Research Article |
Corresponding author: Hannes Baur ( baur.hannes@gmail.com ) Academic editor: Michael S. Engel
© 2015 Hannes Baur.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Baur H (2015) Pushing the limits – two new species of Pteromalus (Hymenoptera, Chalcidoidea, Pteromalidae) from Central Europe with remarkable morphology. ZooKeys 514: 43-72. https://doi.org/10.3897/zookeys.514.9910
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Two new species, Pteromalus briani sp. n. and P. janstai sp. n., with unusual characters are described from the Central Plateau and the Alps in Switzerland, respectively. P. briani sp. n. is remarkable in that it has the metatibia quite abruptly expanded before the middle. This type of modification of the hind tibia is unique within the Pteromalidae and probably also the entire Chalcidoidea. It is also very rare in other parasitic wasps, where it is suspected to be associated with pheromone glands. The species is a gregarious endoparasitoid of pupae of Vanessa atalanta (Linnaeus) and Aglais urticae (Linnaeus), two common butterflies (Lepidoptera: Nymphalidae) in Europe. It is furthermore a koinobiont parasitoid ovipositing in an early larval stage of the host. The other species, P. janstai sp. n., shows a flattened mesosoma. A dorsoventrally depressed body is a unique feature within the genus Pteromalus, but known from a number species in unrelated genera and subfamilies. The two records demonstrate that it is possible to discover entirely new species with extraordinary characters even in one of the taxonomically most thoroughly explored parts of the world.
Systematics, taxonomy, thorax, morphometry, distance measurements, P. apum , P. bifoveolatus , P. cassotis , P. puparum , P. squamifer , P. vanessae , Pteromalinae , Pireninae , Papilionidae , Pieridae
In Western Europe the vast majority of newly discovered insect species usually belong to complexes of cryptic species. Morphologically, such new species are therefore quite similar to known species and in many cases even rather difficult to separate from those (e.g.,
The genus Pteromalus contains 485 species world wide, with the majority (371 species) having been described from Europe (
Specimens are deposited in the following collections (acronyms mostly according to
Geographical coordinates on data labels of type specimens are indicated as WGS 84 latitude and longitude.
Nomenclature and classification of Chalcidoidea follow
Abbreviation, name, definition, and magnification of the 41 measurements used in the description (see Material and methods for further information).
Abbreviation | Character name | Definition | Magnification in pixel/mm* |
---|---|---|---|
ant.l | Pedicel plus flagellum length | Combined length of pedicel plus flagellum, outer aspect ( |
1742 |
clv.b | Clava breadth | Greatest breadth of clava, outer aspect | 3910 |
clv.l | Clava length | Greatest length of clava, outer aspect | 3910 |
eye.b | Eye breadth | Greatest breadth of eye, lateral view | 3910 |
eye.d | Eye distance | Shortest distance between eyes, dorsal view | 1742 |
eye.h | Eye height | Greatest length of eye height, lateral view | 2549 |
eye.l | Eye length | Length of eye, dorsal view ( |
888 |
fl3.b | First funicular segment breadth | Greatest breadth of first funicular segment (= third flagellar segment), outer aspect | 3910 |
fl3.l | First funicular segment length | Greatest length of first funicular segment (= third flagellar segment), outer aspect | 3910 |
fl8.b | Sixth funicular segment breadth | Greatest breadth of sixth funicular segment (= eighth flagellar segment), outer aspect | 3910 |
fl8.l | Sixth funicular segment length | Greatest length of sixth funicular segment (= eighth flagellar segment), outer aspect | 3910 |
fm3.b | Metafemur breadth | Greatest breadth of metafemur, outer aspect | 3910 |
fm3.l | Metafemur length | Length of metafemur, from distal end of trochanter to tip of metafemur, measured along midline, outer aspect | 1742 |
fwi.b | Fore wing breadth | Greatest breadth of fore wing, measured at about right angle to marginal and postmarginal veins | 1742/1089 |
fwi.l | Fore wing length | Greatest length of fore wing, measured from end of humeral plate to tip of wing | 1089 |
gst.b | Gaster breadth | Greatest breadth of gaster, distance between the outermost lateral edges of the gaster, dorsal view | 1742/2549 |
gst.l | Gaster length | Length of gaster along median line from posterior edge of nucha to tip of ovipositor sheath, dorsal view | 1089 |
hea.b | Head breadth | Greatest breadth of head, dorsal view | 1742 |
hea.h | Head height | Distance between anterior margin of clypeus and anterior edge of anterior ocellus, frontal view | 1742 |
hea.l | Head length | Length of head, dorsal view ( |
888 |
lof.h | Lower face height | Distance between anterior margin of clypeus and lower margin of torulus | 2549 |
mav.l | Marginal vein length | Length of marginal vein, distance between the point at which the submarginal vein touches the leading edge of the wing and the point at which stigmal vein and postmarginal vein unite ( |
2549 |
msc.b | Mesoscutum breadth | Greatest breadth of mesoscutum just in front of level of tegula, dorsal view | 1742 |
msc.l | Mesoscutum length | Length of mesoscutum along median line from posterior edge of pronotum to posterior edge of mesoscutum, dorsal view | 2549 |
msp.l | Malar space | Distance between the point where malar sulcus enters mouth margin and malar sulcus enters lower edge of eye, lateral view ( |
3910 |
mss.l | Mesosoma length | Length of mesosoma along median line from anterior edge of pronotum collar to posterior edge of nucha, dorsal view | 1089 |
ool.l | OOL | Shortest distance between posterior ocellus and eye margin, dorsal view ( |
3910 |
pdl.b | Pedicel breadth | Greatest breadth of pedicel, outer aspect | 3910 |
pdl.l | Pedicel length | Length of pedicel, outer aspect | 3910 |
plc.d | Plica distance | Greatest distance between upper edge of anterior plica | 2549 |
pmv.l | Postmarginal vein length | Length of postmarginal vein ( |
2549 |
pol.l | POL | Shortest distance between posterior ocelli, dorsal view ( |
3910 |
ppd.l | Propodeum length | Length of propodeum measured along median line from anterior edge to posterior edge of nucha, dorsal view | 2549 |
scp.b | Scape breadth | Greatest breadth of scape, outer aspect | 3910 |
scp.l | Scape length | Length of scape exclusive of radicle, outer aspect ( |
2549 |
sct.l | Scutellum length | Length of scutellum along median line from posterior edge of mesoscutum to posterior edge of scutellum, dorsal view | 2549 |
stv.l | Stigmal vein length | Length of stigmal vein, distance between the point at which stigmal vein and postmarginal vein unite apically, and the distal end of the stigma ( |
2549 |
ta3.l | Metatarsus length | Length of metatarsus, including pretarsus | 2549 |
tb3.b | Metatibia breadth | Apical breadth of metatibia, outer aspect | 3910 |
tb3.l | Metatibia length | Length of metatibia, measured along midline, outer aspect | 1742 |
tmp.l | Temple length | Length of temple, dorsal view ( |
888 |
Most characters were measured on photographs taken by Lisa Wilmsmeier with a Leica DFC425 camera mounted on a Leica M16 stereomicroscope. Photographs were taken at different magnifications depending on the size of the character, in order to reduce measurement error for the smaller ones. For all measurements, it was ensured that the points of reference were in perfect focus and that the diaphragm of the lens was fully open. The distances were finally measured using ImageJ, version 1.46v (
I measured three characters, eye length, head length, and temple length, on a single stack photograph taken with a Keyence VHX 2000 digital photomicroscope and a VH-Z20R/W zoom lens at a magnification of 200× (i.e., 1000 µm corresponded to 888 pixels, see Table
I also used the Keyence microscope for making stack-images of qualitative character states. A 4-digit individual code including the notion “Baur” (e.g., “Baur 2410”) was provided for specimens that have been measured or photographed, or used as reference specimens for comparison with newly described species.
