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Trigonalini Cresson 1887
Type genus: Trigonalys Westwood 1835
Diagnosis
Body shape variable but never appearing slender and delicate. Antennae generally elongate and filiform; tyloids present, elongate-linear or oval to round, usually on more than 5-6 flagellomeres; genal carina reaching or ending immediately before mandibular base; propodeal foramen 'V' shaped, sharply angled at apex; metasomal plates strongly sclerotised; female armature, if present, on sternum II or III, only rarely both; male sternum II either flattened, concave, or convex, if flattened or convex may have a small bump or projection anteriorly; parameres variable but generally elongate.
Discussion
The Trigonalini includes Taeniogonalos and Trigonalys,
and tentatively, Ischnogonalos. These taxa are generally
stout bodied and males have tyloids, and at least in the groundplan,
female metasomal armature. Though host data are very limited,
none are known to parasitise aculeate wasps. Taeniogonalos
and Trigonalys are easy to distinguish from each
other by the shapes of their heads (Figs 22-23). Ischnogonalos
may be distinguished from them by its petiolate metasoma and thickened
flagellomeres (Schulz, 1907a, 1908). Several genera, previously
differentiated by variable characters, are synonymised under Taeniogonalos
and Trigonalys.
Ischnogonalos Schulz
Ischnogonalos Schulz 1907a: 12. Type species Trigonalys dubia Magretti (by monotypy).
Diagnosis (From Schulz, 1907a, 1908; Magretti, 1897)
Body small, 6 mm long, stature petite, with dense and coarse wrinkles,
low-lustre sheen except vertex which is glossy; pubescence erect;
SAE slightly pyramid-shaped, not connected at midline; middle
flagellomeres swollen, with linear tyloids. Male with bidentate
'armature' on second sternum.
Discussion
Schulz (1908) described the genus as having thin elongate tyloids, and from his illustration it appears typical of Taeniogonalos, except for the narrow petiole and swollen flagellomeres. This species is apparently only known from the holotype, which was apparently deposited in the MCSN and lost (V. Raineri, pers. comm.), and the status of this genus is unclear.
Distribution
Possibly Burma.
Biology
Unknown.
Species included
Ischnogonalos dubia (Magretti
1897).
Taeniogonalos Schulz
Taeniogonalos Schulz 1906b: 212. Type species Trigonalys maculata Smith (by monotypy).
Labidogonalos Schulz 1906b: 207. Type species: Trigonalys ornata Smith (by monotypy). Syn. n.
Poecilogonalos Schulz 1906b: 212. Type species Trigonalys thwaitesii Westwood (by monotypy). Syn. n.
Lycogastroides Strand 1912: 129. Type species L. gracilicornis Strand (by monotypy). Syn. n.
Lycogonalos Bischoff 1913a: 155. Type species L. flavicincta Bischoff (by monotypy). Syn. n.
Nanogonalos Schulz 1906b: 211. Type species N. enderleini De Santis 1980 (by monotypy). Syn. n.
Taiwanogonalos Tsuneki 1991: 35. Type species T. alishana Tsuneki by original designation. Syn. n.
Diagnosis
Entire body punctate, including gena and vertex; length 4.3-13 mm, FW 3.8-11.5 mm. Genal carina ending near outside edge of mandibular base, often dividing or fading into sculpturing at the end; antenna filiform, tyloids linear and parallel-sided, with distinct edges; SAE present but reduced, forming triangular brow over each torulus, not connected to each other nor forming ledge above clypeus; frons between antennae flattened, only slightly angled in side view; intertorulus distance about same as shortest distance from torulus to eye; wings often with smoky or dark markings; metasomal plates strongly sclerotised, meeting laterally with little overlap; female sternum II expanded and often with armature that may or may not be bifid, armature usually does not extend posteriorly over more than half of sternum III, except in New Guinea and Australian species; sternum III without projection or ledge; male sternum II often flattened; parameres usually elongate.
Discussion
This genus is characterised by the parallel-sided, usually elongate tyloids, and reduced SAE that often has a light-coloured mark. The intertorulus area is relatively flattened and the dorsal edge of each torulus is slightly raised: in anterior view the dorsal edge forms a 45-60° angle to the horizontal plane and in dorsal view the dorsal edge of the torulus forms a small projecting triangle (Fig. 17). The metasomal plates are always strongly sclerotised. The armature, if present, is located on the 2nd sternum, and not on the third. Taeniogonalos flavocincta, tentatively placed in this genus, is exceptional in that its main armature is on the third sternum with a distinct protrusion on the second. The frons is not angled either in lateral or dorsal view. The gena is punctate, as is the rest of the body in most specimens.
