Chrysididae
CUCKOO WASPS
Chrysidid wasps
Life   Insecta   Hymenoptera   Chrysidoidea

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Pseudospinolia
Native Bee Inventory and Monitoring Lab; Photographer: Josh · 12
Pseudospinolia

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Pseudospinolia
Native Bee Inventory and Monitoring Lab; Photographer: Josh · 12
Pseudospinolia
Pseudospinolia, tail
Native Bee Inventory and Monitoring Lab; Photographer: Josh · 12
Pseudospinolia, tail

Chrysura cobaltina
© Copyright source/photographer · 10
Chrysura cobaltina
Chrysis angustula
D. Morgan, 1984 · 9
Chrysis angustula

Chrysis ignata
D. Morgan, 1984 · 9
Chrysis ignata
Chrysis ignita
L. S. Kimsey and R. M. Bohart, 1990 · 9
Chrysis ignita

Chrysis florissanticola
Native Bee Inventory and Monitoring Lab; Photographer: Erika Tucker · 9
Chrysis florissanticola
Chrysis florissanticola
Native Bee Inventory and Monitoring Lab; Photographer: Erika Tucker · 9
Chrysis florissanticola

Chrysis florissanticola
Native Bee Inventory and Monitoring Lab; Photographer: Erika Tucker · 9
Chrysis florissanticola
Chrysis florissanticola
Native Bee Inventory and Monitoring Lab; Photographer: Erika Tucker · 9
Chrysis florissanticola

Chrysis archboldi
R. M. Bohart and L. S. Kimsey, 1982 · 9
Chrysis archboldi
Chrysis equidens
R. M. Bohart and L. S. Kimsey, 1982 · 9
Chrysis equidens

Chrysis florissanticola
R. M. Bohart and L. S. Kimsey, 1982 · 9
Chrysis florissanticola
Chrysis nisseri
R. M. Bohart and L. S. Kimsey, 1982 · 9
Chrysis nisseri

cuckoo wasp
© Copyright Tom Crider, 2009 · 0
cuckoo wasp
Overview
Taken from: R. M. Bohart and L. S. Kimsey. 1982. A Synopsis of the Chrysididae in America North of Mexico.
INTRODUCTION
The beautiful coloration of chrysidids has been a source of admiration for all who have observed them. In addition to the metallic blues, greens and purples, many species exhibit magenta, violet, copper, gold, and bright red., Sometimes, most of these colors may be present on a single individual. Furthermore, the impact of the above interference colors is intensified by the dramatic sculpture common in the family: polished integument, coarse punctation, and sharp carinae. Some species have dense silvery pubescence, particularly on the face, and a few genera, such as Argochrysis, Neochrysis and Parnopes, have white ornamentation. It might be thought that in consideration of their spectacular appearance that the gold wasps, ruby wasps, or cuckoo wasps, as they are commonly called, would be a favorite for taxonomic study. This has not been the case and four possible reasons are: (1) difficulty of obtaining material; (2) intraspecific variation, which may result from host influence; (3) the fact that female chrysidids tend to be structurally conservative, as in many other parasite groups; and (4) the unfortunate circumstance that most early descriptions were based on color, a rather unreliable character. An estimate of the number of described species in the Chrysididae, with allowance for probable synonymy is 3, 000. Perhaps an equal number remain to be discovered. We recognize 227 species in America north of Mexico, and this includes 37 described as new. In making this study we have had about 50, 000 specimens available to us. More of these were females than males, perhaps because collectors more often see females as they are searching for a host nest. In support of this idea is the observation that about equal numbers of the sexes show up in Malaise traps. Holotype depositions are indicated in the descriptions and paratypes will be distributed to cooperating institutions as far as possible. In the interest of brevity, collector’s names for paratypes and other material, as well as specific collection dates, are not usually given, except for holotypes.
Taken from: L.S. Kimsey and R.M. Bohart, 1990. The Chrysidid Wasps of the World.


INTRODUCTION

Gold wasps or cuckoo wasps are common names often applied to the family Chrysididae. Their frequently metallic coloration justifies the former, and cleptoparasitic habits refer to the latter. This brilliant aspect is best observed on a sunny day as the female wasp is searching a nesting site of a favoured wasp host. The sunlight reflects from the parasite and seems to accentuate the metallic colours-blue, green, purple, red, copper, brass, and gold-in various combinations. Most chrysidids are small and, although not all are brilliant, the tiny ones often make up for in colour what they lack in size. The sight of a Hedychridium moving erratically over the sand like a drop of pure gold never fails to astonish the collector.

In the Western Hemisphere most chrysidids are metallic blue, green, and purple, in various combinations. In the old world, especially in the southern USSR and Africa, these wasps are often more colourful. A single specimen may be green, blue, purple, copper, gold, and red; all somewhat iridescent. These are interference colours. The true pigment shades of red, brown, and white are not so common. In addition to the often bright integument are the myriads of tiny impressions or punctures which are nearly always present. These modify and often enhance the coloration.

Some 3000 apparently valid species of Chrysididae have been named. These are now arranged in 84 genera and 4 subfamilies. In all probability a thousand or more species remain to be found.

Chrysidids are usually considered by collectors to be rarely encountered; yet they can be abundant. General net-sweeping of grass and low shrubs, or fields of flowers such as Eriogonum (wild buckwheat), will often yield dozens of specimens, mostly small ones. Trapping has been quite successful in recent years. Stick traps, such as those used by Krombein (1967) and Parker and Bohart (1966, 1968), have produced many chrysidids, as well as information on their hosts. Flight traps have been used successfully in the past few decades, and are particularly effective in catching males.

Chrysidids are distributed world-wide, but southern Asia, Africa, and the Middle East have not been fully explored. Only two world revisions have been attempted previously. The earlier authors were Dahlbom (1854) and Mocsáry (1889). In retrospect these valuable publications are seen to be quite incomplete. In particular, they suffer from the absence of a clear generic concept. A more recent work (Linsenmaier 1959 a, b), treating the European fauna, was more positive in this respect but still quite conservative. None of these three workers treated the Amiseginae or Loboscelidiinae. Even now, these subfamilies of small, obscure wasps need much more work by systematists.

Despite the attractive nature of these wasps, there has been little major revisionary work. Most studies have concentrated on faunas of restricted geographic regions, rather than revisions of specific taxa. As a result, many of the characteristics used to define genera, tribes, and subfamilies are only reliable at a regional level. There are also serious problems with homonymy and synonymy in this family, particularly for the Western Palaearctic fauna. Furthermore, no previous study has ever been made of phylogeneric relationships within the Chrysididae.

We have attempted to reorganize the higher classification of the Chrysididae from the ground up, by studying all the generotype species and as many other species as we could locate. Genetic groupings were evaluated and retained if we found them to be discrete, definable units. Unfortunately, several major groups, including the subfamily Cleptinae and the genus Chrysis, may be paraphyletic. We have found few derived characters that define these groups. However, little can be gained by merging them with other taxa; Cleptinae is clearly the sister-group of the rest of the Chrysididae, and Chrysis already accounts for half of the species in the family. Further intensive study of these groups may provide useful derived characteristics. Clearly, chrysidids have nor always evolved in the neat, tidy units that taxonomists prefer to study.

Our re-evaluation of this family has led to an elimination of subgenera and considerable changes in the generic, tribal, and subfamilial classification. We have also attempted to provide as much information about these taxa as possible. Our goals are:

(1) to provide a world-wide overview of the family, with a reclassification of the generic and higher taxa;

(2) summarize previously published information;

(3) indicate problem areas in need of further study; and

(4) give a compilation of detailed synonymic species lists for each genus.

The family Chrysididae is considered co be part of the Aculeata, or stinging wasps and bees. However, chrysidids have a highly reduced sting and the terminal abdominal segments are invaginated. Thus, an external appearance of only 2 to 4, or 5, abdominal segments is created. This situation is unique in the Aculeata. The actual number of external segments is a characteristic of subfamilies and tribes within the Chrysididae.

Synonymic species lists are an important part of any generic revision. Without them the revision has limited value. We have spent much time and effort in making the lists as complete and accurate as possible. However, we are the first to admit that further study based on original types, which we have been unable to see, and on more material, to give a better idea of intraspecific variation, will refine and alter the lists. Our treatment of subspecies names may be controversial. Since most such names have been based on minor colour variations or inadequate study of geographical variation, we have elected not to rule on the validity of most of the names. Hence, they are simply listed under the recognized species.

We hope that this study will provide the groundwork and inspiration for more detailed revision. There is doubtless considerable undetected synonymy remaining in the Chrysididae, and many new species to be described.

Kinds
Taken from: R. M. Bohart and L. S. Kimsey. 1982. A Synopsis of the Chrysididae in America North of Mexico.

FAMILY CHRYSIDIDAE Our concept of the family is in agreement with Krombein (1970) in considering Cleptes, Amisega, Elampus, Chrysis, and Parnopes as representatives of subfamilies. There seems little basis for separating Cleptes related genera as a family, especially since the amisegines are an annectant group. All of the five subfamilies indicated above occur in America north of Mexico but the great majority of species are in the Elampinae and Chrysidinae.
Key to subfamilies of Chrysididae 1. Abdominal venter convex, four or more exposed terga …..2 Abdominal venter flat to concave, three well developed exposed terga (except Parnopinae males)…..3 2. T-I-II dorsally subequal to or shorter than T-III-IV, upper frons with groove descending from midocellus, ovipositor long and robust, tenthredinoid parasites ……Cleptinae p. 12 T-I-II dorsally much longer than length of T-III-IV, upper frons without median groove, ovipositor slender and needlelike, phasmatid egg parasites….. Amiseginae p. 17 3. Tongue long, exserted, reaching at least midcoxae; apical tergum with two subapical foveae and apical margin denticulate; male with four well developed external abdominal segments, females with three ……Parnopinae p.20l Tongue short, folded beneath head; apical tergum simple or with subapical pit row and/or two to six teeth or lobes; both sexes with three well developed external abdominal segments…..4 4. Forewing with ES stub sclerotized for less than half marginal cell length (if vein is extended by its traces and wing crease to wing margin); discoidal and cubital cell veins mostly lacking; T-III without subapical pit row or groove, never more than two apical teeth …..Elampinae p.21 Forewing with ES stub sclerotized for more than half marginal cell length (if vein is extended by its traces and wing crease to wing margin); discoidal and cubital cell veins usually sclerotized and complete; T-III with subapical pit row or groove (except a few Neochrysis which have four teeth on T-III)….. Chrysidinae p. 86

Identification
Taken from: R. M. Bohart and L. S. Kimsey. 1982. A Synopsis of the Chrysididae in America North of Mexico.

CHARACTER STATES: 1. Primitive: antenna with 13 articles; derived: antenna with fewer than 13 articles. 2. Primitive: tongue structure lying flat in oral fossa; derived: tongue with a basal angle or fold, protruding from fossa. 3. Primitive: cardo a small, narrow strip alongside stipes or absent; derived: cardo an elongate rod or plate, frequently almost as long as stipes. 4. Primitive: maxillary palpus 6-segmented, derived: (a) 5- segmented, or (b) 1-segmented. 5. Primitive: labial palpus 4-segmented; derived: palpus (a) 3- segmented, or (b) absent. 6. Primitive: prothorax hinged to mesothorax, freely moveable; derived: prothorax fused or at least closely articulated to mesothorax so that it is no longer readily moveable. 7. Primitive: tarsal claws with a single subsidiary tooth; derived: claws (a) with more than one tooth, or (b) edentate. 8. Primitive: forewing with complete discoidal cell; derived: discoidal cell incomplete. 9. Primitive: forewing marginal cell practically closed, or RS extending at least half marginal cell length (determined by following wing crease and vein remnants); derived: RS stub less than half as long as marginal cell. 10. Primitive: fore wing with a stigma; derived: stigma absent. 11. Primitive tegula covering brewing base; derived: tegula covering bases of both forewing and hindwing. 12. Primitive: metanotum laterally evenly rounded; derived: metanotum with lateral angle or tooth adjacent to propodeal tooth. 13. Primitive: propodeum dorsally elongate and box shaped; derived: propodeum not dorsally elongate, subtriangular. 14. Primitive: number of well formed external terga sexually dimorphic; derived: number of terga equal in the sexes. 15. Primitive: females with six visible and well developed terga; derived: (a) four terga, (b) three terga, (c) two terga. 16. Primitive: males with seven well formed external terga; derived: (a) five terga, (b) four terga, (c) three terga, (d) two terga. 17. Primitive: abdominal sterna convex; derived: sterna flat or concave. 18. Primitive: females with true sting; derived: sting reduced to lancets and sheaths in an ovipositor tube. 19. Primitive: ovipositor tube large and robust, consisting of four to six pairs of overlapping plates; derived: ovipositor tube slender and needlelike, consisting of fewer than four pairs of plates. 20. Primitive: free margin of T-IJI evenly curved to margin; derived: T-III with a subapical pit row and/or groove separating an apical rim from rest of tergum. 21. Primitive: digitus forming an apposable lobe on cuspis; derived: digitus elongate and hinged to base of cuspis, or absent. 22. Primitive: parasitize “harmless’ hosts, ie. prepupal or free- living larvae (moth, beetle or sawfly larvae or prepupae); derived: parasitize “harmful” hosts, ie. larvae or prepupae in nests prepared and often defended by adults (wasps and bees).
Using the above list of characteristics, it is possible to describe a hypothetical ancestor for the Chrysididae. This progenitor had relatively complete wing venation with a closed marginal cell, and complete submarginal, discoidal and cubital cells. The tongue was unmodified and bethylidlike, resting flat in the oral fossa, with six segmented maxillary palpi and four segmented labial palpi. The thorax had the prothorax hinged to the mesothorax, tarsal claws with a single subsidiary tooth, metanotum rounded, and propodeum dorsally elongate and boxlike without lateral teeth. The abdomen had a convex venter, seven visible segments in males and six in females, the latter had a true sting, the apical tergum simple and unmodified, and male genitalia had the digitus forming a lobe on the cuspis. The ancestor of the chrysidids was probably a parasite of freeliving coleopteran or lepidopteran larvae. No existing family exhibits all or nearly all of the primitive states listed above. However, more of them are found with the Bethylidae than any other family we have studied.
GLOSSARY (Fig. 2 illustrates external structural features) brow: foreheadlike swelling between ocelli and scapal basin. cuspis: volsellar element that is setose at least distally. digitus: movable volsellar element that is not setose but usually bars denticles and is often broadly sword shaped. EH: eye height as seep in front view. F-I, F-Il, F-Ill, etc.: flagellar articles or flagellomeres. GC: genal carina. IAD: interantennal distance. IOD: interocellar distance measured between lateral ocelli. LOD: greatest diameter of lateral ocellus. Malar space: shortest distance from compound eye to mandible base. MOD: greatest diameter of median ocellus. n., s., sw., etc.: compass directions abbreviated in lower case. OOD: ocellocular distance measured from lateral ocellus to compound eye. paramere: “clasper’ or main lateral organ of male genitalia. Attached at its base are volsella and median organ or aedeagus. pit row: curving row of pits subapically on T-III in most Chrysidinae. PD: puncture diameter. propodeal tooth: lateral projection of propodeum. RS stub: sclerotized basal part of radial sector in forewing. S-I, S-II, S-III, etc.: apparent abdominal sterna, male subgenital plate is S-V III. Scalpel basin: area of frons covered by rotation of scapes. States or Provinces of New World: generally given in upper case as ONTARIO, NEVADA, DURANGO, etc. subantennal distance: measured from antennal socket to clypeal apex just inside lateral clypeal notch. T-I, T-II, T-III, etc.: apparent abdominal terga. volsella: sublateral organ of male genitalia, sometimes simple, sometimes divided into cuspis and digitus.



Taken from: L.S. Kimsey and R.M. Bohart, 1990. The Chrysidid Wasps of the World.

MORPHOLOGY

To prevent confusion and duplicative terminology we have tried to determine structural homologies among chrysidids and other aculeates and to apply the most appropriate names to these structures. In a few instances the modifications found in the Chrysididae appear to be unique in this family, at least to the extent that they have not been named by anyone previously. We have relied almost exclusively on Bohart and Menke (1976) for the terminology of sculptural features of the mesopleuron, many of which appear to be homologous with those found in Sphecidae. In addition, we have referred to the following studies to develop a morphological nomenclature for the Chrysididae: Snodgrass (1910) and Richards (1956).

The chrysidid abdomen presents a variety of nomenclatural difficulties. First, although the propodeum is actually the first abdominal segment, we refer to the first gastral segment as the beginning of the functional abdomen. Therefore our thorax equals thorax propodeum, or mesosoma, of Michener (1944) and abdomen (actual segment II onwards) equals the metasoma of Michener. Secondly, the chrysidid abdomen consists of two functional parts: the external, apparent abdomen comprising segments I, II, III, IV, or V depending on the subfamily and sex, and the invaginated segments, which form a telescoping genital (males) or ovipositor (females) tube. The external segments are often referred to as the ‘abdomen’, although all abdominal segments (I-IX) are well developed in this family.

Head

The basic structure of the chrysidid head differs little from other members of the Chrysidoidea; although even the most primitive chrysidids such as Cleptes tend to have large, bulging, well-developed eyes.

In most chrysidids, the back of the head tends to be relatively flat or shallowly convex, without distinct carinae or sulci. However, Chrysidini and some Parnopini have a broad indentation across the back of the head above the occipital foramen, adjacent to a more dorsal, transverse swelling. A diagnostic feature of Chrysidini is the formation of this indentation into a transverse carina or welt, similar to that seen in some Ceramius (Masaridae), except that in chrysidines it terminates laterally in a tooth-like projection or hook (Fig. 2a). This sulcus is called the preoccipital carina and the lateral projection the preoccipital hook. In Chrysis ehrenbergi and in Brugmoia this hook is represented only by the sharply curved apex of the carina. In all chrysidids the occipital suture is reduced, extending only from the midventral line to the posterior tentorial pits. The post-occiput is projecting and collar-like. The hypostoma is, at most, indicated by a crease or an elevated region adjacent to the oral fossa. The occipital region in Loboscelidia projects strongly posteriorly, the cervical projection (Fig. 2c), with an odd lateral lamella.

Laterally, the chrysidid head consists primarily of the compound eye. The length of the malar space, the area between the ventral eye margin and the mandibular insertion, is of taxonomic importance at the species, species group, and generic levels. Many amisegine genera are distinguished by the presence of a malar sulcus traversing the malar space from eye to mandible. The sculpture of the gena is also taxonomically important. A characteristic uniting the Chrysidini, Allocoeliini, and Parnopini is the presence of a carina extending through the gena from the mandible along the posterior eye margin (Fig. 2b). Exochrysis and some other genera have a large subgenal area enclosed by a ventral carina below the genal carina (see Fig.66b).

