- Email: [email protected]
Extending Afrotropical endemism: Discovery of Prosapanesia Kieffer (Hymenoptera: Bethylidae) from the Oriental region with the description of two new species
Extending Afrotropical endemism: Discovery of Prosapanesia Kieffer (Hymenoptera: Bethylidae) from the Oriental region with the description of two new species S. Santhosh, A.P. Ranjith PII: DOI: Reference:
S1226-8615(16)30167-4 doi: 10.1016/j.aspen.2016.07.006 ASPEN 820
To appear in:
Journal of Asia-Pacific Entomology
Received date: Accepted date:
14 April 2016 5 July 2016
Please cite this article as: Santhosh, S., Ranjith, A.P., Extending Afrotropical endemism: Discovery of Prosapanesia Kieffer (Hymenoptera: Bethylidae) from the Oriental region with the description of two new species, Journal of Asia-Pacific Entomology (2016), doi: 10.1016/j.aspen.2016.07.006
This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
ACCEPTED MANUSCRIPT Extending Afrotropical endemism: Discovery of Prosapanesia Kieffer (Hymenoptera: Bethylidae) from the Oriental region with the description of two new species
A
PT
S. SanthoshA, B & A. P. RanjithA Department of Zoology, Malabar Christian College, Kozhikode (Affiliated to University of
Corresponding author. email: [email protected]
SC
B
RI
Calicut), 673001, Kerala, India.
NU
Abstract. Pristocerinae is one of the five subfamilies of Bethylidae exhibiting high degree of sexual dimorphism. Prosapanesia Kieffer, a rarely collected pristocerine genus is recorded for
MA
the first time from the Oriental region. Two new species based on females, Prosapanesia emarginata sp. nov. and P. orientalia sp. nov. are described with world key to the known female species. The generic diagnosis for both male and female are revised. The distribution
D
of Prosapanesia and its congeners is commented.
TE
Key words: Prosapanesia, Oriental region, autapomophic characters, key, new distribution
AC CE P
record Introduction
Bethylidae is one of the widely distributed families of Chrysidoidea, with about 100 valid genera and about 2,600 valid species around the world (Azevedo et al. 2015), with 44 genera recorded from Oriental region (Santhosh unpublished). Within Bethylidae some genera exhibit a strong endemism towards Oriental region like, Caloapenesia Terayama, Foenobethylus Kieffer, Galadoxa Nagy and Neoapenesia Terayama (Terayama 2003; Várkonyi and Polaszek 2007, Liu et al. 2011; Savergnini and Azevedo 2013). Terayama (1995a, 1996a, 1998) discovered Glenosema Kieffer, Parascleroderma Kieffer and Protisobrachium Benoit from Oriental region. Azevedo and Barbosa (2010) recently recorded Pararhabdepyris Gorbatovskii from Oriental region. The subfamily Pristocerinae, consists of 736 species in 24 genera around the world, of which 179 species from the Afrotropical region, three from the Australian region, 70 from the Nearctic region, 350 from the Neotropical region, 70 from the Oriental region, 71 from the 1
ACCEPTED MANUSCRIPT Palaearctic region and two from fossil specimens (Lim et al. 2011). Pristocerinae are mostly the ectoparasitoids of coleopteran larvae living in concealed habitat (Terayama 2006) and exhibit profound sexual dimorphism; males are fully winged with ocelli and tegulae, whereas
PT
females completely apterous without ocelli and compound eyes small or absent (Evans 1964; Lim et al. 2011). Some species of females in the genera Dissomphalus and Pseudisobrachium
RI
are often found to be associated with ant nests, so it is suggested that they are parasitoids of
SC
ants (Evans 1961, 1964) or myrmecophilous beetles (Finammore and Gauld 1995). Tachikawa (1985) summarized the families Curculionidae, Bostrychidae, Cleridae and Elateridae as the
NU
coleopteran hosts of the Pristocerinae.
Prosapanesia Kieffer is previously known from six species in which five species are
MA
described only from male specimens one from female specimen (Kieffer 1910, 1914; Benoit 1981; Krombein 1989; Terayama 2004). Terayama (1995b) transferred two Neusakosia Benoit species into Prosapanesia viz., Prosapanesia princeps and P. schoutedeni as there are
D
no concrete characters other than intrageneric differences present in Neusakosia in the
TE
retention of this genus. The original description of the genus Usakosia by Kieffer (1914) suggests that they closely resemble Prosapanesia, but it is unable to draw any conclusion
1981).
AC CE P
about the status of Usakosia as the type material was lost during the first world war (Benoit
Pristocerine females are hard to collect, only eight genera are known so far from females (Terayama 2003) and the only possibility to get the female specimens is that females are being carried by the males during the mating flight (Krombein 1989). Phoretic copulation is well recorded from Apenesia Westwood (Evans 1963, 1969; Gordh 1990). According to Terayama (1996b) the group including Afgoiogfa, Disepyris, Parascleroderma and Prosapanesia constitutes a monophyletic group defined by the absence of the metacarpus vein in the forewing and characters like large, broad pterostigma (in males) and anteriorly truncated clypeus (in females) support the monophyly of Prosapanesia. Prosapanesia displays autapomorphic characters within Pristocerinae and can be separated from its congeners in having medial clypeal carina terminates at a narrow, truncate, thickened apical lobe in females and males by the following combinations of characters, 1) head obliquely truncated anteriorly, 2) pterostigma remarkably large and broad, 3) radial vein of 2
ACCEPTED MANUSCRIPT fore wing weak and thin, 4) posterior margin of subgenital plate a remarkable process (Kieffer 1910, 1914; Krombein 1989; Terayama 2004).
PT
In this paper, we report Prosapanesia from the Oriental region for the first time with the descriptions of two new species based on female specimens. Until now, the species of the
RI
genus are known only from the Afrotropical region.
SC
Materials and methods
Specimens were collected from the scrub jungles using sweep net. The terms for integument
NU
sculpture follow Harris (1979) and most of measurements and morphological terms adopted in this paper follow Evans (1964) and Terayama (2006) and the abbreviation as follows EV,
MA
maximum distance from top of the eye to posterior margin of vertex in lateral view; HE, maximum height of eye in full face view; LH, maximum length of head from clypeal margin to vertex in full dorsal view; LM, maximum diagonal length of mesosoma excluding pronotal
D
collar in lateral view; LP, length of propodeum measured along the midline including the
TE
posterior declivity in dorsal view; LPD, length of propodeal disc measured along the midline excluding the posterior declivity in dorsal view; OOL, ocello-ocular line; SI, profemur
AC CE P
sphericity index; WF, minimum width between eyes in full face view; WH, maximum width of head including eyes in full dorsal view; WOT, width of ocellar triangle; WPD, maximum width of propodeal disc excluding anterior portion from the level of propodeal spiracles in dorsal view.
Specimens were imaged using an Leica M 205A microscope with automated multiple image capture at preset focal levels using an Leica DMC 2900 camera, and image combination using the Leica Application Suite image processing system. All images were edited using Photoshop CS8 (Version 6.1) (Adobe Inc.). A map with the distribution of all species was generated using SimpleMappr (Shorthouse 2010). Holotype and paratypes are deposited in Zoological Survey of India, Kozhikode (ZSIK). Results Prosapanesia Kieffer (Figs 1–3) 3
ACCEPTED MANUSCRIPT Prosapanesia Kieffer, 1910: 42–43; Kieffer, 1914: 424; Krombein, 1989: 627; Terayama, 1995: 889; Terayama, 2004: 31. Neusakosia Benoit, 1981: 839–840. Type species: Neusakosia schoutedeni Benoit, 1981.
PT
Synonymized by Terayama (1995).
RI
Diagnosis.
SC
Female.
Head. Head longer than wide (0.70–0.95×) as long as wide in full face view, moderately to
NU
distinctly coriaceous, slightly compressed in lateral view (1.80–2.0×) as long as high in lateral view, sparsely punctate; frons, vertex coriaceous, sparsely punctate with erect setae; vertex
MA
straight (in P. lacteipennis and in holotype of P. orientalia sp. nov.) or slightly emarginated medially (in paratypes of P. orientalia sp. nov.); clypeal margin thickened anteriorly, distinctly truncate, concave medially (P. emarginata sp. nov.), medial longitudinal carina
D
thickened with or without a distinct, wide arch shaped groove basally (absent in P. emarginata
TE
sp. nov.), area between groove and mid longitudinal carina smooth to faintly coriaceous, lower margin of truncation slightly or distinctly emarginate; mandible bidentate (in P.
