Research Article |
Corresponding author: Alain Roques ( alain.roques@orleans.inra.fr ) Academic editor: Michael Engel
© 2016 Alain Roques, Robert S. Copeland, Laurent Soldati, Olivier Denux, Marie-Anne Auger-Rozenberg.
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:
Roques A, Copeland RS, Soldati L, Denux O, Auger-Rozenberg M-A (2016) Megastigmus seed chalcids (Hymenoptera, Torymidae) radiated much more on Angiosperms than previously considered. I- Description of 8 new species from Kenya, with a key to the females of Eastern and Southern Africa. ZooKeys 585: 51-124. doi: 10.3897/zookeys.585.7503
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A survey of seed chalcids from woody plants in Kenya revealed 12 species belonging to the genus Megastigmus Dalman, 1820, and has increased to 16 the number of Megastigmus species presently recorded from the Afrotropical Region, of which at least 13 are seed feeders. A key to female Megastigmus of the Afrotropical Region is provided. Eight new species are described from morphological evidence: M. lanneae Roques & Copeland, M. laventhali Roques & Copeland, M. ozoroae Roques & Copeland, and M. smithi Roques & Copeland in seeds of species of the family Anacardiaceae, M. copelandi Roques & Copeland and M. grewianae Roques & Copeland in seeds of Malvaceae, M. helinae Roques & Copeland in seeds of Rhamnaceae, and M. icipeensis Roques & Copeland for which no host is known. These collections include the first records of Malvaceae and Rhamnaceae as hosts of Megastigmus seed chalcids, which appear to have radiated in Angiosperms much more than previously considered. Analyses of the mitochondrial (cytochrome oxidase subunit one – COI) and nuclear DNA (28S ribosomal region) could be carried out on 8 of the 16 African species of which 5 were newly described ones. The species associated with Anacardiaceae always clustered together in phylogenies, confirming the existence of a strong and ancestral monophyletic clade, unlike the ones associated with Malvaceae and Rhamnaceae, whose position remains unclear. All holotypes are deposited in the National Museums of Kenya.
Chalcidoidea, Megastigmus, seed, Anacardiaceae, Malvaceae, Rhamnaceae, new species, Africa, plant hosts, fruit
Globalization, the accelerating worldwide movement of goods and people, has led to an exponential increase in the transport of alien organisms across geographical barriers (
Seeds of woody plants can be infested by larvae of a number of arthropod groups, including, among others, dipteran seed midges in the family Cecidomyiidae (e.g., Plemeliella, Mayetiola), coleopteran seed beetles in the family Bruchidae, and mites in the genus Trisetacus (Nalepellidae) (
Data from these recent studies indicate that, worldwide, the genus Megastigmus currently includes 135 species, 70 of which are seed feeders, 45 develop in galls, and 20 for which behavioral data is lacking (
To date, relatively few species have been reported from subtropical and tropical areas. In the Afrotropical region, only seven species have been recognized. Four species develop as phytophages in angiosperm seeds. Three of these, Megastigmus hypogeus (Hussey), M. thomseni (Hussey), and M. transvaalensis (Hussey) (= M. rhusi [Hussey]) attack seeds of Anacardiaceae (
The lower diversity of Megastigmus seed chalcids observed in angiosperms and in subtropical/ tropical areas may be due to a more limited radiation process in these hosts and regions or simply reflect more limited investigations in the tropics compared to those carried out in conifers of the Holarctic region in relation to afforestation programs and establishment of seed orchards (
Over 700 insect species were reared from fruits and much of the data from the two projects has been published previously (e.g.
During the 1999–2004 wild-fruit survey, 3839 fruit collections were made throughout much of Kenya. An additional 347 wild-fruit collections were made during the 2011–2012 survey in the Taita Hills, bringing the total number of fruit samples to 4186, the great majority from woody plants. Over the course of the two surveys approximately 930 species of plants were sampled, representing 122 families. Details of fruit collection and insect rearing methods are available in
During transport, damage to fruits was minimised by placing each fruit sample in a separate polythene bag and suspending the bag above the bottom of a two-litre plastic storage container. A tight-fitting plastic lid fixed the bag and its contents in place. A large rectangular piece had been cut out of the lid and replaced with fine mesh cloth to allow for ventilation of the fruits. In the laboratory, fruits were removed from transport bags and placed within one-litre, rectangular plastic containers that had small elliptical holes cut out of the bottom. Each one-litre container (also provided with a mesh-covered lid) was nested within a fresh two-litre container, the bottom of which had a layer of heat-treated sand. Fruits were usually held for up to two months. Emerged adult insects were held for 1–3 days before being killed.
Beentje (1994) and Agnew and Agnew (1994) were used for preliminary plant identifications. Identifications made by the second author were confirmed or corrected by Quentin Luke of the East African Herbarium. Plant names and authors correspond to those found in The
Independent of the wild-fruit surveys, at various times between 2005 and 2014, the second author surveyed Kenyan insects using Townes-style Malaise traps (
Adult morphology was examined using a MZ12 Leica stereomicroscope equipped with a Leica IC A camera. When more than three specimens were obtained per species and sex, one individual of each sex was dissected and wings of both sexes and male genitalia were mounted on glass slides using Dimethyl hydantoin formaldehyde (DMHF;
In a few specimens, head, antenna and thoracic sculpture were examined using a Cambridge StereoscanB 90 scanning electron microscope equipped with OrionB® image processing software. When less than three specimens were available per species and sex, these specimens were kept intact and mounted on triangular cardboards. Photographs and measurements of body and body parts (wings, antenna, head, thorax, gaster and ovipositor) were made as above without dissecting the mounted specimens. Morphological terminology follows
• DNA extraction, molecular markers, amplification and sequencing
In order to include the Kenyan species in an existing phylogeny of Megastigmus, we used mtDNA sequences previously obtained by
In addition to the COI gene, a nuclear fragment, the D2 region of the 28S ribosomal subunit (rDNA), was used to build a phylogenetic tree of the studied Megastigmus species. Nuclear primers, D1F (5’-ACCCGCTGAATTTAAGCATAT-3’) and D3R (5’-TAGTTCACCATCTTTCGGGTC-3’), previously used for reconstructing a molecular phylogeny of Megastigmus spp. on conifers (
• Phylogenetic analyses
For all datasets, sequences were aligned using Clustal W (
Phylogenetic analyses were performed using maximum likelihood (ML) inference with PhyML v3.0 (
Abbreviations used for insect specimen depositories or private collections cited in the study are as follows:
NMKE National Museums of Kenya
RSC Robert Copeland personal collection, ICIPE, Nairobi, Kenya
ARC Alain Roques personal collection, INRA Zoologie Forestière Orléans, France
ICIPE International Centre of Insect Physiology and Ecology, Nairobi, Kenya
SAMC IZIKO South African Museum, Capetown, South Africa
BNHM British National History Museum, London, UK
Megastigmus wasps were obtained from 31 of the fruit samples (0.74%- Table
Fruit and Malaise trap samples producing Megastigmus species in Kenya.