Face

The front of the head, or the face, is distinctive in chrysidids. The antennae insert low on the face on the dorsal margin of the clypeus, except in Loboscelidiines. The clypeus is usually short and broad, with an apical truncation and the dorsal margin tends to extend around, and partly enclose, the antennal sockets (Fig. 2b). In Cleptinae the clypeus almost appears to be divided into three parts by the antennal sockets (see Fig. 1 2e). However, in some genera, such as Stilbum, the clypeus may be longer than it is broad. The labrum is a small rounded lobe often hidden by the mandibles and clypeus. In all subfamilies except the Cleptinae and Loboscelidiinae, the face has two discrete regions: the upper frons or brow, and the scapal basin (Fig. 2b). The frons has a transverse carina, or TFC, in most Chrysidini, two amisegine genera, and a few species of Hedychridium. This transverse carina may also have branches extending towards, and sometimes encircling, the mid ocellus (Fig. 2b), as in Sri/bum. In most species the scapal basin has punctation different from that of the frons and may also have a zone of fine medial cross-ridging and/or an impunctate and polished medial stripe. The principle exception to this is Chrysura, where the face is nearly flat and densely punctate granulose throughout (see Fig. 1 16a). A different situation occurs in loboscelidiines where the antennae insert near the middle of the face on a shelf-like medial projection (Fig. 2c).

Chrysidid mandibles are usually simple, with one or two subapical teeth. Cleptine mandibles tend to be robust with two or three large subapical teeth. Some elampines, chrysidines, and male amisegines have simple edentate mandibles. Male Adelphe have very odd foliaceous mandibles (see Fig.18a, b).

Chrysis ehrenbergi is unusual in another respect. Females exhibit a variety of allometric forms. Some females are very similar to males, others have a greatly enlarged head, with a number of features exaggerated including the swollen genal region and large, ventrally toothed mandibles (Fig. 21, g).

Antenna

All chrysidids, male or female, have a scape, pedicel, and 11 flagellomeres (Fig. 3). The flagellomeres are usually cylindrical in cross-section. In a variety of species, however, the male (or less commonly the female) flagellum may be broad and flattened. Sexual dimorphism in flagellar shape is relatively common in chrysidids. In amisegines the female flagellomeres are generally short and wide; in males they are greatly elongate and setose. The basal male flagellomeres of one species, Pleurochrysis bruchi, are broad and flabellate (see Fig. 1 32b).

There is also considerable variation in the relative lengths of the first three flagellomeres. In Holopyga and Ceratochrysis the first flagellomere is usually three or more times as long as it is broad. The third flagellomere is usually shorter than the first, at least in females, but in Chrysis stilboides, and other species once placed in the genus Pyria, it is the longest flagellomere (with the first much reduced). Another modification of these flagellomeres occurs in the males of some species of Chrysura, where the segments are lobulate beneath.

Mouthparts

The development of the mouthparts in Chrysididae varies from the most primitive form seen in Cleptes to the most specialized in some Parnopes (Fig. 2d, e; see Fig. 1 56c). The mouthparts of Cleptinae, Amiseginae, and Loboscelidiinae are similar to those in other chrysidoid families. In these groups the maxillae and labium lie flat in the oral fossa, with relatively long palpal segments. In all chrysidids, except some Parnopes, there are five maxillary and three labial palpal segments. The tongue in Elampini is not particularly modified, except that the cardines, stipes, and prementum are elongate and the tongue protrudes from the oral fossa. In Allocoelia the tongue length is sexually dimorphic with the male cardines, stipes, and prementum considerably longer than those of females. Both sexes of Allocoelia have an elongate glossa and galea, covered with microtrichia. Chrysidini all have a somewhat elongate tongue that protrudes from the oral fossa. As in the elampines the cardo, stipes, and prementum are considerably enlarged. The most unusual tongue modification can be seen in many Parnopini. In these species the glossa and galea are greatly elongate, forming a tubular proboscis (see Fig. 1 56c). The palpi are virtually absent in Parnopes. In Cephaloparnops the proboscis is elongate but the palpi are generally normal with five maxillary and three labial palpal articles (see Fig.152c).

Thorax

The simplest and least modified thoracic structure can be seen in the Cleptinae and winged Amiseginae. In this respect these subfamilies do not differ much from other winged chrysidoids, particularly Berhylidae. The more specialized chrysidids often have an elaborately modified thorax, with complex sculpturing and punctation.

Pronotum

The pronotum is separate from, and hinged to, the propleura; except in Loboscelidia where they appear to be fused. It is relatively box-like with a lateral lobe which covers the spiracle (see Fig.48). This lobe touches or nearly touches the tegula in Cleptinae, Amiseginae, and Loboscelidiinae. The two are widely separated in the Chrysidinae.

In most chrysidids the pronotum has an anterior shelf, or collar, followed by a large, elevated quadrate region. These two regions may be separated by submedial carinae, as in Parnopes; a foveate groove, as in Cleptidea; or a vertical lateral carina, as in a variety of Chrysidini. Many chrysidines and elampines have 2-4 deep, submedial, or one medial, pit at the base of the pronotal shelf. These pits correspond with a short, broad internal apodeme. The pronotum may also have a lateral carina, particularly noticeable in Ipsiura and some Praestochrysis, or have a variety of sculpturing, such as the medial and transverse grooves, pits, and foveae, seen in the Cleptinae.

Propleura and sterna

The propleura are usually relatively simple and unmodified. Some species of Allocoelia have a lateral propleural tooth. In the majority of chrysidids the propleural plates meet ventromedially for 50 per cent, or less, of the distance from the anterior margin to the posteromedial junction (Fig. 4 and 5). Where this is the case the prosternum is large, exposed, and somewhat diamond-shaped, and medially longer than the length of the juncture between the propleura. Amisegines and loboscelidiines, however, differ from this form. In these subfamilies the propleura meet for 60-80 per cent of the total distance and the prosternum is narrow and largely obscured by the pleura (Fig. 46, c). The exposed part of the prosternum is considerably shorter than the juncture between the propleura. This condition is much more typical of members of the other chrysidoid Families, except Scolebythidae. Scolebythids also have a very large exposed prosternum.

The chrysidid prosternum is generally medially concave, with two large subapical lobes which extend into the body cavity. These lobes are not visible unless the coxae are removed.

Scutum

The scutum is a broad, somewhat convex plate, divided into three subequal parts by the notauli. Laterally, between the notaulus and the tegula, is the parapsidal line, extending from the posterior scutal margin, anteriorly, about halfway across the scutum, parallel to the notauli. Originating at the anterior scutal margin are two short, submedial admedian lines. Notauli are usually present but their absence or incomplete nature can be a diagnostic feature, as for example in Omalus glotneratus or Loboscelidia. Most species have a rather coarsely and evenly punctate scutum. However, in Omaha and related genera they are either impunctate and very shiny or the punctures tend to be clumped along or between the notauli.

Scutellum

Usually shorter than the scutum, the scutellum is generally unmodified in chrysidids. One exception to this occurs in Muesebeckidium where the scutellum is sharply declivitous anteriorly (Fig. 74d).

Tegula

Tegular shape is an important characteristic at the species and subfamily levels. Parnopes and Loboscelidia have huge ovoid tegulae, which cover both wing bases. In a few species of Parnopes the tegulae are somewhat comma-shaped. Loboscelidia have an unusual arrangement, in which the outer edge of the tegula snaps into a groove on the mesopleuron, effectively enclosing and protecting the wing bases (see Fig.48). Allocoelia have unusually small tegulae, which in the case of capensis are nearly hidden by the elevated and bulging scutum.

Mesopleuron

One of the most complex parts of the thorax is the mesopleuron. The sculpturing of this part of the body is of particular importance at the species, species group, and generic level (see Figs 4 and 5). Some of the grooves and carinae found on the chrysidid mesopleuron appear to be homologous with those found in the Sphecidae and so we have adopted the nomenclature developed by Bohart and Menke (1976), where possible. This is the only complete terminology available since these structures have not been consistently named in studies of other chrysidoid families.

All chrysidids have a signum, scrobe, and subalar fossa (Figs 4 and 5). The signum is a short sublateral line located on the ventral surface of the mesopleuron. The scrobe is a deep pit located about halfway between the fore wing and mid coxa near the meso-metapleural suture. Located just below the wing base is the subalar fovea, a deep, well-marked pit or groove; or in the case of Loboscelidia a ridge.

Anteriorly, the chrysidid mesopleuron has a small ovoid plate set off by a suture from the rest of the pleuron, called the epicnemium.

Most elampines (Fig. 5a) have a relatively simple mesopleuron. However, in several genera, including Muesebeckidium and some Holopyga, the dorsal half of the mesopleuron is greatly expanded both laterally and anteriorly (see Fig.73). In these species the scrobal sulcus extends nearly vertically immediately below the scrobal carina. This catina originates at the meso-metapleural suture above the scrobe and ends at the omaulus. The omaulus originates on the epicnemium and extends ventrally. Just anterior to the mid coxae is the precoxal carina, which extends from the mid-line to the sternaulus. In Omalus, and related genera, a carina extends from the verticaulus along the signum to its apex.

Allocoeliines (Fig. 5b) ate not particularly unusual except that they have a carina that runs from the fore wing base, alongside the scrobe, and ends just above the mid coxa. The dorsal part, above the scrobe, is the episternal carina, and the ventral half is the verticaulus.

Chrysidines (Fig. 5c) can have the most highly sculptured mesopleuron of any of the chrysidids. Many species have a large, well-developed scrobal sulcus, which divides the thorax in half horizontally, and the episternal suds, which divides it in half vertically. The verticaulus and omaulus form a ventral U or V-shaped loop. In Brugmoia, Stilbum, and a number of Chrysis these two carinae may be modified into one or more large teeth or knobs. Probably the most extreme modification of the mesopleuron occurs in Stilbum.

Parnopini (Fig. 5d) are somewhat intermediate between Elampini and Chrysidini in terms of their mesopleural sculpture; the subalar fovea is very long and extends posteriorly almost to the scrobe. In all Parnopes, however, the scrobal carina and omaulus form the edge of a broad, relatively flat epimeral plate.

Mesosternum

The mesosternum in chrysidids is reduced and barely visible. It consists of a narrow, transverse sclerite forming a declivity between the posterior margin of the mesopleuron and the mid coxae.

Metanotum

The metanotum can be the most highly modified of all the dorsal plates. The metanotum in Elampus and Parnopes has a large, dorsally flattened, blade-like medial projection (see Figs 53 and 156f, g). The whole metanotum may be conical, as in some Philoctetes. Numerous species, in a wide variety of genera, have a large or small medial tooth, or prong, on the metanotum. Conversely, a number of amisegine genera have a narrow and much reduced metanotum, or the metanotum may be indistinguishably fused to the propodeum. In addition, one of the diagnostic features of the Chrysidini and Parnopini is the presence of a large, clearly defined lateral metanotal tooth, or angle, immediately adjacent to the propodeal tooth (Fig. 3).

Metapleuron

The metapleuron is obscurely indicated, The metapleural.-propodeal suture is distinct until it reaches a scrobe-like pit, approximately halfway between the hind wing and the hind coxa. Below this point a certain amount of fusion with the propodeum has occurred and the suture may or may not be indicated by a groove or carina. In species with a metanotal tooth there is typically a transpleural carina (Fig. 3), which extends from adjacent to the scrobe to the tip of the tooth, often appearing to form a continuous arc from the ventral end of the verticaulus to the tip of the propodeal tooth.

Propodeum

In the Chrysididae the propodeum occurs in two basic forms: somewhat box-like with a long dorsal surface in the Cleptinae and most Amiseginae; and in the remaining subfamilies abruptly declivitous posteriorly, without a dorsal surface. The condition seen in cleptines and amisegines is typical for most aculeates. However, unlike other aculeates the chrysidid propodeum is expanded laterally and is broadly visible in ventral view. This modification occurs in even the most primitive chrysidids, Cleptes, which are otherwise relatively unspecialized.

All chrysidids, except a few amisegine genera, such as Amisega, have a lateral propodeal tooth or angle (Figs 3, 4 and 5). This tooth can be very long and acute, as in Cleptidea aurora; deeply notched posteriorly and truncate, as in Spintharina (Fig. 1421, j); posteroventrally lobed, as in some Chrysis; or fan-like and lobulate, as in Allocoelia (Fig. 89d, e). In some Holophris this tooth may be represented by an angulate carina. Most chrysidids have this tooth located more than halfway above the middle of the propodeum, generally in a horizontal plane with the spiracle. However, in Allocoelia the ‘tooth’ is located below the middle and oriented vertically.

The sculpture and punctation of the propodeum is important at the species and genus levels. All Exochrysis and some species of Pleurochrysis and Chrysis have a basomedial tooth or projection. In Allocoelia members of the capensis species group lack punctation in a medial triangular area of the propodeum, or propodeal enclosure. In another example the presence of a medial ridge distinguishes species of Hedychrum (see Fig.64l) from Hedychridium. The lack of carinae or enclosures is diagnostic for Adelopyga, Haba (see Fig.58b), Prochridium, and Minymischa.

Legs

Chrysidid legs are shaped for a fossorial habit and are generally unmodified; there are, however, some exceptions to this. The fore femur often has a basoventral angle or projection and/ or carina in many elampines and Parnopes. Female Muesebeckidium (see Fig. 74e), Minymischa (Fig. 72b), and a few other elampine genera have broad, flattened tarsomeres which appear prehensile. Many Female Parnopes have elongate rake spines on the fore tarsomeres, and in one species, P. grandior, these spines occur on the hind tarsomeres as well.

The tarsal claws of many species have a subapical tooth, or teeth. A specialized feature of the Chrysidini and Parnopini is the absence of these teeth. Cleptinae, Allocoeliini, Loboscelidiinae, and most Amiseginae all have a single, submedial perpendicular tooth on each claw. Elampines have a variety of conditions. Several genera, including Hedychridium (see Fig. 62c), have a single perpendicular tooth. Holopyga, Omalus, Philoctetes, Holophris, Pseudomalus, Muesebeckidium, and Elampus have between one and four subsidiary teeth depending on the species (as in Fig.6&1, e). In Minymischa and Adelopyga, the number and arrangement of the subsidiary teeth is sexually dimorphic. The remaining elampine genera, except Xerochrum and Prochridium, have a subparallel tooth making the claw appear almost bifid. Unlike the rest of this tribe these two genera have simple, edentate claws.

Wings

As in the other chrysidoid families, except Plumariidae, the chrysidid wing venation is much reduced (Fig. 6). The fore wing has, at most, six closed cells: costal, radial, medial, cubital, discoidal, and submedial. The hind wing has no closed cells, and at most has indications of the following veins: costa (C), subcosta radius (Sc R), first anal vein (Al), media cubitus (M Cu), and radial sector (us). The most complete wing venation is found in the Chrysidini (Figs 6 and 70. Further reductions in venation are important taxonomic characteristics. In the chrysidine forewing, variations occur in the length and shape of Rs and the presence or absence of Rs M, Cu, and m-cu. Part of this reduction correlates with size; the smaller the individual the more reduced its wing venation tends to be, within limits. Many species of Chrysidea, Caenochrysis, and Primeuchroeus have the discoidal cell incomplete or absent, particularly due to reduction or loss of Rs M and m - Cu.

Other chrysidids, except Loboscelidia, have lost the scierotized Rs M, rn-cu, and Cul veins in the fore wing, and Rs tends to be less than half the length of the marginal cell (if this vein were extended to the wing rnargin (Fig. 7). In Allocoelia the presence or absence of a sclerotized Rs M is an important species characteristic. Amisegines vary in the shape of Rs in a number of genera, including Ade/phe and Atoposega. Here, Rs bends sharply and reaches the costal margin long before the wing tip, as indicated by the wing crease.

A number of different venations occur in Elampini. In Holopyga, Omaha, Holophris, Philoctetes, Pseudomalus, Elampus, Muesebeckidium, and some Hedychridium the M vein is strongly arched, nearly forming a right angle. The relative positions of M and cu-a when they join M Cu also varies, with M meeting M Cu well before cu-a, parallel with, or after cu-a. The most restricted venation occurs in Microchridium, where the venation is limited to the basal fourth or fifth of the wing and the veins are not clearly indicated (see Fig.69)

Loboscelidia and Rhadinoscelidia have the most reduced venation of all the chrysidid genera. These wasps have no costal, Rs M, m-cu, Cu, or stigmal veins (Fig. 7c).

The hind wing of all chrysidids has a well-developed anal lobe, at least stubs of C, Sc, and Al (except loboscelidiines), and no anal lobe. Again, the most complete hind wing venation occurs in the Chrysidini (Fig. 6).

Abdomen

External segments The number of segments in the visible chrysidid abdomen varies from subfamily to subfamily. Amiseginae, Cleptinae, and Loboscelidiinae all have five segments in males and four in females. Parnopini have the male abdomen four segmented, and the female three-segmented. Allocoeliines are unusual because they have two terga and three sterna visible. The remaining tribes have three-segmented abdomens in both sexes; although male Exochrysis, Neochrysis, Ipsiura, and Pleurochrysis have the fourth sternum visible.

The segments of the visible abdomen ate generally heavily sclerotized and weakly intermusculated. Cleptinae, some amisegines, and loboscelidiines can flex the abdomen to some extent. In these subfamilies the abdominal venter is convex and not generally separated from the dorsum by a sharp edge. In the Chrysidinae the abdomen is virtually inflexible. The abdominal venter is flat or concave and clearly separated from the dorsum by a sharp lateral edge. Members of this subfamily assume a defensive posture by rolling up into a ball, completely covering the ventral half of the head and thorax with the abdomen. Where the abdominal venter is set off from the dorsum, the tergal plates have a sublateral weakening or joint, separating a plate called the laterotergite. This is the part of the tergum that wraps around the ventrolateral fold in the abdomen.

Abdominal segments I-V have spiracles. In allocoeliines, chrysidines, parnopines, and a few amisegines the spiracles on segments II-IV are located on the laterotergite. The spiracles are located either on the tergal fold, or on the tergite adjacent to the tergal fold in the remaining chrysidids.

Segment I has some diagnostic features in a few genera. In Exochrysis, for example, the anterolateral corners of the tergum are sharp. Allocoelia capensis has a vertical carina on the anterolateral corner. Other genera, such as Hedychrum may have carinulae or ridges radiating up the frontal declivity from the petiolar insertion. The sternum is divided by a sulcus into a narrow, anterior petiolar region and a broad, flat posterior plate. This anterior region is very narrow with a pronounced medial keel in amisegines. In the other subfamilies the petiolar region is broader and flattened, with two large submedial pits or depressions.

