Ageniellini

Taxonomy of Ageniellini

The tribe Ageniellini was established by Banks (1912) (see also Day 1981, p. 5). He classified the Pepsinae into two tribes, Ageniellini and Pepsini. This system was followed by subsequent using the name Macromerini ( Townes 1957) or Auplopodini ( Evans 1973) for Ageniellini . However, logical ( Shimizu 1994; Pitts et al. 2006; Shimizu et al. 2010) and molecular phylogenetic analyses (

et al. 2015; Rodriguez et al. 2016) corroborated that Ageniellini is monophyletic, but Pepsini is

HISTORY tribal name Ageniellini , because its monophyly is strongly supported by Waichert et al. (2015) preliminary phylogenomic dataset of UCEs.

Ageniellini has been morphologically defined by the combination of the following features (1994): (1) the female labial prementum has very long, curved, erect setae ( Shimizu 1994); (2) the metasomal tergum 1 is petiolate basally (tergum 1 is narrower immediately behind the articulation width at the articulation itself; Day 1988); (3) the male metasomal tergum 1 is attenuated basally, sided or petiolate ( Haupt 1927, p. 23); and (4) the female metasomal tergum 1 lacks a lateral crease (1957). It should be noted, however, that these morphological characteristics have changed to other secondarily or showed reversals in multiple lineages of Ageniellini ( Shimizu et al. 2010, fig. 11). This that the above features cannot characterise the Ageniellini strictly, although these are important characteristics of the tribe.

Evans and Shimizu (1996) classified Ageniellini into two subtribes: the Ageniellina , which is morphologically and behaviourally generalised group of Ageniellini ; and Auplopodina , which is morphologically and behaviourally advanced group of Ageniellini , based on Shimizu’s (1994) Ageniellina , the females usually make short burrows and terminal cells from pre-existing holes or the soil. After carrying their prey spider into the cell and laying an egg on the spider’s abdomen, they the burrow by pulling twigs, pebbles, and soil particles, holding them in the mandibles and backwards, and stuff them into the entrance of the burrow ( Kurczewski and Kurczewski 1987; Evans Shimizu 1996). Thus, wasps of Ageniellina do not display the building-type nesting behaviour, and water carriers or mud daubers. A potential exception to this is one member of Ageniellina , flavipennis Banks View in CoL , which is known to utilise aerial mud cells, as mentioned immediately ‘Summaries of the communal behaviour of eight species’. It is questionable, however, whether the is a true water carrier and mud dauber and whether it belongs to Ageniellina ; see the discussion ‘ nesting behaviour in Ageniellini’.

In contrast, in Auplopodina , the females exhibit unusual nest-constructing behaviour. They drink away from the nest and carry the water in their crop to clayey ground or their own nest (hence, wasps group are named ‘water carriers’; Evans and Shimizu 1996). There they regurgitate water to moisten and remove the soil in the form of mud pellets with their mandibles. They then hold the pellets mouthparts (specifically in the ‘mud basket’, ie the space between the posterior face of the labial and long, stout, curved bristles rising from the prementum; Shimizu 1986, figs 37, 38, 1994, figs Shimizu et al. 2010, p. 110). In burrowing subterranean nests, the females excavate a short burrow ground, using water. The mud pellets generated during this process are discarded away from the entrance or beside it. In contrast, in building aerial nests, the wasps carry the pellets to their nest cell they plaster them onto the cell with the dorsal tip of the metasoma (ie the pygidial area; Shimizu 40–46), bending the metasoma forwards beneath the mesosoma within the cell ( Williams 1919, Shimizu 1986, fig. 39; Evans and Shimizu 1996, fig. 6). In any case, the wasps close the nest cells with other soft materials. All the species of Auplopodina exhibit the building-type nesting behaviour, and are considered water carriers and mud daubers.

Summaries of the communal behaviour of eight species

Almost all species of Pompilidae are solitary. Only eight species have been known to be practically potentially communal. These species are the Southeast Asian (1) Macromeris violacea Lepeletier (1919) , (2) Paragenia argentifrons (Smith) ( Williams 1919; Figure 2) and (3) Macromerella honesta (Barthélémy and Pitts 2012) ; (4) Japanese Machaerothrix tsushimensis Yasumatsu ( Shimizu 2004) (Figure South African Auplopus femoralis (Arnold) ( Weaving 1994) , (6) Panamanian A. esmeralda (Banks) (1980) , (7) Costa Rican A. semialatus Dreisbach ( Wcislo et al. 1988) , and (8) Brazilian Lissagenia

( Santos et al. 2017). Next, we present a summary of their communal behaviour.

