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Behaviour and development of Pseudopontia gola Sáfián & Mitter, 2011 (Lepidoptera: Pieridae)

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Abstract

Information on adult behaviour and development of Pseudopontia gola Sáfián & Mitter, 2011 and foodplant records are presented in this paper, along with a short morphological description of its pre-imaginal stages. Although imagos within the genus Pseudopontia could not be distinguished based on macro-morphological features, there are small but clear morphological differences between the larva and pupa of P. gola and P. zambezi. From the limited number of records, larvae of P. gola seem to utilise a single foodplant species in the Opiliaceae, which differ from, but is related to those of P. paradoxa and P. zambezi. These differences also serve as further evidences of the specific status of P. gola. Resumen En este trabajo se presenta información sobre el comportamiento y desarrollo de los adultos de Pseudopontia gola Sáfián & Mitter, 2011 y sus plantas nutricias, así como una breve descripción de la morfología de los estados preimaginales. Aunque los imagos del género Pseudopontia no pueden distinguirse sobre la base de las características macro-morfológicas, hay pequeñas pero claras diferencias morfológicas entre las larvas y crisálidas de P. gola y P. zambezi. Desde un limitado número de registros, las larvas de P. gola parece que utilizan una sola planta nutricia de Opiliaceae, que es diferente, pero relativamente próxima a las utilizadas por P. paradoxa y P. zambezi. Estas diferencias también sirven como pruebas adicionales del estatus específico de P. gola.
85
Behaviour and development of Pseudopontia gola
Sáfián & Mitter, 2011
(Lepidoptera: Pieridae)
Sz. Sáfián
Abstract
Information on adult behaviour and development of Pseudopontia gola Sáfián & Mitter, 2011 and foodplant records are
presented in this paper, along with a short morphological description of its pre-imaginal stages. Although imagos within the
genus Pseudopontia could not be distinguished based on macro-morphological features, there are small but clear
morphological differences between the larva and pupa of P. gola and P. zambezi. From the limited number of records, larvae
of P. gola seem to utilise a single foodplant species in the Opiliaceae, which differ from, but is related to those of P. paradoxa
and P. zambezi. These differences also serve as further evidences of the specific status of P. gola.
KEY WORDS: Lepidoptera, Pieridae, Pseudopontia, foodplant, egg, larva, pupa.
Comportamiento y desarrollo de Pseudopontia gola Sáfián & Mitter, 2011
(Lepidoptera: Pieridae)
Resumen
En este trabajo se presenta información sobre el comportamiento y desarrollo de los adultos de Pseudopontia gola
Sáfián & Mitter, 2011 y sus plantas nutricias, así como una breve descripción de la morfología de los estados preimaginales.
Aunque los imagos del género Pseudopontia no pueden distinguirse sobre la base de las características macro-morfológicas,
hay pequeñas pero claras diferencias morfológicas entre las larvas y crisálidas de P. gola y P. zambezi. Desde un limitado
número de registros, las larvas de P. gola parece que utilizan una sola planta nutricia de Opiliaceae, que es diferente, pero
relativamente próxima a las utilizadas por P. paradoxa y P. zambezi. Estas diferencias también sirven como pruebas
adicionales del estatus específico de P. gola.
PALABRAS CLAVE: Lepidoptera, Pieridae, Pseudopontia, planta nutricia, huevo, larva, pupa.
Introduction
The butterfly subfamily Pseudopontiinae (Pieridae) was believed to be monophyletic for over 150 years
from the original description of the type species Pseudopontia paradoxa (FELDER & FELDER, 1869). A
distinct subspecies (ssp. australis) was, however, long ago recognised and was formally described by DIXEY
(1923). Based on specimens from throughout its range in the tropical forest zone of Africa, the status of P.
paradoxa was revised by MITTER et al. (2011). Three additional species were recognised: P. mabira Mitter &
Collins, 2011, P. zambezi Mitter & De Prins, 2011 and P. gola Sáfián & Mitter, 2011; while P. australis was
elevated to species rank, separating them from “real” paradoxa mainly on the basis of differences in their DNA.