A number of other taxa can be confused with members of Taeniogonalos. Trigonalys maculifrons superficially resembles the sympatric Taeniogonalos ornata in shape and colour pattern. The shape of the gena distinguishes the two genera (Figs 22, 23). In addition, in T. ornata the basal two-thirds of tergum I is dark brown and the remainder is yellow, the vertex and frons are evenly punctate, and a dark longitudinal line completely divides scutellum into two yellow lateral sides. In Trigonalys maculifrons the basal third or less of tergum I is dark brown and the remainder is yellow, the vertex is smooth, the frons above the antennae is textured, and the dark central line does not reach the anterior edge of the scutellum (i.e. both sides and the anterior third of the scutellum are yellow). Xanthogonalos species are also superficially similar to Taeniogonalos ornata. However, T. ornata females lack armature while Xanthogonalos females have armature on sternum II, and T. ornata males have tyloids, which are lacking in Xanthogonalos males. These species probably all mimic Agelaia species (Vespidae).
As discussed in the introduction, previous workers believed in the stability of armature as a taxonomic character and divided genera into subfamilies based in part on the presence or absence of armature. Poecilogonalos (without armature) was therefore separated from Taeniogonalos (with armature). We are synonymising Poecilogonalos and Taeniogonalos after finding that the armature can vary not only within closely related species, but also within a species, and that the species previously separated into these two genera form part of a monophyletic group that does not warrant subdivision. Taeniogonalos thwaitesii is found from India to Malaya with closely related or the same species found in the Philippines and New Guinea. Specimens from west of Laos have a swollen sternum II but no armature, and east of Laos have sternum II with armature. Both forms are found in Laos (specimens in BPBM). Nanogonalos is also being synonymised under Taeniogonalos. Nanogonalos had been separated from Taeniogonalos by the absence of armature and an elongate body shape. However, these characteristics are insufficient to distinguish genera and Nanogonalos clusters with Taeniogonalos in the phylogenetic analysis.
Tsuneki (1991) described Taiwanogonalos as closely allied to Orthogonalys, but having tyloids. He based this relationship on the similarity of the SAE to that of Orthogonalys, without considering the form of the gena, genal carina, propodeal foramen, or the form of the metasomal terga. Tsuneki (1991) described the seven species of his Taiwanogonalos from eight specimens that Teranishi had identified as Taeniogonalos maga. Five of these specimens, representing four of Tsuneki's species, were collected at the same time and location. We are following Teranishi's determination.
The female of Taeniogonalos flavocincta was previously unknown. Tentatively placed in Taeniogonalos, T. flavocincta shares the critical synapomorphies with the group: tyloids present (Teranishi, 1929), narrow, punctate genae, and typical reduced supra-antennal elevation, but is distinct because its main armature is on sternum III, with only a small projection on sternum II.
There are other sources of confusion in this group. When Schulz (1906b) described the new genus Nanogonalos he used a specimen from Bolivia. In the description he said he initially planned to give the specimen the species name 'enderleini' but before publishing decided the specimen was the same as the Mexican Trigonalys fasciatipennis Cameron. Schulz (1906b) then described Nanogonalos based on the Bolivian specimen, and due to mistaken identity, Cameron's species became the type of the genus. Later, when Schulz (1907b) was able to examine Cameron's syntypes, he realised that it was not the same genus as his Bolivian specimen. Schulz's response was to call the Bolivian specimen Nanogonalos fasciatipennis Schulz. This decision violates article 49 of the International Code of Zoological Nomenclature, which prohibits the re-use of a species-group name wrongly applied through misidentification even if the species are later assigned to different genera. To correct Schulz's error, De Santis (1980) replaced Nanogonalos fasciatipennis Schulz with N. enderleini Schulz. However, as enderleini was at that time a nomen nudum and unavailable De Santis (1980) becomes the author as he has given the species the replacement name. Furthermore, Taeniogonalos fasciatipennis (Cameron 1897) was described from two specimens from separate locations. Since the female is actually T. gundlachii, we are designating the male as lectotype. Females of T. fasciatipennis do not have armature, and the males have a convex sternum.
Synonymy of Lycogastroides under Taeniogonalos is based on the examination of the holotype of the type species, Lycogastroides gracilicornis Strand, and the study of paratypes of Benoit's two species in that genus, all of which are unambiguously Taeniogonalos. Benoit (1950) correctly described Lycogastroides maynei as having the armature on sternum II, but later (1951) incorrectly stated that in Lycogastroides sternum III is armed. Synonymy of the genus Lycogonalos, described from a single male specimen (Bischoff, 1913a), is based on the examination of that specimen. Although this specimen is missing the metasoma, Bischoff (1913a) described the "armature" as located anterior of a flattened area on sternum II, a feature known only from male Taeniogonalos.
We are synonymising T. costalis under
T. gundlachii. Specimens of 'T.
costalis' from North and central America have less
extensive yellow markings than T. gundlachii
from Cuba, but specimens from Florida are intermediate. This is
the only species of Taeniogonalos known in the U.S.
and Canada. In the drawing of the T. gundlachii
(=costalis) head in Townes (1956), the antennae
are too far apart and the SAE is reduced, so that except for the
distinct clypeal suture, the head resembles that of Lycogaster.
Schulz (1907a) synonymised the female Trigonalys
costalis Cresson with the male of the completely
unrelated species Trigonalys pulchella
Cresson, and placed pulchella in the genus Tapinogonalos.
