Phylogenetics, integrative taxonomy and systematics of the Sesamia cretica species group (Lepidoptera: Noctuidae: Apameini: Sesamiina), with the description of 21 new species from the Afrotropical region

Summary In this study, 31 species of noctuid stemborers belonging to the genus Sesamia Guenée, 1852 (Lepidoptera: Noctuidae: Noctuinae: Apameini: Sesamiina) are reviewed. All these species are assigned to the Sesamia cretica group sensu Tams & Bowden (1953). Based on genitalic characters, several subgroups are hereby defined. Nine species belong to a species complex defined as the Sesamia albivena Hampson, 1902 subgroup; it consists of S. albivena, S. mocoensis Tams & Bowden, 1953, n. stat., S. sudanensis Tams & Bowden, 1953, n. stat. S. taenioleuca (Wallengren, 1863), and five new species that are described (S. aethiopica Le Ru n. sp. from Ethiopia, S. kafulo Le Ru n. sp. from Botswana and Zambia, S. kavirondo Le Ru n. sp. from Kenya and Uganda, S. maloukou Le Ru n. sp. from Republic of Congo, and S. soyema Le Ru n. sp. from Ethiopia). Four species belong to a species complex defined as the Sesamia cretica subgroup; this encompasses S. cretica, S. rufescens Hampson, 1910, and two new species that are described (S. ihambane Le Ru n. sp. from Mozambique and Tanzania and S. kikuyuensis Le Ru n. sp. from Kenya); two new synonyms are introduced for Sesamia cretica: Nonagria uniformis Dudgeon, 1905 n. syn. and Sesamia griselda Warren, 1913, n. syn. Ten species belong to a species complex defined as the Sesamia fuscifrontia Hampson, 1914 subgroup; this includes S. fuscifrontia, S. geyri (Strand, 1915) and eight new species that are described (S. babati Le Ru n. sp. from Tanzania, S. babessi Le Ru n. sp. from Cameroon and Zambia, S. mabira Le Ru n. sp. from Uganda, S. nangaensis Le Ru n. sp. from Cameroon and Republic of Congo, S. rungwa Le Ru n. sp. from Tanzania, S. simillima Le Ru n. sp. from Benin, Cameroon, Kenya and Uganda, S. taveta Le Ru n. sp. from Kenya and S. ulaukae Le Ru n. sp. from Ethiopia). One species belongs to a species complex defined as the Sesamia salama Le Ru n. sp. subgroup; this consists of S. salama Le Ru n. sp. from Kenya and another undescribed Sesamia species from South Africa. One species belongs to a species complex defined as the Sesamia viettei Rungs, 1954 subgroup. Six species belong to a species complex defined as the Sesamia wiltshirei Rungs, 1963 subgroup; this groups S. wiltshirei and five new species that are described (S. djenoensis Le Ru n. sp. from Republic of Congo, S. inexpectata Le Ru n. sp. from South Africa and Zambia, S. lefini Le Ru n. sp. from Republic of Congo, S. echinochloa Le Ru n. sp. from Botswana, Kenya, Mozambique, South Africa, Tanzania and Zambia and S. rindini Le Ru n. sp. from Tanzania). A supplemental description of the previously described species is also provided. Novel host plant records are also provided for 11 species of the S. cretica group. To complement the morphological study, both phylogenetic and molecular species delimitation analyses were carried out on a multimarker (four mitochondrial and two nuclear genes) molecular dataset encompassing 144 specimens representing 35 species (including 25 species from the S. cretica group). Molecular analyses provide a well-supported phylogenetic framework for the species of interest, which are all recovered monophyletic. Molecular species delimitation analyses also support the species status of almost all sampled species. Interestingly, the inferred tree indicates that the S. cretica group and the S. fuscifrontia subgroup are both paraphyletic; this indicates that, while highly informative, the chosen genitalic characters in Sesamia are not all synapomorphies.

The taxonomy and systematics of speciose insect groups is generally complex, especially within groups lacking useful external diagnostic characters. In such groups, genitalia (especially male genitalia) often constitute a major source of information to morphologically discriminate, classify and identify species (e.g. Tuxen 1970). However, genitalic characters can be sometime misleading because they often evolve rapidly due to intense sexual selection (Song & Bucheli 2010;Simmons 2014). The latter could lead to high levels of homoplasy, even when contrasting taxa are distantly unrelated. In this context, it is worth making use of molecular phylogenies to better understand the evolution of genitalia and investigate their taxonomic utility (e.g. Kergoat & Alvarez 2008). Regarding species boundaries, integrative taxonomy approaches can also clarify the status of closely related species, even when differences in genitalia are subtle. However, caution must be exercised, because the availability of a plethora of molecular species delimitation (SD) approaches (see e.g. Carstens et al. 2013 for a review) and differences in data treatment can lead to incongruent results (e.g. Astrin et al. 2012;Renner et al. 2017), which are often difficult to disentangle and interpret. To clarify the results of SD analyses, several authors (Miralles & Vences 2013;Ahrens et al. 2016) proposed new metrics that allow the comparison of species partitions generated by all kind of SD approaches. A tool to simplify this procedure was also recently released (Ducasse et al. 2020), which greatly facilitates the estimation of the corresponding indexes.
In this study both morphological and molecular information are used to revise a species group belonging to the noctuid stemborer genus Sesamia Guenée, 1852 (Lepidoptera: Noctuidae: Noctuinae: Apameini: Sesamiina). In its current definition the genus Sesamia consists of about 50 species that are mostly distributed in the Afrotropical region. Based on genitalic characters, the genus was first divided by Tams & Bowden (1953) into the S. cretica and S. nonagrioides (Lefebvre 1827) groups; however, they also underlined that several species (noticeably S. coniota Hampson, 1802 andS. jansei Tams &Bowden, 1953) could not be assigned to any of the two groups because they presented an intermediate morphology. The latter species were recently assigned to a novel species groupthe S. coniota groupby Le Ru et al. (2020c). The genus Sesamia is the second most diversified genus among the Sesamiina with at least 25% of the known species (Moyal 2006;Moyal et al. 2011;Kergoat et al. 2015;Le Ru et al. 2020c). Yet, recent investigations into this genus indicate that its diversity is likely greatly underestimated. About a decade ago, Moyal et al. (2011) described three new species related to S. epunctifera Hampson, 1902. Further, Kergoat et al. (2015 reassessed the S. nonagrioides species complex and described six new species. In parallel, molecular phylogenetics studies by Toussaint et al. (2012) and Kergoat et al. (2018) suggested a substantial number of other new Sesamia species, which led us to progressively undertake a global revision of the genus. However, given the large number of Sesamia species it was decided to tackle previously defined species groups one by one. As such, a first study by Le Ru et al. (2020c) only focused on the S. coniota species group, for which three new species were described.
In the present study, a focus is set on the S. cretica group sensu Tams & Bowden (1953), which was defined based on the following genitalic characters: in males (i) valve with sacculus and cucullus fused, (ii) juxta with a medial projection, and (iii) strongly spinose manica; in females: (i) more pointed bursa, (ii) broad ostium, (iii) frequently heavily sclerotized anteostial pad, and (iv) weakly sclerotized ostial segment. To revise this group, a morphological study was carried out on specimens (including type material) from all known species of the S. cretica group as well on other Sesamia specimens that could correspond to new species.

Material and methods
To summarize, the morphological study of the S. cretica group is based on a total of 728 adult specimens belonging to 31 species collected in 149 localities from the following 31 countries: Algeria, Angola, Benin, Botswana, Cameroon, Croatia, Democratic Republic of Congo, Egypt, Eritrea, Ethiopia, France, Greece, India, Iraq, Kenya, Libya, Madagascar, Malawi,   Mozambique, Namibia, Republic of Congo, South Africa, Somalia, South Sudan, Sri Lanka, Tanzania, Thailand, Turkey, Uganda, Zambia and Zimbabwe. Genitalia of newly collected specimens were dissected after immersion of the end of the abdomen in a boiling 10% potash bath for a few minutes, then cleaned, immersed in absolute alcohol for a few minutes and mounted on slides in Euparal (after separating the aedeagus from the rest of the genitalia in males). Collected insects were identified by comparison with holotypes, paratypes and specimens housed in MCSN, MNHN, MRAC, NHM, NRM, TMSA and ZMB. Types for 10 of the new species were deposited in MNHN (N.B. types of the 13 other new species were already housed in Museum collections) whereas paratypes were deposited in MNHN and NMK.

DNA extraction and sequencing
For the molecular dataset 143 specimens from 34 species were selected. This sampling consists of 142 Sesamia specimens,  Ru, 2020) and six species from the S. nonagrioides group (S. calamistis Hampson, 1910, S. epunctifera Hampson, 1902, S. natalensis Le Ru, 2015, S. nonagrioides, S. oriaula, Tams & Bowden, 1953and S. pennipuncta Moyal, 2011. Based on the results of Kergoat et al. (2018), a representative of the closely related genus Buakea Moyal et al., 2011(Buakea kaeuae Moyal et al., 2011 was used to root all phylogenetic trees. Except for the specimen of S. viettei, all specimens were collected by our research group (mostly by B. Le Ru). DNA was extracted from hind legs using Qiagen DNAeasy tissue kits (Qiagen, Hilden, Germany). Polymerase chain reaction (PCR) amplifications were conducted for four mitochondrial gene fragments, a 658 bp region of the cytochrome oxidase I (COI), 992 bp of the cytochrome b (Cytb), 364 bp of the ribosomal 12S RNA (12S), and 534 bp of the ribosomal 16S RNA (16S). Two nuclear gene regions were also sequenced, 1240 bp of the elongation factor-1a (EF1a), and 842 bp of the 28S ribosomal DNA (28S). For both genes, the primers and settings detailed in Kergoat et al. (2012) were used. Resulting PCR products were processed by the French sequencing centre Genoscope using a BigDye v3.1 sequencing kit and Applied 3730xl sequencers. Both strands were sequenced for all specimens to minimize PCR artefacts and ambiguities. Sequences of complementary strands were edited and reconciled using Geneious R9 software (available at: www.geneious.com/).
All PCR amplifications failed for the only specimen of S. viettei processed. As a leg from the male holotype collected in the early 1950s was used, this outcome was expected, so additional molecular work was carried out and the corresponding DNA extract was further processed with high throughput sequencing (HTS). Four mitochondrial gene fragments were targeted for HTS amplicon sequencing. The primer couple SRJ-14233/SRN-14588 of Simon et al. (1994) was used for the 12S and the primer couple ChiarF/ChiarR of Marquina et al. (2019) was used for the 16S. For the COI, two overlapping fragments were targeted; the fragment "FC" (primer couple "Ill_LCO1490" and "Ill_C_R") of Shokralla et al. (2015) and the fragment amplified by the primer couple "FwhF2" (Vamos et al. 2017) and "Foldegen-rev" (Yu et al. 2012). Amplicon libraries were further constructed following Galan et al. (2018). In comparison with the settings of Galan et al. (2018), the following changes were made to reduce the proportion of chimeric fragments: for the first PCR step the number of cycles was set to 40, and for the fragment FC the extension duration was set to 120 s for the second PCR step. The final library was paired-end sequenced on an Illumina MiSeq flowcell using a MiSeq Reagent Kit v2 at the AGAP laboratory (Montpellier, France). Illumina reads were processed using the FROGS pipeline (http://frogs.toulouse.inra.fr/; Escudié et al. 2018) on the Genotoul Galaxy server (Toulouse, France) using demultiplexing, pre-processing, clustering, chimera removal and filtering tools (in that order). Remaining contaminants were further detected using the BLAST tool (available at: https://blast.ncbi.nlm.nih.gov/Blast.cgi) and removed manually. Two genes were successfully sequenced for S. viettei: a 313 bp fragment of the COI (corresponding to the "FwhF2" "Foldegen-rev" fragment) and a 404 bp fragment of the 16S. Finally, GenBank data was used to add the sampling S. grisescens Warren, 1911, which was hypothesized to belong to the S. coniota group (see Le Ru et al. 2020c for more details).
The sequences of the three ribosomal genes (12S, 16S and 28S) were variable in length. Their alignment was accomplished using MAFFT v7 (Katoh & Standley 2013) with default option settings. For all protein-coding genes, Mesquite v3.61 (Maddison & Maddison 2018) was used to check the coding frame for possible errors or stop codons. All corresponding sequences were deposited in GenBank (see Appendix S1 for all accession numbers). The combination of the six gene fragments resulted in a combined matrix of 144 specimens and 4630 aligned characters (with c. 18% of missing data).
Phylogenetic analyses IQ-TREE v2. 1.3 (Minh et al. 2020) was used to carry out phylogenetic analyses under maximum likelihood (ML). The concatenated dataset was divided a priori into 12 partitions, with three partitions (one per codon position) defined for each coding gene fragment (COI, Cytb and EF1a) and one partition defined for each non-coding gene fragment (12S, 16S and 28S). To infer individual gene trees, additional analyses were also conducted on each gene fragment, with three partitions implemented for the coding genes. Finally, ML analyses were conducted on the concatenated nuclear dataset (EF1a, 28S) to explore the phylogenetic signal of the nuclear dataset only. Best-fit substitution models and partition schemes were selected using the Bayesian information criterion implemented in IQ-TREE (see Appendix S2).
Maximum likelihood trees were obtained using heuristic searches implementing 500 random-addition replicates with the following settings: random-starting tree, hill-climbing nearest neighbour interchange (NNI) search (-allnni option), a perturbation strength set to 0.5 (-pers 0.5 option), partition-resampling strategy (-sampling GENE option), best partition scheme allowing the merging of partitions (-m MFP+MERGE option). Clade support for all analyses was assessed using 1000 replicates for both SH-like approximate likelihood ratio tests (SH-aLRT; Guindon et al. 2010) and ultrafast bootstraps (uBV;Minh et al. 2013). Nodes supported by ultrafast bootstrap values SH-aLRT values ≥ 80% and (uBV) ≥ 95% were considered as strongly supported following authors' recommendations.

Molecular species delimitation analyses
Distinct molecular SD approaches and settings were used in this study to better assess the reliability and repeatability of the proposed SDs (see e.g. Astrin et al. 2012;Luo et al. 2018).
First, two tree-based SD methods were carried out: the treebased Poisson-tree-process (PTP) approach of Zhang et al. (2013) and the General Mixed Yule Coalescent (GMYC) model of Pons et al. (2006). Those SD methods were applied on the best-scoring ML tree resulting from the analyses of the concatenated dataset; they were also implemented on the COI and Cytb gene trees, which were well resolved. Analyses were carried out with default settings on a dedicated webserver (https://species.h-its. org/). For the GMYC model, both the default single threshold approach (results referred to as "GMYCs") of Pons et al. (2006) and the more parameter-rich approach (results referred to as "GMYCm") of Monaghan et al. (2009) were implemented, which allows the use of multiple thresholds to account for the potential heterogeneity of evolutionary rates among lineages. As inputs GMYC approaches require ultrametric trees (where all tips are equidistant from the root), which were generated using treePL (Smith & O'Meara 2012) with default settings.
Second, two distance-based methods were used: the Automatic Barcode Gap Discovery (ABGD) model of Puillandre et al. (2012) and the Assemble Species by Automatic Partitioning (ASAP) model of Puillandre et al. (2021). Those approaches were only used on COI and Cytb individual gene fragments for which there was enough intraspecific variability. ABGD analyses were performed on a dedicated webserver (https://bioinfo.mnhn.fr/abi/public/abgd/abgdweb. html) using default settings, a standard Kimura 2-parameter model (K80) and recursive partitioning strategies. ASAP analyses were carried out on a dedicated webserver (https://bioinfo.mnhn.fr/abi/ public/asap/#) using default settings and a K80 model. Although the ASAP approach is similar to ABGD, it includes a specific scoring system to identify the best-fitting set of putative species. Here the partition with the best asap-score (results referred to as "ASAP") as well as the partition with best W-score (results referred to as "ASAP-W") were considered, following the authors' recommendations.
Third, a multi-locus coalescent-based SD approach was used, as implemented in the program tr2 ). This method uses both gene trees and a guide tree as inputs, and relies on a Bayesian model comparison framework with rooted triplets. All six gene trees were used and the topology resulting from the analyses of the concatenated dataset was used as a guide tree.
Lastly, LIMES (Ducasse et al. 2020), an automatic calculation tool which compares partitions of SD analyses, was implemented. A SPART file (Miralles et al. 2022) was used (see Appendix S3) including all the results from SD analyses, the monophyletic clades inferred by the concatenated nuclear phylogeny, as well as previously defined morphospecies to rank them depending on the mean congruence (mCtax) obtained. Following authors' recommendations, specimens with missing gene fragments were excluded (e.g. Sesamia viettei and S. djenoensis).

Taxonomy
Genus Sesamia Guenée, 1852 Sesamia cretica species group Based on the results of the morphological studies, the Sesamia cretica group is split into the following subgroups: S. albivena subgroup, S. cretica subgroup, S. fuscifrontia subgroup, S. salama subgroup, S. viettei subgroup and S. wiltshirei subgroup. The S. albivena subgroup consists of S. albivena and of eight morphologically related species (seven of which are new to science). The S. cretica subgroup consists of S. cretica and of three morphologically related species (two of which are new to science). The S. fuscifrontia subgroup consists of S. fuscifrontia and of nine morphologically related species (eight of which are new to science). The S. salama subgroup consists of S. salama n. sp. and of one related species that could not be described due to the genitalia loss of the unique known specimen. The S. viettei subgroup consists of S. viettei species only. The S. wiltshirei subgroup consists of S. wiltshirei and of five morphologically related species new to science. In all, this study includes the description of 23 species which have been cross-checked against all Sesamia types preserved in museums. A supplemental description is also provided for the eight previously described species belonging to the S. albivena, S. cretica, S. fuscifrontia, S. viettei and S. wiltshirei subgroups. To facilitate species identification, identification keys are also provided for the five species subgroups based on male and female genitalia (see the identification keys at the end of the description paragraph of each species subgroup). A key to the Sesamia cretica species subgroups based on male and female genitalia is also provided just below along with a simplified tree that highlights the five key characters that have been used in the identification keys ( Figure 1).