The genetic data were also supported by the biogeography and in some cases minor morphological differences
were also revealed (MITTER et al., 2011). In the extensive literature of the Afrotropical butterflies only a few
notes were found about the larval foodplants and development of P. paradoxa (on the population from Ikelenge,
Zambia, which is now recognised as P. zambezi) (HEATH, 1977), and another short mention of a laying female
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in Sierra Leone (which pertains to P. gola) (OWEN, 1971). During extensive fieldwork in Sierra Leone and
Liberia the author studied the life-cycle of the newly described P. gola, including adult behaviour, pre-imaginal
stages and larval foodplant preferences; his field observations are compiled in this paper.
Material and methods
S
TUDY AREA AND HABITATS
Most field observations were documented in the wet lowland rainforests of Gola Forests, Western Liberia
and Eastern Sierra Leone. The various forest blocks of Gola Forests lie on both sides of the Moro River, which
forms the state-border between the two countries. These protected forest blocks are linked through a non-
protected ‘transboundary corridor’ area, which is a mixture of farmland, patches of younger and older secondary
forest and primary rainforest. This corridor is under preparation to be sustained in agreement between local
communities and the governments as a community nature reserve. The former Gola East, Gola West, Gola
North and Extension Two Forest Reserves are now merged under Gola Rain Forest National Park covering
approximately 75,000 hectares, while the size of forest on the Liberian side, the Gola National Forest, reaches
almost 100,000 hectares. The governments of Sierra Leone and Liberia wish to merge the protected forests by
forming the third largest protected forest in West Africa, west of the Dahomey Gap, next to Taï National Park
(3,000 km
2
) in Ivory Coast and Sapo National Park (1,800 km
2
) in Liberia.
D
OCUMENTATION
All details of adult behaviour of P. gola were observed under natural conditions in the field (the locations
and habitats are described above) by following selected individuals. For breeding, eggs and larvae were
collected with young shoots and leaves of foodplant; the caterpillars were kept in plastic food-storage boxes.
Various aspects of the life-cycle and behaviour of the species were documented by Canon EOS 40D digital
photo camera, equipped with 85-150 mm Canon zoom lens. The foodplant was identified by William
Hawthorne (Oxford University, UK) and Karel Jongkind (Wageningen University, The Netherlands) from
photographs, taken of both live individuals and pressed herbarium samples.
Results
A
DULT BEHAVIOUR
Imagos of P. gola fly very slowly low down in the forest interior, including dark places, similarly to P.
paradoxa as described in LARSEN (2005). They often congregate around a few flowering plants and bushes
available in the forest, sucking nectar as their food source (Figure 1). A number of specimens could often be
observed flying around individuals of the larval foodplant, especially near the ones with young shoots, when
they sometimes ascend above head-height. In fact, almost the entire life-cycle of the species is restricted to the
foodplant individuals, including even mating, occasionally.
Two different ways of courtship were observed in Liberia and western Sierra Leone. When the abundance
of P. gola is locally high, male individuals find freshly hatched females easily even in the dark understorey,
possibly from the reflection of UV light from the wings, as shown in MITTER et al. (2011). On a few occasions
males were observed competing for the acceptance of freshly hatched, still soft-winged females, who were
sitting and drying themselves on the pupal skeleton, still unable to fly. Sometimes three or four males were
aggressively pushing away each other from the female for quite a long time before succeeding. Successful
mating with still soft-winged females was also observed (Figure 2). When abundance is generally low and
females have a chance to dry themselves before mating, the courtship usually begins with a joint flight, during
which the male is possibly gathering information on the receptiveness of the female, also giving signs of
attempts at mating. If the female is ready to accept the approach of the male, it settles on low plants in the dense
undergrowth with wings horizontally open (Figure 3), while the male starts a long-lasting flapping flight right
SZ. SÁFIÁN
86 SHILAP Revta. lepid., 43 (169), marzo 2015
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above the female (it often lasts over 10 minutes). This flight will eventually turn into mating but males
sometimes lose direction during this “hovering in slow motion”, since they were also seen being chased away
by competitors. In these cases the females remain still for a couple of minutes keeping their wings completely
horizontal, and if the male does not find its way back, they simply fly away.
No information could be collected about the time of egg-maturation after mating.
The eggs are usually laid singly only on the very fresh shoots of foodplant, with no or with very small
young leaves. Terminal shoots are preferred, even if they are higher (up to 2-3 metres) from the ground.
Foodplants with only older leaves and shoots are almost completely avoided. When no fresh shoots are
available, still relatively young leaves are also accepted, in which case the females lay on the underside of the
leaf on veins. It is often the case that multiple females lay on the same plant.