I compiled all morphological data in a FileMaker Pro 12®, version 12.0v5, database, of which natural language descriptions as well as ranges of body ratios were generated using the FileMaker script language. Because this is commercial software, a qualitative and a quantitative data matrix (raw values in µm) were exported as comma separated values (CSV) files made available at the BMNH data portal at DOI: https://doi.org/10.5519/0056966. The repository furthermore contains all photographs used for measurements, photographs of reference specimens (sometimes provided by other institutions, see acknowledgments) and of labels of the holotypes of the newly described species.
Holotype ♀ Switzerland, Canton Bern, Köniz, Niederwangen, 570 m, 46.92361°N, 7.37266°E, leg. Jacqueline Grosjean, 28-ii-2004, ex pupa 16-iii-2004, host Vanessa atalanta (Linnaeus, 1758) (Lepidoptera: Nymphalidae), deposited in NMBE (Baur 2129). The host pupa was collected in sheltered cavity of a pedestrian underpass beneath the highway and railway line in Niederwangen. Paratypes 46 ♀ 2 ♂, emerging from the same host pupa as the holotype, deposited in: 2 ♀ BMNH, 2 ♀ CNC, 2 ♀ ETHZ, 5 ♀ JGC, 2 ♀ LUZN, 2 ♀ MHNG, 2 ♀ NHMV, 19 ♀ (Baur 2408, 2414, 2416, 2418–2421, 2423–2426) 2 ♂ (Baur 2139, 2415) NMBE, 2 ♀ SMNS, 2 ♀ UCR, 2 ♀ USNM, 2 ♀ VVC, 2 ♀ ZFMK. Paratypes 6 ♀ Switzerland, Canton Bern, Reichenbach, Kien, 560 m, 46.6132°N, 7.6854°E, v-2008, leg. Rahel Schnidrig, reared from pupa of Aglais urticae (Linnaeus, 1758) (Lepidoptera: Nymphalidae), deposited in: 1 ♀ CNC, 5 ♀ NMBE. According to Schnidrig (pers. comm.), the host was collected as a larva (size about 2.5 mm) and afterwards reared under protected conditions. A total of 40–50 specimens emerged from the pupa but only the paratypes were preserved.
Color: Head and mesosoma: green to blue-green with metallic luster; setae on head and mesosoma: whitish, inconspicuous; tegula: testaceous; setae on callus of propodeum: whitish.
Scape: testaceous; pedicel: testaceous, slightly infuscate dorsally; flagellum: brown.
Fore wing: hyaline; fore wing venation: testaceous; setae on fore wing: fuscous; hind wing: hyaline.
Coxae: green; trochanters: testaceous; femora: testaceous; tibiae: testaceous; tarsi: testaceous with fifth segment slightly infuscate; pretarsi: slightly infuscate.
Petiole: green with purplish tinge; gaster: green; gastral terga: one to five with strong purplish tinge.
Sculpture: Head in frontal view: finely reticulate with relatively high dividing septa; clypeus: finely striate (Fig.
A, D, G Pteromalus briani sp. n. holotype ♀, B, C, E, F, H paratype ♀. A gena, anterolateral view B head, dorsal view C left antenna, outer aspect D mesoscutum, dorsal view E fore wing venation F left metatibia, outer aspect G propodeum, dorsal view H gaster, dorsal view. Arrows mark important character states; scale bars 0.5 mm.
Mesoscutum: finely reticulate, meshes rather high, areoles small and only moderately enlarged in posterior part of sclerite (Fig.
Pro- and mesocoxa: finely alutaceous, metacoxa: finely reticulate.
Median area of propodeum: evenly reticulate, as strong as on mesoscutum (Fig.
Petiole in dorsal view: smooth; gastral terga: smooth and shining, sixth tergum and syntergum alutaceous (Fig.
Shape and structure: Head in frontal view: subtrapezoid; gena in frontal view: rounded; temple in dorsal view: obtuse (Fig.
Antenna (Fig.
Mesosoma in lateral view: moderately strongly bent; propodeum in lateral view sloping with respect to dorsal plane of mesoscutum and scutellum at an angle of: 45 degrees; pronotum breadth with respect to mesoscutum breadth: distinctly narrower; pronotum collar: horizontal, well defined, its length with respect to mesoscutum length: one sixth, its anterior margin: rounded edge; pronotum posterior margin: thin, shiny strip; notaulus: extremely superficial, hardly traceable, reaching: about half along mesoscutum (Fig.
Fore wing (Fig.
Femora: moderately slender; metatibia: quite abruptly expanded before the middle (Fig.
Petiole in dorsal view: conical, in ventral view: open; gaster in dorsal view: ovate, obtusely pointed (Fig.
Length and body ratios: Body length: 2.3–2.9 mm; mesoscutum breadth: 591–806 µm.
Head breadth to height: 1.2–1.41; head breadth to length: 2.02–2.08; head breadth to mesoscutum breadth: 1.26–1.34; lower face height to head height: 0.5–0.58; POL to OOL: 0.76–0.87; eye height to breadth: 1.3–1.36; eye distance to height: 1.74–1.88; temple length to eye length: 0.35–0.44; malar space to eye height: 0.68–0.76.
Pedicel plus flagellum length to head breadth: 0.72–0.87; scape length to eye height: 0.99–1.04; scape length to breadth: 5.24–5.82; pedicel length to breadth: 1.22–1.54; pedicel length to first funicular segment length: 0.84–1.13; first funicular segment length to breadth: 0.91–1.33; sixth funicular segment length to breadth: 0.85–1.04; first funicular segment breadth to clava breadth: 0.85–1.06; clava length to breadth: 2.01–2.57.