Identification key to the Sesamia cretica species subgroups
Key based on internal morphology of the adult males (A) and females (B)     The Sesamia albivena subgroup consists of S. albivena, S. aethiopica n. sp., S. kafulo n. sp., S. kavirondo n. sp., S. maloukou n. sp., S. mocoensis n. stat., S. soyema n. sp., S. sudanensis n. stat. and Sesamia taenioleuca; it is characterized by the following combination of characters: (i) tegumen with prominent peniculi, vinculum with a small saccus; (ii) valve with sacculus and cucullus fused; costa short, costal spine broad, the internal side less sclerotized than the external side, spoon-shaped upwardly, most of the time produced downwardly into a sharp process; sacculus broad, well sclerotized with a strong narrowing below a more or less flared apex, inner side with a weak and only outlined harpe; cucullus not sclerotized, small and flat lying at a flatter angle on the sacculus, with scattered and papillated hairs; (iii) juxta trapezoidal with a very large laterally compressed medial projection; (iv) uncus angled and stout at base then thin tapering to a fine point, tufted with long hair on upper side; (v) aedeagus short and stout; manica with a carina crest produced into paired lateral lobes strongly spinose and most of the time with the presence of one or two elongated scobinate lobes; (vi) ostium bursae slightly Figure 1. Simplified phylogenetic tree highlighting the five subgroups belonging to the Sesamia cretica group (for the S. fuscifrontia subgroup, this simplified view does not show the paraphyletic nature of the subgroup). Based on the two identification keys, we identified five homologous characters that can be polarized through a parsimony analysis carried out under Mesquite (ancestral state in white, derived state in black; grey is used to underline a character that cannot be polarized). The corresponding characters are the following. Male characters: 1a, juxta without a large medial projection; 1b, juxta with a large medial projection; 2a, cucullus not sclerotized; 2b, cucullus sclerotized; 3a, inner side of sacculus with a clasper; 3b, inner side of sacculus without a clasper (note that for the S. cretica subgroup one of the four known species -S. rufescens-has a clasper); 4a, shape of the apex of costa not ampulla-like or not anvilshaped or not with teeth; 4b, shape of the apex of costa ampulla-like or anvil-shaped or with teeth. Female character: 5a, anterior margin of ostium bursae cup-shaped; 5b, anterior margin in the form of a wavy bead; 5c, anterior margin of ostium bursae funnelshaped, bilobate or villi like; 5d, anterior margin of ostium bursae with row or villi like; 5e, anterior margin of ostium bursae beanshaped, ribbon-shaped or villi like; 5f, anterior margin of ostium bursae kidney-shaped.

Diagnosis
See also the identification key to species of the albivena subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: tegumen with small prominent rounded peniculi; vinculum v-shaped at the bottom margin; base of costa small without indentation, internal side of costal spine curved, apex of the external side of the costal spine produced downwardly into a sharp pointed triangular process; apex of the sacculus very flared, inner side with a weak and only outlined harpe, slightly curved, 5.0 times longer than wide; apex of the cucullus very small and slightly rounded; juxta with a very large, thin and elongated, laterally compressed medial projection; posterior margin of ostium bursae strongly sclerotized, very flattened heart-shaped in the middle, barely concave, with rows of villi like, anterior margin cup-shaped with curved pointed tip on each side, ovipositor lobes curved, 2.1 times longer than wide.

Redescription
Figure 2a-f. Adults from both sexes have been described by Janse (1939) and Tams and Bowden (1953) based on specimens from Kenya, South Africa, Uganda and Zimbabwe. The specimens from Kenya and Uganda are considered to belong to a different species (i.e. S. kavirondo n. sp.), which can explain why their description does not fit well with those of the holotype preserved in the NHM. The latter still holds true even if the colours of old specimens could have become ochraceous with time, in contrast with the buff-pink and flesh-pink colours recorded by Janse (1939) and Tams & Bowden (1953), respectively. Our redescription is only based on specimens from the South Africa and Zimbabwe.
General shape of the female's forewings more elongated at the apex than those of the male. Antennae ochraceous, serrate in the male, filiform in the female, flagellum adorned dorsally with pale ochraceous scales; palpus ochraceous in male, buff in female; eyes dark brown. Wing pattern similar in both sexes, thorax and wing colour pale ochraceous; head and thorax covered with long pale ochraceous hairs. Forelegs pale ochraceous, suffused with ochraceous-pink, fore tibia infuscate. Forewings pale ochraceous tinged with pinkish-buff more densely along costal, apex and termen areas, darkened to rufous at termen in some specimens; no transverse markings, veins streaked with white, some fuscous antemedial and postmedial markings more or less visible; a diffuse longitudinal fuscous fascia along lower margin of cell, partly within, partly without cell from base of cell extending almost to termen; a dark fuscous-black marking at lower angle of cell followed by a smaller one, indicating the reniform; outer margin adorned with fuscous-black elongated markings between the veins; fringe infuscate with a white basal line. Hind wings white without markings, fringe white with a basal fuscous line. Underside of forewings heavily suffused with pinkish-buff or buff in costa, apex and termen areas and below the cell; outer margin adorned with fuscous-black elongated markings between the veins, fringe fuscous with a white basal line, suffusion is more pronounced in male. Underside of hind wings white, costa and apex slightly suffused with buff and fuscous, fringe white with a fuscous basal line. Forewings length: males 26.5 mm (min-max 24-28 mm, N = 8); female 30 mm (N = 1). Male genitalia (Figure 3a, h). Tegumen with small prominent rounded peniculi; vinculum narrow with a small saccus, v-shaped at the bottom margin, w-shaped at the top margin with an indentation. Valve with sacculus and cucullus fused; costa short, the base small without indentation, costal spine large, internal side curved, less sclerotized than the external side, apex rounded upwardly, external side slightly wavy, apex produced downwardly into a sharp pointed triangular process; sacculus broad, well sclerotized, with a strong narrowing below a very flared apex, inner side with a weak and only outlined harpe, slightly curved, heavily sclerotized, 5.0 times longer than wide; cucullus narrow lying at a flatter angle on the sacculus, base bead-shaped, slightly sclerotized, apex very small and slightly rounded, not sclerotized, with scattered and papillated hairs; juxta trapezoidal, with a very large, thin and elongated, laterally compressed medial projection, dentated at apex; uncus thin, curved at middle, tapering near apex to a fine curved point, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose, two elongated scobinate lobes, one straight, the other one strongly curved. Female genitalia (Figure 3o). The genitalia slide No. 1295 preserved in the NHM is the only female genitalia o, female genitalia; p, S. aethiopica, female genitalia; b, i, q, S. kafulo: b, male genitalia; i, male aedeagus; q, female genitalia; c, j, S. kavirondo: c, male genitalia; j, male aedeagus; d, k, S. maloukou: d, male genitalia; k, male aedeagus; e, l, S. mocoensis: e, male genitalia; l, male aedeagus; r, S. soyema, female genitalia; f, m, s, S. sudanensis: f, male genitalia; m, male aedeagus; s, female genitalia; g, n, t, S. taenioleuca: g, male genitalia; n, male aedeagus; t, female genitalia. Scale bars: male genitalia, 1 mm for valves and 0.5 mm for aedeagus; female genitalia, 1 mm. preparation available. When comparing it with other female genitalia preparations of related species of the S. cretica group (i.e. S. rufescens and S. sudanensis n. stat. preserved in the NHM and S. aethiopica n. sp., S. kafulo n. sp., S. soyema n. sp. and S. taenioleuca preserved in the MNHN and prepared by B. Le Ru), it was suspected that this particular specimen was poorly prepared as indicated by the reversal of the posterior lip of the anteostial pad. In order to harmonize all descriptions, the female genitalia of S. albivena is redescribed. Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin strongly sclerotized, very flattened heart-shaped in the middle, barely concave, with rows of villi like, cup-shaped with curved pointed tip on each side; ductus bursae short, not sclerotized; corpus bursae short, ovoid, without signa; ovipositor lobes curved, 2.1 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores.

Remarks
Sesamia albivena is widely distributed throughout South Africa and Zimbabwe. In addition to the specimens listed in the other material section, 70 male and female specimens are preserved in the TMSA. These specimens were collected from multiple localities in South Africa (

Diagnosis
See also the identification key to species of the albivena subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the female genitalia: posterior margin of ostium bursae, strongly sclerotized, heartshaped, slightly convex in the middle with rows of villi like, pointed base extending towards the ductus bursae, anterior margin cup-shaped with very sharp curved ends on each side; ovipositor lobes curved, 2.2 times longer than wide. Sesamia aethiopica n. sp. female genitalia are very similar to those of S. kafulo n. sp.; however, posterior margin is less concave in the middle than in S. kafulo n. sp. and the pointed base extending towards the ductus bursae is wider than in S. aethiopica n. sp.
Description Figure 2g, h. Antennae pinkish-buff, filiform, flagellum adorned dorsally with pale beige scales; palpus ochraceous; eyes dark brown. Head and thorax covered with long pale beige hairs, abdomen pink beige. Legs ochraceous, suffused with beige. Forewings pinkish-buff tinged with buff more densely along costal, apex and termen areas, darkened to rufous at termen; no transverse markings, veins streaked with white, some brown antemedial and postmedial markings; a diffuse longitudinal pale pink fascia along lower margin of cell, partly within, partly without cell from base of cell extending almost to termen; a dark fuscous-black marking at lower angle of cell followed by a smaller one, indicating the reniform; outer margin adorned with fuscous-black elongated markings between the veins; fringe infuscate with a white basal line. Hind wings pinkish-white without markings, fringe white with a basal fuscous line. Underside of forewings heavily suffused with fuscous in costa, apex, termen areas and below the cell; outer margin adorned with fuscous-black elongated markings between the veins, fringe infuscate with a white basal line. Underside of hind wings white slightly suffused with fuscous, fringe white with a fuscous basal line. Forewings length: female 22.0 mm (N = 1). Female genitalia (Figure 3p). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin strongly sclerotized, heart-shaped slightly convex in the middle with rows of villi like, pointed base extending towards the ductus bursae, anterior margin cup-shaped with very sharp curved ends on each side; ductus bursae short, slightly sclerotized on ostium side; corpus bursae short, ovoid, without signa; ovipositor lobes curved, 2.2 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores.

Etymology
Named after Ethiopia, the country where the species has been collected, treated as an adjective.

Distribution
Ethiopia. The recorded locality is from East African evergreen and semi-evergreen bushland and thicket (mosaic #38) vegetation mosaic (White 1983) ( Figure 5).

Remarks
Although morphologically close to S. kafulo n. sp., S. aethiopica n. sp. female genitalia are easily distinguished with characters of the posterior margin of the anteostial pad. The only known specimen of this species was collected at 9 pm, 3 h after sunset, at the same time as the only known specimen of S. soyema n. sp.

Diagnosis
See also the identification key to species of the albivena subgroup below. This species can be distinguished from all other known species of the subgroup by the following combination of characters of the male and female genitalia: tegumen with small rounded peniculi; vinculum v-shaped at the bottom margin; base of costa large with indentations, internal side of costal spine flat, apex rounded upwardly, cap-shaped at the end, external side of costal spine produced downwardly into a sharp pointed triangular process; apex of the sacculus very flared, inner side with a weak and only outlined harpe, 6.0 times longer than wide; apex of the cucullus small and slightly rounded; juxta with a large, robust and thick, medial projection; posterior margin of ostium bursae heart-shaped, slightly convex in the middle, anterior margin cup-shaped with straight gutter-shaped ends on each side; ovipositor lobes curved, 2.0 times longer than wide. Sesamia kafulo n. sp. female genitalia are very similar to those of S. aethiopica n. sp.; however, the anteostial pad is more concave in the middle than in S. aethiopica n. sp. and the pointed base extending towards the ductus bursae is narrower than in S. aethiopica n. sp.
Description Figure 2i-l. Antennae ochraceous, serrate in the male, filiform in the female, flagellum adorned dorsally with buff scales; palpus buff; eyes dark brown. Wing pattern similar in both sexes, thorax and wing colour buff; head and thorax covered with long pale buff hairs. Forelegs pale brown, otherwise buff. Forewings pale buff tinged with dark buff more densely along costal, apex and termen areas; no transverse markings; a diffuse longitudinal ochraceous fascia along lower margin of cell, partly within, partly without cell from base of cell extending almost to termen; a dark fuscous-black marking at lower angle of cell indicating the reniform; outer margin adorned with fuscous-black elongated markings between the veins; fringe infuscate with a white basal line. Hind wings white slightly suffused with pale buff in costa, apex and termen areas, fringe concolour with a basal white line. Underside of forewings heavily suffused with dark-buff in costa, apex and termen areas; outer margin adorned with fuscous-black elongated markings between the veins, fringe infuscate with a white basal line. Underside of hind wings white, costa and apex suffused with buff, fringe infuscate with a white basal line. Forewings length: males 27.5 mm (min-max 24-31 mm, N = 11); female 29.0 mm (min-max 19-26 mm, N = 11).
Male genitalia (Figure 3b, i). Tegumen with small rounded peniculi; vinculum narrow with a small saccus, v-shaped at the bottom margin, w-shaped at the top margin with an indentation. Valve with sacculus and cucullus fused; costa short, base large with indentations, costal spine large, internal side flat, less sclerotized than the external side, apex rounded upwardly, cap-shaped at the end, external side slightly wavy produced downwardly into a sharp pointed triangular process; sacculus broad, well sclerotized with a strong narrowing below a very flared apex, inner side with a weak and only outlined harpe, slightly curved, 6.0 times longer than wide; cucullus narrow lying at a flatter angle on the sacculus, base bead-shaped, slightly sclerotized, the apex small and slightly rounded, not sclerotized, with scattered and papillated hairs; juxta trapezoidal, with a large, robust and thick, medial projection, slightly dentated at apex; uncus thin, curved at middle, tapering near apex to a fine curved point, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose, two elongated scobinate lobes, one straight, the other one strongly curved. Female genitalia (Figure 3q). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin strongly sclerotized, heart-shaped, slightly convex in the middle with rows of villi like, pointed base extending towards the ductus bursae, anterior margin cup-shaped with straight guttershaped ends on each side; ductus bursae short, slightly sclerotized on ostium side; corpus bursae short, ovoid, without signa; ovipositor lobes curved, 2.0 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores.

Etymology
Named after Kafulo, the town close to the place where the species was collected; treated as a noun in apposition.

Bionomics
Biology unknown. The moths were caught with light traps in grasslands inhabited with various Poales species belonging to the following genera: Andropogon, Cymbopogon, Hyparrhenia, Megathyrsus, Panicum L., 1753 and Sporobolus.

Diagnosis
See also the identification key to species of the albivena subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male genitalia: tegumen with small flat peniculi; vinculum v-shaped at the bottom margin; base of costa large without indentation, internal side of costal spine curved, apex rounded, upwardly cap-shaped at the end, external side of the costal spine produced downwardly into a large triangular process; apex of the sacculus very flared, inner side with a weak and only outlined harpe, slightly curved, 4.0 times longer than wide; apex of the cucculus very small, almost flat; juxta with a very large, thin and elongated, laterally compressed medial projection.
Description Figure 2m-o. Antennae ochraceous, serrate, flagellum adorned dorsally with pale buff scales; palpus ochraceous; eyes dark brown. Thorax pale buff, wing pale ochraceous; head and thorax covered with long pale buff hairs. Forelegs dark ochraceous, otherwise pale ochraceous. Forewings pale ochraceous tinged with pinkish-buff more densely along costal, apex and termen areas; no transverse markings, some fuscous antemedial and postmedial markings more or less visible; a diffuse longitudinal fuscous fascia along lower margin of cell, partly within, partly without cell from base of cell extending almost to termen; a dark fuscous marking at lower angle of cell followed by a smaller one, indicating the reniform; outer margin adorned with fuscousblack elongated markings between the veins; fringe infuscate with a white basal line. Hind wings white without markings, fringe white with a basal fuscous line. Underside of forewings suffused with fuscous and pinkishbuff in costa, apex and termen areas and around the cell; outer margin adorned with fuscous-black elongated markings between the veins, fringe fuscous with a white basal line. Underside of hind wings white, costa and apex slightly suffused with fuscous, fringe white with a fuscous basal line. Forewings length: males 24.0 mm (min-max 20-28 mm, N = 4) Male genitalia (Figure 3c, j). Tegumen with small flat peniculi; vinculum narrow with a small saccus, v-shaped at the bottom margin, w-shaped at the top margin with an indentation. Valve with sacculus and cucullus fused; costa short, base large without indentation, costal spine large, internal side curved, less sclerotized than the external side, apex rounded, upwardly cap-shaped at the end, external side slightly wavy, produced downwardly into a large triangular process; sacculus broad, well sclerotized with a strong narrowing below a very flared apex, inner side with a weak and only outlined harpe, slightly curved, 4.0 times longer than wide; cucullus narrow lying at a flatter angle on the sacculus, base bead-shaped, apex very small, almost flat, not sclerotized, with scattered and papillated hairs; juxta trapezoidal, with a very large, thin and elongated, laterally compressed medial projection, dentated at apex; uncus thin, curved at middle, tapering near apex to a fine curved point, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose, two elongated scobinate lobes, one straight, the other one strongly curved.