L
ARVAL FOODPLANT
Larval foodplant records were collected both as photos and plant samples for further identification from
three different localities: 1. Money Camp, Gola National Forest, Liberia, 2. Mogaima, Corridor Area between
Gola North and the Liberian border in Sierra Leone, 3. Trail from Madina to Waama, buffer zone to Gola North,
Sierra Leone. In all three localities, P. gola utilised Urobotrya congolana ssp. afzelii (Engl.) P.Hiepko as larval
foodplant, a rainforest interior-dwelling shrub endemic to West Africa west of the Dahomey Gap (Figure 4). Its
distribution covers largely the Liberian biogeographical sub-region (including parts of Guinea and Sierra Leone,
almost the whole of Liberia and Western Ivory Coast, just penetrating to South-Western Ghana in Ankasa
National Park as a disjunct occurrence), which largely overlaps the known range of P. gola and grows in the
undergrowth of wet rainforests (HIEPKO, 2008, Hawthrorne pers. comm., Jongkind pers comm.).
Previously, the larval foodplant was believed to be Pseuderanthemum tunicatum (Afzel.) Milne-Redh.
Based on egg-laying observations by OWEN (1971), this now seems very unlikely, as this plant belongs to
Acanthaceae, while other species in the genus Pseudopontia also utilize members of the Opiliaceae family:
Pentarhopalopilia marquesii (Engl.) Hiepko by P. zambezi (HEATH, 1977) and Rhopalopilia pallens Pierre by
P. paradoxa (LEES, 1989).
E
GG
The eggs are oval in shape and slightly conical at the tip; their colour is shiny white. The size is 1 mm. The
eggs hatch in 2-4 four days after laying.
L
ARVA
The freshly hatched larva is about a millimetre long, almost transparent light green. The larva changes its
colour to light grass-green in the second instar, and it remains very similar in colour and pattern until pupation.
The fully developed larva is 18 mm in length, the body’s ground colour is grass-green with a white lateral stripe,
which stretches from the head to the last segment, including a pair of short protuberances on the last segment,
which are also white. The legs and all pro-legs are glossy light green. The dorsal surface is creased transversely.
The entire body is densely scattered with short setae, including the head and the pro-legs (Figures 5-6). A lateral
line is present, but is pale yellow on the fully grown larvae of P. zambezi, the protuberances and the anal pro-
legs are bright yellow and are bare in P. zambezi (HEATH, 1977).
P
UPA
The shape of the pupa is very similar to those of Colotis (mentioned also for P. zambezi (=Paradoxa
australis) (HEATH, 1977) with the characteristic rounded wingcase on the abdominal side, although it is
significantly smaller than many Colotis species (the length is 12-14 mm). The colour is glossy grass-green
(Figure 6). This glossiness was not mentioned in the description of P. zambezi (P. paradoxa australis) (HEATH,
1977), neither was a blackish lateral line (stretches from the head to the cremaster along the inner edge of the
BEHAVIOUR AND DEVELOPMENT OF PSEUDOPONTIA GOLA SÁFIÁN & MITTER, 2011
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wings) on the pupa, in which it differs from P. zambezi (Figures 2, 6). The pupa of P. gola also lacks the short
setae of that of P. zambezi. The recorded pupal stage was 7 days (6 days was recorded by HEATH (1977) for P.
zambezi in captivity).
Discussion
Imagos within the genus Pseudopontia cannot be distinguished based on macromorphological features.
The adult behaviour too, does not differ much from those observed for other species (no information about
courtship is available for other species). However the larval foodplant of the newly described P. gola differs
from though related to those of P. zambezi and P. paradoxa. The larva and pupa of P. gola and P. zambezi (the
only other species, where pre-imaginal stages are known) also differ in various small, but important features of
colour and pattern as well as in the lack of setae in the pupal stage in P. gola. These differences serve as further
evidence of the specific status of P. gola, which could not be presented at the time of description.
Acknowledgement
The author is grateful to the “Across the River” a - Transboundary Peace Park for Sierra Leone and Liberia
project for supporting Lepidoptera research in the Gola Forests. Dr. Annika Hillers was not only a great research
coordinator but she sacrificed herself, carrying shoots of foodplant with P. gola eggs for over ten kilometres
during trekking out from Mogbaima. Thanks to all research technicians, namely Mohamed Lumeh, Jerry C.