Mesosoma length to mesoscutum breadth: 1.5–1.6; mesoscutum breadth to length: 1.57–1.76; mesoscutum length to scutellum length: 1.03–1.1; propodeum length to scutellum length: 0.57–0.62; plica distance to propodeum length: 1.21–1.31.
Fore wing length to breadth: 2–2.18; marginal vein length to stigmal vein length: 1.51–1.68; postmarginal vein length to stigmal vein length: 0.78–1.01.
Metafemur length to breadth: 3.27–4.47; metatibia length to breadth: 5.61–7.82; metatarsus length to metatibia length: 0.65–0.89.
Gaster length to breadth: 1.17–1.62; gaster length to mesosoma length: 0.82–1.11.
Color: Head and mesosoma: bright green to blue-green with metallic luster; setae on head and mesosoma: whitish, inconspicuous; tegula: testaceous; setae on callus of propodeum: whitish.
Scape: testaceous; pedicel: testaceous, slightly infuscate dorsally; flagellum: testaceous, slightly infuscate dorsally.
Fore wing: hyaline; fore wing venation: testaceous; setae on fore wing: fuscous; hind wing: hyaline.
Coxae: green; trochanters: testaceous; femora: testaceous; tibiae: testaceous; tarsi: testaceous with fifth segment slightly infuscate; pretarsi: slightly infuscate.
Petiole: green with purplish tinge; gaster: green; gastral terga: one to three with an indistinct yellowish spot.
Sculpture: Head in frontal view: finely reticulate with relatively high septae; clypeus: finely striate; area between clypeus and malar sulcus: meshes conspicuously enlarged (Fig.
A, B Pteromalus briani sp. n. paratype ♂, C–H Pteromalus squamifer Thomson ♀, from Sweden. A head, ventral view B left antenna, outer aspect C gena, anterolateral view D head, dorsal view E mesoscutum, dorsal view F left metatibia, outer aspect G propodeum, dorsal view H gaster, dorsal view. Arrows mark important character states; scale bars 0.5 mm.
Mesoscutum: finely reticulate, meshes rather high, areoles small and only moderately enlarged in posterior part of sclerite; scutellum: reticulate, meshes about as strong and coarse as on posterior part of mesoscutum, but with a narrow band of smaller areoles in anterior half of median longitudinal line; frenum: reticulate, meshes of similar size to those on scutellum; axilla: reticulate, about as strong as on central part of scutellum; prepectus upper triangular area: reticulate; upper mesepimeron: anteriorly smooth, posterior corner distinctly alutaceous; upper mesepisternum: reticulate, about as strong as on mesoscutum; metapleuron: reticulate, about as strong as on mesepisternum.
Pro- and mesocoxa: finely alutaceous, metacoxa: finely reticulate.
Median area of propodeum: evenly reticulate, as strong as on mesoscutum; inner corner of anterior plica: with a depression, weakly reticulate; nucha: reticulate, as strong as on median area of propodeum; callus of propodeum: reticulate; paraspiracular sulcus: reticulate with few transverse costulae.
Petiole in dorsal view: smooth; gastral terga: smooth and shining, sixth tergum and syntergum alutaceous.
Shape and structure: Head in frontal view: subtrapezoid; gena in frontal view: rounded; temple in dorsal view: obtuse; forming an angle with occiput of: 120 degrees; occipital carina: absent; torulus position with respect to lower ocular line: distinctly above; lower face in lateral view: flat, receding with respect to upper face: weakly, forming an angle of: 35 degrees; scrobe: narrow, rather shallow; malar sulcus: superficial, but traceable; clypeus, anterior margin: widely and shallowly emarginate, without a median depression above emarginate edge; gena near mouth: terete; tentorial pit: distinctly visible; mouth extension: not conspicuously enlarged (Fig.
Antenna (Fig.
Mesosoma in lateral view: moderately strongly bent; propodeum in lateral view sloping with respect to dorsal plane of mesoscutum and scutellum at an angle of: 50 degrees; pronotum breadth with respect to mesoscutum breadth: distinctly narrower; pronotum collar: horizontal, well defined, its length with respect to mesoscutum length: one sixth, its anterior margin: rounded edge; pronotum posterior margin: thin, shiny strip; notaulus: extremely superficial, hardly traceable, reaching: about half along mesoscutum; scutellum in lateral view: moderately convex; scutellum in posterior view: moderately convex; scutellum posterior margin projection: level of anterior margin of dorsellum; scutellum posterior margin in posterior view: narrowly emarginate in the middle; frenal line: finely indicated, especially on sides; prepectus upper triangular area: separated by a fine oblique carina; upper mesepimeron: strongly narrowing below, not reaching base of mesopleuron; anterior plica: bent inwards in anterior two fifths and strong; posterior plica: present, joining anterior plica; orientation of posterior plicae: almost parallel; median carina of propodeum: weakly indicated, irregular; nucha: elevated but not clearly differentiated from median area of propodeum; spiracle: oval, size: small, separated from anterior margin of propodeum by: shortest diameter; callus pilosity: numerous long setae; paraspiracular sulcus: narrow and deep.
Fore wing apex with respect to apex of gaster when folded back: distinctly exceeding; basal cell number of setae: 6; basal setal line: complete, with: 8 setae; cubital setal line: incomplete, with: 4 setae; costal cell pilosity on dorsal side: bare; costal cell pilosity on lower side: numerous setae in distal half and a complete setal line extending to base; speculum on upper side: bare, widely open below; fore wing disc: rather thickly pilose; marginal setae: present, short; stigma: subcircular, small; uncus: short.
Femora: moderately slender; metatibia: quite abruptly expanded before the middle; metacoxa pilosity, dorsally: bare.
Petiole in dorsal view: conical, in ventral view: open; gaster in dorsal view: ovate; gastral terga: weakly sunken.
Length and body ratios: Body length: 2.7 mm; mesoscutum breadth: 682–684 µm.
Head breadth to height: 1.46–1.47; head breadth to length: 2.02–2.03; head breadth to mesoscutum breadth: 1.3; lower face height to head height: 0.59–0.6; POL to OOL: 0.89–0.96; eye height to breadth: 1.29–1.3; eye distance to height: 1.78; temple length to eye length: 0.39–0.43; malar space to eye height: 0.58–0.61.
Pedicel plus flagellum length to head breadth: 0.84; scape length to eye height: 0.97–0.98; scape length to breadth: 4.89–5.15; pedicel length to breadth: 1.37; pedicel length to first funicular segment length: 0.85–0.96; first funicular segment length to breadth: 1.27–1.57; sixth funicular segment length to breadth: 1.02–1.05; first funicular segment breadth to clava breadth: 0.91–0.98; clava length to breadth: 2.44–3.13.
