hirsute; 1st subdiscal cell with central and distal parts sparsely hirsute. Hind wing with 7-8 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa bowed; first abscissa of Cu1 unusual in that it is bowed, and joining cu-a at an angle of less than 90°; cu-a unusually long, 0.40.5 times the length of the first abscissa of Cul (Fig. 142). OPHIONINAE OF TROPICAL MESOAMERICA 283 Fore leg with tibia distinctly flattened, with either very few, or without obvious spines on outer surface. Mid leg with longer tibial spur 1.4-1.5 times length of the shorter. Hind leg with coxa in profile 1.9-2.0 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.3—2.5 times as long as broad; claws of female moderately long, evenly curved, with close short pectinae, those of male similar. Gaster long and slender; tergite 2 in profile more than 6 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 2.53.5 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 7-9 bearing fine semidecumbent pubescence; gonosquama apically evenly rounded. Colour generally from pale brownish yellow to medium yellowish brown, with central lobe of meso- scutum weakly infuscate and with gaster indistinctly infuscate posteriorly; interocellar area yellowish; antenna yellowish brown; pterostigma from yellowish brown to quite dark brown; wings hyaline. VARIATION. The Cuban specimen is paler yellow in colour than the others. REMARKS. This species is named in honour of Henry and Anne Howden in recognition of their collecting efforts in Central America. Enicospilus howdenorum and E. sondrae form a distinctive sister-species pair. They are characterized by having cu-a in the hind wing 0.4—0.5 times as long as a bowed first abscissa of Cul (Fig. 142). In most other species this abscissa of Cu1 is straight, and almost invariably cu-a is much less than 0.25 times its length. Both E. howdenorum and E. sondrae have somewhat similar central sclerites, lack any trace of a distal sclerite and have cu-a in the fore wing virtually opposite the base of Rs&M. The two species may easily be separated by the characters given in the key (see also p. 284). E. howdenorum is only known to occur on the older larger Caribbean islands of Cuba, Hispaniola and Jamaica and its sister-species E. sondrae has a similar but slightly more extensive distribution (it occurs in Florida and may also occur on Dominica). This suggests that this species-complex may have arisen on the old Caribbean islands. BIOLOGICAL INFORMATION. Enicospilus howdenorum is only recorded from Cuba, Jamaica and Hispaniola where it has been collected at altitudes between 1050 and 1350 m. Nothing is otherwise known of its habitat preferences or its hosts. MATERIAL EXAMINED Holotype 9, Jamaica: Hardwar Gap, 1300 m, vii.1966 (Howden & Becker) (CNC). Paratypes. Cuba: 1 9, Sierra Maestra, 1050-1350 m, vii.1922 (Ballou & Bruner) (USNM). Haiti: 1 9, Desbarriére, La Hotte, 1300 m, x.1934 (Darlington) (MCZ). Jamaica: 3 0’, 1 9, Hardwar Gap, 1300 m, vii.1966 (Howden & Becker) (BMNH, CNC). Enicospilus sondrae sp. n. (Figs 146, 333) DEscriPTIOn. Mandibles moderately long, proximally strongly narrowed, distally weakly narrowed, api- cally twisted 30—40°; upper mandibular tooth quite slender, subcylindrical, 1.5—1.7 times as long as the lower tooth; outer mandibular surface sparsely pubescent, distally flat, proximally with a shallow weak concavity. Labrum 0.2 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.3—1.4 times as broad as long, with margin subtruncate apically. Lower face 0.74—0.76 times as broad as long, weakly polished, finely punctate with a little wrinkling centrally. Head in dorsal view with genae constricted behind eyes; posterior ocellus close to eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.8 times the basal mandibular width away from mandible. Antenna long and slender, with 50-57 flagellar segments; 20th segment 1.7—2.1 times as long as broad. Mesoscutum weakly polished, finely punctate, in profile evenly rounded; notauli vestigial. Mesopleuron polished, the upper part punctate to punctostriate, the lower part punctostriate to striate; epicnemial carina inclined towards anterior margin of pleuron, with upper end evanescent. Scutellum in profile moderately convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, smooth anteriorly, posteriorly with fine longitudinal striae. Metapleuron convex, punctate with diagonal rugae at least present posterodorsally (Fig. 146); submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina more or less complete, posterior transverse carina absent; anterior area steep, striate; spiracular area moderately long, finely punctate; posterior area irregularly rugose- 284 IAN D. GAULD coriaceous, usually with a median longitudinal wrinkle; lateral longitudinal carina weak but more or less complete, joined to spiracular margin by a short carina. Fore wing length 15-17 mm; discosubmarginal cell as in Fig. 333; AI = 1.00-1.38; CI = 0.31-0.40; ICI = 0.38-0.54; SDI = 1.10-1.25; cu-a subopposite to the base of Rs&M; marginal cell proximally very sparsely hirsute; 1st subdiscal cell with isolated hairs anterodistally. Hind wing with 5—7 hamuli on R1; Ist abscissa of Rs straight, 2nd abscissa almost straight; cu-a unusual in being 0.40.5 times as long as the bowed first abscissa of Cu1. Fore leg with tibia flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.5— 1.7 times length of the shorter. Hind leg with coxa in profile 1.7—1.8 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 1.7—1.8 times as long as broad; claws of female long, distally very abruptly curved through about 100°, with close short pectinae; claws of male similar. Gaster long and slender; tergite 2 in profile 5.0-6.0 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 4-5 times its own length. Ovipositor straight, its sheath narrow. Male with sternites 7-9 bearing highly modified pubescence — sternite 7 centrally glabrous with lateral longitudinal rows of long erect hairs, sternite 8 more or less entirely glabrous (partly obscured in material at hand so I could not see if the hair rows are present anteriorly), and sternite 9 with short fine decumbent pubescence; gonosquama quite short, apically rounded. Colour generally yellowish brown, head and scutellum bright yellow, mesoscutum with dark longitudi- nal vittae; interocellar area yellow; antenna golden; pterostigma golden; wings hyaline. VARIATION. The female from ‘Dominica’ has the metapleuron regularly punctate, without the rugae that characterize the other specimens. ReMaRKS. This species is dedicated to Sondra Ward in appreciation for her great help in preparing this manuscript. Enicospilus sondrae is closely related to E. howdenorum (see that species) and is most easily recognized by the centrally incrassate Rs+2r and the proximally sparsely hirsute marginal cell. The males of E. sondrae differ strikingly from those of E. howdenorum in the distribution of the hairs on the terminal sternites. In particular, E. howdenorum has scattered fine semidecumbent hairs present on sternite 7, whilst E. sondrae has long erect hairs arranged in rows, and the sternite is glabrous centrally. A similar hair pattern occurs in E. lacsa and E. echeverri, and possibly these two taxa may be related to E. sondrae. The distribution of these characters presents a considerable classificatory problem, and amply demonstrates the difficulties one has trying to group species of Enicospilus. E. howdenorum and E. sondrae share several striking autapomorphic features (hind wing venation; alar sclerites) which are not shared by E. lacsa and E. echeverri. However, E. sondrae, E. lacsa and E. echeverri all have males with very similar and highly specialized hair patterns on sternites 7-9 (see Fig. 200). These two suites of characters are incompatible; either E. sondrae evolved the hair pattern independently from the other two species, or E. howdenorum and E. sondrae evolved very similar fore and hind wings independently. The distribution of these species suggests the former, but clearly more evidence of relationship is required. BIOLOGICAL INFORMATION. Enicospilus sondrae is only known to occur on some Caribbean Islands and on the Florida Keys. A single damaged female in the BMNH is labelled as having been collected in ‘Dominica’. Whether this refers to the island of Dominica between Guadeloupe and Martinique in the Lesser Antilles, or to the Dominican Republic in eastern Hispaniola is uncertain. Nothing is known of the habitat preferences or host range of this species. MATERIAL EXAMINED Holotype 9, U.S.A.: Florida, Paradise Key, iv.1951 (Townes & Townes) (TC). Paratypes. Dominica: 1 9, no further locality data, 1901 (Nicholls) (BMNH). Dominican Republic: 1 9, Dajabon Prov., 13 km S. Loma de Cabrera, 400 m, v.1973 (Davis & Davis) (USNM). Jamaica: 1 @, Trelawney, Good Hope, vii.1966 (Howden) (CNC). U.S.A.: Florida: 2 9, 1 0’, Paradise Key, iv.1951 (Townes & Townes) (TC). Enicospilus oduberi sp. n. (Figs 123, 334) Description. Mandibles quite short, rather evenly tapered, but with a distinct proximoventral lobe, the apex of the mandible twisted about 30°; upper mandibular tooth very slender, depressed, about 1.8 times as long as the lower tooth; outer mandibular surface finely and sparsely pubescent, centrally slightly concave, and with a small deeper concavity on proximoventral lobe. Labrum 0.3 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin flat, slightly blunt; OPHIONINAE OF TROPICAL MESOAMERICA 285 clypeus in front view 1.3—1.4 times as broad as long, with margin subtruncate apically. Lower face 0.62— 0.63 times as broad as long, centrally polished, with indistinct punctures. Head in dorsal view with genae constricted behind eye; posterior ocellus very close to eye; FI = 68-72%; occipital carina mediodorsally complete, ventrally angled to join hypostomal carina about 0.8—0.9 times the basal mandibular width away from mandible. Antenna very long and slender, with 58—60 flagellar segments; 20th flagellar segment 3.1— 3.2 times as long as broad. Mesoscutum polished with isolated inconspicuous punctures, in profile abruptly rounded; notauli absent. Mesopleuron polished, the upper part with isolated punctures, the lower part similar, but with a little irregular wrinkling also; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for all of its length; scutellum in dorsal view 1.5 times as long as anteriorly broad, smooth with isolated wrinkles posteriorly. Metapleuron weakly convex, with close fine punctures; submetapleural carina fairly evenly anteriorly broadened; posterior transverse carina of meso- sternum complete. