Anace perpusilla, WALKER, 1856: 1720
Przybyłowicz, Łukasz & Tarcz, Sebastian, 2015, Strong sexual dimorphism unraveled by DNA analysis - towards a better understanding of Pseudothyretes classification (Lepidoptera: Erebidae: Arctiinae), Zoological Journal of the Linnean Society 173 (1), pp. 22-54 : 44-47
treatment provided by
Holotype: Male (no locality) ( BMNH).
Additional material (233 ♂♂, 16 ♀♀): Guinea: ♂ ‘ Konakry, Forêt classée de Diéckié, 7–16/5/2004, leg. Erik Vingerhoedt’ ( RMCA) ; ♂ ‘ Guinée Forestière, 25 km südlich Boula, Forêt Classée du Mont Béro, 8°10′53.76″N, 8°42′28.2″W, 680 m, Lichtfang , 6.VI.2013, leg. M. Ochse’ ( ZSM) GoogleMaps ; 9♂♂ Guinée Forestière, 19 km südlich Lolo, Institut de Recherches Environmentales de Bossou , 7°38′59.30″N, 8°30′21.22″W, 587 m, Lichtfang , 10.VI.2013, leg. M. Ochse’ ( ZSM) GoogleMaps . Sierra Leone: 22♂♂ ‘ Kenema , IX/ X.1974, don A. Allaer’ ( RMCA) ; ♂ same data, but ‘VI/ VII 1974 ’. Liberia: 2♂♂ ‘ Grand Gedeh County, FDA Camp, Putu Jawodee, Bufferzone to Sapo National Park , 20–26.XI. 2012 ′ ( SZS) ; 3♂♂ ‘ Nimba County, Nimba Berge, 4 km südöstlich Yekepa, 7°33′39.31″N, 8°30′23.67″W, 622 m, Lichtfang , 11.VI.2013, leg. M. Ochse’ ( ZSM) GoogleMaps ; ♂ same data, but ‘ 13.06.2013 GoogleMaps ’; ♂ same data, but ‘ 14.06.2013 GoogleMaps ’; ♀ ‘ Nimba County, Nimba Berge, 5 km südöstlich Yekepa, 7°33′40.20″N, 8°30′10.62″W, 705 m, Lichtfang , 13.VI.2013, leg. M. Ochse’ ( ZSM) GoogleMaps . Ivory Coast: 3♂♂ ‘ Forêt du Banco , 2/ 3.X.1964, P. Griveaud’ ( RMCA) ; ♂ same data, but ‘ IX.1963 ’; ♂ ‘ Lamto , 11.01.1969, R. Vuattoux’ ( RMCA) ; ♂ same data, but no date ; ♂ same data, but ‘ 10.09.1966 ’, Cl. Girard ; 2♂♂ ‘ Station Orstom, Adiopodoume , IX.1963, P. Griveaud’ ( RMCA) . Ghana: 2♂♂ ‘ Eastern, Atewa Hills , 800 m, 7 km W Sagyimase, 22–24.III.2010, K. Larsen & W. Kubasik’ ( AMU) ; 3♂♂ ‘ Ashanti, Bobiri , 240 m, 4 km N Kubeasi, 18–20.III.2010, K. Larsen & W. Kubasik’ ( AMU) ; ♂ same data, but 9–12.III.2010, ( AMU) ; ♂ ‘ Central Kakum NP – Entrance , 5°21′N 22°W, 220 m, 31.V.2011, leg. J.