Bischoff (1938) removed costalis from synonymy with
pulchella and placed costalis in Lycogaster.
Townes (1951) moved costalis into Poecilogonalos.
The presence of armature in the female and tyloids in the male
places this species in Taeniogonalos under Schulz's
system as well as in the current classification. Schulz (1907a)
left Trigonalys gundlachii as a species
whose generic placement was doubtful, and Bischoff (1938) and
Weinstein & Austin (1991) placed the species in Lycogaster.
Alayo (1974), recognising the close relation of gundlachii
to costalis, followed Townes (1956) and placed the
species in Poecilogonalos.
Distribution
Taeniogonalos, is the most widely distributed of all trigonalid genera, and is found everywhere trigonalids occur, except Europe and western North America. Most species are from eastern Asia and South America.
Biology
Taeniogonalos species have been reared from a variety
of tachinid and ichneumonid parasitoids of Lepidoptera. Taeniogonalos
thwaitesii has been reared from an ichneumonid parasitoid
of Lygropia quaternalis (Pyralidae)
(USNM) and from cocoons of Henicospilus rufus
(Ichneumonidae) collected in the soil of a tea plantation (USNM,
LACM) (Clausen, 1929). Taeniogonalos gundlachii
has been reared from a tachinid, Nilea lobeliae,
parasitising a noctuid Acronicta lobeliae
(Riley & Howard, 1891; Schulz, 1907a) and another tachinid,
Lespesia sp. from a noctuid, Phosphila
turbulenta (Carlson, 1979). It has also been reared
from unidentified tachinids from Anisota senatoria
(Saturniidae)(TAMU) and Megalopyge opercularis
(Megalopygidae)(NCSU). A specimen from North Carolina (NCSU) has
the label information that it was reared from a larva of Apantesis
anna (Arctiidae). In Costa Rica T.
gundlachii has been reared from both tachinid (Lespesia,
Blepharipa, Winthemia, Zizyphomyia,
and Drino) and ichneumonid (Enicospilus
and ?Trogus) parasitoids of various large bodied
Lepidoptera (D.H. Janzen, pers. comm.). In Kansas this species
was reared from the tachinid, Allophorocera arator,
parasitising a detritivore tipulid, Tipula ?flavoumbrosa
(Gelhaus, 1987). Taeniogonalos maynei
was reared from pupae of Latoia albipunctata
(Limacodidae), presumably an intermediate host (Benoit, 1950).
In Australia, Taeniogonalos maculata
and T. venatoria have been reared
directly from pergid sawfly hosts including Perga
condiei, P. dorsalis,
P. nemoralis, and P.
affinis (Carne, 1969; Raff, 1934; Weinstein &
Austin, 1991, 1995a, b). In some cases this parasitism was apparently
host density-dependent, and caused a significant reduction of
the pest sawfly numbers (Carne, 1969). Riek (1962b) reared an
unidentified Taeniogonalos from an ichneumonid parasitising
an anthelid (Lepidoptera) and a male and female T.
?maculata were reared from Panacela
lewini (Eupterotidae, Lepidoptera)(ANIC). The report
of Taeniogonalos directly parasitising geometrid
and tortricid cocoons (Weinstein & Austin, 1991) was not confirmed
(A. Austin, pers. comm.; I.D. Naumann, pers. comm.).
Species Included
Taeniogonalos chadwicki Riek 1954.
Taeniogonalos enderleini (De Santis 1980). Transferred from Nanogonalos. Comb. n.
Examined 58 males, 36 females from South America (AEIC, CNCI, MCZC, OSUO, NHMW, UCDC, BMNH, PORT, IMLA, CUIC, ISNB, CDFA, ZMHB*).
Taeniogonalos fasciata (Strand 1913). Transferred from Poecilogonalos. Comb. n.
=Poecilogonalos magnifica Teranishi 1929. Syn. n.
Examined 2 males, 18 females from TAIWAN, JAPAN, CHINA, INDONESIA, MALAYSIA (AEIC, NHMW, RMNH, BMNH, CNCI, ZMUC, CARM, CMNH, TARI).
Taeniogonalos fasciatipennis (Cameron 1897). Type material examined: desig. lectotype n.:
Female syntype of T. fasciatipennis examined, = T. gundlachii. (BMNH). Examined: 11 males, 9 females from MEXICO (BMNH*, CUIC, CNCI, EMUS). Same or closely related species from COSTA RICA and HONDURAS (4 females; USNM, FSCA, PORT, CARM).
Taeniogonalos flavicincta (Bischoff 1913a). Transferred from Lycogonalos. Comb. n.
Holotype examined, male, no locality (ZMHB*).
Taeniogonalos flavocincta (Teranishi1929). Tentative placement, transferred from Nanogonalos. Comb. n.
Examined male, female, KOREA (USNM, HNHM).
Taeniogonalos formosana (Bischoff 1913a). Transferred from Poecilogonalos. Comb. n.
Taeniogonalos fulvoscutellata (Ayyar 1919). Transferred from Poecilogonalos. Comb. n.