Cleptinae, Amiseginae, and Loboscelidiinae all have relatively smooth, tapering abdomens. Most amisegines have a lateral edge or fold extending through the second and, sometimes, third segment. Duckeia cyanea has S-II produced prow-like anteromedially. Aside from patterns of punctation in some amisegines, and coloration, the external abdomen has few useful characteristics in these subfamilies.

In Allocoelia T-II is the apical dorsal segment, although S-III is the apical ventral plate. T-II is elongate and may terminate in 2-5 apical teeth or angles, and the apical margin may also be thickened and/or rolled under (Figs 88 and 89c).

Elampines have relatively unmodified abdomens. Segment IT is the longest of the three and it may be swollen posteriorly as in Hedychridium crebrum. The greatest variation occurs in T-III. In Elampus it is produced and snout-like apically (see Figs 53 and 54h-l). Omaha often have a transparent rim and an apicomedial notch (see Fig.76b, c). In Philoctetes the apex of T-III may even become snout-like in a few species (see Fig. 80c). A very few Palaearctic and African Hedychrum and Hedychridium have the apical margin of T-III with 2-6 teeth or a medial notch (see Figs 62 and 64).

The structure of T-III varies most in Chrysidini. The dominant genus Chrysis can have from 0-8 apical teeth, although, except for a few rate, five-toothed forms, the number of teeth is always a multiple of two. Other genera have odd numbers of teeth, either 0, 1, or 3: Chrysidea, Primeuchroeus (Fig. 136), Trichrysis (Fig. 150), and Caenochrysis (Fig. 98); or 5: Praestochrysis (Fig. 134) and Pentachrysis (Fig. 130). A few unusual species of Chrysis from Africa and southern Europe have a strongly produced, and often laterally pronged, apicomedial truncation on T-III. Brugmoia (Fig. 95), some Spinolia (Fig. 139), and Gaze/lea (Fig. 121) have multiple, irregular, and usually asymmetrical apical teeth. The remaining chrysidine genera have smooth and edentate apical rims, a medial notch (often guarded laterally by an angular projection), or 2-8 teeth.

A subapical pit row, which normally consists of a series of large deep, circular or ovoid pits in a sunken transverse, groove is also typical of the Chrysidini. This pit row is often preceded by a transverse swelling of various forms. The secondary loss of the pit row is characteristic of a few species in a variety of genera, for example Spinoia theresiae (Fig. 140f) and, typically, in the genus Neochrysis (Fig. 125).

In all Parnopines, even the primitive Isadelphia, T-III (or T-IV in males) is thick and more or less rounded apically with numerous small, irregular denticles, apically and subapically on the posterior margin (Fig. 156g). Parnopines also have a pair of sublateral foveae or transverse depressions on the apical tergum.

Internal segments

Internal abdominal segments form the ovipositor tube in females (Fig. 8). The basal segment is composed of two relatively short and broad sclerites. The more apical sclerites are much more elongate with long slender apodemes. These segments are strongly curved laterally and form a sub-cylindrical telescoping tube. In Cleptinae this tube is large and robust, composed of segments V-VIJI (Fig. 8J). Amiseginae and Loboscelidiinae have a slender, needle-like tube also formed by segments V-VIII (Fig. 8e). In the Chrysidinae the tube is formed by segments IV-VII (Fig. 86). However, in Allocoeliini T-III is also involved.

The chrysidid sting is quite reduced and essentially non- functional, a feature unique in the Aculeata. It functions more as an egg guide than a defensive structure. The valvulae are long and rod-like (Fig. 8a). The third valvula is apically setose and can be confused with the male gonocoxa when only the tip is showing.

The internal abdominal sclerites have not had much detailed study for species or generic distinctions because of the time involved in removing, clearing, separating, and slide-mounting these plates. There are two exceptions to this: segments IV and V in females, which may be quite distinctive and can be seen on a partly exserted ovipositor (Fig. 8b, d), and the sub genital plate (S-VIII) in males (Fig. 9b).

In many Chrysidini the females have abdominal segments IV and V heavily sclerotized. One of the best examples is in female Stilbum cyanurum (Fig. 86). T-IV has a series of heavy lateral ridges or partial annuli and terminates in a tooth-like structure. S-IV and T-V are heavily sclerotized apically and have one or two apicomedial teeth. S-V may also be heavily sclerotized and be either sharp or blunt apically. The degree of these modifications of the basal ovipositor tube varies, with its most extreme development occurring in Stilbum. The function of this structure is discussed in chapter 4.

The ovipositor tube in most other Chrysidinae is unmodified, although it may attain extreme length in some species groups of Chrysis.

Males also have a genital tube composed of internal telescoping segments (Fig. 9). This tube is somewhat reduced in Loboscelidiinae and Amiseginae, with segments VII and VIII reduced to narrow transverse sclerites.

As in other aculeates, S-VIII forms the subgenital plate, lying against the ventral surface of the genital capsule. T-VIIII is a membranous, somewhat U-shaped flap lying against the dorsum of the gonobase. S-VIII is generally an unremarkable transverse sclerite in most of the Chrysididae. However, in the Chrysidinae S-VIII is often highly modified, particularly in the Chrysidini. The shape of S-VIII may be a useful generic or species group character in this tribe. In most species S-VIII is somewhat bell-shaped in outline. However, in some Chrysis for example, it may be cruciform, subtriangular, or quadrate.

Male genitalia

The genital capsule of chrysidids consists of a broad collar-like gonobase, gonocoxa, volsella, and aedeagus (Fig. 9a). Unlike most chrysidids the structure of the gonocoxa, volsella, and aedeagus shows little variation between taxa in the Amiseginae and Loboscelidiinae, although further study may reveal useful characters in these groups. The shape of the gonocoxa is a useful species or group character in the Cleptinae and Chrysidinae. The gonocoxa may be lobulate apically or along the inner margin, or actually have an articulated gonostyle, as in Caenocbrysis (Figs 9a and 98d) and parnopines (Fig. 156b).

Throughout the Chrysididae the volsella is divided at the base into a slender digitus and cuspis. In some elampine genera, including Minymischa (Fig. 72e) and Muesebeckidium (Fig. 74f), the volsella is undivided, or the digitus has been lost secondarily. Parnopines have a broad, undivided, membranous volsella (Fig. 156b). The structure of the volsella is one feature that immediately separates cleptines from other chrysidids. In the Cleptinae the digitus is located medially or subapically on the cuspis, as it is in Bethylidae and most other aculeates (Figs 12f and 14e). The aedeagus is generally lanceolate, without lobes or other subsidiary structures. However, in the Cleptinae the aedeagus has a large subapical hook or row of teeth (Figs 1 2f and 14e). Exallopyga has a large spinose sub-basal lobe (see Fig.56f). Many species of Hedychridium have a sub-basal tuft of setae on the outer surface of the aedeagus.

Distinction of the sexes

We are commonly asked how to distinguish males from females in the Chrysididae. This subject has been given rather cursory treatment in the past and so we have decided to discuss this problem at some length.

Distinguishing the sexes is relatively simple in the Cleptinae, Amiseginae, Loboscelidiinae, and Parnopini. In these groups, females have one less external abdominal segment than males. Therefore, in the first three subfamilies males have five-segmented abdomens and females four. In the Parnopini males have four segments and females three. In addition, females often have the ovipostor tube partly exserted.

Determining the sex of a specimen in the Chrysidini, Elampini, and Allocoeliini without extracting the genitalia is much more difficult. Some genera are sexually dimorphic to some extent. Male Allocoelia have much longer tongues than females. In Neochrysis, Pleurochrysis, Ipsiura, and Exochrysis male S-IV is clearly visible, protruding beyond S-III for at least one-quarter of the length of S-III. S-IV is not visible in females. In Hedychrum the female S-III has a sub-basal transverse carinae and usually has an apicomedial tooth or small projection of some sort (see Fig.64c-e). Female Elampus have a row of short, erect, closely placed setae along the genal region behind the eye in lateral view (see Fig.54c). Exallopyga males have a stripe of dense appressed setae extending down the mid-line of T-III; and T-III is slightly notched apicomedially (see Fig.56b).

Otherwise, in these three subfamilies the shape of S-III differs between the sexes. In males S-III is completely flat, and the membranous apex of S-IV is often visible. S-III in females generally has a triangular swelling, apicomedially. Females can also be recognized if the ovipositor is exserted. However, if only the tip of the ovipositor is showing, this structure can be confused with the partly exserted male genitalia and vice versa. By close examination it is possible to see the slender, needle-like first valvulae between the second and third valvulae in females. Although the apices of the gonocoxae can resemble those of the third valvulae, and the apex of the aedeagus resembles that of the second valvulae, there is never a slender needle-like medial structure showing in males.

Coloration

Chrysidids are generally colourful insects. The majority of species are metallic blue or green, but a wide variety of other colours may be present, also. Certain distinct colour patterns occur within each subfamily, many of which have an interesting geographic significance.

Loboscelidiines and Allocoeliines are exceptional in a number of ways, including their complete absence of metallic coloration. All members of these subfamilies are brown, black, or reddish. In addition, a few Allocoelia may have whitish markings. Amiseginae are generally brown to black with metallic green to blue highlights on the face and thorax. The abdomen is usually non-metallic except in such rare genera as Duckeia.

The coloration of Cleptinae varies considerably from species to species. Several, including Cleptidea scutellaris and Cleptes townesi are non-metallic black. Cleptidea tend to be either black and orange, or blue, black, and red with some white markings. Cleptes may be entirely metallic, as in fritzi or purpuratus, or more commonly the head and thotax may be brightly metallic and the abdomen predominantly brown or black. In several species groups the abdomen may be non-metallic red to black. There is also some degree of sexual dimorphism in many Cleptes; females tend to be more brassy or bronzy.

Chrysidini and Elampini are always metallic (except Microchridium) and a variety of distinct colour patterns can be seen in some taxa. Many of these patterns show an interesting geographic distribution. In most of the world these wasps are blue, green, or purplish, with few exceptions. A very different situation occurs in the Palaearctic Region, particularly in Europe. Here the dominant colour is blue or green on the head and thorax, with a coppery red to brassy abdomen. One of the most striking examples of this can be seen in Chrysis. The Palaearctic ignita group is structurally the same as the Nearctic nitidula group, and in fact the species ignita and nitidula are nearly indistinguishable, except that ignite and the other species in the ignita group are brightly bicoloured, with a coppery or brassy abdomen. Members of the nitidula group are concolorous blue or green. Species in southern Spain, North Africa, and the Middle East tend to be completely brassy or coppery. In southern Africa the majority of species are blue or green but some are brightly marked with purple, orange, red, and yellow. A very odd colour pattern occurs in the Philippine Islands; species of Chrysis, Praestochrysis, and Stilbum are purplish with a bright red head. In tropical Asia a number of species of Chrysis, Praestochrysis, Primeuchroeus, and Trichrysis are green except for a large brassy or coppery spot on either side of tergum-IT. Certain Holarctic species of Hedycbridium, including the roseum group in the Palaearctic, semirufum of North America, and Xerochrum rubeum have an essentially non-metallic red abdomen.

There are relatively distinct zoogeographic differences among species of Parnopes. The African and American forms all tend to be metallic blue, green, or purple, with the head, thorax, and abdomen concolorous. In all Palaearctic species the abdomen is often quite differently coloured from the rest of the body; tending to be non-metallic red, or, particularly in the Middle East and North Africa, the entire body is quite reddish.

White markings occur on numerous species in all the subfamilies and tribes except the Loboscelidiinae and Elampini. The presence of these markings may indicate species group relationships but are never present on all the members of a genus. Female Adelphe in the Amiseginae may have the outer surface of F-I and F-II white. About one-third of Cleptidea have white markings on the face, thorax, and abdomen. Two species of Allocoelia, bidens and mocsaryi, have white on the thorax and abdomen. In Chrysidini white is found on the mandibles, F-I and F-II, femoral and tibial apices, and T-III; particularly in Spintharina, Spintharosoma, and Argocbrysis. There are two basic patterns of white markings on T-TII in this tribe; either the entire apical rim is white or at least the teeth, as in Brugmoia, Gaullea, Argocbrysis, and Spintharina, or T-III has a basolateral spot, as in Neochrysis, Pleurochrysis, Ipsiura, and Exochrysis. Many Parnopes have whitish markings on the tegulae, propodeal teeth, posterolateral angles of the terga, and even the apical denticles of T-I-III or T-IV.

Leg colour varies to some extent in the Chrysididae from black, to metallic blue or green, to yellowish. In the Elampini, Chrysidini, and Parnopini the femora and tibiae are usually metallic. The absence of metallic coloration is a useful species characteristic.

Wing coloration is generally unremarkable in chrysidids. Chrysidinae have evenly clear or brown-stained wings. Only the extremes in wing colour, either water clear as in Philoctetes telfordi or very dark brown as in Chrysis angolensis, are distinctive. Most Cleptes also have evenly stained wings; except for a few, like semiauratus, in which the wings are faintly brown banded. The wings of most Cleptidea have two relatively dark brown bands, one basal and one subapical. A few, as in the xantha group, have only a single subapical band, and scutellaris has unbanded, lightly stained wings. Loboscelidiines generally have mottled wings with spots of brown to yellow. In Amiseginae wing colour is not generally distinctive, with the membrane evenly tinted various shades of brown.

The colour of the tegulae and abdominal sternum is of particular importance in many elampine and parnopine genera. The presence or absence of metallic coloration on the tegulae and S-II and S-III is an important species characteristic in these groups, particularly in Parnopes, Hedychridium, and Hedychrum.

One cautionary note: the hue of metallic coloration can be altered, particularly if specimens are killed and/or preserved in solvent. In addition, rehydrating specimens can temporarily change the colour from purple to green, or green to coppery.

Sternal spots

In the Chrysidini most species have two more or less, round, flat black spots on S-II. We discuss these spots in this section because, despite their distinctive nature, their function is unknown. However, they may be secretory. The size and shape of the sternal spots, the distance between them, or their absence are all useful taxonomic characters. They are often sexually dimorphic. A species of Stilbichrysis, aurovirens (see Fig. l46c), and a few species of Brugmoia may have these spots also on S-III.

Vestiture

In general vestiture is not particularly conspicuous or useful as a diagnostic feature in most chrysidids. The overall colour of the erect setae on the head and thorax of some Cleptes, Hedychrum, and Chrysis may be an important species character. Many amisegines have long ocular setulae. The lack of microtrichia on the wing membrane in the fore wing medial cell is a diagnostic feature for the genera related to Omalus.

There is also a sexual component in patterns of setae. Many Hedychridium, Brugmoia, and Chrysis have dense appressed setae on the face, but it tends to be much denser in males. In the Amiseginae many genera have a long slender setose male flagellum. The length of the setae is a species character in these groups. Male Exallopyga have a stripe of appressed setae extending down the mid-line of T-III. In addition, a few male Hedychridium and Ceratochrysis may have a tuft of long setae on the femur or tibia of the mid and/or hind leg.

Terminology

Many of the following terms may be unfamiliar to other workers in Hymenoptera, and require definition.

Brow. Swelling often found above scapal basin and in front of ocelli.

Epimeral plate. Broad, flat, dorsal mesopleural plate delimited ventrally by omaulus and scrobal carina in Parnopini.

Episternal sulcus. Vertical groove extending from below fore wing to, or towards, the scrobal sulcus.

F-I-II, etc. Antennal articles (flagellomeres) following scape and pedicel.

LID. Least interocular distance.

L/w: Length versus width.

Malar space. Shortest distance from eye to mandible base.

Malar sulcus. Vertical sulcus extending from ocular margin to mandibular socket in Amiseginae.

Midocellar area. Usually depressed area in front of mid ocellus and sometimes delineated by carinules (mid ocellar carinae) arising from TFC.

Midocellar lid. Eyelid-like integumental fold behind mid ocellus.

MOD. Middle ocellus diameter.

Omaulus. Ridge or carina originating below pronotal lobe, descending obliquely and posteriorly to venter of mesopleuron.

PD: puncture diameter.

Pit row. Transverse row of pits located subapically on T-IJI in Chrysidini.

Preoccipital book. Hook-like or tooth-like projection beneath head at end of preoccipital carina in Chrysidini.

Preoccipital carina. Transverse carina or welt located above occiput in Chrysidini.

Pronotal length. Submedian length of dorsal surface; usually compared with length of scutellum.

Propodeal angle. Lateral and often hook-like angle or projection of propodeum.

Rs stub. Radial sector when greatly shortened.

S-I-II, etc. Gastral sterna.

S-II spots. Two, often large, dark spots on S-II, sometimes coalesced.

Scapal basin. Usually concave area covered by rotation of scapes; area below TFC and brow.

Scrobal carina. Carina extending along dorsal margin of scrobal sulcus.

Scrobal sulcus. Longitudinal groove extending from scrobe either horizontally, or obliquely towards venter, across mesopleuron.

Subantennal space. Distance from lowest edge of antennal socket straight down to free edge of clypeus.

Subgenal area. Roughly triangular area below genal carina, defined ventrally by ridges or carinae.

T-I-II, etc. Gastral terga beginning at the base.

TFC. Transverse frontal carina located on or just above brow.

Transpleural carina. Carina extending across metapleuron from adjacent to scrobe usually to apex of propodeal angle, and then down to hind coxa.

Verticaulus. Vertical ridge extending ventrally from scrobal sulcus near scrobe, often joining omaulus to delimit lower part of mesopleuron.


Names
Taken from: R. M. Bohart and L. S. Kimsey. 1982. A Synopsis of the Chrysididae in America North of Mexico.

HISTORICAL REVIEW The oldest names among North American chrysidids, Omalus auratus (Linnaeus 1758) and Cleptes semiauratus (Linnaeus l761 were based on European specimens. These species were probably introduced into eastern United States, auratus in the 16th or 17th centuries and semiauratus rather recently. In the case of auratus a likely means of introduction was in nests of Pemphredon lethifer (Shuckard) in rose cuttings from Europe. Cleptes semiauratus may have been brought over by plane to New Jersey. The first species endemic to North America to be described were Chrysis nitidula Fabricius (1775) and C. smaragdula Fabricius (1775). For nearly a century after Linnaeus’ landmark work a scattering of species were named by numerous authors, including in part Fabricius (1775, 1804) Say (1824, 1828, 1836), Lepeletier (1825, 1846), Guerin (1842), and Brulle (1846). Not until Dahlbom (1845, 1854) was a serious attempt made to include our species in a revisionary work. There were further miscellaneous descriptions after this until Aaron (1885) wrote the next revision, and the first to concentrate on American species. At this point there were about 61 named species known from North America, a little more than a fourth of the 227 species we are now considering. The next important study was the world revision by Mocsary (1889), followed by the world species catalog of Dalla Torre (1892). In the first half of the 20th century there were many more species descriptions. Then the synonymic catalog by Bodenstein (1951) set the stage for a series of generic revisions. These treated the Amiseginae (Krombein 1957, 1960, 1969), Omalus (Bohart and Campos 1960), Parnopes (Telford 1964), Neochrysis (Bohart 1966) Muesebeckidium (Krombein 1969), Elampus (Huber and Pengelly 197), Hedychridium (Bohart and Kimsey 1978), several monotypic genera (Bohart 1980), Pseudolopyga and Minymischa (Kimsey 1981) and Cleptes (Kimsey 1981). Knowledge of the chrysidid fauna of the world is at best partial. The only exotic areas extensively studied are the Palearctic Subregion and the southern part of the Ethiopian Region.