&W. De Prins’ ( RMCA) GoogleMaps ; ♂ ‘ Bia Cons. Area, Bongo Camp , 23.01.2009, leg. U. Dall’Asta’ ( RMCA) ; ♂ same data, but ‘ Camp 15, 21.01.2009 ’ ( RMCA) ; ♀ same data, but ‘ Camp 15, 12.01.2009 ’ ( RMCA) ; 1♂♂ 3♀♀, ‘ Volta Region, Likpe Bakua , 16–18.X.2011, leg. Safian Sz., Pühringer F. & Pöll N. ’ ( SZS) ; 2♂♂ 2♀♀ same data, but ‘ XI.2011 ’ ( SZS) ; 7♂♂ ‘ Ashanti Region, Bonkro, Nyamibe Bepo Forest Reserve , 26–28.X.2011, leg. Safian Sz., Pühringer F. & Pöll N. ’ ( SZS) ; ♂ ‘ Eastern Region, Bunso Arboretum , 10–18.IX.2010, leg. Safian Sz., Dall’Asta U. ’ ( ISEA) ; 2♂♂ ♀ same data, but ‘ 19– 20.X.2011 ’, leg. Safian Sz., Pühringer F. & Pöll N. ’ ( ISEA) ; ♂ ♀ same data, but ‘ 12–20.X.2009, leg. Safian Sz., Walker A., Davey S., Onstein R. ’ ( ISEA) ; ♂ same data, but ‘ 22–30.X.2009, leg. Safian Sz., Walker A., Davey S., Onstein R. ’ ( ISEA) ; ♂ ‘ Central Region, Kakum Guesthouse, Kakum National Park , 23–26.X.2011, leg. Safian Sz., Pühringer F. & Pöll N. ’ ( ISEA) ; 2♂♂ 2♀♀ same data, but ‘ XII.2011, leg. Safian Sz. ’ ( ISEA) ; 5♂♂ same data, but 01–03.XII.2011, leg. Safian Sz. ( ISEA) ; ♂ ‘ Central Region, 30 km north of Cape Coast, 1 km north of Abrafo, Kakum National Park , Visitor Center , 1°24′W, 5°21′N, 240 m, light attraction, 18.X.2003, leg. M. Ochse’ ( ZSM) GoogleMaps ; 2♂ same data, but Canopy Walkway , 1°24′W, 5°21′N, 260 m, light attraction, 20.X.2003, leg. M. Ochse’ ( ZSM) GoogleMaps ; ♀ same data, but 22.III.2003 GoogleMaps ; ♂ ‘ Western Region, 16 km east of Elubo, Ankasa National Park, 1 km south-east of Nkwanta camp, Exploration Base , 103 m, light attraction 5°16′29.2′N, 2°38′46.0″W, February 24th 2006, leg. M. Ochse’ ( ZSM) ; ♂ ‘ Western Region, 16 km east of Elubo, Ankasa National Park, 7.3 km south of Nkwanta camp, Visitor Center , 5°13′2.3′N, 2°38′5.4″W, 58 m, light attraction, 6.V.2005, leg. M. Ochse’ ( ZSM) ; ♂ ‘ Western Region, 11 km east of Elubo, Ankasa National Park, Exploration Base , 5°27′N, 2°04′W, 100 m, light attraction, 15.X.2003, leg. M. Ochse’ ( ZSM) GoogleMaps ; ♀ ‘ Western Region, 16 km east of Elubo, 1 km southeast of Nkwanta camp, Ankasa National Park , Exploration Base , 103 m, light attraction, 5°16′29.246.0′W, 24.II.2006, leg. M. Ochse ; ♂ ‘ Western Region, 16 km east of Elubo, Ankasa National Park, 500 m northwest of Nkwanta camp, 5°17′09.1′N, 2°38′29.1″W, 92 m, light attraction, 4.V.2005, leg. M. Ochse’ ( ZSM) ; ♀ ‘ Western Region, 3 km E of Elubo , 5°17′N 2°45′W, 30 m, 8.VI.2011, leg. J.