Etymology
Named after Kavirondo, the name of the region along the north-east coast of Victoria lake where the holotype was collected; treated as a noun in apposition.

Distribution
Kenya and Uganda. The recorded localities are from two different vegetation mosaics: lowland rain forest and secondary grassland (mosaic #11a), and East African evergreen bushland and secondary Acacia wooded grassland (mosaic #45) (White 1983) ( Figure 5).

Diagnosis
See also the identification key to species of the albivena subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male genitalia: tegumen with small flat peniculi; vinculum u-shaped at the bottom margin; base of costa large with indentations, internal side of costal spine slightly convex, apex rounded upwardly, produced downwardly into a small blunt triangular process; apex of the sacculus slightly flared, inner side with a weak and only outlined harpe, 5.0 longer than wide; apex of cucullus almost flat; juxta with a large, robust and thick, medial projection in dorsal position.
Description Figure 2p, q. Antennae buff serrate, flagellum adorned dorsally with pale buff scales; palpus pale buff; eyes brown. Head covered with long dark buff hairs, thorax covered with long pale buff hairs, abdomen pale buff. Forelegs pale brown, otherwise buff. Forewings dark buff; no transverse markings, veins streaked with pale buff; a diffuse longitudinal fuscous fascia along lower margin of cell, partly within, partly without cell from base of cell extending almost to termen; a fuscousblack marking at lower angle of cell indicating the reniform; outer margin adorned with fuscous-black elongated markings between the veins; fringe infuscate with a white basal line. Hind wings pinkish white, fringe concolour with a basal white line. Underside of forewings heavily suffused with dark-buff in costa, apex and termen areas; outer margin adorned with fuscous-black elongated markings between the veins, fringe infuscate with a white basal line. Underside of hind wings pinkish white, costa and apex slightly suffused with buff, fringe concolour. Forewings length: males 25.0 mm (min-max 24-26 mm, N = 2). Male genitalia (Figure 3d, k). Tegumen with small flat peniculi; vinculum narrow with a small saccus, u-shaped at the bottom margin, w-shaped at the top margin with an indentation. Valve with sacculus and cucullus fused; costa short, base large with indentations, costal spine large, internal side slightly convex, less sclerotized than the external side, apex rounded upwardly, external side slightly wavy produced downwardly into a small blunt triangular process; sacculus broad, well sclerotized with a strong narrowing below a slightly flared apex, inner side with a heavily sclerotized bead, 5.0 times longer than wide; cucullus narrow, lying at a flatter angle on the sacculus, base bead-shaped, apex small, almost flat, not sclerotized, with scattered and papillated hairs; juxta trapezoidal, with a large, robust and thick, medial projection in dorsal position, slightly dentated at apex; uncus thin, curved at middle, tapering near apex to a fine curved point, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose, one elongated strongly curved scobinate lobe.

Etymology
Named after Maloukou, the name of the village along the Congo river where the only two known specimens have been collected; treated as a noun in apposition.

Bionomics
Biology unknown. The moths were caught with a light trap in grasslands inhabited with various Poales species belonging to the following genera: Cymbopogon, Hyparrhenia, Megathyrsus, Panicum and Sporobolus.

Distribution
Republic of Congo. The recorded localities are from lowland rain forest and secondary grassland (mosaic #11a) vegetation mosaic (White 1983) ( Figure 5).

Remarks
There may be another elongated straight scobinate lobe like in all related species but it is not visible because of the poor genitalia preparation. Tams

Diagnosis
See also the identification key to species of the albivena subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male genitalia: tegumen with rounded peniculi; vinculum narrow with a saccus u-shaped at the bottom margin; base of costa large without indentation, internal side of costal spine flat, apex rounded, upwardly cap-shaped, external side of costal spine produced downwardly into a sharp triangular process; apex of the sacculus slightly flared, inner side with a weak and only outlined harpe, 6.0 times longer than wide; apex of the cucullus slightly rounded; juxta with a very large, thin and elongated, laterally compressed medial projection.

Redescription
Figure 6a-f. Antennae ochraceous, serrate in the male, filiform in the female, flagellum adorned dorsally with pale buff scales; palpus ochraceous; eyes dark brown. Head and thorax covered with long pale buff hairs, abdomen pale buff. Forelegs ochraceous, otherwise pale ochraceous. Wing pattern similar in both sexes. Forewings pale ochraceous tinged with buff more densely along costal, apex and termen areas, darkened to rufous at termen in some specimens; no transverse markings, veins streaked with white; a diffuse longitudinal buff fascia along lower margin of cell, partly within, partly without cell from base of cell extending almost to termen; a dark fuscous-black marking at lower angle of cell followed by a smaller one, indicating the reniform; outer margin adorned with more or less visible fuscous-black elongated markings between the veins; fringe infuscate with a white basal line. Hind wings white without markings, fringe white with a basal buff line. Underside of forewings heavily suffused with buff in costa, apex and termen areas; outer margin adorned with fuscous-black elongated markings between the veins, fringe infuscate with a white basal line. Underside of hind wings white, costa and apex slightly suffused with buff and fuscous, fringe white, slightly infuscate in apex, a fuscous basal line. Forewings length: males 28.5 mm (min-max 26-31 mm, N = 11); females 31.7 mm (min-max 30-32 mm, N = 3). Male genitalia (Figure 3e, l). Tegumen with small rounded peniculi; vinculum narrow with a small saccus, u-shaped at the bottom margin, w-shaped at the top margin with an indentation. Valve with sacculus and cucullus fused; costa short, base large without indentation, costal spine large, internal side flat, less sclerotized than the external side, apex rounded upwardly cap-shaped, external side slightly wavy, produced downwardly into a sharp triangular process; sacculus broad, well sclerotized with a strong narrowing below a slightly flared apex, inner side with a weak and only outlined harpe, slightly curved, 6.0 times longer than wide; cucullus narrow, lying at a flatter angle on the sacculus, base bead-shaped, sclerotized, apex slightly rounded, not sclerotized, with scattered and papillated hairs; juxta trapezoidal, with a very large, thin and elongated, laterally compressed medial projection, dentated at apex; uncus thin, curved at middle, tapering near apex to a fine curved point, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose, two elongated scobinate lobes, one straight, the other one strongly curved.

Remarks
Tams & Bowden (1953) noticed the great morphological resemblance between S. albivena and S. mocoensis, which led them to consider S. mocoensis as a subspecies of S. albivena. However, when comparing male genitalia of the two taxa, notable differences are observed; in the male the most discriminating characters are the costa, the internal side of the costal spine, the inner side of the apex of the sacculus and the apex of the cucullus (see also the identification key to species of the albivena subgroup below). Unfortunately, without any available female genitalia preparation, it is not possible to describe it.

Diagnosis
See also the identification key to species of the albivena subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the female genitalia: posterior margin of ostium bursae rounded heart-shaped with a pointed base extending towards the ductus bursae, anterior margin cupshaped with straight gutter-shaped ends on each side; ovipositor lobes curved, 2.0 times longer than wide. Figure 6g, h. Antennae ochreous, filiform, flagellum adorned dorsally with pale beige scales; palpus buff; eyes dark brown. Head and thorax covered with long ochreous buff hairs, abdomen pink beige. Legs ochraceous buff, suffused with beige. Forewings pale-buff tinged with ochreous buff more densely along costal, apex and termen areas, darkened to rufous at termen; no transverse markings, veins streaked with white; a diffuse longitudinal fuscous fascia along lower margin of cell, partly within, partly without cell from base of cell extending almost to termen; a fuscous marking at lower angle of cell, indicating the reniform; outer margin adorned with fuscous-brown markings between the veins; fringe infuscate with a white basal line. Hind wings white without markings, fringe white with a basal fuscous line. Underside of forewings pale buff heavily suffused with buff in costa, apex, termen areas; outer margin adorned with fuscous-brown markings between the veins, fringe infuscate with a white basal line. Underside of hind wings white slightly suffused with fuscous in costa and apex areas, fringe white, slightly infuscate in apex, with a fuscous basal line. Forewings length: female 30.0 mm (N = 1). Female genitalia (Figure 3r). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin strongly sclerotized, rounded heart-shaped, slightly convex in the middle, with rows of villi like, a pointed base extending towards the ductus bursae, anterior margin cup-shaped with straight gutter-shaped ends on each side; ductus bursae short, slightly sclerotized on ostium side; corpus bursae short, ovoid, without signa; ovipositor lobes curved, 2.0 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores.

Etymology
Named after Soyema, the town close to the place where the species was collected; treated as a noun in apposition.

Bionomics
Biology unknown. The adults were caught with a light trap in grasslands surrounding banks of a river inhabited with various Poales species belonging to the following genera: Andropogon, Cymbopogon, Hyparrhenia, Megathyrsus and Sporobolus.

Distribution
Ethiopia. The recorded locality is from East African evergreen and semi-evergreen bushland and thicket (mosaic #38) vegetation mosaic (White 1983) ( Figure 5). Tams

Diagnosis
See also the identification key to species of the albivena subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: tegumen with triangular peniculi; vinculum with a saccus u-shaped at the bottom margin; base of costa small without indentation, internal side of costal spine flat, apex rounded upwardly, cap-shaped at the end, apex of external side of costal spine produced downwardly into one or two small spikes; apex of the sacculus very flared, inner side with a weak and only outlined harpe, curved, 8.0 times longer than wide; apex of the cucullus large, rounded; juxta with a very large, thin and elongated, laterally compressed medial projection; posterior margin of ostium bursae strongly sclerotized, heartshaped, strongly convex in the middle, square base extending towards the ductus bursae, anterior margin cup-shaped with very sharp straight ends on each side; ovipositor lobes curved, 2.7 times longer than wide.
Redescription Figure 6i-n. Male ochraceous, female pale buff; antennae ochraceous, serrate, cilia short, in the male, pale buff, filiform in the female, flagellum adorned dorsally with ochraceous scales in male, with pale buff in female; palpus pale buff; eyes dark brown. Head and thorax covered with long ochraceous buff hairs in male, long pale buff hairs in female. Legs ochraceous buff, suffused with beige. Forewings buff tinged with ochraceous buff more densely along costal, apex and termen areas, darkened to rufous at termen, more pronounced in male; no transverse markings, veins streaked with white; some fuscous postmedial markings more or less visible; a diffuse longitudinal ochraceous fascia along lower margin of cell, partly within, partly without cell from base of cell extending almost to termen; a fuscous marking at lower angle of cell, indicating the reniform; outer margin adorned with elongated fuscous-brown markings between the veins; fringe fuscous in male, infuscate in female, with a pale buff basal line. Hind wings white slightly suffused with buff in male, without markings, fringe infuscate in male, white, slightly infuscate in costa in female, basal line fuscous. Underside of forewings buff, heavily suffused with dark buff in costa, apex, termen areas and below the cell in male, slightly suffused in female; outer margin adorned with elongated fuscous-brown markings between the veins, fringe infuscate with a pale buff basal line. Underside of hind wings white suffused with fuscous in costa, apex and termen areas in male, slightly suffused in costa and apex in female, fringe infuscate in male, slightly infuscate in apex in female, with a fuscous basal line. Forewings length: males 28.0 mm (min-max 28 mm, N = 2); female 31.0 mm (N = 1). Male genitalia (Figure 3f, m). Tegumen with triangular peniculi; vinculum narrow with a small saccus, u-shaped at the bottom margin, w-shaped at the top margin with an indentation. Valve with sacculus and cucullus fused; costa short, base small without indentation, costal spine large, internal side flat, less sclerotized than the external side, apex rounded upwardly, cap-shaped at the end, external side slightly wavy, apex produced downwardly into one or two small spikes; sacculus broad, well sclerotized with a strong narrowing below a very flared apex, inner side with a weak and only outlined harpe, slightly curved, heavily sclerotized, 8.0 times longer than wide; cucullus narrow lying at a flatter angle on the sacculus, base bead-shaped slightly sclerotized, apex large rounded, not sclerotized, with scattered and papillated hairs; juxta trapezoidal, with a very large, thin and elongated, laterally compressed medial projection, dentated at apex; uncus thin, curved at middle, tapering near apex to a fine curved point, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose, two elongated scobinate lobes, one straight, the other one strongly curved. Female genitalia (Figure 3s). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin strongly sclerotized, heart-shaped strongly convex in the middle with rows of villi like, square base extending towards the ductus bursae, anterior margin cup-shaped with very sharp straight ends on each side; ductus bursae short, slightly sclerotized on ostium side; corpus bursae short, ovoid, without signa; ovipositor lobes curved, 2.7 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores.

Remarks
Tams & Bowden (1953) noticed the great morphological resemblance between S. albivena and S. sudanensis, which led them to consider S. sudanensis as a subspecies of S. albivena. However, when comparing both male and female genitalia of the two taxa, notable differences are observed; in the males the most discriminating characters are the internal side and the apex of the external side of the costal spine, the inner side of the apex of the sacculus, the apex of the cucullus and the vinculum; in the females the posterior margin of the anteostial pad (see also the identification key 2 below). Tams and Bowden (1956)  See also the identification key to species of the albivena subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: tegumen with triangular peniculi; vinculum with a saccus v-shaped at the bottom margin; base of costa large with indentations, internal side of costal spine straight, apex rounded spoon-shaped upwardly, external side of costal spine widening strongly towards the apex, produced downwardly into a very large triangular process; apex of the sacculus slightly flared, inner side with a weak and only outlined harpe, curved, 6.0 times longer than wide; apex of cucullus small and rounded; juxta with a large, robust and thick, laterally compressed mediodorsal projection; posterior margin of ostium bursae heart-shaped with a rounded base, anterior margin cup-shaped with very sharp slightly curved ends on each side; ovipositor lobes curved, 2.8 times longer than wide.

Description
Figure 6o-r. The male of this species has been described in detail by Janse (1939) and Tams & Bowden (1953). The female is recorded here for the first time: antennae filiform, otherwise the general shape of the female's forewings is more elongated at the apex than those of the male but the wing pattern and colour are similar in both sexes. Forewings length: males 29.7 mm (minmax 27-31 mm, N = 8); female 32.5 mm (min-max 32-33 mm, N = 2). Male genitalia (Figure 3g, n). Tegumen with small triangular peniculi; vinculum narrow with a small saccus, v-shaped at the bottom margin, w-shaped at the top margin with an indentation. Valve with sacculus and cucullus fused; costa short, base large with indentations, costal spine large, internal side straight, less sclerotized than the external side, apex rounded spoon-shaped upwardly, external side slightly wavy, widening strongly towards the apex, produced downwardly into a very large triangular process; sacculus broad, well sclerotized with a strong narrowing below a slightly flared apex, inner side with a weak and only outlined harpe, slightly curved, heavily sclerotized, 6.0 times longer than wide; cucullus narrow lying at a flatter angle on the sacculus, base bead-shaped, slightly sclerotized, apex small and rounded, not sclerotized, with scattered and papillated hairs; juxta trapezoidal, with a large, robust and thick, laterally compressed mediodorsal projection, slightly dentated at apex; uncus thin, curved at middle, tapering near apex to a fine curved point, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose, two elongated scobinate lobes, one straight, the other one strongly curved. Female genitalia (Figure 3t). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin strongly sclerotized with rows of villi like, slightly convex in the middle, base rounded, anterior margin cup-shaped with very sharp slightly curved ends on each side; ductus bursae short, slightly sclerotized on ostium side; corpus bursae short, ovoid, without signa; ovipositor lobes curved, 2.8 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores.  (White 1983) ( Figure 5).

Remarks
In addition to the specimens recorded in the additional material section, 14 male and female specimens are preserved in the TMSA, collected from one locality in South Africa (Limpopo: Duivelskloof) and from several localities in Zimbabwe (Bulawayo, Griffin, Emangeni, Mazoe, Salisbury (Harare), Sawmills, Shabana).
Identification key to species of Sesamia albivena subgroup The Sesamia cretica subgroup consists of S. cretica, S. ihambane n. sp., S. kikuyuensis n. sp. and S. rufescens; it is characterized by the following combination of characters: (i) tegumen with prominent peniculi, vinculum with a small saccus; (ii) valve with sacculus and cucullus fused; costa short, costal spine broad, the internal side less sclerotized than the external side, spoon-shaped upwardly, produced downwardly into a sharp process, sometimes a strong clasper; sacculus broad, well sclerotized; cucullus not sclerotized, broadly rounded with scattered and papillated hairs; (iii) juxta heart-shaped or trapezoidal, with a laterally small compressed medial projection; (iv) uncus angled and stout at base then thin tapering to a fine point, tufted with long hair on upper side; (v) aedeagus short and stout; manica with a carina crest produced into paired lateral lobes strongly spinose, presence of one or two elongated scobinate lobes or a tongue-shaped flat cornutus at apex of vesica; (vi) ostium bursae strongly transverse, posterior margin narrow bandshaped, anterior margin funnel-shaped, bilobate, with pointed tip on each side or villi like.  1975-372, 22 (NHM).