Family / Host plant species | Megastigmus species | Region | Location | Latitude N | Longitude E | Elevation1 (m) |
---|---|---|---|---|---|---|
Anacardiaceae | ||||||
Lannea cf. schimperi (A.Rich.) Engl. | M. smithi Roques & Copeland, sp. n. | Eastern Arc Mountains | Sagalla Mountain | -3,4786 | 38,5746 | 1090 |
Lannea rivae (Chiov.) Sacl. | M. lanneae Roques & Copeland, sp. n. | Central highlands | Machakos | -1,5609 | 37,2338 | 1586 |
Lannea schweinfurthii (Engl.) Engl. | M. laventhali Roques & Copeland, sp. n. | Eastern Arc Mountains | Base of Kasigau Mountain | -3,8130 | 38,6405 | 619 |
Ozoroa insignis Del. subsp. reticulata (Bak.f.) Gillett | M. ozoroae Roques & Copeland, sp. n. | Western mid-altitude | Gembe Hills | -0,4894 | 34,2433 | 1362 |
O. insignis subsp. reticulata | M. ozoroae | Western mid-altitude | Sindo-Mbita Road | -0,4849 | 34,1765 | 1205 |
O. insignis subsp. reticulata | M. hypogeus (Hussey) | Western mid-altitude | Sindo-Mbita Road | -0,4849 | 34,1765 | 1205 |
Ozoroa obovata (Oliv.) R. & A. Fernandes | M. hypogeus | Coastal forests | Arabuko-Sokoke Forest | -3,3103 | 39,9978 | 55 |
O. obovata | M. hypogeus | Coastal forests | Arabuko-Sokoke Forest | -3,3716 | 39,8949 | 55 |
O. obovata | M. hypogeus | Coastal forests | Arabuko-Sokoke Forest | -3,2030 | 39,9271 | 98 |
O. obovata | M. hypogeus | Coastal forests | Shimba Hills | -4,1349 | 39,4680 | 285 |
Pistacia lentiscus L. subsp. emarginata (Engl.) AL-Saghir | M. pistaciae Walker | Western highlands | nr. Narok on main road | -1,1023 | 36,0192 | 1919 |
P. lentiscus subsp. emarginata | M. pistaciae | Western highlands | nr. Narok on main road | -1,1243 | 35,9868 | 1830 |
P. lentiscus subsp. emarginata | M. pistaciae | Northern highlands | Mt Kulal | 2,6290 | 36,9278 | 1639 |
P. lentiscus subsp. emarginata | M. pistaciae | Western highlands | Mai Mahiu-Narok Road | -1,1121 | 35,9765 | 1900 |
Rhus natalensis Krauss | M. transvaalensis (Hussey) | Central Highlands | Kikuyu Escarpment | -1,0290 | 36,6025 | 2100 |
R. natalensis | M. transvaalensis | Central Highlands | Kikuyu Escarpment | -1,0717 | 36,6033 | 2100 |
R. natalensis | M. transvaalensis | Central Highlands | Burguret | -0,1128 | 37,0375 | 1960 |
R. natalensis | M. transvaalensis | Northern mid-altitude | south of Marsabi town | 2,2659 | 37,8976 | 1200 |
R. natalensis | M. transvaalensis | Western mid-altitude | Gembe Hills | -0,4894 | 34,2433 | 1370 |
R. natalensis | M. transvaalensis | Eastern Arc Mountains | Ronge-Nyika, mixed shrub- and grassland | -3,3963 | 38,4309 | 1050 |
R. natalensis | M. transvaalensis | Eastern Arc Mountains | Kasigau forest | -3,8194 | 38,6619 | 1280 |
Rhus vulgaris Meikle | M. transvaalensis | Central Highlands | Kikuyu escarpment | -1,0780 | 36,6020 | 2200 |
Schinus molle L. | M. transvaalensis | Central Highlands | Nairobi | -1,2211 | 36,8963 | 1610 |
Schinus terebinthifolius Raddi | M. transvaalensis | Central highlands | Nairobi | -1,2212 | 36,8963 | 1610 |
S. terebinthifolius | M. transvaalensis | Central highlands | Nairobi | -1,2501 | 36,7835 | 1750 |
Cupressaceae | ||||||
Juniperus procera Endl. | M. somaliensis (Hussey) | Central highlands | Mt. Kenya Forest | -0,0308 | 37,1230 | 2130 |
J. procera | M. somaliensis | Northern highlands | Mt. Kulal | 2,6319 | 36,9282 | 1650 |
Rhamnaceae | ||||||
Helinus integrifolius (Lam.) Kuntze | M. helinae Roques & Copeland, sp. n. | Northern highlands | Mathews Range | 1,1777 | 37,3141 | 1340 |
Malvaceae | ||||||
Grewia stuhlmannii K. Schum. | M. grewianae Roques & Copeland, sp. n. | Coastal forests | Arabuko-Sokoke Forest | -3,2997 | 39,9869 | 60 |
Grewia tephrodermis K. Schum. | M. grewianae | Eastern savanna | Tsavo | -2,6760 | 38,3325 | 640 |
Grewia tephrodermis | M. copelandi Roques & Copeland, sp. n. | Eastern Arc Mountains | Mwatate area, on small farm | -3,4894 | 38,3330 | 1050 |
n.a., Malaise trap | M. grewianae | Eastern savanna | Kasaala | -2,0749 | 38,2250 | 740 |
n.a., Malaise trap | M. grewianae | Eastern Arc Mountains | Kasigau Mountain Forest | -3,8270 | 38,6488 | 1070 |
n.a., Malaise trap | M. icipeensis Roques & Copeland, sp. n. | Central Highlands | Nairobi, ICIPE | -1,2232 | 36,8965 | 1600 |
n.a., Malaise trap | M. laventhali | Coastal island | Funzi Island | -4,5778 | 39,4413 | 0 |
A total of 12 Megastigmus species were found during the survey, of which eight were new to science (Table
Megastigmus chalcids were reared from fruits of seven genera of shrubs and trees native to tropical Africa (Table
Anacardiaceae host fruits of Megastigmus spp. in Kenya. 3 Lannea cf. schimperi 4 L. rivae 5 Lannea schweinfurthii 6 Ozoroa insignis subsp. reticulata 7 O. obovata 8 Pistacia lentiscus subsp. emarginata 9 Rhus natalensis (note the chalcid hole in a fruit).
Other host fruits of Megastigmus spp. in Kenya. 10 Helinus integrifolius (Rhamnaceae) 11 Grewia tephrodermis (Malvaceae) 12 G. stuhlmannii 13 Juniperus procera (Cupressaceae).
Host-plant genera of Megastigmus species in Kenya.
Family | Genus | Total no. of Kenyan species | No. of Kenyan species sampled | No. of sampled Kenyan species yielding Megastigmus (%) | Total number of Afrotropical species |
---|---|---|---|---|---|
Anacardiaceae | Lannea 1 | 10 | 5 | 3 (60) | 29 |
Ozoroa1 | 2 | 2 | 2 (100) | 44 | |
Pistacia 1,2 | 1 | 1 | 1 (100) | 2 | |
Rhus 1 | 6 | 4 | 2 (50) | 106 | |
Schinus 1,5 | 25 | 25 | 2 (100) | 25 | |
Cupressaceae | Juniperus 1 | 1 | 1 | 1 (100) | 4 |
Rhamnaceae 3 | Helinus 1,2,4 | 2 | 2 | 1 (50) | 3 |
Malvaceae 3 | Grewia 1,2,4 | 28 | 14 | 2 (14) | 122 |
Of the host families, Anacardiaceae was most important. Including the collections from two exotic Schinus species, Anacardiaceae contributed 10 (67%) of the host fruit species (n=15) from which Megastigmus were reared. Among these plant hosts, Lannea was the most important genus with three of five species sampled producing Megastigmus. Each Lannea species was attacked by a different, previously undescribed chalcid: Lannea cf. schimperi (A.Rich.) Engl. (Figure
Before our study the host of M. hypogeus was an unresolved issue. When transcribing label information in his description of M. hypogeus from specimens then available to him,
Among non-Anacardiaceae host families, two new Megastigmus species were obtained from fruits of two species of Malvaceae in the genus Grewia. Megastigmus grewianae Roques & Copeland, sp. n. emerged from both Grewia tephrodermis K. Schum. (Figure
Table
Host fruit infestation indices for Megastigmus species. Only samples whose fruits were counted are included; no samples of Juniperus procera fruits were counted.