&W. De Prins’ ( RMCA) GoogleMaps ; ♂ ‘Akwapim Togo Range, Volta Region, Biakpa Mount Gemi Area, Mountain Paradise , 6°50′57.4′N, 0°25′11.8″E, 459 m, light attraction, November 23rd 2011, leg. M. Ochse’ ( ZSM) ; ♂ same data, but ‘ 24.XI.2011 ’; 5♂♂ ‘ Umg. Yamfo , 15–20.X.1993, leg. L. Kühne’ ( ZMHB) ; 2♂♂ ‘ Kumasi , 15.IX.1967, Endrödy Yunga’ ( ISEA) ; ♂ ‘ Tafo , 10.V.1969, Endrödy Yunga’ ( ISEA) ; ♂ ‘ Busua , 12.IV.1969, Endrödy Yunga’ ( ISEA) ; ♂ ‘ Kwadso , 14.VII.1969, Endrödy Yunga’ ( ISEA) ; ♂ ‘ Ghana, 6.VI.1967, Endrödy Yunga’ ( ISEA) ; ♂ ‘Ashanti- Region, Kumasi-Kwadaso , 300 m, 4.70, leg. D. Schröder’ ( ZSM) ; ♂ ‘ Ajenjua Bepo Forest Reserve , 24– 30.08.2006, site 1, N06°22′2.3′W01°01′58.6″, Elevation: 300 m a.s.l., leg. M. Ba ˛kowski’ ( ISEA) . Nigeria: ♂ ‘ Ibadan, Iita Forest , 21–30.IX.2010, leg. Sz. Safian & A. Horvath’ ( SZS) ; 1♂♂ ‘ Cercopan Research and Conservation Area, Rhoko Forest , Iko Esai , Cross River Loop , 18.V.2012. 6°39′27.20′N 8°15′42.16″E leg. Safian Sz. ’ ( SZS) ; 4♂♂ ‘ Ikom , 26.V.1974, leg. H. Politzar’ ( ZSM) ; 2♂♂ same data, but ‘ 26–28.V.1974 ’; 2♂♂ same data, but ‘ 22– 24.XII.1970 ’; 2♂♂ ‘ Jemaa , 16.X.1971, leg. H. Politzar’ ( ZSM) ; 2♂♂ ‘ Okomu , april 1992, A.A. Knoop’ ( RMCA) ; 3♂♂ ‘ Bendel State, Okomu Forest , 12.II.1984, leg. J. Wojtusiak’ ( ISEA) ; ♂ same data, but ‘ 9.06.1986 ’; ♂ ‘ Anambra State, Nsukka F. Res., 10.10.1982, leg. J. Wojtusiak’ ( ISEA) ; ♂ same data, but ‘17– 25.12.1982 ’; ♂ ‘ Ibadan , ca. Jan.-Juni 1954, H. Stenholt Clausen’ ( ISEA) . Cameroon: ♂ ‘ Eloumden , 27/ 28.10.1992, Th. Bouyer’ ( RMCA) ; ♂ ‘ Cameroun, 18.II.1962 ’ ( ISEA) ; ♂ ‘ Kounden , 27.12.1975, F. Puylaert’ ( RMCA) ; ♂ ‘Mount Cameroun, Mapanja , 9 8.4E 4 5.4N, 03/02/89, night 19.00–06.30, 1150 m, SE side Mt GoogleMaps Cameroon, 3,5 km NW of Mapanja. Trap on bottom of valley. Sub-mont forest , scrubby low veget.’ ( RMCA) ; 2♂♂ ‘Mount Cameroun, Bonjongo , 9 11.0E 4 3.8N, 30/01/89, night 19.15- 06.30, 460 m, Patch work of gardens of mixed crops abandoned gardens & berbaceous second growth. Trees from or. for.’ ( RMCA) GoogleMaps ; ♂ ‘ Mokundange , 16–30.VI.05., Tessmann S.G. ’ (type of Metarctia rubicundula ab. quadrisignatula Strand, 1912) ( ZMHB) ; ♂ ‘ Buea , 1–10.XI.10, 1000–1200 m, E. Hintz’ ( ZMHB) ; 2♂♂ ‘ Johann-Albrechtshöhe , 1/6.96, L. Conradt’ ( ZMHB) ; ♂ Südwest Provinz, Kupeberg, 2 km östlich Nyasoso , 1283 m, 4°49′28.8″N 9°42′6.6″O, 21 May 2009, Lichtfang, leg. M. Ochse’ ( ZSM); 8♂♂ ‘Südwest Provinz, Kupeberg, Nyasoso, Ortslage, 846 m, 4°50′5.4′N 9°40′52.9″O, 15.VII.