Phylogeny
Taken from: R. M. Bohart and L. S. Kimsey. 1982. A Synopsis of the Chrysididae in America North of Mexico.

PHYLOGENY To understand the relationships within the Chrysididae we studied various genera in each of the subfamilies and arrived at a list of 22 characteristics. A primitive and a derived state can be found for each of these characteristics in the Chrysidoidea, as discussed below. Our ideas on the interrelationships are summarized in fig. 1 in which each number indicates the derived state. The Sclerogibbidae and Scolebythidae have been left out of this treatment because we have not had adequate representation of these families. For several characteristics a number of derived states are listed. These are ordered from a to d based on the degree of modification in which d is the most highly derived.



Taken from: L.S. Kimsey and R.M. Bohart, 1990. The Chrysidid Wasps of the World.


GENERAL SYSTEMATICS

The superfamily Chrysidoidea consists of the families Scolebythidae, Plumariidae, Sclerogibbidae, Dryinidae, Bethylidae, Embolemidae, and Chrysididae. The majority of these families are highly specialized, and all are parasitoids. A variety of phylogenetic studies have been made of the Aculeata (Brothers 1975; Koenigsmann 1978; Rasnitsyn 1980) or Chrysidoidea specifically (Carpenter 1986). Based on these studies, and particularly that of Carpenter, rhe Chrysididae is the sister-group of Bethylidae. The other relationships among these families are discussed in detail by Carpenter (1986). Chrysididae Bethylidae is the sister-group of the Sclerogibbidae (Embolemidae Dryinidae).

The family Chrysididae is a diverse group of parasitic wasps. Although the majority of species commonly encountered in the field are brilliantly coloured, it would be erroneous to assume that this is a family characteristic because several subfamilies and tribes are consistently non-metallic. However, there are a number of modifications which do clearly distinguish chrysidids from other wasps.

The most obvious characteristic of the Chrysididae, which is unique for this family, is the reduction of the number of external abdominal segments and the formation of an ovipositor or genital tube by invaginated distal segments. As a result, male chrysidids have five or fewer visible gastral segments and females four or fewer. Other diagnostic features are: flagellomeres with 11 articles in both sexes; labial palpus three-segmented and maxillary palpus five-segmented (rarely fewer segments); prosternum large and exposed (except in Amiseginae Loboscelidiinae); fore wing venation with five closed cells (medial, submedial, costal, discoidal, and marginal) or fewer; hind wing without closed cells or jugal lobe; and propodeum usually with a lateral tooth or angle.

The higher classification of the Chrysididae has varied considerably through the years. Dahlbom (1854) made the first attempt to divide what he termed the Chrysidiformium into groups. These groups were the Cleptidae, Elampidae, Hedychridae, Chrysididae, Euchroeidae, and Parnopidae. This system was followed by Radoszkowski (1877). Aaron (1885) gave a somewhat different classification, dividing the Chrysididae into the Cleptinae, Elampinae, Chrysidinae, and Parnopinae. Mocsáry (1889) divided the family into Cleptinae, Amiseginae, Allocoeliinae, Ellampinae, Hedychrinae, Chrysidinae, and Parnopinae. He was the originator of the spelling of Ellampinae with a double 1’. In addition, he was the first to recognize the Allocoeliinae and Amiseginae (1889), and later (1890) the Adelphinae. The first species in these groups were described in 18746 by Smith (Allocoelia), in 1888 by Cameron (Amisega), and in 1890 by Mocsáry (Adeiphe). Dalla Torre (1892) used the classification of MocsIry. (1889). Buysson initially followed Aaron’s groupings, except that he split the Cleptinae off as a separate family in 1896, and then placed them back in the Chrysididae in 1899. In 1901 he made a major change in terminology and divided the family into ‘tribes’, Cleptidae, Heteronychidae, Euchrysidae, and Parnopidae. Bischoff (1913) Further altered this latter system of Buysson, using the Heteronychinae, including Ellampini and Hedychrini, and Holonychinae, including Pseudochrysidini, Parnopini, Allocoeliini, and Euchrysidini. In his treatment of the chrysidids of southern Africa, Edney (1944- 1956) followed Bischoffs classification. Linsenmaier (1959a), in his treatment of the European chrysidids, used a simplified version of Mocsáry’s groupings, dividing the family into the Cleptinae, Chrysidinae, Parnopinae, and Allocoeliinae.

The position of Loboscelidiinae has varied considerably. Westwood (1874) placed Loboscelidia in the Diapriidae. Ashmead (1902) considered these wasps to be a tribe of Cynipidae. Maa and Yoshimoto (1961) treated them as family Loboscelidiidae in the Chrysidoidea. Loboscelidia was then placed back in the Proctotrupoidea by Riek (1970). Finally, Day (1978) made a detailed morphological study of Loboscelidia and determined that it belonged in the Chrysididae, as the sister-group of the Amiseginae.

The classification ofBohart and Kimsey (1982) is the system used in this study, with considerable modifications. Principal among these is the treatment of the Elampini, Allocoeliini, Chrysidini, and Parnopini as tribes in the subfamily Chrysidinae. The reasoning behind the change in status of these groups is given in more detail later, but is based on the large number of shared derived characters among them. In addition, we are following Kimsey (1986a) in viewing the Allocoeliini as a sister-group of the Chrysidini Parnopini.

Cleptinae includes the genera Cleptes and Cleptidea. These are the most primitive chrysidids. Despite extensive studies we have been able to distinguish few derived characteristics for this subfamily that are not diagnostic for the family as a whole. Cleptinae is the sister-group of the rest of Chrysididae.

The number of amisegine taxa has grown tremendously since Mocsáry first proposed the subfamily in 1889, and now comprise 30 genera. It is the sister-group of the Loboscelidiinae, based on the highly modified ovipositor plates and the obscured prosternum. In addition, both are parasites of phasmatid eggs.

Loboscelidiinae includes two genera, Loboscelidia and Rhadinoscelidia. The so-called higher chrysidids (Chrysidinae) are closely related and are united by a large number of derived characteristics, including the position of the scrobal sulcus, reduced number of visible gastral segments, flat or concave gastral sternum, and absence of an occipital catina. Furthermore, all are nest parasites of wasps and bees, with the exception of Praestochrysis.

PHYLOGENETIC DISCUSSION

The phylogenetic relationships among subfamilies and rribes are represented by Fig.10. Derived characteristics used to construct this tree are discussed below.

1. Scapal basin. The presence of a flat or concave, differentially sculptured, scapal basin (Fig. 2b) occurs throughout the Chrysididae, except in the Cleptinae and Loboscelidiinae. The lack of a scapal basin in cleptines is considered to be primitive, since the facial structure in this group closely resembles that of bethylids and other non-chrysidid chrysidoids. In loboscelidiines the absence of a scapal basin is secondarily derived as a result of the dorsal elongation of the clypeus and formation of a frontal projection which surrounds the antennal sockets (Fig. 2cr).

2. Malar sulcus. Throughout the Chrysididae only the Amiseginae have a vertical sulcus that traverses the malar space from the posterior mandibular condyle to the ocular margin (Fig. 16e). In a few other groups in the Chrysidinae a line of appressed setae can be found in this position. The presence of this sulcus is considered derived.

3. Antennal position. Another of the peculiarities of the loboscelidiines is the position of the antennae. In all other chrysidids the clypeus is short, and the antennal sockets, located on the dorsal clypeal margin, insert on the lower third or fourth of the face. This position is considered primitive in chrysidids. Loboscelidiinae have an elongate clypeus and shortened frons. As a result the antennae insert mid face and are located on a bilobate or trilobate shelf or projection, with the sockets oriented horizontally (Fig. 2c).

4. Genal carina. Most chrysidid genera have an ecarinate gena, which is typical of cleptines and is considered the primitive condition in this family. However, in the Chrysidinae there is one, or occasionally two, carinae extending from the mandibular socket dorsally along the ocular margin (Fig. 3); a derived condition.

5. Transfrontal carina. The presence of a transverse carina across the frons, or TFC (Fig. 2b), is a diagnostic feature of the Chrysidini, and occurs only rarely in other groups. It has been secondarily lost in Chrysura, Spinolia, and a few species in other genera.

6. Cervical projection. Loboscelidiines have the occipital region of the head produced posteriorly into a large fringed lobe (Fig. 2c). This condition is unique in the Chrysididae and is considered derived.

7. Preoccipital carina. In the majority of genera the back of the head is evenly rounded or somewhat indented above the occipital foramen. This condition is primitive. Chrysidini have a transverse preoccipital carina (Fig. 2a), considered derived. This carina often ends laterally in a hook-like projection.

8-9. Tongue length. Bethylids, cleptines, amisegines, and loboscelidiines have the tongue lying flat in the oral fossa. The cardo is short and broad, and the galea and glossa are short lobes. In the derived condition, in Chrysidinae, the tongue base protrudes from the oral fossa, the cardo is relatively long and slender, and the galea and glossa are elongate (8), attaining extreme lengths in Parnopes. A further derivation occurs in Allocoeliini. Tongue length is sexually dimorphic in these wasps, with the male tongue considerably longer than the female (9).

10. Pronotal shape. Homologies between the shape of the chrysidid pronotum and that of other Chrysidoidea are difficult to ascertain. However, the quadrate pronotum with a short, flared anterior collar seems to be the least specialized form in this family, and most closely resembles the shape typical of other chrysidoids. The derived condition can be seen in the Cleptinae where the pronotum is narrowed submedially with an anteriorly arcuate, submedial sulcus, and a broad, flared, and often foveolate collar (Fig. 12).

11. Pronotal lobe position. Despire the specialized modification of the cleptine pronotum, the proximity of the pronotal Lobe to the tegula is primitive. In this subfamily, as well as Loboscelidiinae, Amiseginae, and Bethylidae, the pronotal lobe is immediately adjacent to the tegula. The derived condition occurs in the Chrysidinae where the lobe is separated from the tegula by at least one-third the length of the tegula.

12. Pronotal pit. Throughout the Chrysididae the pronotum is sculptured in a variety of ways. However, only in the Amiseginae is there a discrete pit at the origin of the pronotal lobe (as in Fig.20). The presence of this pit is considered derived; its function is unknown.

13. Prosternal shape. Unlike other chrysidoid families, except Scolebythidae, most chrysidids have a large, exposed prosternum. The propleura meet ventrally for less than half the distance from the apex of the collar to the prosternum. This is a derived condition characteristic of the family. However, within the Chrysididae the amisegines and loboscelidiines have rhe prosternum largely obscured by the propleura (Fig. 4), as it is in most other aculeates. The condition of the prosternum in Amiseginae and Loboscelidiinae could therefore be taken as a primitive feature retained by these subfamilies. Considering the specialized nature of these groups, a more parsimonious explanation would be that the short, hidden prosternum is actually a secondary derivation in the Chrysididae. Therefore, we are treating this modification as the derived condition.

14. Hinged pronotum. Throughout the Chrysidoidea the pronotum is freely hinged to the mesothorax. This is the typical situation in chrysidids. However, in the Loboscelidiinae the pronotum is immovably fused to the mesothorax; a derived condition.

15. Tegular clip. Another unusual and unique feature of loboscelidiines is the presence of a linear ridge on the mesopleuron, which fits over the outer edge of the tegula, holding the tegula down over the wing bases (as in Fig.48). The presence of this ‘regular clip’ is considered derived.

16. Scrobal sulcus. Although most Chrysidinae have a scrobal sulcus, the orientation and extent of the sulcus is a group character. In the primitive configuration found in Elampini and Parnopini the scrobal sulcus is short and generally shallow, extending obliquely towards the venter from the scrobal pit (Fig. 5). The derived condition occurs in the Chrysidini where the scrobal sulcus is broad and often deep, extending horizontally from the scrobe to the epicnemium, effectively bisecting the mesopleuron. Most allocoeliines lack a scrobal sulcus. In the two species that have one, latinota and capensis, it is a broad irregular pit that extends anteriorly a short distance from the scrobe.

17. Mesopleural plate. Modifications of the chrysidid mesopleuron provide useful group characters, Unlike other chrysidids, parnopines have the epimeron clearly set off from the rest of the mesopleuron, as a broad, flat plate (Fig. 5). This epitneral plate has a posterior lobe above the scrobe and is delimited ventrally by the scrobal carina and omaulus. The formation of this plate is considered derived.

18-20. Tegula size. In the majority of chrysidids the tegula is subovoid and covers the base of the fore wing. However, three different derived conditions occur in this group. The loboscelidiine tegula is enlarged and subrectangular, completely covering both wing bases and held in place by a ridge on the mesopleuron (18). Parnopines also have an enlarged tegula that extends posteriorly over both wing bases, but it is subovoid and not clipped laterally against the mesopleuron (19). Finally, in allocoeliines the tegula is much reduced and partly obscured by a lateral bulge of the scutum (20)

21. Scutellar lobe. The scutellum anterolaterally extends smoothly into the fore wing fossa in all chrysidids except the Chrysidini and the elampine genus Exallopyga. They have an anterolateral scutellar lobe, the ‘scutellar tubercle’ of French (1985), which projects posteriorly into the wing fossa (as in Fig.56e). The presence of a scutellar lobe is derived.

22. Metapleural carina. The metapleuron is relatively unmodified and ecarinate in most chrysidoids; this is the primitive condition. However, Chrysidinae have a transverse metapleural carina that extends from near the scrobe to the propodeal projection (Fig. 3). The carina may terminate in a large lobe, particularly in the Chrysidini.

23. Propodeal tooth. Throughout the Chrysididae the propodeal angles are more or less located in a horizontal plane with the spiracle. This orientation is the primitive condition in this family. In allocoeliines the propodeal angle is oriented obliquely between the spiracle and hind coxa. In addition, the propodeal angles are broad and usually lobulate. 24. Propodeal shape. In the Chrysidoidea the propodeum is generally box-like with a long dorsal surface. A specialized form occurs in the Chrysidinae, where the propodeum is essentially wedge-shaped (Fig. 3), with no dorsal surface, and abruptly declivitous posteriorly.

25-26. Claw dentition. Most chrysidoids have a single, subperpendicular tooth on all of the tarsal claws. This is also the condition seen in Cleptinae, Amiseginae, Loboscelidiinae, Allocoeliini, and primitive Elampini. However, two specialized conditions occur in the Chrysidinae. The Chrysidini Parnopini have edentate tarsal claws (25). The majority of Elampini have more than one subsidiary tooth, typically 2-5 (26).

27-28. Forewing venation. A major change in wing venation occurs between Cleptinae and other chrysidids. In cleptines Rs M originates considerably below the stigma, often one-half to two-thirds of the way along the media (Fig. 7a). This is also the most common position in other chrysidoids and is taken here to be the primitive condition. In the remaining chrysidid subfamilies Rs M originates at or near the apex of M, just below the stigma (Fig. 7) (27).

Although all chrysidid wing venation is reduced, the most extreme reduction occurs in loboscelidiines where the hind wings have no sclerotized veins, and their fore wings are without a stigma. They also lack costal and cu-a veins (Fig. 7c) (28).

29-31. Gastral segments. Aculeates in general have six visible gastral segments in females and seven in males. The primitive condition in chrysidids is four in females and five in males and is found in Cleptinae, Amiseginae, and Lobocelidiinae. Several derived conditions occur in this family. The first (29) is the general reduction of one or more gastral segments to three or fewer in females, and four or fewer in males (retained in Parnopini). Allocoeliines are further reduced with two apparent segments in both sexes, although S-Ill is clearly visible (30). Both Elampini and Chrysidini have three-segmented abdomens, with no sexual dimorphism (31). The condition in allocoeliines and parnopines suggests that the three-segmented abdomen has in fact evolved twice, particularly since no other abdominal characters indicate a close relationship between elampines and chrysidines.

32. Gastral sternum. A derived feature of the Chrysidinae is the flat or concave sternum. This abdominal characteristic is unusual in the Aculeata. Coupled with the flat sternum, the terga have a discrete lateral lobe, or laterotergite, which is sharply folded under ventrally.

33. Abdominal spiracles. All chrysidids have some indication of a laterotergite, and even in some cleptines it is at least indicated by a fold. A laterotergite was not found in the other non-chrysidid chrysidoids examined. This fold becomes a sharp crease or ridge in other chrysidids. The position of the spiracles on segments II-IV are assumed to be primitive in Cleptinae, Amiseginae, and Loboscelidiinae, where they are located more dorsally on the tergum proper (Fig. 9c). Within the Chrysidinae the spiracles are in the primitive position in Elampini. In the remaining chrysidine tribes the spiracles are located on the laterotergite (Fig. 9b), a derived condition.

34-36. Apical tergum. In the Chrysidinae the simplest and least specialized form of T-III is seen in elampines where the apical margin is smooth and evenly rounded. Several derived conditions occur in this subfamily. Chrysidines have a pit row that extends subapically along the margin (Fig. 3) (34). This pit row is found in all but a few species and in the genus Neochrysis. It is apparently a secondary reduction in these groups, since traces of the pit row can still be found. In Chrysidini the apical rim of T-III is usually dentate or at least strongly angulate (35). Parnopines have the hind margin of the apical tergum rolled under and thickened, with numerous irregular denricles and two oblique subapical foveae, one on either side of the mid-line (as in Fig. 156g) (36).

37. Sternum II. A unique, derived feature of the Chrysidini is the presence of two dark spots on S-II. These spots vary from broadly rounded, to comma-shaped, to contiguous.

38-39. Volsellar structure. Typically, in Aculeara the volsella has both a digitus and cuspis, and the digitus forms a sort of opposable lobe on the cuspis. This arrangement, with the digitus articulated with the apical half of the cuspis, also occurs in cleptines (as in Fig.12f), and is assumed here to be the primitive condition. In the Amiseginae Loboscelidiinae Chrysidinae the digitus and cuspis articulate basally and both are generally long and slender (Fig. 9a) (38). A further modification of the volsella occurs in Parnopini, where the volsella is broad and membranous without a digitus (Fig. 156b) (39).