Garteh, Daniel K. Harris, Gbessay E. S. Momoh, Mohamed Koroma who ensured that the efficiency of the
research was always at its maximum. William Hawthorne (Oxford University, UK) and Karel Jongkind
(Wageningen University, The Netherlands) helped with the identification of foodplant samples and with
providing further information about U. congolana ssp. afzelii.
BIBLIOGRAPHY
DIXEY, F. A., 1923.– Pseudopontia paradoxa; its affinities, mimetic relations and geographic races.– Transactions of the
Entomological Society of London, 1922: 61-64.
HEATH, A., 1977.– Early stages of a remarkable species of African Pieridae.– Arnoldia Zimbabwe, 8(16): 1-3.
HIEPKO, P., 2008.– Opiliaceae, Species Plantarum.– Flora of the World, 12: 71 pp.
LARSEN, T. B., 2005.– Butterflies of West Africa: 595 pp + 135 plates. Apollo Books, Svendborg.
LEES, D. C., 1989.– A feasibility study into butterfly farming for the Korup National Park, Cameroon. WWF UK / WWF
Korup.
MITTER, K. T., LARSEN, T. B., DEPRINS, W., DE PRINS, J., COLLINS, S. C., VANDE WEGHE, G., SÁFIÁN, SZ.,
ZAKHAROV, E., HAWTHORNE,D. J., REGIER, J. C. & KAWAHARA, A. Y., 2010.– Genetic diversity in
Pseudopontia (Lepidoptera: Pieridae: Pseudopontiinae): Evidence for multiple species.– Systematic Entomology DOI:
10.1111/j.1365-3113.2010.00549.x.
OWEN, D. F., 1971.– Pseudopontia paradoxa, a unique butterfly of the rain forest of West and Central Africa.– Biological
Conservation, 4(1): 73-74.
Sz. S.
Institute of Silviculture and Forest Protection
University of West Hungary
Bajcsy-Zsilinszky utca, 4
H-9400 Sopron
HUNGRÍA / HUNGARY
E-mail: safian@bcghana.org
(Recibido para publicación / Received for publication 17-XI-2013)
(Revisado y aceptado / Revised and accepted 28-II-2014)
(Publicado / Published 30-III-2015)
SZ. SÁFIÁN
88 SHILAP Revta. lepid., 43 (169), marzo 2015
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BEHAVIOUR AND DEVELOPMENT OF PSEUDOPONTIA GOLA SÁFIÁN & MITTER, 2011
SHILAP Revta. lepid., 43 (169), marzo 2015 89
Figs. 1-6.– 1-2. The main food source of P. gola is nectar from various flowers and flowering shrubs (Photo:
Szabolcs Sáfián, left). Violent courting resulted in successful mating with a (still soft-winged) freshly hatched
female (Photo: Szabolcs Sáfián, right). 3-4. During the long-lasting courting females sit still in the dark
undergrowth with horizontally open wings (Photo: Annika Hillers, left). Urobotrya congolana ssp. afzelii -
foodplant of P. gola in flower (Photo: Szabolcs Sáfián, right). 5-6. L4 stage larva of P. gola (Photo: Szabolcs
Sáfián, left). Pre-pupation stage larva and pupa of P. gola (Photo: Szabolcs Sáfián, right).
1 2
3 4
5 6
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... Pseudopontiinae is a monotypic subfamily of Ethiopian distribution with five described species (Mitter et al. 2011, Sáfián 2015, two of which are used in this paper (appendix), P. paradoxa and P. gola ANTENNAE: The antenna is very short compared to the costal margin of the forewing, length ca. 2.3 mm (2267 µm), and thin. ...
... Based on these characters, Pseudopontia cannot be basal to the rest of Pieridae, the sister-group to Pieridae, or the sister-group to Dismorphiini. It appears to be a more recent and specialized branch of Coliadinae based on the type of pupa (Sáfián 2015, Dias et al. 2016). Pseudopontia may have arisen earlier than Coliadinae and Pierinae (Wahlberg et al. 2014) as the ancestor of both subfamilies. ...