Plant family/ Plant species | No. of fruits | Megastigmus species | No. of reared Megastigmus | % of infestation by Megastigmus |
---|---|---|---|---|
Anacardiaceae | ||||
Lannea cf. schimperi | 786 | M. smithi | 3 | 0,4 |
Lannea rivae | 338 | M. lanneae | 11 | 3,3 |
Lannea schweinfurthii | 168 | M. laventhali | 1 | 0,6 |
Ozoroa insignis subsp. reticulata | 2806 | M. ozoroae | 226 | 8,1 |
Ozoroa obovata | 1505 | M. hypogeus | 42 | 2,8 |
Ozoroa obovata | 698 | M. hypogeus | 1 | 0,1 |
Ozoroa obovata | 2442 | M. hypogeus | 22 | 0,9 |
Pistacia lentiscus subsp. emarginata | 2603 | M. pistaciae | 12 | 0,5 |
Pistacia lentiscus subsp. emarginata | 1019 | M. pistaciae | 1 | 0,1 |
Rhus natalensis | 674 | M. transvaalensis | 41 | 6,1 |
Rhus natalensis | 524 | M. transvaalensis | 6 | 1,1 |
Rhus natalensis | 2394 | M. transvaalensis | 14 | 0,6 |
Rhus natalensis | 894 | M. transvaalensis | 36 | 4,0 |
Rhus natalensis | 412 | M. transvaalensis | 6 | 1,5 |
Rhus vulgaris | 769 | M. transvaalensis | 113 | 14,7 |
Schinus molle | 535 | M. transvaalensis | 79 | 14,8 |
Rhamnaceae | ||||
Helinus integrifolius | 82 | M. helinae | 8 | 9,8 |
Malvaceae | ||||
Grewia tephrodermis | 448 | M. copelandi | 3 | 0,7 |
Grewia tephrodermis | 100 | M. grewianae | 13 | 13,0 |
Grewia stuhlmannii | 1056 | M. grewianae | 29 | 2,7 |
The four Malaise trap samples containing Megastigmus yielded a total of 10 specimens. Given our estimate of 13,740 Malaise trap sampling days, Megastigmus were captured at the very low rate of approximately one specimen per 1374 sampling days. Despite this, a total of three species of Megastigmus were collected, all of them representing previously undescribed species. Single specimens of Megastigmus grewianae Roques & Copeland, sp. n. were collected in Malaise traps set in dry savanna in Eastern Kenya (Kasaala) and in a small campsite clearing in Kasigau Forest (Table
Our fruit collections bring to six the number of Afrotropical plant families that are hosts of Megastigmus, including Rosaceae which are attacked by the introduced species, M. aculeatus. The listing, in a summary table, by
Table
MtDNA was sequenced for nine of the 12 African species defined morphologically, including five of the eight new species and the four previously described ones. Using the longest mitochondrial marker (“Clyde-Bonnie”), 810 bp- long COI sequences were obtained for M. helinae (1 specimen), M. grewianae (4 specimens), M. lanneae (1 specimen), M. pistaciae (2 specimens), M. somaliensis (1 specimen) and M. zebrinus (1 specimen). In the case of M. hypogeus and M. ozoroae, six specimens per species were extracted but we obtained only a short fragment of 417 bp with the internal primer “Jerry” for one individual of M. ozoroae, and for two of M. hypogeus. Similarly, we obtained this short fragment for the sole specimen of M. icipeensis. DNA from the other species was amplified with the different sets of primers and, although we obtained data from more species with the shorter fragment than with the longer, we studied and compared the two datasets because some short sequences showed the presence of overlapping fragments (for less than 2% of the short sequence) which could be due to heteroplasmy or numt’s (nuclear copies of mitochondrial DNA), even if they were of the correct length and reading frame and contained no stop codons. These trends were observed in M. lanneae and M. ozoroae and their occurrence seemed to be species specific. These problems (no amplification and/or overlapping fragments) could be due to the preservation quality of the specimens. For M. transvaalensis, we used sequences from specimens previously obtained from Morocco, because we failed to amplify any Kenyan specimens. Megastigmus copelandi, M. laventhali and M. smithi were not analyzed molecularly because of the limited number of available specimens necessary to define holotype and paratypes. For the nuclear marker 28S, one specimen per species was successfully amplified for all the species analyzed in the COI dataset except M. icipeensis, and an alignment of 924 bp including gaps was obtained.
The inclusion of the nine Kenyan species in an already existing molecular phylogeny (
Mean genetic distances within and between groups of Megastigmus sequences. For each cell: COI long fragment before backslash (/), and COI short fragment after backslash (in this case, the group “unknown” is corresponding to the only specimen of M. icipeensis, caught in malaise trap and only sequenced in short fragment).
Pinaceae | Cupressaceae | Taxodiaceae | Rosaceae | Tiliaceae | Rhamnaceae | Anacardiaceae | Gall-former | unknown | outgroup | |
---|---|---|---|---|---|---|---|---|---|---|
Pinaceae | 0.061\0.057 | |||||||||
Cupressaceae | 0.088\0.073 | 0.060\0.046 | ||||||||
Taxodiaceae | 0.081\0.074 | 0.084\0.074 | -\- | |||||||
Rosaceae | 0.085\0.074 | 0.093\0.080 | 0.086\0.077 | 0.079\0.068 | ||||||
Tiliaceae | 0.098\0.086 | 0.089\0.080 | 0.094\0.083 | 0.102\0.087 | -\- | |||||
Rhamnaceae | 0.113\0.095 | 0.112\0.105 | 0.107\0.096 | 0.108\0.088 | 0.111\0.097 | -\- | ||||
Anacardiaceae | 0.111\0.096 | 0.112\0.099 | 0.099\0.104 | 0.108\0.100 | 0.111\0.095 | 0.107\0.078 | 0.068\0.063 | |||
Gall-former | 0.102\0.092 | 0.100\0.085 | 0.094\0.096 | 0.100\0.086 | 0.106\0.080 | 0.094\0.086 | 0.105\0.092 | -\- | ||
Unknown | -\0.094 | -\0.089 | -\0.096 | -\0.094 | -\0.096 | -\0.083 | -\0.082 | -\0.083 | ||
Outgroup | 0.157\0.127 | 0.151\0.123 | 0.158\0.135 | 0.157\0.134 | 0.161\0.132 | 0.140\0.113 | 0.146\0.115 | 0.154\0.121 | -\0.113 | -\- |
Between groups, similar values were observed between Holarctic groups, as well as between the species related to Malvaceae and the Holarctic groups, whereas pairwise K2P distances showed higher values for the other host families. In the COI-long fragment dataset, the values between M. somaliensis and the two other species related to the genus Juniperus (Cupressaceae) were 3.5%. The two specimens identified as M. pistaciae (one from Greece and one from Kenya) diverged by 4.5%, suggesting that the two are probably sibling species.
The use of maximum likelihood and Bayesian inference methods resulted in similar information about phylogenetic relationships within the genus Megastigmus. Therefore, we present only the phylogenetic trees built with the Bayesian inference method (Figure
Bayesian-likelihood inference phylogenies based on cytochrome oxidase I (COI) and ribosomal DNA (28S) sequences in seed-specialized wasps of the Megastigmus genus. A Short fragment COI (34 taxa, 417bp) B Long fragment COI (31 taxa, 810pb) C Nuclear fragment (33 taxa, 924pb). Torymus azureus Boheman, 1834 was used as an outgroup. Posterior probability values are indicated at each node. New taxa described in this paper are indicated in red and bold. Branches of taxa associated with Anacardiaceae as host plant family are in blue, and branches of taxa associated with Cupressaceae are in green.
In the different trees, robust clades (Pinaceae, Cupressaceae, Rosaceae, Anacardiaceae) included several species, whereas the positions of single species varies depending of the marker. Whatever the marker and method, the Megastigmus species associated with Anacardiaceae clustered together in a highly supported monophyletic clade (Figure
Despite amplifying for all markers used, the phylogenetic position of M. zebrinus remains uncertain. According to
The mtDNA sequences and nuclear sequences determined in the course of this study are registered under the following GenBank accession numbers: KU984677 to KU984706.