Examined 3 males, 7 females, INDIA (USNM, RMNH, BMNH, OSUO).
Taeniogonalos gracilicornis (Strand 1912). Transferred from Lycogastroides. Comb. n.
Holotype examined, female, EQUATORIAL GUINEA (ZMHB*).
Taeniogonalos gundlachii (Cresson 1865). Transferred from Poecilogonalos. Comb. n.
=Trigonalys costalis Cresson 1867. Transferred from Poecilogonalos. Described in Trigonalys and also placed in Lycogaster. Syn. n.
Numerous specimens from CANADA, COSTA RICA, CUBA, UNITED STATES (MEMU, NCSU, INBIO, TAMU, USNM, ZSMC).
Taeniogonalos henicospili (Rohwer 1929). Transferred from Poecilogonalos. Comb. n.
Taeniogonalos intermedia (Chen 1949). Transferred from Poecilogonalos. Comb. n.
Taeniogonalos javana (Bischoff 1933). Transferred from Poecilogonalos. Comb. n.
Holotype examined, female, INDONESIA (ZMHB).
Taeniogonalos jucunda (Westwood 1868). Examined 2 females, 'Amazons', COLOMBIA (OXUM*, BMNH).
Taeniogonalos kerala (Ayyar 1919). Transferred from Poecilogonalos. Comb. n.
Taeniogonalos lugubris (Westwood 1868). Examined female, 'Amazons' (OXUM*).
Taeniogonalos maculata (Smith 1851). Examined 19 specimens from Australia (ANIC). Type repository, BMNH.
Taeniogonalos maga (Teranishi 1929). Tentative generic placement, transferred from Poecilogonalos. Comb. n.
=Poecilogonalos yuasai Teranishi 1938 (syn. by Tsuneki 1991).
Holotype examined, female, JAPAN (Ibaraki*).
=Taiwanogonalos alishana Tsuneki 1991
Holotype examined, female, TAIWAN (OMNH*).
=Taiwanogonalos alticola Tsuneki 1991
=Taiwanogonalos claripennis Tsuneki 1991
=Taiwanogonalos laeviceps Tsuneki 1991
=Taiwanogonalos minima Tsuneki 1991
=Taiwanogonalos satoi Tsuneki 1991
=Taiwanogonalos similis Tsuneki 1991
Examined 82 specimens from JAPAN and TAIWAN (AEIC, CNCI, TARI).
Taeniogonalos maschuna (Schulz 1907a). Tentative placement, transferred from Tapinogonalos. Comb. n.
Holotype examined, female, missing head, ZIMBABWE (ZMHB*).
Taeniogonalos maynei (Benoit 1950). Transferred from Lycogastroides. Comb. n.
Examined male paratype, ZAIRE (MRAC).
Taeniogonalos ornata (Smith 1860). Transferred from Labidogonalos. Comb. n.
Examined 19 males, 21 females from COSTA RICA and MEXICO (BMNH*, ZMHB, AEIC, UMMZ, EMUS, UCDC, TMSA).
Taeniogonalos pictifrons (Smith 1860). Described in "Trygonalys" [sic] and here transferred from Lycogaster. Comb. n.
Examined 2 females, INDONESIA [OXUM (syntype); RMNH].
Taeniogonalos raymenti New replacement name.
Taeniogonalos tricolor Rayment 1952
Taeniogonalos rufofasciata (Chen 1949). Transferred from Poecilogonalos. Comb. n.
Taeniogonalos sauteri Bischoff 1913a.
=Poecilogonalos flavoscutellata Chen 1949 (syn. by Tsuneki 1991).
=Taeniogonalos pictipennis Strand 1914 (syn. by Tsuneki 1991).
Examined 7 females from PHILIPPINES, TAIWAN (AEIC, USNM, CASC).
Taeniogonalos schulzi (Bischoff 1933). Transferred from Nanogonalos. Comb. n.
Taeniogonalos semibrunnea (Bischoff 1951). Tentative identification, 3 males, 10 females from AUSTRALIA (MAMU).
Taeniogonalos taihorina (Bischoff 1914). Transferred from Nanogonalos. Comb. n.
Taeniogonalos tenebrosa Riek 1954.
Taeniogonalos thwaitesii (Westwood 1874). Described in Trigonalys and here transferred from Poecilogonalos. Comb. n.
Examined 9 males, 30 females from INDIA, SRI LANKA, THAILAND, LAOS, MALAYSIA, PAPUA NEW GUINEA, TAIWAN (BMNH, BPBM, USNM, LACM, OSUO, AEIC, OXUM*).
Taeniogonalos tricolor (Chen 1949). Transferred from Poecilogonalos. Comb. n.
Examined 6 females, CHINA, THAILAND, KOREA (USNM, HNHM).
Taeniogonalos venatoria Riek 1962. 215 specimens from AUSTRALIA (ANIC, AEIC).
Taeniogonalos zairensis (Benoit 1950). Transferred from Lycogastroides. Comb. n.
Examined female paratype, ZAIRE (MRAC).