... Based on antennae, our results indicate that Pseudopontia exhibit more derived features than Dismorphiinae genera, a conclusion that was hinted at by Jordan (1898). The venation, pupa, some genes, and host plants showed or implied that, usually, Pseudopontia characters are not plesiomorphic with respect to Dismorphiinae, but apomorphic (Grote 1900, Klots 1931-1933, Clench 1955, Jong et al. 1996, Vane-Wright 2003, Wahlberg et al. 2014, and Sáfián 2015. Derived character states of Pseudopontia include almost completely scaleless antennae, the presence of different sulci, and a large, continuous outline in all antennomeres the antennal club. ...
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We examined antennal ultrastructure in species of Dismorphiinae and Pseudopontiinae (Pieridae) using scanning electron microscopy (SEM). We examined two species of Pseudopontia (Pseudopontiinae) and more than 30 species in seven genera of two tribes in the subfamily Dismorphiinae: Leptidea, Enantia, Pseudopieris, Lieinix, Moschoneura, Dismorphia, and Patia. We focused on the scaleless terminal area of the antenna where there are many types of sensilla, some of which are clustered together and constitute specialized organs. We measured, described, and illustrate at different magnifications structures including the antennal club, antennomeres, sulci, pseudosulci, and chaetic, trichoid, coeloconic, basiconic, and auriculate sensilla, as well as other previously unnamed sensilla. From these antennal features, we created a matrix of characters that allowed us to recognize divergence between the tribes Leptideini and Dismorphiini. The antennae of Leptideini have fewer scaleless antennomeres in the antennal club than those of Dismorphiini, a greater number of pseudosulci disaggregated or dispersed (in Leptidea), and fewer types of sensilla and microtrichia (a more homogeneous antennal morphology), as well as a reduction in the density of sensilla. In Leptidea the antennal form is more specialized: it is shorter in comparison to genera of other Papilionoidea families. We also created a matrix of general morphological characters of Dismorphiinae and Pieridae from the taxonomic literature. This matrix confirms the marked character divergence between the tribes and allows for a more meaningful discussion regarding the relationships between Dismorphiinae and the other subfamilies of Pieridae (i.e., Pseudopontiinae, Coliadinae, and Pierinae). We argue that Pseudopontiinae cannot be considered the least derived subfamily among Pieridae because pupal features, wing venation, and antennal characters exhibit a combination of primitive and specialized states. In addition, we discuss our results from the perspective of patterns of food plant usage in pierid subfamilies, specifically the diversification of several Dismorphiinae genera on Hologalegina and Ingeae (Fabaceae).
... A complete analysis can be found in Castro-Gerardino & Llorente-Bousquets (2017). Taxonomic implications of the studies that include Pseudopontia (e.g., Mitter et al. 2011, Sáfián 2015 provide further evidence that Pseudopontiinae is more derived than Dismorphiinae. Chorionic studies in species of Pseudopontia, such as those conducted in Coliadinae by Nieves-Uribe et al. (2016a, b, c), could be of conbsiderable phylogenetic interest. ...
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We present the results of the first examination and comparative analysis of the ultrastructure of the antennae in seven species of Leptidea Billberg, 1820 (Pieridae). The results add further support to the hypothesis that the subfamily Dismorphiinae is composed of the tribes Leptideini and Dismorphiini. We summarize the results in a table and discuss the ultrastructure similarities and differences between the two tribes. Following a previously proposed framework, we trace the progression of character states and discuss the significance of the results in the context of varying phylogenetic hypotheses for Leptidea, with special emphasis on biogeography. We include a brief description of the terms we employ to described the main characters and types of sensilla of the antennal club of Leptidea. Remarkable antennal sexual dimorphism is noted in L. gigantea.