AC CE P
emarginata sp. nov. and P. lacteipennis) or tetradentate (in P. orientalia sp. nov.) concave basally, narrower medially with lower tooth more acute and larger than the rest, upper two teeth blunt (in P. orientalia sp. nov.), transverse groove distinct in between two lower teeth, sparsely setose; eye absent; ocelli absent; antenna 13 segmented, antenna 1.64–2.05× as long as head; scape 3.85–4.70× as long as its maximum width, 3.70–4.45× as long as pedicel; pedicel 1.37–1.50× as long as its maximum width, 1.55–1.70× as long as first flagellomere; flagellomere 1–11 wider than long; terminal flagellomere acute apically, 1.68–2.40× as long as its maximum width. Mesosoma. Mesosoma slightly to distinctly coriaceous, punctures smaller and sparser than head (in P. lacteipennis) or same as head (in P. orientalia sp. nov.); pronotal collar slightly transverse anteriorly; pronotum convex anteriorly, not carinate, slightly widened or converged posteriorly, coriaceous, sparsely punctate with long erect setae, glabrous medially; notauli, paraspidal furrows absent; mesonotum narrower than pronotum basally, 1.03–1.16× as long as wide basally; scutellum coriaceous, sparsely setose, distinctly constricted posteriorly; 4
ACCEPTED MANUSCRIPT propleuron coriaceous with a distinct depression antero-medially, not extending to the posterior margin in lateral view; mesopleuron with large dorsal area rounding gradually to lateral surface, coriaceous with erect setae; propodeum 1.49–1.95× as long as maximum width,
PT
minimum width 0.33–0.35× maximum width, propodeal formula 9–12:7–16.25:20–29, sublateral posterior margin with few long erect setae, propodeal declivity glabrous; wings absent;
RI
fore femur 2.4–3.0× as long as its maximum width, mid femur swollen; mid tibia with long
SC
erect spines; hind tibia with long setae.
Metasoma. Metasoma sessile, coriaceous and polished except otherwise mentioned; 2.2–2.6×
NU
as long as its maximum width, 1.2× as long as mesosoma, 0.81× as long as head plus mesosoma; T1 smooth with erect setae; T2 smooth or punctate; T2 and T3 sparsely setose,
MA
glabrous basally; rest of the tergite sparsely setose. Male.
D
Head. Head wider than long, 1.12–1.17× as wide as long (in P. lacteipennis) with or without
TE
distinct bilobed process antero-medially; frons, vertex with small to large shallow sparse punctures, interspaces weakly microreticulate to smooth and shing; vertex straight in smaller
AC CE P
specimens and strongly grooved medially near to anterior ocellys (in P. lacteipennis) and moderately convex; mandible narrow to broad, robust, triangular with 4–5 teeth, apical tooth acute and large followed by blunt teeth; clypeus narrow, anterior margin concave or slightly produced; WF 1.2–2.13× HE, 0.56–0.66× WH, with small to large punctures; eyes slightly protuberant laterally, EV 0.9–1.79× HE; ocular setae lacking; ocelli small to large, OOL 0.56– 0.86× WOT; antenna short, densely setose with erect hairs; ratio of first four flagellomeres 11–36: 4–9: 6–19: 5.5–19; third flagellomere 1.5–2.38× as long as wide. Mesosoma. Pronotum smooth to weakly microreticulate, with scattered punctures, not carinate anteriorly, convex antero-medially; notauli, paraspidal furrows distinct; scutellum with anterior transverse groove, sparsely punctate; metanotum with small, shallow pit medially, small fovea laterally; propodeum as long as wide with a weak to strong medial longitudinal carina (medial longitudinal carina present at basal 3/5 in P. longimandibulata) and weak lateral carina (lateral carina absent in P. sesriemensis) transverse carina absent, basal
5
ACCEPTED MANUSCRIPT triangular area distinct or indistinct, propodeal declivity rugulose; fore wing with broad pterostigma; mid tibia with spines at outer margin.
PT
Metasoma. Metasoma smooth to weakly microreticulate; subgenital plate bilobed apically; genitalia with aedeagus massive, paramere subtruncate to slightly rounded apically, deeply
RI
concave ventrally.
SC
Distribution
NU
Afrotropical and Oriental regions. Biology
MA
Unknown.
Key to the female species of Prosapanesia Kieffer Head and mesosoma distinctly coriaceous; mandible tetradentate; scape 3.3–3.7× as
D
1.
maximum
TE
long as its maximum width; WH 0.7× LH; propodeum 1.3–1.5× as long as its width;
propodeal
disc
constricted
-
AC CE P
anteriorly………………………………………………Prosapanesia orientalia sp. nov. Head and mesosoma moderately coriaceous; mandible bidentate; scape 4.2–4.7× as long as its maximum width; WH 0.87–0.96× LH; propodeum 1.65–1.95× as long as its maximum width; propodeal disc parallel anteriorly ……………………………….…..2 2.
Anterior clypeal margin distinctly emarginate; pronotum diverging posteriorly; medial clypeal carina with inditinct groove basally; lateral clypeal margin without denticles; propodeum 1.65× as long as its maximum width; fore femur 2.4× as long as wide; antennae, legs yellow…………………………….....Prosapanesia emarginata sp. nov.
-
Anterior clypeal margin not emarginated; pronotum converging posteriorly; meadial clypeal carina with distinct groove basally; lateral clypeal margin with denticles; propodeum 1.95× as long as its maximum width; fore femur 3× as long as wide; antennae, legs light red………………...……………Prosapanesia lacteipennis Kieffer Descriptive taxonomy Prosapanesia emarginata, Santhosh and Ranjith sp. nov. 6
ACCEPTED MANUSCRIPT (Figs 1, 3A, B, E) Material Examined sea level, sweep net, 20.v.2015. coll. Ranjith, A.P.”
PT
Holotype. Female, “INDIA: Kerala, Wayanad, Kalladi, scrub jungle, 4612 ft. above
RI
Measurements: Body length 2.21 mm; LH 0.43 mm; WH 0.32 mm; LM 0.76 mm; LP 0.34
SC
mm; LPD 0.27 mm; WPD 0.17 mm.
Colour. Head black; antennae, mandibles and clypeus yellow; median clypeal ridge and
NU
anterior truncated margin dark brown; mesosoma black except dark brown pronotal collar and propodeum, posterior margin of pronotum, mesonotum, propleuron and mesopleuron brown,
MA
legs yellow except fore coxa brownish yellow, metasoma brownish yellow except darker T1, dorso medially black at base.
Head. Head sub-rectangular in full face view, polished, moderately coriaceous, WH 0.74× LH;
D
compressed in lateral view, 1.87× as long as high; frons, vertex moderately coriaceous,
TE
sparsely punctate with erect setae; vertex convex in dorsal view; in full face view, anterior clypeal margin emarginate, truncate, without denticles are on either side of the median clypeal
AC CE P
carina, median clypeal carina weak, arched, basal groove indistinct; mandible with long erect setae, bidentate; eye and ocelli absent; antenna 2.05× as long as head; scape 4.70× as long as its maximum width, 3.5× as long as pedicel; pedicel 1.44× as long as its maximum width, 1.64× as long as first flagellomere; flagellomere wider than long except terminal one, terminal flagellomere acute apically, 1.77× as long as its maximum width. Mesosoma. Sparsely coriaceous with sparse suberect setae; pronotal collar anteriorly carinate, setose; pronotum 1.20× as long as its maximum width, convex anteriorly, widened posteriorly; mesonotum strongly constricted posteriorly, 1.03x as long as basal width; notauli and parapsidal furrows absent; propleuron with a shallow distinct depression antero-medially, not extending to the posterior margin in lateral view; propodeal disc constricted, parallel anteriorly with two diverging processes in the anterior end joining posterior apex of mesonotum medially, propodeum 1.65× as long as its maximum width, minimum width 0.34x maximum width, propodeal formula 7.8:7.2:17.5, sublateral posterior margin without long erect setae, propodeal declivity glabrous; wings absent; forefemur 2.4× as long as its 7
ACCEPTED MANUSCRIPT maximum width, mid femur swollen; mid tibia with long erect spines; hind tibia with long setae.
