Introduction
Ongoing Projects
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Revision of the Genera and Species of the Australian Stephanidae
Reclassification of the North American Species
Catalog of the World Species (same link as to the "World Species Catalog")
A New Stephanidae Species from Europe
Phylogenetic Revision
Revision of the Genus Hemistephanus, with methodological considerations
In Progress
Mexican Species
Central American Species
Megischus and Foenatopus from South America
Stephanidae in the World
[Clickable world map!]
North America
Central America
South America
Europe
Africa
Middle East
Far Orient
Philippines
Australia
New Caledonia
Taxonomy & Images
Classification
Subfamilies
Genera
Subfamily Schlettererinae Orfila, 1949
Schlettererius Ashmead, 1900
Subfamily Stephaninae Enderlein, 1905
Stephanus Jurine, 1801
Megischus Brullè, 1846
Hemistephanus Enderlein, 1906
?Parastephanellus Enderlein, 1906
Subfamily Foenatopinae Enderlein, 1905
Diastephanus Enderlein, 1905
Neostephanus Kieffer, 1904
Foenatopus Smith, 1861
Collecting Stephanids
World Species Catalog
Explanation, list of literature I still do not have
Acknowledgements to people who contributed (w/ links to their names)
Access to Database
Bibliography
Aguiar & Sharkov (link)
Aguiar (Hemistephanus) (link)
Funding
CAPES
INTRODUCTION
Stephanid wasps are unusual insects, with a slender and richly sculptured body, highly modified hing legs (leg figure), and a somewhat spherical head, set out on a long neck, and bearing a crown of about five teeth around the median ocellus (head figure).
Overal size ranges from three millimeters, for some Australian species, to almost 10 centimeters, as in the Neotropical Megischus macullipenis (photo).
They have been part of the scientific literature for the past 200 years, but are still considered a poorly known group of wasps. The last revision of the family was done by Elliott (1922). This revision is an excellent compilation of the work done on stephanids up to 1922, but its highly heterogeneous descriptions and highly simplified keys are of very limited use in recognizing stephanid species.
Part of the obscurity that surrounds the family is probably due to the fact that stephanids are not easily spotted in their natural habitat, and are only rarely collected by most of the usual collection methods (click here to further details). Consequently, they are also rare in most collections.
Many collectors have observed stephanids flying around dead trees, and the name of some plant species from which they emerged are eventually mentioned in the label of some museum specimens. However, it is rare to find a host association. The biology is well known only to Schlettererius cinctipes (Cresson), from west USA (Taylor 1967a). This species develops on the larvae of Sirex noctilio (Siricidae), the pine wasp. The discovery of this association caused some surprise, since the most common assumption was that stephanids were parasitoids on Coleoptera (Roman, 1917; Rood, 1951; Townes in Muesebeck et al. 1951). Some time later, Kirk (1975) reported other Sirex species as hosts to S. cinctipes. Nonetheless, reports of Coleoptera hosts have also been published. Braza (1989), searching for parasitoids of the beetle Agrillus sexsignatus (Buprestidae), also a pest in pine trees, found, among others, a species of Foenatopus, for which he does not give further information. Townes (1949) mentions Agrillus kalshoveni as a host to the Philippine Diastephanus leucosticus, and Blüthgen (1953) reports some Xylotrechus (Cerambycidae) species as hosts of the European Stephanus serrator. Fragmentary information is available for a few other stephanids, but to the great majority nothing is known. Oviposition behaviour and general aspect of the larva were described and illustrated by Rodd (1951) for an unidentified species of Parastephanellus.
The only attempt of using stephanids in biological control programs was performed by Taylor (1967b and 1976), in Tasmania. Taylor used S. cinctipes (together with other non-stephanid parasitoids) against Sirex noctilio, the pine wasp, with good results.
STEPHANIDAE IN THE WORLD
The distribution of the Stephanidae has not yet been comparatively discussed, although some degree of endemism for the different genera does seem to occur. Only the genus Megischus seems to have a widespread distribution, while all other genera seem to be more or less restricted to some areas (e.g. Hemistephanus in South America, Parastephanellus in India and Australia, etc).
North America
Basic Reference (s) Townes, 19..
Taxonomy
Valid Genera and Species
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Genera Described Undescribed
Schlettererius ? ?
Megischus ? ?
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Comments. There is a single North American species of Schlettererius (link), which is very easy to recognize.