Taeniogonalos zimmeri (Bischoff 1933). Transferred from Lycogaster based on male having tyloids (Bischoff 1933). Comb. n.
Unplaced specimens: Taeniogonalos sp. 1: ARGENTINA,
2 males, female,
(IMLA, AEIC). Taeniogonalos sp. 2: male,
PAPUA NEW GUINEA (UCDC). Taeniogonalos sp. 3: female,
INDONESIA (RMNH). Taeniogonalos sp. 4: 21 males,
16 females, PAPUA NEW GUINEA (ANIC).
Trigonalys Westwood
Trigonalys Westwood 1835: 52. Type species T. melanoleuca Westwood (by monotypy). Trigonalos Schulz 1906a, emended without justification; Trigonalis Spinola 1840, incorrect subsequent spelling.
Stygnogonalos Schulz 1907b:305. Type species Trigonalys championi Cameron (by monotypy). Syn. n.
Discenea Enderlein 1905: 200. Type species Trigonalys natalensis Kreichbaumer (by monotypy). Syn. n.
Lycogastrula Strand 1912: 130. Type species L. micanticeps Strand (by monotypy).
Stygnogonaloides Strand 1912: 128. Type species S. crassiceps Strand (by monotypy).
Diagnosis
Length 6.2-13 mm, commonly 8-10 mm, FW 4.9-10.6 mm. Occiput strongly
excavated all the way to mandibular base; genal carina extending
straight and unbranched to lateral edge of mandibular base without
getting smaller; genal carina on genal angle (Fig. 22), area around
the end of carina smooth, not rough; gena wide in side view and
shining behind lower part of eye; malar space short; antennae
elongate and filiform; tyloids small, rounded and often glistening;
intertorulus distance about equal or a little less than distance
from torulus to inner margin of eye; lower face above clypeal
suture and area between antennae not punctate, usually shining;
frons sharply angled in side view, with slight carina between
antennae; female armature present or absent, if present generally
on sternum III, rarely on II or II & III; female tergum V
often hoodlike, with flange over posterior segments; male metasoma
rounded ventrally.
Discussion
Members of this genus are the only trigonalids that have the genal carina located on the genal angle, and have the occiput strongly excavated all the way to the mandibular base (Fig. 22). The SAE join together forming a ledge between the base of the antennae and the area between the ledge and the clypeal suture is shiny, as is the gena.
A number of Trigonalys species resemble those of other genera. Trigonalys flavescens, with its smooth metasoma, orange colour and elongate shape, superficially resembles Orthogonalys. However, in T. flavescens the genal carina ends at the mandible base while in Orthogonalys the genal carina fades before reaching the hypostomal carina, and in Trigonalys the head is more hemispherical. Trigonalys maculifrons can be separated from the superficially similar Taeniogonalos ornata by the generic differences in head shape and the slight differences in colour patterns described in the discussion of Taeniogonalos, the key to New World genera, and figures 22 and 23.
We are synonymising Discenea and Lycogastrula with Trigonalys. These genera were erected for species known only from Africa, but these species share the generic characteristics of Trigonalys. Bischoff (1951) argued for the synonymy of Lycogastrula within Discenea while Benoit (1950, 1951) placed Lycogastrula in the subfamily Lycogastrinae and erected the subfamily Disceneinae. Weinstein & Austin (1991) recognised Benoit's subfamily Disceneinae but not Benoit's acceptance of the genus Lycogastrula.
Trigonalys melanoleuca is represented by two syntypes [BMNH, OXUM]. We are designating the BMNH specimen, a female, as lectotype. The wings of the lectotype are spread and it is missing the left hind leg, both middle legs, and the tips of both antennae.
Sharp (1895) illustrated Trigonalys maculifrons with the caption "Trigonalys maculifrons Cam. i.l. Mexico." prior to Cameron's (1897) description of the species. Since the illustration clearly depicts the holotype specimen described in Cameron (1897), Sharp is therefore the legitimate author of this species rather than Cameron.
We have seen three undescribed species allied to Trigonalys championi Cameron represented by four specimens (CNCI, BMNH, EMUS, PAGL). The similar looking Trigonalys melanoleuca has tergum I with the apex rounded (Fig. 39) and the female without armature. Trigonalys championi has tergum I with the apex nearly straight across (Fig. 40) and the female with armature on sternum III. Additional variation within the genus can be seen comparing Figs 41-42.
The figure in Schulz's monograph (1907a) labelled Labidogonalos ornata is actually Trigonalys sanctaecatharinae from South America. In T. sanctaecatharinae, some females have one to three extremely reduced tyloid-like structures, instead of the normal 4-8 tyloids per antenna found in males.
Distribution
Trigonalys is known from South and Central America, subsaharan Africa, and Asia (India and Philippines).
Biology
Although T. melanoleuca is one of the most commonly collected trigonalids, its host is unknown. Trigonalys natalensis has been reared from an unidentified lepidopteran pupa (Schulz, 1910) and T. ?micanticeps has been recorded from a larva of Achaea catacoloides (Noctuidae) (BMNH).
Species included
Trigonalys championi Cameron 1897. Restored from Stygnogonalos. Comb. revived.