40. Ovipositor tube. Formation of an ovipositor tube is a derived character unique to the Chrysididae in the Aculeata. Within this family the tube is generally robust with all of the involved segments well developed. The ovipositor of the Amiseginae Loboscelidiinae is considered derived. In this group, segments V-VII are highly elongate and slender, appearing needle-like (Fig. Sc).

41-43. Hosts. Assuming that chrysidids evolved from a scolebythid-like ancestor, then the primitive host would be a prepupal moth or beetle or similar inactive, harmless, and cryptic form. Therefore, cleptines which parasitize prepupal sawfly larvae, attack a specialized type of host (41). Two different specializations occur in chrysidids. The first involves parasitism of phasmarid eggs in the Loboscelidiinae Amiseginae (42). The second group, Chrysidini, are nest parasites of non- social wasps and bees, placing their eggs in the host cells, and either feeding on the host provisions, or the prepupal host (43).

The Chrysididae consists of three major phylogenetic lines, the Cleptinae, Amiseginae Loboscelidiinae, and Chrysidinae. Cleptinae is the sister-group of the other three subfamilies. The Loboscelidiinae, Amiseginae, and Chrysidinae constitute a monophyletic group based on the subapical origin of Rs M on M, digitus and cuspis joined basally, and formation of a scapal basin. The Amiseginae Loboscelidiinae lineage is characterized by elongation and reduction of the ovipositor plates, reduction of the prosternum, and parasitism of walking stick eggs.

KEY TO SUBFAMILIES AND TRIBES OF CHRYSIDIDAE

1. Gaster with 5 external segments in males and 4 in females; sternum strongly convex; propodeum generally box-like in profile, with some horizontal dorsal surface 2

Gaster with 4 or fewer segments in males and 3 or fewer in females; sternum strongly concave or flat; propodeum abruptly declivitous posteriorly, short and somewhat wedge shaped, without dorsal surface (Chrysidinae) 4

2. Face above antennal sockets convex, without indication of scapal basin; clypeus deeply emarginate below each antennal socket and with a protruding medial truncation (Fig. 12e); pronotum narrowed submedially, bisected by transverse groove (Fig. 12a-c); female ovipositor robust (Fig. 8f); male volsella with digitus attached near apical third of cuspis (Fig. 12f) Cleptinae, p.52

Face above antennal sockets flat or concave with some indication of scapal basin (Fig. 26), or antennae insert on frontal projection (Fig. 2c); clypeus not deeply emarginate below each antennal socket, without protruding medial truncation (Fig. 2b) 3

3. Antennal sockets horizontal and located mid face on projection (Fig. 2c); head with large cervical projection; tegula large covering both wing bases and held in place by ridge on mesopleuron (Fig. 48); fully winged, fore wing venation reduced without costal vein or stigma (Fig. 7c) Loboscelidiinae, p.l41

Antennal sockets vertical and located on lower fourth of face; head without cervical projection; tegula small, covering only fore wing base and without associated ridge on mesopleuron; fore wing with costal vein and stigma (Fig. 76), or strongly brachypterous Amiseginae, p.71 4.

4. Gaster with 2 visible terga; body without metallic coloration; tegula small and partly hidden by notum; propodeal tooth originating just above hind coxa Allocoeliini, p.272

Gaster with 3 or 4 terga; body partly or entirely metallic (except Microebridium); tegula at least normal-sized, not obscured by notum; propodeal tooth originating adjacent to metanotum, well above hind coxa 5

5. Tegula large, covering both wing bases; apical abdominal tergum with 2 subapical foveae, and apically thickened with numerous small irregular teeth or denticles (as in Fig.156) Parnopini, p.574

Tegula normal-size, only covering fore wing base; apical abdominal tergum without 2 subapical foveae, at most slightly thickened apically, apical rim simple, dentate, denticulate, serrate, or medially notched 6

6. T-III with subapical pit row (sometimes faint) (Fig. 3), and tarsal claws edentate; occiput with transverse welt or carina above foramen, often ending in a hook’ (Fig. 2a) (rarely absent); mesopleuron with scrobal sulcus horizontal (rarely absent) Chrysidini, p.276

T-III without pit row and tarsal claws dentate (rarely edentate); occiput without welt, carina, or hook’; mesopleuron with scrobal sulcus oblique Elampini, p.152


Taken from: L.S. Kimsey and R.M. Bohart, 1990. The Chrysidid Wasps of the World.


GENERAL SYSTEMATICS

The superfamily Chrysidoidea consists of the families Scolebythidae, Plumariidae, Sclerogibbidae, Dryinidae, Bethylidae, Embolemidae, and Chrysididae. The majority of these families are highly specialized, and all are parasitoids. A variety of phylogenetic studies have been made of the Aculeata (Brothers 1975; Koenigsmann 1978; Rasnitsyn 1980) or Chrysidoidea specifically (Carpenter 1986). Based on these studies, and particularly that of Carpenter, rhe Chrysididae is the sister-group of Bethylidae. The other relationships among these families are discussed in detail by Carpenter (1986). Chrysididae Bethylidae is the sister-group of the Sclerogibbidae (Embolemidae Dryinidae).

The family Chrysididae is a diverse group of parasitic wasps. Although the majority of species commonly encountered in the field are brilliantly coloured, it would be erroneous to assume that this is a family characteristic because several subfamilies and tribes are consistently non-metallic. However, there are a number of modifications which do clearly distinguish chrysidids from other wasps.

The most obvious characteristic of the Chrysididae, which is unique for this family, is the reduction of the number of external abdominal segments and the formation of an ovipositor or genital tube by invaginated distal segments. As a result, male chrysidids have five or fewer visible gastral segments and females four or fewer. Other diagnostic features are: flagellomeres with 11 articles in both sexes; labial palpus three-segmented and maxillary palpus five-segmented (rarely fewer segments); prosternum large and exposed (except in Amiseginae Loboscelidiinae); fore wing venation with five closed cells (medial, submedial, costal, discoidal, and marginal) or fewer; hind wing without closed cells or jugal lobe; and propodeum usually with a lateral tooth or angle.

The higher classification of the Chrysididae has varied considerably through the years. Dahlbom (1854) made the first attempt to divide what he termed the Chrysidiformium into groups. These groups were the Cleptidae, Elampidae, Hedychridae, Chrysididae, Euchroeidae, and Parnopidae. This system was followed by Radoszkowski (1877). Aaron (1885) gave a somewhat different classification, dividing the Chrysididae into the Cleptinae, Elampinae, Chrysidinae, and Parnopinae. Mocsáry (1889) divided the family into Cleptinae, Amiseginae, Allocoeliinae, Ellampinae, Hedychrinae, Chrysidinae, and Parnopinae. He was the originator of the spelling of Ellampinae with a double 1’. In addition, he was the first to recognize the Allocoeliinae and Amiseginae (1889), and later (1890) the Adelphinae. The first species in these groups were described in 18746 by Smith (Allocoelia), in 1888 by Cameron (Amisega), and in 1890 by Mocsáry (Adeiphe). Dalla Torre (1892) used the classification of MocsIry. (1889). Buysson initially followed Aaron’s groupings, except that he split the Cleptinae off as a separate family in 1896, and then placed them back in the Chrysididae in 1899. In 1901 he made a major change in terminology and divided the family into ‘tribes’, Cleptidae, Heteronychidae, Euchrysidae, and Parnopidae. Bischoff (1913) Further altered this latter system of Buysson, using the Heteronychinae, including Ellampini and Hedychrini, and Holonychinae, including Pseudochrysidini, Parnopini, Allocoeliini, and Euchrysidini. In his treatment of the chrysidids of southern Africa, Edney (1944- 1956) followed Bischoffs classification. Linsenmaier (1959a), in his treatment of the European chrysidids, used a simplified version of Mocsáry’s groupings, dividing the family into the Cleptinae, Chrysidinae, Parnopinae, and Allocoeliinae.

The position of Loboscelidiinae has varied considerably. Westwood (1874) placed Loboscelidia in the Diapriidae. Ashmead (1902) considered these wasps to be a tribe of Cynipidae. Maa and Yoshimoto (1961) treated them as family Loboscelidiidae in the Chrysidoidea. Loboscelidia was then placed back in the Proctotrupoidea by Riek (1970). Finally, Day (1978) made a detailed morphological study of Loboscelidia and determined that it belonged in the Chrysididae, as the sister-group of the Amiseginae.

The classification ofBohart and Kimsey (1982) is the system used in this study, with considerable modifications. Principal among these is the treatment of the Elampini, Allocoeliini, Chrysidini, and Parnopini as tribes in the subfamily Chrysidinae. The reasoning behind the change in status of these groups is given in more detail later, but is based on the large number of shared derived characters among them. In addition, we are following Kimsey (1986a) in viewing the Allocoeliini as a sister-group of the Chrysidini Parnopini.

Cleptinae includes the genera Cleptes and Cleptidea. These are the most primitive chrysidids. Despite extensive studies we have been able to distinguish few derived characteristics for this subfamily that are not diagnostic for the family as a whole. Cleptinae is the sister-group of the rest of Chrysididae.

The number of amisegine taxa has grown tremendously since Mocsáry first proposed the subfamily in 1889, and now comprise 30 genera. It is the sister-group of the Loboscelidiinae, based on the highly modified ovipositor plates and the obscured prosternum. In addition, both are parasites of phasmatid eggs.

Loboscelidiinae includes two genera, Loboscelidia and Rhadinoscelidia. The so-called higher chrysidids (Chrysidinae) are closely related and are united by a large number of derived characteristics, including the position of the scrobal sulcus, reduced number of visible gastral segments, flat or concave gastral sternum, and absence of an occipital catina. Furthermore, all are nest parasites of wasps and bees, with the exception of Praestochrysis.

PHYLOGENETIC DISCUSSION

The phylogenetic relationships among subfamilies and rribes are represented by Fig.10. Derived characteristics used to construct this tree are discussed below.

1. Scapal basin. The presence of a flat or concave, differentially sculptured, scapal basin (Fig. 2b) occurs throughout the Chrysididae, except in the Cleptinae and Loboscelidiinae. The lack of a scapal basin in cleptines is considered to be primitive, since the facial structure in this group closely resembles that of bethylids and other non-chrysidid chrysidoids. In loboscelidiines the absence of a scapal basin is secondarily derived as a result of the dorsal elongation of the clypeus and formation of a frontal projection which surrounds the antennal sockets (Fig. 2cr).

2. Malar sulcus. Throughout the Chrysididae only the Amiseginae have a vertical sulcus that traverses the malar space from the posterior mandibular condyle to the ocular margin (Fig. 16e). In a few other groups in the Chrysidinae a line of appressed setae can be found in this position. The presence of this sulcus is considered derived.

3. Antennal position. Another of the peculiarities of the loboscelidiines is the position of the antennae. In all other chrysidids the clypeus is short, and the antennal sockets, located on the dorsal clypeal margin, insert on the lower third or fourth of the face. This position is considered primitive in chrysidids. Loboscelidiinae have an elongate clypeus and shortened frons. As a result the antennae insert mid face and are located on a bilobate or trilobate shelf or projection, with the sockets oriented horizontally (Fig. 2c).

4. Genal carina. Most chrysidid genera have an ecarinate gena, which is typical of cleptines and is considered the primitive condition in this family. However, in the Chrysidinae there is one, or occasionally two, carinae extending from the mandibular socket dorsally along the ocular margin (Fig. 3); a derived condition.

5. Transfrontal carina. The presence of a transverse carina across the frons, or TFC (Fig. 2b), is a diagnostic feature of the Chrysidini, and occurs only rarely in other groups. It has been secondarily lost in Chrysura, Spinolia, and a few species in other genera.

6. Cervical projection. Loboscelidiines have the occipital region of the head produced posteriorly into a large fringed lobe (Fig. 2c). This condition is unique in the Chrysididae and is considered derived.

7. Preoccipital carina. In the majority of genera the back of the head is evenly rounded or somewhat indented above the occipital foramen. This condition is primitive. Chrysidini have a transverse preoccipital carina (Fig. 2a), considered derived. This carina often ends laterally in a hook-like projection.

8-9. Tongue length. Bethylids, cleptines, amisegines, and loboscelidiines have the tongue lying flat in the oral fossa. The cardo is short and broad, and the galea and glossa are short lobes. In the derived condition, in Chrysidinae, the tongue base protrudes from the oral fossa, the cardo is relatively long and slender, and the galea and glossa are elongate (8), attaining extreme lengths in Parnopes. A further derivation occurs in Allocoeliini. Tongue length is sexually dimorphic in these wasps, with the male tongue considerably longer than the female (9).

10. Pronotal shape. Homologies between the shape of the chrysidid pronotum and that of other Chrysidoidea are difficult to ascertain. However, the quadrate pronotum with a short, flared anterior collar seems to be the least specialized form in this family, and most closely resembles the shape typical of other chrysidoids. The derived condition can be seen in the Cleptinae where the pronotum is narrowed submedially with an anteriorly arcuate, submedial sulcus, and a broad, flared, and often foveolate collar (Fig. 12).

11. Pronotal lobe position. Despire the specialized modification of the cleptine pronotum, the proximity of the pronotal Lobe to the tegula is primitive. In this subfamily, as well as Loboscelidiinae, Amiseginae, and Bethylidae, the pronotal lobe is immediately adjacent to the tegula. The derived condition occurs in the Chrysidinae where the lobe is separated from the tegula by at least one-third the length of the tegula.

12. Pronotal pit. Throughout the Chrysididae the pronotum is sculptured in a variety of ways. However, only in the Amiseginae is there a discrete pit at the origin of the pronotal lobe (as in Fig.20). The presence of this pit is considered derived; its function is unknown.

13. Prosternal shape. Unlike other chrysidoid families, except Scolebythidae, most chrysidids have a large, exposed prosternum. The propleura meet ventrally for less than half the distance from the apex of the collar to the prosternum. This is a derived condition characteristic of the family. However, within the Chrysididae the amisegines and loboscelidiines have rhe prosternum largely obscured by the propleura (Fig. 4), as it is in most other aculeates. The condition of the prosternum in Amiseginae and Loboscelidiinae could therefore be taken as a primitive feature retained by these subfamilies. Considering the specialized nature of these groups, a more parsimonious explanation would be that the short, hidden prosternum is actually a secondary derivation in the Chrysididae. Therefore, we are treating this modification as the derived condition.

14. Hinged pronotum. Throughout the Chrysidoidea the pronotum is freely hinged to the mesothorax. This is the typical situation in chrysidids. However, in the Loboscelidiinae the pronotum is immovably fused to the mesothorax; a derived condition.

15. Tegular clip. Another unusual and unique feature of loboscelidiines is the presence of a linear ridge on the mesopleuron, which fits over the outer edge of the tegula, holding the tegula down over the wing bases (as in Fig.48). The presence of this ‘regular clip’ is considered derived.

16. Scrobal sulcus. Although most Chrysidinae have a scrobal sulcus, the orientation and extent of the sulcus is a group character. In the primitive configuration found in Elampini and Parnopini the scrobal sulcus is short and generally shallow, extending obliquely towards the venter from the scrobal pit (Fig. 5). The derived condition occurs in the Chrysidini where the scrobal sulcus is broad and often deep, extending horizontally from the scrobe to the epicnemium, effectively bisecting the mesopleuron. Most allocoeliines lack a scrobal sulcus. In the two species that have one, latinota and capensis, it is a broad irregular pit that extends anteriorly a short distance from the scrobe.

17. Mesopleural plate. Modifications of the chrysidid mesopleuron provide useful group characters, Unlike other chrysidids, parnopines have the epimeron clearly set off from the rest of the mesopleuron, as a broad, flat plate (Fig. 5). This epitneral plate has a posterior lobe above the scrobe and is delimited ventrally by the scrobal carina and omaulus. The formation of this plate is considered derived.

18-20. Tegula size. In the majority of chrysidids the tegula is subovoid and covers the base of the fore wing. However, three different derived conditions occur in this group. The loboscelidiine tegula is enlarged and subrectangular, completely covering both wing bases and held in place by a ridge on the mesopleuron (18). Parnopines also have an enlarged tegula that extends posteriorly over both wing bases, but it is subovoid and not clipped laterally against the mesopleuron (19). Finally, in allocoeliines the tegula is much reduced and partly obscured by a lateral bulge of the scutum (20)

21. Scutellar lobe. The scutellum anterolaterally extends smoothly into the fore wing fossa in all chrysidids except the Chrysidini and the elampine genus Exallopyga. They have an anterolateral scutellar lobe, the ‘scutellar tubercle’ of French (1985), which projects posteriorly into the wing fossa (as in Fig.56e). The presence of a scutellar lobe is derived.

22. Metapleural carina. The metapleuron is relatively unmodified and ecarinate in most chrysidoids; this is the primitive condition. However, Chrysidinae have a transverse metapleural carina that extends from near the scrobe to the propodeal projection (Fig. 3). The carina may terminate in a large lobe, particularly in the Chrysidini.

23. Propodeal tooth. Throughout the Chrysididae the propodeal angles are more or less located in a horizontal plane with the spiracle. This orientation is the primitive condition in this family. In allocoeliines the propodeal angle is oriented obliquely between the spiracle and hind coxa. In addition, the propodeal angles are broad and usually lobulate. 24. Propodeal shape. In the Chrysidoidea the propodeum is generally box-like with a long dorsal surface. A specialized form occurs in the Chrysidinae, where the propodeum is essentially wedge-shaped (Fig. 3), with no dorsal surface, and abruptly declivitous posteriorly.

25-26. Claw dentition. Most chrysidoids have a single, subperpendicular tooth on all of the tarsal claws. This is also the condition seen in Cleptinae, Amiseginae, Loboscelidiinae, Allocoeliini, and primitive Elampini. However, two specialized conditions occur in the Chrysidinae. The Chrysidini Parnopini have edentate tarsal claws (25). The majority of Elampini have more than one subsidiary tooth, typically 2-5 (26).

27-28. Forewing venation. A major change in wing venation occurs between Cleptinae and other chrysidids. In cleptines Rs M originates considerably below the stigma, often one-half to two-thirds of the way along the media (Fig. 7a). This is also the most common position in other chrysidoids and is taken here to be the primitive condition. In the remaining chrysidid subfamilies Rs M originates at or near the apex of M, just below the stigma (Fig. 7) (27).

Although all chrysidid wing venation is reduced, the most extreme reduction occurs in loboscelidiines where the hind wings have no sclerotized veins, and their fore wings are without a stigma. They also lack costal and cu-a veins (Fig. 7c) (28).