Mesosoma length to mesoscutum breadth: 1.63–1.64; mesoscutum breadth to length: 1.48–1.5; mesoscutum length to scutellum length: 1.08–1.12; propodeum length to scutellum length: 0.55–0.59; plica distance to propodeum length: 1.21–1.39.
Fore wing length to breadth: 2–2.02; marginal vein length to stigmal vein length: 1.39–1.56; postmarginal vein length to stigmal vein length: 0.84–0.93.
Metafemur length to breadth: 4.23–4.62; metatibia length to breadth: 7.16–7.35; metatarsus length to metatibia length: 0.72–0.74.
Gaster length to breadth: 1.68–1.71; gaster length to mesosoma length: 1–1.01.
Close examination of the expanded metatibia under a stereomicroscope did not reveal any distinctive characteristics compared to the “normal”, i.e. unexpanded, metatibia of the other Pteromalus species. It should be noted that for some of the specimens reared from Aglais urticae the expansion is slightly less abrupt than shown in Fig.
The female of P. briani sp. n. keys out in
Below the most important differences are given for those species with which P. briani sp. n. might be most easily confounded. Because of the difficulty to identify some of them, a rather large number of taxa either related to P. puparum or with similar hosts (Lepidoptera: Papilionidae, Nymphalidae or Pieridae) has been considered.
P. apum (Retzius, 1783): female femora infuscate; reticulation between clypeus and malar sulcus without enlarged meshes; POL greater than OOL; tentorial pit indistinct; antenna inserted less high on face, lower edge of torulus below the middle between anterior margin of clypeus and anterior edge of anterior ocellus; mesoscutum with areoles small and only moderately enlarged in posterior part of sclerite; scutellum in lateral view flattened; metatibia gradually widening towards apex; female gaster acuminate, often more than 1.6 times as long as broad. Source of information: 2 ♀ 2 ♂ from Switzerland in NMBE (Baur 2517–2520), also compared with the key by
P. bifoveolatus (Förster, 1861): female femora infuscate; reticulation between clypeus and malar sulcus without enlarged meshes; POL slightly greater than OOL; tentorial pit indistinct; antenna high on face, lower edge of torulus at about the middle between anterior margin of clypeus and anterior edge of anterior ocellus; mesoscutum with areoles small and only moderately enlarged in posterior part of sclerite; scutellum in lateral view moderately convex; metatibia gradually widening towards apex; female gaster acuminate, often more than 1.6 times as long as broad. In addition, the male of P. bifoveolatus is special in that the mouth is very wide, so that the malar space is much less than 0.1 times as long as eye height (0.58–0.61 in P. briani sp. n.). Source of information: syntype ♂ in NHMV, 2 ♀ 2 ♂ (Baur 2521–2524) from Switzerland in NMBE.
P. cassotis Walker, 1847 (syn. P. archippi Howard, 1889: 1891): female legs except coxae testaceous; reticulation between clypeus and malar sulcus without enlarged meshes; POL about as great as OOL; tentorial pit indistinct; antenna high on face, lower edge of torulus at about the middle between anterior margin of clypeus and anterior edge of anterior ocellus; mesoscutum with areoles small and only moderately enlarged in posterior part of sclerite; scutellum in lateral view moderately convex; metatibia gradually widening towards apex; female gaster acuminate, about 1.25 times as long as broad. Source of information: photographs of lectotype ♀ in BMNH, provided by N. Dale-Skey Papilloud; lectotype ♀ of P. archippi in USNM.
P. fuscipes (Provancher, 1881): The lectotype is deposited in the Laval University, Quebec, Canada (
P. luzonensis Gahan, 1925: female femora infuscate; reticulation between clypeus and malar sulcus without enlarged meshes; POL about as great as OOL; tentorial pit indistinct; antenna high on face, lower edge of torulus at about the middle between anterior margin of clypeus and anterior edge of anterior ocellus; mesoscutum with areoles small and only moderately enlarged in posterior part of sclerite; scutellum in lateral view moderately convex; metatibia gradually widening towards apex; female gaster obtusely pointed, 1.4–1.6 times as long as broad. Source of information: photographs of a syntype ♀ from Luzon, Mount Makiling, provided by the USNM Chalcidoidea type catalog. 5 ♀ 5 ♂ from Assam and Nepal, in BMNH, compared with the original description by
P. melitaeae Dzhanokmen, 1998: female femora infuscate; reticulation between clypeus and malar sulcus without enlarged meshes; POL greater than OOL; tentorial pit indistinct; antenna less high on face, lower edge of torulus slightly below the middle between anterior margin of clypeus and anterior edge of anterior ocellus; mesoscutum with areoles small and only moderately enlarged in posterior part of sclerite; scutellum in lateral view moderately convex; metatibia gradually widening towards apex; female gaster acuminate, about 2.3 times as long as broad. Source of information: 2 ♀ from Switzerland in NMBE (Baur 2525, 2526), compared with a paratype 1 ♀ in BMNH and the English version of the original description by
P. platensis Brèthes in Massini, 1913 (syn. P. caridei Brèthes, 1913: 93, synonymized by
P. platyphilus Walker, 1874: female femora infuscate; reticulation between clypeus and malar sulcus without enlarged meshes; POL greater than OOL; tentorial pit indistinct; antenna less high on face, lower edge of torulus distinctly below the middle between anterior margin of clypeus and anterior edge of anterior ocellus; mesoscutum with areoles small and only moderately enlarged in posterior part of sclerite; scutellum in lateral view moderately convex; metatibia gradually widening towards apex; female gaster obtusely pointed, about 1.3 times as long as broad. Source of information: 1 ♀ from Morocco in NMBE (Baur 2527), det. Z. Bouček 1996.
P. puparum (Linnaeus, 1758): female femora infuscate; reticulation between clypeus and malar sulcus without enlarged meshes; POL slightly greater than OOL; tentorial pit indistinct; antenna high on face, lower edge of torulus at about the middle between anterior margin of clypeus and anterior edge of anterior ocellus; mesoscutum with areoles small and only moderately enlarged in posterior part of sclerite; scutellum in lateral view moderately convex; metatibia gradually widening towards apex; female gaster obtusely pointed, rarely more than 1.6 times as long as broad. Source of information: 3 ♀ 2 ♂ from Switzerland in NMBE (Baur 2528–2531, 2549).
P. puparum vanessae Howard, 1889:
P. semotus (Walker, 1834): female femora infuscate; reticulation between clypeus and malar sulcus without enlarged meshes; POL distinctly greater than OOL; tentorial pits indistinct; antenna less high on face, lower edge of torulus slightly below the middle between anterior margin of clypeus and anterior edge of anterior ocellus; mesoscutum with areoles small and only moderately enlarged in posterior part of sclerite; scutellum in lateral view moderately convex; metatibia gradually widening towards apex; female gaster acuminate, distinctly more than twice as long as broad. Source of information: 1 ♀ from Switzerland in NMBE (Baur 2532), compared with the lectotype ♀ in BMNH.