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area short, steep, rugose-striate; spiracular area long, smooth; posterior area with weak transverse rugae which are more irregular and difficult to discern near transverse carina; lateral longitudinal carina complete, joined to spiracular margin by a short carina. Fore wing length 10-11 mm; discosubmarginal cell as in Fig. 334; AI = 1.32-1.63; CI = 0.25-0.27; ICI = 0.35—-0.39; SDI = 1.15-1.22; cu-a from subopposite to distal to the base of Rs&M from by about its own thickness to by 0.1 times its own length; marginal cell proximally broadly glabrous; 1st subdiscal cell fairly evenly hirsute except for proximal end which is glabrous. Hind wing with 4 hamuli on R1; 1st abscissa of Rs very slightly curved, 2nd abscissa distinctly arcuate. Fore leg with tibia subcylindrical, with scattered spines on outer surface. Mid leg with longer tibial spur 1.7-2.0 times length of the shorter. Hind leg with coxa in profile 1.9 times as long as deep; trochantellus dorsally 0.2 times as long as broad; 4th segment of tarsus 2.1—2.2 times as long as broad; claws of female long, apically evenly curved, with quite short, widely spaced pectinae. Gaster slender; tergite 2 in profile 6-7 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 4.5—5.0 times its own length. Ovipositor apically slender, slightly decurved, its sheath narrow. Male unknown. Colour generally dirty yellow, with area between antennal bases, mesoscutal vittae, posterior margin of mesopleuron from upper corner to below level of episternal scrobe (Fig. 123), and anterior part of metapleuron dark-marked; gaster yellowish with tergites 4+ weakly infuscate; interocellar area yellowish, but infuscate between posterior ocelli; antenna golden, slightly infuscate apically; pterostigma yellow; wings hyaline. VARIATION. None remarkable. REMARKS. This species is named in honour of Daniel Oduber, in recognition of his efforts to establish the national parks of Costa Rica. Enicospilus oduberi is a small, delicate insect that resembles species in the E. marini species-complex in its general appearance, coloration and sculpture, though it differs in having the mandible much less strongly twisted. E. oduberi may easily be recognized by its colour pattern, glabrous marginal cell, very unequal mid tibial spurs and usually having cu-a slightly distal to the base of Rs&M. BIOLOGICAL INFORMATION. Only two females of Enicospilus oduberi have been collected. They were taken in Santa Rosa National Park, Costa Rica, at the end of the dry season. Nothing else is known about their biology. MATERIAL EXAMINED Holotype 2, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, iv.1983 (Janzen & Hallwachs) (BMNH). Paratype. Costa Rica: Guanacaste Prov.: 1 9, Santa Rosa National Park, 300 m, iv.1984 (Janzen & Hallwachs) (BMNH). Enicospilus gallegosi sp. n. (Figs 129, 139, 335) DescriPTION. Mandibles moderately long, proximally evenly narrowed, distally almost parallel-sided, apically twisted 20—30°; upper mandibular tooth from subcylindrical to slightly depressed, 1.6-1.8 times as long as the lower tooth; outer mandibular surface bearing fine sparse long hair, distally more or less flat, proximally with a shallow concavity. Labrum 0.2 times as long as broad; malar space 0.3-0.4 times as long as basal mandibular width. Clypeus in profile flat, margin thin; clypeus in front view 1.3—1.6 times as broad 286 IAN D. GAULD as long, with margin truncate apically. Lower face 0.70-0.75 times as broad as long, centrally smooth and polished with fine, inconspicuous punctures. Head in dorsal view with genae constricted behind eyes; posterior ocellus more or less contiguous with eye; FI = 73-78%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6—0.7 times the basal mandibular width away from mandible. Antenna long and slender, with 51-54 flagellar segments; 20th segment 2.0—2.3 times as long as broad. Mesoscutum polished, finely punctate, in profile abruptly rounded; notauli vestigial. Mesopleuron polished, the upper part punctate or punctostriate, the lower part from punctostriate to striate; epicnemial carina curved towards, but not reaching, anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.4-1.5 times as long as anteriorly broad, polished, anteriorly smooth but posteriorly with scattered longitudinal wrinkles. Meta- pleuron moderately convex, diagonally rugoso-striate or striate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina usually weak at extreme lateral ends, sometimes complete, posterior transverse carina absent; anterior area short, steep irregularly rugose; spiracular area quite long, smooth or finely punctate; posterior area transversely wrinkled, with the wrinkles tending to be concentric except often with an irregularity along median line; lateral longitudinal carina complete, not joined to spiracular margin by a short carina. Fore wing length 9-12 mm; discosubmarginal cell as in Fig. 335; AI = 1.20—-1.80; CI = 0.20-0.37; ICI = 0.33-0.50; SDI = 1.20-1.36; cu-a proximal to the base of Rs&M by 0.1—0.3 times its own length; marginal cell proximally broadly glabrous; 1st subdiscal cell fairly evenly but sparsely hirsute, proximally glabrous. Hind wing with 5-6 hamuli on R1; 1st abscissa of Rs straight (Fig. 129), 2nd abscissa more or less straight. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.4-1.5 times length of the shorter. Hind leg with coxa in profile 1.7-1.9 times as long as deep; trochantellus dorsally 0.40.6 times as long as broad (Fig. 139); 4th segment of tarsus 2.5—2.7 times as long as broad; claws of female rather short, evenly curved with stout sparse pectinae, those of male similar but with pectinae shorter. Gaster slender; tergite 2 in profile more than 6 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 3 times its own length. Ovipositor short, straight, its sheath narrow. Male with sternites 7-9 bearing scattered long stout, slightly curved erect hairs; gonosquama apically subacute. Colour generally yellowish, with three dark brown longitudinal mesoscutal vittae, and sometimes with an indistinct dark mark near upper hind corner of mesopleuron; interocellar area yellow, but sometimes with a slight trace of infuscation between the posterior ones; antenna golden; pterostigma yellow; wings hyaline. VARIATION. The Brazilian specimens have the alitrunk laterally smoother and more polished than Central American specimens. REMARKS. This species is named in honour of Luis Roberto Gallegos for his imaginative interactions with Guanacaste National Park in its formative days. Enicospilus gallegosi resembles several other Central American species, especially E. hemicrescellae (see p. 294). However, E. gallegosi may be distinguished from all others by the following combination of characters: the distal sclerite is distinctly separated from the proximal sclerite; the proximal corner of the marginal cell of the fore wing is broadly glabrous (Fig. 335); the hind trochantellus projects dorsally well beyond the distal end of the trochanter; the lateral longitudinal carina of the propodeum is complete and not joined to the spiracular margin; AI is relatively large. BIOLOGICAL INFORMATION. Enicospilus gallegosi is a rather rarely collected species that has a range which extends from about 17°N in southern Mexico south to about 14°S in Brazil. A single damaged specimen in the BMNH is labelled as ‘St. Dom’ and is believed to have originated from the Dominican Republic, although no other specimens of this species are known from the Caribbean. In Santa Rosa National Park isolated specimens of E. gallegosi have been collected during the first half of the wet season. Nothing is known of its biology. MATERIAL EXAMINED Holotype ©’, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, vii.1982 (Janzen & Hallwachs) (BMNH). Paratypes. Brazil: 2 9, Bahia, Encruzilhada, 960 m, iii & xi.1972 (Alvarenga) (TC); 1 0’, 3 9, same locality and collector, 980 m, xi.1974 (TC). Costa Rica: Guanacaste Prov.: 1 0’, 1 9, Santa Rosa National OPHIONINAE OF TROPICAL MESOAMERICA 287 Park, 300m, viii.1982, vi.1984 (Janzen & Hallwachs) (BMNH). Dominican Republic: 1 2, ‘St Dom.’ no further data (BMNH). Mexico: Chiapas: 2 2, 32 km N. Huixtla, 1000 m, vi.1969 (CNC). Panama: 1 9, Barro Colorado Island, 120 m, vi.1985 (Wolda) (BMNH). Enicospilus parkeri sp. n. (Figs 210, 336) Description. Mandibles quite long and stout, proximally strongly narrowed, distally almost parallel-sided, apically twisted 40—50°; upper mandibular tooth subcylindrical, 1.4-1.7 times as long as the lower tooth; outer mandibular surface sparsely pubescent, distally concave, and with a broad shallow proximal con- cavity. Labrum 0.2-0.3 times as long as broad; malar space 0.3-0.4 times as long as basal mandibular width. Clypeus in profile flat, margin subacute; clypeus in front view 1.5—1.7 times as broad as long, with margin subtruncate apically. Lower face 0.71—0.80 times as broad as long, centrally polished, punctate. Head in dorsal view with genae slightly inflated behind eyes; posterior ocellus very close to or contiguous with the eye; FI = 70-75%); occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.50.7 times the basal mandibular width away from mandible. Antenna long and quite slender, with 61-68 flagellar segments; 20th segment 1.8-2.1 times as long as broad. Mesoscutum from polished and finely punctate to matt, inconspicuously granulate, in profile abruptly rounded; notauli vestigial. Mesopleuron polished, the upper part punctate to punctostriate, the lower part punctostriate to striate, often with longitudinal wrinkling near ventral corner; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for 0.6 or more of its length; scutellum in dorsal view 1.4-1.5 times as long as anteriorly broad, punctate, though often with irregular rugosities present. Metapleuron ventrally flattened, punctate to puncto-coriaceous or even almost striate, and postero-dorsally with a ridge that usually bears rugae (Fig. 210); submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile abruptly declivous; anterior transverse carina complete except at lateral extremities, posterior transverse carina absent or present as a sublongitudinal crest laterally; anterior area short, steep, bearing isolated striae; spiracular area moderately long, smooth; posterior area irregularly rugose, usually with rugosities centrally tending to form longitudinal striae; lateral longitudinal carina present anteriorly, joined to spiracular margin by a short weak carina. Fore wing length 17-22 mm; discosubmarginal cell as in Fig. 336; AI = 1.06-1.18; CI = 0.40-0.62; ICI = 0.42-0.52; SDI = 1.39-1.53; cu-a proximal to the base of Rs&M by 0.2-0.4 times its own length; marginal cell proximally broadly glabrous; 1st subdiscal cell with distal end and anterior 0.4 hirsute. Hind wing with 8-9 hamuli on R1; 1st abscissa of Rs more or less straight, 2nd abscissa weakly bowed. Fore leg with tibia subcylindrical, with scattered spines on outer surface. Mid leg with longer tibial spur 1.3-1.5 times length of the shorter. Hind leg with coxa in profile 1.7—1.9 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.5—2.8 times as long as broad; claws of female short, apically abruptly curved, with fine close pectinae; those of male very similar but with pectinae closer. Gaster slender; tergite 2 in profile 6-7 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 4.04.5 times its own length. Ovipositor slender, straight, its sheath narrow. Male with sternites 7-9 bearing dense long erect pubes- cence; gonosquama long, apically rounded. Colour generally pale yellowish, though often with indistinct darker brownish marks, especially on mesoscutum; gaster with dorsal margins of most tergites infuscate or blackish, often more extensively dark- marked; interocellar area usually