29-31. Gastral segments. Aculeates in general have six visible gastral segments in females and seven in males. The primitive condition in chrysidids is four in females and five in males and is found in Cleptinae, Amiseginae, and Lobocelidiinae. Several derived conditions occur in this family. The first (29) is the general reduction of one or more gastral segments to three or fewer in females, and four or fewer in males (retained in Parnopini). Allocoeliines are further reduced with two apparent segments in both sexes, although S-Ill is clearly visible (30). Both Elampini and Chrysidini have three-segmented abdomens, with no sexual dimorphism (31). The condition in allocoeliines and parnopines suggests that the three-segmented abdomen has in fact evolved twice, particularly since no other abdominal characters indicate a close relationship between elampines and chrysidines.

32. Gastral sternum. A derived feature of the Chrysidinae is the flat or concave sternum. This abdominal characteristic is unusual in the Aculeata. Coupled with the flat sternum, the terga have a discrete lateral lobe, or laterotergite, which is sharply folded under ventrally.

33. Abdominal spiracles. All chrysidids have some indication of a laterotergite, and even in some cleptines it is at least indicated by a fold. A laterotergite was not found in the other non-chrysidid chrysidoids examined. This fold becomes a sharp crease or ridge in other chrysidids. The position of the spiracles on segments II-IV are assumed to be primitive in Cleptinae, Amiseginae, and Loboscelidiinae, where they are located more dorsally on the tergum proper (Fig. 9c). Within the Chrysidinae the spiracles are in the primitive position in Elampini. In the remaining chrysidine tribes the spiracles are located on the laterotergite (Fig. 9b), a derived condition.

34-36. Apical tergum. In the Chrysidinae the simplest and least specialized form of T-III is seen in elampines where the apical margin is smooth and evenly rounded. Several derived conditions occur in this subfamily. Chrysidines have a pit row that extends subapically along the margin (Fig. 3) (34). This pit row is found in all but a few species and in the genus Neochrysis. It is apparently a secondary reduction in these groups, since traces of the pit row can still be found. In Chrysidini the apical rim of T-III is usually dentate or at least strongly angulate (35). Parnopines have the hind margin of the apical tergum rolled under and thickened, with numerous irregular denricles and two oblique subapical foveae, one on either side of the mid-line (as in Fig. 156g) (36).

37. Sternum II. A unique, derived feature of the Chrysidini is the presence of two dark spots on S-II. These spots vary from broadly rounded, to comma-shaped, to contiguous.

38-39. Volsellar structure. Typically, in Aculeara the volsella has both a digitus and cuspis, and the digitus forms a sort of opposable lobe on the cuspis. This arrangement, with the digitus articulated with the apical half of the cuspis, also occurs in cleptines (as in Fig.12f), and is assumed here to be the primitive condition. In the Amiseginae Loboscelidiinae Chrysidinae the digitus and cuspis articulate basally and both are generally long and slender (Fig. 9a) (38). A further modification of the volsella occurs in Parnopini, where the volsella is broad and membranous without a digitus (Fig. 156b) (39).

40. Ovipositor tube. Formation of an ovipositor tube is a derived character unique to the Chrysididae in the Aculeata. Within this family the tube is generally robust with all of the involved segments well developed. The ovipositor of the Amiseginae Loboscelidiinae is considered derived. In this group, segments V-VII are highly elongate and slender, appearing needle-like (Fig. Sc).

41-43. Hosts. Assuming that chrysidids evolved from a scolebythid-like ancestor, then the primitive host would be a prepupal moth or beetle or similar inactive, harmless, and cryptic form. Therefore, cleptines which parasitize prepupal sawfly larvae, attack a specialized type of host (41). Two different specializations occur in chrysidids. The first involves parasitism of phasmarid eggs in the Loboscelidiinae Amiseginae (42). The second group, Chrysidini, are nest parasites of non- social wasps and bees, placing their eggs in the host cells, and either feeding on the host provisions, or the prepupal host (43).

The Chrysididae consists of three major phylogenetic lines, the Cleptinae, Amiseginae Loboscelidiinae, and Chrysidinae. Cleptinae is the sister-group of the other three subfamilies. The Loboscelidiinae, Amiseginae, and Chrysidinae constitute a monophyletic group based on the subapical origin of Rs M on M, digitus and cuspis joined basally, and formation of a scapal basin. The Amiseginae Loboscelidiinae lineage is characterized by elongation and reduction of the ovipositor plates, reduction of the prosternum, and parasitism of walking stick eggs.

KEY TO SUBFAMILIES AND TRIBES OF CHRYSIDIDAE

1. Gaster with 5 external segments in males and 4 in females; sternum strongly convex; propodeum generally box-like in profile, with some horizontal dorsal surface 2

Gaster with 4 or fewer segments in males and 3 or fewer in females; sternum strongly concave or flat; propodeum abruptly declivitous posteriorly, short and somewhat wedge shaped, without dorsal surface (Chrysidinae) 4

2. Face above antennal sockets convex, without indication of scapal basin; clypeus deeply emarginate below each antennal socket and with a protruding medial truncation (Fig. 12e); pronotum narrowed submedially, bisected by transverse groove (Fig. 12a-c); female ovipositor robust (Fig. 8f); male volsella with digitus attached near apical third of cuspis (Fig. 12f) Cleptinae, p.52

Face above antennal sockets flat or concave with some indication of scapal basin (Fig. 26), or antennae insert on frontal projection (Fig. 2c); clypeus not deeply emarginate below each antennal socket, without protruding medial truncation (Fig. 2b) 3

3. Antennal sockets horizontal and located mid face on projection (Fig. 2c); head with large cervical projection; tegula large covering both wing bases and held in place by ridge on mesopleuron (Fig. 48); fully winged, fore wing venation reduced without costal vein or stigma (Fig. 7c) Loboscelidiinae, p.l41

Antennal sockets vertical and located on lower fourth of face; head without cervical projection; tegula small, covering only fore wing base and without associated ridge on mesopleuron; fore wing with costal vein and stigma (Fig. 76), or strongly brachypterous Amiseginae, p.71 4.

4. Gaster with 2 visible terga; body without metallic coloration; tegula small and partly hidden by notum; propodeal tooth originating just above hind coxa Allocoeliini, p.272

Gaster with 3 or 4 terga; body partly or entirely metallic (except Microebridium); tegula at least normal-sized, not obscured by notum; propodeal tooth originating adjacent to metanotum, well above hind coxa 5

5. Tegula large, covering both wing bases; apical abdominal tergum with 2 subapical foveae, and apically thickened with numerous small irregular teeth or denticles (as in Fig.156) Parnopini, p.574

Tegula normal-size, only covering fore wing base; apical abdominal tergum without 2 subapical foveae, at most slightly thickened apically, apical rim simple, dentate, denticulate, serrate, or medially notched 6

6. T-III with subapical pit row (sometimes faint) (Fig. 3), and tarsal claws edentate; occiput with transverse welt or carina above foramen, often ending in a hook’ (Fig. 2a) (rarely absent); mesopleuron with scrobal sulcus horizontal (rarely absent) Chrysidini, p.276

T-III without pit row and tarsal claws dentate (rarely edentate); occiput without welt, carina, or hook’; mesopleuron with scrobal sulcus oblique Elampini , p.152


Geographic distribution
Taken from: R. M. Bohart and L. S. Kimsey. 1982. A Synopsis of the Chrysididae in America North of Mexico.

BIOGEOGRAPHY Western United States with its great diversity in elevation and climate has lead to the development of a considerable number of endemic genera of predaceous Hymenoptera as well as a great many species. It is not surprising, then, to find a similarly large proportion of western parasites. The 100th meridian has frequently been cited as a dividing line between the “humid east” and the “arid west”. This is an oversimplification, but the fact remains that a slightly irregular 20 mile corridor which roughly follows the 100th meridian is an important biogeographical division. We have selected five of the larger chrysidid genera to emphasize the point. In Chrysis there are 34 species exclusively to the west, 9 exclusively to the east, and 25 occurring on both sides. Comparable figures for Hedychridium are 27:1:3, for Ceratochrysis 22:1:2, for Argochrysis 15:0:0, and for Chrysura 7:0:3. In summary for the 148 species in these five genera, the figures are 105:11:33 or 71 percent exclusively west of the 100th meridian, 7 percent east of it, and 22 percent ranging widely on both sides (see map). The disproportion in numbers of genera is also marked but less dramatically so. The 24 nearctic genera that we recognize are distributed as follows: seven genera exclusively west of the 100th meridian, three to the east, and 14 ranging on both sides. The great majority of New World chrysidids are colored in various shades of green, blue and purple. The same can be said for the fauna of the Oriental, Ethiopian and Australasian Regions. On the other hand, the palearctic species are for the most part marked with bright red or shades of coppery and gold, hence the common names for the family: ruby wasps or gold wasps. In many cases the nearctic and palearctic species are closely related, as in Chrysis coerulans and ignita, yet quite different in coloration. The reasons for this disparity in coloration are not known. It is true that a blue or green chrysidid left overnight in a humidifier will often become a little coppery and this indicates that there is some relationship between moisture and interference colors. However, most of the few red-marked nearctic species are restricted to arid regions, so there is no obvious correlation with climatic humidity.

Natural history
Taken from: R. M. Bohart and L. S. Kimsey. 1982. A Synopsis of the Chrysididae in America North of Mexico.

Biology of Chrysididae Chrysididae are usually thought to be a derivative of a bethylidlike ancestor. Bethylidae, which are parasitic on moth and beetle larvae, have six abdominal segments in the female and a true sting. In the supposed transition to chrysidids the host list has expanded, the number of abdominal segments has reduced, and a tubular ovipositor has been developed. Although hosts are known for fewer than one fourth of North American chrysidids, some generalizations can be made. First, type of host is practically a subfamilial character in Chrysididae. Second, there seems to be a direct relationship between morphology of the parasite, (and hence classification) and the type of host, particularly its ability to defend its nest. Each of these points, all with associated habits, is discussed briefly below. In the most primitive subfamilies, Cleptinae parasitize tenthredae sawfly larvae and Amiseginae attack walking stick (Phasmatidae). All of the remaining subfamilies specialize on various aculeate families including Sphecidae, Megachilidae, Eumenidae and Masaridae. In the chrysidine genus, Praestochrysis the theoretical aculeate host has been bypassed and Lepidoptera larvae are attacked directly (D. Pa,1936). Each chrysidid genus or species group (in the case of Chrysis) attack a different host stage. Biological information on additional species may show this to be more a specific than a group trait. The host stage most commonly attacked is the resting prepupa. This is accomplished in one of two ways. First, the female chrysidid may chew a hole in the host’s cocoon and oviposit through it onto the prepupa. This method is used by Cleptes (Clausen 1940) and Praestochrysis (D. Parker 1936). Similarly, Chrysis fuscipennis apparently chew hole through the mud nest wall and cocoon of Sceliphron or Eumene and oviposits on the prepupa (Stage 1960). The second and more common mode is used by Chrysis(at least the nisseri group), Chrysura, Parnopes, and Argochrysis. Here the chrysidid oviposits in the host cell while it is being provisioned, a Chrysura pacifica (Krombein 1967) or after provisioning is complete and the cell is sealed, as in Stilbum cyanurum (Förster) (Moczar 1961). In either case the chrysidid egg hatches and the first instar parasite attaches itself to the host larva. Instead of immediately beginning to feed, the chrysidid larva waits until the host reaches its prepupa stage. At this point the parasite goes through its development as it consumes the host. There is some evidence that the chrysidid may take in a small quantity of fluid from the host before the latter reaches the prepupal stage (Krombein 1967), but no growth occurs. In other chrysidid taxa the larva feeds on the host egg or early instar, after which the provisions are eaten. This behavior has been recorded for Pseudolopyga, Neochrysis, Trichrysis, and Chrysis (at least in the coerulans group). The chrysidid egg once placed h the host cell, hatches immediately and the larva begins feeding on everything in the cell. As in the case of prepupal feeders there are two ways that the chrysidid adult gets its egg into the host cell. The commonest mode is by direct oviposition. The second method is reminiscent of that used by Trigonalidae (Clausen 1940). Pseudolopyga carrilloi oviposits on Nysius nymphs (Lygaeidae), which are in turn provisioned by sphecid wasps of the genus Solierella. The entire sequence was studied in the laboratory by Carrillo and Caltagirone (1970). Pseudolopyga eggs on Nysius apparently do not hatch unless the bugs are provisioned by Solierella. There is a positive correlation between chrysidid morphology a nature of the host. Those subfamilies of Chrysididae which parasitize harmless or helpless hosts, such as tenthredinoid larvae or walking stick eggs, have four often lightly sclerotized abdominal segments in the female. Furthermore, the venter is convex and the wasp is unable to roll up into a ball. Included in this group are the Cleptinae, Amiseginae and Loboscelidiinae (an Old World group). The remaining subfamilies, Elampinae, Chrysidinae, Parnopinae and Allocoeliinae (an African group), parasitize hosts which are able to defend their nests with sting and mandibles. In these cases the adult female chrysidid may have to protect herself while ovipositing in the nest of the host. These parasites have only three or rarely two usually heavily sclerotized and punctate external abdominal segments and a flat or concave venter. Thus, they can roll up into a relatively impervious ball when threatened. The question of host-parasite specificity has not been studied in detail. It is apparent that many types of correlation exist. For instance, one species of chrysidid, Chrysis coerulans, attacks a family of hosts, Eumenidae. A number of cases are known in which a species may attack a genus of hosts, as in Chrysurissa densa versus Pseudomasaris (Masaridae). Several situations are genus to genus: Argochrysis to Ammophila (Sphecidae), and Hedychrum to Cerceris (or the closely related Eucerceris) (Sphecidae). Other arrangements known are species group to genus, as in the Chrysis lauta group to the bee genus Anthidium (Megachilidae), or genus to family, as in Chrysura to several genera of Megachilidae. Obviously, there are many sorts of interactions. The one not absolutely proven is species to species, although such may occur in the absence of related forms of parasite and host.

Taken from: L.S. Kimsey and R.M. Bohart, 1990. The Chrysidid Wasps of the World.




HISTORICAL OVERVIEW

The history of chrysidid nomenclature parallels that of insects in general. The earliest work began with Linnaeus (1758). Most of the published studies of the following 100 years were purely descriptive, especially at the species level. Towards the end of the nineteenth century occasional efforts were made to bring information together in a revisional form. The quantity of taxonomic work was certainly influenced in a negative way by periods of national conflict. The Napoleonic Wars (1804-15) were one example. The effect of the First World War (1914-18) was drastic, and can be measured by the slim size of Zoological Record volumes during those years. The Second World War had a similar effect.

Some 140 workers have published about 4000 chrysidid names, and some have written catalogues, reviews, synopses, or monographs. The complete list of authors is given in the bibliography. Only the more significant of these are discussed below.

The earliest species and genus names were established by Linnaeus, DeGeer, Pallas, Scopoli, and J.R. Förster. Their chrysidid descriptions were usually part of ambitious encyclopaedic publications, and little attempt was made to define genera or higher categories. Count Maximillian Spinola (1805-41) published a series of papers, and Thomas Say (1824-36) provided some early descriptions of American chrysidids.

Andrea Dahlbom, in several papers, including early ones in 1829, 1831, and culminating in his monograph of 1854, was the first author to make a concerted attempt to pull together all information on chrysidids, and to make the group understandable with keys to genera and species. He contributed more than 150 specific names and his descriptions were models for the time. He also took varietal differences into account.

Some years after Dahlbom, but ranking with him in importance, was Alesandro (Sandor) Mocsáry (1878-1914). His Monographie Chrysididarum in 1889 was a landmark and brought together everything known at that time. Mocsáry himself confined his collecting activities to Hungary but a great deal of material came to him from other parts of the world. His descriptions and redescriptions of older species were detailed and useful. The only negative comments that can be made about this remarkable worker are that the generic concepts were somewhat weak, and the series of papers from 1890 to his death in 1914 were not quite as good as his earlier work. However, a majority of his 650, or more, new species names are still valid.

There were more than 50 other active chrysidid taxonomists who were, in a sense, competing with Mocsáry. Of these Robert du Buysson (1887-1913) was the dominant figure, describing nearly 250 species and subspecies. His collection at the Natural History Museum in Paris is one of the five best in the world.

During this period Bischoff provided 92 new specific names, many subsequently synonymized by others, and created six new generic names, Eurychrysis, Pseudogonochrysis, Pseudotetrachrysis, Pseudobexachrysis, Stilbichrysis, and Cephaloparnops. Fortunately, his awkward ‘pseudo’ names are no longer in use. Bischoff followed the earlier work of J. Lichtenstein (1876) who all too simply arranged the generic names of the Chrysis group according to the number of teeth on T-III (tergum-III). These were Olochrysis (0), Gonochrysis (indistinct), Monochrysis (1), Dichrysis (2), Trichrysis (3), Tetrachrysis (4), Pentachrysis (5), and Hexachrysis (6). Of these only Trichrysis and Pentachrysis are still in use. It is interesting that Lichtenstein used Tetrachrysis in place of Chrysis Linnaeus (1758). He might be excused because the International Rules of Nomenclature of 1901 had not appeared. However, Bischoff and others who subsequently used Tetrachrysis had no such excuse.

General d’Artillerie O. Radozskowski (sometimes spelled Radozskowsky) (1866-91) was the first Russian to publish extensively on Chrysididae, providing about 104 specific names. He was the first to point out the taxonomic significance of male genitalia, an anatomical feature largely ignored by most later workers. Andreas Semenov (1891-1912) continued the work on Central Asian chrysidids, and between 1891 and 1967 published about 350 specific names and 32 generic or subgeneric ones. Many of these appeared in manuscripts published after his death in 1942, as discussed later. The majority of his types are in the Zoological Institute, Leningrad.

Elzeár Abeille de Perrin (1877-79) studied in Paris under Perris. He is responsible for some 39 specific names, at least half of which are known to be synonyms. Buysson (18876) published some of Abeille’s manuscript names under authorship of ‘Ab.’ but the descriptions were clearly rewritten and must be attributed to Buysson. Since the interchange of types was practically non-existent in Mocsáry’s time, and with competition between more than 50 describers, much synonymy resulted. The contribution of Dalla Torre (1892) in presenting the chrysidid part of his Catalogus Hymenopterorum was a much needed forward step. Although he did not provide many new names he corrected misidentifications, placing them in the proper synonymy.

The period between the First and Second World Wars was not especially productive. Some 25 workers contributed new species. Bodenstein (19396) advanced the study of genera with his The genotypes of the Chrysididae.

After the Second World War a number of large regional monographs were published. Balthasar (1943-53) wrote extensively on the fauna of south-eastern Europe, but created considerable synonymy. Linsenmaier (1951) produced an important, but preliminary, work on European chrysidids. Nikol’skaya (1950-54) published several of Semenov’s manuscripts posthumously, with some of the new species attributed to both authors. Apparently, descriptions assigned solely to Semenov were dictated to Nikol’skaya after 1932.