PT
Metasoma. Faintly coriaceous and punctate; 2.50× as long as its maximum width, 1.42× as long as mesosoma, 0.97× as long as head plus mesosoma; T1 smooth with erect setae and pair
RI
of distinct pits laterally; T2 punctate, with erect hairs, glabrous basally; T3 punctate, sparsely
SC
setose, glabrous basally; rest of the tergite sparsely setose. Male
NU
Unknown.
MA
Etymology
The species name refers to the emarginated structure of anterior clypeal margin.
AC CE P
Remarks
TE
India-Kerala (Fig 4).
D
Distribution
Prosapanesia emarginata sp. nov. differs from the remaining female Prosapanesia spp. in having anterior clypeal margin concave, scape more slender (4.7× as long as its maximum width) and propodeum less than 1.7× as long as its maximum width. This species comes close to P. lacteipennis in having mandible bidentate, head moderately smooth. It differs in the following characters, antennae dark yellow (light red in P. lacteipennis), fore femur 2.4× as long as wide (3.0× as long as wide in P. lacteipennis). It differs from P. orientalia sp. nov. in having head and mesosoma moderately coriaceous (strongly coriaceous in P. orientalia sp. nov.), mandible bidentate (tridentate in P. orientalia sp. nov.), lateral clypeal margin without denticles (with denticles in P. orientalia sp. nov.), medial clypeal carina without groove basally (medial clypeal carina with carinate groove basally in P. orientalia sp. nov.), scape distinctly slender, 4.7× as long as its maximum width (3.26× as long as its maximum width in P. orientalia sp. nov.), propodeal disc parallel anteriorly (distinctly constricted anteriorly in P. orientalia sp. nov.), sub lateral posterior margin of propodeum glabrous (sub lateral posterior margin of propodeum with few long erect setae in P. orientalia sp. nov.), metasoma 2.5× as 8
ACCEPTED MANUSCRIPT long as its maximum width (2.2× as long as its maximum width in P. orientalia sp. nov.), T2 punctate (T2 smooth in P. orientalia sp. nov.).
PT
Prosapanesia orientalia, Ranjith and Santhosh sp. nov. (Figs 2, 3C, D, F)
RI
Material Examined
Holotype. Female, “INDIA: Kerala, Calicut, Janakikkadu, scrub jungle, sweep net,
SC
23.xii.2014. coll. Ranjith, A.P.”
NU
Paratypes. 2 Females, “INDIA: Kerala, Malappuram, Calicut University Botanical Garden 11°7’N 75° 5’E, Sweep net, 26.v.2004. coll. S. Santhosh”
mm; LPD 0.35 mm; WPD 0.28 mm.
MA
Measurements. Body length 2.88 mm; LH 0.59 mm; WH 0.46 mm; LM 1.02 mm; LP 0.43
D
Colour. Head black; antennae yellow; mandibles and clypeus reddish brown; median clypeal
TE
ridge and anterior truncated margin dark brown; mesosoma black except brown pronotal collar, posterior margin of pronotum, mesonotum, propleuron and mesopleuron brown, legs
black.
AC CE P
yellow except forecoxa reddish brown, metasoma brown except base of T1 dorso medially
Head. Head sub-rectangular in full face view, coriaceous, WH 0.78× LH; moderately compressed in lateral view, 1.83× as long as high; frons, vertex strongly coriaceous, sparsely punctate with erect setae; vertex straight in dorsal view; anterior clypeal margin thickened, distinctly truncate, three dark brown colored denticles are on either side of the truncated clypeus, median clypeal carina strong, with arched groove with carinate margins basally; mandible with long erect setae, tetradentate with innermost two teeth smaller and blunt; eye and ocelli absent; antenna 1.58× as long as head; scape 3.26× as long as its maximum width, 3.7× as long as pedicel; pedicel 1.32× as long as its maximum width, 1.64× as long as first flagellomere; flagellomere wider than long except terminal one, terminal flagellomere acute apically, 1.53× as long as its maximum width. Mesosoma. Strongly coriaceous with sparse erect setae except otherwise mentioned; pronotal collar anteriorly carinate, asetose; pronotum 1.03x as long as its maximum width, convex 9
ACCEPTED MANUSCRIPT anteriorly, slightly widened posteriorly; mesonotum strongly constricted posteriorly, 1.03x as long as basal width; notauli and parapsidal furrows absent; propleuron with a distinct depression antero-medially, not extending to the posterior margin in lateral view; propodeal
PT
disc constricted anteriorly with two diverging processes in the anterior end joining posterior apex of mesonotum medially, propodeum 1.49× as long as its maximum width, minimum
RI
width 0.33x maximum width, propodeal formula 12:10:29, sub lateral posterior margin with
SC
few long erect setae, propodeal declivity glabrous; wings absent; fore femur 2.7× as long as its maximum width, mid femur swollen; mid tibia with long erect spines; hind tibia with long
NU
setae.
Metasoma. Coriaceous and polished except otherwise mentioned; 2.2× as long as its
MA
maximum width, 1.2× as long as mesosoma, 0.81× as long as head plus mesosoma; T1 smooth with erect setae and a pair of distinct pits laterally; T2 & T3 sparsely setose, glabrous
D
basally; rest of the tergite sparsely setose.
TE
Male
Etymology
AC CE P
Unknown.
The name refers to the distribution of the species, the only Prosapanesia known from the Oriental region. Distribution
India-Kerala (Fig 4). Variations Mandibles and clypeus yellow with mandible base and teeth dark brown; median clypeal ridge and anterior truncated margin brown; pronotal collar, posterior margin of pronotum, mesonotum, propleuron and mesopleuron paler, legs yellow except darker fore coxa; terminal flagellomere 1.76× as long as its maximum width; vertex slightly emarginate in full face view; propodeum 1.3× as long as its maximum width.
10
ACCEPTED MANUSCRIPT Remarks Prosapanesia orientalia sp. nov. comes close to P. lacteipennis in having head sparsely
PT
punctate, lower tooth more acute and longer, pronotum 1.3× as long as its apical width, mesonotum narrower than pronotum, mesopleuron with large dorsal area. But P. orientalia sp.
RI
nov. differs from P. lacteipennis in having head distinctly coriaceous (head moderately coriaceous in P. lacteipennis), antennae yellow (light red in P. lacteipennis), mandible
SC
tetradentate (bidentate in P. lacteipennis), lower margin of truncation distinctly emarginated (slightly emarginate in P. lacteipennis), pronotum distinctly convex (pronotum somewhat
NU
straight anteriorly in P. lacteipennis), propodeum not rounded posteriorly (evenly rounded posteriorly in P. lacteipennis), scape distinctly robust, 3.78–3.85× as long as its maximum
MA
width (scape slender, 4.2× as long as wide in P. lacteipennis), legs except fore coxa yellow (legs light red in P. lacteipennis), fore femur slightly robust, 2.6–2.7× as long as its maximum
D
width (slender 3× as long as wide in P. lacteipennis), metasoma dark yellow, 2.3–2.9× as long as its maximum width (metasoma light red, 2.6× as long as its maximum width), T3+
AC CE P
Discussion
TE
coriaceous (shiny in P. lacteipennis).
Distribution of Prosapanesia and its congeners Though, the non pristocerine genera shows strong endemism towards the Neotropical and Palearctic regions (Gordh and Móczár 1990), many pristocerine wasps are found endemic to the Afrotropical region (Gordh and Móczár 1990) and according to Terayama (2003) eight genera are endemic to the Afrotropics. Female pristocerine wasps except Afgoiogfa and Prosapanesia are widely distributed in the tropical and temperate regions (Gordh and Móczár 1990; Terayama 2003). Species of Prosapanesia are distributed in the southwest region of Africa (Kieffer 1910, 1914; Benoit 1981; Krombein 1989; Terayama 2004). Even though the non pristocerine genera like Trachepyris Kieffer and Pycnomesitius Móczár have shared Afrotropical and Oriental endemism in their distribution (Terayama 2003), no pristocerines are so far found to have their distribution restricted to both the regions. 11
ACCEPTED MANUSCRIPT The discovery of Oriental range extension of Prosapensia throws light on a strong possibility of existence of other endemic Afrotropical pristocerines in the Oriental region. The above possibility is supported by the previous discoveries of Protisobrachium (Terayama 1995), and
PT
Parascleroderma Kieffer (Terayama 1998), which are the congeners of Prosapanesia. The unidentified species of Protisobrachium (Santhosh unpublished) support the extension of
RI
distribution status of the genus, which is found to be endemic to Afrotropical region earlier.
