Andesipolis Whitfield and Choi 2004

Mitio, Shimbori Eduardo, Souza, Souza-Gessner Carolina Da Silva, Maria, Penteado-Dias Angelica & Richard, Shaw Scott, 2017, A revision of the genus Andesipolis (Hymenoptera: Braconidae: Mesostoinae) and redefinition of the subfamily Mesostoinae, Zootaxa 4216 (2), pp. 101-152 : 105-110

Andesipolis Whitfield and Choi 2004

Andesipolis Whitfield et al. 2004: 3

Type species Andesipolis masoni Choi and Suh.

Diagnosis. In addition to the diagnostic characters cited by Whitfield et al. (2004) (e.g. presence of vein 2a on fore wing; propodeum with cross bridge dividing areola, rarely areola completely absent; presence of midpit on mesoscutum), several other characters are useful to distinguish Andesipolis , especially when compared with Rhysipolis . The pronotum of Andesipolis has a transverse carina dorsally, dividing the notum in two parts, the anterior part frequently at lower level than posterior part, crenulate and/or with mid-longitudinal carina. Hypostomal carina very weak or absent (hardly visible even under scanning electron microscope), and occipital carina always complete and meeting the base of mandibles. Epicnemial carina incomplete ventrally, present at least laterally and frequently extending ventrally; in most species the epicnemial carina ends in a curve ventrally. Propodeum relatively short, always shorter than petiole. Veins (RS+M)b and r-m spectral. Metasoma beyond petiole varies from membranose to evenly weakly sclerotized. Head and mesoscutum smooth and polished; propodeum, metapleuron and petiole variable: smooth to rugose, polished to finely and irregularly strigulate (similar to a very fine fingerprint-like sculpturing, not visible under light microscope but the reflection is clearly distinct from a polished surface). Hind tibial spurs short, less than 1/3 length of basitarsus. Spiracles of TII and TIII clearly on laterotergites. Ovipositor sheaths covered with long and dense setae.

Biology. One species is known to be a gregarious koinobiont parasitoid of shelter-building Pyralidae (Lepidoptera) feeding on leaves of Phenax rugosus and Boehmeria bullata (Urticaceae) ( Townsend & Shaw 2009).

Etymology. The genus was named for its distribution, originally thought to be restricted to the Andes mountain chain (Andes-), and the morphological resemblance with the genus Rhysipolis (- sipolis) ( Whitfield et al. 2004).

Discussion. With the newly described species, Andesipolis becomes the most speciose genus in the Mesostoinae , with 26 described species. Hydrangeocola is the second with 11 species, but apparently with a much more restricted distribution. An ongoing review of Aspilodemon is showing that this genus might be as species-rich as Andesipolis . On the other hand, all other Mesostoinae occurring in Australian region have no more than a handful of species described, although many more genera (three in South America vs. 10 in Australian Region). Whether this is a result of biological processes or of the arbitrariness of the taxonomic treatments is uncertain.

Some features of the Andesipolis species surveyed in this work have proven helpful for comparison with other mesostoine genera. The state of occipital and hypostomal carinae, as discussed by Townsend & Shaw (2009), is an important character to distinguish Rhysipolinae from other Hormiinae s.l. (Wharton 1997). As observed by Whitfield el al. (2004), the hypostomal carina is weak or completely absent in Andesipolis (see Figures 12 and 23 in Whitfield et al., 2004), a condition also found in the Mesostoinae s.s. ( Quicke & Huddleston 1989). Therefore, this is another argument in favor of a closest relation with Mesostoinae rather than Rhysipolinae , where the hypostomal carina is well defined and close but clearly separated from occipital carina ventrally. In Andesipolis the spiracles of the TII and TIII are positioned on the laterotergites. This is another important feature defining Mesostoinae s.s. and Rhyssalinae ( Quicke & Huddleston 1989; Wharton 1993). One of the newly described species has the dorso-ventrally compressed mesosoma (as in Mesostoa ) and also the mesoscutum projected anteriorly over the pronotum, considered a diagnostic character for Mesostoinae by Quicke & Huddleston (1989), but not found in the South American species until now (i.e. Aspilodemon and Hydrangeocola ). In Andesipolis the labial palpi is 3- segmented as in Aspilodemon and Hydrangeocola . These two genera also present the fingerprint-like microsculpturing in some species (e.g. with the right light incidence, reflection reveals a microsculpturing covering most of body, except metasoma beyond TI; it can be characterized only under Scanning Electron Microscope), providing a mat aspect to the body surface as seen in most Andesipolis . The relatively long pronotum with a distinct transverse carina, seen in Andesipolis , is identical to that of Neptihormius , the only other genera sharing this type of pronotum within Mesostoinae . Neptihormius differs from Andesipolis in having a complete epicnemial carina (a plastic character within Mesostoinae , varying from completely absent to complete, although a carina interrupted ventrally seems to be the plesiomorphic common state), and vein 2a absent. In Hormiitis these two characters have the same state as in Andesipolis , but the short pronotum covered dorsally by protruding mesoscutum is of a typical Mesostoinae s.s. The midpit and short notauli, more or less directed to the mid pit posteriorly, are present in all genera of Mesostoinae (Mesostoini, Hydrangeocolini, Austrohormiini and Avgini) distributed in Austral lands (i.e. Australasia and South America). The notauli short and directed to a midpit is a putative synapomorphy for Mesostoinae , as well as the weak or absent hypostomal carina. The lateral position of the spiracles on TII and TIII is also a diagnostic character, although possibly a plesiomorphic one (see discussion in Wharton 1993).

