Ropalidia guttatipennis (DE SAUSSURE) 1853, 1918

Ropalidia guttatipennis (DE SAUSSURE) 1853 View in CoL

Icaria guttatipennis DE SAUSSURE, 1853

Icaria politica DE SAUSSURE, 1854

Icaria tricinctella GRIBODO, 1895 View in CoL [loc typ. Maputo, Mozambique], syn. nov.

Type material. Four females from Senegal (deposited in MNHN) were treated as syntypes (J. M. Carpenter, 1999). Another specimen with a similar label type from MHNG was assigned a lectotype status, while the four MNHN specimens were assigned a status of paralectotypes (J. Kojima, 2001) .

Icaria politica DE SAUSSURE. The species description is partial, and de Saussure himself stated that “ I would not be surprised if this was a variety of Icaria guttatipennis” (H. De Saussure, 1853) , which was also mentioned by Bequaert ( Bequaret, 1918). The examination of the types from NHM indeed supports this claim; the type is conspecific with R. guttatipennis (DE SAUSSURE) , specifically the TT cluster.

Icaria tricinctella GRIBODO. Gribodo View in CoL described Icaria tricinctella View in CoL from Mozambique, which agrees with the redescription of I. cincta View in CoL by de Saussure; mainly yellow pronotum, large yellow spot on the mandible and thick yellow band on T2 ( Gribodo, 1895). He also mentioned bicolorous terminal antennal article in males ( Gribodo, 1895), which is reported in another of de Saussure’s papers (description of R. cincta View in CoL male) (H. De Saussure, 1863), but missing from Bequaert; Bequaert mentions this taxon as the synonym of R. cincta ( Bequaret, 1918) View in CoL . The examination of the type photos of Icaria tricinctella GRIBODO View in CoL shows that it is conspecific with R. guttatipennis (DE SAUSSURE) , TT cluster.

Comments. This is the most variable African Ropalidia species, which should be treated as a species complex based on the combination of the phenotypic and genetic differences. It is characterized by the moderately developed superior propodeal carina, distally narrowing propodeum and complete lack of or weakly developed inferior carina; in addition, females have strongly punctate inner orbit, while males have almost a clavate antenna and bicolorous terminal flagellomere dorsally. The scope of colour variation in females includes almost the entire scope of colour variation of the non- capensis -group. Due to extreme variation, numerous problems may arise in the separation of several species.

Morphology and variation. The overall morphological features of this species are relatively constant, with moderate variability of the T1 and propodeum shape.We defined eight clusters of this species based on morphological, colour, and genetic features.

1) n colour cluster ( nominal form, Figure 92a View FIGURE 92 ). Holotype colour pattern, characterized by the reddish-brown basal colour with yellow colour confined to the head in females, or with a thin yellow line on pronotum and, at most, a thin yellow band on T2 . This cluster is common in the West to central Africa ( Senegal to the Central African Republic).

2) tT cluster ( Figure 92b View FIGURE 92 ). This cluster is characterized by the thin yellow line on pronotum and a wider posterior yellow band on T2 (therefore, tT). Two specimens from this cluster have an extremely thick yellow band on T2 , which occupies more than half of the entire segment surface, prompting Giordani Soika to consider a new subspecies, which he labelled as R. cincta ssp. villersi . Both specimens were labelled, but the name was not published, therefore remaining invalid. This cluster is present from Guinea to Uganda.

3) TT cluster ( Figure 92c View FIGURE 92 ). This cluster has the most yellow colour on the body, including a thick band on T2 with a predominantly yellow pronotum or a thick yellow line on the pronotum. This cluster is a predominant colour form in the South Africa and Mozambique, although it occurs across entire Sub-Saharan Africa. This form was commonly referred to as R. cincta (LEPELETIER) in previous papers and museum collections.

4) b cluster ( Figure 92d View FIGURE 92 ). This cluster is characterized by the dark brown or even black basal body (abbreviated from black), including an entirely black antenna from above, black tarsi and strong blackening of the anterior part of the fore wing. This cluster is common in Benin and Cameroon.

5) d cluster ( Figure 92e View FIGURE 92 ). This is a likely cline of the b cluster towards South, with a mainly darker colour (abbreviated from the dark); specimens from this cluster have dark brown basal colour, but lightly coloured antenna, gena and tarsi. This cluster is predominant in Angola.

6) sm cluster ( Figure 92f View FIGURE 92 ). Brown basal colour, but with interestingly coloured antenna; female antenna resemble males, as they are blackened from above, but terminal flagellomere is more lightly coloured dorsally, while males have entirely black antenna from above (in contrast to the nominal form, whose males have ferruginous scape-AF4 from above and blackened remaining segments).Also, the average size of these specimens is substantially smaller than the nominal form, up to nearly half of the body size of specimens from Senegal (abbreviated from smaller). This cluster also has a separate 28s rDNA sequence, suggesting that this might be the most distinctive cluster and could be regarded as a separate taxon if additional support is gained through larger-scale genetics analysis. This cluster is common from Cote d’Ivoire to Nigeria.

7) e cluster. This cluster is represented by a single examined specimen from Ethiopia. Morphologically, it is a mix of R. guttatipennis (DE SAUSSURE) and R. dondo sp. nov., with predominantly brown body colour and reduced yellow markings. Although the COI gene analysis suggests that this is a separate lineage more closely related to R. dondo sp. nov. than R. guttatipennis (DE SAUSSURE) , the pubescence of this specimen is short and silvery, which is in direct opposition to R. dondo sp. nov. In addition, the specimen shares the same 28s rDNA sequence as the main cluster of R. guttatipennis (DE SAUSSURE) .

8) r cluster. This is by far the most complex cluster, based on only four genetically verified and remote specimens (abbreviated from remote). The common feature of these specimens is their abundant yellow area (present in three of four specimens), but they seem to form a very distant genetic group, separated substantially from the nominal form (by at least a few more separate and sufficiently distinctive species). The main morphological problem of this cluster is the inconsistency of inner orbit punctures, since two specimens have punctures, one has poorly defined and barely visible punctures, while one specimen has impunctate inner orbit. Furthermore, not even the geographical information provides a consistent pattern, since the three mostly yellow specimens (that would resemble the TT cluster; equivalent to Figure 92c View FIGURE 92 ) are from Tanzania and Kenya, while the last specimen (with less yellow colour) is from the Gambia. Finally, one of these specimens (from the Island of Pemba in Tanzania) has a different 28s rDNA sequence, suggesting a separate lineage from most other specimens of this species. Arguably , this cluster might satisfy the support towards elevation to an independent species, but with the absence of males and more specimens, we retained it as R. guttatipennis (DE SAUSSURE) .

Genetics. This species was the most frequently sequenced among all examined specimens; 35 sequences were used in the analysis. The results of the COI analysis had indicated numerous distinctive clusters, suggesting an extreme pattern of genetic variability of this species. In addition, we assigned two remote clusters to the same morphospecies; notably, this is not an ideal situation, but in the absence of more data, there was not enough support to resolve this issue any further. Comparing genetic data and observed colour patterns did not suggest any meaningful and reliable pattern. The most complicated situation in this entire diversity was the analysis of 28s rDNA, which was nearly wholly invariable for this entire cluster. Two small separate clusters were recorded, one belonging to the sm cluster, while the second belonged to an island population from Tanzania, within the r cluster (this sequence did not seem to represent a separate lineage aligned with the COI gene). One of the most interesting findings was the sympatric occurrence of multiple clusters in the same micro-location, suggesting a very complex population structure and possibly hybridization between different lineages.