SC
Acknowledgements
NU
This work is part of a research project funded by Department of Science and Technology, Government of India under SERB-Young Scientist Scheme (SB/YS/LS-42/2014). We are
MA
indebted to the authorities of Malabar Christian College, Calicut for providing the facilities. References
D
Azevedo, C. O., Barbosa, D. N., 2010. Two new species Pararhabdepyris Gorbatovskii
TE
(Hymenoptera, Bethylidae) from Australia and Thailand. Zootaxa. 2668, 55–62.
AC CE P
Azevedo, C. O., Garcia, R., Gobbi, F. T., Noort, S. van, 2015. Insecta, Hymenoptera, Bethylidae: range extension and filling gaps in Central African Republic. Check List. 11 (2), 1–4. doi:10.15560/11.2.1606 Benoit, P. L. G. , 1981. Bethylidae africains (Hymenoptera). La Tribu Usakosiini. Revue de Zoologie Africaine. 95 (4), 833–842. Evans, H. E., 1963. A revision of the genus Apenesia in the Americas (Hymenoptera, Bethylidae). Bulletin of the Museum of Comparative Zoology. 130, 251–359. Evans, H. E., 1969. Phoretic copulation in Hymenoptera. Entomological News. 80,113–124. Evans, H. E., 1961. A revision of the genus Pseudisobrachium in North and Central America (Hymenoptera, Bethylidae). Bulletin of the Museum of Comparative Zoology. 126, 211–318. Evans, H. E., 1964. A synopsis of the American Bethylidae (Hymenoptera: Aculeata). Bulletin of the Museum of Comparative Zoology. 132 (1), 1–222. 12
ACCEPTED MANUSCRIPT Finammore, A. T., Gauld, I. D., 1995. Chrysididae, in: Hanson, P.E. and Gauld, I.D. (Eds.), The Hymenoptera of Costa Rica. Oxford: Oxford University Press, pp. 479–488.
PT
Gordh, G., 1990. Apenesia evansi sp. n. (Hymenoptera: Bethylidae) from Australia with comments on phoretic copulation in Bethylids. Journal of the Australian
RI
Entomological Society. 29, 167–170. doi:10.1111/j.1440-6055.1990.tb00341.x
SC
Gordh, G., Móczár, L., 1990. A catalog of the world Bethylidae (Hymenoptera: Aculeata). Memoirs of the American Entomological Institute. 46, 1–364.
NU
Harris, R. A., 1979. A glossary of surface sculpturing. Occasional Papers in Entomology. 28,
MA
1–31.
Kieffer, J. J., 1910. Description de nouveaux Béthylides (Hymén.). Annales de la Société Entomologique de France. 79, 31–56.
D
Kieffer, J. J., 1914. Bethylinae, in: Das Tierreich. Vol. 41. R. Friedlander und Sohn, Berlin, pp.
TE
228–595.
AC CE P
Krombein, K. V., 1989. Systematic notes on some Bethylidae from Botswana: Pristocerinae (Hymenoptera: Aculeata). Proceedings of the Entomological Society of Washington. 91: 620–631.
Lim, J., Terayama, M., Koh, S., Lee, J., Lee, S., 2011. A taxonomic review of the subfamily Pristocerinae (Hymenoptera: Chrysidoidea: Bethylidae) from Korea with descriptions of
two
new
species.
Journal
of
Natural
History.
45
(7-8),
435–460.
doi:10.1080/00222933.2010.534191 Liu, J., Chen, H., Xu, Z., 2011. Two new species of genus Foenobethylus Kieffer, 1913 (Hymenoptera, Bethylidae) from China with a key to the known species. Zootaxa. 2806, 53–59. Savergnini, J.R., Azevedo, C.O. 2013. Taxonomy of Foenobethylus Kieffer (Hymenoptera, Bethylidae) with description of two new species. Journal of Asia-Pacific Entomology. 16, 433–441. doi:10.1016/j.aspen.2013.06.004
13
ACCEPTED MANUSCRIPT Shorthouse, D.P. 2010. SimpleMappr, an online tool to produce publication-quality point maps. Available from http://www.simplemappr.net (accessed on 2 February 2016).
PT
Tachikawa, T.,1985. On the bethylid wasps (II). Forest Pests. 34(9), 161–169 (in Japanese). Terayama, M., 1995a. Discovery of the genus Protisobrachium Benoit 1957 from the Oriental
RI
region (Hymenoptera: Bethylidae). Journal of Entomological Science. 30 (4), 443–446.
SC
Terayama, M., 1995b. Caloapenesia and Neoapenesia, new genera of the family Bethylidae (Hymenoptera, Chrysidoidea) from the Oriental region, with proposals of two new
NU
synonymies of genera. Japanese Journal of Entomology. 63 (4), 881–891.
MA
Terayama, M., 1996a. Discovery of the Genus Glenosema from the Oriental Region with Descriptions of Three New Species (Hymenoptera: Bethylidae). The Florida Entomologist. 79 (4), 591–595.
D
Terayama, M., 1996b. The phylogeny of the bethylid wasp subfamily Pristocerinae
TE
(Hymenoptera, Bethylidae). Japanese Journal of Entomology. 64, 587–601.
AC CE P
Terayama, M., 1998. Discovery of the genus Parascleroderma Kieffer from the Oriental region, with descriptions of four new species (Hymenoptera: Bethylidae). Entomological Science. 1(1), 129–132. Terayama, M., 2003. Phylogenetic systematic of the family Bethylidae (Insecta: Hymenoptera). Part II. Keys to subfamilies, tribes and genera in the world. Academic Reports, Faculty of Engineering, Tokyo Polytechnic University. 26 (1), 16–29. Terayama, M., 2004. Descriptions of new taxa and distribution records of the family Bethylidae (Insecta, Hymenoptera) I. Subfamily Pristocerinae. Academic Reports, Faculty of Engineering, Tokyo Polytechnic University. 27 (1), 22–38. Terayama, M., 2006. The Insects of Japan. Vol. I, Bethylidae (Hymenoptera). Fukuoka, (Japan): Entomological Society of Japan. 1–319. Várkonyi, G., Polaszek, A., 2007. Rediscovery and revision of Foenobethylus Kieffer, 1913 (Hymenoptera, Bethylidae). Zootaxa. 1546, 1–14. 14
ACCEPTED MANUSCRIPT
FIGURE CAPTIONS
PT
figure 1. Prosapanesia emarginata sp. nov., female, holotype (A) Habitus, lateral view; (B) Head, dorsal view; (C) Mesosoma, dorsal view; (D) Propodeum, dorsal view; (E) Propodeum
RI
& metasoma (in part), dorsal view; (F) metasoma, lateral view.
SC
figure 2. Prosapanesia orientalia sp. nov., female, holotype (A) Habitus, lateral view; (B) Head, dorsal view; (C) Mesosoma, dorsal view; (D) Propodeum, dorsal view; (E) Propodeum
NU
& metasoma (in part), dorsal view; (F) metasoma, lateral view.
MA
figure 3. (A, B) Prosapanesia emarginata sp. nov., head oblique views; (C, D) P. orientalia sp. nov.; (E) P. emarginata sp. nov., metasoma, dorsal view; (F) P. orientalia sp. nov., metasoma, dorsal view.
AC CE P
TE
D
figure 4. Distribution map of Prosapanesia species
15
NU
SC
RI
PT
ACCEPTED MANUSCRIPT
AC CE P
TE
D
MA
Graphical abstract
16
AC CE P
TE
D
MA
NU
SC
RI
PT
ACCEPTED MANUSCRIPT
17
ACCEPTED MANUSCRIPT
AC CE P
TE
D
MA
NU
SC
RI
PT
Fig. 1
18
AC CE P
TE
D
MA
NU
SC
RI
PT
ACCEPTED MANUSCRIPT
19
ACCEPTED MANUSCRIPT
AC CE P
TE
D
MA
NU
SC
RI
PT
Fig. 2
20
AC CE P
TE
D
MA
NU
SC
RI
PT
ACCEPTED MANUSCRIPT
21
ACCEPTED MANUSCRIPT
AC CE P
TE
D
MA
NU
SC
RI
PT
Fig. 3
22
AC CE P
Fig. 4
TE
D
MA
NU
SC
RI
PT
ACCEPTED MANUSCRIPT
23
ACCEPTED MANUSCRIPT Highlights
PT
Rarely collected bethylid genus Prosapanesia is discovered from Oriental region Prosapanesia emarginata sp. nov. and P. orientalia sp. nov. are described
RI
A world key for Prosapanesia is provided
SC
Male and female generic diagnosis of Prosapanesia are redefined
AC CE P
TE
D
MA
NU
Oriental presence of more endemic Afrotropical genera is suggested
24
S1226-8615(16)30167-4 doi: 10.1016/j.aspen.2016.07.006 ASPEN 820
To appear in:
Journal of Asia-Pacific Entomology
Received date: Accepted date:
14 April 2016 5 July 2016
Please cite this article as: Santhosh, S., Ranjith, A.P., Extending Afrotropical endemism: Discovery of Prosapanesia Kieffer (Hymenoptera: Bethylidae) from the Oriental region with the description of two new species, Journal of Asia-Pacific Entomology (2016), doi: 10.1016/j.aspen.2016.07.006
This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
ACCEPTED MANUSCRIPT Extending Afrotropical endemism: Discovery of Prosapanesia Kieffer (Hymenoptera: Bethylidae) from the Oriental region with the description of two new species
A
PT
S. SanthoshA, B & A. P. RanjithA Department of Zoology, Malabar Christian College, Kozhikode (Affiliated to University of
Corresponding author. email: [email protected]
SC
B
RI
Calicut), 673001, Kerala, India.