In some Megastigmus species, males, unlike females, are highly variable in size and colour. This is observed especially in species associated with Anacardiaceae (e.g., M. transvaalensis;
Therefore, we provide a key to females only, including the species associated with rose hips, conifers, and galls that were not collected during the surveys (i.e. M. aculeatus, M. thomseni, M. pretorianensis and M. zebrinus). We also include M. asir Ghramh & Shati recorded from seeds of Juniperus procera in a region of Saudi Arabia facing Eritrea and Ethiopia (
1 | Ovipositor sheaths shorter (0.4–0.9×) than gaster length (Figures |
2 |
– | Ovipositor sheaths longer (1.1×–1.8×) than gaster length (Figures |
5 |
2 | Ovipositor sheaths much shorter (0.4–0.6×) than gaster length (Figures |
3 |
-– | Ovipositor sheaths 0.8–0.9× as long as gaster length (Figures |
4 |
3 | Body colour nearly entirely orange-yellow; only a narrow black band along anterior suture of pronotum (Figures |
M. helinae Roques & Copeland, sp. n. |
- | Body colour with a large longitudinal, median rufous band extending from pronotum to scutellum (Figure |
M. smithi Roques & Copeland, sp. n. |
4 | Pilosity on head entirely pale (Figure |
M. copelandi Roques & Copeland, sp. n. |
– | Pilosity pale on face but dark on other parts of head (Figure |
M. ozoroae Roques & Copeland, sp. n. |
5 | Minute species, body length less than 1.5 mm; Thorax pilosity pale; 2 pairs of hairs on scutellum (Figure |
6 |
– | Larger species, body length more than 2.5 mm; Thorax pilosity dark; from 3 to 8 pairs of hairs on scutellum (Figures |
7 |
6 | Body colour mostly black | M. pretorianensis Doǧanlar |
– | Body colour nearly completely orange-yellow (Figure |
M. icipeensis Roques & Copeland, sp. n. |
7 | Thorax colour orange or pale yellow without extensive dark markings (Figures |
8 |
– | Thorax with extensive dark markings (Figures |
12 |
8 | Ovipositor sheaths less than 1.2× as long as gaster length (Figure |
9 |
– | Ovipositor sheaths at least 1.4× as long as gaster length (Figures |
10 |
9 | Forewing with an elongate stigma, 1.7× as long as wide | M. thomseni (Hussey) |
– | Forewing with an oval- rounded stigma, 1.2–1.3 × as long as wide (Figure |
M. pistaciae Walker |
10 | Ovipositor sheaths at least 1.8 × as long as gaster length and 0.8× as long as body length (Figure |
M. laventhali Roques & Copeland, sp. n. |
– | Ovipositor sheaths at most 1.6× as long as gaster length and 0.6× as long as body length (Figure |
11 |
11 | Forewing stigma oblong, 1.7 × as long as wide, with a very short stigmal vein, at most 0.2× as long as stigma length; gaster tergites pale yellow with translucent median longitudinal stripes, transversely infumate | M. zebrinus Grissell |
– | Forewing stigma elongate-oval, 2.0 × as long as wide, with an elongate stigmal vein, 0.4 × as long as stigma length (Figure |
M. transvaalensis (Hussey) |
12 | Ovipositor sheaths never longer than 1.5× gaster length (Figures |
13 |
– | Ovipositor sheaths at least 1.8× longer than gaster length (Figures |
15 |
13 | Pronotum with a large median dark brown band progressively decreasing in size from the anterior to posterior suture (Figure |
M. hypogeus (Hussey) |
– | Pronotum yellow or brownish, without median dark band; scutellum with 7–8 pairs of hairs; in seeds of Juniperus procera | 14 |
14 | Propodeum smooth; forewing stigma elongate, more than 2× as long as broad | M. asir Ghramh & Shati |
– | Propodeum rugose, with reticulate striae (Figure |
M. somaliensis Hussey |
15 | Ovipositor sheaths at least 1.2× longer than body length. In Rosa seeds | M. aculeatus Swederus |
– | Ovipositor sheaths never longer than body length. In Malvaceae or Anacardiaceae seed | 16 |
16 | Pronotum orange with a conspicuous longitudinal black stripe on each side (Figures |
M. grewianae Roques & Copeland, sp. n. |
– | Pronotum without lateral black stripes but with a large, nearly rectangular dark brown band medially extending from anterior to posterior suture (Figures |
M. lanneae Roques & Copeland, sp. n. |
Megastigmus helinae Roques & Copeland, sp. n. female. 15 dorsal view of the body 16 lateral view of body 17 dorsal view of thorax 18 front view of head 19 electroscan of antenna 20 electroscan of dorsal view of thorax 21 forewing.
Megastigmus helinae Roques & Copeland, sp. n. male. 22 dorsal view of the body 23 lateral view of body 24 dorsal view of thorax 25 front view of head 26 electroscan of antenna 27 electroscan of dorsal view of thorax 28 forewing 29 genitalia.
Megastigmus smithi Roques & Copeland, sp. n. female. 30 dorsal view of the body 31 lateral view of body 32 dorsal view of thorax 33 front view of head 34 antenna 35 forewing.
Megastigmus copelandi Roques & Copeland, sp. n. female. 36 dorsal view of the body 37 lateral view of body 38 dorsal view of thorax; 39 front view of head 40 antenna 41 forewing.
Megastigmus ozoroae Roques & Copeland, sp. n. female. 42 dorsal view of the body 43 lateral view of body 44 dorsal view of thorax 45 front view of head 46 antenna 47 forewing.
Megastigmus ozoroae Roques & Copeland, sp. n. male pale form. 48 dorsal view of the body 49 lateral view of body 50 dorsal view of thorax; 51 front view of head 52 antenna 53 genitalia 54 forewing.
Megastigmus ozoroae Roques & Copeland, sp. n. male dark form. 55 dorsal view of the body 56 lateral view of body 57 dorsal view of thorax; 58 front view of head 59 antenna 60 genitalia 61 forewing.
Megastigmus icipeensis Roques & Copeland, sp. n. female. 62 dorsal view of the body 63 lateral view of body 64 dorsal view of thorax 65 front view of head 66 antenna 67 forewing.
Megastigmus icipeensis Roques & Copeland, sp. n. male. 68 dorsal view of the body 69 lateral view of body 70 dorsal view of thorax 71 front view of head 72 antenna 73 forewing 74 genitalia.
Megastigmus laventhali Roques & Copeland, sp. n. female. 75 dorsal view of the body 76 lateral view of body 77 dorsal view of thorax 78 front view of head 79 antenna 80 forewing.
Megastigmus grewianae Roques & Copeland, sp. n. female. 81 dorsal view of the body 82 lateral view of body 83 dorsal view of thorax 84 electroscan of dorsal view of thorax 85 front view of head 86 electroscan of antenna 87 forewing.
Megastigmus grewianae Roques & Copeland, sp. n. male. 88 dorsal view of the body 89 lateral view of body 90 dorsal view of thorax 91 front view of head 92 electroscan of antenna 93 electroscan of dorsal view of thorax 94 forewing 95 genitalia.
Megastigmus lanneae Roques & Copeland, sp. n. female. 96 dorsal view of the body 97 lateral view of body 98 dorsal view of thorax 99 front view of head 100 antenna 101 forewing.
Megastigmus lanneae Roques & Copeland, sp. n. male pale form. 102 dorsal view of the body 103 lateral view of body 104 dorsal view of thorax 105 front view of head; 106 antenna 107 genitalia 108 forewing.
Megastigmus lanneae Roques & Copeland, sp. n. dark form. 109 dorsal view of the body 110 lateral view of body 111 dorsal view of thorax 112 front view of head; 113 antenna 114 forewing 115 genitalia.