yellowish, sometimes weakly infuscate; antenna golden, but generally infuscate apically; pterostigma yellowish; wings hyaline or weakly infumate. VARIATION. The most striking variation in this species is in the colour of the gaster. All specimens have at least the dorsal margin of tergites 3-6 blackish, and often the lateral parts of the same tergites very pallid. Some individuals have the posterior and ventral parts of the tergites infuscate so the darker patterning is somewhat scalariform in lateral view. A possibly conspecific specimen from Brazil, that has been excluded from the paratype series, is rather uniformly yellowish brown. REMARKS. This species is named in honour of Pam Parker for her tireless efforts in conservation biology. Enicospilus parkeri belongs to the E. dispilus species-complex, an exceptionally difficult species group to resolve as several species show great variation. E. parkeri may be separated from most other species with a central sclerite and golden antennae by the fact that the central sclerite is crescentic, the fenestra is rather small and the posteromedial part of Rs+2r has a weak swelling opposite the anterior end of the proximal sclerite (Fig. 336). In these features it resembles E. georginae (Fig. 338) and some northern American 288 IAN D. GAULD specimens of E. dispilus, and care is required to separate these three taxa. Their critical features are compared below. dispilus parkeri georginae anterior propodeal transverse carina incomplete complete complete Cl <0.40 0.40-0.62 0.59-0.79 metapleuron posterodorsally punctate rugose/ rugose/ striate striate marginal cell proximally proximally more or less of fore wing glabrous glabrous hirsute 1st abscissa of Rs in hind wing straight straight bowed In Costa Rica most specimens of E. parkeri have only the dorsum of the gaster narrowly black and the female subgenital plate infuscate. E. georginae always has the gaster mottled. BIOLOGICAL INFORMATION. Enicospilus parkeri is a Mesoamerican species that is known to occur from about 19°N in southern Mexico south to 8°N in northern Panama, and I have seen a single, possibly conspecific male from Brazil. It has been collected in lower montane rainforests at altitudes between 600 and 1500 m, though large numbers of individuals have never been collected at any one site. At Monteverde, Costa Rica, where most specimens have been taken, isolated individuals have been collected in January, May, July and August. Despite intensive collecting E. parkeri has never been taken in lowland sites such as Santa Rosa and Barro Colorado. The hosts of this species are not known. MATERIAL EXAMINED Holotype 9, Costa Rica: Guanacaste Prov., Estacion Mengo, SW. side of Volcan Cacao, 1100 m, vi.1987 (Janzen) (BMNH). Paratypes. Costa Rica: Alajuela Prov.: 1 0’, Finca San Gabriel, 16 km E. Quebrada Grande, 630 m, iii.1983 (Janzen & Hallwachs) (BMNH): Cartago Prov.: 1 CO’, Tapanti, Rio Grande de Orosi, 13-1400 m, i.1985 (Janzen & Hallwachs) (BMNH): Puntarenas Prov: 1 2, Monteverde, 1300 m, vii.1981 (Janzen & Hallwachs) (BMNH); 1 co’, same locality, v.1984 (Fogden) (BMNH); 2 9, same locality, viii.1985, 1.1986 (Haber) (BMNH): San José Prov.: 1 2, Estacion Zurqui (el Tunel), Braulio Carrillo National Park, 1500 m, xi.1985 (Chacon & Chacon) (BMNH). Mexico: Oaxaca: 1 9, 19 km S. Valle Nacional, 1000 m, v.1971 (Howden) (TC). Panama: 4 9, Chiriqui, Fortuna, 1050 m, ii, iv & v.1978 (Wolda) (RNH). Non-paratypic material. Brazil: 1 9, Santa Catarina, Nova Teutonia, i.1939 (Plaumann) (BMNH). Enicospilus ulfstrandi sp. n. (Fig. 337) Description. Mandibles moderately long, distally evenly tapered, apically twisted 15—25° with upper tooth depressed, 1.3-1.6 times as long as the lower; outer mandibular surface slightly concave with a weak proximal concavity bearing fine hairs. Labrum 0.2-0.3 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile almost flat, margin blunt; clypeus in front view 1.2-1.3 times as broad as long, apically more or less truncate. Lower face 0.65—0.70 times as broad as long, polished, centrally punctostriate, laterally punctate. Head in dorsal view with genae constricted; posterior ocellus contiguous with eye; FI = 65-70%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.7 times the basal mandibular width away from mandible. Antenna slender, with 53-56 flagellar segments; 20th segment 2.1—2.3 times as long as broad. Mesoscutum polished, punctate, in profile abruptly rounded; notauli absent. Mesopleuron weakly polished, the upper part punctostriate to striate, the lower part rugose-striate; epicnemial carina curved towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for at least 0.5 of its length, sometimes completely carinate; scutellum in dorsal view 1.4-1.6 times as long as anteriorly broad, anteriorly smooth, posteriorly wrinkled. Metapleuron weakly evenly convex, from striate to finely irregularly rugose; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area steep, rugose-striate; spiracular area moderately long, OPHIONINAE OF TROPICAL MESOAMERICA 289 smooth; posterior area finely irregularly reticulate, sometimes tending to be longitudinally striate medi- ally; lateral longitudinal carina complete anteriorly, posteriorly rather irregular, sometimes absent, the anterior part sometimes joined to spiracular margin by a weak short carina. Fore wing length 11-13 mm; discosubmarginal cell as in Fig. 337; AI = 0.52-0.91; CI = 0.12-0.25; ICI = 0.38-0.53; SDI = 0.95-1.08; cu-a from opposite to proximal to the base of Rs&M by about 0.2 of its own length; marginal cell proximally very slightly more sparsely hirsute than centrally; 1st subdiscal cell with anterior and distal parts hirsute. Hind wing with 5-6 hamuli on R1; 1st abscissa of Rs very weakly bowed, 2nd abscissa straight. Fore leg with tibia strongly flattened, with isolated spines on outer surface. Mid leg with longer tibial spur 1.4-1.6 times length of the shorter. Hind leg with coxa in profile 1.9-2.0 times as long as deep; trochantellus dorsally 0.10.2 times as long as broad; 4th segment of tarsus 2.7—2.8 times as long as broad; claws of female quite long, apically evenly curved, with long close pectinae, those of male with pectinae slightly shorter. Gaster slender; tergite 2 in profile more than 5 times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by about 4 times its own length. Ovipositor slender, straight, its sheath narrow. Males with sternites 7-9 bearing scattered long erect, rather fine hairs; gonosquama evenly rounded. Colour generally yellowish brown; mesoscutum usually slightly darker anteriorly on central lobe, sometimes with darker longitudinal vittae; interocellar area yellow; antenna yellowish with distal few segments slightly infuscate; pterostigma golden; wings hyaline. VARIATION. The Mexican specimens have the meso- and metapleurae more uniformly and finely striate than the Costa Rican ones which always have a rugose metapleuron, and sometimes have similar sculpture on the lower part of the mesopleuron. The Ecuadorian specimen is morphologically very similar to the Mexican ones, but has dark mesoscutal vittae and has segments 3+ of the gaster infuscate. REMARKS. This species is named in honour of Staffan Ulfstrand in recognition of his deep interest in Costa Rican biology and conservation. Enicospilus ulfstrandi may be distinguished from other Mesoamerican species by the characteristically small circular central sclerite and by the faint, but generally wide, distal sclerite (Fig. 337). The sculpture, venation and form of the mandibles suggest that E. ulfstrandi belongs to the E. dispilus species-group, but no other Central American species in this complex has such a small central sclerite. BIOLOGICAL INFORMATION. Enicospilus ulfstrandi is a rather widespread species whose range extends from southern Florida and the extreme south of Texas southwards throughout Mexico to Venezuela and Ecuador (Map 31). In Costa Rica most individuals have been collected in the seasonally dry forests of Santa Rosa National Park. The cumulative distributional data for this site for 1982-86 are: Lok MA Mete) Als ON D ee ee These data indicate that E. ulfstrandi is present during the dry season, and the collecting peak (March/ April) corresponds with the driest time of the year when no hosts apparently are present. E. ulfstrandi also has never been collected in wet forest sites such as Monteverde, though a single individual was collected at light on Barro Colorado Island. Quite what the hosts of this species are in Guanacaste remains a mystery. MATERIAL EXAMINED Holotype 9, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, iii.1984 (Janzen & Hallwachs) (BMNH). Paratypes. Costa Rica: Guanacaste Prov.: 1 9, Cerro el Hacha, 3-400 m, xi.1986-i.1987 (Janzen & Hallwachs) (BMNH); 1 9, Rincén de la Vieja, Mirador, 900 m, iii.1984 (Janzen, Hallwachs & Gauld) (BMNH); Santa Rosa National Park, 300 m, 1 9, iii.1982, 1 0’, 1.1983, 1 9, iii.1983, 1 9, iv.1983, 107, 11.1984, 2 9, iii.1984, 2 9, v.1984 (Janzen & Hallwachs) (BMNH); 1 Q, same locality, iv.1984 (Gauld) (BMNH): San José Prov.: 1 9, San Antonio de Escazt, 1300 m, v-vi.1981 (Eberhard) (UMC). Ecuador: 1 2, Rio Leén, 2100 m, xi.1970 (Pena) (TC). Mexico: Baja California: 2 9, 8.5 km N. La Paz on highway 1, xil. 1978 (Weismann et al.) (CAS); 1 0’, San Domingo, vii.1938 (Michelbacher & Ross) (CAS): Guerrero: 1 Q, Tepetlapa, 100m, x.1904 (Smith) (BMNH): Nuevo Leon: 2 9, 8kmS. Monterrey, vii.1963 (Howden) (CNC); 1 2, Monterrey, Huasteca Canyon, vii.1963 (Howden) (CNC): Chihuahua: 1 0’, 1 9, Presidio, 1904 (Godman-Salvin) (BMNH): San Luis Potosi: 1 9, El Salto Falls, vi.1963 (Woodruff) (FSCA): Tamaulipas: 1 0’, Mesa de Llera, nr Cuidad Victoria, vi.1977 (Porter & Cerbone) (FSCA). Panama: 1 CO’, Barro Colorado Island, 120 m, vii.1983 (Wolda) (BMNH). U.S.A.: Florida: 1 9, Highlands Co., vi.1967 IAN D. GAULD 290 ‘poyda][oo usaq sey ipuvais{jn snjidsoouy yorum ye sanyesoqT Ee dey] OPHIONINAE OF TROPICAL MESOAMERICA 291 (Heinrich) (CNC); 1 9, same locality, vi.1979 (Weems & Webber) (FSCA); 1 2, same locality, i.1980 (Weems & Carrel) (FSCA); 1 0’, Levy Co., Williston, xii.1949 (Townes) (TC); Pinellas Co., 1 &’, Tarpon Springs, xii.1949 (Townes) (TC): Texas: 1 9, Cameron Co., palm jungle near Southmost, vi.1948 (Mason) (CNC). Venezuela: 2 2, Aragua, Portachuelo Pass, nr Rancho Grande, iv.1960 (Test) (UMC); 1 9, Aragua, Rancho Grande, iv.1960 (Test) (UMC). Enicospilus georginae sp. n. (Figs 144, 338) DEscriPTION. Mandibles quite long and stout, proximally strongly narrowed, distally almost parallel-sided, apically twisted 30—-50°; upper mandibular tooth subcylindrical, 1.4-1.7 times as long as the lower tooth; outer mandibular surface sparsely pubescent, distally concave, and with a broad shallow proximal con- cavity. Labrum 0.3 times as long as broad; malar space 0.3 times as long as basal mandibular width. Clypeus in profile weakly convex, margin blunt; clypeus in front view 1.41.5 times as broad as long, with margin subtruncate apically. Lower face 0.69-0.72 times as broad as long, centrally polished, punctate. Head in dorsal view with genae constricted behind eyes; posterior ocellus very close to or contiguous with the eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.7 times the basal mandibular width away from mandible. Antenna long and quite stout, with 63-67 flagellar segments; 20th segment 1.9-2.0 times as long as broad. Mesoscutum polished and finely punctate, in profile evenly rounded; notauli vestigial. Mesopleuron polished, the upper part punctate to punctostriate, the lower part striate, with longitudinal wrinkling near ventral corner; epicnemial carina curved towards anterior margin of pleuron. Scutellum in profile moder- ately convex, laterally carinate for 0.60.8 of its length; scutellum in dorsal view 1.2-1.3 times as long as anteriorly broad, punctate. Metapleuron ventrally flattened, punctate and postero-dorsally with a ridge that bears weak rugae or striations; submetapleural carina evenly anteriorly broadened; posterior trans- verse carina of mesosternum complete. Propodeum in profile abruptly declivous; anterior transverse carina complete, posterior transverse carina present as a sublongitudinal crest laterally; anterior area short, steep, bearing isolated striae; spiracular area moderately long, smooth; posterior area irregularly rugose, with rugosities centrally tending to form longitudinal striae; lateral longitudinal carina complete, joined to spiracular margin by a short discontinuous carina. Fore wing length 21-23 mm; discosubmarginal cell as in Fig. 338; AI = 0.86—1.11; CI = 0.59-0.79; ICI = 0.43-0.50; SDI = 1.28—1.42; cu-a proximal to the base of Rs&M by 0.3-0.5 times its own length; marginal cell proximally only very slightly more sparsely hirsute than it is centrally, with a very narrow glabrous area adjacent to Rs+2r; 1st subdiscal cell with anterior 0.4 hirsute. Hind wing with 7-10 hamuli on R1; Ist abscissa of Rs distinctly bowed (Fig. 144), 2nd abscissa from almost straight to weakly arcuate. Fore leg with tibia subcylindrical, with scattered spines on outer surface. Mid leg with longer tibial spur 1.4-1.6 times length of the shorter. Hind leg with coxa in profile 1.8—-1.9 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.0—2.2 times as long as broad, unusual in that it bears denser, long fine pubescence than many species; claws of female quite short, apically abruptly curved, with fine, close pectinae; claws of male similar. Gaster slender; tergite 2 in profile about 7 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 3.0 times its own length. Ovipositor slender, straight, its sheath moderately stout. Male with sternites 7-9 bearing dense long erect pubescence; gonosquama apically rounded. Colour generally pale yellowish brown, with head, pronotum, mesoscutal stripes, scutellum, upper and lower hind parts of mesopleuron, metanotum laterally, and metapleuron in part bright yellow; gaster with ventral and posterior margins of tergites 3-6 infuscate, anterocentral parts of same tergites pale yellow so the gaster appears mottled; interocellar area yellowish; antenna golden; pterostigma yellowish; wings moderately strongly infumate, yellowish. VARIATION. None remarkable. REMARKS. This species is named after Pensiédn Georgina, on the Cerro de la Muerte, Costa Rica, for providing food and shelter for many a cold tropical biologist. Enicospilus georginae is a large species that is most easily distinguished by the mottled coloration of the gaster, the large value of CI, the very obtuse posterodistal corner of the 2nd discal cell, the yellowish wings and the crescent-shaped central sclerite (Fig. 338). Structurally it is very similar to E. parkeri (see p. 287). BIOLOGICAL INFORMATION. In Mesoamerica, Enicospilus georginae is found at higher altitudes than almost any other species in the genus. It has only been collected above 2000 m in tall oak forest in Costa Rica, and specimens have been taken in a similar habitat in northern Panama. Its hosts are not known. 292 IAN D. GAULD MATERIAL EXAMINED Holotype 9, Panama: Chiriqui Prov., Guadalupe Arriba, 2200 m, vi.1984 (Wolda) (BMNH). Paratypes. Costa Rica: San José Prov.: 1 0’, 1 9, San Gerardo de Dota, Cerro de la Muerte, 2430 m, xii.1981 (Janzen & Hallwachs) (BMNH). Panama: 3 9, Chiriqui, Guadalupe Arriba, 2200 m, iii-vi.1984/5 (Wolda) (BMNH). Enicospilus masoni sp. n. (Figs 141, 339) Description. Mandibles moderately long, distally fairly evenly narrowed, apically twisted 35—45°; upper mandibular tooth subcylindrical, 1.5—1.8 times as long as the lower tooth; outer mandibular surface sparsely pubescent, flat, with a weak proximal concavity. Labrum 0.3—0.4 times as long as broad; malar space 0.3-0.4 times as long as basal mandibular width. Clypeus in profile flat or very weakly convex, margin blunt; clypeus 1.3—1.5 times as broad as long, margin apically subtruncate. Lower face 0.67-0.71 times as broad as long, polished, centrally punctate. Head in dorsal view with genae rounded behind eye; posterior ocellus very close to eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6 times the basal mandibular width away from mandible. Antenna long and slender, with 63-65 flagellar segments; 20th segment 2.4~2.5 times as long as broad. Mesoscutum polished, punctate, in profile abruptly rounded; notauli vestigial. Mesopleuron polished, the upper and lower parts punctate, with a few irregular weak rugae radiating posteriorly from the epicnemial carina; epicnemial carina curved towards anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.4-1.5 times as long as anteriorly broad, punctate with isolated rugae posteriorly. Metapleuron evenly weakly convex, punctate with isolated rugae; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina sinuous, usually complete, sometimes lateromedially interrupted, posterior transverse carina absent; anterior area short, steep, bearing irregular striae; spiracular area short, punctate; posterior area irregularly rugose-reticulate; lateral longitudinal carina present anteriorly, joined to spiracular margin by a short carina. Fore wing length 15-17 mm; discosubmarginal cell as in Fig. 339; AI = 0.86-1.07; CI = 0.61-0.83; ICI = 0.43-0.50; SDI = 1.31-1.50; cu-a proximal to Rs&M by 0.1-0.3 times its own length; marginal cell proximally uniformly hirsute; 1st subdiscal cell with anterior and distal parts broadly hirsute. Hind wing with 7 hamuli on R1; 1st abscissa of Rs more or less straight (Fig. 141), 2nd abscissa arcuate. Fore leg with tibia very slightly flattened, with scattered stout spines on outer surface. Mid leg with longer tibial spur 1.3—1.5 times length of the shorter. Hind leg with coxa in profile 1.8—1.9 times as long as deep; trochantellus dorsally <0.1 times as long as broad; claws of female moderately long, evenly curved, with close, rather short pectinae; claws of male similar. Gaster long and slender; tergite 2 in profile more than 5 times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by about 3-4 times its own length. Ovipositor straight, apically very slender, its sheath quite narrow. Male with sternites 7-9 bearing exceptionally long stout erect hairs; gonosquama evenly rounded apically. Colour generally yellowish, with the central mesoscutal vitta slightly to moderately infuscate, and with the posterior tergites of the gaster from not to moderately strongly infuscate; interocellar area yellowish; antenna brownish yellow, often infuscate distally; pterostigma yellowish brown; wings hyaline. VARIATION. The female from Monteverde, Costa Rica, differs from other specimens in having the base of the antennae black, the upper part of the mesopleuron punctostriate and the lower part rugose, the metapleuron exceptionally rugose, the lateral longitudinal carina complete and tergites 3+ of the gaster black. This Costa Rican specimen is only tentatively regarded as conspecific, and is excluded from the paratype series. REMARKS. This species is named in honour of the distinguished Canadian entomologist, Dr Bill Mason, who collected a large number of ophionines in tropical Mexico. Enicospilus masoni can most easily be recognized by the form of the alar sclerites and by the large CI (Fig. 333). The form of the mandibles and its general sculpture suggest it belongs to the E. dispilus species- group. BIOLOGICAL INFORMATION. Enicospilus masoni has most commonly been collected in Mexico where occa- sional individuals have been taken from about 24°N in Durango south to southern Chiapas (15°N). A single, possibly conspecific individual from Costa Rica suggests it may occur throughout Central America. Nothing is known of the hosts of this insect. OPHIONINAE OF TROPICAL MESOAMERICA 293 MATERIAL EXAMINED Holotype 2, Mexico: Atoyac, v.1904 (Smith) (BMNH). Paratypes. Mexico: Chiapas: 1 2, Muste, nr Huixtla, 440 m, ix.1970 (Welling) (CNC): Durango: 1 0’, 10 km S. Durango, 2000 m, vii.1964 (Mason) (CNC); 1 9, 48 km W. La Cuidad, 2100m, vil964 (Mason) (CNC): Sinaloa: 1 9, 42 kmN. of Pericos, viii.1960 (Arnaud, Ross & Rentz) (CAS): Veracruz: 1 2 Fortin de las Flores, Sumidero, Cerveceria Ing. D. Rabago Res., 700-1000 m, v.1965 (Weems) (FSCA). Non-paratypic material. Costa Rica: 1 2, Puntarenas Prov., Monteverde, ii.1984 (Cameron) (WC). Enicospilus kleini sp. n. (Figs 212, 340) DEscrIPTION. Mandibles moderately long, proximally strongly narrowed, distally weakly narrowed, api- cally twisted 25—35S°; upper mandibular tooth slender, depressed, 1.2—1.5 times as long as the lower tooth; outer mandibular surface bearing fine pubescence, centrally slightly concave, proximally more or less flat. Labrum 0.2-0.3 times as long as broad; malar space 0.3—-0.4 times as long as basal mandibular width. Clypeus in profile almost flat, margin blunt; clypeus in front view 1.3—1.5 times as broad as long, with margin truncate or subtruncate apically. Lower face 0.63—0.71 times as broad as long, polished, punctate, centrally sometimes tending to punctostriate. Head in dorsal view with genae constricted behind eye; posterior ocellus very close to eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.7—0.9 times the basal mandibular width away from mandible. Antenna long and slender, with 54-59 flagellar segments; 20th segment 2.1—2.4 times as long as broad. Mesoscutum polished, finely punctate, in profile quite steeply rounded; notauli vestigial. Mesopleuron polished, the upper part generally sparsely punctate, sometimes ventrally tending to punctostriate, the lower part usually punctostriate, occasionally striate or even punctate; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.41.6 times as long as anteriorly broad, from polished and finely punctate to bearing transverse rugae. Metapleuron weakly convex, diagonally striate (Fig. 212), the striae some- times coarse and rather irregular; submetapleural carina generally rather narrow and anteriorly rather abruptly broadened into a small rounded lobe; posterior transverse carina of mesosternum complete. Propodeum quite long, in profile evenly declivous; anterior transverse carina complete except for lateral extremities, posterior transverse carina absent; anterior area short, steep striate; spiracular area long, smooth; posterior area irregularly rugose, sometimes with the rugae tending to be somewhat concentric; lateral longitudinal carina generally only present anteriorly, joined to spiracular margin by a short carina. Fore wing