NU
Abstract. Pristocerinae is one of the five subfamilies of Bethylidae exhibiting high degree of sexual dimorphism. Prosapanesia Kieffer, a rarely collected pristocerine genus is recorded for
MA
the first time from the Oriental region. Two new species based on females, Prosapanesia emarginata sp. nov. and P. orientalia sp. nov. are described with world key to the known female species. The generic diagnosis for both male and female are revised. The distribution
D
of Prosapanesia and its congeners is commented.
TE
Key words: Prosapanesia, Oriental region, autapomophic characters, key, new distribution
AC CE P
record Introduction
Bethylidae is one of the widely distributed families of Chrysidoidea, with about 100 valid genera and about 2,600 valid species around the world (Azevedo et al. 2015), with 44 genera recorded from Oriental region (Santhosh unpublished). Within Bethylidae some genera exhibit a strong endemism towards Oriental region like, Caloapenesia Terayama, Foenobethylus Kieffer, Galadoxa Nagy and Neoapenesia Terayama (Terayama 2003; Várkonyi and Polaszek 2007, Liu et al. 2011; Savergnini and Azevedo 2013). Terayama (1995a, 1996a, 1998) discovered Glenosema Kieffer, Parascleroderma Kieffer and Protisobrachium Benoit from Oriental region. Azevedo and Barbosa (2010) recently recorded Pararhabdepyris Gorbatovskii from Oriental region. The subfamily Pristocerinae, consists of 736 species in 24 genera around the world, of which 179 species from the Afrotropical region, three from the Australian region, 70 from the Nearctic region, 350 from the Neotropical region, 70 from the Oriental region, 71 from the 1
ACCEPTED MANUSCRIPT Palaearctic region and two from fossil specimens (Lim et al. 2011). Pristocerinae are mostly the ectoparasitoids of coleopteran larvae living in concealed habitat (Terayama 2006) and exhibit profound sexual dimorphism; males are fully winged with ocelli and tegulae, whereas
PT
females completely apterous without ocelli and compound eyes small or absent (Evans 1964; Lim et al. 2011). Some species of females in the genera Dissomphalus and Pseudisobrachium
RI
are often found to be associated with ant nests, so it is suggested that they are parasitoids of
SC
ants (Evans 1961, 1964) or myrmecophilous beetles (Finammore and Gauld 1995). Tachikawa (1985) summarized the families Curculionidae, Bostrychidae, Cleridae and Elateridae as the
NU
coleopteran hosts of the Pristocerinae.
Prosapanesia Kieffer is previously known from six species in which five species are
MA
described only from male specimens one from female specimen (Kieffer 1910, 1914; Benoit 1981; Krombein 1989; Terayama 2004). Terayama (1995b) transferred two Neusakosia Benoit species into Prosapanesia viz., Prosapanesia princeps and P. schoutedeni as there are
D
no concrete characters other than intrageneric differences present in Neusakosia in the
TE
retention of this genus. The original description of the genus Usakosia by Kieffer (1914) suggests that they closely resemble Prosapanesia, but it is unable to draw any conclusion
1981).
AC CE P
about the status of Usakosia as the type material was lost during the first world war (Benoit
Pristocerine females are hard to collect, only eight genera are known so far from females (Terayama 2003) and the only possibility to get the female specimens is that females are being carried by the males during the mating flight (Krombein 1989). Phoretic copulation is well recorded from Apenesia Westwood (Evans 1963, 1969; Gordh 1990). According to Terayama (1996b) the group including Afgoiogfa, Disepyris, Parascleroderma and Prosapanesia constitutes a monophyletic group defined by the absence of the metacarpus vein in the forewing and characters like large, broad pterostigma (in males) and anteriorly truncated clypeus (in females) support the monophyly of Prosapanesia. Prosapanesia displays autapomorphic characters within Pristocerinae and can be separated from its congeners in having medial clypeal carina terminates at a narrow, truncate, thickened apical lobe in females and males by the following combinations of characters, 1) head obliquely truncated anteriorly, 2) pterostigma remarkably large and broad, 3) radial vein of 2
ACCEPTED MANUSCRIPT fore wing weak and thin, 4) posterior margin of subgenital plate a remarkable process (Kieffer 1910, 1914; Krombein 1989; Terayama 2004).
PT
In this paper, we report Prosapanesia from the Oriental region for the first time with the descriptions of two new species based on female specimens. Until now, the species of the
RI
genus are known only from the Afrotropical region.
SC
Materials and methods
Specimens were collected from the scrub jungles using sweep net. The terms for integument
NU
sculpture follow Harris (1979) and most of measurements and morphological terms adopted in this paper follow Evans (1964) and Terayama (2006) and the abbreviation as follows EV,
MA
maximum distance from top of the eye to posterior margin of vertex in lateral view; HE, maximum height of eye in full face view; LH, maximum length of head from clypeal margin to vertex in full dorsal view; LM, maximum diagonal length of mesosoma excluding pronotal
D
collar in lateral view; LP, length of propodeum measured along the midline including the
TE
posterior declivity in dorsal view; LPD, length of propodeal disc measured along the midline excluding the posterior declivity in dorsal view; OOL, ocello-ocular line; SI, profemur
AC CE P
sphericity index; WF, minimum width between eyes in full face view; WH, maximum width of head including eyes in full dorsal view; WOT, width of ocellar triangle; WPD, maximum width of propodeal disc excluding anterior portion from the level of propodeal spiracles in dorsal view.
Specimens were imaged using an Leica M 205A microscope with automated multiple image capture at preset focal levels using an Leica DMC 2900 camera, and image combination using the Leica Application Suite image processing system. All images were edited using Photoshop CS8 (Version 6.1) (Adobe Inc.). A map with the distribution of all species was generated using SimpleMappr (Shorthouse 2010). Holotype and paratypes are deposited in Zoological Survey of India, Kozhikode (ZSIK). Results Prosapanesia Kieffer (Figs 1–3) 3
ACCEPTED MANUSCRIPT Prosapanesia Kieffer, 1910: 42–43; Kieffer, 1914: 424; Krombein, 1989: 627; Terayama, 1995: 889; Terayama, 2004: 31. Neusakosia Benoit, 1981: 839–840. Type species: Neusakosia schoutedeni Benoit, 1981.
PT
Synonymized by Terayama (1995).
RI
Diagnosis.
SC
Female.
Head. Head longer than wide (0.70–0.95×) as long as wide in full face view, moderately to
NU
distinctly coriaceous, slightly compressed in lateral view (1.80–2.0×) as long as high in lateral view, sparsely punctate; frons, vertex coriaceous, sparsely punctate with erect setae; vertex
MA
straight (in P. lacteipennis and in holotype of P. orientalia sp. nov.) or slightly emarginated medially (in paratypes of P. orientalia sp. nov.); clypeal margin thickened anteriorly, distinctly truncate, concave medially (P. emarginata sp. nov.), medial longitudinal carina
D
thickened with or without a distinct, wide arch shaped groove basally (absent in P. emarginata
TE
sp. nov.), area between groove and mid longitudinal carina smooth to faintly coriaceous, lower margin of truncation slightly or distinctly emarginate; mandible bidentate (in P.