Examined 2 females, GUATEMALA, COSTA RICA (BMNH*, USNM).
Trigonalys crassiceps (Strand 1912). Transferred from Discenea. Comb. n.
Examined female, EQUATORIAL GUINEA (ZMHB*).
Trigonalys flavescens (Bischoff 1951). Transferred from Labidogonalos. Comb. n.
Examined 3 males, MEXICO (BMNH*, EMUS, PORT).
Trigonalys lachrymosa Westwood 1874. Restored from Lycogaster. Comb. revived. Examined female, PHILIPPINES (USNM).
Trigonalys maculifrons Sharp 1895. Restored from Labidogonalos. Comb. revived. Examined 5 males, 19 females, MEXICO, COSTA RICA, HONDURAS, GUATEMALA (BMNH*, AMNH, ANSP, LACM, EMUS, MCZC, FSCA, UCDC, USNM).
Trigonalys melanoleuca Westwood 1835. Type material examined: Desig. lectotype n.: female, BRAZIL (BMNH). Condition: missing very tip of both antennae, wings spread, missing left hind and middle legs. 1 female paralectotype, no collection data (OXUM). Also examined 183 specimens from South America including ARGENTINA, BOLIVIA, BRAZIL, ECUADOR, GUYANA, PARAGUAY, PERU, TRINIDAD, VENEZUELA (MCZC, IMLA, BMNH, FSCA, AEIC, CNCI, UCDC, CDFA, and others).
Trigonalys micanticeps (Strand 1912). Transferred from Discenea. Comb. n.
Examined, tentative identification, 6 males, 13 females from CAMEROON, GHANA, IVORY COAST, GABON, ZAIRE, ANGOLA (BMNH, FSAG, MRAC, CNCI).
Trigonalys natalensis Kreichbaumer 1894. Restored from Discenea. Comb. revived. Examined 2 males, 10 females from SOUTH AFRICA, MADAGASCAR, KENYA, ZIMBABWE, ANGOLA (AEIC, BMNH, ZMHB, CNCI, UCDC, MRAC).
Trigonalys rufiventris Magretti 1897. Restored from Lycogaster. Comb. revived. Holotype not found in MCSN (V. Raineri, pers. comm.). Examined male, 2 females, INDIA (MCZC, OSUO).
Trigonalys sanctaecatharinae (Schulz 1907a). Transferred from Labidogonalos. Comb. n.
=Xanthogonalos fasciatus Bertoni 1912, Syn. n.
Xanthic form of T. sanctaecatharinae.
=Xanthogonalos severini Schulz 1907a, Syn. n.
Examined X. fasciatus paratype female, PARAGUAY (USNM). Also examined 81 males, 25 females, 9? from BRAZIL, ARGENTINA, PARAGUAY (MCZC, CNCI, AEIC, UCDC, OSUC, BMNH and others).
Genera not assigned to Tribes (within Trigonalinae)
Jezonogonalos Tsuneki
Jezonogonalos Tsuneki 1991: 32. Type species J. marujamae Tsuneki (by monotypy), repository: OMNH. Described from a single female (not male) without antennae. Tentative placement.
Diagnosis (From Tsuneki, 1991)
Head from above with SAE strongly produced anteriorly, at least as long as wide at base; SAE directly rising from frons (not from indentation); gaster robust, slightly less than twice as long as wide, forewing weakly fasciated.
Discussion
Tsuneki (1991) distinguished Jezonogonalos from Taeniogonalos maga by the slightly more anteriorly produced SAE. The single specimen on which the genus is based is inadequate, and more specimens in better condition are needed to evaluate its generic status.
Distribution
The specimen described by Tsuneki (1991) is from Japan.
Biology
Unknown.
Species included
Jezonogonalos marujamae Tsuneki 1991
J. marujamanae Tsuneki 1991 (variant spelling in original work, J. marujamae chosen by Tsuneki (1992).
Examined female from JAPAN (OMNH*).
Mimelogonalos Schulz
Mimelogonalos Schulz 1907a: 8. Type species M. bouvieri Schulz (by monotypy).
Diagnosis
Body punctate and shiny black, with yellow clypeus and parts of
legs, sometimes spots on the scutellum, dorsellum, and metasoma;
length 5-9 mm, forewing 4.2-7.9 mm. Genal carina meeting mandibular
base; malar space long; antennae with 22-25 segments, almost as
long as body; tyloids broad and elongate, with a dull lead-like
lustre, surface keel shaped; toruli close together: shortest distance
between toruli 0.6-0.9 times less than the shortest distance between
the inner eye margin and the torulus; wings hyaline or yellowed
but without dark markings; mesosoma and metasoma compact, not
elongate; tergum II about as long as following terga together;
parameres are truncate apically; aedeagus rectangular, in side
view broader and more squared at the tip than in any other known
trigonalid. Female without armature, terminal sternum modified
into cylindrical tube that often points straight down out of metasoma.