length 12-16 mm; discosubmarginal cell as in Fig. 340; AI = 0.86-1.11; CI = 0.21-0.29; ICI = 0.47-0.57; SDI = 1.12-1.31; cu-a proximal to the base of Rs&M by 0.1-0.3 times its own length; marginal cell proximally slightly more sparsely hirsute than it is centrally; 1st subdiscal cell with anterior and distal margins hirsute. Hind wing with 6-8 hamuli on R1; Ist abscissa of Rs straight, 2nd abscissa straight. Fore leg with tibia slightly flattened, with scattered inconspicuous spines on outer surface. Mid leg with longer tibial spur 1.5—1.6 times length of the shorter. Hind leg with coxa in profile 1.8—1.9 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.4-2.9 times as long as broad; claws of female moderately long, apically evenly curved with long fairly widely interspaced pectinae; claws of male similar but with pectinae a little finer, shorter and closer together. Gaster long and slender; tergite 2 in profile 5.5—6.5 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical to oval and separated from anterior margin of tergite by 3-5 times its own length. Ovipositor slightly decurved, its sheath quite narrow. Male with sternites 7-9 bearing numerous long stout erect hairs; gonosquama apically evenly rounded. Colour generally brownish yellow, with central and sometimes also the lateral mesoscutal vittae darker brown; interocellar area yellowish, very rarely slightly infuscate between the posterior ocelli; antenna yellowish; pterostigma yellowish brown; wings hyaline. VARIATION. None remarkable. REMARKS. This species is named in honour of William Klein for his many contributions to the development of Guanacaste National Park, Costa Rica. Enicospilus kleini is most easily recognized by the characteristic form of the central sclerite in the fore wing (Fig. 340). Structurally it is very similar to an undescribed species from southern Brazil (BMNH, CNC) but differs in having Rs+2r more sinuous and erect hairs on the posterior sternites of the male gaster. BIOLOGICAL INFORMATION. Enicospilus kleini is a widespread Neotropical species whose range extends from 18°N in southern Mexico, south to about 23°S in Brazil. It seems to be associated with dry or semidry areas, 294 IAN D. GAULD and in Costa Rica it has only been collected in the seasonally dry parts of Guanacaste Province. In Santa Rosa National Park most specimens have been collected in the dry season. The cumulative seasonal distribution for 1980-85 is: Ji-3? Boi Me: AORM Ly goer 82 TMUANIGS -enGe TN AD 4 3 AL get oreo J 2" Nothing is known of the natural history of this species. MATERIAL EXAMINED Holotype 2, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, iii.1983 (Janzen & Hallwachs) (BMNH). Paratypes. Brazil: 1 O, 34 9, Bahia, Encruzilhada, 960-980 m, xi. 1972-4 (Alvarenga) (TC); 2 9, Ceara, Barbalha, 400 m, v.1969 (Alvarenga) (TC); 8 2, Minas Gerais, Aguas Vermelhas, 800 m, xii.1983 (Alvarenga) (TC); 3 2, Minas Gerais, Pedra Azul, 800 m, xi.1971-2 (Alvarenga & Seabra) (TC); 1 9, Sao Paulo, Boracea Salesopolis, xii.1969 (Campbell) (CNC). Colombia: 1 2 , Amazon River, Monkey Island, nr Leticia, ix.1979 (Tidwell) (USNM). Costa Rica: Guanacaste Prov.: 1 2, W. of Carmona Nicoya, 600— 700 m, viii.1982 (Janzen & Hallwachs) (BMNH); Santa Rosa National Park, 300 m, 1 9, vii.1980, 1 Co’, Hallwachs) (BMNH); 1 9, same locality, vi.1985 (Gauld) (BMNH); 1 9, Volcan Orosi, Casa Mariksa (= Maritza), 800 m, vii.1986 (Gauld) (BMNH). Ecuador: 1 9, Playas de Montalro, iv.1928 (Clarke & McIntyre) (TC). Honduras: 1 9 , Morazan, Zamorano, oak quebrada, 1000 m, viii.1948 (Hubbell) (UMC). Mexico: Tabasco: 1 0’, Teapa, ii.1904 (Smith) (BMNH). Venezuela: 2 2, Aragua, Portachelo Pass, nr Rancho Grande, iv.1960 (Test) (UMC). Enicospilus hemicrescellae sp. n. (Fig. 342) Description. Mandibles moderately long, proximally strongly narrowed, distally more weakly narrowed, apically twisted 30-40°; upper mandibular tooth depressed, 1.3—1.4 times as long as the lower tooth; outer mandibular surface bearing fine sparse pubescence, distally more or less flat, proximally with a shallow ventrobasal concavity. Labrum 0.2 times as long as broad; malar space 0.2-0.3 times as long as basal mandibular width. Clypeus in profile flat, margin blunt; clypeus in front view 1.4~1.5 times as broad as long, with margin slightly convex apically. Lower face 0.65—0.68 times as broad as long, centrally smooth and polished with fine, inconspicuous punctures. Head in dorsal view with genae constricted behind eyes; posterior ocellus more or less contiguous with eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6—0.7 times the basal mandibular width away from mandible. Antenna long and slender, apically incomplete; 20th segment 2.5—2.6 times as long as broad. Mesoscutum polished, virtually impunctate, in profile abruptly rounded with anterior margin out- turned; notauli vestigial. Mesopleuron polished, the upper part punctate grading to punctostriate, the lower part striate; epicnemial carina curved towards, but not reaching, anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.4~1.5 times as long as anteriorly broad, polished, anteriorly smooth but posteriorly with scattered longitudinal wrinkles. Metapleuron moderately convex, longitudinally striate; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile evenly declivous; anterior transverse carina complete or weak at extreme lateral ends, posterior transverse carina absent; anterior area short, steep irregularly rugose; spiracular area quite long, smooth or finely punctate; posterior area transversely wrinkled, with an irregular median longitudinal rugosity; lateral longitudinal carina incomplete posteriorly, joined to spiracular margin by a short carina. Fore wing length 10-12 mm; discosubmarginal cell as in Fig. 342; AI = 0.78-1.00; CI = 0.22-0.29; ICI = 0.43-0.47; SDI = 1.14-1.34; cu-a proximal to the base of Rs&M by 0.1 times its own length; marginal cell proximally slightly more sparsely hirsute than it is centrally; 1st subdiscal cell fairly evenly but sparsely hirsute. Hind wing with 5—6 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa more or less straight. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.4-1.5 times length of the shorter. Hind leg with coxa in profile 1.7-1.9 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.5—2.7 times as long as broad; claws of female quite long, evenly curved with stout sparse pectinae; claws of male similar but with pectinae shorter. OPHIONINAE OF TROPICAL MESOAMERICA 295 Gaster slender; tergite 2 in profile more than 6 times as long as posteriorly deep, laterotergite folded under, thyridia elli4 times its own length. Ovipositor short, straight, its sheath narrow. Male with sternites 6-9 bearing scattered long stout, slightly curved erect hairs; gonosquama apically subacute. Colour generally pale yellowish, with three dark brown longitudinal mesoscutal vittae; terminal seg- ments of gaster very weakly infuscate; interocellar area yellow; antenna golden; pterostigma yellow; wings hyaline. VARIATION. None remarkable. REMARKS. This species is named after the resilient agouti, Small Half Grown, in recognition of her contributions to tropical rodent biology. Enicospilus hemicrescellae is very similar to E. gallegosi in general appearance, and in particular in the form of the alar sclerites (cf. Figs 335 and 342). The two species may be confused unless some care is taken. Their critical features are compared below. hemicrescellae gallegosi mandibles more evenly tapered, twisted 30-40° mandibles distally parallel-sided, twisted 20—30° flagellum very slender, 20th flagellar segment 2.5— flagellum less slender, 20th flagellar segment 2.0— 2.6 times as long as broad 2.3 times as long as broad hind trochantellus short hind trochantellus elongate marginal cell of fore wing sparsely pubescent marginal cell of fore wing glabrous proximally proximally BIOLOGICAL INFORMATION. Enicospilus hemicrescellae is only known to occur in north-western Costa Rica. It has occasionally been collected in Santa Rosa National Park where isolated specimens have been taken during the dry season. Nothing is known about its biology. MaTERIAL EXAMINED Holotype ©’, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, iv.1983 (Janzen & Hallwachs) (BMNH). Paratype. Costa Rica: Guanacaste Prov.: 1 9, Santa Rosa National Park, 300 m, i.1983 (Janzen & Hallwachs) (BMNH). Enicospilus ceciliae sp. n. (Fig. 344) Description. Mandibles moderately long, proximally strongly narrowed, with a basal lobe, distally more weakly tapered, apically twisted 20—30°; upper mandibular tooth compressed, 1.3-1.6 times as long as the lower tooth; outer mandibular surface centrally almost flat, sparsely pubescent, and with a strong proximal concavity. Labrum 0.2-0.3 times as long as broad; malar space 0.30.4 times as long as basal mandibular width. Clypeus in profile almost flat, margin blunt; clypeus in front view 1.3—1.4 times as broad as long, the margin apically almost truncate; lower face 0.69-0.71 times as broad as long, finely punctate. Head in dorsal view with genae evenly narrowed behind eyes; posterior ocellus very close to eye; FI = 65-70%; occipital carina mediodorsally complete, ventrally curved to almost join or join hypostomal carina about 0.7-0.8 times the basal mandibular width away from mandible. Antenna slender, with 56-59 flagellar segments; 20th segment 2.1—2.2 times as long as broad. Mesoscutum polished, with vestigial punctures, in profile fairly evenly rounded. Mesopleuron polished, the upper part punctate grading to punctostriate; lower part punctostriate with a coarser rugose-striate band of sculpture extending from epicnemial carina to lower hind corner of mesopleuron; epicnemial carina quite weak, curved towards anterior margin of pleuron. Scutellum in profile moderately convex, laterally carinate for 0.8 or more of its length; scutellum in dorsal view 1.41.5 times as long as anteriorly broad, smooth, but posteriorly with some longitudinal wrinkles. Metapleuron convex, punctate; sub- metapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile quite long and evenly declivous; anterior transverse carina complete, posterior transverse carina absent; anterior area moderately long, rugose; spiracular area quite short, smooth; posterior area reticulate; lateral longitudinal carina more or less complete, usually joined to spiracular margin by a short carina. Fore wing length 12-14 mm; discosubmarginal cell as in Fig. 344; AI = 0.95-1.10; CI = 0.18-0.29; ICI = 0.45—0.62; SDI = 1.14~1.18; cu-a from subopposite to proximal to the base of Rs&M by 0.2 times its own length; marginal cell proximally narrowly sparsely pubescent close to Rs+2r; 1st subdiscal cell broadly 296 IAN D. GAULD but rather sparsely hirsute. Hind wing with 5-6 hamuli on R1; 1st abscissa of Rs straight, 2nd abscissa almost straight. Fore leg with tibia subcylindrical, with scattered slender spines on outer surface. Mid leg with longer tibial spur 1.4-1.5 times length of the shorter. Hind leg with coxa in profile 1.8-1.9 times as long as deep; trochantellus dorsally <0.1 times as long as broad; 4th segment of tarsus 2.5—2.7 times as long a broad; claws of female rather long, distally abruptly but