AC CE P
emarginata sp. nov. and P. lacteipennis) or tetradentate (in P. orientalia sp. nov.) concave basally, narrower medially with lower tooth more acute and larger than the rest, upper two teeth blunt (in P. orientalia sp. nov.), transverse groove distinct in between two lower teeth, sparsely setose; eye absent; ocelli absent; antenna 13 segmented, antenna 1.64–2.05× as long as head; scape 3.85–4.70× as long as its maximum width, 3.70–4.45× as long as pedicel; pedicel 1.37–1.50× as long as its maximum width, 1.55–1.70× as long as first flagellomere; flagellomere 1–11 wider than long; terminal flagellomere acute apically, 1.68–2.40× as long as its maximum width. Mesosoma. Mesosoma slightly to distinctly coriaceous, punctures smaller and sparser than head (in P. lacteipennis) or same as head (in P. orientalia sp. nov.); pronotal collar slightly transverse anteriorly; pronotum convex anteriorly, not carinate, slightly widened or converged posteriorly, coriaceous, sparsely punctate with long erect setae, glabrous medially; notauli, paraspidal furrows absent; mesonotum narrower than pronotum basally, 1.03–1.16× as long as wide basally; scutellum coriaceous, sparsely setose, distinctly constricted posteriorly; 4
ACCEPTED MANUSCRIPT propleuron coriaceous with a distinct depression antero-medially, not extending to the posterior margin in lateral view; mesopleuron with large dorsal area rounding gradually to lateral surface, coriaceous with erect setae; propodeum 1.49–1.95× as long as maximum width,
PT
minimum width 0.33–0.35× maximum width, propodeal formula 9–12:7–16.25:20–29, sublateral posterior margin with few long erect setae, propodeal declivity glabrous; wings absent;
RI
fore femur 2.4–3.0× as long as its maximum width, mid femur swollen; mid tibia with long
SC
erect spines; hind tibia with long setae.
Metasoma. Metasoma sessile, coriaceous and polished except otherwise mentioned; 2.2–2.6×
NU
as long as its maximum width, 1.2× as long as mesosoma, 0.81× as long as head plus mesosoma; T1 smooth with erect setae; T2 smooth or punctate; T2 and T3 sparsely setose,
MA
glabrous basally; rest of the tergite sparsely setose. Male.
D
Head. Head wider than long, 1.12–1.17× as wide as long (in P. lacteipennis) with or without
TE
distinct bilobed process antero-medially; frons, vertex with small to large shallow sparse punctures, interspaces weakly microreticulate to smooth and shing; vertex straight in smaller
AC CE P
specimens and strongly grooved medially near to anterior ocellys (in P. lacteipennis) and moderately convex; mandible narrow to broad, robust, triangular with 4–5 teeth, apical tooth acute and large followed by blunt teeth; clypeus narrow, anterior margin concave or slightly produced; WF 1.2–2.13× HE, 0.56–0.66× WH, with small to large punctures; eyes slightly protuberant laterally, EV 0.9–1.79× HE; ocular setae lacking; ocelli small to large, OOL 0.56– 0.86× WOT; antenna short, densely setose with erect hairs; ratio of first four flagellomeres 11–36: 4–9: 6–19: 5.5–19; third flagellomere 1.5–2.38× as long as wide. Mesosoma. Pronotum smooth to weakly microreticulate, with scattered punctures, not carinate anteriorly, convex antero-medially; notauli, paraspidal furrows distinct; scutellum with anterior transverse groove, sparsely punctate; metanotum with small, shallow pit medially, small fovea laterally; propodeum as long as wide with a weak to strong medial longitudinal carina (medial longitudinal carina present at basal 3/5 in P. longimandibulata) and weak lateral carina (lateral carina absent in P. sesriemensis) transverse carina absent, basal
5
ACCEPTED MANUSCRIPT triangular area distinct or indistinct, propodeal declivity rugulose; fore wing with broad pterostigma; mid tibia with spines at outer margin.
PT
Metasoma. Metasoma smooth to weakly microreticulate; subgenital plate bilobed apically; genitalia with aedeagus massive, paramere subtruncate to slightly rounded apically, deeply
RI
concave ventrally.
SC
Distribution
NU
Afrotropical and Oriental regions. Biology
MA
Unknown.
Key to the female species of Prosapanesia Kieffer Head and mesosoma distinctly coriaceous; mandible tetradentate; scape 3.3–3.7× as
D
1.
maximum
TE
long as its maximum width; WH 0.7× LH; propodeum 1.3–1.5× as long as its width;
propodeal
disc
constricted
-
AC CE P
anteriorly………………………………………………Prosapanesia orientalia sp. nov. Head and mesosoma moderately coriaceous; mandible bidentate; scape 4.2–4.7× as long as its maximum width; WH 0.87–0.96× LH; propodeum 1.65–1.95× as long as its maximum width; propodeal disc parallel anteriorly ……………………………….…..2 2.
Anterior clypeal margin distinctly emarginate; pronotum diverging posteriorly; medial clypeal carina with inditinct groove basally; lateral clypeal margin without denticles; propodeum 1.65× as long as its maximum width; fore femur 2.4× as long as wide; antennae, legs yellow…………………………….....Prosapanesia emarginata sp. nov.
-
Anterior clypeal margin not emarginated; pronotum converging posteriorly; meadial clypeal carina with distinct groove basally; lateral clypeal margin with denticles; propodeum 1.95× as long as its maximum width; fore femur 3× as long as wide; antennae, legs light red………………...……………Prosapanesia lacteipennis Kieffer Descriptive taxonomy Prosapanesia emarginata, Santhosh and Ranjith sp. nov. 6
ACCEPTED MANUSCRIPT (Figs 1, 3A, B, E) Material Examined sea level, sweep net, 20.v.2015. coll. Ranjith, A.P.”
PT
Holotype. Female, “INDIA: Kerala, Wayanad, Kalladi, scrub jungle, 4612 ft. above
RI
Measurements: Body length 2.21 mm; LH 0.43 mm; WH 0.32 mm; LM 0.76 mm; LP 0.34
SC
mm; LPD 0.27 mm; WPD 0.17 mm.
Colour. Head black; antennae, mandibles and clypeus yellow; median clypeal ridge and
NU
anterior truncated margin dark brown; mesosoma black except dark brown pronotal collar and propodeum, posterior margin of pronotum, mesonotum, propleuron and mesopleuron brown,
MA
legs yellow except fore coxa brownish yellow, metasoma brownish yellow except darker T1, dorso medially black at base.
Head. Head sub-rectangular in full face view, polished, moderately coriaceous, WH 0.74× LH;
D
compressed in lateral view, 1.87× as long as high; frons, vertex moderately coriaceous,
TE
sparsely punctate with erect setae; vertex convex in dorsal view; in full face view, anterior clypeal margin emarginate, truncate, without denticles are on either side of the median clypeal
AC CE P
carina, median clypeal carina weak, arched, basal groove indistinct; mandible with long erect setae, bidentate; eye and ocelli absent; antenna 2.05× as long as head; scape 4.70× as long as its maximum width, 3.5× as long as pedicel; pedicel 1.44× as long as its maximum width, 1.64× as long as first flagellomere; flagellomere wider than long except terminal one, terminal flagellomere acute apically, 1.77× as long as its maximum width. Mesosoma. Sparsely coriaceous with sparse suberect setae; pronotal collar anteriorly carinate, setose; pronotum 1.20× as long as its maximum width, convex anteriorly, widened posteriorly; mesonotum strongly constricted posteriorly, 1.03x as long as basal width; notauli and parapsidal furrows absent; propleuron with a shallow distinct depression antero-medially, not extending to the posterior margin in lateral view; propodeal disc constricted, parallel anteriorly with two diverging processes in the anterior end joining posterior apex of mesonotum medially, propodeum 1.65× as long as its maximum width, minimum width 0.34x maximum width, propodeal formula 7.8:7.2:17.5, sublateral posterior margin without long erect setae, propodeal declivity glabrous; wings absent; forefemur 2.4× as long as its 7
ACCEPTED MANUSCRIPT maximum width, mid femur swollen; mid tibia with long erect spines; hind tibia with long setae.
PT
Metasoma. Faintly coriaceous and punctate; 2.50× as long as its maximum width, 1.42× as long as mesosoma, 0.97× as long as head plus mesosoma; T1 smooth with erect setae and pair
RI
of distinct pits laterally; T2 punctate, with erect hairs, glabrous basally; T3 punctate, sparsely
SC
setose, glabrous basally; rest of the tergite sparsely setose. Male
NU
Unknown.