Megastigmus hypogeus Hussey female. 116 dorsal view of the body 117 lateral view of body 118 dorsal view of thorax 119 front view of head 120 antenna 121 forewing.
Megastigmus hypogeus Hussey male pale form. 122 dorsal view of the body 123 lateral view of body 124 dorsal view of thorax 125 front view of head; 126 antenna 127 genitalia 128 forewing.
Megastigmus hypogeus Hussey male dark form. 129 dorsal view of the body 130 lateral view of body 131 dorsal view of thorax 132 front view of head; 133 antenna 134 genitalia 135 forewing.
Megastigmus pistaciae Walker female. 136 dorsal view of the body 137 lateral view of body 138 dorsal view of thorax 139 front view of head 140 antenna 141 forewing.
Megastigmus pistaciae Walker male. 142 dorsal view of the body (pale form) 143 lateral view of body (pale form) 144 dorsal view of thorax(pale form) 145 lateral view of body (dark form) 146 front view of head (pale form); 147 antenna (pale form) 148 genitalia (pale form) 149 forewing (pale form).
Megastigmus transvaalensis Hussey female. 150 dorsal view of the body 151 lateral view of body 152 dorsal view of thorax 153 front view of head 154 antenna 155 forewing.
Megastigmus transvaalensis Hussey male. 156 dorsal view of the body 157 lateral view of body 158 dorsal view of thorax 159 front view of head; 160 antenna 161 forewing 162 genitalia.
Megastigmus somaliensis Hussey. 163 lateral view of female body 164 electroscan of front view of male head 165 electroscan of male antenna 166 electroscan of dorsal view of male thorax 167 male forewing.
Updated synthesis of biological habits of the world Megastigmus species recorded to date with a detail of the species present in Eastern and Southern Africa. The four bars on the top detail the chalcid feeding regimes. The following ones present the host plant families colonized by the phytophagous seed chalcids, splitted into angiosperms and conifers.
Holotype ♀, Kenya, Scandent climber, Rift Valley Prov. Matthews Range, 1.1777°N, 37.3141°E, 1342m, 16 Jan 2004, ex. Helinus integrifolius fruits, A&M coll. N°2692, R. Copeland leg. (NMKE).
Paratypes. 2♀♀, same collection data as holotype (ARC, 1 destroyed); 2♂♂, same collection data as holotype (NMKE, ICIPE).
Holotype ♀. A small species with body length 3.2 mm (without ovipositor); length of ovipositor sheaths 0.8 mm. Body colour nearly entirely orange-yellow (Figures
Head transverse (Figure
Male. Body colour similar to that of female (Figures
Head rectangular (Figure
Variation. The transverse brown band at the anterior part of mid-lobe of mesoscutum is less marked in female paratypes. In males, the lateral black-brown spots could be absent on T5–T7.
Helinus integrifolius (Rhamnaceae). Probably a seed feeder, based on its place in the molecular phylogeny of Megastigmus spp. (Figure
Known only from the Matthews Range, Kenya where it was observed to emerge from 9.8 % of the collected fruits (Table
Named after its host plant.
This is the only species of Megastigmus found to date to emerge from fruits of the Rhamnaceae. Females are easily distinguished from those of other species present in the region by the very small ovipositor associated with an orange body colour. Genitalic characters differentiate males M. helinae by the presence of only two teeth on the digitus whereas the other species bear three teeth except M. icipeensis but the phallobase of the latter species is less elongate (Figure
Holotype ♀, Kenya, Coast Province, Sagalla Mountain, 3.47864°S, 38.57463°E, 1090m, 22 Sep 2014, ex. Lannea cf. schimperi fruits, CHIESA coll. N°343, R. Copeland leg. (NMKE).
Paratypes 2♀♀, same collection data as holotype (1♀ ICIPE; 1♀ARC).
Holotype ♀. Medium-sized species, body length (without ovipositor) 4.6 mm; length of ovipositor sheaths 0.8 mm. Body colour orange-yellow and brown (Figures
Forewing stigma brown without infuscation; basal cell indistinct, lacking setae, no setae on basal setal line and costal setal line (Figure
Head transverse (Figure
Variation: Body length from 4.5–4.9 mm, ovipositor sheaths from 0.8–0.9 mm. No variation was observed in colour or pilosity.
No males reared.
Lannea cf. schimperi (Anacardiaceae). Like other species also reared from Anacardiaceae fruits, e.g., M. pistaciae and M. transvaalensis, it is probably a seed chalcid.
Known only from Sagalla Mountain, Taita Hills, Kenya. Specimens emerged from only 0.4% of sampled fruits (Table
Named in honour of Robert J. Smith, noted musicologist and exceptional left fielder.
Its comparatively very short ovipositor, nearly 2 × smaller than gaster length, allows to separate easily this species from the other ones developing in Anacardiacae fruits and showing resembling body colour patterns but a comparatively much longer ovipositor, quite as long as gaster length (M. ozoroae) or longer (M. lanneae; M. hypogeus).
Holotype ♀: Kenya, Coast Province, Taita hills, Mwatate area, 3.48940°S, 38.33298°E, 1050m, 4 Feb 2012, ex. Grewia tephrodermis seeds, CHIESA coll. N°98, R. Copeland leg. (NMKE)
Paratypes: 2 ♀♀, same collection data as holotype (1 ♀ RSC; 1 ♀ ARC)
Holotype ♀. Small species with body length (without ovipositor) 2.6 mm; length of ovipositor sheaths 1.1 mm. Body colour entirely orange to dark orange except a few black patterns including contours of each ocellus, a spot at wing insertions, contour of propodeal spiracle, rounded spots on the lateral parts of T3, a lunule on the middle of T4, and 2 elongated blackspots on the lateral sides of T5 (Figures
Head rounded, width: height ratio: 1.3×; POL: OOL 1.0; inter-antennal area as broad as torulus width; scrobe short, ca. 2.6 × as long as wide. Scape 1.2 × as long as combined length of pedicel, anellus, and F1 (Figure
Variation. Females range in length from 2.6–2.7 mm. In one paratype, gaster is dark brown. Wing chaetotaxy is variable with basal setal line with 5–6 long setae and costal setal line with 3–6 setae.
No males reared.
Grewia tephrodermis (Malvaceae).
Known only from the Mwatate area, Taita Hills, Kenya. Megastigmus copelandi emerged from 0.7% of collected fruits (Table
Named after Dr. Robert Copeland, who collected the specimen.
The combination of an almost entirely orange body, pale pilosity on the head, and an ovipositor a bit shorter than gaster length readily distinguishes this species from other Afrotropical Megastigmus, and from M. grewianae, in particular, which also attacks Grewia fruits. The latter species is much larger (4.1 mm vs. 2.6–2.7) and has an ovipositor ca. 1.8 × longer than gaster length (Figure
Holotype. ♀, Kenya, Nyanza Province, Sindo-Mbita Road, 0.4849°S, 34.1765°E, 1205m, 11 Dec 2004, A&M coll. N°3064, ex. Ozoroa insignis ssp reticulata, R. Copeland leg. (NMKE)
Paratypes. 8 ♀♀, same collection data as holotype; 6♂♂, same collection data as holotype, except 19 Dec 2004, A&M coll. N°3077 (2♀♀, 1♂, NMKE; 2♀♀, 1♂ RSC; 2♀♀, 1♂ ARC, 1♀, 1♂ ICIPE, 1♀, 1♂ SAMC)
23♀♀, 11♂♂ same collection data as holotype, except 19 Dec 2004, A&M coll. N°3077 (11♀♀, 5♂♂ RSC; 12♀♀, 6♂♂ ARC)
Holotype ♀. Large species, body length (without ovipositor) 4.8 mm; length of ovipositor sheaths 1.5 mm. Body colour yellow and black (Figures
Legs pale yellow except claws brownish; coxae with numerous pale hairs extending from small black dots, especially on hind coxa; femora with small black hairs; tibiae with 3 rows of long black hairs. Forewing stigma brown without infuscation; basal cell only partly closed, the disc with 5 very small setae; basal line with 2 setae; no setae on costal line; 4 setae in coastal cell (Figure
Head about 1.1 × as wide as long in front view (Figure
Variation. Females range in length from 3.5 to 4.8 mm. Five of the examined specimens out of 14 have lateral lobes of mesoscutum and scutellum pale yellow. The black spots on pronotum and mid-lobe of mesoscutum as well as the other black markings are absent in one specimen, except at base of eye, on the suture of the lateral lobe of mesoscutum and on the central part of propodeum. The general colour of this specimen is orange, with the gaster brownish and not blackish in its middle part, the brownish colour beginning only at T5. Another specimen has the central part of propodeum completely shining black. The relative length of the ovipositor sheaths varies between 0.7 to 0.8 × the gaster length, and 0.3–0.4 × the body length.