Diagnosis
See also the identification key to species of the cretica subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: tegumen with small flat peniculi; vinculum u-shaped at the bottom margin; internal side of the costal spine convex, apex rather triangular; apex of the sacculus slightly flared, inner side with a weak and only outlined harpe; cucullus broadly rounded; juxta with a small laterally compressed medial projection; posterior margin of ostium bursae ribbon-shaped slightly convex in the middle ending in a point on each side; anterior margin funnel-shaped with a small medial depression; ovipositor lobes curved, 2.2 times longer than wide.
Description Figure 7a-d. Both sexes were described in great details by Tams (1938) and Tams & Bowden (1953). Forewings length: males 26.3 mm (min-max 24-28 mm, N = 7); female 30.4 mm (min-max 29-32 mm, N = 7). Male genitalia (Figure 8a, e). Tegumen with small flat peniculi; vinculum narrow with a small saccus, u- shaped at the bottom margin, w-shaped at the top margin; valve with sacculus and cucullus fused; costa short, costal spine large, internal side convex, apex rather triangular; sacculus broad, well sclerotized with a strong narrowing below a slightly flared apex, inner side with a weak and only outlined harpe, heavily sclerotized; cucullus not sclerotized, broadly rounded, lying at a flatter angle on the sacculus, with scattered and papillated hairs; juxta trapezoidal, with a small laterally compressed medial projection, dentated at apex; uncus angled and stout at base, narrowing in distal part, tapering near apex to a fine curved point, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose.
Female genitalia (Figure 8i). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin ribbon-shaped, slightly convex in the middle ending in a point on each side, anterior margin funnel-shaped with a small medial depression on the posterior side, anterior side concave, strongly sclerotized; ductus bursae very short, slightly sclerotized on the ostium side; corpus bursae short, ovoid, without signa; ovipositor lobes curved, 2.2 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores without spatulate tips.

Bionomics
This is a common pest species of maize, sorghum and sugarcane (Saccharum officinarum L., 1753). The species  is also recorded from wild grasses, especially Panicum repens L., 1762; however, considering that this species is difficult to identify without genitalia dissection, other host plant records need to be considered with caution.

Distribution
Sesamia cretica is widely distributed in the Palearctic (mostly in the western Palearctic) and in the Oriental region; it is also found in the Afrotropics (mostly in eastern Africa, but also in North Cameroon and Mauritania). In Africa this species is generally found in dry vegetation mosaicsundifferentiated Sudanian and Ethiopian woodlands (mosaics #29), Somalia-Masaï Acacia-Commiphora deciduous bushland and thicket (mosaics #42), Sahel Acacia wooded grassland and deciduous bushland (mosaics #43)and even in very dry vegetation mosaicssemi-desert grassland and shrubland (mosaics #54a, 54b, 55) (White 1983).

Remarks
Sesamia (Nonagria) cyrnaea Mabille, 1866 from Corsica has been considered erroneously as a synonym of S. cretica by Hampson (1910), then by Poole (1989) and finally by Zilli et al. (2005). Indeed, the male antennae of S. cyrnaea is longly bipectinate, where in S. cretica it is serrate; this suggests that S. cyrnaea is most likely a synonym of S. nonagrioides.

Diagnosis
See also the identification key to species of the cretica subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: tegumen with small flat peniculi; vinculum with rectangular saccus; internal side of the costal spine flat, apex rather rectangular upwardly ending in a point, produced downwardly into a sharp triangular process; sacculus broad, tapering sharply towards the apex, inner side with a weak harpe, only outlined; cucullus broadly spatulated, in the prolongation of the sacculus; juxta diamond-shaped, with a very small laterally compressed medial projection; vesica with a small tongue-shaped flat cornutus villi like; posterior margin of ostium bursae very open v-shaped, widening on each side; anterior margin bilobate with a v-shaped area between them, each lobe cup-shaped, with sharp curved ends on external side; ovipositor lobes curved, 2.4 times longer than wide.
Description Figure 7e-h. The general shape of the female's forewings is more elongated at the apex than those of the male. Antennae pinkish-buff, serrate in the male, filiform in the female, flagellum adorned dorsally with pinkish scales; palpus grey suffused with pinkish; eyes brown. Head and thorax covered with long pale pinkish hairs, abdomen white suffused with pinkish. Forelegs grey, otherwise pinkish. Wing pattern similar in both sexes. Forewings pinkish-buff suffused with brown more densely along costal, apex, termen and tornus areas; no transverse markings, some brown antemedial and postmedial markings, a longitudinal grey brown fascia along lower margin of cell, partly within, partly without cell from base of cell extending almost to termen; outer margin adorned with small black markings between the veins; fringe infuscate with a wide dark buff basal line. Hind wings pinkish-white to white, without markings, fringe pinkish-white with a basal pinkish line. Underside of forewings pinkish-buff heavily suffused with pinkish beige and brown in costa, apex and termen areas; outer margin adorned with black markings between the veins, fringe infuscate with a wide buff basal line. Underside of hind wings white to pinkish white slightly suffused with brown in costa and apex areas, fringe infuscate in the apex, otherwise white with a fuscous basal line. Forewings length: males 19.8 mm (min-max 17-22 mm, N = 10); female 23.5 mm (min-max 19-26 mm, N = 10). Male genitalia (Figure 8b, f). Tegumen with small flat peniculi; vinculum narrow with a small rectangular saccus, with a bulge in the middle at the bottom margin, w-shaped at the top margin; valve with sacculus and cucullus fused; costa short, costal spine large, internal side flat, apex rather rectangular, upwardly ending in a point, produced downwardly into a sharp triangular process strongly sclerotized; sacculus broad, well sclerotized, tapering sharply towards the apex, inner side with a weak and only outlined harpe; cucullus not sclerotized, broadly spatulated, in the prolongation of the sacculus with scattered and papillated hairs; juxta diamondshaped, with a very small laterally compressed medial projection; uncus angled and stout at base, tapering near apex to a fine point, tufted with hair on upper side; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose, vesica with a small tongue-shaped flat cornutus villi like. Female genitalia (Figure 8j). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin very open v-shaped, slightly sclerotized, widening on each side, anterior margin bilobate with a v-shaped area between them, each lobe cup-shaped, with sharp curved ends on external side, strongly sclerotized; ductus bursae very short, slightly sclerotized on the ostium side; corpus bursae short, ovoid, without signa; ovipositor lobes curved, 2.4 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores without spatulate tips.

Etymology
Named after the Inhambane region in Mozambique, which is the south part of the Zanzibar-Inhambane coastal forest mosaic also known as the Swahili coastal forests and woodlands; treated as a noun in apposition.

Bionomics
The larvae were collected in stems of Rottboellia cochinchinensis in patches of grasslands in Zanzibar-Inhambane vegetation mosaic inhabited with various Poales species belonging to the following genera: Cymbopogon, Hyparrhenia, Megathyrsus and Panicum.

Diagnosis
See also the identification key to species of the cretica subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: tegumen with small flat peniculi; vinculum with a rectangular saccus; internal side of costal spine flat, apex rather rectangular, upwardly ending in a point, produced downwardly into a sharp pointed process strongly sclerotized; sacculus with a slightly flared apex, inner side with a weak and only outlined harpe, heavily sclerotized; cucullus broadly ovoid, in the prolongation of the sacculus; juxta bilobate, each lobe diamondshaped; vesica with a small tongue-shaped flat cornutus villi like; posterior margin of ostium bursae slightly convex, anterior margin bilobate with a convex area between them, each lobe cup-shaped with sharp curved ends on external side; ovipositor lobes curved, 2.0 times longer than wide, ventral surface with a pronounced indentation.
Description Figure 7i-l. Antennae ochreous, shortly serrate in male, filiform in the female, flagellum adorned dorsally with white and buff scales, palpus white suffused with buff; eyes dark brown. Body colour and wing pattern similar in both sexes. Head, thorax and abdomen white buff with long white and brown hairs, eyes dark brown. Legs buff with brown scales, fore femur and tibia infuscate on inner surface. Forewings ochraceous buff suffused with dark ochraceous and brown scales, more heavily on the costa, apex and termen. A longitudinal grey ochraceous fascia from base along lower margin of cell, partly within, partly without the cell, limited to the first two thirds of the wing; some few grey ochraceous markings around the cell; reniform not visible; outer margin adorned with dark brown spots between the veins, fringe infuscate with a narrow basal pale buff line. Hind wings white without markings, fringe whitish iridescent with a pale buff basal line. Underside of forewings with ground colour pale buff heavily suffused with fuscous and dark buff on costa, apex and along the cell, more heavily in male than in female. Underside of hind wings white, costa slightly suffused with fuscous scales, fringe white, slightly infuscate in apex, with a fuscous basal line. Forewings length: males 16.1 mm (min-max 15-19 mm, N = 13); females 19.2 mm (min-max 17-22 mm, N = 6). Male genitalia (Figure 8c, g). Tegumen with small flat peniculi; vinculum narrow with a small rectangular saccus, w-shaped at the top margin; valve with sacculus and cucullus fused; costa short, costal spine large, internal side flat, apex rather rectangular upwardly ending in a point, produced downwardly into a sharp pointed process strongly sclerotized; sacculus broad, well sclerotized with a strong narrowing below a slightly flared apex, inner side with a weak and only outlined harpe, heavily sclerotized; cucullus not sclerotized, broadly ovoid, in the prolongation of the sacculus, with scattered and papillated hairs; juxta bilobate, each lobe diamond-shaped; uncus angled and stout at base, tapering near apex to a fine point, tufted with hair on upper side; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose, vesica with a small tongue-shaped flat cornutus villi like. Female genitalia (Figure 8k). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium large, posterior margin slightly convex, slightly sclerotized, anterior margin bilobate with a convex area between them, each lobe cup-shaped, with sharp curved ends on external side strongly sclerotized; ductus bursae very short, slightly sclerotized on the ostium side; corpus bursae short, ovoid, without signa; ovipositor lobes curved, 2.0 times longer than wide, ventral surface with a pronounced indentation, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores without spatulate tips.

Etymology
Named after Kikuyu, the name of the tribe of Central Kenya where the holotype has been collected; specific epithet treated as an adjective.

Bionomics
One larva was collected in stems of Rottboellia cochinchinensis in a grassland in Zanzibar-Inhambane vegetation mosaic at the bottom of Shimba Hills inhabited with various Poales species belonging to the following genera: Cymbopogon, Hyparrhenia, Megathyrsus and Panicum. Otherwise adults were caught with light traps in the same habitat as the larva and in a grassland of Central Kenya, north of Nairobi, inhabited with various Poales species belonging to the following genera: Hyparrhenia, Megathyrsus, Panicum, Setaria P. Beauv., 1812 and Sporobolus.

Distribution
Kenya. The recorded localities are from two different vegetation mosaics: East African coastal (mosaic #16a) and East African evergreen bushland and secondary Acacia wooded grassland (mosaic #45) (White 1983) (Figure 5).

Remarks
The couple collected by D. Sevastopulu in 1956 in Mombasa on the Kenyan Coast was studied by Nye and provisionally named Sesamia nov. sp. "marina"; Nye considered it as a species related to S. rufescens, which our study confirms. Hampson, 1910 (

Diagnosis
See also the identification key to species of the cretica subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: tegumen with flat peniculi; vinculum with a saccus ushaped at the bottom margin; internal side of costal spine swollen at base, apex expanded upwardly in the shape of a small spoon irregularly edged with teeth, a strong hooked clasper below the costal spine; apex of the sacculus slightly flared, inner side with weak and only outlined harpe, 4.0 times longer than wide; apex of the cucullus rounded; juxta with a small elongated and blunt medial projection; posterior margin of ostium bursae convex in the middle with lateral expansion arm-shaped raised backwards, anterior margin thin and concave with villi like tip on each side; ovipositor lobes curved, 2.2 times longer than wide, ventral face strongly curved towards the apex.
Description Figure 7m-r. Both sexes of the species have been accurately described by Tams & Bowden (1953). Forewings length: males 28.0 mm (min-max 26-32 mm, N = 9); female 28.0 mm (N = 1). Male genitalia (Figure 8d, h). Tegumen with small flat peniculi; vinculum narrow with a small saccus, u-shaped at the bottom margin, w-shaped at the top margin with an indentation; valve with sacculus and cucullus fused; costa short, apex expanded upwardly in the shape of a small spoon irregularly edged with teeth, a strong hooked clasper below the costal spine; sacculus broad, well sclerotized with a strong narrowing below a slightly flared apex, inner side with a weak and only outlined harpe, slightly curved, heavily sclerotized, 4.0 times longer than wide; cucullus lying at a flatter angle on the sacculus, base bead-shaped, slightly sclerotized, apex rounded, not sclerotized, with scattered and papillated hairs; juxta trapezoidal, with a small elongated and blunt medial projection; uncus thin, curved at middle, tapering near apex to a fine curved point, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose, two slightly curved elongated scobinate lobes. Female genitalia (Figure 8l). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae small, posterior margin, strongly sclerotized, with lateral expansion armshaped raised backwards, anterior margin thin and concave with villi like tip on each side; ductus bursae short, slightly sclerotized on ostium side; corpus bursae short, ovoid, without signa; ovipositor lobes curved, 2.2 times longer than wide, ventral face strongly curved towards the apex, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores.

Remarks
In addition to the specimens listed in the other material section, 16 male and female specimens are preserved in the TMSA. These specimens were collected from four localities in South Africa (Eastern Cape: Kleinmonde, Umtata; Kwazulu-Natal: New Hanover, Humgababa).
Identification key to species of Sesamia cretica subgroup Key based on internal morphology of the adult male (A) and female (B) genitalia. The Sesamia fuscifrontia subgroup consists of S. babati n. sp., S. babessi n. sp., S. fuscifrontia, S. geyri, S. mabira n. sp., S. nangaensis n. sp., S. rungwa n. sp., S. simillima n. sp., S. taveta n. sp. and S. ulaukae n. sp.; it is characterized by the following combination of characters: (i) tegumen with small drooping peniculi, vinculum with a small saccus; (ii) valve with sacculus and cucullus fused; costa short, costal spine broad, the internal side less sclerotized than the external side, spoon-shaped or rounded upwardly, produced downwardly into a sharp process; sacculus broad, well sclerotized, with a strong narrowing below a more or less flared apex; cucullus small, not sclerotized, broadly rounded, in the extension of the sacculus or lying at a flatter angle on the sacculus with scattered and papillated hairs; (iii) juxta trapezoidal, with a small laterally compressed medial projection; (iv) uncus stout, angled at base, apex tapering to a fine point or blunt, tufted with long hair on upper side; (v) aedeagus short and stout; manica with a carina crest strongly spinose; (vi) ostium bursae large, posterior margin like a bead not or slightly sclerotized, anterior margin bilobate, each lobe bean-shaped or ribbon-shaped, sclerotized or villi-shaped, more or less sclerotized.

Diagnosis
See also the identification key to species of the fuscifrontia subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male genitalia: vinculum narrow with a small saccus, v-shaped at the bottom margin, wshaped at the top margin without indentation; costal spine strongly curved inwards, apex with two pointed teeth; sacculus with a strong narrowing below a narrow apex; cucullus small, narrow, lying at a flatter angle on the sacculus, base bead-shaped, apex rounded; juxta trapezoidal, with a small, robust and thick, laterally compressed mediodorsal projection, slightly dentated at apex; uncus blunt at apex. Figure 9a, b. Antennae buff suffused with brown, shortly serrate, flagellum adorned dorsally with brown scales; palpus pale buff; eyes brown. Head and thorax covered with long ochraceous hairs, abdomen pale buff suffused with brown. Forelegs brown, otherwise pale buff suffused with fuscous. Forewings buff suffused with brown from base to medial areas, termen heavily suffused with dark ochraceous; a dark brown marking below the cell, a line of brown markings along the upper margin of the cell; orbicular spot indistinct, reniform spot visible; a longitudinal dark buff fascia between lower and upper margins of cell, from base of cell to the termen; one postmedial and one subterminal series of brown markings on the veins; outer margin adorned with brown spots between the veins; fringe concolour with a basal ochraceous line. Hind wings white at base, pale buff at termen, suffused with brown, four elongated fuscous markings on the veins connected by a diffuse zigzagging fuscous line, outer margin adorned with brown spots between the veins, fringe light buff suffused with fuscous, with a basal white line. Underside of forewings fuscous around the cell, otherwise pale buff suffused with brown, reniform spot visible, outer margin adorned with brown spots between the veins; fringe concolour at the base otherwise fuscous. Underside of hind wings white, suffused with fuscous in costa, apex and termen areas, discal spot visible, outer margin adorned with brown spots between the veins, fringe white at the base, otherwise pale buff. Forewings length: males 24.0 mm (N = 2). Male genitalia (Figure 10a, b, m). Tegumen with flat drooping peniculi; vinculum narrow with a small saccus, v-shaped at the bottom margin, w-shaped at the top margin without indentation. Valve with sacculus and cucullus fused; costa short, costal spine large strongly curved inwards, apex with two pointed teeth; sacculus broad, well sclerotized with a strong narrowing below a narrow apex; cucullus small, narrow, lying at a flatter angle on the sacculus, base bead-shaped, apex rounded, not sclerotized, with scattered and papillated hairs; juxta trapezoidal, with a small, robust and thick, laterally compressed mediodorsal projection, slightly dentated at apex; uncus stout and short, curved at middle, blunt at apex, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest strongly spinose.