P. smaragdus Graham, 1969: female legs except coxae bright testaceous [this is in contrast to the original description, where it is stated on p. 494 that the legs have the same color as P. procerus (Graham, 1969) which is said to have the femora infuscate (p. 493)]; reticulation between clypeus and malar sulcus without enlarged meshes; POL slightly greater than OOL; tentorial pit indistinct; antenna high on face, lower edge of torulus at about the middle between anterior margin of clypeus and anterior edge of anterior ocellus; mesoscutum with areoles small and only moderately enlarged in posterior part of sclerite; scutellum in lateral view moderately convex; metatibia gradually widening towards apex; female gaster acuminate, about 1.3 times as long as broad. Source of information: photographs of holotype ♀ in BMNH, provided by N. Dale-Skey Papilloud.
P. squamifer (Thomson, 1878): female legs except coxae testaceous (Fig.
P. varians (Spinola, 1808): female femora varying from infuscate to testaceous; reticulation between clypeus and malar sulcus without enlarged meshes; POL distinctly greater than OOL; tentorial pits indistinct; antenna high on face, lower edge of torulus at about the middle between anterior margin of clypeus and anterior edge of anterior ocellus; mesoscutum with areoles small and only moderately enlarged in posterior part of sclerite; scutellum in lateral view moderately convex; metatibia gradually widening towards apex; female gaster acuminate, distinctly more than twice as long as broad. Source of information: 4 ♀ 1 ♂ from France, Moldavia, and Switzerland in NMBE (Baur 2534–2539), compared with lectotypes of synonyms of P. varians, that is, ♀ P. grandis Walker, 1835 and ♀ P. latipennis Walker, 1835 in BMNH.
P. vopiscus Walker, 1839: female femora infuscate; reticulation between clypeus and malar sulcus without enlarged meshes; POL slightly greater than OOL; tentorial pit indistinct; antenna high on face, lower edge of torulus at about the middle between anterior margin of clypeus and anterior edge of anterior ocellus; mesoscutum with areoles small and only moderately enlarged in posterior part of sclerite; scutellum in lateral view moderately convex; metatibia gradually widening towards apex; female gaster acuminate, often more than 1.6 times as long as broad. Source of information: 2 ♀ from Switzerland, in NMBE (Baur 2540, 2541). Identification originally based on Graham’s (1995) redescription of the species, however, the specimens were later also compared with specimens from Southern France in BMNH identified by Graham himself.
Following the suggestion of the collector of the new species, Jacqueline Grosjean, Pteromalus briani sp. n. is named after Brian Jones, since the V. atalanta pupa was collected on his birthday. The name “briani” is a noun in the genitive case and need not agree in gender with the generic name.
Pteromalus briani sp. n. is a gregarious, primary endoparasitoid of pupae of Nymphalidae (Lepidoptera). Currently, Vanessa atalanta and Aglais urticae are known as hosts but the species is likely to attack pupae of other nymphalids or possibly of related families. About 58–60 specimens emerged from the overwintering pupa of V. atalanta (only 51 ♀, 2 ♂ preserved). According to Rahel Schnidrig (pers. com.) about 40–50 specimens emerged from the pupa of Aglais urticae but only 6 ♀ were preserved. The investigation of Schnidrig suggests a koinobiont life history strategy, because the host was collected in an early larval stage (body length 2.5 mm), which was afterwards protected from further parasitization during captive rearing.
Holotype ♀ Switzerland, Canton Wallis, Kippel, Zend, 2100 m, 46.4069°N, 7.7494°E, 15.07.2005, leg. P. Jansta & H. Baur, 15-vii-2005, on Larch (Larix decidua Mill.), NMBE (Baur 2410). Paratype 1 ♀, same data as holotype, BMNH (Baur 2411). Paratype 1 ♂ Switzerland, Canton Grisons, Samedan, Blais Granda, 2100 m, 46.4412°N, 9.86456°E, 10-viii-1998, leg. H. Baur, NMBE (Baur 2412).
Color: Head and mesosoma: green to blue-green with metallic luster; setae on head and mesosoma: fuscous, inconspicuous; tegula: green; setae on callus of propodeum: whitish.
Scape: fuscous with basal third testaceous; pedicel: fuscous; flagellum: fuscous.
Fore wing: hyaline; fore wing venation: brownish; setae on fore wing: fuscous; hind wing: hyaline.
Coxae: green; trochanters: slightly greenish, testaceous at tips; pro- and mesofemur: green, testaceous in apical quarter, metafemur: green, testaceous in apical sixth; protibia: testaceous, meso- and metatibia: testaceous, medially slightly infuscate; tarsi: testaceous, apical segments slightly infuscate; pretarsi: slightly infuscate.
Petiole: dark purplish; gaster: green to blue-green with metallic luster; gastral terga: one to five with strong purplish tinge.
Sculpture: Head in frontal view: finely reticulate with moderately high septae; clypeus: striate; area between clypeus and malar sulcus: finely reticulate.
Mesoscutum: finely reticulate, meshes moderately high, areoles small and not enlarged in posterior part of sclerite; scutellum: reticulate, meshes about as strong and coarse as on posterior part of mesoscutum; frenum: reticulate, meshes larger than those on scutellum; axilla: reticulate, about as strong as on central part of scutellum; prepectus upper triangular area: weakly reticulate; upper mesepimeron: anteriorly smooth, posterior corner distinctly alutaceous; upper mesepisternum: reticulate, about as strong than on mesoscutum; metapleuron: weakly reticulate, less strong as on mesepisternum.
Coxae: weakly reticulate.
Median area of propodeum: uniformly reticulate, as strong as on mesoscutum; inner corner of anterior plica: with a smooth depression and transverse carinae; nucha: reticulate, as strong as on median area of propodeum; callus of propodeum: weakly reticulate; paraspiracular sulcus: smooth with few transverse costulae.
Petiole in dorsal view: smooth; gastral terga: smooth and shining, third to fifth tergum anteriorly, sixth tergum and syntergum wholly alutaceous.
Shape and structure: Head in frontal view: subtrapezoid (Fig.
A–G Pteromalus janstai sp. n. paratype ♀, H paratype ♂. A head, frontal view B head, dorsal view C left antenna, outer aspect ♀ D mesosoma, lateral view E fore wing venation F propodeum, dorsal view G gaster, lateral view H left antenna, outer aspect ♂. Arrows mark important character states; scale bars 0.5 mm.
Antenna (Fig.
Mesosoma in lateral view: rather flattened; propodeum in lateral view sloping with respect to dorsal plane of mesoscutum and scutellum at an angle of: 20 degrees (Fig.