Males. They are highly variable in colour, and two extreme forms can be distinguished with intermediates.
Pale form. Body length from 4.3 to 5.7 mm. Body colour pale yellow and black (Figures
Legs yellow expect anterior part of coxae with large black spots (extending to the posterior part in fore coxa), and small brownish spots at femur insertion, 3 rows of large hairs on tibia. Forewing stigma brown without infuscation (Figure
Head about 1.2 × as wide as long in front view. POL: OOL: 2.0; inter-antennal area as broad as torulus width; scrobe short, ca. 1.5 × as long as wide. Scape 1.3 × as long as combined length of pedicel, anellus, and F1 (Figure
Dark form. Larger than pale form, with body length 5.0 to 6.5 mm. Body colour mostly black (Figures
Intermediates. In some specimens, black patterns on thorax are limited to a sharp median line on pronotum and mid-lobe of mesoscutum, and to the sutures of lateral panel of pronotum, prepectus, and tegula. These specimens also show a grey spot on lateral lobes of mesoscutum and a brownish median line on scutellum. Gaster is mostly black with a few yellow patterns as follows: an annelation at the base of T3, the sides of T4, a small lateral spot on T5 and the last two segments. Forewing stigma not infuscated in these individuals. In some others, the lateral parts of thorax are black (panel of pronotum, prepectus, mesepisternum, and upper mesepimeron) except lower mesepimeron and metapleuron and callus yellow (with black spots on callus), and scutellum is black at the base with the remainder dark brown. In these individuals, the forewing stigma is infuscated. Pilosity may include 5–6 pairs of setae on mid-lobe of mesoscutum and 3–4 pairs on scutellum.
Ozoroa insignis (Anacardiaceae). Its position in the molecular phylogeny is close to those Megastigmus species known to be seed feeders in Anacardiaceae, strongly suggesting it is also a seed feeder. Although the presence of numts was noticed in the analyzed specimen of M. ozoroae it belonged whatever the marker (short mitochondrial or nuclear fragment) to the “Anacardiaceae clade” confirming the existence of a strong monophyletic clade, the most divergent from the others (Figure
Known only from western Kenya, near Lake Victoria. In one collection, 8.1% of fruits were infested (Table
Named after the genus of the host plant.
Females are easily distinguished from those of other species observed to emerge from fruits of Anacardiaceae by the relatively small ovipositor, nearly as long as gaster length whereas it is at least 1.2 × the gaster length in M. hypogeus (Figure
Holotype ♀, Kenya, Nairobi Province, ICIPE Campus, Kasarani, 1.2232°S,36.8965°E, 1600m elevation, Malaise trap, meadow in degraded shrub-/grassland, 24 Feb to 3 Mar 2014, R. Copeland leg. (NMKE)
Paratypes 4♀♀, the same as holotype (1♀ NMKE; 1♀ RSC; 1♀ ARC; 1♀ ICIPE).
3♂♂, collected together with the four females (2♂♂ RSC; 1♂ ARC).
Holotype ♀. Very small species with body length (without ovipositor) 1.2 mm; length of ovipositor sheaths 0.9mm. Body colour nearly entirely orange-yellow (Figures
Head about 1.2 × as wide as long in front view (Figure
Variation. Body ranges 1.2–1.3 mm with ovipositor up to 1.2 mm. Ovipositor length varies from 0.7 to 0.9 × the body length, and from 1.6 to 2.0 × the gaster length. No variation in colour and pilosity.
Male. In the absence of molecular analyses, the males caught in Malaise traps along with the four female M. icipeensis could not be attributed with certainty to this species but noticeable convergences in morphology make it probable. Body length 1mm. Same body colour as in female of M. icipeensis (Figures
Legs yellow with the two last tarsal segments and claws black. Pilosity pale on head and thorax, showing the same setal pattern as in female M. icipeensis, including the interocellar brisles and 2 pairs of pale hairs on scutellum (Figure
Head transverse, about 1.3 × as wide as long in front view (Figure
Unknown. In the molecular study, M. icipeensis did not group with other taxa, but the phylogenetic position (Figure
Only known from the collection site in a degraded meadow near a small stream.
Named after the institution on whose grounds the species was collected, the campus of the International Centre of Insect Physiology and Ecology (ICIPE), Nairobi.
The species can be easily distinguished from other tiny Megastigmus spp. known from the Afrotropical region by the pale bristles on the thoracic dorsum whereas they are mostly black in M. zebrinus (Grissell, 2006), and by its orange body whereas M. pretorianensis is mostly black with brown areas (
Holotype ♀, Kenya, Coast Province, Base of Kasigau Mountain, 3.81301°S, 38.64050°E, 619m, 5 Mar 2012, ex. Lannea schweinfurthii fruits, CHIESA coll. N°141, R. Copeland leg. (NMKE);
Paratype 1♀ Kenya, Coast Province, Funzi island, 4.57776°S, 39.44127°E, near sea level, Malaise trap in mixed grass and woodland, 24 to 28 Jul 2012, R. Copeland leg. (ICIPE).
Holotype ♀. Medium-sized species with body length (without ovipositor) 4.0 mm; length of ovipositor sheaths 3.4 mm. Body colour predominantly orange (Figures
Head rounded, width: height ratio: 1.0 (Figure
Variation. In the paratype, the basal cell presents two long setae on disc, and the basal setal line has 1 seta.
No males reared.
Lannea schweinfurthii (Anacardiaceae). Probably a seed feeder but it could not be integrated in the molecular analysis because too few specimens were available.
Known from the base of Kasigau Mountain, a Taita Hills outlier. Only 1 of 168 fruits (0.6%) was infested (Table
Named in honour of Leo Laventhal, historian, Yiddish scholar and union man.
Females can be separated easily from those of other species associated with fruits of Anacardiaceae by the combination of a predominantly orange body and a relatively long ovipositor, at least 1.8 × as long as the gaster. In most of the other species, body colour combines dark and yellow patterns (M. hypogeus, M. ozoroae, M. lanneae, M. smithi). For the remaining species that have a predominantly orange body colour, the ovipositor length is at most 1.4 × gaster length (M. pistaciae, M. thomseni and M. transvaalensis).
Holotype ♀, Kenya, Coast Province, Arabuko-Sokoke forest, 3.2997°S, 39.9869°E, 55 m, 17 Feb 2000, ex. Grewia stuhlmanii fruits, Coll. N° Kip-356, R Copeland leg. (NMKE)
Paratypes: Kenya, 1♀, 1♂, same collection data as holotype (RSC); 2♀♀, 1♂, same collection data as holotype (ARC); 1♀, 7♂♂, Eastern Province, Tsavo, 2.6760°S, 38.3325°E, 638 m, 20 Feb 2000, ex. fruits of Grewia tephrodermis, A&M Coll. N°514, R. Copeland leg. (1♀, 3♂♂ NMKE, 4♂♂ ICIPE); 1♀, Coast Province, Kasigau Mountain, 3.82700°S, 38.64875°E, 1065 m, Malaise trap in campsite clearance, 19 May to 2 Jun 2011, R. Copeland leg. (SAMC); 1♀, Eastern Province, Kasaala area, 2.07486°S, 38.22530°E, 741 m, Malaise trap, 28 Nov. to 4 Dec 2013, leg. J. Bukhebi & R. Copeland (ICIPE).