–18.VIII.2008, Lichtfang, leg. Francis Ajebe Ngeh’ ( ZSM) ; ♂ ♀ in copula ‘ Südwest Provinz, Kupeberg, Nature Trail , Cave , 2 km östlich Nyasoso , 990 m, 4°49′24.4′N 9°41′15.4″O, 23.V.2009, Lichtfang, leg. Michael Ochse’ ( ZSM); ♀, Südwest Provinz, Zwischen Burutu und Echom, 244 m, Lichtfang, 4°46′51.0″N 9°36′13.6″O, 3 Dezember 2008, leg. M. Ochse’ ( ZSM) ; Equatorial Guinea: ♂ ‘ Insel Bioco ( Fernando Po), Ruiche 5 km S Luba, nördlicher Ortsrand bei 2. Schule , ca 740 mNN, verbuschende Bananenplantage , 3°24′50′ n. Br. 8°35′ ö.L., 21.VIII.1994, LF (160 W), leg. T. Karisch’ (TK) . DRC: ♂ ‘ P.N. A., Secteur Nord, Village Tunguru , 840 m, 13.VI.1953, P. Vanschuytbroeck & J. Kekenbosch’ ( RMCA) ; ♂ same data, but Mutwanga , 1200 m, 13.VI.1954, à la lumiere ; ♂ ‘ P.N. A., Massif Ruwenzori, Kalonge, 2060 m, riv. Katauleko affl. Butahu , 9.XII.1957, P. Vanschuytbroeck’ ( KBIN) ; ♂ ‘ Kibali-Ituri, Nioka , 8.VII.1952, J. Hecq’ ( RMCA) ; ♂ same data, but ‘ 8.VI.1953 ’; ♂ same data, but ‘ 18.X.1953 ’; ♂ ‘ Uele, Paulis , 8.X.1954, Dr M. Fontaine’ ( RMCA) ; ♂ same data, but ‘ 21.III.1957 ’; ♂ same data, but ‘ 14.12.1957 ’; ♂ ‘ N. Lac Kivu, Rwankwi , IV.1948, Mme J.V. Leroy’ ( RMCA) ; ♂ same data, but ‘ II.1958 ’; ♂ same data, but ‘ IV. 1958 ’; ♂ same data, but ‘ 25.II.1951 ’; ♂ ‘ Sankuru, Katako-Kombe , 14.VIII.1952, Dr M. Fontaine’ ( RMCA) ; ♂ ‘ Flandria , 1935, J. Ghesquière’ ( RMCA) ; ♂ ‘ Coquilhatville , 7.X.1913, L. Burgeon ( RMCA) ; ♂ ‘ Bambessa , 26.XI.1930, J. Vrydagh’ ( RMCA) ; ♂ ‘ Kafko, N. f. Rutshuru, O. Kongo , 13.XII.1946 ’ ( ISEA) ; 2♂♂ ‘ Prov. Equateur, Kalamba 55 km südl. Mbandaka , 3 km östlich Ort , 450 mNN, 0°25 s. Br. ; 18°19′ ö. L., 10.VIII.1991, LF 125 W HQL, T. Karisch legit’ (TK) ; Angola: 7♂♂ ‘ Prov. Nordcuanza, Canzele, 30 km nördl. Quiculungo , 18.X.1957, leg. G. Heinrich ( ZSM) . Rwanda: ♂ ‘ Nyungwe NP, 1800 m, 11 km N Uwinka, 2°25′S 29°09′E, 03.VIII.2008, leg. J.&W. De Prins’ ( RMCA) GoogleMaps . Burundi: 4♂♂ ‘ Kitega , 23.V.1963, 18.XI.1963, 13.XII.1963, 10.I.1965 leg. Dr M. Fontaine’ ( RMCA) . Uganda: 2♂♂ ‘ Kallinzu Rorest , 7–10.I.65, leg. J. Scheven’ ( ZSM) ; ♂ ‘ Gulu, 270 km N of Kampala , 18.02.1976, leg. K. Strzałka’ ( ISEA) ; Kenya: 3♂♂ Kakamega Forest , different dates and collecting places, (8♂♂ ISEA, 3♂♂ ZMHB, 13♂♂ RMCA, 6♂♂ ZSM, ♂ NHMO) . Ethiopia: ♂ ‘ Arba-Minch , 20.XI.1973, P.- C. Rougeot ( MNHN) .