Critiques of more recently active authors (1982-to present day) are not appropriate. However, a special note should be made about the work of Walter Linsenmaier of Luzern (Ebikon), Switzerland. His 1959 revision of the European species was another landmark. Here, a species group concept was formalized for the first time, with descriptions and keys to various categories. Linsenmaier also created a number of generic and subgeneric names. Although we do not agree with the status of all of these, many are quite useful. Most of Linsenmaier’s types are in his private collection and have been unavailable for study.

E. Berry Edney wrote an impressive series of papers on chrysidids of southern Africa, giving keys and illustrations. Since he restricted his field to South Africa and Zimbabwe, some synonymy resulted with wide-ranging species in the Afrotropical Region. Most of his types are at Cape Town, Pretoria (Transvaal Museum), or London. Karl V. Krombein (1956-1990) of the Smithsonian Institute published a series of papers on Amiseginae, Elampinae, and Chrysidinae. He is especially noted for his detailed work on the Amiseginae, with many new genera and species. His types are nearly all at Washington, DC.

Lázlo Moczar, former Curator at the Hungarian Museum, has been the outstanding recent authority on Cleptinae, initiating six new subgenera of Cleptes in 1962, and describing several new species.

Finally, some general comments can be made. The most prolific describers (50 or more species) have been Mocsáry, Semenov, Buysson, Linsenmaier, Radoszkowski, Edney, Krombein, Bohart, and Kimsey. Descriptions at the species level are basically important even though some synonymy may result. On the other hand, a different sort of credit should be given to those whose work was truly revisional: such early authors as Dahlbom, Mocsáry, Aaron, Semenov, and Bingham; and later workers such as Linsenmaier, Bohart, Zimmermann, Moczár, Edney, Campos (with Bohart), Krombein, Telford, Huber and Pengelly, Horning, Kimsey, and French.

A survey of newly proposed generic and subgeneric names reveals some interesting statistics. In the period between 1758 and 1784 only Chrysis Linnaeus (1761) appeared. From 1785 to 1804 three more generic names were presented: Hedychrum Latreille, Omaha Panzer, and Parnopes Latreille. In the early l800s 18 more genera were named, and by 1914 57 new generic names were given, many of which have since been synonymized. After 1914 the total number of new generic and subgeneric names was 96, allotted to the various subfamilies as follows: Cleptinae (8), Amiseginae (25), Loboscelidiinae (1), Elampini (25), Chrysidini (37).

A chrysidid worker soon finds it necessary to visit or borrow material from important world collections. On the basis of both numbers of chrysidid types and overall size, the five most outstanding museums are at Budapest (Mocsáry collection), Leningrad (Semenov collection), Paris (Buysson collection), London (various contributors), and Luzern (Linsenmaier collection). Runners-up will be found in Pretoria, Philadelphia, Davis, Washington, Vienna, East Berlin, and Copenhagen. At one time or another we have been able to visit each of these museums.

A considerable number of chrysidid species remain to be described, based on our examination of unidentified material from desert regions. At the same rime it seems likely that nearly as many are still lurking in species lists, and only diligent study will help to place them as synonyms. It is to be hoped that in future work we will see; (1) a decrease in naming of subspecies and subgenera; and (2) naming of new species only in connection with generic revisions or large geographical studies.



MATERIALS AND METHODS

We have attempted to provide as much information on the Chrysididae as possible in this study. Due to space limitations we have used a variety of notations and abbreviations which require explanation. In addition, we feel that it is important to describe as fully as possible our approach to this study, as well as the facts which have led us to certain decisions. This will enable others to evaluate our conclusions for themselves.

For each major taxonomic group we give a variety of information. Discussions for each tribe and subfamily include ancestral characteristics, phylogenetically important characters and a corresponding cladogram, keys to genera, and relationships among taxa. Generic discussions include generic synonymy and diagnostic features, relationships to other genera, and detailed species lists.

Species lists

A generic study, such as this one, would only be half complete without synonymic species lists. It has taken us thousands of hours to collect the relevant information, and for various reasons it is still not complete. We have examined some 2000 type specimens through borrowing and visits to the major museum depositories. Since nineteenth-century authors, before the establishment of the International Rules of Nomenclature, described many species from a series of syntypes, in many instances it has been necessary to designate lectotypes, both for valid species and for synonyms. On the whole, curators have been most co-operative in lending types and other specimens, and our appreciation has been expressed in the Acknowledgements. Some of the older types are no longer in existence and we have indicated this by ‘lost?’ or destroyed.’

In order to be helpful to future workers, we have furnished as much detail as possible in the checklists. A typical, but fictitious, listing might be: Chrysis nonedita (Smith) 1873:26 (Tetrachrysis). Lectotype male (desig. Bohart herein); Spain: Madrid (VIENNA). (ignita group).* in explanation, this theoretical species of Chrysis was published by Smith in 1873 on page 26 (reference in the Bibliography). It was originally placed in the wrong genus (Tetrachrysis), so Smith is placed in parentheses. He described it from several specimens (syntypes), so Bohart here designates a single one of them as a lectotype. The original locality for the lectotype was Madrid, Spain. The type is now in the Vienna Natural History Museum (All museums containing type specimens are denoted by the city name in the lists. The full museum name is given in the Acknowledgements under the relevant city.). The species belongs in the Chrysis ignita group. When the repository is supposed to be a certain museum, but this has not been verified, we have followed the name of the museum with a question mark. Also if we have seen the primary type we give an asterisk (*) or if only secondary types then (**) at the end of the species entry.

Some authors have used the designations f. (form) and ab. (aberration) for observed variations, along with new Latin names. These are invalid according to the International Rules of Nomenclature. However, they could possibly be elevated to the subspecies or species rank, and this has occurred several times. Consequently, we have listed these names in synonymy. When an elevation in rank has taken place, we have attributed the name to the reviser, with their publication date.

We have not indicated new combinations in these lists because this would be cumbersome. In some genera, for example Brugmoia, nearly every species would be a new combination. In others, like Allochrysis, the specific and generic names are new combinations but the generic name was previously treated as a subgenus.

Each entry in the list gives the valid species name and author, and distributional information. In the species distributions we first give the overall zoogeographic region (see Fig. 1), followed by progressively mote precise locations. For example: Afrotropical: South Africa (Cape Prov.). In addition, the symbols: n - northern, s - southern, w - western etc., are used. They do nor refer to specific place names but to general directional areas within distributions. We have used the zoological regions because most distributions are within a single zoogeographic region.

A final word about the checklists. We ate only too aware that errors may have gone unnoticed in spite of our precautions. A simple calculation of some 4000 entries, and perhaps 10 bits of information each, gives 40000 possibilities for error. We hope that users of the lists will be charitable!

Missing types

For perhaps five per cent of the listed names, the type specimens have been lost or their present location is unknown to us. Some of these pertain to older authors, particularly those who published before 1800, such as Rossi, Scopoli, Pallas, and Forster. After 1800 some types have simply disappeared, sometimes through neglect and insect or mould damage, such as those of Say, Harris, and Shuckard. In mote modern species we have been unable to account for those types of l3uysson (in Andre) (1891-1901), and many of the Radoszkowski types presumably deposited in Krakow, Poland. A small number of types were lost during the Second World War at Hamburg and Dresden through bombing and subsequent fire.

Sub genera and subspecies Our treatment of subgenera and subspecies must be considered quite conservative. With respect to subgenera, our opinion is that species groups are more flexible and nomenclature is not cluttered by them. Therefore we have avoided the formal sub- generic names. In the case of subspecies, many authors have proposed them quite freely in the past. For instance, Chrysis ignita embraces 34 infraspecific names. Some of these are simple synonyms; some are local colour varieties; a few may represent dma1 variation in colour, size, or punctation; some may be bona fide geographical races; and a few may be valid species which have not yet been recognized as such. On this last point, Linsenmaier (1959a) recognized as species the following, which were originally named by various authors as subspecies or varieties of ignites: rutiliventris, japanensis, sinensis, scuipturata, valida, chinensis, pseudobrevitarsis, sparsepunctata, solida, longula, and obtusidens. Other species in the ignita group are nearly as complicated and many other chrysidids are in a similar situation

Taking ignita and its species group as an example, the present confused state can be solved only by study of long series from many localities, abundant rearing recotds, and detailed examination of male genitalia. In the absence of such informarion, we have simply listed all infraspecific names under the particular species without ruling on their validity. The whole subject of synonymy will be a challenge to future workers.



BIOLOGY

The biology of only a small percentage of chrysidid species has been investigated, and even these cases deal mostly with the identity of the host. All known chrysidids are parasites and their presence nearly always results in the death of the host. Therefore they fall into the categories of parasitoids or cleptoparasites. The latter name is appropriate when provisions by the adult host for its larva are ‘stolen’ and consumed.

Bethylids, the sister-group of the chrysidids, are parasitic on moth and beetle larvae. Chrysidids may feed on both of these, but only in a secondary way. The primary host list is much broader: walking stick insects (eggs), sawflies, silk moths, eumenids, bees, sphecids, and masarids. If provisions are included, we can add spiders, true bugs, aphids, thrips, and dead insects (Microbembex provisions). The nature of the primary food source is practically a subfamilial character in Chrysididae. Thus, Amiseginae and Loboscelidiinae attack walking stick eggs. Cleptinae parasitize sawfly prepupae, and Chrysidinae (except Praestochrysis, see below) use aculeate wasp and bee larvae. The secondary items of food (provisions) are much less significant.

As pointed out by Bohart and Kimsey (1982), there is a strong correlation between chrysidid morphology and the nature of the host. Those subfamilies which use harmless or helpless hosts, such as walking stick eggs or sawfly larvae, have four or five flexible, and often rather lightly sclerotized, abdominal segments. The Chrysidinae parasitize hosts which are able to defend their nests with sting and mandibles. In such cases the female chrysidid may have to protect herself while ovipositing in the host nest. These parasites have only three, four, or rarely two, relatively inflexible, and usually heavily sclerotized abdominal segments, and a flat or concave venter. This abdominal structure allows them to roll up into a relatively impervious ball when threatened.

There are two basic host stages attacked by chrysidids. Most species studied oviposit on, or adjacent to, the host prepupa. The egg hatches immediately and the resulting larva consumes the host and provisions, or the larva attaches to the host and only begins to feed when the host molts to the prepupal stage. The second method is apparently the more primitive of the two strategies used by chrysidids. The host has already eaten and physiologically processed the provisions. As a result, the chrysidid probably requires less digestive specialization than it would if it ate the host egg or larvae and provisions, which could be bugs, spiders, caterpillars, pollen balls, and so on.

Chrysidids exhibit a variety of parasitic behaviour, most of which involve oviposition, feeding habits of the larvae, and host selection. These are briefly summarized by subfamily in Table 1.

Cleptinae are parasites of prepupal sawflies in the families Diprionidae and Tenthredinidae. Based on the few studies of their behaviour (Clausen 1940; Gauss 1964; Dahlsten 1961, 1967) certain generalizations can be made. Cleptes search for their hosts cocoons in leaf litter or loose soil. Once a cocoon is located, the female chews a hole in it with her heavy mandibles. The long, robust ovipositor is inserted and the egg is placed on the host. When oviposition is complete, the hole in the cocoon wall is closed with mucilaginous material. The chrysidid larva spins a cocoon within the host cocoon.

Both Amiseginae and Loboscelidiinae are parasites of walking stick (Phasmatidae) eggs. Males are most commonly collected because they frequent low vegetation, often taking up exposed positions on leaves. Females are infrequently seen, probably because they are more likely to be found in leaf litter or in other sheltered areas searching for host eggs. What biological information we have has been clearly summarized by Krombein (1983a). Females nip a small hole in the host egg chorion with their slender mandibles and use the needle-like ovipositor to place an egg inside the host egg.

Major behavioural and structural changes have occurred in the Chrysidinae. These wasps, with one exception, (Praestochrysis) are nest parasites of wasps and bees. They actively enter nests, whether the host is present or not. The adult host is a capable fighter, equipped with powerful mandibles and sting. However, chrysidine morphology allows them to roll up into an impenetrable ball if they happen to be attacked by the host. The rigid cup- like abdomen effectively covers vulnerable intersegmental membranes and leg joints.

As discussed below, host nests can be above or below ground. Chrysidine females penetrate the host cell and either oviposit directly on the host or elsewhere within the cell. Some chrysidids, such as Stilbum cyanurum, Chrysis angolensis, and C. lincea appear to be nest-type specialists and parasitize a wide range of wasp taxa that all build similar mud nests in exposed areas. Others, such as Chrysurissa and Pseudolopyga are apparently host-taxon specific. In a further specialization, Pseudolopyga oviposits on the bugs provisioned by the host, rather than in the nest (as discussed below). Finally, the majority of Praestochrysis are direct parasites of prepupal moths, a secondary reversion to bethylid-like behaviour.

Some ‘case histories’ warrant further detail. These are given below.

A typical amisegine life history was given by D. J. Pirone (in Krombein 1960) who reared Adeiphe anisomorphae from the ova of the walking stick insect, Anisomorpha ferrugitsea (Beauvois), in Georgia. An adult female oviposited in an egg of Anisomorpha after gnawing a hole in the chorion. The puncture was sealed by coagulation of the egg contents. The wasp embryo developed within the fluid of the egg, pupated, and finally emerged by popping off the operculum of the host egg. Oviposition by unmated chrysidid females resulted, as expected, in males only.

Carrillo and Caltagirone (1970) made detailed observations on host-parasite relationships between two species of sphecids, Solierella peckhami (Ashmead) and S. plenoculoides Fox, and a chrysidid, Pseudolopyga carrilloi. The studies were conducted at several sites in central California, and then repeated under laboratory conditions. Results were quite surprising since it was found that the chrysidid oviposited on free-living nymphal bugs of the genus Nysths (Hemiprera Lygaeidae). Two Nysths species were involved, N. raphanus Howard and N. tenellus Barker. The Solierella were seen reconstruct nests in hollow twigs, holes in the ground, or in almond hulls, and provisioned each cell with 4-10 paralyzed Nysius nymphs. Only those nests with one or more bugs bearing Pseudolopyga eggs produced chrysidids. The parasite females searched for and oviposited in first - or second-instar nymphs of Nysius, attaching the egg to the hind gut of the bug. Apparently, the parasite developed beyond the first instar larva only in bugs paralyzed by Solierella and then used as prey. In nests of the wasp the first instar parasite molted to the second instar and emerged from the Nysius. It then sought out the Solierella egg or first instar larva, destroyed it, then fed on its host bug and others adjacent to it. Female Pseudolopyga examined Nysius nymphs before ovipositing. Those previously attacked were released, thus ensuring only a single parasite per bug. This is the only reported case of a chrysidid attacking a free-living host which is secondarily used as prey by a wasp.

An interesting source of competition for Pseudolopyga comes from the sympatric chrysidid, Hedychridium solierellae. This species, also studied by Carrillo and Caltagirone (1970), parasitizes the same species of Solierella as Pseudolopyga. However, it oviposirs directly in the Solierella cells, and the parasite larva eats the host provisions and host, or perhaps the Pseudolopyga egg or larva.

Mocsár (1961) reported observations in Hungary on oviposition by Stilburn cyanurum into the closed mud cell of Sceliphron destillatorium (Illiger). The female wasp was seen to moisten a spot on the dry mud with a droplet from its mouthparts, then probe the spot with its ovipositor. Repeated moistenings and probings finally penetrated the thick mud wall and an egg was deposited inside the Sceliphron cocoon. As the ovipositor was withdrawn, the softened mud closed over the hole, leaving only a wide hollow. The stout and multidentate Stilbum ovipositor has been assumed to be an efficient boring tool and this has not been entirely discredited. However, Móczar advanced the theory that the teeth were used primarily to give ‘adequate support to the ovipositor sunk into the cell’, rather than for boring per se. Berland and Berland (1938) listed hosts of cyanurum as Sceliphron, Eumenes, Chalicodoma, and Megachile, all making exterior nests of mud or plant products. The variety of hosts, from large to small, probably accounts for the great range in size of the parasite.

Piel (1933) studied the biology of Praestochrysis sbanghaiensis in eastern Asia, where it is a direct parasite of the oriental moth, Monema flavescens Walker. The habits of this chrysidid appear to be typical for most, but not all, members of the genus. The wasp attacks the silken cocoon of the prepupal caterpillar when it has already become quite hard. A hole is made by biting out a small piece to allow penetration by the ovipositor. One or more eggs are laid loosely inside the cocoon. Then, the wasp scrapes material from the outside of the cocoon, moistens it with saliva, and plugs the hole. If for some reason the hole is not filled, mould may attack the contents of the cocoon with fatal results. The wasp egg hatches in about two days and the first- stage larva begins to feed on the host caterpillar. In cases where several eggs hatch, battles ensue, and only one larva survives. The mature wasp larva forms its own cocoon inside that of its host, goes through a prepupal stage, and after 6-10 days enters the pupal stage, emerging as an adult several weeks later. Mating takes place soon after. Males live for only about two weeks but females may be active for a month or more.

Bordage (1913) reported on the habits of Praestochrysis lusca as a parasite of Sceliphron. The parasite egg is deposited in the nearly complete mud cell shortly after the Sceliphron egg. After the cell is closed, the first instar larvae of the two engage in combat, after which the survivor feeds on the provisioned spiders. However, if the Sceliphron egg fails to hatch, the Praestochrysis larva dies without touching the food supply!

An ongoing, detailed study of the habits of Ammophila dysmica Menke is being made by). A. Rosenheim (1987) in the Sagehen Creek area of Nevada County, California. Mr Rosenheim has kindly given us permission to summarize some of his findings relating to Argochrysis armilla. The construction of many nests by the host wasp was observed. Frequent abandonment of nests occurred in the presence of armilla, especially when this cleptoparasite was active in the presence of the Ammophila. In a total of 275 nests examined, 71 (25.8 per cent) yielded armilla, with one to as many as six individuals per nest. In two cases both parasite and host fed on the provisioned catetpillars and developed successfully. When both the sarcophagid fly, Hilarella hilarella, and armilla attacked a nest, only the former finally emerged. Oviposition by armilla females took place when they followed the host into the nest during provisioning. These parasites were also commonly observed digging into nest closures, but only rarely did they reach the cell. Nesting Ammophila occasionally chased armilla females but the heavily punctate integument of the latter seemed impervious to attack, and the parasites resumed observation of nesting activities.