Kenya, 7♀♀, 1♂, same as holotype (ARC); 1♀, Eastern Province, Tsavo, 2.6760°S; 38.3325°E, 638 m, 20 Feb 2000, ex. fruits of Grewia tephrodermis, Coll. #514, R. Copeland leg. (ARC)
Holotype ♀. Body length (without ovipositor) 4.1mm; length of ovipositor sheaths 3.0 mm. Body colour orange with some darker patterns (Figures
Legs entirely pale yellow except claws brown, coxae with conspicuous hair dots. Forewing stigma brown without infuscation; basal cell closed, with 13 setae on disc ; basal setal line with 6 long setae; costal setal line with 16 small setae, costal cell with 11 setae in 2 rows (Figure
Head rounded, width: height ratio: 1.1 × (Figure
Male. Body length 4.2 mm; Body colour, pilosity and black patterns similar to female (Figures
Head subquadrate, width: height ratio: 1.1 × (Figure
Variation. Females range in length from 3.5 to 4.0mm. The number of lateral spots on gaster varies from 6 (with two on T3) to 2 (only the large ones remaining), the 3 last lateral spots sometimes fused into a line. Pilosity on thorax paler in one female paratype. Males range in length from 4.1 to 4.5 mm. The spots on gaster sometimes larger, the pairs of lateral hairs on scutellum ranging up to 6 in both sexes.
Grewia stuhlmannii, Grewia tephrodermis (Malvaceae). Probably a seed feeder, based on its place in the molecular phylogeny of Megastigmus spp. (Figure
Known from dry Acacia/Commiphora savanna, moderately wet mid-altitude mountain forest, and bush associated with costal forest. Adults emerged from 2.7% of G. stuhlmannii fruits and 13% of G. tephrodermis fruits (Table
Named after the genus of its host plant.
Females are easily separated from those of M. copelandi, the other species reared from Grewia fruits, by their significantly larger size (4.1 mm vs. 2.6–2.7) and the relatively longer ovipositor (ca. 1.8 × longer than gaster length vs. 0.9 × in M. copelandi). Females and males are also easily distinguished from the other species of Afrotropical Megastigmus by the unique pattern of a longitudinal black stripe extending 7/8 of the length of the side of the pronotum (Figures
Holotype ♀: Kenya, Eastern Province, Machakos, 1.5609°S, 37.2338°E, 1586m elevation, 3 May 2005, ex. Lannea rivae fruits, Coll. N°3109, R. Copeland leg. (NMKE)
Paratypes 8♀♀, 12♂♂, same collection data as holotype (2♀♀, 2♂♂ NMKE; 2♀♀, 4♂♂ RSC; 2♀♀, 2♂♂ ARC; 1♀, 2♂♂ ICIPE; 1♀, 2♂♂ SAMC);
Kenya; 7 ♀♀ same collection data as holotype; 2♀♀, same collection data as holotype except 25 Nov. 2004, ex. Lannea rivae seeds, Coll. N°3042, R. Copeland leg. (ARC).
Holotype ♀. Large species, body length (without ovipositor) 5.4 mm; length of ovipositor sheaths 4.2 mm. Body colour yellow and brownish (Figures
Face quadrate, width: height ratio: 1.0 (Figure
Variation. Length of female body varies from 4.8 to 5.4 mm. The relative length of the ovipositor sheaths varies between 1.6 and 1.8 × the gaster length, and between 0.6 and 0.7 × the body length. In three out of the eleven specimens examined, the brownish bands on thorax and abdomen are closer to to dark orange, becoming less distinct. One specimen, with two setae on the basal setal line, a second specimen with four setae.
Males. Like in most other Megastigmus species related to Anacardiaceae, males are highly variable in colour, and two extreme forms can be distinguished with intermediates.
Pale form. Body length from 4.2 to 6.2 mm. Body colour mostly orange-yellow (Figures
Head about 1.1 × as wide as long in front view (Figure
Dark form. Larger than pale form, with body length 5.7 to 6.4 mm. Body colour substantially black (Figures
Intermediates. Some specimens show a slightly infuscated stigma along with head and thorax mostly orange but with more blackish patterns than in the pale form: a black-brown band between eye margin and torulus, a brownish median band on thorax from anterior suture of pronotum to posterior suture of mid lobe of mesoscutum which prolongates in a fuzzy median band on scutellum, and a brown longitudinal band on upper and lower part of mesepimeron. Legs yellow with a large black spot on fore coxa, a smaller spot on mid-coxa, and a brownish infuscation on hind coxa. Propodeum with a large median black band, black spots around spiracles, and a brownish infuscation at suture with callus. Pilosity is similar to the one of pale from but some individuals have 3 pairs of hairs on scutellum.
Lannea rivae (Anacardiaceae). Probably a seed feeder. Although the presence of numts was noted in the COI sequences of the only specimen of M. lanneae which amplified for DNA analysis, this species clearly clustered within the “Anacardiaceae clade” (Figure
Known only from farmland tree in Machakos area. Adults emerged from 3.3% of the collected fruits (Table
Named after the genus of its host plant.
Females can be separated easily from those of other species associated with fruits of Anacardiaceae by the combination of an ovipositor at least 1.8 × as long as gaster length and a yellow and brownish body. In other species having dark and yellow patterns, the ovipositor length is either shorter than gaster length (M. ozoroae- Figure
Holotype ♀ “Bred ex. oil seed, Nairobi; 5–37; Kenya, Corydon Museum; Pres. by Com. Inst. Ent., B.M. 1957–41; B.M. TYPE HYM. 5.1653; NHMUK010263947” (BNHM); Paratypes: 3♀♀, 4♂♂, same collection data as holotype (BNHM);
11♀♀, 14♂♂, Kenya, Coast Province, Shimba Hills, 4°08.096'S, 39°28.082'E, 285m elevation, 25 Nov. 2001, ex. Ozoroa obovata seeds A&M coll. N°1609, R. Copeland leg.; 1♀, Nyanza Province, Sindo-Mbita Road, 0°29.091'S, 34°10.592'E alt 1205m, 11 Dec 2004, ex. Ozoroa insignis ssp reticulata seeds, A&M coll. N°3064, R. Copeland leg.; 11♀♀, 8♂♂, Arabuko-Sokoke forest, 3.3716°S, 39.8949°E, 55m elevation, 18 Feb 2000., coll. N°Kip-372, ex. Ozoroa obovata seeds, R. Copeland leg; 2♀♀, 3♂♂, Arabuko-Sokoke forest, 3.3716°S, 39.8949°E, 55m elevation, 18 Feb 2000, ex. Ozoroa obovata seeds, A&M coll. N°509, R. Copeland leg.; 1♀, Arabuko-Sokoke forest, 3.2030°S, 39.9271°E, 98m elevation, 17 Mar 2001, ex. Ozoroa obovata seeds, A&M coll. N°1103, R. Copeland leg. (all material in RSC except 5♀♀, 5♂♂ from Arabuko-Sokoke forest in ARC)
This species has been first described by
Female. Females range in length from 3.9 to 5.1 mm (without ovipositor), length of ovipositor sheaths from 2.1 to 2.5 mm. Body colour usually yellow and black (Figures
Thorax yellowish-brown with blackish patterns (Figures
Face subquadrate, width: height ratio in front view: 1.2×; POL: OOL 1.6×; inter-antennal area as broad as torulus width; scrobe rather short, ca. 2.9 × as long as wide. Scape 1.3 × as long as combined length of pedicel, anellus, and F1 (Figure
Males. They are highly variable in colour, and two extreme forms can be distinguished with intermediates. Paratypes ♂ include three specimens with pale patterns and one with dark patterns whereas our collection included nine specimens of pale form, 13 of dark form, and three of intermediate color.