(1) In Macromeris violacea, Williams (1919) observed one to four females at one time on a single consisting ultimately of 15 cells arranged in more or less vertical rows in a tree hollow (Williams

(2) Paragenia argentifrons build groups of mud cells arranged vertically in contact with one another stout bamboo canes ( Williams 1919; Figure 2). One nest consisted of 24 mud cells. In or about up to eight wasps existed at one time. Williams (1919) described their ‘peaceful’ interaction communal nesting by multiple females, including concurrent respective cell construction.

(3) Macromerella honest built multicellular nests (up to 15–16 cells) on either side of broad leaves, or trees (Barthélémy and Pitts 2012). Nests were composed of two rows of exposed ovoid made of a plastic mud mixture (clay and sand grains). At least two females of successive remained on a nesting site for a week (Barthélémy and Pitts 2012). There was, however, no communal nesting observed by the authors.

(4) Machaerothrix tsushimensis usually build a cluster of mud cells covered with an outer mud envelope in narrow spaces concealed and sheltered from rain ( Shimizu 2004, figs 1A–C, H largest nest contained about 10 cells and up to five cohabiting females, although usually two coexisted on a nest ( Shimizu 2004; Figure 3 View Figure 3 ). The most dominant female performed normal activities, including transporting prey into cells, while the subordinate females scarcely brought to the nest and were apt to spend much time in resting on the nest ( Shimizu 2004, fig. 4).

(5) Auplopus femoralis arranged 2–28 mud cells inside the cavity of trap-nests (bundles of canes) (1994). Two trap-nests in a bundle of three canes were nearly full of cells. A female was seen one and entering the other with no interaction with the other two females. Weaving (1994) stated these females may have belonged to one ‘family’ of wasps, although he did not confirm communal nesting.

(6) In A. esmeralda , four females were seen flying in and out of the base of a fallen palm frond (1980). Inside the frond, there was a nest consisting of 95 mud cells arranged in an elongate mass the curved inner wall of the frond base ( Kimsey 1980, fig. 1a, b). Many of the adults had emerged, and most cells were empty.

(7) In A. semialatus , two to eight females built and maintained groups of up to 21 mud cells ( Wcislo 1988). The cells were on the underside of the trunks of leaning palm trees. Cohabiting females usually tolerant of one another and defended their nests against natural enemies, but often intensely competitive when spiders were brought to the nests. Construction of new brood the reconditioning of old cells for reuse, was usually cooperative in that more than one participated.

(8) One to three females of Lissagenia flavipennis were observed on the same mud nests, which located on the house walls or in the crevices of the walls. Usually, one female stayed outside crevice, while the other female(s) remained on the nest ( Santos et al. 2017, fig. 1a). however, two females were present on the same nest ( Santos et al. 2017, fig. 1b). A female a prey spider ( Enoploctenus cyclothorax (Bertkau)) in a nest ( Santos et al. 2017, fig. 1c). None were observed building their nests and there was no evidence of communal nesting by the females.

Previous phylogenetic analyses

The phylogeny of the Pompilidae was first analysed using morphological information by Shimizu (1994 Pitts et al. (2006). Their results were similar in that Auplopodina ( Machaerothrix Haupt , Phanagenia Auplopus Spinola , Macromerella Banks , Macromeris Lepeletier ) were associated into a monophyletic but the phylogenetic relationships of other members of Ageniellini ( Meragenia Banks , Poecilageniella Ishikawa, Priocnemella Banks , Phanochilus Banks , Atopagenia Wasbauer , Ageniella Banks ) were not analysed.

Shimizu et al. (2010) performed maximum parsimony and Bayesian inference analyses based on morphological characters, coded for 41 exemplar species, including five outgroup species,

almost all the genera and subgenera of Ageniellini . This study explored the evolution of nest behaviour based on the phylogeny. The strict consensus tree shows that the Ageniellini is

HISTORY

Haupt, Paragenia Bingham , Macromerella , and Macromeris . However, the relationships of the rest Ageniellini were not sufficiently resolved.

More recently, Waichert et al. (2015) reconstructed a molecular phylogeny of the world Pompilidae Bayesian and maximum likelihood analyses of four nuclear DNA markers. The resulting consensus showed that the sampled Ageniellini were grouped into a monophyletic group sister to Cyphononyx

Turner. Within Ageniellini , Machaerothrix sp. ( Auplopodina ) was the most basal clade. The Ageniellini diverged into two monophyletic groups, one comprising Macromeris and ( Auplopodina ) and the other comprising Phanagenia (Auplopodina) , Ageniella Banks and Eragenia However , these last two genera belong to Ageniellina . Thus, Waichert et al.’s (2015) tree did not support two subtribes, Ageniellina and Auplopodina , of the tribe Ageniellini .