The male of P. perpusilla is superficially very similar to the other taxa with black, transverse bands on the abdomen. From P. obscurus sp. nov., P. erubescens , and P. carnea it can be separated by the undivided blotchlike m 3 ( Fig. 2G–I View Figure 2 ). Furthermore, P. obscurus sp. nov. is characterized by the significant and easy to notice difference in the size of blotches m 4 and am 4, which are much smaller and often completely absent. Both blotches are more or less the same size in P. perpusilla . Pseudothyretes kamitugensis is also rather different from P. perpusilla . It is much larger and is characterized by a significantly more contrasting background coloration of the forewing. Pseudothyretes nigrita , which seems to be the most similar to P. perpusilla (also in morphology of labial palpus, Fig. 6G View Figure 6 ) never bears red or pink scales on the scapus, which is always concolorous with the uniformly coloured head. In case of doubt the male genitalia should be analysed ( Fig. 4G View Figure 4 ). The prominent, elongate lobes of the uncus (more than 2.5 times longer than wide) immediately separates P. perpusilla from all other taxa except P. nigrita . The uncus lobes of P. nigrita are not widened at the tip (they are narrow and rounded), and instead bear small processes in the middle. In contrast the lobes of P. perpusilla are slightly widened at the tip and do not have any protrusion in the middle. The phallus is devoid of cornuti ( Fig. 5H View Figure 5 ).
Description of female
Head: Frons and vertex rusty brown to buff; labial palpus concolorous, short, projecting slightly upwards, last segment bullet shaped ( Fig. 6L View Figure 6 ); scapus and basal segment of flagellum rusty brown; flagellum blackish, serrate, dorsal scales tawny, except for terminal six or seven segments on which they are white.
Thorax: Vestiture unicolorously rusty brown to buff; legs with intensive orange tinge from distal part of femur, sometimes extended to entire leg.
Abdomen: Dorsal and ventral segments unicolorous, pinkish buff to rusty; terga 2–6 with narrow black distal margins.
Forewing ( Fig. 3E, F View Figure 3 ): Length 20.0– 23.2 mm (average 21.8 mm, N = 11); upper side background colour buff to rusty, more intensive and darker around transparent blotches; pattern consisting of seven blotches with strongly marked margins (m 2 large, subsquare, in terminal region of DC, usually with concave outer margin; m 3 medium sized, elongate, between CuA 2 and 1A+2A in middle of dorsum, sometimes with separated am 3 in basal part; fm 3 small, triangular; m 4 medium sized, usually ovate; am 4 large, quadrangular; m 5 rather small, triangular; m 6 large, discoidal); fringe scales pale rusty brown; ventral background colour concolorous with dorsum, markings similar.
Hindwing: Background colour of dorsum and venter buff to pinkish rust; blotch d rather small, irregular, with diffuse margins, occupying distal portion of inner area of wing.
Female genitalia ( Fig. 4L View Figure 4 ): Papillae anales rectangular, covered with numerous setae along distal and dorsal margins; apophyses anteriores and posteriores sclerotized, of similar length; dorsal pheromone glands prominent, in form of paired irregular pouches with common opening, terminated in several irregular tapelike processes; ventral pheromone glands usually larger, in form of single wide pouch bearing several irregular tape-like processes; ostium bursae wide, discoidal; ductus bursae short, asymmetrical, directed slightly to the left, divided into three parts (anterior section shallow, weakly sclerotized, and fused with lamella postvaginalis; middle section indistinct, membranous, without plicae, convergent towards narrowed centre; posterior section membranous, very short); lamella antevaginalis invisible; lamella postvaginalis weakly sclerotized but forming distinct longitudinal ridge between paired invaginations located posterolaterally from ostium bursae; corpus bursae oval, membranous, without any trace of sclerotization or distinct plicae; signum absent; ductus seminalis slender, originating in basal half of corpus bursae.