Pale form. Body length from 4.3 to 5.4 mm. Body colour mostly orange-yellow (Figures
Thorax mostly yellow- orange with a few black patterns (Figure
Face rounded, width: height ratio in front view: 1.0× (Figure
Dark form. Body length slightly longer than in pale form, from 5.0 to 5.5 mm. Body substantially black (Figures
Relative dimensions of head and antenna similar as in pale form; sculpture of thorax and propodeum same as in pale form. Forewing stigma nearly rectangular, 1.3 × as long as wide (Figure
Body length as in pale form, from 4.6 to 5.4 mm. Some specimens with infuscate stigma show lighter coloration patterns than in the extreme dark form, with the median band on pronotum smaller, the lateral lobes of mesoscutum and axilla yellow, the scutellum with the median band limited to the anterior part, and the gaster with narrow yellow bands from T4 to T6.
Ozoroa obovata, Ozoroa insignis ssp reticulata (Anacardiaceae). Probably a seed feeder, based on its place in the molecular phylogeny of Megastigmus spp. where it clusters with the other species developing in seeds of Anacardiaceae (Figure
Mixed bush-/grassland in western Kenya near Lake Victoria and in similar habitat in coastal areas. Adults emerged from up to 2.8% of the fruits in some collections (Table
The key from
Genitalic characters also allow separation of M. hypogeus males from those of other species developing in seeds of Anacardiaceae, the aedeagus part above digitus being significantly less elongated (1.2 × as long as digitus length) than in M. ozoroae (1.8×; Figures
3♀♀, 3♂♂, Kenya, Mount Kulal, 2.6290°S, 36.9278°E, 1640m elevation, 11 Dec 2008, ex. Pistacia lentiscus subsp. emarginata seeds, R. Copeland leg. (2♀♀, 1♂ RSC; 1♀, 1♂ ARC; 1♂ destroyed)
Male and female specimens fit the detailed description of the species by
In the molecular phylogeny of Megastigmus, the Kenyan specimen clusters with the other specimen of M. pistaciae from Southern Europe (Figure
Pistacia lentiscus subsp. emarginata seeds.
Reared from samples collected in highland areas (1640–1920 m above sea level) of the Rift Valley, in northern and southwestern Kenya. Few fruits (0.6%) were infested (Table
Females can be separated from those of other species with a predominantly orange colour associated with fruits of Anacardiaceae by the shape of the forewing stigma and the relative length of ovipositor. The stigma is oval in M. pistaciae (1.2–1.3× as long as wide; Figure
The genitalia allow one to separate males from those of other species identified in seeds of Anacardiaceae. The aedeagus part above digitus is comparatively longer than in M. transvaalensis (0.8 × vs. 0.5 × as long as digitus length; Figures
3♀♀, 7♂♂, Kenya, Kikuyu Escarpment, Central Province, 1.0290°S, 36.6025°E, 2100 m, coll. 85, ex. Rhus vulgaris seeds, 29 Apr 1999, R. Copeland leg.; 2♀♀, 3♂♂, Kenya, Burguret, Central Province, 0.1128°S, 37.0375°E, coll. 2162, ex. Rhus natalensis seeds, 16 Aug 2002, R. Copeland leg.; 3♀♀, 7♂♂, Kenya, Nairobi Province, 1.2212°S, 36.8963°E, 1610m, coll. 2787, ex. Schinus terebinthifolius seeds, 28 Apr 2004, R. Copeland leg. (4♀♀, 10♂♂ RSC; 4♀♀, 7♂♂ ARC)
Male and female specimens fit the description of the species by
Schinus molle, Schinus terebinthifolius, Rhus natalensis, R. vulgaris (Anacardiaceae). A seed feeder.
Adults emerged from 14.8% of the fruits of S. molle, 14.7% of those of R. vulgaris and up to 6.1% of R. natalensis fruits (Table
Females can be distinguished from these of other species associated with fruits of Anacardiaceae and showing a predominantly orange body by the relative size of the ovipositor and the shape of the forewing stigma. In M. transvaalensis, the ovipositor is 1.4 × as long as gaster length (Figure
Genitalia allows the separation of males from those of other species reared from seeds of Anacardiaceae. The aedeagus part above digitus is conspicuously shorter than digitus length (0. 5×; Figure
Holotype ♀ “Bristish Somaliland; ex. seeds of Juniperus procera; coll. i.1954, em. 25.ii.1954, J.T. Lawrie; Brit. Mus. 1956–294; Com. Inst. Ent. coll. 13661; B.M. TYPE HYM. 5.1623a; NHMUK010263946 5.1623a”(BNHM). Paratypes 4♀♀, 2♂♂, same collection data as holotype (BNHM);
1♀ “Abyssinia, Mulu, above Muger Valley; circa 8000 ft., 18–23. 12. 1926, Dr. H. Scott” (BNHM); 1♀ “Abyssinia, Mt. Chillálo Digula; circa 9500 ft., 27.11.1926, from foliage giant juniper, Dr. H. Scott” (BNHM); 3♀♀, 2♂♂, Kenya, Central Prov., Mt. Kenya Forest, 0.0308°S, 37.1230°E, 2125m, coll. 3034, ex. Juniperus procera seeds, 2 Nov. 2004, R. Copeland leg. (2♀♀, 1♂, RSC; 1♀ ARC; 1♂ destroyed).
Male and female specimens obtained from Kenya fit the general description of the species by
Juniperus procera. A seed feeder clustering in the molecular phylogeny of Megastigmus with the other species developing in seeds of Juniperus in Europe (Figure
Sampled in highland forest and woodland in central and northwestern Kenya (Figure
This species, as well as the related M. asir which also attacks Juniperus procera (
Is the radiation on Angiosperms more important than previously considered?
Our results increase to 16 the number of Megastigmus species presently known from the Afrotropical region, of which at least 13 are seed feeders. The results also increase to 28 the number of species shown to be associated worldwide with angiosperm seeds vs. 48 with conifers, and to 7 the number of angiosperm families hosting Megastigmus seed chalcids (Figure
This work was partly supported by grants awarded to R. Wharton, Texas A&M University, USA, from USAID (no. PCE-G-00-98-0048-00) and USDA/CSREES/IFAFS (no. 00-52103-9651) in collaboration with the International Centre for Insect Physiology and Ecology (ICIPE) and its African Fruit Fly Initiative. The authors are grateful to the government of Finland for partial funding of our research through a grant to ICIPE (the CHIESA Project; Climate Change Impacts on Ecosystem Services and Food Security in Eastern Africa). Many thanks to the Kenya Wildlife Service (KWS), and in particular to Dr Richard Bagine, the KWS Head of Research, for permission to sample in Kenyan national parks and reserves. Thanks also to the director, Kenya Forest Service for permission to sample in the forests of Taita Hills. Quentin Luke helped with plant identifications. Juliet Muruiki, Millicent Okumu and Peris Machera provided excellent technical support in the field and in the laboratory. Mwadime Mjomba and Eric Mwakajana helped with fruit collection in the Taita Hills. Jackson Kimani of ICIPE’s Earth Observation Unit made the excellent maps. Claudine Courtin, INRA, gave valuable help in sequencing the insects. We thank very much Natalie Dale-Skey Papilloud, Curator of Chalcidoidea at the Natural History Museum in London for providing access to the type material from Hussey and to the Museum collections of Megastigmus from Africa. We also thank Simon Van Noort (IZIKO) for supplying data about the presence of rose seed chalcids in South Africa, and John Noyes and Fernando Farache for critical reviews.