MA
Etymology
The species name refers to the emarginated structure of anterior clypeal margin.
AC CE P
Remarks
TE
India-Kerala (Fig 4).
D
Distribution
Prosapanesia emarginata sp. nov. differs from the remaining female Prosapanesia spp. in having anterior clypeal margin concave, scape more slender (4.7× as long as its maximum width) and propodeum less than 1.7× as long as its maximum width. This species comes close to P. lacteipennis in having mandible bidentate, head moderately smooth. It differs in the following characters, antennae dark yellow (light red in P. lacteipennis), fore femur 2.4× as long as wide (3.0× as long as wide in P. lacteipennis). It differs from P. orientalia sp. nov. in having head and mesosoma moderately coriaceous (strongly coriaceous in P. orientalia sp. nov.), mandible bidentate (tridentate in P. orientalia sp. nov.), lateral clypeal margin without denticles (with denticles in P. orientalia sp. nov.), medial clypeal carina without groove basally (medial clypeal carina with carinate groove basally in P. orientalia sp. nov.), scape distinctly slender, 4.7× as long as its maximum width (3.26× as long as its maximum width in P. orientalia sp. nov.), propodeal disc parallel anteriorly (distinctly constricted anteriorly in P. orientalia sp. nov.), sub lateral posterior margin of propodeum glabrous (sub lateral posterior margin of propodeum with few long erect setae in P. orientalia sp. nov.), metasoma 2.5× as 8
ACCEPTED MANUSCRIPT long as its maximum width (2.2× as long as its maximum width in P. orientalia sp. nov.), T2 punctate (T2 smooth in P. orientalia sp. nov.).
PT
Prosapanesia orientalia, Ranjith and Santhosh sp. nov. (Figs 2, 3C, D, F)
RI
Material Examined
Holotype. Female, “INDIA: Kerala, Calicut, Janakikkadu, scrub jungle, sweep net,
SC
23.xii.2014. coll. Ranjith, A.P.”
NU
Paratypes. 2 Females, “INDIA: Kerala, Malappuram, Calicut University Botanical Garden 11°7’N 75° 5’E, Sweep net, 26.v.2004. coll. S. Santhosh”
mm; LPD 0.35 mm; WPD 0.28 mm.
MA
Measurements. Body length 2.88 mm; LH 0.59 mm; WH 0.46 mm; LM 1.02 mm; LP 0.43
D
Colour. Head black; antennae yellow; mandibles and clypeus reddish brown; median clypeal
TE
ridge and anterior truncated margin dark brown; mesosoma black except brown pronotal collar, posterior margin of pronotum, mesonotum, propleuron and mesopleuron brown, legs
black.
AC CE P
yellow except forecoxa reddish brown, metasoma brown except base of T1 dorso medially
Head. Head sub-rectangular in full face view, coriaceous, WH 0.78× LH; moderately compressed in lateral view, 1.83× as long as high; frons, vertex strongly coriaceous, sparsely punctate with erect setae; vertex straight in dorsal view; anterior clypeal margin thickened, distinctly truncate, three dark brown colored denticles are on either side of the truncated clypeus, median clypeal carina strong, with arched groove with carinate margins basally; mandible with long erect setae, tetradentate with innermost two teeth smaller and blunt; eye and ocelli absent; antenna 1.58× as long as head; scape 3.26× as long as its maximum width, 3.7× as long as pedicel; pedicel 1.32× as long as its maximum width, 1.64× as long as first flagellomere; flagellomere wider than long except terminal one, terminal flagellomere acute apically, 1.53× as long as its maximum width. Mesosoma. Strongly coriaceous with sparse erect setae except otherwise mentioned; pronotal collar anteriorly carinate, asetose; pronotum 1.03x as long as its maximum width, convex 9
ACCEPTED MANUSCRIPT anteriorly, slightly widened posteriorly; mesonotum strongly constricted posteriorly, 1.03x as long as basal width; notauli and parapsidal furrows absent; propleuron with a distinct depression antero-medially, not extending to the posterior margin in lateral view; propodeal
PT
disc constricted anteriorly with two diverging processes in the anterior end joining posterior apex of mesonotum medially, propodeum 1.49× as long as its maximum width, minimum
RI
width 0.33x maximum width, propodeal formula 12:10:29, sub lateral posterior margin with
SC
few long erect setae, propodeal declivity glabrous; wings absent; fore femur 2.7× as long as its maximum width, mid femur swollen; mid tibia with long erect spines; hind tibia with long
NU
setae.
Metasoma. Coriaceous and polished except otherwise mentioned; 2.2× as long as its
MA
maximum width, 1.2× as long as mesosoma, 0.81× as long as head plus mesosoma; T1 smooth with erect setae and a pair of distinct pits laterally; T2 & T3 sparsely setose, glabrous
D
basally; rest of the tergite sparsely setose.
TE
Male
Etymology
AC CE P
Unknown.
The name refers to the distribution of the species, the only Prosapanesia known from the Oriental region. Distribution
India-Kerala (Fig 4). Variations Mandibles and clypeus yellow with mandible base and teeth dark brown; median clypeal ridge and anterior truncated margin brown; pronotal collar, posterior margin of pronotum, mesonotum, propleuron and mesopleuron paler, legs yellow except darker fore coxa; terminal flagellomere 1.76× as long as its maximum width; vertex slightly emarginate in full face view; propodeum 1.3× as long as its maximum width.
10
ACCEPTED MANUSCRIPT Remarks Prosapanesia orientalia sp. nov. comes close to P. lacteipennis in having head sparsely
PT
punctate, lower tooth more acute and longer, pronotum 1.3× as long as its apical width, mesonotum narrower than pronotum, mesopleuron with large dorsal area. But P. orientalia sp.
RI
nov. differs from P. lacteipennis in having head distinctly coriaceous (head moderately coriaceous in P. lacteipennis), antennae yellow (light red in P. lacteipennis), mandible
SC
tetradentate (bidentate in P. lacteipennis), lower margin of truncation distinctly emarginated (slightly emarginate in P. lacteipennis), pronotum distinctly convex (pronotum somewhat
NU
straight anteriorly in P. lacteipennis), propodeum not rounded posteriorly (evenly rounded posteriorly in P. lacteipennis), scape distinctly robust, 3.78–3.85× as long as its maximum
MA
width (scape slender, 4.2× as long as wide in P. lacteipennis), legs except fore coxa yellow (legs light red in P. lacteipennis), fore femur slightly robust, 2.6–2.7× as long as its maximum
D
width (slender 3× as long as wide in P. lacteipennis), metasoma dark yellow, 2.3–2.9× as long as its maximum width (metasoma light red, 2.6× as long as its maximum width), T3+
AC CE P
Discussion
TE
coriaceous (shiny in P. lacteipennis).
Distribution of Prosapanesia and its congeners Though, the non pristocerine genera shows strong endemism towards the Neotropical and Palearctic regions (Gordh and Móczár 1990), many pristocerine wasps are found endemic to the Afrotropical region (Gordh and Móczár 1990) and according to Terayama (2003) eight genera are endemic to the Afrotropics. Female pristocerine wasps except Afgoiogfa and Prosapanesia are widely distributed in the tropical and temperate regions (Gordh and Móczár 1990; Terayama 2003). Species of Prosapanesia are distributed in the southwest region of Africa (Kieffer 1910, 1914; Benoit 1981; Krombein 1989; Terayama 2004). Even though the non pristocerine genera like Trachepyris Kieffer and Pycnomesitius Móczár have shared Afrotropical and Oriental endemism in their distribution (Terayama 2003), no pristocerines are so far found to have their distribution restricted to both the regions. 11
ACCEPTED MANUSCRIPT The discovery of Oriental range extension of Prosapensia throws light on a strong possibility of existence of other endemic Afrotropical pristocerines in the Oriental region. The above possibility is supported by the previous discoveries of Protisobrachium (Terayama 1995), and
PT
Parascleroderma Kieffer (Terayama 1998), which are the congeners of Prosapanesia. The unidentified species of Protisobrachium (Santhosh unpublished) support the extension of
RI
distribution status of the genus, which is found to be endemic to Afrotropical region earlier.
SC
Acknowledgements
NU
This work is part of a research project funded by Department of Science and Technology, Government of India under SERB-Young Scientist Scheme (SB/YS/LS-42/2014). We are
MA
indebted to the authorities of Malabar Christian College, Calicut for providing the facilities. References
D
Azevedo, C. O., Barbosa, D. N., 2010. Two new species Pararhabdepyris Gorbatovskii
TE
(Hymenoptera, Bethylidae) from Australia and Thailand. Zootaxa. 2668, 55–62.