Fore wing (Fig.
Femora: slender; metatibia: gradually widening towards apex; metacoxa pilosity, dorsally: bare.
Petiole in dorsal view: conical, in ventral view: open; gaster in dorsal view: very elongate and acuminate; gastral terga: strongly convex; posterior margin of first gastral tergum: entire; first gastral tergum reaching: one fourth of gaster; tip of hypopygium reaching: almost three fifths of gaster (Fig.
Length and body ratios: Body length: 3.9–4 mm; mesoscutum breadth: 815–829 µm.
Head breadth to height: 1.35–1.39; head breadth to length: 2.08–2.1; head breadth to mesoscutum breadth: 1.18; lower face height to head height: 0.42–0.44; POL to OOL: 1.19–1.2; eye height to breadth: 1.54–1.65; eye distance to height: 1.4–1.46; temple length to eye length: 0.36–0.45; malar space to eye height: 0.45–0.48.
Pedicel plus flagellum length to head breadth: 1.05–1.07; scape length to eye height: 0.81–0.86; scape length to breadth: 6.05–6.39; pedicel length to breadth: 1.51; pedicel length to first funicular segment length: 0.64–0.68; first funicular segment length to breadth: 1.77–1.83; sixth funicular segment length to breadth: 1.06–1.1; first funicular segment breadth to clava breadth: 0.8–0.84; clava length to breadth: 2.09–2.24.
Mesosoma length to mesoscutum breadth: 1.7–1.71; mesoscutum breadth to length: 1.44–1.52; mesoscutum length to scutellum length: 1.13–1.22; propodeum length to scutellum length: 0.59–0.61; plica distance to propodeum length: 1.2–1.36.
Fore wing length to breadth: 2.24–2.3; marginal vein length to stigmal vein length: 1.7–1.78; postmarginal vein length to stigmal vein length: 0.93–0.99.
Metafemur length to breadth: 3.88–4.47; metatibia length to breadth: 7.19–7.29; metatarsus length to metatibia length: 0.8.
Gaster length to breadth: 5.04–5.35; gaster length to mesosoma length: 1.51–1.52.
Color: Head and mesosoma: bright green to blue-green with metallic luster; setae on head: whitish, inconspicuous, on mesosoma: whitish, inconspicuous; tegula: green; setae on callus of propodeum: whitish.
Scape: fuscous with basal two fifths testaceous; pedicel: fuscous; flagellum: brown.
Fore wing: hyaline; fore wing venation: brownish testaceous; setae on fore wing: fuscous; hind wing: hyaline.
Coxae: green; pro- and mesotrochanter: slightly infuscate, metatrochanter: fuscous; pro- and mesofemur: infuscate, testaceous in apical third, metafemur: green, testaceous on tips; tibiae: testaceous; protarsus: slightly infuscate, meso- and metatarsus: testaceous, apical segments slightly infuscate; pretarsi: slightly infuscate.
Petiole: dark purplish; gaster: green; gastral terga: basal terga with large dark yellow spot.
Sculpture: Head in frontal view: finely reticulate with moderately high septae; clypeus: striate; area between clypeus and malar sulcus: finely reticulate.
Mesoscutum: finely reticulate, meshes moderately high, areoles small and not enlarged in posterior part of sclerite; scutellum: weakly reticulate, meshes less strong and coarse than on posterior part of mesoscutum; frenum: weakly reticulate, meshes larger than those on scutellum; axilla: reticulate, about as strong as on lateral part of scutellum; prepectus upper triangular area: weakly reticulate; upper mesepimeron: anteriorly smooth, posterior corner distinctly alutaceous; upper mesepisternum: reticulate, about as strong as on mesoscutum; metapleuron: weakly reticulate, less strong than on mesepisternum.
Pro- and mesocoxa: finely alutaceous, metacoxa: finely reticulate.
Median area of propodeum: uniformly reticulate, as strong as on mesoscutum but with smaller meshes; inner corner of anterior plica: with a smooth depression and transverse carinae; nucha: reticulate, as strong as on median area of propodeum; callus of propodeum: weakly reticulate; paraspiracular sulcus: smooth with few transverse costulae.
Petiole in dorsal view: smooth; gastral terga: smooth and shining, second to sixth tergum and syntergum alutaceous.
Shape and structure: Head in frontal view: subtrapezoid; gena in frontal view: buccate; temple in dorsal view: obtuse; occipital carina: absent; torulus position with respect to lower ocular line: distinctly above; lower face in lateral view: rather flat, receding with respect to upper face: weakly, forming an angle of: 35 degrees; scrobe: narrow, moderately deep; malar sulcus: superficial, but traceable; clypeus, anterior margin: widely and shallowly emarginate, medially slightly inclined above anterior edge; gena near mouth: terete; tentorial pit: indistinct; mouth extension: not conspicuously enlarged; mandibular formula: ?3-4 (the mandibles are in the single male concealed, but the mandibular formula is most likely the same as in females).
Antenna (Fig.
Mesosoma in lateral view: rather flattened; propodeum in lateral view sloping with respect to dorsal plane of mesoscutum and scutellum at an angle of: about 25 degrees; pronotum breadth with respect to mesoscutum breadth: distinctly narrower; pronotum collar: horizontal, well defined, its length with respect to mesoscutum length: one sixth, its anterior margin: slightly elevated edge, medially carinate; pronotum posterior margin: thin, shiny strip; notaulus: superficial, reaching: two thirds along mesoscutum; scutellum in lateral view: almost flat; scutellum in posterior view: almost flat medially; scutellum posterior margin projection: level of anterior margin of dorsellum; scutellum posterior margin in posterior view: narrowly emarginate in the middle; frenal line: finely indicated, especially on sides; prepectus upper triangular area: ? (the lower part of the prepectus is concealed in the single male, but the character state is likely to be the same as for the females); upper mesepimeron: strongly narrowing below, not reaching base of mesopleuron; anterior plica: present, almost straight in anterior part; posterior plica: present, joining anterior plica; orientation of posterior plicae: almost parallel; median carina of propodeum: anteriorly indicated, effaced posteriorly; nucha: elevated but not clearly differentiated from median area of propodeum; spiracle: oval, size: small, separated from anterior margin of propodeum by: shortest diameter; callus pilosity: relatively sparsely setose; paraspiracular sulcus: narrow and deep.
Fore wing apex with respect to apex of gaster when folded back: not exceeding; basal cell number of setae: with up to 10 setae in distal part; basal setal line: complete, with: 11 setae; cubital setal line: incomplete, with: 4 setae; costal cell pilosity on dorsal side: bare; costal cell pilosity on lower side: with numerous setae in distal half and a complete setal line extending to base; speculum on upper side: bare, widely open below; fore wing disc: moderately thickly pilose; marginal setae: present, short; stigma: subrectangular, small; uncus: short.