Etymology
Named after the name of Babati town in Manyara province in Tanzania; treated as a noun in apposition.
Sesamia babati n. sp. is a markedly hygrophilous species inhabiting lakesides. Larvae were collected on young stems of antelope grass Echinochloa pyramidalis; typically, stems exhibiting signs of infestation by S. babati n. sp. larvae have dry leaves and shoots (dead hearts).

Diagnosis
See also the identification key to species of the fuscifrontia subgroup below. This species can be distinguished  from other known species of the subgroup by the following combination of characters of the male genitalia: vinculum with a small saccus, u-shaped at the bottom margin, flat at the top margin without indentation; internal side of the costal spine convex in the middle, apex ampulla-like, rounded, slightly triangular upwardly, markedly triangular downwardly without tooth at the base; sacculus with a strong narrowing below a slightly flared apex; cucullus narrow lying at a flatter angle on the sacculus, base bead-shaped, apex rounded; juxta trapezoidal, with a small, robust and thick, laterally compressed mediodorsal projection, slightly dentated at apex; uncus blunt at apex.

Description
Figure 9c, d. The wings of the unique specimen of this species are heavily rubbed, also the wing pattern cannot be described precisely. Antennae dark buff shortly serrate, flagellum adorned dorsally with dark buff scales; palpus dark buff; eyes brown. Head and thorax covered with long dark buff hairs, abdomen dark buff. Legs brown suffused with dark buff. Forewings uniformly fuscous; fringe concolour with a basal white line. Hind wings suffused with fuscous more heavily in costa, apex and termen areas; fringe dark buff with a basal white line. Underside of forewings and hind wings uniformly suffused with fuscous, more heavily in costa and apex areas, fringe concolour. Forewings length: male 19.0 mm (N = 1). Male genitalia (Figure 10c, n). Tegumen with flat drooping peniculi; vinculum with a small saccus, u-shaped at the bottom margin, flat at the top margin without indentation. Valve with sacculus and cucullus fused; costa short, costal spine large, internal side convex in the middle, less sclerotized than the external side, apex ampulla-like, rounded, slightly triangular upwardly, markedly triangular downwardly; sacculus broad, well sclerotized with a strong narrowing below a slightly flared apex; cucullus narrow, lying at a flatter angle on the sacculus, base bead-shaped, apex rounded, not sclerotized, with scattered and papillated hairs; juxta trapezoidal, with a small, robust and thick, laterally compressed mediodorsal projection, slightly dentated at apex; uncus stout and short, curved at middle, blunt at apex, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest strongly spinose.

Etymology
Named after Babessi, the name of the village where the species has been collected; treated as a noun in apposition.

Bionomics
Biology unknown. The moths were caught with a light trap in a grassland inhabited with various Poales species belonging to the following genera: Andropogon, Cymbopogon, Hyparrhenia, Megathyrsus, Panicum and Sporobolus.

Diagnosis
See also the identification key to species of the fuscifrontia subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: vinculum with a small saccus, u-shaped at the bottom margin, flat at the top margin without indentation; costal spine convex inwards, apex with two pointed teeth at the tip; sacculus with a strong narrowing below a narrow apex; cucullus narrow, lying at a flatter angle on the sacculus, apex rounded; uncus pointed at apex; posterior margin of ostium bursae convex, sclerotized on each side, anterior margin bilobate in the middle, each lobe bean-shaped with a curved pointed end, strongly sclerotized, sides curved, slightly sclerotized; ductus bursae very short, slightly sclerotized; corpus bursae very short; ovipositor lobes short and curved, 2.0 times longer than wide, ventral surface with a pronounced indentation.

Redescription
Figure 9e-j. The species was originally described succinctly from the male holotype by Hampson (1914 (Figure 10d, e). Tegumen with flat drooping peniculi; vinculum with a small saccus, u-shaped at the bottom margin, flat at the top margin without indentation. Valve with sacculus and cucullus fused; costa short, costal spine large, convex inwards, apex with two pointed teeth at the tip; sacculus broad, well sclerotized with a strong narrowing below a narrow apex; cucullus narrow, lying at a flatter angle on the sacculus, base bead-shaped, apex rounded, not sclerotized, with scattered and papillated hairs; juxta not seen; uncus stout and short, curved at middle, pointed at apex, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest strongly spinose. Female genitalia (Figure 11a). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin convex, sclerotized on each side, anterior margin bilobate in the middle, each lobe bean-shaped with a curved pointed end, strongly sclerotized, sides curved, slightly sclerotized; ductus bursae very short, slightly sclerotized; corpus bursae very short, ovoid, without signa; ovipositor lobes short and curved, 2.0 times longer than wide, ventral surface with a pronounced indentation, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores without spatulate tips.

Bionomics
Biology unknown. The female was caught with a light trap in grasslands surrounding banks of marshes and wetlands inhabited with various Poales species belonging to the following genera: Cymbopogon, Cyperus L., 1753, Echinochloa, Hyparrhenia and Sporobolus.

Remarks
Without molecular data for the two male specimens preserved in the NHM it is not possible to determine whether the female from Keundwe belongs to S. fuscifrontia species; however, the wing pattern and the ecological characteristics suggests it could be the female. The specimen from Keundwe could have been described as a new species, but in a conservative way it was decided to consider that this specimen belongs to S. fuscifrontia.

Diagnosis
See also the identification key to species of the fuscifrontia subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: vinculum narrow with a small saccus, v-shaped at the bottom margin, flat at the top margin without indentation; costal spine large, strongly curved inwards, apex anvil-shaped with a sharp tooth at the tip; sacculus with a strong narrowing below a flared apex; cucullus small, narrow, lying at a flatter angle on the sacculus, base bead-shaped, apex rounded; uncus blunt at apex; anterior margin of ostium bursae bilobate in the middle, each lobe bean-shaped with a curved blunt end, strongly sclerotized; ovipositor lobes short and curved, 2.0 times longer than wide, ventral surface with a pronounced indentation.

Redescription
Figure 9k-p. The species was originally described from the male holotype by Strand (1915). This species is redescribed based on the additional male and female specimens collected in Uganda. The general shape of the female's forewings is more elongated at the apex than those of the male. Antennae ochraceous, shortly serrate in the male, filiform in the female, flagellum adorned dorsally with pale buff scales; palpus pale buff; eyes brown. Head covered with long ochraceous hairs, thorax covered with long pale buff hairs, abdomen pale buff slightly suffused with fuscous. Legs pale buff suffused with fuscous. Wing pattern similar in both sexes, but male darker. Forewings buff suffused with fuscous; a dark brown marking below the cell, two brown markings along the upper margin of the cell; orbicular spot and reniform indistinct; a longitudinal ochre brown fascia, between lower and upper margins of cell, from base of cell to the termen; one postmedial and one subterminal series of brown markings, more or less visible, on the veins; outer margin adorned with brown spots between the veins; fringe ochre brown. Hind wings white slightly suffused with fuscous, four elongated fuscous markings on the veins, outer margin adorned with brown spots between the veins, fringe white slightly suffused with fuscous. Underside of forewings ochre brown around the cell in male, otherwise pale buff suffused with fuscous, reniform spot visible, outer margin adorned with brown spots between the veins, fringe ochre brown. Underside of hind wings pale buff suffused with dark fuscous around the cell up to the apex and termen in male, white slightly suffused with fuscous in costa and apex areas in female, outer margin adorned with brown spots between the veins, fringe white slightly suffused with fuscous in female, pale buff suffused more markedly with fuscous in male. Forewings length: male 27.0 mm (N = 1); male 28.0 mm (N = 1). Male genitalia (Figure 10f, o). Tegumen with flat drooping peniculi; vinculum narrow with a small saccus, vshaped at the bottom margin, flat at the top margin without indentation. Valve with sacculus and cucullus fused; costa short, costal spine large strongly curved inwards, apex anvil-shaped with a sharp tooth at the tip; sacculus broad, well sclerotized with a strong narrowing below a flared apex; cucullus small, narrow, lying at a flatter angle on the sacculus, base bead-shaped, apex rounded, not sclerotized, with scattered and papillated hairs; juxta not observed; uncus stout and short, curved at middle, blunt at apex, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest strongly spinose. Female genitalia (Figure 11b). The genitalia preparation is of poor quality, as all the parts could not be observed. Ostium bursae large, posterior margin not seen, anterior margin bilobate in the middle, each lobe bean-shaped with a curved blunt end, strongly sclerotized; ductus bursae and corpus bursae not seen; ovipositor lobes short and curved, 2.0 times longer than wide, ventral surface with a pronounced indentation, dorsal surface bearing short and stout setae.

Bionomics
The larvae were collected in stems of Echinochloa pyramidalis and Megathyrsus maximus.

Distribution
Republic of the Sudan and Uganda. Recorded from two localities only from Guineo-Congolian lowland rain forest and secondary grassland (mosaic #11a) and semi-desert grassland and shrubland northern Sahel (mosaic #54a) vegetation mosaics (White 1983) (Figure 12).

Remarks
Surprisingly the two recorded host plants have quite different ecologies: Megathyrsus maximus (commonly known as Guinea grass) is an invasive grass mostly found along roads in rather dry habitats whereas Echinochloa pyramidalis (commonly known as antelope grass) is typical of flooded regions and frequently found floating on the surface of water beside lakes.

Diagnosis
See also the identification key to species of the fuscifrontia subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: vinculum with a small saccus, u-shaped at the bottom margin, flat at the top margin without indentation; internal side of the costal spine convex in the middle, apex ampulla-like, slightly triangular upwardly, produced downwardly into a large and sharp triangular process, strongly sclerotized; sacculus with a slightly flared apex; cucullus narrow lying at a flatter angle on the sacculus, base bead-shaped, apex rounded; juxta trapezoidal, with a small, robust and thick, laterally compressed mediodorsal projection, slightly dentated at apex; uncus pointed at apex; posterior margin of ostium bursae like a domed bead, more sclerotized on each side, anterior margin villi like, x-shaped, slightly sclerotized, slightly flared on external side; ductus bursae short; corpus bursae very short, ovoid, without signa; ovipositor lobes short and curved, 2.0 times longer than wide, ventral surface with an indentation.
Description Figure 13a-d. The general shape of the female's forewings is more elongated at the apex than those of the male. Antennae ochraceous, shortly serrate in the male, filiform in the female, flagellum adorned dorsally with pale buff scales; palpus pale buff. Head covered with long ochraceous hairs, thorax with pale ochraceous hairs, abdomen pale buff suffused with fuscous. Forelegs brown, otherwise pale buff suffused with fuscous. Wing pattern similar in both sexes, but markings more pronounced in the male. Forewings pale buff suffused with fuscous; a dark brown marking below the cell at base; orbicular spot indistinct, reniform spot visible; a longitudinal fuscous fascia between lower and upper margins of cell, from base of cell to the termen; one postmedial and one subterminal series of brown markings, more or less visible, on the veins; outer margin adorned with small brown spots between the veins; fringe pale buff suffused with fuscous. Hind wings white slightly suffused with fuscous in costa, apex and termen areas, four elongated fuscous markings on the veins, outer margin adorned with more or less visible brown spots between the veins, fringe white slightly suffused with buff. Underside of forewings buff suffused with fuscous around the cell in male, buff suffused with ochraceous and fuscous along the costa in female, outer margin adorned with brown spots between the veins, fringe pale buff suffused with fuscous. Underside of hind wings white suffused with fuscous, four elongated fuscous markings on the veins in male, discal spot visible in male, outer margin adorned with brown spots between the veins, fringe white slightly suffused with buff. Forewings length: males 24.1 mm (minmax 22-26 mm, N = 7); females 27.3 mm (min-max 26-30 mm, N = 11). Male genitalia (Figure 10g, p). Tegumen with flat drooping peniculi; vinculum with a small saccus, u-shaped at the bottom margin, flat at the top margin without indentation. Valve with sacculus and cucullus fused; costa short, costal spine large, internal side convex in the middle, apex ampulla-like, slightly triangular upwardly, produced downwardly into a large and sharp triangular process, strongly sclerotized; sacculus broad, well sclerotized with a strong narrowing below a slightly flared apex; cucullus narrow lying at a flatter angle on the sacculus, base bead-shaped, apex rounded, not sclerotized, with scattered and papillated hairs; juxta trapezoidal, with a small, robust and thick, laterally compressed mediodorsal projection, slightly dentated at apex; uncus stout and short, curved at middle, slightly pointed at apex, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest strongly spinose. Female genitalia (Figure 11c). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin like a domed bead, more sclerotized on each side, anterior margin villi like, x-shaped, slightly sclerotized, slightly flared on external side; ductus bursae short, slightly sclerotized; corpus bursae very short, ovoid, without signa; ovipositor lobes short and curved, 2.0 times longer than wide, ventral surface with an indentation, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores without spatulate tips.

Etymology
Named after Mabira Forest, the forest where this species was collected for the first time; treated as a noun in apposition.

Bionomics
Larvae were collected on young stems and shoots of Echinochloa pyramidalis growing in wetland inhabited with various Poales species belonging to the following genera: Cyperus, Eriochloa, Typha L., 1753 and Urochloa P. Beauv. 1812.

Diagnosis
See also the identification key to species of the fuscifrontia subgroup below. This species can be distinguished from all other known species of the subgroup by the following combination of characters of the male genitalia: vinculum with a small saccus, v-shaped at the bottom margin, wshaped at the top margin with an indentation; internal side of costal spine slightly concave with a bulge at the base, apex ampulla-like upwards with some kind of tooth in the middle, flattened triangle shape downwards; sacculus with a strong narrowing below a flared apex; cucullus small, broadly rounded, lying at a flatter angle on the sacculus; juxta trapezoidal, with a small laterally compressed medial projection, dentated at apex; uncus tapering near apex to a fine curved point.

Description
Figure 13e, f. Antennae dark ochraceous, shortly serrate, flagellum adorned dorsally with fuscous scales; palpus fuscous; eyes brown. Head and thorax covered with long fuscous hairs, abdomen pale buff suffused with fuscous. Forelegs fuscous, otherwise pale buff. Forewings dark ochraceous suffused with fuscous in costa, apex and termen areas, irrorated with brown along the outer margin; no transverse markings; orbicular spot and reniform not visible; a longitudinal fuscous fascia, between lower and upper margins of cell, from base of cell to the termen; fringe brown. Hind wings pale buff suffused with fuscous, irrorated with dark ochraceous along outer margin, veins streaked with dark ochraceous, fringe concolour with a basal pale buff line. Underside of forewings buff suffused with fuscous, irrorated with brown along outer margin, fringe brown. Underside of hind wings pale buff, suffused with fuscous, fringe pale buff suffused with fuscous, a basal pale buff line. Forewings length: males 17.5 mm (min-max 16-19 mm, N = 9). Male genitalia (Figure 10g, q). Tegumen with small flat peniculi; vinculum small with a small saccus, v-shaped at the bottom margin, w-shaped at the top margin with an indentation; valve with sacculus and cucullus fused; costa short, costal spine large, internal side slightly concave with a bulge at the base, apex ampulla-like upwards with some kind of tooth in the middle, weakly sclerotized, flattened triangle shape downwards, strongly sclerotized; sacculus broad, well sclerotized with a strong narrowing below a flared apex, a weak and only outlined harpe; cucullus small, not sclerotized, broadly rounded, lying at a flatter angle on the sacculus, with scattered and papillated hairs; juxta trapezoidal, with a small laterally compressed medial projection, dentated at apex; uncus angled and stout at base, narrowing in distal part, tapering near apex to a fine curved point, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest strongly spinose. Figure 13. Adults of the Sesamia fuscifrontia species subgroup. a-d, S. mabira: a, b, male (a, upper side; b, underside); c, d, female (c, upper side; d, underside); e, f, S. nangaensis, male: e, upper side; f, underside; g, h, S. rungwa, female: g, upper side; h, underside; i-l, S. simillima: i, j, male (i, upper side; j, underside); k, l, female (k, upper side; l, underside); m-p, S. taveta: m, n, male (m, upper side; n, underside); o, p, female (o, upper side; p, underside); q-t, S. ulaukae: q, r, male (q, upper side; r, underside); s, t, female (s, upper side; t, underside). Scale bars: 8 mm.

Etymology
Named after Lake Nanga, the lake shore where this species was collected for the first time; specific epithet treated as an adjective.

Bionomics
Biology unknown. The moths were caught with a light trap in grasslands surrounding banks of marshes and wetlands inhabited with various Poales species belonging to the following genera: Cymbopogon, Cyperus, Echinochloa, Hyparrhenia, Megathyrsus, Sporobolus and Typha.

Distribution
Cameroon and Republic of Congo. The two recorded localities are from Guineo-Congolian lowland rain forest and secondary grassland (mosaic #11a) vegetation mosaic (White 1983) (Figure 12).