AC CE P
Azevedo, C. O., Garcia, R., Gobbi, F. T., Noort, S. van, 2015. Insecta, Hymenoptera, Bethylidae: range extension and filling gaps in Central African Republic. Check List. 11 (2), 1–4. doi:10.15560/11.2.1606 Benoit, P. L. G. , 1981. Bethylidae africains (Hymenoptera). La Tribu Usakosiini. Revue de Zoologie Africaine. 95 (4), 833–842. Evans, H. E., 1963. A revision of the genus Apenesia in the Americas (Hymenoptera, Bethylidae). Bulletin of the Museum of Comparative Zoology. 130, 251–359. Evans, H. E., 1969. Phoretic copulation in Hymenoptera. Entomological News. 80,113–124. Evans, H. E., 1961. A revision of the genus Pseudisobrachium in North and Central America (Hymenoptera, Bethylidae). Bulletin of the Museum of Comparative Zoology. 126, 211–318. Evans, H. E., 1964. A synopsis of the American Bethylidae (Hymenoptera: Aculeata). Bulletin of the Museum of Comparative Zoology. 132 (1), 1–222. 12
ACCEPTED MANUSCRIPT Finammore, A. T., Gauld, I. D., 1995. Chrysididae, in: Hanson, P.E. and Gauld, I.D. (Eds.), The Hymenoptera of Costa Rica. Oxford: Oxford University Press, pp. 479–488.
PT
Gordh, G., 1990. Apenesia evansi sp. n. (Hymenoptera: Bethylidae) from Australia with comments on phoretic copulation in Bethylids. Journal of the Australian
RI
Entomological Society. 29, 167–170. doi:10.1111/j.1440-6055.1990.tb00341.x
SC
Gordh, G., Móczár, L., 1990. A catalog of the world Bethylidae (Hymenoptera: Aculeata). Memoirs of the American Entomological Institute. 46, 1–364.
NU
Harris, R. A., 1979. A glossary of surface sculpturing. Occasional Papers in Entomology. 28,
MA
1–31.
Kieffer, J. J., 1910. Description de nouveaux Béthylides (Hymén.). Annales de la Société Entomologique de France. 79, 31–56.
D
Kieffer, J. J., 1914. Bethylinae, in: Das Tierreich. Vol. 41. R. Friedlander und Sohn, Berlin, pp.
TE
228–595.
AC CE P
Krombein, K. V., 1989. Systematic notes on some Bethylidae from Botswana: Pristocerinae (Hymenoptera: Aculeata). Proceedings of the Entomological Society of Washington. 91: 620–631.
Lim, J., Terayama, M., Koh, S., Lee, J., Lee, S., 2011. A taxonomic review of the subfamily Pristocerinae (Hymenoptera: Chrysidoidea: Bethylidae) from Korea with descriptions of
two
new
species.
Journal
of
Natural
History.
45
(7-8),
435–460.
doi:10.1080/00222933.2010.534191 Liu, J., Chen, H., Xu, Z., 2011. Two new species of genus Foenobethylus Kieffer, 1913 (Hymenoptera, Bethylidae) from China with a key to the known species. Zootaxa. 2806, 53–59. Savergnini, J.R., Azevedo, C.O. 2013. Taxonomy of Foenobethylus Kieffer (Hymenoptera, Bethylidae) with description of two new species. Journal of Asia-Pacific Entomology. 16, 433–441. doi:10.1016/j.aspen.2013.06.004
13
ACCEPTED MANUSCRIPT Shorthouse, D.P. 2010. SimpleMappr, an online tool to produce publication-quality point maps. Available from http://www.simplemappr.net (accessed on 2 February 2016).
PT
Tachikawa, T.,1985. On the bethylid wasps (II). Forest Pests. 34(9), 161–169 (in Japanese). Terayama, M., 1995a. Discovery of the genus Protisobrachium Benoit 1957 from the Oriental
RI
region (Hymenoptera: Bethylidae). Journal of Entomological Science. 30 (4), 443–446.
SC
Terayama, M., 1995b. Caloapenesia and Neoapenesia, new genera of the family Bethylidae (Hymenoptera, Chrysidoidea) from the Oriental region, with proposals of two new
NU
synonymies of genera. Japanese Journal of Entomology. 63 (4), 881–891.
MA
Terayama, M., 1996a. Discovery of the Genus Glenosema from the Oriental Region with Descriptions of Three New Species (Hymenoptera: Bethylidae). The Florida Entomologist. 79 (4), 591–595.
D
Terayama, M., 1996b. The phylogeny of the bethylid wasp subfamily Pristocerinae
TE
(Hymenoptera, Bethylidae). Japanese Journal of Entomology. 64, 587–601.
AC CE P
Terayama, M., 1998. Discovery of the genus Parascleroderma Kieffer from the Oriental region, with descriptions of four new species (Hymenoptera: Bethylidae). Entomological Science. 1(1), 129–132. Terayama, M., 2003. Phylogenetic systematic of the family Bethylidae (Insecta: Hymenoptera). Part II. Keys to subfamilies, tribes and genera in the world. Academic Reports, Faculty of Engineering, Tokyo Polytechnic University. 26 (1), 16–29. Terayama, M., 2004. Descriptions of new taxa and distribution records of the family Bethylidae (Insecta, Hymenoptera) I. Subfamily Pristocerinae. Academic Reports, Faculty of Engineering, Tokyo Polytechnic University. 27 (1), 22–38. Terayama, M., 2006. The Insects of Japan. Vol. I, Bethylidae (Hymenoptera). Fukuoka, (Japan): Entomological Society of Japan. 1–319. Várkonyi, G., Polaszek, A., 2007. Rediscovery and revision of Foenobethylus Kieffer, 1913 (Hymenoptera, Bethylidae). Zootaxa. 1546, 1–14. 14
ACCEPTED MANUSCRIPT
FIGURE CAPTIONS
PT
figure 1. Prosapanesia emarginata sp. nov., female, holotype (A) Habitus, lateral view; (B) Head, dorsal view; (C) Mesosoma, dorsal view; (D) Propodeum, dorsal view; (E) Propodeum
RI
& metasoma (in part), dorsal view; (F) metasoma, lateral view.
SC
figure 2. Prosapanesia orientalia sp. nov., female, holotype (A) Habitus, lateral view; (B) Head, dorsal view; (C) Mesosoma, dorsal view; (D) Propodeum, dorsal view; (E) Propodeum
NU
& metasoma (in part), dorsal view; (F) metasoma, lateral view.
MA
figure 3. (A, B) Prosapanesia emarginata sp. nov., head oblique views; (C, D) P. orientalia sp. nov.; (E) P. emarginata sp. nov., metasoma, dorsal view; (F) P. orientalia sp. nov., metasoma, dorsal view.
AC CE P
TE
D
figure 4. Distribution map of Prosapanesia species
15
NU
SC
RI
PT
ACCEPTED MANUSCRIPT
AC CE P
TE
D
MA
Graphical abstract
16
AC CE P
TE
D
MA
NU
SC
RI
PT
ACCEPTED MANUSCRIPT
17
ACCEPTED MANUSCRIPT
AC CE P
TE
D
MA
NU
SC
RI
PT
Fig. 1
18
AC CE P
TE
D
MA
NU
SC
RI
PT
ACCEPTED MANUSCRIPT
19
ACCEPTED MANUSCRIPT
AC CE P
TE
D
MA
NU
SC
RI
PT
Fig. 2
20
AC CE P
TE
D
MA
NU
SC
RI
PT
ACCEPTED MANUSCRIPT
21
ACCEPTED MANUSCRIPT
AC CE P
TE
D
MA
NU
SC
RI
PT
Fig. 3
22
AC CE P
Fig. 4
TE
D
MA
NU
SC
RI
PT
ACCEPTED MANUSCRIPT
23
ACCEPTED MANUSCRIPT Highlights
PT
Rarely collected bethylid genus Prosapanesia is discovered from Oriental region Prosapanesia emarginata sp. nov. and P. orientalia sp. nov. are described
RI
A world key for Prosapanesia is provided
SC
Male and female generic diagnosis of Prosapanesia are redefined
AC CE P
TE
D
MA
NU
Oriental presence of more endemic Afrotropical genera is suggested
24