Femora: slender; metatibia: gradually widening towards apex; metacoxa pilosity, dorsally: bare.
Petiole in dorsal view: conical, in ventral view: open; gaster in dorsal view: elongate, obtuse; gastral terga: weakly sunken; posterior margin of first gastral tergum: entire; first gastral tergum reaching: slightly less than one third of gaster.
Length and body ratios: Body length: 3.1 mm; mesoscutum breadth: 732 µm.
Head breadth to height: 1.44; head breadth to length: 2.06; head breadth to mesoscutum breadth: 1.17; lower face height to head height: 0.51; POL to OOL: 1.33; eye height to breadth: 1.39; eye distance to height: 1.46; temple length to eye length: 0.38; malar space to eye height: 0.44.
Pedicel plus flagellum length to head breadth: 1.3; scape length to eye height: 0.84; scape length to breadth: 5.42; pedicel length to breadth: 1.28; pedicel length to first funicular segment length: 0.54; first funicular segment length to breadth: 2.08; sixth funicular segment length to breadth: 1.41; first funicular segment breadth to clava breadth: 0.97; clava length to breadth: 3.25.
Mesosoma length to mesoscutum breadth: 1.65; mesoscutum breadth to length: 1.44; mesoscutum length to scutellum length: 1.23; propodeum length to scutellum length: 0.64; plica distance to propodeum length: 1.1.
Fore wing length to breadth: 2.11; marginal vein length to stigmal vein length: 1.75; postmarginal vein length to stigmal vein length: 0.91.
Metafemur length to breadth: 4.3; metatibia length to breadth: 7.04; metatarsus length to metatibia length: 0.79.
Gaster length to breadth: 3.38; gaster length to mesosoma length: 1.19.
The dorsoventrally compressed mesosoma and the shape of the propodeum allowed an easy association of the females with the male even though they were collected in separate localities (about 160 km as the crow flies).
P. janstai sp. n. is distinguished from all known species of Pteromalus species by the following combination of characters: mesosoma strongly flattened; female gaster elongate, laterally strongly compressed, more than 5 times as long as broad.
The female keys out in
Most similar are the following species but they differ – among many other characters mentioned in the description – by a rather more strongly bent mesosoma and a much less elongate female gaster:
P. cyniphidis (Linnaeus, 1758) (syn. P. capreae (Linnaeus, 1761)), P. dispar (Curtis, 1827), P. dolichurus (Thomson, 1878), P. fasciatus (Thomson, 1878), P. pontaniae (Askew, 1985) and P. tereus Walker, 1839. Source of information, beside the keys of
Pteromalus janstai sp. n. is named after Petr Jansta, who collected the female specimens. The name “janstai” is a noun in the genitive case and need not agree in gender with the generic name.
Host unknown. The females of Pteromalus janstai sp. n. were swept on some isolated Larch trees (Larix decidua) in an Alpine meadow. The male was swept in a similar habitat, but it cannot be determined whether it was swept from trees.
Although the two new species are clearly placed within the genus Pteromalus, their morphology and some life history traits are remarkable and merit discussion. The most notable morphological feature concerns the metatibia of P. briani sp. n. Its abrupt expansion in proximal half is unique, not only within the genus but also within the family Pteromalidae and – as far as I can judge – the entire Chalcidoidea. Expansions of tibiae are known from some Pteromalidae, but here they look quite different. For instance, in Spathopus (Pireninae) the metatibia is conspicuously but very uniformly swollen only in males. Furthermore, the mesotibia of males of some Mesopolobus, Pegopus, and Spaniopus (Pteromalinae) differs in that the expansion is accompanied by a flattening or at least lateral compression of the tibia (
The expansion of the metatibia is a very rare phenomenon in parasitoid wasps.
The rearing of the host larva under protected condition suggests that P. briani sp. n. develops as a gregarious, koinobiont endoparasitoid, since the host was allowed to continue its development after oviposition in an early larval stage and was only killed in the pupal stage. This is in contrast to some related gregarious endoparasitoids of Lepidoptera pupa. For instance, P. puparum, a widespread parasitoid of Papilionidae and Pieridae, immobilizes the pupal stage of its host on which the development also takes place (
The other species, P. janstai sp. n., is unique within Pteromalus because of its flattened mesosoma. This trait is reported from species of a number of other genera of Pteromalidae, for instance Macroglenes (Pireninae), Anogmus, Guancheria, Monoksa, Pachyneuron, Platypteromalus, Psilonotus, Rakosina, Syntomopus, and Zdenekiana (Pteromalinae) (
I would specially like to thank Jacqueline Grosjean (JGC) who discovered the new species and presented the majority of specimens to the NMBE. I thank Rahel Schnidrig (Spiez, Switzerland) who provided the specimens of P. briani sp. n. reared from Aglais urticae and information from her unpublished high school exam thesis (“Maturaarbeit, 9. Maturität D”, Gymnasium Thun Seefeld, 2008). I am also grateful to Lisa Wilmsmeier (Bern, Switzerland) for taking measurements of most of the specimens. Petr Jansta (Prague, Czech Republic) discovered the two females of P. janstai sp. n. during a very pleasant joint excursion in 2005 in Valais (Switzerland). Richard Askew (Saint-Marcel-du-Périgord, France), Roger Burks (UCR), Natalie Dale-Skey Papilloud (BMNH), Michael Gates (USNM), Gary Gibson (CNC), Christer Hansson (LUZM), John Huber (CNC), and Veli Vikberg (VVC) checked specimens or provided images of specimens from their collections, which is herewith gratefully acknowledged. I finally thank Elsa Obrecht (NMBE), Roger Burks (UCR), and two reviewers for critical reading of the manuscript and many useful suggestions.