shortly curved, with close, short pectinae; those of male similar but with pectinae closer together. Gaster slender; tergite 2 in profile 6-7 times as long as posteriorly deep, laterotergite turned under, thyridia elliptical and separated from anterior margin of tergite by about 2-3 times its own length. Ovipositor slender, more or less straight, its sheath slender. Male with sternites 7-9 bearing scattered, long, erect slightly curved hairs; gonosquama distally evenly rounded. Colour generally pale yellowish on head and alitrunk, with gaster yellowish brown; legs and antennae golden; interocellar area yellowish; pterostigma yellowish brown; wings hyaline. VaRIATION. Enicospilus ceciliae is a morphologically and chromatically uniform species. Remarks. Enicospilus ceciliae is apparently closely related to E. dispilus from which it differs consistently in the form of the alar sclerites (cf. Figs 344, 347-352). The mandibles are slightly more slender than those of E. dispilus and most specimens of E. dispilus have dark mesoscutal markings; such vittae are never present in E. ceciliae. BIOLOGICAL INFORMATION. Enicospilus ceciliae has only been collected in north-estern Costa Rica at two sites 14 km apart. These localities, at altitudes between 300 and 400 m, support quite mature but disturbed, seasonally dry forest. Surprisingly, E. ceciliae has not been collected in Santa Rosa National Park. Nothing is known about its natural history. MATERIAL EXAMINED Holotype ©’, Costa Rica: Alajuela Prov., Rio Meno crossing, W. of Santa Cecilia, 350 m, vii.1986 (Gauld) (BMNH). Paratypes. Costa Rica: Alajuela Prov.: 2 &’, Rio Meno Xing, W. of Santa Cecilia, 350 m, vii.1986 (Gauld) (BMNH): Guanacaste Prov.: 1 2, Cerro el Hacha, 300-400 m, i-iv.1987 (Janzen & Hallwachs) (BMNH). Enicospilus dispilus (Szépligeti) (Figs 140, 143, 213, 214, 347-352) Ophion (Eniscopilus) [sic] arcuatum Felt, 1902: 307. Syntypes O’, 9, U.S.A. (Albany). [Junior primary homonym of Ophion arcuatus Brullé, 1846. ] Henicospilus dispilus Szépligeti, 1906: 145. Holotype 2, BRAZIL (TM) [examined]. Enicospilus fuscipennis flavostigma Hooker, 1912: 58. Holotype 2, FRENCH GUIANA (USNM) [exam- ined]. Syn. n. Enicospilus purgatus arcuatus (Felt) Hooker, 1912: 76. Enicospilus dispilus (Szépligeti) Hooker, 1912: 87. Enicospilus arcuatus (Felt) Wolcott, 1923: 65. Enicospilus flavostigma Hooker; Townes & Townes, 1966: 178. Description. Mandibles moderately long, generally fairly evenly narrowed, though in some larger indivi- duals they may be more or less parallel-sided distally, apically twisted 20—45°; upper mandibular tooth subcylindrical, 1.3-1.8 times as long as the lower tooth; outer mandibular surface with fine scattered pubescence, almost always centrally very slightly concave, and with at most a weak proximoventral concavity. Labrum 0.2-0.4 times as long as broad; malar space 0.2-0.3 times as long as basal mandibular width. Clypeus in profile from almost flat to weakly concave, margin blunt; clypeus in front view 1.3-1.6 times as broad as long, with margin from subtruncate to slightly convex apically. Lower face 0.65—0.80 times as broad as long, centrally usually finely and sparsely punctate. Head in dorsal view with genae generally slightly inflated, but constricted behind eyes; posterior ocellus close to but usually not touching eye; FI = 70-80%; occipital carina mediodorsally almost invariably complete, ventrally curved to join hypostomal carina about 0.40.7 times the basal mandibular width away from mandible. Antenna quite long and slender, with 55-63 flagellar segments; 20th segment 1.7—2.1 (or rarely up to 2.4) times as long as broad. Mesoscutum polished, finely punctate, in profile fairly abruptly rounded; notauli weak but discernible. Mesopleuron polished, the upper part always finely punctate, the lower part from punctate to finely punctostriate; epicnemial carina inclined towards anterior margin of pleuron. Scutellum in profile weakly OPHIONINAE OF TROPICAL MESOAMERICA 297 to moderately convex, laterally carinate for 0.5 or more (usually most) of its length; scutellum in dorsal view 1.3—1.5 times as long as anteriorly broad, generally finely punctate with isolated striae posteriorly. Metapleuron moderately convex, from regularly punctate to rugulose or even almost striate (Fig. 213); submetapleural carina usually evenly anteriorly broadened; posterior transverse carina of mesosternum usually complete, rarely centrally interrupted. Propodeum in profile abruptly declivous; anterior trans- verse carina from complete to (rarely) completely absent, posterior transverse carina usually absent, sometimes represented by a weak crest laterally; anterior area from smooth to striate, quite deep; spiracular area rather short, more or less smooth; posterior area wrinkled to reticulate, generally with one to several longitudinal rugae, very occasionally either almost smooth or concentrically rugose; lateral longitudinal carina usually virtually absent, very rarely more complete, not joined to spiracular margin by a distinct short carina. Fore wing length 13-22 mm; discosubmarginal cell as in Figs 347-352; AI = 0.75-—1.30; CI = 0.25-0.35; ICI = 0.40-0.70; SDI = 1.14~1.29; cu-a proximal to the base of Rs&M by 0.1-0.5 times its own length; marginal cell proximally usually rather sparsely hirsute, sometimes broadly glabrous; 1st subdiscal cell with anterior and distal margins narrowly to quite broadly hirsute. Hind wing with 5-8 hamuli on R1; 1st abscissa of Rs straight (Fig. 143), 2nd abscissa usually straight. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.3-1.5 times length of the shorter. Hind leg with coxa in profile 1.6-1.8 times as long as deep; trochantellus dorsally 0.1 or less times as long as broad (Fig. 140); 4th segment of tarsus 2.1—2.4 times as long as broad; claws of female long, bearing long fine pectinae, apically abruptly curved; claws of male similar but with pectinae a little shorter. Gaster long and slender; tergite 2 in profile more than 6 times as long as posteriorly deep, laterotergite folded under, thyridia oval to elliptical and separated from anterior margin of tergite by about 34 times its own length. Ovipositor slender, its sheath narrow. Male with sternites 6-9 bearing dense long stout erect pubescence (Fig. 214); gonosquama apically subacute. Colour generally brownish yellow, usually with head paler yellow; mesoscutum sometimes with dark vittae, terminal segments of gaster from brownish yellow to blackish; interocellar area yellow; antenna golden; pterostigma yellowish to brownish; wings hyaline to weakly infumate. VARIATION. This species shows exceptional variation in a range of features. The most obvious is in the form of the alar sclerites (Figs 347-352), though the proximal sclerite is always more or less regularly triangular and the distal sclerite, if present, is confluent with its distal corner. The most frequently encountered configuration is for the fenestra to be moderately long, the distal sclerite to extend around about two-thirds of its periphery, and for the central sclerite to be more or less oval to D-shaped and positioned closer to the distal margin of the fenestra than it is to the proximal sclerite (Fig. 350). The holotype of Enicospilus flavostigma has such an alar configuration. A few individuals are similar but have the distal sclerite very reduced (Fig. 351). A large number of individuals have the central sclerite strong and almost circular, and also have a strong complete distal sclerite (Fig. 352). Frequently, in such individuals, the fenestra may be exceptionally short, so the central sclerite appears to be positioned almost centrally (Fig. 349). The holotype of Enicospilus dispilus has such an alar configuration. A number of specimens have the distal sclerite obliquely D-shaped, with the long straight side directed posteromedially; other specimens have a weak crescent-shaped central sclerite (Figs 347-348). Coloration is also variable. Typically this is a rather pale yellowish species, and over much of its range it is not dark-marked. However, many specimens in tropical Central and South America have well-defined dark mesoscutal vittae, and I have seen one population (from Virgen de Socorro, Costa Rica) which have more or less the entire mesoscutum black. Many dark-marked individuals also have the posterior segments of the gaster infuscate and a few have tergites 3+ entirely black. Sculpture is also variable. Typically the posterior area of the propodeum is irregularly wrinkled to rugose, with one to several median longitudinal rugae. In some specimens the entire area is concentrically rugose, whilst in the north of its range many have an almost smooth propodeum. The anterior transverse carina is usually complete, but in some individuals from the eastern United States, and from Central America, this carina can be extremely reduced. Some such individuals also have the posterior transverse carina of the mesosternum centrally incomplete. Specimens with finer propodeal sculpture usually have the metapleuron punctate, as do many other individuals, but some have coarse rugae present anterodorsally. The general shape of the mandibles is quite uniform though those of some individuals are stout and weakly twisted apically (ca 20°), whilst others have a slender, strongly twisted mandible (ca 40—45°). A group of specimens from Guanacaste Province, Costa Rica (with a vestigial anterior transverse propodeal carina, and a concentrically rugose posterior area) are remarkable in having extremely long erect hairs on the posterior gastral sternites. Those on sternite 8 are almost as long as the sternite. 