Some of the species in Megischus can be recognized with Townes (19..), but there are two undescribed species which escaped his attention. These undescribed species are related to M. bicolor, and will run to the respective couplet in Townes' key.
Available illustrations.
S. cinctipes (mating couple, female)
M. bicolor (pronotum)
Central America
Basic Reference (s) Townes, 19..
Taxonomy
Valid Genera and Species
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Genera Described Undescribed
Schlettererius ? ?
Megischus ? ?
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Comments.
Available Illustrations.
South America
Basic Reference (s) Townes, 19..
Taxonomy
Valid Genera and Species
---------------------------------------------------------------------
Genera Described Undescribed
Megischus ? ?
Hemistephanus ? ?
Foenatopus ? ?
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Comments.
Available Illustrations.
Europe
Basic Reference (s) Townes, 19..
Taxonomy
Valid Genera and Species
---------------------------------------------------------------------
Genera Described Undescribed
Stephanus 1 ?
Megischus 0 1
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Comments.
Available Illustrations.
Africa
Middle East
Far Orient
Philippines
Australia
New Caledonia
CLASSIFICATION
The only point of consensus about stephanid classification seem to be its consideration as a distinct family, albeit its relationships with other Hymenoptera families are still obscure. Nonetheless, even the establishment of the family as such was not immediate. The first Stephanidae in the literature was described as "Ichneumon no.193", as a member of the family Ichneumonidae (Zschat 1788), and renamed later as Ichneumon serrator by Fabricius (1798). Fabricius would soon transfer this species to Bracon (1804), thus changing also its family status to Braconidae. Jurine (1807), studying this same species, erected the genus Stephanus (from the Greek stephanus, crown, in reference to the strong spines in the coronal area). The "group", then composed by only two known species, was first considered to be a distinct family by Leach (1815). However, the limits of the family were not precisely defined until the publication of Elliott's revision (1922), who removed some Braconidae species usually considered as Stephanidae by some authors (e.g. Kieffer, 1908).
Since the family was erected, many ideas have been proposed to explain its relationships with other Hymenoptera. Eventually, almost all of them were discarded as more detailed studies appeared in the literature, leaving the correct placement of the family among the Hymenoptera as a problem that is still in need of a solution. Nonetheless, many past and contemporary authors insisted in the idea that the Stephanidae would be part of the Ichneumonoidea (e.g. Townes, 1969; Carlson, 1979; DeSantis, 1980), especially in function of the strong overal similarity with some species of Braconidae and Ichneumonidae. Sharkey & Wahl (1992), however, have demonstrated that the Stephanidae share only one of the autapomorphies of the Ichneumonoidea (loss of 2r-m on the front wing), which also occurs, by convergence, in all major groups of Apoidea, showing that there is no evidence to suport Stephanidae as part of Ichneumonoidea. Alternatively, some authors included Stephanidae in the Megalyroidea (with Megalyridae), on the basis of similarities on head and hind-leg structure (Riek in CSIRO, 1973; Gillott, 1980). This last idea is not new, and was first considered by Ashmead (1900), but is still waiting for a more persuasive proof. Currently, the major tendency is to classify the Stephanidae in an isolated superfamily, the Stephanoidea (e.g. Gauld & Bolton, 1988; Naumann, 1992; Goulet & Huber, 1993).
SUBFAMILIES
Stephanidae subfamilies were originally proposed by Enderlein (1905), and adopted by Orfila (1949, 1956), who respectively erects a new subfamily to place the genus Schlettererius, and provides a key to the recognition of the subfamilies (translated below). Although each subfamily shows a considerably homogeneous group of genera, this taxonomc level has been ignored in most works about Stephanidae, probably because there are not enough genera in each subfamily to make its use practical.
Subfamily Schlettererinae Orfila, 1949
Schlettererius Ashmead, 1900
Subfamily Stephaninae Enderlein, 1905
Stephanus Jurine, 1801
Megischus Brullè, 1846
Hemistephanus Enderlein, 1906
?Parastephanellus Enderlein, 1906
Subfamily Foenatopinae Enderlein, 1905
Diastephanus Enderlein, 1905
Neostephanus Kieffer, 1904
Foenatopus Smith, 1861
GENERA
Discussions about the relationships among the genera of the Stephanidae have been inconclusive, and a precise definition for each of them is still not available. All genera in this family have been defined almost exclusively on the basis of characters from the wing venation, while many external characters and nearly all internal characters of the family were never investigated, making the group a promising source of discoveries. In the list below, click on taxon of interst for more detailed information.