Article
The Afrotropical butterfly subfamily Pseudopontiinae (Pieridae) was traditionally thought to comprise one species, with two subspecies (Pseudopontia paradoxa paradoxa Felder & Felder and Pseudopontia paradoxa australis Dixey) differing in a single detail of a hindwing vein. The two subspecies also differ in their known geographic distributions (mainly north of versus south of the equator). Unlike most butterflies, Pseudopontia is white with no visible wing or body markings. We now report that males of P. paradoxa australis have an area of ultraviolet-reflecting scales along the anal vein of the forewing, whereas males of P. paradoxa paradoxa and all females do not. A total of 21 individuals of the northern subspecies, which were collected in three localities south of the equator, were found in the collection of the Royal Museum for Central Africa, indicating sympatry of the two traditional subspecies in the Congo River basin. To determine if additional cryptic species might be present, we sequenced three nuclear genes (CAD, DDC and wingless) as well as cytochrome oxidase I (COI), examined amplified fragment-length polymorphisms, and re-examined wing and genitalic morphology, using recently collected specimens from several regions of Africa. Phylogenetic analyses of the COI sequences and amplified fragment-length polymorphism data concur, and indicate the existence of at least five monophyletic, non-interbreeding populations, with a particularly deep divergence between three populations of P. paradoxa paradoxa and two of P. paradoxa australis. Despite the slow rate of evolution of the nuclear genes studied, individual gene trees and a concatenated three-gene tree demonstrate, with high bootstrap support, clear divergence among the five populations of Pseudopontia. In addition, consistent variations in details of wing vein stalks were found among four of the genetically distinct populations, which supports the hypothesis of multiple species. Division of Pseudopontia into five phylogenetic species is proposed, including the elevation of ssp. australis to species rank and the description of Pseudopontia mabira, Pseudopontia gola and Pseudopontia zambezi
Early stages of a remarkable species of African Pieridae
  • A Heath
HEATH, A., 1977.-Early stages of a remarkable species of African Pieridae.-Arnoldia Zimbabwe, 8(16): 1-3.
– Butterflies of West Africa: 595 pp + 135 plates– A feasibility study into butterfly farming for the Korup National Park, Cameroon
  • P Hiepko
  • Apollo Books
  • Svendborg
  • D C Lees
  • Wwf Uk / Wwf Korup
  • K T Mitter
  • T B Larsen
  • W Deprins
  • J De Prins
  • S C Collins
  • G Vande Weghe
  • Sz Sáfián
  • E Zakharov
  • D J Hawthorne
  • J C Regier
  • A Y Kawahara
HIEPKO, P., 2008.– Opiliaceae, Species Plantarum.– Flora of the World, 12: 71 pp. LARSEN, T. B., 2005.– Butterflies of West Africa: 595 pp + 135 plates. Apollo Books, Svendborg. LEES, D. C., 1989.– A feasibility study into butterfly farming for the Korup National Park, Cameroon. WWF UK / WWF Korup. MITTER, K. T., LARSEN, T. B., DEPRINS, W., DE PRINS, J., COLLINS, S. C., VANDE WEGHE, G., SÁFIÁN, SZ., ZAKHAROV, E., HAWTHORNE, D. J., REGIER, J. C. & KAWAHARA, A. Y., 2010.– Genetic diversity in Pseudopontia (Lepidoptera: Pieridae: Pseudopontiinae): Evidence for multiple species.– Systematic Entomology DOI: 10.1111/j.1365-3113.2010.00549.x. OWEN, D. F., 1971.– Pseudopontia paradoxa, a unique butterfly of the rain forest of West and Central Africa.– Biological Conservation, 4(1): 73-74.
Mohamed Koroma who ensured that the efficiency of the research was always at its maximum The Netherlands) helped with the identification of foodplant samples and with providing further information about U. congolana ssp– Pseudopontia paradoxa; its affinities, mimetic relations and geographic races
  • Daniel K Garteh
  • Harris
  • E S Gbessay
  • Uk Momoh
  • Karel Jongkind
Garteh, Daniel K. Harris, Gbessay E. S. Momoh, Mohamed Koroma who ensured that the efficiency of the research was always at its maximum. William Hawthorne (Oxford University, UK) and Karel Jongkind (Wageningen University, The Netherlands) helped with the identification of foodplant samples and with providing further information about U. congolana ssp. afzelii. BIBLIOGRAPHY DIXEY, F. A., 1923.– Pseudopontia paradoxa; its affinities, mimetic relations and geographic races.– Transactions of the Entomological Society of London, 1922: 61-64.
Institute of Silviculture and Forest Protection University of West Hungary Bajcsy-Zsilinszky utca
  • S Sz
Sz. S. Institute of Silviculture and Forest Protection University of West Hungary Bajcsy-Zsilinszky utca, 4
Pseudopontia paradoxa; its affinities, mimetic relations and geographic races
  • F A Dixey
DIXEY, F. A., 1923.-Pseudopontia paradoxa; its affinities, mimetic relations and geographic races.-Transactions of the Entomological Society of London, 1922: 61-64.
  • P Hiepko
HIEPKO, P., 2008.-Opiliaceae, Species Plantarum.-Flora of the World, 12: 71 pp.