298 IAN D. GAULD REMARKS. For many years this species has been called Enicospilus arcuatus (Felt) (in, for example, Townes & Townes, 1966; Carlson, 1979) despite the fact that this name is unavailable as it is a homonym. The earliest available name I have found that definitely applies to this species is dispilus (Szépligeti, 1906). However, I have not been able to locate the type of bifoveolatus (Brullé, 1846), a North American species, and it is possible that this name may apply to this species. Enicospilus dispilus is one of the most extraordinarily variable species of the genus that I have seen. There are a large number of fairly distinctive morphotypes, but I have chosen not to regard these as separate species for the following reasons: 1) intermediates can be found between these morphotypes; 2) although at any one site one to several morphotypes may be recognizable (for convenience call them A and B), at a different locality other morphotypes may exist which combine some ‘distinctive’ features of A with B and B with A to give other (and at that site separable) morphotypes; 3) any key to the morphotypes is unlikely to be workable as collecting at new sites often yields another morphotype with another combina- tion of features; 4) every series of more than a few individuals from any one locality contains two or more morphotypes; 5) to call the several morphotypes ‘species’ obscures a real biological phenomenon which requires further study. BIOLOGICAL INFORMATION. Enicospilus dispilus is an extremely widely distributed species (Map 32) whose range extends from southern Ontario and Quebec (47°N) south to Argentina and Cautin Province, Chile (c. 39°S). In Central America, although it has never been found in large numbers at any one site, it is overall one of the most frequently collected species of Enicospilus. It has been taken at almost every site sampled from sea-level up to about 2000 m. It seems to be most frequently encountered in habitats which have suffered agricultural disturbance, and it is relatively uncommon in cloud forest sites. For example, despite intensive collecting only a few individuals have been taken in cloud forest at Monteverde, Costa Rica (4 specimens). In Santa Rosa National Park it is quite frequently collected. Most specimens have been taken at the beginning and end of the wet season, and only isolated examples have been collected in the driest months of the year. Its cumulative seasonal distribution for 1977-86 is: eee INE A NE MIR TPES NG) SINGS LE eS AS) ES 2m AO In Costa Rica E. dispilus has been collected in the following localities: Finca Campana at 5 km NW. of Dos Rios, Finca San Gabriel at 16 km ENE. Quebrada Grande, San Ramon Forest Reserve and Virgen de Socorro, in Alajuela Province; Turrialba in Cartago Province; Finca Biesnan at 11 km E. Quebrada Grande, Rinc6én National Park, Santa Rosa National Park and Casa Mariksa (= Maritza) on the lower slopes of Volcan Orosi in Guanacaste Province; Monteverde in Puntarenas Province; Braulio Carrillo National Park in San José Province. On Barro Colorado Island, Panama relatively few individuals have been collected, but the cumulative data for 1978-85 are: 7c Mer AveMen) daesAy (SytO' Ne tbD =i otter 22 ncowSs2). whe @8rine= mee wlth Enicospilus dispilus is an endoparasitoid of the larvae of a variety of species of Noctuidae and Notodon- tidae that eat the leaves of large trees. In Costa Rica I have seen one female of this species reared in Santa Rosa National Park from the larva of a noctuid (rearing reference number 84-SRNP-1149) found feeding on oak (Quercus oleoides). The host larva began to spin up on the 15th June and the parasitoid emerged less than one month later on 18th July. A second female was reared in the same locality from an unidentified species of notodontid found feeding on Chlorophora (87-SRNP-901). In North America, judging from the incidence of reared individuals in museum collections, E. dispilus is a fairly common larval endoparasitoid of the notodontids Heterocampa guttivitta (Walker) and Schizura concinna (Smith). I have also seen individuals reared from Hippia packardi Morrison, Nadata gibbosa (Smith) (in USNM) and Heterocampa mantea (in CNC) (Lepidoptera: Notodontidae) and a single specimen allegedly reared from Cirphis unipuncta (Haworth) (Lepidoptera: Noctuidae). E. dispilus has also been recorded as a parasitoid of the noctuid Scoliopteryx libatrix (L.) (Carlson, 1979), and in southern South America it is recorded as a parasitoid of Alabama argillacea (Hiibner) (Lepidoptera: Noctuidae) (DeSantis, 1941; DeSantis & Esquivel, 1966). MATERIAL EXAMINED Holotype 9 (Henicospilus dispilus Szépligeti), Brazil: Minas Gerais, 1897 (TM). Holotype 9 (Enicospilus fuscipennis flavostigma Hooker), French Guiana: Cayenne (USNM). 370 CO, 486 9, from the following localities: Argentina (Misiones, Tucuman); Bahamas (Grand Bahama); Belize; Bolivia (Chapare, Cochabamba, Santa Cruz); Brazil (Bahia, Mato Grosso, Rio de OPHIONINAE OF TROPICAL MESOAMERICA 299 Map 32 Localities at which Enicospilus dispilus has been collected. Janeiro, Santa Catarina); Canada (British Columbia, Ontario, Quebec); Chile (Cautin); Colombia (Valle); Costa Rica (Alajuela, Cartago, Guanacaste, Puntarenas, San José); Dominican Republic; Ecuador (Pichincha); El Salvador; Guatemala; Guyana; Mexico (Chiapas, Durango, Guerrero, Jalisco, Mexico State, Neuvo Le6n, Quintana Roo, San Luis Potosi, Sonora, Veracruz); Panama; Peru (Huanuco, Lima, Loreto); Surinam; U.S.A. (Arizona, Arkansas, California, Colorado, Connecticut, Delaware, District of Columbia, Florida, Georgia, Illinois, Iowa, Kentucky, Louisiana, Maine, Maryland, Mas- sachusetts, Michigan, Mississippi, Missouri, Nevada, New Hampshire, New Mexico, New York, North Carolina, Ohio, Pennsylvania, Rhode Island, South Carolina, Tennessee, Texas, Virginia). Venezuela (Aragua). (In BMNH, CAS, CNC, FSCA, TC, UMC, USNM.) 300 IAN D. GAULD Enicospilus echeverri sp. n. (Figs 319, 346) Description. Mandibles moderately long, fairly evenly tapered, apically twisted 35—-45°; upper mandibular tooth subcylindrical to slightly depressed, 1.4-1.6 times as long as the lower tooth; outer mandibular surface bearing isolated pubescence, distally flat, proximally with a weak shallow concavity. Labrum 0.2— 0.3 times as long as broad; malar space 0.2-0.3 times as long as basal mandibular width. Clypeus in profile very weakly convex, margin blunt to subacute; clypeus in front view 1.3—-1.5 times as broad as long, with margin subtruncate apically. Lower face 0.71—0.76 times as broad as long, centrally sparsely punctate, sometimes with punctures elongate and tending to punctostriate. Head in dorsal view with genae rounded behind eyes; posterior ocellus close to eye; FI = 70-75%; occipital carina mediodorsally complete, ventrally curved to join hypostomal carina about 0.6—0.9 times the basal mandibular width away from mandible. Antenna long and slender, with 53—56 flagellar segments; 20th segment 2.0—2.2 times as long as broad. Mesoscutum polished, finely punctate, in profile abruptly rounded; notauli vestigial. Mesopleuron polished, the upper part rather sparsely punctate, the lower part punctate, occasionally tending to punctostriate; epicnemial carina curved towards but not reaching anterior margin of pleuron. Scutellum in profile weakly convex, laterally carinate for 0.9 or more of its length; scutellum in dorsal view 1.5—1.6 times as long as anteriorly broad, punctate and usually with some longitudinal striae posteriorly. Metapleuron weakly to moderately convex, punctate, occasionally with isolated striae posterodorsally; submetapleural carina evenly anteriorly broadened; posterior transverse carina of mesosternum complete. Propodeum in profile fairly abruptly declivous; anterior transverse carina complete, usually very strong laterally, pos- terior transverse carina usually represented by vestigial lateral crests; anterior area steep, centrally striate; spiracular area long, smooth or very finely punctate; posterior area reticulate; lateral longitudinal carina present anteriorly, sometimes complete, joined to spiracular margin by a short carina. Fore wing length 15-17 mm; discosubmarginal cell as in Figs 319, 346; AI = 1.04-1.25; CI = 0.25-0.36; ICI = 0.39-0.43; SDI = 1.25-1.41; cu-a usually proximal to the base of Rs&M by its own thickness, rarely by up to 0.2 times its own length; marginal cell proximally slightly more sparsely hirsute than it is centrally; 1st subdiscal cell usually rather sparsely hirsute. Hind wing with 6-7 hamuli on R1; 1st abscissa of Rs very slightly bowed, 2nd abscissa almost straight. Fore leg with tibia slightly flattened, with scattered spines on outer surface. Mid leg with longer tibial spur 1.5—1.7 times length of the shorter. Hind leg with coxa in profile 1.7-1.9 times as long as deep; trochantellus dorsally 0.1 times as long as broad; 4th segment of tarsus 2.0—2.2 times as long as broad; claws of female quite long, bearing fine pectinae, apically evenly curved; claws of male similar but more finely pectinate. Gaster long and slender; tergite 2 in profile more than 6.0 times as long as posteriorly deep, laterotergite folded under, thyridia elliptical and separated from anterior margin of tergite by about 3-4 times its own length. Ovipositor slender, its sheath narrow. Male with sternite 7 and extreme anterior part of sternite 8 bearing two rows on short, stout closely packed hairs; remainder of sternite 8 and sternite 9 with fine decumbent pubescence; gonosquama distally rounded. Colour generally reddish brown, head and usually also the scutellum paler yellow, mesoscutum with three dark longitudinal vittae, gaster with tergites 3+ infuscate; interocellar area yellowish; antenna brownish to almost black; pterostigma brownish; wings weakly infuscate. VARIATION. Specimens from Brazil tend to have the alitrunk more brightly yellowish than Central Amer- ican specimens. REMARKS. This species is named in honour of Gustavo Echeverri for his contributions to the Guanacaste National Park, Costa Rica. Enicospilus echeverri is apparently very closely related to E. lacsa. Both species are similar in general appearance and have identical and highly modified male terminal gastral sternites (Fig. 200). E. echeverri can most easily be distinguished from E. lacsa by the distally more acute central sclerite (cf. Figs 345, 346), its stouter flagellar segments, coloration and rather broader face. BIOLOGICAL INFORMATION. Enicospilus echeverri is a fairly widely distributed species whose range extends from about 19°N in southern Mexico, south to Rio de Janeiro, Brazil (23°S). It has been collected from near sea-level up to about 1400 m. In Santa Rosa National Park only isolated specimens have been collected between March and August. Nothing is known of its host range. MATERIAL EXAMINED Holotype 9, Costa Rica: Guanacaste Prov., Santa Rosa National Park, 300 m, vi.1984 (Janzen & Hallwachs) (BMNB). OPHIONINAE OF TROPICAL MESOAMERICA 301 Paratypes. Brazil: 1 0’, 15 2, Bahia, Encruzilhada, 960-980 m, xi.1972—4 (Alvarenga) (TC); 1 2, Minas Gerais, Pedra Azul, 800 m, xi.1972 (Alvarenga & Seabra) (TC); 1 2, Rio de Janeiro, Itatiaia National Park, x.1969 (Otero) (TC). Costa Rica: Guanacaste Province: 1 2, Santa Rosa National Park, 300 m, viii.1977 (Janzen) (TC); same locality, 1 9, vi.1982, 1 Q, iii.1983 (Janzen & Hallwachs) (BMNH); 1 @, Volcan Cacao, Finca La Luz [= Estacion Mengo], 1100 m, viii. 1986 (Janzen & Hallwachs) (BMNH); 1 @, same locality, vi.1987 (Janzen) (BMNH): Puntarenas Prov.: 1 9, Monteverde, 1300-1400 m, vii.1982 (Janzen & Hallwachs) (BMNH). Mexico: Chiapas: 1 2 , 32 km N Huixtla, 1000 m, vi.1969 (CNC): Oaxaca: 1 9,19 km S. Valle Nacional, 1000 m, v.1971 (Howden) (TC): Veracruz: 1 2, Cordoba, v.1906 (Knab) (USNM). Panama: 1 ©’, 2 9, Barro Colorado Island, 120 m, vi-vii. 1984-5 (Wolda) (BMNH). Acknowledgements Iam grateful to the Director of the National Parks Services of Costa Rica, and to his staff, for allowing me to work in Santa Rosa National Park. This work would not have been accomplished had it not been for the encouragement and enthusiastic support that I received in the field from Dan Janzen and Winnie Hallwachs. Dr Janzen additionally supplied a great deal of information concerning hosts and allowed me to use his unpublished rearing records; he was also responsible for suggesting the majority of dedications for species names. I thank the following people for collecting specimens for me: Henk Wolda, Bill Haber, Abelardo Chacon, Isidro Chacon, Mike Fogden, Tricia Fogden and Joe Eger. Roberto and Alvaro Espinosa have helped by rearing material and servicing Malaise traps. I am particularly grateful to Henry and Marjorie Townes, Lubomir Masner, Mike Sharkey and David Wahl for their hospitality on a recent trip to the U.S.A., and I thank them and all the curators who lent me valuable material from their collections. Many other people provided help in a variety of ways; their assistance has been acknowledged by naming species after them. I am especially grateful to Sondra Ward for all her help in organizing work for me whilst I was temporarily incapacitated, and for the care with which she prepared the stereoscan photographs. I also thank the staff of the photographic unit and stereoscan unit of the British Museum (Natural History). My colleagues Mike Fitton, Mick Day and Mark Shaw have, over the years, offered many insights into hymenopteran biology. My wife, Pam Mitchell, helped as always in innumerable ways. References Agassiz, J. L. R. 1846. Nomenclator zoologicus, Index Universalis 1135 pp. Soloduri. Ashmead, W. H. 1890. Descriptions of new Ichneumonidae in the collection of the U.S. National Museum. Proceedings of the United States National Museum 12: 387-451. 