Diagnosis
See also the identification key to species of the fuscifrontia subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the female genitalia: ostium bursae large, posterior margin like a domed bead, slightly sclerotized, anterior margin bilobate, each lobe in the shape of a large curved ribbon slightly flared towards the median part bead-shaped, slightly sclerotized; ductus bursae short; corpus bursae very short, ovoid; ovipositor lobes short and curved, 2.0 times longer than wide.
Description Figure 13g, h. Antennae pinkish-buff suffused with fuscous, filiform, flagellum adorned dorsally with pinkish-buff scales; palpus light ochraceous; eyes brown. Head and thorax covered with long pinkish-buff hairs, abdomen pinkish-buff suffused with fuscous. Legs light ochraceous suffused with fuscous. Forewings pinkish-buff suffused with fuscous; no transverse markings, a dark brown marking below the cell at base; orbicular and reniform spots not visible; one postmedial and one subterminal series of brown markings on the veins; outer margin adorned with brown spots between the veins; fringe fuscous with a basal buff line. Hind wings white, four elongated light fuscous markings on the veins, fringe white, with a basal pinkish-buff line. Underside of forewings buff suffused with fuscous, apex of the cell with two fuscous areas, outer margin adorned with brown spots between the veins; fringe fuscous. Underside of hind wings white, suffused with fuscous in costa and apex areas, four barely visible elongated light fuscous markings on the veins, fringe white. Forewings length: female 20.0 mm (N = 1). Female genitalia (Figure 11d). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin like a domed bead, slightly sclerotized, anterior margin bilobate, each lobe in the shape of a large curved ribbon slightly flared towards the median part bead-shaped, slightly sclerotized; ductus bursae short, slightly sclerotized; corpus bursae very short, ovoid, without signa; ovipositor lobes short and curved, 2.0 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores without spatulate tips.

Etymology
Named after Rungwa, a village on the main road between Singida and Mbeya, in Singida Region; treated as a noun in apposition.

Bionomics
Biology unknown. The only known specimen was caught with a light trap in grasslands surrounding banks of wetlands inhabited with various Poales species belonging to the following genera: Cymbopogon, Cyperus, Echinochloa, Hyparrhenia and Sporobolus.

Diagnosis
See also the identification key to species of the fuscifrontia subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: vinculum with a small saccus, u-shaped at the bottom margin, w-shaped at the top margin without indentation; internal side of costal spine w-shaped, apex ampulla-like, rounded upwardly, with a sharp triangular process downwardly and a small tooth on the top; cucullus narrow, lying at a flatter angle on the sacculus, base bead-shaped, apex rounded, not sclerotized; juxta trapezoidal, with a small, robust and thick, laterally compressed mediodorsal projection, slightly dentated at apex; uncus truncate at apex; posterior margin of ostium bursae like a domed bead, more sclerotized on each side, anterior margin villi like, shaped like two opposing triangles with rounded angles, base of triangle shorter than height, slightly sclerotized; ovipositor lobes 2.0 times longer than wide.

Description
Figure 13i-l. The general shape of the female's forewings is more elongated at the apex than those of the male. Antennae ochraceous, shortly serrate in the male, filiform in the female, flagellum adorned dorsally with pale buff scales; palpus fuscous; eyes brown. Head and thorax covered with long ochraceous or pale buff hairs, abdomen pale buff suffused with fuscous. Legs fuscous suffused with brown. Wing pattern similar in both sexes. Forewings buff more or less suffused with ochraceous and fuscous; a dark brown marking below the cell; orbicular and reniform spots visible; a longitudinal ochraceous fascia between lower and upper margins of cell, from base of cell to the termen; one postmedial and one subterminal series of brown markings on the veins; outer margin adorned with brown spots between the veins; fringe fuscous with a basal buff line. Hind wings white suffused with fuscous in costa and apex areas, four elongated fuscous markings on the veins, outer margin adorned with brown spots between the veins, fringe white suffused with fuscous, with a basal white line. Underside of forewings pale buff suffused with dark fuscous around the cell up to the apex and termen in male, pale buff slightly suffused with fuscous in costa, apex and termen areas in female, orbicular spot more or less visible, outer margin adorned with brown spots between the veins, fringe fuscous. Underside of hind wings white, suffused with fuscous in costa, apex and termen areas, four elongated fuscous markings on the veins, discal spot visible, outer margin adorned with brown spots between the veins, fringe white suffused with pale buff or fuscous. Forewings length: males 23.6 mm (min-max 20-26 mm, N = 12); females 27.3 mm (min-max 23-30 mm, N = 12). Male genitalia (Figure 10i, j, r). Tegumen with flat drooping peniculi; vinculum with a small saccus, u-shaped at the bottom margin, w-shaped at the top margin without indentation. Valve with sacculus and cucullus fused; costa short, costal spine large, internal side w-shaped less sclerotized than the external side, apex ampulla-like, rounded upwardly, with a sharp triangular process downwardly and a small tooth on the top; sacculus broad, well sclerotized with a strong narrowing below a flared apex, inner side with a weak and only outlined harpe, 4.5 times longer than wide; cucullus narrow lying at a flatter angle on the sacculus, base bead-shaped, apex rounded, not sclerotized, with scattered and papillated hairs; juxta trapezoidal, with a small, robust and thick, laterally compressed mediodorsal projection, slightly dentated at apex; uncus stout and short, curved at middle, truncate at apex, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest strongly spinose. Female genitalia (Figure 11e). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin like a domed bead, more sclerotized on each side, anterior margin villi like, shaped like two opposing triangles with rounded angles, base of triangle shorter than height, slightly sclerotized; ductus bursae very short, slightly sclerotized; corpus bursae very short, ovoid, without signa; ovipositor lobes short and curved, 2.0 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores without spatulate tips. Larvae -L5 instar (Figure 4c). Length, 30-35 mm, width, 4.0 mm; head smooth, brown, prothoracic shield light brown; body with ground colour salmon pink suffused with dark pink, pinnacula and caudal plate brown. Young larvae are very similar in appearance to mature ones.

Etymology
Named simillima, nominative feminine singular of simillimus, because it was first expected to be the same species as Sesamia echinochloa; treated as an adjective.

Diagnosis
See also the identification key to species of the fuscifrontia subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: vinculum with a small saccus, u-shaped at the bottom margin, w-shaped at the top margin without indentation; internal side of costal spine swollen at the base, apex ampulla-like, rounded upwardly, with a sharp triangular process downwardly without tooth at the base; cucullus narrow, lying at a flatter angle on the sacculus, apex rounded, not sclerotized; juxta trapezoidal, with a small, robust and thick, laterally compressed mediodorsal projection, slightly dentated at apex; uncus stout and short; posterior margin of ostium bursae like a domed bead, more sclerotized on each side, anterior margin villi like, shaped like two opposing triangles, base of triangle longer than height, slightly sclerotized; ovipositor lobes short and curved, 2.0 times longer than wide.

Description
Figure 13m-p. The general shape of the female's forewings is more elongated at the apex than those of the male. Antennae pale buff, shortly serrate in the male, filiform in the female, flagellum adorned dorsally with pale buff scales; palpus pale buff. Head and thorax covered with long pale ochraceous hairs, abdomen pale buff suffused with fuscous. Forelegs fuscous, otherwise pale buff suffused with fuscous. Wing pattern similar in both sexes, but male darker. Forewings buff, heavily suffused with fuscous and ochraceous in male, slightly suffused with fuscous and ochraceous in female; a brown marking below the cell at base; orbicular spot indistinct, reniform spot visible; a longitudinal fuscous fascia between lower and upper margins of cell, from base of cell to the termen; one postmedial and one subterminal series of brown markings, on the veins, the markings of the postmedial series are connected by a diffuse zigzagging fuscous line; outer margin adorned with brown spots between the veins; fringe fuscous. Hind wings white slightly suffused with fuscous in costa, apex and termen areas, four elongated fuscous markings on the veins, outer margin adorned with brown elongated spots between the veins, fringe white slightly suffused with fuscous. Underside of forewings pale buff suffused with dark fuscous around the cell up to the apex and termen in male, pale buff slightly suffused with fuscous in costa, apex and termen areas in female, outer margin adorned with brown spots between the veins, fringe fuscous. Underside of hind wings white, suffused with fuscous in costa, apex and termen areas, four elongated fuscous markings on the veins, outer margin adorned with brown spots between the veins, fringe white suffused with pale buff and fuscous. Forewings length: males 24.4 mm (min-max 23-26 mm, N = 5); females 27.0 mm (min-max 26-28 mm, N = 4). Male genitalia (Figure 10k, s). Tegumen with flat drooping peniculi; vinculum with a small saccus, u-shaped at the bottom margin, w-shaped at the top margin without indentation. Valve with sacculus and cucullus fused; costa short, costal spine large, internal side swollen at the base, less sclerotized than the external side, apex ampulla-like, rounded upwardly, with a sharp triangular process downwardly, heavily sclerotized; sacculus broad, well sclerotized with a strong narrowing below a slightly flared apex; cucullus narrow, lying at a flatter angle on the sacculus, apex rounded, not sclerotized, with scattered and papillated hairs; juxta trapezoidal, with a small, robust and thick, laterally compressed mediodorsal projection, slightly dentated at apex; uncus stout and short, curved at middle, slightly pointed at apex, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest strongly spinose. Female genitalia (Figure 11f). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin like a domed bead, more sclerotized on each side, anterior margin villi like, shaped like two opposing triangles, base of triangle longer than height, slightly sclerotized; ductus bursae very short, slightly sclerotized; corpus bursae very short, ovoid, without signa; ovipositor lobes short and curved, 2.0 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores without spatulate tips.

Etymology
Named after Taveta town in Eastern Region in Kenya, close to the Tanzanian border, at the bottom of Mount Kilimanjaro; treated as a noun in apposition.

Bionomics
Larvae were collected on young stems of Echinochloa pyramidalis. Sesamia taveta n. sp. is a markedly hygrophilous species found in wetlands inhabited by various Poales species belonging to the following genera: Cyperus, Eriochloa and Typha.

Remarks
The female genitalia of S. taveta n. sp. look very similar to those of S. simillima n. sp.; the main difference is the height of the two opposing triangles of the anterior margin of the anteostial pad, which is shorter in S. taveta n. sp.

Diagnosis
See also the identification key to species of the fuscifrontia subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: internal side of costal spine w-shaped, apex ampulla-like, rounded upwardly, with a sharp triangular process downwardly and a small tooth on the top, heavily sclerotized; cucullus narrow, lying at a flatter angle on the sacculus, base bead-shaped, apex ovoid, not sclerotized; carina crest with two lateral lobes with shorter and robust cornuti in ventral position; posterior margin of ostium bursae like a domed bead, anterior margin villi like, shaped like two opposing funnels, slightly sclerotized. Figure 13q-t. The general shape of the female's forewings is more elongated at the apex than those of the male. Antennae ochraceous, shortly serrate in the male, filiform in the female, flagellum adorned dorsally with pale buff scales; palpus fuscous; eyes brown. Head and thorax covered with long ochraceous hairs, abdomen pale buff suffused with fuscous. Forelegs fuscous, otherwise pale buff suffused with fuscous. Wing pattern similar in both sexes but female darker. Forewings buff in male, ochraceous in female suffused with fuscous, another one in the cell; a dark brown marking below the cell; orbicular not visible reniform spots visible; a longitudinal more or less visible ochraceous fascia, between lower and upper margins of cell, from base of cell to the termen; one postmedial and one subterminal series of brown markings, more or less visible, on the veins; outer margin adorned with brown spots between the veins; fringe fuscous. Hind wings white suffused with fuscous in costa, apex and termen areas, four elongated fuscous markings on the veins in male, outer margin adorned with brown spots between the veins, fringe white suffused with buff, with a basal white line. Underside of forewings pale buff suffused with dark fuscous around the cell up to the apex and termen in male, pale buff slightly suffused with fuscous in costa, apex and termen areas in female, outer margin adorned with brown spots between the veins, fringe fuscous. Underside of hind wings white, suffused with fuscous in costa, apex and termen areas, four elongated fuscous markings on the veins in male, outer margin adorned with brown spots between the veins, fringe white suffused with pale buff or fuscous. Forewings length: males 25.5 mm (min-max 25-26 mm, N = 2); female 30.0 mm (N = 1). Male genitalia (Figure 10l, t). Tegumen with flat drooping peniculi; vinculum with a small saccus, u-shaped at the bottom margin, w-shaped at the top margin without indentation. Valve with sacculus and cucullus fused; costa short, costal spine large, internal side w-shaped, less sclerotized than the external side, apex ampulla-like, rounded upwardly, with a sharp triangular process downwardly and a small tooth on the top, heavily sclerotized; sacculus broad, well sclerotized with a strong narrowing below a flared apex, inner side with a weak and only outlined harpe, 5.0 times longer than wide; cucullus narrow lying at a flatter angle on the sacculus, base bead-shaped, apex ovoid, not sclerotized, with scattered and papillated hairs; juxta trapezoidal, with a small, robust and thick, laterally compressed mediodorsal projection, slightly dentated at apex; uncus stout and short, curved at middle, pointed at apex, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose, two of them with shorter and robust cornuti in ventral position. Female genitalia (Figure 11g). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin like a domed bead, anterior margin villi like, shaped like two opposing funnels, slightly sclerotized; ductus bursae very short, slightly sclerotized; corpus bursae very short, ovoid, without signa; ovipositor lobes short and curved, 2.0 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores without spatulate tips.

Etymology
Named after Ula Ukae a village close to Jimma in Oromia Region in Ethiopia; treated as a noun in apposition.

Bionomics
Larvae were collected on young stems of Echinochloa pyramidalis. Sesamia ulaukae n. sp. is a markedly hygrophilous species found in wetlands inhabited with various Poales species belonging to the following genera: Cyperus, Eriochloa and Typha.
Identification key to species of Sesamia fuscifrontia subgroup Key based on internal morphology of the adult male (A) and female (B) genitalia. The Sesamia salama subgroup consists of S. salama n. sp.
The results of molecular phylogenetic analyses (see below) also indicate that this subgroup includes another new species (referred to as Sesamia "n. sp. 35") that cannot be described in the absence of genitalia.
It is characterized by the following combination of characters: (i) tegumen with small drooping peniculi, vinculum with a very small saccus; (ii) valve with sacculus and cucullus fused; costa short, costal spine broad, ending with a thick shoulder without inner point, sacculus broad, well sclerotized, with a strong narrowing below a slightly flared apex; cucullus small, as an extension of the sacculus, ovoid at apex; (iii) juxta very large, trapezoidal, without medial projection; (iv) aedeagus short and stout, manica with two elongated lobes strongly spinose at apex, vesica with two large and flat beak-shaped cornuti; (vi) ostium bursae large, posterior margin convex in the middle, barely sclerotized, spoon-shaped on each side with a pointed apex, heavily sclerotized, anterior margin cup-shaped, slightly sclerotized; ductus bursae long.

Diagnosis
This species can be distinguished from other known species of the S. cretica and S. fuscifrontia subgroups by the following combination of characters of the male and female genitalia: vinculum narrow with a small saccus, u-shaped at the bottom margin, u-shaped at the top margin with a pronounced indentation; valve with sacculus and cucullus fused; internal side of costal spine concave, apex with a thick shoulder without inner point; cucullus as an extension of the sacculus, ovoid at apex; juxta very large and trapezoidal, the base concave, the sides slightly pointed, the distal part convex; uncus tapering near apex to a fine curved point; manica with two elongated lobes strongly spinose at apex, vesica with two large beakshaped cornuti; posterior margin of ostium bursae convex in the middle, barely sclerotized, spoon-shaped heavily sclerotized on each side with a pointed apex, anterior margin cup-shaped, slightly sclerotized; ductus bursae long; ovipositor lobes curved, 2.7 times longer than wide, ventral surface with a pronounced indentation.
Description Figure 14a-d. The two known males are much lighter than the 14 known females; it is not possible to determine whether it is a gender difference, part of the colour range of the species or because the forewings are slightly rubbed. The general shape of the forewings is similar in both sexes. Antennae pale buff and shortly serrate in the male, ochraceous and filiform in the female, flagellum adorned dorsally with pale buff scales; palpus pale buff. Head and thorax covered with long pale buff to pale ochraceous hairs, abdomen pale buff heavily suffused with fuscous. Forelegs fuscous, otherwise pale buff suffused with fuscous. Wing pattern similar in both sexes. Forewings buff suffused with fuscous and brown; a dark brown marking below the cell at base; orbicular spot not visible, reniform clearly visible; a longitudinal fuscous fascia, between lower and upper margins of cell, from base of cell to the termen; one postmedial series of brown fuscous markings, one subterminal series of brown fuscous markings on the veins; veins streaked with white; tornus and dorsum adorned with a brown fuscous fascia; outer margin adorned with large brown spots between the veins; fringe with alternating brown and fuscous areas. Hind wings white; outer margin adorned with brown fuscous spots between the veins; fringe white slightly suffused with fuscous. Underside of forewings pale buff suffused with fuscous in costa, apex and termen areas; reniform visible, outer margin adorned with brown fuscous spots between the veins; fringe with alternating brown and fuscous areas. Underside of hind wings white, slightly suffused with fuscous in costa, apex and termen areas; discal spot visible; outer margin adorned with brown fuscous spots between the veins; fringe white slightly suffused with fuscous. Forewings length: male 22.0 mm (N = 1); females 22.6 mm (minmax 20-26 mm, N = 14). Male genitalia (Figure 14e, f). Tegumen with small flat peniculi; vinculum narrow with a small saccus, u-shaped at the bottom margin, u-shaped at the top margin with a pronounced indentation; valve with sacculus and cucullus fused; costa short, costal spine large, internal side concave, apex with a thick shoulder; sacculus broad, well sclerotized, with a strong narrowing below a slightly flared apex, inner side with a weak and only outlined harpe, heavily sclerotized; cucullus not sclerotized, as an extension of the sacculus, ovoid at apex, with scattered and papillated hairs; juxta very large and trapezoidal, the base concave, the sides slightly pointed, the distal part convex; uncus angled and stout at base, narrowing in distal part, tapering near apex to a fine curved point, tufted with long hair on upper side; aedeagus short and stout, manica with two elongated lobes strongly spinose at apex, vesica with two large and flat beakshaped cornuti.
Female genitalia (Figure 14g). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin convex in the middle, barely sclerotized, spoon-shaped on each side with a pointed apex, heavily sclerotized, anterior margin cup-shaped, slightly sclerotized; ductus bursae long, slightly sclerotized on the ostium side; corpus bursae ovoid, without signa; ovipositor lobes curved, 2.7 times longer than wide, dorsal surface bearing short and stout setae, ventral surface with a pronounced indentation; apophyses posteriores slenderer and longer than apophyses anteriores without spatulate tips.