Discussion
This genus can be recognised by its black body with yellow markings, and shiny metasoma without ventral modifications. Taeniogonalos, the only other genus in Australia, has a dull-coloured and punctate integument, and males have flattened metasomal sterna while females have armature. Taeniogonalos is often reported as black with lighter markings (Chen, 1949; Riek, 1954; Teranishi, 1929; Townes, 1956) but the 'black' is actually fuscous. In Mimelogonalos the metasoma is either all black or black with two dorsolateral yellow spots, while in Taeniogonalos the metasoma is usually transversely striped.
The holotype of Mimelogonalos bouvieri Schulz 1907a was reported as lost and the species redescribed by Riek (1954) but a neotype not designated (article 75, International Code of Zoological Nomenclature). Berland recorded that the holotype was loaned to Schulz and not returned (J. Casevitz-Weulersse, pers. comm.).
Mimelogonalos contained only M. bouvieri until Rayment (1952) described a second species, and Riek (1954) added four others. All six appear closely related. Riek (1954) separated species using characters including the number of antennal segments and the extent of colour markings. These characters vary within a single species in other genera. Since Riek based his species on one or two specimens, and since he was only able to examine two specimens of M. bouvieri, he was unable to assess the extent to which the species varied. Thus, many specimens now available do not fit his key. Based on our study of Riek's Mimelogonalos type material, we have synonymised M. punctulata and M. partiglabra under M. bouvieri. Additional specimens of this rarely collected genus are required to evaluate the status of the remaining species.
Two undescribed females (ANIC, ZMHB) of the smallest trigonalids we have seen may belong to Mimelogonalos. They are from Australia and are about 3 mm long (forewing 2.7 mm). These differ from other Mimelogonalos in that the genal carina ends inside of the mandibular base instead of bending at the end toward the mandibular base, and their mandibles are symmetrical. These differences could be due to their small size. More specimens, especially males, are needed before their relationships can be confirmed.
Distribution
Mimelogonalos species are known only from eastern Australia including Tasmania.
Biology
Unknown.
Species included
Mimelogonalos bouvieri Schulz 1907a.
=Mimelogonalos partiglabra Riek 1954 (syn. n.).
=Mimelogonalos punctulata Riek 1954 (syn. n.).
Examined holotype M. punctulata, female (ANIC); holotype M. partiglabra, female (ANIC) and 10 males, 11 females, all from AUSTRALIA (ANIC, AEIC).
Mimelogonalos minuta (Rayment 1952). Examined male, 7 females, AUSTRALIA (ANIC, BPBM).
Mimelogonalos nigricauda Riek 1954. Examined male, AUSTRALIA (ANIC*).
Mimelogonalos nigrithorax Riek 1954. Examined female, AUSTRALIA (ANIC*).
Undescribed, small species, from Australia, tentatively placed,
2 females (ANIC).
Pseudogonalos Schulz
Pseudogonalos Schulz 1906b: 209. Type species Trigonalis
[sic] hahnii Spinola (by monotypy), repository:
MRSN (Scaramozzino & Pagliano, 1989).
Diagnosis
Body slender, elongate, shiny black; metasoma and head sparsely
punctate, mesosoma punctate to rugose; length 5.5-13.9 mm, FW
4.3-9.4 mm. Genal carina meeting hypostomal carina with occiput
deeply excavated; malar space short; SAE large vertical ligulate
lobes whose upper and mesal parts are rounded, smooth and shiny
while lower and distal portions form the torulus; antennae very
close together: shortest distance between toruli 0.4-0.7 X the
shortest distance between inner eye margin and torulus; tyloids
present, round to slightly ovoid; terga slightly overlapping sterna
(not meeting laterally); tergum II longer than tergum III but
not as long as all the following terga together; paramere as wide
as basiparamere and 1.4 times as long as wide; basiparamere slightly
longer (1.3X) than paramere; aedeagus similar to Orthogonalys,
cylindrical with a capitate tip; female terminalia without armature
and sternum not expanded; terminal sternum in female pointing
backwards, not strongly sclerotised or sharply pointed.
Discussion
The SAE in the shape of a thin anteriorly projecting lobe mesad of the torulus and the fasciate forewing are diagnostic for this genus. The genal carina ending at the hypostomal carina, with the occiput deeply excavated at the genal carina posteriorly but flattened near the hypostomal carina are also distinctive features. The relationships of the genus are obscure: the tyloids are closest in shape to Trigonalys, but that genus has the SAE meeting at the midline and the genal carina going straight to the mandibular base. The terga are thickened and do not wrap around ventrally as is characteristic of Orthogonalys.
Oehlke (1983) recognised the name Trigonalis Spinola
1840 as a generic name distinct from Trigonalys
Westwood 1835, making Pseudogonalos a junior synonym
of Trigonalis. Scaramozzino & Pagliano (1989)
pointed out that Trigonalis was a misspelling of
Trigonalys. Spinola (1840) attributed Trigonalis
to Klug, indicating his intent was not to describe a new genus,
and later noted his error (Spinola, 1841). Trigonalis
is an "incorrect subsequent spelling" (International
Commission on Zoological Nomenclature, 1985, article 33 section
c) and is not an available name. Thus the name Pseudogonalos
stands.