1894. Some parasitic Hymenoptera from Lower California, Mexico. Proceedings of the California Academy of Sciences (2) 4: 122-129. 1895. Some parasitic Hymenoptera from Baja California and Tepic, Mexico. 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Actias luna 167 Agapema galbina 152 Agrotis ipsilon 38 Alabama argillacea 178, 298 Amyna octo 192 Antheraea polyphemus 167, 168 Arctiidae 13, 130 Automeris io 167, 168 Automeris tridens 13, 163 Automeris rubrescens 163 Callosamia promethea 167, 168 Callosamia securifera 167, 168 Ceratocampinae 12, 61 Cirphis unipuncta 298 Composia sp. 130 Copaxa moinieri 13, 124 Dasychira basiflava 130 Diacrisia virginica 130 Eulepidotis sp. 277, 282 Faronta albilinea 178 Gonodonta nutrix 199 Gondonta sp. 235 Helicoverpa zea 178 Heliothis zea 38 Hemerocampa leucostigma 130 Index to hosts Hemerocampa sp. 130 Hemileuca juno 152 Hemileuca magnifica 152 Hemileuca maia 152 Hemileuca oliviae 152 Hemileuca peigleri 152 Hemileuca tricolor 152 Heterocampa mantea 298 Heterocampa guttivitta 298 Hippia packardi 298 Hyalophora cecropia 167 Hyalophora euryalis 167 Hylesia lineata 13, 139 Hylesia sp. 136 Hypercompe alibicornis 130 Hypercombe icasia 130 Hypercompe suffusa 130 Hyphantria cunea 130 Lasiocampidae 142 Lophocampa maculata 130 Lymanitriidae 13, 130, 154 Malacosoma americana 154 Malacosoma disstria 154 Mocis repanda 38 Mythimna unipuncta 178 Nadata gibbosa 298 Noctuidae 13, 15, 38, 178, 209, 298 Notodontidae 15, 178, 254, 298 Index OPHIONINAE OF TROPICAL MESOAMERICA Orgyia sp. 130 Othorene purpurascens 61 Pachylia ficus 59 Peridromia saucia 39, 178 Perigonia lusca 60 Pseudaletia unipuncta 39 Ptiloscola dargei 60 Rothschildia arethusa 168 Rothschildia erycina 13 Rothschildia lebeau 12, 13, 161 Rothschildia lebeau forbesi 161 Rothschildia maurus 168 Rothschildia orizaba 167 Samia cynthia 167 307 Saturniidae 12, 13, 146, 152, 154, 167, 178 Schizura concinna 298 Scoliopteryx libatrix 298 Sphingidae 12, 59, 60, 178 Spodoptera eridania 39 Spodoptera frugiperda 38, 178 Spodoptera ornithogalli 178 Syssphinx molina 61 Tolype pauperata 168 Xylophanes anubis 63 Xylophanes turbata 60 Invalid names are in italics; principal references are in bold. abelardoi 22, 79 145, 157 affinis 55 Agathophiona 9, 14, 27, 48 aktites 15, 81, 147, 155, 156 Alophophion 14, 15 alvaroi 22, 77, 131 americanus 12, 13, 14, 15, 20, 21, 75, 83, 146, 161, 163, 164, 166, 169, 170, 172 ancyloneura 29, 37 angulatus 129 antefurcalis 178 antomelas 233 appendiculatum 189 apricus 18, 55, 56, 61 arctiae 128 arcuatus 296 arida 52 arribai 18, 22, 33, 45 Athyreodon 9, 54, 55 atricolor 55, 61 atrivensis 12, 15, 55, 56 attritus 272 Aulophion 9, 72 baltodanorum 22, 87, 201 barbarae 95, 248 biangularis 37 bicarinatus 18, 72, 74 bicolor (Enicospilus) 74, 216 bicolor (Thyreodon) 55 bima 95, 214, 247 bozai 13, 18, 77, 123, 125, 132 brasiliensis 189 brenesiae 22, 97, 265 brevinervis 190 brevis 79, 136 brullei 189 burgosi 99, 102, 266, 267 cacaoi 22, 31, 40, 42 calcator 134 calliope 22, 31, 34, 35 cameronii 81, 143, 146, 147, 151, 167 capensis 178 carlota 85, 185, 191, 195 carri 89, 202 catemacoi 18, 95, 251, 253 ceciliae 22, 102, 295 cecropiae 166 cepillo 93, 225, 226 chaconi 15, 20, 22, 77, 125 chiriquensis 79, 123, 133, 134 clarkorum 77, 126, 128 clio 18, 22, 31, 39 colini 89, 202, 203 columbianus 15, 75, 85, 180 concolor (Enicospilus) 230 concolor (Ophion) 37 corcovadoi 18, 91, 220 cornifuscus 95, 250 cressoni 18, 87, 89, 196, 198 cubensis 15, 85, 91, 183 cubitalis 128 curvinervis 147 cushmani 81, 153 dajaboni 85, 186, 191, 195 devriesi 93, 239, 243, 273 dichromus 216 dimidiator 126 dirzoi 18, 87, 191, 194 308 dispilus 15, 22, 75, 101, 102, 287, 288, 289, 296 diversus 37 donahuei 18, 93, 242 doylei 83, 173, 178 druryi 166 duckworthi 93, 227, 238 echeverri 97, 102, 263, 300 elegans (Enicospilus) 233 elegans (Thyreodon) 55 elongatus 18, 29 Enicospilus 9, 10, 11, 14, 15, 18, 20, 22, 23, 27, 29, 49, 55, 74 enigmus 77, 123, 124, 125 erasi 18, 99, 267, 269, 272 erato 18, 33, 43, 44 Eremotylus 10, 14, 27, 52 erythrocera 55, 56, 62 estradarum 22, 89, 202, 213 euterpe 18, 31, 34, 35 excarinatus 72 excubitalis 129 exoticus 15, 77, 87, 91, 125, 126, 215, 216 fenestralis 18, 20, 51, 240 fernaldi (Enicospilus) 15, 93, 220, 228, 231 fernaldi (Thyreodon) 55 ferrugineus 18, 55, 62 flammiger 55 flammipennis 11, 53, 54 flavarius 230 flaviceps 175 flavidus 11, 13, 15, 29, 31, 37 flavoorbitalis 18, 29, 34, 47 flavoscutellatus 15, 20, 99, 179, 206, 240, 272 flavosignatus 237 flavostigma 296 flavus 15, 93, 102, 230 fogdenorum 22, 99, 266 forsythei 18, 99, 262, 265 fosteri 18, 95, 243, 249, 254 fulvescens 55 fulvicornis 18, 49 fuscicornis 250 fuscipennis 233 gabrieli 22, 95, 252 galilea 99, 267, 270, 272 gallegosi 101, 285, 295 gamezi 22, 81, 150 georginae 18, 22, 102, 287, 288, 291 glabratus 12, 15, 75, 77, 128, 167 gomezpompai 18, 79, 81, 157 gonzalezi 22, 66, 68, 70 grandis 55 grenadensis 56 guatemalensis 15, 89, 206 guindoni 95, 256, 258 guyanensis 230 haberi 22, 97, 261 hacha 91, 223 IAN D. GAULD halffteri 18, 81, 152, 154, 157 hallwachsae 79, 81, 146, 157 hemicrescellae 22, 102, 294 hookeri 191 howdenorum 18, 101, 282 hubbelli 18, 99, 267, 270, 271 Janzophion 14, 22, 27, 49 jesicae 91, 99, 215, 217, 218 karrensis 18, 89, 209 kelloggae 95, 97, 201, 259, 263 kleini 102, 238, 293 lacsa 97, 102, 260, 262, 300 lateralis 175 laticinctus 12, 15, 55, 56, 62 latilineatus 147 latus, 15, 20, 64 laurenae 15, 87, 199, 201 lebophagus 11, 12, 13, 18, 81, 160, 167 lemairei 22, 66, 70, 71 leoni 97, 257 liesneri 15, 101, 221, 276 lovejoyi 91, 221, 223, 277 luisi 95, 250, 253, 255, 260 lupemejia 85, 188, 191 luquillo 18, 83, 179 maculiceps 189 maculipennis (Enicospilus 89, 205 maculipennis (Thyreodon) 12, 18, 55, 56, 59, 60, 61 maculithorax 15, 66, 68, 69 madrigalae 93, 227, 236 major 77, 126, 127 marini 101, 221, 278, 280 maritzai 91, 215, 217, 219 martae 18, 85, 180, 181 masneri 87, 195 masoni 18, 97, 102, 292 mayi 18, 79, 132, 134 melanostigma 18, 29, 65 melpomene 22, 33, 44 mengoi 22, 97, 260 mexicanus 15, 20, 79, 142, 148, 167 monticola 15, 20, 93, 97, 233, 258 morosus 15, 55, 56, 63 nebosus 18, 49, 51, 52, 240 neotropicus 83, 172, 178 niger 18, 55 nigricauda 189 nigrifrons 68 nigriventris 68 oduberi 22, 101, 284 Ophiogastrella 9, 10, 11, 14, 18, 29, 65, 66 Ophion 9, 10, 11, 12, 14, 15, 18, 20; 23, 27, 29; 52,55; 240 opleri 15, 95, 254 ornatipennis 55 ornatus 57 orosii 15, 93, 246 pamelae 18, 89, 211 parkeri 18, 102, 287, 291 parvifasciatus 205 peigleri 83, 170 persimilis 93, 227 pescadori 101, 221, 280, 281 pleuralis 51 politior 37 polyhymniae 22, 31, 36, 45 porteri 22, 85, 187, 191 Prethophion 14, 27, 64 principalis 62 pulchricornis 63 purgatus 13, 15, 21, 75, 83, 175 quintanai 83, 165 ramidulus 75, 178, 191 randalli 87, 89, 196, 199 renovatus 148 Rhynchophion 9, 14, 21, 23, 27, 53 rivinae 12, 18, 55, 56, 60, 61 robertoi 13, 79, 137, 140 robur 18, 55 rufothorax 56 rugosus 166 sanchezi 101, 279, 281, 282 santarosae 12, 22, 55, 56, 61 sarukhani 18, 81, 155, 157 saxis 18, 49, 50 scuintlei 79, 137, 139, 140 selenaction 63 Sicophion 10, 14, 18, 22, 27, 51 simoni 93, 243 Simophion 14, 29, 65 sondrae 18, 101, 283 species 1 22, 56, 60 sphacelatus 189 Stauropoctonus 10, 14, 18, 21, 27, 72 stevensi 89, 203, 210, 212 stilesi 22, 66, 70, 71 striatus 189 subfuliginosus 18, 52 tenuigena 15, 20, 83, 167, 168 teodorae 91, 214, 248 terpsichore 22, 33, 44 texanus 13, 18, 81, 151, 154, 167 thaliae 22, 34, 42, 46 thoracicus (Enicospilus) 272 thoracicus (Thyreodon) 56 Thyreodon 9, 10, 11, 12, 14, 15, 18, 23, 27, 54 trilineatus 11, 15, 75, 85, 185, 186, 187, 188, 189, 194, 195, 276 trimaculatus 272 trispilus 272 Trophophion 14, 15 tropicus 18, 52 ugaldei 13, 18, 83, 161 ulfstrandi 15, 102, 288 ultor 18, 55 OPHIONINAE OF TROPICAL MESOAMERICA 309 umanai 83, 164 uraniae 18, 22, 33, 40, 41, 46, 47 vecors 189 vegai 87, 200 venezuelanus 83, 169 vidor 178 vilmari 22, 89, 212 volubilis 175 woldai 99, 275 xanthocarpus 15, 20, 93, 237 xanthostigma 91, 225, 226 zonatus 63 agile ti >. ie We, oy = re oe 1 ge’ me of. en F eeacepien®. 111, in, (4-19 A, ai > Re os in 438 rug? ve. | se iat wrest 1F ai i ay 18 7 a! it wttiorabae (i, Ma : te ane“. olathe pas 4 > ee jrope te, we, 39, a] Ve = i » ia 1e ad? —_ 7e =) a >= ve e »- << @ ‘-- - _ 7 ad “. a ‘ = > ) Li? = | “si. Paps rand © AGL 41; Ly British Museum (Natural History) Milkweed butterflies: their cladistics and biology P. R. Ackery & R. I. Vane-Wright The Danainae, a subfamily of the Nymphalidae, contains only some 150 species, yet aspects of their biology have stimulated far more attention than can be justified by species numbers alone. In recent years, an expansive literature has grown, considering aspects of their courtship and pre-courtship behaviour, migration, larval hostplant associations, mimicry and genetics. The popularity of danaines among biologists can certainly be attributed to this combination, within one small group, of so many of the factors that make butterflies such an interesting group to study. The obvious need to place this wealth of biological data within an acceptable systematic framework provided the impetus for this volume. Started in 1976 within the conventions of evolution by natural selection and Hennig’s phylogenetic systematics, the book is now largely about natural history (what the animals have and do, where they live and how they develop) and natural groups — as revealed by a form of analysis approaching that practised by the new school of ‘transformed cladistics’. The authors have prepared a handbook that will appeal to a wide range of biologists, from museum taxonomists to field ecologists. 425 pp, 12 pp colour, 73 b/w plates, line and graphic illustrations, maps, extensive bibliography. ISBN 0 565 00893 5. 1984. Price £50. Titles to be published in Volume 57 A survey of the Ophioninae (Hymenoptera: Ichneumonidae) of tropical Mesoamerica with special reference to the fauna of Costa Rica By Ian D. Gauld A taxonomic revision of Alabagrus (Hymenoptera: Braconidae) By Michael J. Sharkey A taxonomic revision of Caryocolum (Lepidoptera: Gelechiidae) By Peter Huemer Photoset by Computerised Typesetting Services Ltd, Redhill, Surrey Printed in Great Britain by Henry Ling Ltd, Dorchester