Etymology
Named after Salama, the name of the town where this species was collected for the first time; treated as a noun in apposition.

Bionomics
Biology unknown. The moths were caught with light traps in grasslands inhabited with various Poales species belonging to the following genera: Andropogon, Cymbopogon, Hyparrhenia, Megathyrsus, Panicum and Sporobolus.

Distribution
Kenya. The recorded localities are from East African evergreen bushland and secondary Acacia wooded grassland (mosaic #45) vegetation mosaic (White 1983) (Figure 15).

Remarks
The SD analyses (see below) suggest that S. salama n. sp. specimens from western and eastern Kenya, on both sides of the Gregory Rift Valley, should belong to different species; however, examination of the corresponding specimens did not yield any robust morphological evidence to support the description of an additional taxa. The S. salama subgroup is also probably more diverse than it seems; indeed, the Pretoria museum is home to at least five undescribed Sesamia species which probably belong to this subgroup.  Rungs (1954) and Viette (1967) are presented.

Sesamia viettei subgroup
Our subset of interest (hereby referred to as the S. viettei subgroup) only consists of Sesamia viettei. It is characterized by the following combination of characters: (i) tegumen with small drooping peniculi, vinculum with a small saccus; (ii) valve with sacculus and cucullus fused; costa short, costal spine broad, the internal side less sclerotized than the external side, rounded upwardly, produced downwardly into a sharp process; sacculus broad, well sclerotized, with a large clasper with two strong sharp teeth; cucullus small, not sclerotized, broadly rounded, in the extension of the sacculus with scattered and papillated hairs; (iii) juxta large and trapezoidal, with a very small mediodorsal projection; (iv) uncus stout, angled at base, apex tapering to a fine point, tufted with long hair on upper side; (v) aedeagus short and stout; manica with a carina crest spinose; (vi) ostium bursae large, posterior margin with two lateral small rounded areas clearly punctuated, anterior margin in the form of a wavy bead, slightly sclerotized.

Diagnosis
This species can be distinguished from other known species of the cretica group by the following combination of characters of the male and female genitalia: internal side of costal spine strongly concave, apex ampulla-like, serrated upwardly, very sharply triangular downwardly, heavily sclerotized; inner side of the sacculus with a large clasper with two strong sharp teeth at right angle; cucullus lying at a flatter angle on the sacculus, base bead-shaped, apex ovoid, not sclerotized; juxta trapezoidal, with a very small mediodorsal projection, the distal part with a short wide neck, with pointed ends; uncus pointed at apex; ostium bursae large, posterior margin with two lateral rounded small areas clearly punctuated, anterior margin in the form of a wavy bead, slightly sclerotized.

Description
Figure 14h-m. The species was originally described from the male holotype by Rungs (1954). It has been redescribed by Viette (1967) with an additional female specimen. The general shape of the female's forewings is similar to that of the male. Forewings length: male 34 mm (N = 1); female 36.0 mm (N = 1).
Male genitalia (Figure 14n-p). The preparation of the genitalia could not be observed; our description is thus based on Rungs's pictures (1954) and Viette's drawing (1967). Tegumen with flat drooping peniculi; vinculum with a small saccus, u-shaped at the bottom margin, ushaped at the top margin without indentation. Valve with sacculus and cucullus fused; costa short, costal spine large, internal side strongly concave, apex ampulla-like, serrated upwardly, very sharply triangular downwardly, heavily sclerotized; sacculus broad, well sclerotized with a weak narrowing below a flared apex, inner side with a large clasper with two strong sharp teeth at right angle; cucullus lying at a flatter angle on the sacculus, base bead-shaped, apex ovoid, not sclerotized, with scattered and papillated hairs; juxta large and trapezoidal, with a very small mediodorsal projection, the base pointed, the sides rounded, the distal part with a short wide neck, with pointed ends; uncus stout and short, curved at middle, pointed at apex, tufted with long hair on upper side; aedeagus short and stout, curved in the middle, manica with a carina crest spinose. Female genitalia (Figure 14q). Apophyses anteriores with spatulate tips; lateral plates of ostial segment large, slightly sclerotized; ostium bursae large, posterior margin with two lateral small rounded areas clearly punctuated, anterior margin in the form of a wavy bead, villi like, slightly sclerotized; ductus bursae short, slightly sclerotized; corpus bursae short, subspheric, without signa; ovipositor lobes short and curved, 2.0 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores without spatulate tips.

Sesamia wiltshirei subgroup
The Sesamia wiltshirei subgroup consists of S. djenoensis n. sp., S. echinochloa n. sp., S. inexpectata n. sp., S. lefini n. sp., S. rindini n. sp., and S. wiltshirei; it is characterized by the following combination of characters: (i) tegumen with small drooping peniculi, vinculum with a small saccus; (ii) valve with sacculus and cucullus fused; costa short, sclerotized, without costal spine; clasper kidneyshaped parallel to the internal side of the valve, strongly toothed at the apex; sacculus broad, well sclerotized, with a strong narrowing below a more or less flared apex, ending with a narrow thickening or globular extension, toothed; cucullus small, sclerotized like a finger curved at apex with scattered and papillated hairs (iii) juxta heart-shaped or trapezoidal, with a small laterally compressed medial projection; (iv) uncus stout, angled at base, apex tapering to a fine point, tufted with long hair on upper side; (v) aedeagus short and stout; manica with a carina crest produced into paired lateral lobes strongly spinose, two of them with shorter and robust cornuti in ventral position; (vi) ostium bursae large, posterior margin like a great barely sclerotized dome, anterior margin bilobate, each lobe kidney or bean-shaped, more or less sclerotized.

Diagnosis
See also the identification key to species of the wiltshirei subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: costa with a marked bulge; clasper 1.5 times longer than wide; sacculus ending with a narrow thickening toothed; cucullus strongly curved at apex, in the shape of a bird's head; juxta trapezoidal, with a small, robust and thick, laterally compressed mediodorsal projection, slightly dentated at apex; carina crest with two lateral lobes strongly spinose and short and robust cornuti in ventral position; anterior margin of ostium bursae bilobate in the middle, each lobe kidney-shaped, strongly sclerotized, the anterior part 2.4 times wider than the posterior part, sides flared, slightly sclerotized; ovipositor lobes short and curved, 2.0 times longer than wide.
Description Figure 16a-d. The general shape of the female's forewings is more elongated at the apex than those of the male. Antennae pale buff, shortly serrate in the male, filiform in the female, flagellum adorned dorsally with pale buff scales; palpus pale buff; eyes brown. Head and thorax covered with long pale buff hairs, abdomen pale buff slightly suffused with fuscous. Legs pale buff slightly suffused with fuscous. Wing pattern similar in both sexes, but male darker. Forewings buff suffused with fuscous; a dark brown marking below the cell at base; orbicular and reniform spots indistinct; a longitudinal ochraceous fascia, between lower and upper margins of cell, from base of cell to the termen; one subterminal series of brown fuscous markings on the veins; outer margin adorned with barely visible brown spots between the veins; fringe dark buff with a basal pale buff line. Hind wings pale buff slightly pinkish, suffused with fuscous in costa and apex areas; fringe white with a basal buff line. Underside of forewings pale buff slightly suffused with fuscous in both sexes, areas around the cell up to the apex and termen fuscous in male; fringe dark buff. Underside of hind wings pale buff, slightly pinkish, suffused with fuscous in costa and apex areas; fringe white with a basal buff line. Forewings length: males 23.5 mm (min-max 23-24 mm, N = 2); female 25.0 mm (N = 2). Male genitalia (Figure 17a, b, i). Tegumen with flat drooping peniculi; vinculum narrow with a small saccus, u-shaped at the bottom margin, flat at the top margin without indentation. Valve with sacculus and cucullus fused; costa short, sclerotized, with a marked bulge, without costal spine; clasper kidney-shaped parallel to the internal side of the valve, 1.5 times longer than wide, strongly toothed at the apex; sacculus with a strong narrowing below a flared apex ending with a narrow thickening toothed; cucullus small, like a short and thick finger curved at the tip in the shape of a bird's head, with scattered and papillated hairs; juxta trapezoidal, with a small, robust and thick, laterally compressed mediodorsal projection, slightly dentated at apex; uncus stout and short, curved at middle, pointed at apex, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose, two of them with shorter and robust cornuti in ventral position. Female genitalia (Figure 17n). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin like a bell-shaped bead, not sclerotized, anterior margin bilobate in the middle, each lobe kidney-shaped, strongly sclerotized, the anterior part 2.4 times wider than the posterior part, sides flared, slightly sclerotized; ductus bursae very short, slightly sclerotized on ostium side; corpus bursae very short, ovoid, without signa; ovipositor lobes short and curved, 2.0 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores without spatulate tips.

Etymology
Epithet from the name of Djeno village in Kouilou department in Republic of Congo where this species was discovered; the specific epithet treated as an adjective.

Diagnosis
See also the identification key to species of the wiltshirei subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: internal side of costa almost flat; clasper kidneyshaped, 2.0 times longer than wide, sacculus ending with a globular extension toothed; cucullus daggershaped; juxta small and flat, trapezoidal; anterior margin of ostium bursae bilobate in the middle, each lobe beanshaped almost triangular with the posterior part wider than the anterior part, strongly sclerotized, sides flared, less sclerotized; ovipositor lobes short and curved, 2.0 times longer than wide.
Description Figure 16e-h. The general shape of the female's forewings is more elongated at the apex than those of the male. Antennae pale buff and shortly serrate in the male, ochraceous and filiform in the female, flagellum adorned dorsally with pale buff scales; palpus pale buff; eyes brown. Head and thorax covered with long pale ochraceous hairs, abdomen pale buff slightly suffused with fuscous. Legs pale buff slightly suffused with fuscous in male, ochraceous suffused with fuscous in female. Wing pattern similar in both sexes, but female darker. Forewings buff suffused with fuscous in male, buff suffused with ochraceous in female; a dark brown marking below the cell at base; orbicular spot not visible, reniform clearly visible; a longitudinal grey fuscous fascia, between lower and upper margins of cell, from base of cell to the termen; two postmedial brown markings more or less visible, one subterminal series of brown fuscous markings on the veins; outer margin adorned with brown spots between the veins; fringe fuscous with a basal pale buff line. Hind wings white slightly suffused with fuscous in costa, apex and termen areas; fringe white slightly suffused with fuscous, a basal pale buff line. Underside of forewings pale buff suffused with fuscous in male, ochraceous in female, areas around the cell up to the apex and termen fuscous in male, outer margin adorned with brown spots between the veins, fringe dark buff in male, dark ochraceous in female. Underside of hind wings white, suffused with fuscous in costa, apex and termen areas, suffusion is more intense in male, fringe white suffused with fuscous, a basal buff line. Forewings length: males 21.9 mm (min-max 18-24 mm, N = 12); females 25.7 mm (min-max 22-29 mm, N = 12). Male genitalia (Figure 17c, j). Tegumen with flat drooping peniculi; vinculum small with a small saccus, ushaped at the bottom margin, w-shaped at the top margin without indentation. Valve with sacculus and cucullus fused; costa short, sclerotized, internal side almost flat, without costal spine; clasper kidney-shaped parallel to the internal side of the valve, 2.0 times longer than wide, strongly toothed at apex; sacculus with a strong narrowing below a globular apex, toothed; cucullus dagger-shaped with scattered and papillated hairs; juxta small and flat trapezoidal, slightly sclerotized; uncus stout and short, curved at middle, blunt at apex, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose, two of them with shorter and robust cornuti in ventral position. Female genitalia (Figure 17o). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin like a bell-shaped bead not sclerotized, anterior margin bilobate in the middle, each lobe beanshaped, almost triangular with the posterior part wider than the anterior part, strongly sclerotized, sides flared, less sclerotized; ductus bursae very short, slightly sclerotized on ostium side; corpus bursae very short, ovoid, without signa; ovipositor lobes short and curved, 2.0 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores without spatulate tips.
Larvae -L5 instar (Figure 4d). Length, 30-35 mm, width, 4.0 mm; head smooth, orange brown, prothoracic shield salmon beige; body with ground colour salmon pink, pinnacula and caudal plate brown. Young larvae are very similar in appearance to mature ones.

Etymology
Named after Echinochloa, the genus name of its main host plant species; treated as a noun in apposition.

Bionomics
Larvae were collected on young stems and shoots of  Figure 15).

Remarks
This species was already treated by Janse (1939) with the nomen nudum of Sesamia perplexa, though no text at all was related to S. perplexa in Janse (1939). Since Tams & Bowden (1953) already described a Sesamia species with the name jansei it was not possible to describe this species as S. jansei.

Diagnosis
See also the identification key to species of the wiltshirei subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: costa short with a slight bulge; clasper 2.3 times longer than wide; sacculus ending with a narrow thickening toothed; cucullus small, like a short and thick finger curved at apex in the shape of a bird's head; anterior margin of the ostium bursae bilobate in the middle, each lobe kidney-shaped with the posterior part as wide as the anterior part, strongly sclerotized, sides flared, slightly sclerotized; ovipositor lobes short and curved, 2.0 times longer than wide.
Description Figure 16i-l. The general shape of the female's forewings is more elongated at the apex than those of the male. Antennae ochraceous, shortly serrate in the male, filiform in the female, flagellum adorned dorsally with pale buff scales; palpus pale buff. Head and thorax covered with long pale buff hairs in male, long ochraceous hairs in female, abdomen pale buff slightly suffused with fuscous. Forelegs brown, otherwise pale buff suffused with fuscous. Wing pattern similar in both sexes, but female darker. Forewings buff, slightly suffused with fuscous in male, buff heavily suffused with fuscous and ochraceous in female; a dark brown marking below the cell at base; orbicular spot indistinct, reniform spot visible; a longitudinal fuscous fascia, between lower and upper margins of cell, from base of cell to the termen; one postmedial and one subterminal series of brown markings, more or less visible, on the veins; outer margin adorned with brown spots between the veins; fringe fuscous in male, dark ochraceous in female. Hind wings white slightly suffused with fuscous in costa, apex and termen areas, fringe white slightly suffused with buff. Underside of forewings buff suffused with fuscous around the cell in male, buff suffused with ochraceous and fuscous in female, outer margin adorned with brown spots between the veins, fringe fuscous in male, ochraceous in female. Underside of hind wings white suffused with fuscous in costa, apex and termen areas, fringe white slightly suffused with buff. Forewings length: males 22.7 mm (min-max 22-23 mm, N = 4); females 23.4 mm (min-max 21-26 mm, N = 5).
Male genitalia (Figure 17d, e, k). Tegumen with flat drooping peniculi; vinculum narrow with a small saccus, u-shaped at the bottom margin, w-shaped at the top margin without indentation. Valve with sacculus and cucullus fused; costa short, sclerotized, with a slight bulge, without costal spine; clasper kidney-shaped parallel to the internal side of the valve, 2.3 times longer than wide, strongly toothed at the apex; sacculus with a strong narrowing below a slightly flared apex ending with a narrow thickening toothed; cucullus small, like a short and thick finger curved at the tip, in the shape of a bird's head; juxta trapezoidal, with a small, robust and thick, laterally compressed mediodorsal projection; uncus stout and short, curved at middle, pointed at apex, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose, two of them with shorter and robust cornuti in ventral position. Female genitalia (Figure 17p). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin like a bellshaped bead not sclerotized, anterior margin bilobate in the middle, each lobe kidney-shaped with the posterior part as wide as the anterior part, strongly sclerotized, sides flared, slightly sclerotized; ductus bursae very short, slightly sclerotized on ostium side; corpus bursae very short, ovoid, without signa; ovipositor lobes short and curved, 2.0 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores without spatulate tips. Larvae -L5 instar (Figure 4e). Length, 30-35 mm, width, 4.0 mm; head smooth, dark brown, prothoracic shield salmon beige; body with ground colour salmon pink, pinnacula and caudal plate pale brown. Young larvae are very similar in appearance to mature ones.

Etymology
Named inexpectata, because at first, before preparing the genitalia, it was thought to be Sesamia echinochloa; specific epithet treated as an adjective.

Bionomics
Larvae were collected on young stems and shoots of Echinochloa pyramidalis growing in wetland inhabited with various Poales species belonging to the following genera: Cyperus, Eriochloa Kunth, 1815, Juncus, Phragmites Adans, 1763 and Typha.

Diagnosis
See also the identification key to species of the wiltshirei subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the female genitalia: anterior margin of the ostium bursae bilobate in the middle, each lobe pea-shaped almost spherical, strongly sclerotized, sides slightly flared, less sclerotized; ovipositor lobes short and curved, 2.0 times longer than wide, ventral surface with a pronounced indentation.  (Figure 17q). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin like a domed bead, not sclerotized, anterior margin bilobate in the middle, each lobe pea-shaped almost spherical, strongly sclerotized, sides slightly flared, less sclerotized; ductus bursae short, slightly sclerotized; corpus bursae very short, ovoid, without signa; ovipositor lobes short and curved, 2.0 times longer than wide, ventral surface with a pronounced indentation, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores without spatulate tips.