Distribution ( Fig. 1). The genus was recorded from Argentina, Brazil, Chile, Colombia, Ecuador and Peru, although the single specimen from Peru could not be identified or described due to its relatively poor condition. The species of Andesipolis are only known from South America, at high latitude and/or altitude. In the higher latitudes the species are found close to sea level, while in low latitudes the elevation is increasingly higher. The southernmost record, in Tierra del Fuego ( Argentina), is at 54°36'S and 60–200 meters above sea level. Nevertheless, some species are found close to sea level in Chile, at 38°47'S. The northernmost record is in Colombia, at 5°25’N and 3,380 meters above sea level. The highest location is also in Colombia, at 3,880 meters and 4°31’N. All the previously cited records are from the western coast of South America, most of them occurring along the Andes mountain chain. However, the Atlantic mountain chains of Brazil, in the eastern coast, also have a considerable diversity of Andesipolis . The Brazilian mountain chains are characterized by medium elevation (rarely higher than 2,000 meters) and mostly covered by subtropical wet forest with high altitude fields close to mountaintops, and usually above 2,000 meters of elevation. The distribution of species of Andesipolis is therefore characterized by mild elevation and mild latitude ranges in Brazil (about 1,000 to 2,200 meters above sea level and 20–26°S), but in the southernmost records, at S 26°19’ and S 24°16’ the elevations are 650 and 850 meters, respectively.

The distribution of Andesipolis , and the results of all phylogenetic studies that include the genus, are good evidence of biogeographic processes, and corroborate the theory of Mesostoinae as a Gondwanan clade, sister of Aphidiinae , which, unlike its sister group (Schlinger 1974; Belshaw et al. 2000), remained restricted to the Austral lands. The widely disrupted Austral (South America and Australia and surrounding islands) distribution has several parallels in fauna and flora, and has been subject of researchers’ interest for centuries. Some of those have very interesting relation with the history of Andesipolis , such as the similarly distributed genus Podonomus ( Chironomidae , Podonominae ), mostly restricted to Southern South America and recently found in inselbergs in Brazil ( Brundin 1966; Roque & Trivinho-Strixino 2004; Trivinho-Strixino et al. 2012).

The high altitude fields, occurring in patches, mostly at mountaintops in Southern Brazil, are biogeographical relicts, where several Andean and Austral-Antarctic taxonomic groups are present ( Safford 1999). As a result of the climatic fluctuations, e.g. the glaciations during the Quaternary, the floristic composition of these fields are highly endemic, with several taxonomic groups present in other Austral regions ( Fiaschi & Pirani 2009; Behling 1997). The history of these areas and the Mesostoinae in Brazil are closely related, particularly for Andesipolis . This genus has been collected and identified in Brazil for the first time in 2001, at São Bento do Sul, SC. After this first record, only a couple of specimens in poor conditions were collected in Teresópolis, RJ. The amount of material was increased only after systematic sampling above 1,000 m of elevation at Itatiaia National Park, as part of projects focused on surveying the fauna of high altitudinal ranges. In our results, we found a strikingly high diversity and endemism in high altitude fields (i.e. at least seven of the 12 Brazilian species are endemic for the fields of Itatiaia ), especially considering the small patchy coverage of this type of vegetation. How the secondary colonization of other adjacent vegetational types occurred, and which physiological and ecological traits lead to the actual distribution of this genus, are interesting questions remaining to be addressed.

Figures

FIGURES 2 – 13. Pronotum, dorsal. 2, Andesipolis atlanticus sp. n.; 3, Andesipolis brasiliensis sp. n.; 4, Andesipolis flamengo sp. n.; 5, Andesipolis soaresi sp. n.; 6, Andesipolis longicollis sp. n.; 7, Andesipolis monteiroi sp. n.; 8, Andesipolis masneri sp. n.; 9, Andesipolis chilensis sp. n.; 10, Andesipolis pseudoscapus sp. n.; 11, Andesipolis supernus sp. n.; 12, Andesipolis brevisulcus sp. n.; 13, Andesipolis whitfieldi sp. n.

FIGURES 14 – 25. Propodeum: 14, Andesipolis atlanticus sp. n.; 15, Andesipolis brasiliensis sp. n.; 16, Andesipolis itatiaia sp. n.; 17, Andesipolis flamengo sp. n.; 18, Andesipolis longicollis sp. n.; 19, Andesipolis monteiroi sp. n.; 20, Andesipolis cacildis sp. n.; 21, Andesipolis bicolor sp. n.; 22, Andesipolis belshawi sp. n.; 23, Andesipolis ona sp. n.; 24, Andesipolis masneri sp. n.; 25, Andesipolis supernus sp. n.