As for all other Pseudothyretes species , the available biological and ecological data are very scarce and random. The food plants of the caterpillars as well as details of ontogenetic development remain unknown. The seasonal occurrence of the imago is also not completely understood, although some preliminary generalizations can be presented. Across most of the range imagoes are detected throughout the year, with the exception of July. This 1-month gap was already noticed by Przybyłowicz (2009). The present count based on more than 80 specimens confirms this pattern. It can be speculated that this period is a gap between generations, and that if imagoes were collected in July it would be exceptional ( Tjönneland, 1962). The higher abundance of moths attracted to light differs accross the regions that were prospected more intensively. Such estimations can also be also somewhat imprecise because of uneven collecting intensity during longer periods of time. Fieldwork in the Kakamega Forest ( Kenya) indicates that the highest occurence of moths is in April and May ( Przybyłowicz & Kühne, 2008). This observation confirms earlier data obtained by Tjönneland (1962) in Jinja (south-east Uganda). His year-round light-trapping efforts produced most specimens from February until July, with a sharp peak in April. In Ghana, however, most moths were collected from October until the end of December, with dominance in October. Tjönneland (1962) showed that moths are most abundant at light sources during the early part of the night, from 1 to 4 h after sunset.
The biotope information retrieved from labels suggests that the species is a generalist connected with different types of woodland and shrubby vegetation. Specimens have been also collected in devastated or highly anthropogenic biotopes such as abandoned plantations and rural gardens.
Pseudothyretes perpusilla is the most common and most widely distributed representative of the genus ( Fig. 7G View Figure 7 ). Its range spreads from Guinea in the west up to Ethiopia, Kenya, and Tanzania in the east. The southernmost localities are in northern Angola and south DRC. The record from Bioko ( Equatorial Guinea) makes it the only Pseudothyretes species colonizing an offshore island. Despite the lack of data from several countries from around the Guinea Bay (e.g. Togo, Benin, and Gabon), it can be assumed that the distribution of P. perpusilla is continuous and the ‘gaps’ result from insufficient penetration of such regions by scientists and collectors. The specimens studied have been collected mostly in the lowlands, as well as in the highlands, with the highest elevations not exceeding 2060 m a.s.l. (Kalonge, Ruwenzori, DRC).
A total of eight COI haplotypes were found among the 24 P. perpusilla specimens studied. The intraspecific haplotype diversity value was Hd = 0.837 and the nucleotide diversity amounted to π = 0.02033. Detailed information concerning the molecular variation of the COI fragment studied is presented in Table 3.
The correct assignment of the female was confirmed by a pair in copula collected at a light trap in Kupeberg, Cameroon, 23 May 2009. The body coloration and the wing pattern of P. perpusilla is the most variable of all members of the genus. Across the whole range various forms differing also in size and wing shape can be found. Unfortunately, the scarcity of the material does not allow for a more precise evaluation of this variability. The identity of all of the specimens studied has been confirmed by an examination of the genitalia, source of the most diagnostically reliable characters; however, further studies are needed to elucidate whether the variability has its origin in seasonal, geographic, or just random genetic variation. The substantial sample from Ghana may suggest a kind of mimicry between the co-occurring P. perpusilla and P. obscurus sp. nov. At the same locality (Guesthouse, Kakum National Park) and time (December) both taxa have been collected. Pseudothyretes perpusilla was present in two forms. One representing the typical pattern and coloration of the species, whereas the other was extremely similar to P. obscurus sp. nov. (rusty coloration, prominent m 1). The proper determination of such specimens was possible by the characteristic male genitalia, undivided m 3 (always divided in P. obscurus sp. nov.), proportion of m 4 to am 4, and results of the genetic analysis. The presence of such a rusty form is further confirmed by two females (Bunso Ar- boretum and surroundings of Elubo, Ghana), which superficially resemble females of P. obscurus sp. nov. more than P. perpusilla . Supportive evidence of heterogeneity within perpusilla is the topography of the phylogenetic tree ( Fig. 9). Samples N59, N67, and N68 form a small cluster of ‘rusty’ form separated from the main assemblage of typical Pseudothyretes samples. This observation should be further investigated with the help of more extensive material collected over a longer period of the year. Another possible explanation to explain this phenomenon is that there is a hidden species that currently falls under P. perpusilla . A way to elucidate the variability would be to analyse many more samples from West Africa. Sorting them by external and internal morphology and correlating this information with molecular characters would provide a more definitive taxonomic solution. Until that time, both ‘forms’ should be treated as one taxon.
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