Schlettererinae Orfila, 1949
Schlettererius Ashmead, 1900
Stephaninae Enderlein, 1905
Stephanus Jurine, 1801
Megischus Brullè, 1846
Hemistephanus Enderlein, 1906
?Parastephanellus Enderlein, 1906
Foenatopinae Enderlein, 1905
Diastephanus Enderlein, 1905
Neostephanus Kieffer, 1904
Foenatopus Smith, 1861
Key to Genera? Probably nah...
One page for each genera:
Genus Schlettererius Ashmead, 1900.
Representative taxon Schlettererius cinctipes (Cameron) (link to image)
Number of known species 2
Distribution Norht America, Korea
Diagnosis Townes (1954)
Species recognition Madl (1991)
Status This is the most primitive genus in the family (Townes, with a
single very characteristic species, S. cinctipes, of Nearctic
distribution...
Research in progress 1. Mapping of USA and Australian species (link)
Collecting tips Large specimens, good flyers. Malaise traps seem to have some
success.
Databasing In progress
Genus Stephanus Jurine, 1801.
Type species Stephanus serrator (Fabricius)
Number of known species 1 (+1 undescribed)
Distribution Europe
Diagnosis
Species recognition .
Status
Research in progress 1. New species from Italy
Collecting tips
Databasing In progress
Genus Megischus Brullé, 1846.
Type species Megischus furcatus (LePeletier & Serville)
Number of known species
Distribution
Diagnosis
Species recognition .
Status
Research in progress
Collecting tips
Databasing
Genus Hemistephanus Enderlein, 1906.
Type species
Number of known species 19 (Aguiar, in litt.)
Distribution
Diagnosis
Species recognition .
Status
Research in progress
Collecting tips
Databasing
Stephanus, Megischus and Hemistephanus. These three genera are closely related to each other.
Stephanus was proposed by Jurine (1807), in a somehow unfortunate way, because it was based in an atypical species for the group. Brullé (1846), noticing the peculiarities of the type species of Stephanus, proposed Megischus to all other species in that taxon, leaving S. serrator (Fabricius, 1798) as the only valid species for the original genus. However, only a few authors agreed with this, and most stephanid species were still being described as Stephanus for more than a century after Brullé's proposition. Megischus would be definitely accepted only after the revision of Townes (1954). In the mean time, Enderlein (1906) proposed two new genera (Hemistephanus and Parastephanellus), both of them supported on the basis of venational characters of the front wing. However, no further investigation was conducted to support Enderlein's ideias, and the discussion remained a matter of personal opinion. Townes (1954), for example, considered Hemistephanus as a synonym for Megischus. Aguiar (1998), however, presents new evidences for supporting the genus Hemistephanus, and, at the same time, points out to the current instability of the genus Megischus, which is still a higly heterogeneous and poorly defined group.
Genus Parastephanellus Enderlein, 1906.
This genus was first proposed as Parastephanus (Enderlein, 1905), including species from the Indo-Australian as well as the Neotropical regions. It was later changed to Parastephanellus (Enderlein 1906) because the original name was preoccupied by Haeckel for Protozoa. Enderlein also transfered the Neotropical species to a new genus, Hemistephanus. Most Parastephanellus specimens can be still recognized by Enderlein's original diagnosis, and thus seem to form a considerably stable genus. Although some species can be eventually confounded with Hemistephanus, the geographic distribution of the two groups are distinct enough to prevent such mistake with labelled specimens. However, whether Hemistephanus is trully related to Parastephanellus or not has never been proved, and the monophily of the genus Parastephanellus itself was never assessed.
Type species
Number of known species
Distribution
Diagnosis
Species recognition .
Status
Research in progress
Collecting tips
Databasing
Genus Diastephanus Enderlein, 1905.
Type species
Number of known species
Distribution
Diagnosis
Species recognition .
Status
Research in progress
Collecting tips
Databasing
Genus Neostephanus Kieffer, 1904.
Representative taxon
Number of known species
Distribution
Diagnosis
Species recognition .
Status
Research in progress
Collecting tips
Databasing
Genus Foenatopus Smith, 1861.
Type species
Number of known species
Distribution
Diagnosis
Species recognition .
Status
Research in progress
Collecting tips
Databasing