The host appeared to be aware of the presence of wasp and/or fly parasites, and undertook exceptional cleaning of the cell once oviposition on the caterpillar prey by armilla, or larviposition in the upper regions of the nest by Hi/arella, had taken place. These cleaning efforts often resulted in ejection of the fly maggots, but were ineffectual in dislodging the armilla egg.

Two interesting points arising from this study contradict previous assumptions. First, when provisions are sufficient, both parasite and host may emerge successfully. Secondly, more than one chrysidid (up to six) may be produced from a single host cell.


Taken from: L.S. Kimsey and R.M. Bohart, 1990. The Chrysidid Wasps of the World.


BIOGEOGRAPHY

The fossil history of the Chrysididae is scant. The only specimens are from upper Cretaceous and lower Tertiary shale and amber, and belong to the Cleptinae, Amiseginae, and Chrysidinae (Elampini) (Table 2). The oldest of these are cleptines. Despite the paucity of fossil information, the data we have supports observations on the biogeography of chrysidids based on extant forms (Table 3), and allows certain generalizations and speculations to be made.


Cleptines are the most primitive and probably the oldest group of chrysidids. They parasitize diprionid sawflies, herbivores on Pinaceae, and nematine tenthredinids, which feed on a wide range of north-temperate plant species. These chrysidids apparently evolved in the Holarctic or Laurasian Region. They managed to penetrate as far south as Argentina, probably in the Tertiary, but were later replaced in the Neotropical Region by Cleptidea. Only a single species of Cleptes, fritzi, remains in South America. For some reason this group never extended into the Afrotropical Region or much of the Orient, presumably because of the absence of appropriate hosts in these areas.

Zoological Region The Amiseginae also originated in the Holarctic Region. They are egg parasites of walking sticks, which must have been widespread in this region during the Cretaceous and early Tertiary. By the Quaternary amisegines were probably well established in the Oriental and Neotropical Regions. Presumably, major changes in climate and repeated glaciation towards the end of the Tertiary and during the Quaternary caused the complete extinction of amisegines in the Palaearctic Region and in western North America.

One of the two most primitive extant genera, Adelphe, occurs in the Americas. However, this scenario does not explain why the other primitive genus, Anachrysis, and the most specialized ones, Afrosega and Leptosega, occur in southern Africa. We have no tecords of Amiseginae in any other part of Africa, including Madagascar. The presence in Africa of one of the most primitive and several of the most specialized genera, and nothing in between, is an enigma that we are unable to explain. Much of the difficulty with this subfamily is caused by the paucity of available information. This group is probably far commoner than indicated by collections. Unfortunately, relatively specialized collecting techniques are required to obtain specimens; ‘pitfall’ and certain types of malaise traps are particularly effective. As a result, general collectors rarely see these wasps.

The Chrysidinae apparently evolved in the early Tertiary. Chrysidines also appear to have had their origins in the Holarctic Region, with the majority of genera still dominant there. Many of the larger genera, such as Chrysis, Chrysura, and Hedychridium have species groups common to both the Palaearctic and Nearctic Regions. In the Western Hemisphere Chrysis is found throughout the Neotropical Region, but in considerably smaller numbers than in the Nearctic. It has been largely replaced by Neochrysis, Ipsiura, Exochrysis, and Pleurochrysis in tropical America. The Afrotropical and Australasian chrysidines appear to have been derived from this Holarctic fauna, with a few notable exceptions which are discussed below.

Madagascan chrysidids are all closely related to those in south-eastern Africa. For example, Stilbum viride is closest to S. cyanurum, Parnopes madecassus to P fisheri, and Chrysis gheudi to C. lincea. This, combined with an apparent lack of endemic genera, suggests that the chrysidid fauna of Madagascar is recently derived. One genus, Chrysidea, is found throughout the Afrotropical, southern Palaearctic, and Oriental Regions. However, the great majority of species occur in Madagascar, where the genus has undergone considerable radiation.

Australia has a remarkably depauperate chrysidid fauna, and the smallest number of taxa of any of the continents. Although the amisegine genera, Myrmecomimesis and Exova, are endemic in northern Australia, they are clearly related most closely to Oriental groups. The same is true of the other Australian genera, although none are endemic. As with Madagascar, there is one genus, Primeuchroeus, that has undergone tremendous speciation in Australia. Primeuchroeus occurs in the Afrotropical and Oriental Regions, but the majority of species, as well as the least specialized ones, occur in Australia. This genus may very well have evolved in Australia and then radiated north and west. It parasitizes sphecid wasps in the genus Pison. Pison is parasitized by a wide variety of chrysidids, particularly Chrysis in the rest of the world, but apparently only by Primeuchroeus in Australia, where Pison is quite diverse.

There is considerable evidence that the Chrysididae evolved, or at least diversified, after the breakup of Gondwanaland. Much of this can be gleaned from the above discussion. There are no close relationships between the Chilean, Argentinean, and Australian chrysidids. Although Chrysis occurs in both places, the Australian species are clearly more closely related to oriental ones than to any others. The American genera, Pleurochrysis, Exochrysis, Caenochrysis, Ipsiura, and Neochrysis, are endemic in the New World, occurring also in Argentina and Chile. Australia has Stilbum, Praestochrysis, and Primeuchroeus; all Old World genera. No amisegines occur in Chile, or in all but the northernmost part of Argentina. The Australian amisegines are most closely related to other oriental genera. In addition, there are no relationships between species in southern Africa and those of Australia or South America, except the enigmatic African Anachrysis, which is closer to Adelphe in the Americas than to any other extant genera.

As discussed above, chrysidids occur in all zoogeographic regions except Antarctica. We give specific details about distribution under each genus, particularly in the species lists However, there are certain patterns that warrant further discussion.

Most species and/or genera occur within specific zoogeographic regions (Fig. 1). These regions are gross generalizations but they are useful to show global distributions. However, the regional limits that we are using are slightly different from the classical ones of Wallace (1876). These differences more closely reflect the distributions of chrysidid species, and are as follows.

Cuba shows more affinities with North America than with Middle or South America, so we are treating it as part of the Nearctic Region. We use the term Afrotropical for the Ethiopian Region as it more clearly describes this area. In addition, the chrysidid faunas of Ethiopia and most of Somalia show the closest affinities to the Palaearctic, even though chrysidids in the area known as Etitrea, along the coast of Ethiopia, show a relationship with other Afrotropical species. The Middle East, including all of Saudi Arabia, both Yemens, and Oman, is clearly Palaearctic and not Afrotropical in affinity. Other major differences are in the extent of the Oriental Region. For chrysidids the western part of the Oriental Region is bounded by the Great Indian Desert and the southern slope of the Himalayas. The boundary between the Oriental and Australian Regions extends between Celebes and New Guinea and Australia and Timor. There are five basic types of distribution in this family. (1) small, locally endemic genera; (2) widespread, ‘weedy’ species, doubtless transported by human activities; (3) relatively large genera restricted to one, or perhaps two, faunal regions; (4) endemic island faunas; and (5) odd and highly disjunct distributions.

The Chrysididae includes quite a number of small, endemic genera. Most of those that attack aculeate Hymenoptera occur in temperate desert regions, undoubtedly reflecting the greater abundance of Apoidea, Sphecidae, and Eumenidae in these areas. Unfortunately, we lack information about the hosts of most of these, but they are probably endemic as well. chrysidid genera found only in the south-western USA and north-western Mexico are Minymircha, Hedychreides, Microchridium, Argochrysis, Chrysurissa, and Xerochrum. Spintharosoma, Parachrum, and Odontocbrydium occur only in southern Africa. Four genera are restricted to the area extending from North Africa and the Middle East to southern USSR, including Haba, Procbridium, Allochrysis, and Adelopyga. There is only one endemic genus, Gaullca, in South America, and it occurs in the arid thorn scrub of Argentina.

The Amiseginae includes many small and seemingly endemic genera. Much of this apparent endemicity may be due to limited and incomplete collecting, rather than restricted host distribution.

Human activities are clearly responsible for the distribution of a number of weedy’ species. The majority of these are parasites of sphecid and eumenid wasps, particularly Sceliphron and Eumenes, that build mud nests above ground. Nests are commonly built on equipment, ships rigging, and other such structures, and are just as often parasitized. Since many of the host genera are virtually cosmopolitan, and the chrysidids are nest-type specialists rather than host-taxon specialists, some chrysidids have been transported and established widely. Two of the best examples of this are Chrysis angolensis and Stilbum cyanurum. Chrysis angolensis is practically cosmopolitan and S. cyanurum occurs throughout the warmer parts of the Eastern Hemisphere. In both cases this has resulted in a multiplicity of synonyms.

A few species in other genera have been introduced into other regions in a different fashion. Omalus aeneus and Pseudomalus auratus parasitize twig-nesting pemphredonine sphecids, which commonly nest in rose and berry canes. These chrysidids have been introduced to North America in plant material, and have managed to become established as parasites of local or introduced pemphredonine species. A similar situation has occurred with Cleptes semiauratus, which parasitizes the prepupae of several Holarctic genera of sawflies. Cocoons containing prepupae are usually located in litter and debris below the host plant and may be transported with horticultural material.

Although most chrysidids are fully winged and are relatively strong fliers, amisegines and loboscelidiines are not. This has resulted in a large number 0f island endemics, particularly in the Caribbean area and the Oriental Region. Nearly every island that has been well sampled has its own endemic species and sometimes even endemic genera. For example, in the Caribbean every island appears to have one or maybe two unique species of Adelphe. In the Oriental Region many species have flightless females with cryptic habits, which further restricts their dispersal. In addition, their phasmatid hosts may feed on plants with restricted distributions.

For some reason the Philippines have several peculiarly coloured endemic species, which are more closely related to other widespread species than to each other. These are Stilbum chrysocephalum, Chrysis diademata, C. laevicollis, C. igniceps, and Praestochrysis luzonensis. All have a strikingly red head and dark purplish or blue body. We have been unable to find any reasonable explanation for the repetition of this unique colour pattern.

Finally, there are a small number of genera with odd, disjunct distributions. One of these, Cleptes, has been discussed above. Pseudospinolia occurs in the Palaearctic, with one species extending into the northern Nearctic, and one in Chile. These wasps are parasites of the eumenids, Odynerus and Paravespa. The Chilean species, P tertrini, is probably a relict left from the time when Odynerus and Pseudospinolia were more widely distributed in the Americas. Although Odynerus is not now found in South America, the related genus Hypodynerus is endemic in southern South America, and abundant in Chile. It may be the host of P. tertrini. Pseudolopyga has three species in western North America and one in Chile. As in the case of Pseudospinolia, the Chilean species may be a relict.


Acknowledgements
Taken from: R. M. Bohart and L. S. Kimsey. 1982. A Synopsis of the Chrysididae in America North of Mexico.

ACKNOWLEDGMENTS This study of North American Chrysididae began in 1960 with a visit by the senior author to type depositories in Europe and United States. Since that time many individuals have sent material to aid in the work of one or more researchers at the University of California, Davis: A. D. Telford, D. S. Horning, Jr., L. D. French, L. S. Mem. Amer. Ent. Inst., no. 33, 1982 Kimsey and R. M. Bohart. In addition, staff, students and former students at this institution have donated personal collections to the UCD Museum. As a result there has been a nucleus of some 25, 000 North American specimens to which must be added an equal number identified and returned to various institutions. We would like to express our appreciation to the many cooperators who are listed below in three categories, and apologize to those who were unwittingly omitted. 1. Type depositories and associated personiel: ANSP: Academy of Natural Sciences, Philadelphia (S. Roback, C. E. Dunn, M. G. Emsley). Berlin: Zoologisc hes Museum der Humboldt- Universitat, Berlin (late E. Konigsmann, A. Christoph). BMN}l: British Museum, Natural History, London (M. C. Day, C. R. Vardy). Brussels: Institut Royal des Sciences Naturelles de Belgique, Brussels (G. Fagel). Budapest: Hungarian Natural History Museum, Budapest (L. Moczar, J. Papp). Buenos Aires: Argentine National Museum, Buenos Aires (M. J. Viana). CAS: California Academy of Sciences, San Francisco (P. H. Arnaud, Jr.). CNC: Canadian National Museum, Ottawa (L. Masner). Copenhagen: Universitetets Zoologiske Museum, Copenhagen (B. Petersen). Cornell: Cornell University, Ithaca (L. L. Pechuman). Geneva: Museum d’Histoire Naturelle, Geneva (C. Besuchet). Genoa: Museo Civico di Storia Naturale, Genova (D. Guiglia). Invrea Coll.: Private Collection of F. Invrea, now in care of Invrea family, Genoa, Italy. KU: University of Kansas, Lawrence (G. W. Byers). Leiden: Ryksmuseum van Natuurlyke Historie, Leiden (I. C. van Achterberg, J. Van der Vecht). Lund: Lunds Universitets Zoologiska Institution, Lund (H. Anderson). MCZ: Museum of Comparative Zoology, Cambridge (M. Thayer). Museo Argentino de Ciencias Naturales, Buenos Aires (M. J. Viana). Paris: Museum National d’Histoire Naturelle, Paris (S. KeinerPillault). Quebec: Laval University, Quebec (J. M. Perron). UCB: University of California, Berkeley (J. Powell, J. Chemsak, E. G. Linsley). UCD: University of California, Davis (R. 0. Schuster). UCR: University of California, Riverside (S. Frommer). USU: Utah State University, Logan (W. Hansen, G. E. Bohart, F. D. Parker). USNM: National Museum of Natural Science, Washington (K. V. Krombein, A. S. Menke). Vienna: Zoologischen Abteilung des Naturhistorisches Museums (M. Fischer). 2. Institutions furnishing material for identification, and cooperating individuals: American Museum of Natural History, New York, N.Y. (J. Rozen, M. Favreau); Arizona State University, Tempe (F. F. Hasbrouk); Bernice P. Bishop Museum, Honululu (G. M. Nishida); Brigham Young University, l1rovo, UT (S. L. Wood); California State Department of Agricuhure, Sacramento (M. Wasbauer). Bohart and Kimsey: Chrysididae 3 Canadian Department of Agriculture, Lethbridge (G. A. Hobbs); Carnegie Museum of Natural History, Pittsburgh, PA (G. Wallace); Chicago Museum of Natural History, Chicago (R. L. Wenzel); Clemson University, Clemson, SC (J. H. Cochran); Colorado State University, Fort Collins (H. E. Evans): Entomology Research Division, Moorestown, NJ (H. W. Allen); FSCA: Florida State Collection of Arthropods, Gainesville (H. V. Weems, Jr., E. E. Grissell, L. A. Stange); Fresno State University, Fresno, CA (D. J. Burdick);Illinois Natural History Survey, Urbana (W. E. LaBerge, M. E. Irwin); Indiana University, Bloomington (F. N. Young); Iowa State University, Ames (J. L. Laffoon); Kansas State University, Manhattan (H. D. Blocker); LACM: Los Angeles County Museum, Los Angeles (B. R. Snelling); Long Beach State University, CA (R. J. Hampton, E. L. Sleeper); Louisiana State University, Baton Rouge (L. D. Newson); Michigan State University, E. Lansing (R. L. Fischer); Montana State University, Bozeman (N. L. Anderson, S. Rose); Nevada State Department of Agriculture (B. C. Bechtel); New Mexico State University, Las Cruces (J. R. Zimmerman, G. S. Forbes); New York State University, Syracuse (F. Kurczewski); North Carolina State University, Raleigh (T. B. Mitchell); Northern Iowa University, Cedar Falls (J. C. Downey); Ohio State University, Columbus (C. A; Tripiehorn); Oklahoma State University, Stiliwater (D. C. Arnold); Oregon State Department of Agriculture, Salem (R. Westcott); Oregon State University, Corvallis (G. Ferguson); Pacific Union College, Angwin, CA (L. E. Eighme); Pennsylvania State University, University Park (K. C. Kim); Purdue University, Lafayette, Indiana (J. N. Stibick); Rutgers University, New Brunswick, NJ (J. B. Schmitt); San Diego Natural History Museum, San Diego, CA (D. K. Faulkner); San Jose State University, San Jose, CA (J. G. Edwards, W. E. ‘erguson); South Dakota State University, Brookings (E. V. Balsbaugh, Jr.); Texas A. and I. University, Kingsville, TX (J. E. Gillaspy); Texas A and M University, College Station (H. R. Burke); University of Alabama University (E. A. Cross); University of Alberta, Edmonton (G. E. Ba1l University of Arizona, Tucson (F. G. Werner); University of Arkansas, Fayetteville (E. P. Rouse); University of British Columbia, Vancouver (S. G. Cannings); University of Colorado (U. Lanham); University of Connecticut, Storrs (D. J. Peckman); University of Delaware, Newark (P. P. Burbutis); University of Georgia, Athens (C. L. Smith, P. E. Hunter); University of Idaho, Moscow (W. F. Barr); University of Kentucky, Lexington (L. H. Townsend); University of Louisville, Louisville, KY (C. V. Covell, Jr.); University of Maine, Orono (E. A. Osgood); University of Massachusetts, Amherst (M. E. Smith); University of Michigan, Ann Arbor (T. C. Moore, D. Corvan); University of Minnesota, St. Paul (E. F. Cook); University of Nebraska, Lincoln (P. C. Peterson); University of New Hampshire, Durham (J. C. Conklin); University of Nevada, Reno (R. Rust); University of Oklahoma, Norman (C. Hopla); University of Southern illinois, Carbondale (J. C. Downey); University of Tennessee, Knoxville (A. C. Cole); University of Wisconsin, Madison (L. T. Bayer); University of Wyoming Laramie (D. W. Ribble); Virginia Polytechnic Institute, Blacksburg (‘M. Kosztarab); Washington State University, Pullman (W. J. Turner); 3. Private collectors who furnished material as gifts or for identification: B. W. Brooks, University of Kansas, Lawrence; 4 Mem. Amer. Ent. Inst., no. 33, 1982 Bohart and Kimsey: Chrysididae K. W. Cooper, University of California, Riverside; R. Coville, University of California, Berkeley; D. G. Denning, Walnut Creek, CA; G. B. Fairchild, Gainesville, FL; T. Griswold, Utah State University, Logan UT; G. F. Knowlton, Logan, UT; J. Lyon, Van Nuys, CA; D. R. Miller, USDA, Beltsville MD; R. C. Miller, Auburn, CA; F. D. Parker, USDA, Logan UT; J. Schuh, Klamath Falls, OR; J. W. Stubblefield, University of California, Riverside; W. H. Tyson, Fremont, CA; J. Wiley, Gainesville, FL.

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Hosts · map
FamilyScientific name @ source (records)
Apiaceae  Daucus carota @ PN- (1)
Asteraceae  Conyza canadensis @ PN- (1)
Lamiaceae  Prunella vulgaris @ PN- (1)

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