Overview of 41 measurements (in µm) of Pteromalus briani sp. n. and P. janstai sp. n., showing minimum, maximum, mean, and standard deviation (except for P. janstai male with n = 1). For character name and definition, see Table
P. briani, females, n = 11 | P. briani, males, n = 2 | P. janstai, females, n = 2 | P. janstai, male, n = 1 | ||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Character | MIN | MAX | MEAN | SD | MIN | MAX | MEAN | SD | MIN | MAX | MEAN | SD | VALUE |
ant.l | 577 | 815 | 749.2 | 73.72 | 743 | 743 | 743.4 | 0.01 | 1033 | 1035 | 1033.9 | 0.93 | 1116 |
clv.b | 65 | 90 | 77.2 | 6.88 | 64 | 75 | 69.3 | 8.23 | 94 | 97 | 95.4 | 1.64 | 72 |
clv.l | 168 | 197 | 184 | 8.31 | 183 | 199 | 190.9 | 10.9 | 201 | 211 | 206.1 | 6.58 | 235 |
eye.b | 237 | 304 | 280.4 | 28.08 | 272 | 274 | 273.4 | 1.47 | 291 | 299 | 295.2 | 5.42 | 282 |
eye.d | 559 | 739 | 673.6 | 79.6 | 631 | 631 | 631.2 | 0.34 | 668 | 674 | 671.2 | 4.3 | 572 |
eye.h | 315 | 408 | 371.3 | 38.59 | 355 | 355 | 355 | 0.01 | 459 | 481 | 470 | 15.25 | 393 |
eye.l | 231 | 301 | 276 | 28.51 | 262 | 267 | 264.9 | 3.57 | 291 | 297 | 293.9 | 4.8 | 280 |
fl3.b | 68 | 84 | 74 | 4.86 | 62 | 69 | 65.4 | 4.82 | 77 | 79 | 78.4 | 1.5 | 70 |
fl3.l | 63 | 110 | 88.4 | 15.64 | 87 | 97 | 92.3 | 6.99 | 140 | 141 | 140.7 | 0.55 | 146 |
fl8.b | 65 | 83 | 75.5 | 5.85 | 64 | 65 | 64.7 | 1.03 | 81 | 85 | 83 | 3.15 | 71 |
fl8.l | 63 | 82 | 73.3 | 6.22 | 65 | 69 | 67 | 2.61 | 85 | 94 | 89.8 | 6.18 | 101 |
fm3.b | 115 | 155 | 138.6 | 17 | 128 | 137 | 132.6 | 5.85 | 151 | 175 | 163.1 | 17.35 | 134 |
fm3.l | 496 | 659 | 581.7 | 70.88 | 578 | 594 | 586.2 | 11.28 | 675 | 681 | 677.7 | 3.99 | 577 |
fwi.b | 794 | 1054 | 961 | 110.3 | 903 | 905 | 903.8 | 1.32 | 1118 | 1142 | 1130.1 | 17.32 | 928 |
fwi.l | 1721 | 2214 | 2002.6 | 208.5 | 1813 | 1825 | 1818.8 | 8.59 | 2564 | 2568 | 2565.9 | 2.79 | 1963 |
gst.b | 622 | 875 | 765.9 | 91.68 | 650 | 673 | 661.4 | 16.26 | 403 | 415 | 409.1 | 8.2 | 424 |
gst.l | 932 | 1111 | 1045.7 | 46.68 | 1113 | 1133 | 1123.1 | 14.28 | 2090 | 2157 | 2123.4 | 47.45 | 1435 |
hea.b | 789 | 1036 | 946.4 | 102.7 | 886 | 889 | 887.8 | 1.96 | 965 | 981 | 972.8 | 11.33 | 859 |
hea.h | 575 | 740 | 688.1 | 60.65 | 604 | 610 | 607.1 | 3.76 | 694 | 727 | 710.4 | 23.5 | 598 |
hea.l | 380 | 507 | 461.5 | 53.02 | 436 | 440 | 437.8 | 2.81 | 459 | 472 | 465.5 | 9.54 | 416 |
lof.h | 333 | 423 | 388.8 | 36.03 | 355 | 367 | 361.1 | 8.45 | 303 | 303 | 303 | 0.5 | 304 |
mav.l | 349 | 475 | 419.8 | 48.08 | 362 | 381 | 371.4 | 12.89 | 533 | 557 | 545 | 16.8 | 433 |
msc.b | 591 | 806 | 731 | 87.75 | 682 | 684 | 683.3 | 1.43 | 815 | 829 | 822 | 9.83 | 732 |
msc.l | 363 | 511 | 451.8 | 61.49 | 456 | 462 | 458.9 | 4.39 | 536 | 577 | 556.7 | 29.11 | 509 |
msp.l | 222 | 299 | 262.4 | 27.02 | 205 | 217 | 211.1 | 8.98 | 214 | 219 | 216.4 | 3.43 | 171 |
mss.l | 940 | 1279 | 1140.1 | 136.82 | 1118 | 1118 | 1118.3 | 0.32 | 1384 | 1419 | 1401.7 | 24.4 | 1208 |
ool.l | 170 | 236 | 211.3 | 25.8 | 187 | 189 | 187.9 | 1.93 | 185 | 188 | 186.3 | 2.03 | 153 |
pdl.b | 53 | 67 | 62.5 | 5.33 | 61 | 61 | 60.9 | 0.52 | 59 | 63 | 61.1 | 2.45 | 61 |
pdl.l | 69 | 97 | 86.5 | 10.33 | 83 | 84 | 83.4 | 0.99 | 90 | 95 | 92.2 | 3.68 | 79 |
plc.d | 256 | 358 | 314.9 | 40.58 | 299 | 318 | 308.5 | 13.17 | 345 | 380 | 362.6 | 25.06 | 294 |
pmv.l | 301 | 414 | 364.9 | 38.59 | 320 | 339 | 329.3 | 13.34 | 518 | 530 | 524.2 | 8.42 | 392 |
pol.l | 142 | 186 | 167.2 | 17.67 | 168 | 179 | 173.5 | 7.83 | 222 | 224 | 222.9 | 0.97 | 202 |
ppd.l | 202 | 286 | 251.1 | 30.68 | 228 | 248 | 238.3 | 14.05 | 279 | 288 | 283.4 | 6.24 | 267 |
scp.b | 59 | 78 | 69.2 | 6.93 | 67 | 71 | 69 | 3.07 | 61 | 65 | 62.8 | 3.12 | 61 |
scp.l | 320 | 412 | 375.9 | 37.81 | 344 | 348 | 346 | 2.7 | 387 | 393 | 390.2 | 4.27 | 329 |
sct.l | 346 | 463 | 421.5 | 51.33 | 413 | 422 | 417.5 | 6.71 | 474 | 475 | 474.3 | 0.55 | 414 |
stv.l | 215 | 293 | 263.8 | 32.12 | 244 | 260 | 251.9 | 11.59 | 314 | 314 | 313.9 | 0.41 | 248 |
ta3.l | 400 | 529 | 485.5 | 48.7 | 468 | 478 | 473.1 | 7.13 | 672 | 675 | 673.4 | 2.51 | 572 |
tb3.b | 78 | 110 | 98.5 | 10.45 | 88 | 91 | 89.6 | 2.31 | 115 | 117 | 116.1 | 0.94 | 102 |
tb3.l | 586 | 778 | 696.5 | 80.08 | 646 | 653 | 649.8 | 4.77 | 840 | 842 | 840.9 | 0.83 | 721 |
tmp.l | 84 | 130 | 108.8 | 18.49 | 103 | 113 | 107.8 | 6.77 | 107 | 130 | 118.6 | 15.93 | 107 |