Distribution
Pseudogonalos hahnii is known from
Europe and parts of Asia including Siberia. Reports of this species
from Japan are actually of the genus Teranishia
(Tsuneki, 1991). Pseudogonalos harmandi
was known from a single specimen from Darjeeling, north-eastern
India (Schulz, 1907a), now apparently lost.
Biology
Pseudogonalos hahnii lays eggs on
the exterior of foliage (Bischoff, 1936b) and has been reared
from Diprion similis (Diprionidae)
(CNCI), and ichneumonids parasitising Lepidoptera (Bischoff, 1938).
Other host records are unconfirmed. Reports that P.
hahnii parasitises yellowjackets (or their parasitoids)
(Clausen, 1940; Thompson, 1958; Weinstein & Austin, 1991)
apparently started with speculation of Dours (1873) that was never
confirmed (Bischoff, 1936a, b; Reichert, 1911). The reports (Rayment,
1948; Weinstein & Austin, 1991) of this palearctic species
as a parasitoid of the neotropical Polistes lanio
are clearly improbable and stem from a misreading of Sharp (1895).
Again, through a misreading of the original text, Weinstein &
Austin's (1991) citation of Popov (1945) reporting that P.
harmandi parasitises ichneumonids is erroneous.
Species included
Pseudogonalos hahnii (Spinola 1840). Examined ca. 90 specimens from Europe and Asia Minor (ZSMC, MNHN, CNCI, FSAG, TMSA, CASC, NHMW, PAGL).
Pseudogonalos harmandi Schulz
1907a.
Teranishia Tsuneki
Teranishia Tsuneki 1991: 15. Type species T. nipponica Tsuneki (by monotypy), repository: OMNH.
Diagnosis
Body slender, elongate, head and metasoma shiny black, thorax
deeply punctate; metasoma fusiform; length 8 mm [10-12 mm (Tsuneki,
1991)] FW 6.8 mm. Genal carina ending before hypostomal carina;
gena shiny with sparse long white pubescence emerging from punctures;
malar space narrower than or approximately equalling width of
first flagellomere; antennae long, filiform, with 25 [24-27 (Tsuneki,
1991)] segments, not banded; toruli closer together than shortest
distance between torulus and inner eye margin; SAE strongly lobed
and pointing anteriorly, dorsal margin slightly indented; FW lightly
fasciated behind stigma, darkest anteriorly at either side of
2r-rs; submarginal cell 2 rarely petiolate; propodeal foramen
'V' shaped with a 'U' shaped carina around it (similar to Pseudogonalos);
terga thin, wrapping laterally but not ventrally; tergum II slightly
longer than III; terminal female sternum tightly folded and pointing
posteriorly, not strongly sclerotised or awl-like.
Discussion
According to Tsuneki (1991), who described Teranishia from a series of 9 males and 7 females, this genus has a lobed SAE on the frons similar to Pseudogonalos but lacks tyloids. We have not seen males in this genus but the females do appear very close to Pseudogonalos hahnii. Females in the genus Teranishia Tsuneki appear to share many primitive traits with Orthogonalys, but more importantly, share derived character states with Pseudogonalos. As males of Teranishia were not available for this study we were only able to tentatively evaluate the phylogenetic position of this genus.
Distribution
The specimens described by Tsuneki (1991) is from Japan.
Biology
Unknown.
Species included
Teranishia nipponica Tsuneki
1991. Examined 2 females from
JAPAN (AEIC, CNCI).
Acknowledgments
The people in the Bohart Museum, University of California, Davis,
(Richard Bohart, Guolin Chen, Steve Heydon, and Phil Ward) helped
us on a daily basis. Early encouragement for this study came from
Lubomir Masner, A. P. Rasnitsyn, and David R. Smith. Seiki Yamane
entrusted his personal types and undescribed specimens. We are
indebted to all the people mentioned earlier who loaned their
specimens. We wish to especially thank those that gave us access
to type specimens: Donald Azuma (ANSP); E. De Coninck (MRAC);
E. Diller (ZSMC); Laraine Ficken, Ian Gauld and Tom Huddleston
(BMNH); Bert Gustafsson (NHRS); F. Koch (ZMHB); Ian Naumann (ANIC);
Chris O'Toole (OXUM); J. Papp (HNHM); David R. Smith (USNM). Micki
Yuval did most of the illustrations. Phil Ward did the scanning-electron
micrographs 10-12 and L. Coote, with support from D.C. Darling
and his NSERC operating grant, did 1-3 and 13. Erika Allen beautifully
translated some of the very difficult older German works. Melissa
Bennett, Mary Berbee, Martin Dieterich, Brian Fisher, Guolin Chen,
and Kei Majima helped with additional translations. Laurence Mound
and Andrew Austin gave important suggestions for improving the
manuscript. H.D. Cameron, Professor of Classical Languages at
University of Michigan, helped with the stem of the family name.
This work was supported in part by University of California, Davis
Jastro-Shields and Natural Reserve System grants.
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