Etymology
Named after Lefini river, on whose banks this species was collected; treated as a noun in apposition.

Bionomics
Biology unknown. The moth was caught with a light trap in grasslands surrounding Lefini riverside inhabited with various Poales species belonging to the following genera: Cyperus, Echinochloa, Eriochloa, Hyparrhenia, Sporobolus, Typha and Vossia.

Distribution
See also the identification key to species of the wiltshirei subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: internal side of costa convex; clasper 1.6 times longer than wide; sacculus ending with a narrow thickening toothed; cucullus like a short and very thick finger curved at the tip, in the shape of a bird's head; carina crest with two lateral lobes with shorter and robust cornuti in ventral position; anterior margin of ostium bursae bilobate in the middle, each lobe kidney-shaped, strongly sclerotized, the anterior part two times wider than the posterior part, sides flared, slightly sclerotized; ovipositor lobes short and curved, 2.4 times longer than wide.
Description Figure 16o-r. The general shape of the female's forewings is more elongated at the apex than those of the male. Male genitalia (Figure 17f, l). Tegumen with flat drooping peniculi; vinculum small with a small saccus, u-shaped at the bottom margin, w-shaped at the top margin without indentation. Valve with sacculus and cucullus fused; costa short, sclerotized, internal side slightly convex, clasper curved sausage-shaped parallel to the internal side of the valve, 1.3 times longer than wide, strongly toothed at the apex; sacculus broad, well sclerotized with a strong narrowing below a slightly flared apex ending with a narrow thickening toothed; cucullus small, sclerotized, like a short and very thick finger curved at the tip at the tip, in the shape of a bird's head, with scattered and papillated hairs; juxta small and flat trapezoidal, with a very small laterally compressed medial projection, slightly sclerotized; uncus stout and short, curved at middle, blunt at apex, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose, two of them with shorter and robust cornuti in ventral position. Female genitalia (Figure 17r). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin domed, the middle part sclerotized, flattened triangleshaped, the side parts not sclerotized, anterior margin bilobate in the middle, each lobe kidney-shaped, strongly sclerotized, the anterior part two times wider than the posterior part, sides flared, slightly sclerotized; ductus bursae very short, slightly sclerotized on ostium side; corpus bursae very short, ovoid, without signa; ovipositor lobes short and curved, 2.4 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores without spatulate tips.
Larvae -L5 instar (Figure 4f). Length, 30-35 mm, width, 4.0 mm; head smooth, orange brown, prothoracic shield salmon beige; body with ground colour salmon pink, pinnacula and caudal plate light brown. Young larvae are very similar in appearance to mature ones.

Etymology
Named after Rindini, a village near Manyoni in Singida Region; treated as a noun in apposition.

Bionomics
Larvae were collected on young stems and shoots of Sporobolus consimilis growing in wetland inhabited with various Poales species belonging to the following genera: Cyperus, Eriochloa, Echinochloa and Juncus.

Diagnosis
See also the identification key to species of the wiltshirei subgroup below. This species can be distinguished from other known species of the subgroup by the following combination of characters of the male and female genitalia: tegumen with rounded peniculi; vinculum with a saccus ushaped at the bottom margin; valve very wide, costa short, slightly swollen, without costal spine; clasper narrow kidney-shaped, three times longer than wide, parallel to the internal side of the valve, strongly toothed at apex; sacculus very broad with a strong narrowing below a slightly flared apex ending with a narrow thickening toothed; cucullus small, sclerotized, like a short and very thick finger curved at the tip in the shape of a bird head, with scattered and papillated hairs; vesica with a tongue-shaped cornutus adorned with longitudinal villi like close to the tip; posterior margin of the anteostial pad flattened heart-shaped, convex on the posterior side, pointed base towards the ductus bursae, anterior margin of the anteostial pad cup-shaped with sharp curved ends on each side; ovipositor lobes very short and sharp, 1.3 times longer than wide.

Description
Figure 16s-x. The male holotype was accurately described in much detail by Rungs (1963). The female is recorded here for the first time; antennae filiform, otherwise the wing pattern and colour are similar in both sexes. Forewings length: males 24.0 mm (min-max 24-25 mm, N = 2), female 27.0 mm (N = 1). Male genitalia (Figure 17g, h, m). The poor quality of the NHM genitalia preparation does not allow us to use it for a precise description; we also could not observe the genitalia preparation #613 by Wiltshire used by Rungs (1963) for his description; hence our description is only based on Rungs's drawing (1963). Tegumen with small rounded peniculi; vinculum narrow with a small saccus, u-shaped at the bottom margin. Valve very wide with sacculus and cucullus fused; costa short slightly swollen, without costal spine; clasper narrow kidney-shaped, three times longer than wide, parallel to the internal side of the valve, strongly toothed at apex; sacculus very broad with a strong narrowing below a slightly flared apex ending with a narrow thickening toothed; cucullus small, sclerotized, like a short and very thick finger curved at the tip in the shape of a bird head, with scattered and papillated hairs; juxta not seen; uncus thin, curved at middle, tapering near apex to a fine curved point, tufted with long hair on upper side; aedeagus short and stout, manica with a carina crest strongly spinose, vesica with a tongue-shaped cornutus adorned with longitudinal villi like close to the tip; cucullus small, sclerotized, like a short and very thick finger curved at the tip in the shape of a bird head, with scattered and papillated hairs; aedeagus short and stout, manica with a carina crest produced into paired lateral lobes strongly spinose. Female genitalia (Figure 17s). Apophyses anteriores with spatulate tips; lateral plates of ostial segment small, slightly sclerotized; ostium bursae large, posterior margin flattened heart-shaped, strongly sclerotized, convex on the posterior side, pointed base towards the ductus bursae, anterior margin strongly sclerotized, cup-shaped with sharp curved ends on each side; ductus bursae very short, slightly sclerotized on ostium side; corpus bursae very short, ovoid, without signa; ovipositor lobes very short and sharp, 1.3 times longer than wide, dorsal surface bearing short and stout setae; apophyses posteriores slenderer and longer than apophyses anteriores.

Bionomics
Larvae were collected in stems of Panicum repens.

Distribution
Egypt. The recorded localities are from regs, hamadas, wadis (mosaic #71) vegetation mosaic (White 1983) (Figure 15). (1963) noticed that, although this small species looks very similar in size and wing pattern to S. coniota, its genitalia morphology brings it closer to S. albivena; our study clearly shows it belongs to the Sesamia cretica group.

Rungs
Identification key to species of Sesamia wiltshirei subgroup Key based on internal morphology of the adult male (A) and female (B) genitalia.

Species delimitation analyses
Depending on the methods and loci analysed between 13 and 33 putative species are inferred by the SD analyses ( Figure 18). The sampled species from the S. cretica group are recovered by at least 60% of the SD analyses, with about one-half recovered by all SD analyses. Noticeably, fewer putative species are found by distance-based methods (19-24 putative species) and when examining the clades inferred in the concatenated nuclear phylogeny (13 putative species), whereas more putative species are found by tree-based methods (23-33 putative species). Based on the mCtax values obtained with LIMES, four SD analyses are considered as equally or more likely than the morphospecies (in mCtax order: Cytb.ASAP-W, Concat.GMYCs, Concat.PTP and Cytb.GMYCs). The three SD analyses considered as more likely (Cytb.ASAP-W, Concat.GMYCs, and Concat.PTP) however fit well with the morphology (Ctax > 0.94) except for S. salama, S. kafulo, S. babati, which are oversplitted (respectively by all three analyses, Concat.GMYCs and Concat.PTP), and the fourth SD analysis (Cytb.GMYCs) fully matches with morphology (Ctax = 1).

Discussion
Morphology, taxonomy and systematics Except for Sesamia taenioleuca whose forewings are purplish-pink, the remaining 30 Sesamia species revised here constitute a morphologically homogeneous group when only considering wing patterns and colours. However, as underlined by the results of the molecular phylogenetics analyses, the S. cretica group is not monophyletic and instead consists of five distinct lineages. When taking Figure 18. Maximum likelihood tree resulting from the analysis of the concatenated dataset, complemented by the results of molecular species delimitation analyses. The tree is shown on the left along with information on clade support for major nodes (S = supported; NS = not supported). Results of molecular analyses are displayed on the right, ranked depending on the mCtax value. The bars are coloured in green, yellow or red, depending on whether a given analysis matches the number of described species, split them or merge them, respectively. Analysis type, mCtax, Ctax (between morphology and a given analysis) and number of species found (Nb species) are summarized in the table at the bottom left.
into account male and female genitalic characters it is feasible to assign all the species revised here to six homogeneous subgroups, which mirror the clades evidenced by the molecular analyses: the S. albivena subgroup, the S. cretica subgroup, the S. fuscifrontia subgroup, the S. salama subgroup, the S. viettei subgroup and the S. wiltshirei subgroup. Specimens can be easily assigned to these distinct subgroups. For male specimens, the characters of the costa and the juxta allow for a clear assignation. For female specimens, the characters of the posterior and anterior margins of the anteostial pad are the most relevant for an easy identification of the subgroup. The identification key 1 presented above allows one to determine to which subgroup of the S. cretica group a species belongs. Four of these subgroups strictly correspond to four of the five clades recovered by the molecular analyses; the last two (S. fuscifrontia and S. wiltshirei subgroups) belong to the same clade, whereas the S. fuscifrontia subgroup is found to be paraphyletic due to the placement of representatives of the S. wiltshirei subgroup. The latter indicates that the combination of genitalic characters retained to define the S. fuscifrontia subgroup likely includes homoplasic characters; it is also the case when considering the S. cretica group as a whole, as underlined by the fact that it consists of five distinct evolutionary lineages.
Further morphological and molecular studies are likely required to increase our understanding of the diversity of the S. cretica group as currently defined, as it will likely be expanded to include other species yet to be described. For instance, Hermann Hacker's personal collection hosts several potential new species belonging to the group: two males from Iringa in Tanzania (genitalia N°24915 and N°2 5004) belong to a new species of the S. albivena subgroup; one male from Ethiopia (genitalia N°25082) belongs to the S. wiltshirei subgroup, and correspond to a new species closely related to S. rindini n. sp.; one male from Tanzania (genitalia N°24985) belong to a new species from the S. cretica subgroup and close to S. rufescens. In the Middle East, two species described in 2002 also belong to the S. cretica group. Sesamia ilonae Hacker, 2002(Hacker 2001 from Israel is a member of the S. fuscifrontia subgroup and S. vanharteni Hacker & Fibiger, 2002(Hacker et al. 2002 from Yemen is almost indistinguishable from S. cretica to the point that it would be interesting to do a molecular study to clarify its status. In addition, regarding S. cretica it is worth highlighting that specimens from this widespread species (distributed in Africa, Asia and Europe) present some geographical variation in the costal spine apex more or less rounded or triangular, which could indicate a potential species complex. In Asia, three species (Acrapex exsanguis Lower, 1902, Sesamia sp. from Australia and S. rungsi Boursin, 1957 from Afghanistan) also belong to the S. cretica group, and most likely to the S. fuscifrontia subgroup.

Molecular species delimitations
In this study, the results of SD approaches generally provide a good support for the newly described species. The five analyses with the highest mCtax values (see Figure 18) were followed to provide the new species descriptions. Among them, only three species (S. babati, S. kafulo and S. salama) are sometimes split when the morphology does not show any difference. For these species, in an iterative manner, morphological studies were further carried out but no notable differences among specimens from the same species were found. Here it can be postulated that these suggested splits are likely artefactual, possibly resulting from the low number of specimens sequenced and potential biases in branch length estimation (due to uneven gene coverage among some specimens), which could have affected the tree-based SD methods. Finally, the results of our SD analyses are also consistent with a general trend reported by other authors (Miralles & Vences 2013;Talavera et al. 2013;Hamilton et al. 2014;Lecocq et al. 2015;Dellicour & Flot 2018;Luo et al. 2018), which is that tree-based approaches (such as GMYC and PTP) generally yield higher numbers of putative species whereas distance-based methods (such as ABGD and ASAP) generate lower numbers of putative species. The latter underlines the importance of using a wide array of methods and settings to critically interpret and reassess the results of SD analyses, while avoiding biases that could result from the reliance on a limited number of approaches.

Ecology, distribution and diversity
Among the 24 collected species belonging to the different S. cretica subgroups, members of the clade grouping the S. fuscifrontia and S. wiltshirei subgroups are generally hygrophilous species inhabiting wetlands; by contrast, species from the S. albivena, S. cretica and S. salama subgroups are rather mesophilous and even xerophilous (especially S. cretica), inhabiting mostly woodland and secondary grasslands. This phylogenetic conservatism in habitat preference echoes the results of Kergoat et al. (2018), who found a similar trend among Sesamiina as a whole when contrasting the preferences for dry or wet environments. With regards to vegetation mosaics, marked preferences within S. cretica subgroups are also revealed. Members of the clade grouping the S. fuscifrontia and S. wiltshirei subgroup are equally found in Guineo-Congolian rain forest and secondary grassland mosaics, or woodland (Miombo, East African Costal, East African evergreen bushland, Sudanian woodland, Somalia-Masai bushland) mosaics. Species from the S. albivena, S. cretica and S. salama subgroups are mostly found in woodland (Miombo, East African coastal, East African evergreen bushland, Sudanian woodland, Somalia-Masai bushland) mosaics, and more seldom in Guineo-Congolian rain forest and secondary grassland mosaics. Two thirds of species (68%) are also recorded from altitudes between 500 and 1500 m asl; four species (13%) are recorded below 500 m asl and six species (19%) are recorded above 1500 m asl (mostly below 2000 m), and only S. mabira is recorded up to 2343 m asl.
Regarding host plants, for the clade grouping the S. fuscifrontia and S. wiltshirei subgroups, new host records are provided for 11 of the 17 known species. Interestingly, nine of them are found on Echinochloa pyramidalis. The predominance of this particular host plant can be likely accounted for by its ecological characteristics. It is a tall semi-aquatic perennial grass, native to tropical and subtropical Africa, which is present everywhere in seasonally inundated grasslands, swamps and banks of river and lakes. Although this plant is geographically widespread it has a very fragmented distribution as well: typically, Echinochloa pyramidalis habitats are frequently separated by several tens of kilometres. Here it can be hypothesized that the fragmented distribution of this host plant could have spurred population structuration (and possibly allopatric speciation) in Sesamia species belonging to the S. fuscifrontia and S. wiltshirei subgroups. For the S. cretica subgroup, two species were reared from Rottboellia cochinchinensis (Andropogoneae). For the S. albivena and S. salama subgroups, all specimens were collected with light traps, as no larvae from these groups were found on their hosts despite dozens of thousands of Sesamiina larvae collected during the field surveys. A possible explanation for this pattern is the fact that the damage caused by Sesamiina larvae is much less visible on some plants (noticeably Andropogon spp., Cymbopopon spp. and Hyparrhenia spp.) than others such as Echinochloa spp., Imperata cylindrica (L.) P. Beauv., 1812, Panicum spp., Pennisetum spp. and Rottboellia cochinchinensis. Considering that all species of the S. albivena and S. salama subgroups were caught in grasslands inhabited mainly by Andropogoneae (Andropogon spp., Cymbopogon spp., Hyparrhenia spp., Imperata cylindrica and Rottboellia cochinchinensis), it can be hypothesized that most species of these subgroups are likely to be associated with Andropogoneae species belonging to the three genera for which it is harder to spot larval damage.
In terms of geographic distribution, like most Sesamiina species already studied (Le Kergoat et al. 2015;Le Ru et al. 2017a, 2017b, 2017c, 2020a, 2020b, 2020c our results show that most species in the S. cretica group have restricted areas of distribution. When looking at the bioregions defined by Linder et al. (2012), in sub-Saharan Africa, 24 of the 31 studied species are only recorded from one bioregion, and 11 of them are only found in one or two localities. That said, the hypothesis that the sampling method (short sampling time in few localities) is partially responsible for this distribution pattern cannot be ruled out. In sub-Saharan Africa, almost half of the known species from the S. cretica group (15 species) are distributed in the Zambezian bioregion. The second region with the highest level of diversity is the Congolian bioregion (eight species), followed by the Southern African bioregion (six species), the Ethiopian bioregion (four species), the Sudanian bioregion (three species), and Madagascar (one species).
Until the early 2000s, only eight species of the S. cretica group species were known in sub-Saharan Africa, from several African bioregions; however, at the time only one species was recorded from the Zambezian bioregion, and none from the Congolian bioregion. Different reasons can be put forward to explain the significant increase in species diversity reported in these two bioregions. First, species discovery benefited from a high sampling effort since the early 2000s (Le Ru et al. 2006aRu et al. , 2006bNdemah et al. 2007;Matama-Kauma et al. 2008;Moolman et al. 2014;Ong'amo et al. 2014;Kankonda et al. 2017;Goftishu et al. 2018;Moeng et al. 2018;Ong'amo et al. 2018), using both light trapping and the collection of visually damaged grasses. In addition, the development and implementation of integrative taxonomy and systematics allowed us to clarify and disentangle at a higher pace the systematics and species boundaries of several species complexes. Such is the case for example of the S. albivena specimens preserved in the MCSN and NHM, which have been shown to actually belong to four different species.